SOURCE OF CURRENT RESEARCH IN Ao EY A ee! : NA Barney L. Lipscomes, Epitor Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, USA 817 332-4441 / 817 332-4112 FAX Electronic mail: sida@brit.org Home page at the URL: http://www.brit.org/sida/ SiDA CONTRIBUTIONS TO BOTANY WAS FOUNDED BY JOHN W. THIERET, ASSOCIATE EDITOR Lovo H. SHINNERS (LEFT) Dept. of Biological Sciences Northern Kentucky Universit IN 1962. INHERITED BY , Highland Heights, KY 41099 USA WILLIAM F. MAHLER (RIGHT), DiRECTOR EMERITUS Feuix LLAMAS, CONTRIBUTING SPANISH EDITOR oF BRIT in 1971, ano Dpto. de Botanica, Facultad de Biologia Universidad de Leon SINCE 1993, IT HAS BEEN E-2471 Ledn, SPAIN PUBLISHED BY BRIT PRESS. Guidelines for contributors are available upon request and on our SIDA home page as well as the last pages of each volume. =) Subscriptions for year 2004: $35. Individual $60. USA Institutions $70. Outside USA numbers issued twice a year OSIDA, CONTRIBUTIONS TO BOTANY VOLUME 21, NumBeR 1, PAGES 1-510. 20 AuGcust 2004 COPYRIGHT 2004 BOTANICAL RESEARCH INSTITUTE OF TEXAS PRINTED IN THE UNITED STATES OF AMERICA r- PRESS ISSN 0036-1488 TABLE OF CONTENTS SYSTEMATICS A new species of Leucophyllum (Scrophulariaceae) and comments on relationships of the genus JAMES HENRICKSON Amaranthus acanthobracteatus (Amaranthaceae) JAMES HENRICKSON Taxonomic reevaluations in North American Erigeron (Asteraceae: Astereae Guy L. Nesom Notes on North American arctic and boreal species of Erigeron (Asteraceae: Astereae) Guy L. Nesom and Davip F Murray Notes on typification in Pluchea (Asteraceae: Plucheae) Guy L. Nesom A review of Crataeg iesR lifoliae (Rosaceae) in western Canada J.B. Puipes and R.J. O7KENNON Croton bigbendensis (Euphorbiaceae), a new species from Trans- Pecos, Texas B.L. TURNER A new species of Centaurium (Gentianaceae) from Trans-Pecos Texas B.L. TURNER A new combination in Dendrocalamus (Poaceae: Bambusoideae) MUKTESH KuMAR, REMESH M., and N. UNNIKRISHNAN Una nueva ie de Struthanthus (1 tl ) para Costa Rica Luts A. GONZALEz y J. FRANCISCO MorRALES The Gonolobus complex (Apocynaceae: Asclepiadoideae) in the southeastern United States ALEXANDER Krincs and (JENNY) QIu-YUN XIANG Polyphyly of the genus Echites (Apocynaceae: Apocynoideae: Echiteae): evidence based on a morphological cladistic analysis JUSTIN K. WILLIAMS Allotoonia, a new neotropical genus of Apocynaceae based on a subgeneric segregate of Echites J. FRANcisco Moraces and Justin K. WILLIAMS Estudios en las ADOC MASEL neotropicales IV: notas taxonomicas en Prestonia (Apocynoid hiteae) con una nueva especie de Ecuador J. FRANCISCO MORALES Estudios en las Apocynaceae neotropicales V: una nueva especie nuevos reportes y nueva sinonimia en las Apocynaceae de Bolivia J. FRANCISCO MorALes y ALFREDO FUENTES b) 93 97 103 117 Pseudostellaria oxyphylla (Caryophyllaceae), a long overlooked species from northern Idaho RONALD L. HARTMAN and RICHARD K. RABELER The genus Prenanthes (Asteraceae: Lactuceae) in Texas Jason R. SiNGHURST, RoBERT J. OKENNON, and WALTER C. HOLMES Two new subspecies of Microseris laciniata (Asteraceae) from the Siskiyou Mountains KENTON L. CHAMBERS New taxa and new combinations in North American Cirsium (Asteraceae: Cardueae Davip J. KEIL Anew name and anew combination in Solidago subsect. Glomeruliflorae (Asteraceae: Astereae RACHEL E. Cook and JOHN C. SEMPLE Taxonomic notes on Krigia (Asteraceae) KENTON L, CHAMBERS New combinations in North America Eremogone (Caryophyllaceae) RONALD L. HartMAN and RICHARD K. RABELER New combinations in the genus Gundlachia and four new genera of Astereae (Asteraceae) from northern Mexico and the southern United States Lowe Lt E. UrsAtscH and ROLAND P. ROBERTS A new species of Calycadenia (Asteraceae) from north central California Ropert L. CARR and GERALD D. Carr Two new varieties of Agoseris (Asteraceae: Lactuceae) Gary |. Bairb Lectotypification of Passiflora affinis (Passif loraceae) and discussion of its geographic range within the United States DouGias H. GOLDMAN Thymophylla setifolia var. greggii (Com positae) JOHN L. STROTHER Two new combinations in the genus Packera (Asteraceae) Desra K. TROCK A new combination in Persicaria (Polygonaceae) Cralc C. FREEMAN An evaluation of Anthenantia (Poaceae) ROBERT KRAL Calamagrostis coahuilensis and C. divaricata (Poaceae: Pooideae: Agrostidinae), two new species from México Paut M. PETERSON, ROBERT J. SORENG, and JESUS VALDES-REYNA 175 ETHNOBOTANY Ethnobotany of Rhodiola rosea (Crassulaceae) in Norway TORBJORN ALM Notes on “coffee” from the Kentucky coffeetree (Gymnocladus dioicus, Fabaceae JOHN P. SpaetTH and JOHN W. THIERET Reflections on the taxonomy and distribution of medicinal flowers of Pakistan MOHAMMAD ATHAR and M. AKMAL SIDDpIQI BOTANICAL HISTORY Cuscuta (Convolvulaceae)—the strength of weakness: a history of its name, uses and parasitism concept during ancient and medieval times Minal Costea and Francois J. TARDIF History and eponymy of the genus name Amsonia (Apocynaceae) JAMES S. PRINGLE FLORISTICS AND ECOLOGY Carbon use by the endangered Texas wild rice (Zizania texana, Poaceae) PauLaA Power and Rosert D. DoyLe The Post Oak Savanna ecoregion: a floristi fOr its uniqueness MicHAEL H. MAcRoserts and BARBARA R. MACROBERTS A quantitative study of the vegetation surrounding yellow lady-slip- per orchid (Cypripedium kentuckiense, Orchidaceae) populations at Fort Polk in west central Louisiana CHARLES M. ALLEN, SARA THAMES, SPENCER TRICHELL, and JEREMY WHITE Changes in baygall vegetation from 1986 to 2001 at Fort Polk in west central Louisiana CHARLES M. ALLEN, JOHN PATE, SARA THAMES, SPENCER TRICHELL, and Lacy EZELL Vascular flora of Hackberry Flat, Frederick Lake, and Suttle Creek, Tillman County, Oklahoma Bruce W. HoAGLanpb, PrisciLLA H.C. CrAwrorb, PHILLIP T. CRAWFORD, and FORREST JOHNSONT Sawtooth oak (Quercus acutissima, Fagaceae) in North America ALAN T. WHITTEMORE Sporobolus coahuilensis (Poaceae): a new record for the U.S.A from Trans-Pecos, Texas B.L. TURNER New reports of Eurybia and Aster s.str. (Asteraceae: Astereae ) from California, Idaho, and Wyoming Luc BROUILLET Notes on the distribution of Pseudognaphalium luteoalbum (Asteraceae: Gnaphalieae) Guy L. Nesom Dichanthelium scoparium and Muhlenbergia glabrifloris: new to the flora of Ohio RicHarp L. GARDNER, JAMES S. McCormac, and Davip MINNEY New records of Erigeron (Asteraceae: Astereae) for Nevada Guy L. NEsoM A putative hybrid of Schoenoplectus saximontanus and S. hallii (Cyperaceae) from Oklahoma MARIAN Situ, PAuL McKenzie, PAIGE METTLER-CHERRY, and GALEN SMITH Additions to and noteworthy records for the vascular flora of West Virginia Cyntuia M. Morton, JOHN Kartesz, BONNIE IsAAc, and ROBERT COXE Phalaris arundinacea (Poaceae: Aveneae) a species new to Texas anda key to Phalaris in Texas STEPHAN L. Hatcu, DALE A. Kruse, and JENNIFER PLUHAR Silene conoidea (Caryophyllaceae) new to Texas MONIQUE DUBRULE REED Noteworthy collections of Cyperus drummondii(Cyperaceae) from Texas Davip J. ROSEN Ruppia cirrhosa (Ruppiaceae) in north central Texas TIFFANY L. MorGAN and Walter C. HOLMES Osmorhiza bipatriata (Apiaceae) in Texas: servation considerations Jun Wen, Porter P. Lowry II, and JAmes C. ZECH status and con- First record of Clitoria mariana (Leguminosae) in western Texas BRIAN P. OswaLb, MOHAMMAD BATAINEH, and AMANDA ROUNTREE 507 Book reviews AND Notices 10, 18, 40,58, 78, 86, 92, 132, 164, 180, 192, 206, 220, 242, 258, 266, 286, 288, 388, 408, 418, 428, 446, 458, 509 Announcements 462, 472 Index to new names and new combinations in Sida 21(1), 2004 Agoseris eer geen var. leptophylla GL. Baird, var. nov.—267 Agoseris heterophylla var. quent GL. aird, var. nov.— Allotoonia J. F ees &) K. Williams, _ nov.— sain es Lae ae Jacq.) J.F es & J.K. Williams, comb. nov.— ae Allotoonia caudata (Woodson) J.F. Morales, comb. nov.—145 amie jdaatsnis (Sessé & Moc.) J.F K. Williams, comb. aa Allotoonia ‘urinal (Woodson) _ F es & J.K. Williams, comb no = Allotoonia tuxtlensis (Standl.) J.F Morales & J.K. Williams, comb. Aneeaniae acanthobracteatus Hen- rickson, sp. nov.— Anthenantia texana R. Kral, sp. nov.—296 Calamagrostis coahuilensis P.M. Peter- son, Soreng & Valdés-Reyna, sp. nov.—312 Calamagrostis divaricata PM. Peterson oreng, sp. nov.—315 Calycadenia micrantha R.L. Carr & . Cart, sp. nov.—261 Centaurium ee B.L. Tur . nov.— Gaiam Uroaicch pat RP. Roberts, Chihuahuana purpusii Sopa ae Urbatsch & R.P. Roberts, comb nov.— Cirsium arizonicum var. bipinnatum (Eastw.) DJ. Keil, comb. nov.—209 a ee var. chellyense (RJ. Ore str ature DJ. Keil, cat et stat. nov.— eesna arizonicum Var. een (A. Gray) DJ. Keil, comb. et stat. ov.—210 Cirsium arizonicum var. tenuisectum . Keil, var. nov.—210 Cirsium clavatum var. americanum (A. Gray) DJ. Keil, comb. nov.—211 Cirsium clavatum var. osterhoutii (Rydb.) — Keil, comb. et stat. nov.— Cirsium cymosum var. canovirens (Rydb.) DJ. Keil, comb. et stat. 2 Cirsium eatonii var. clokeyi (S.F Blake) DJ. Keil, comb. et stat. nov.—212 Cirsium eatonii var. eriocephalum (A. Nelson) DJ. Keil, comb. nov.—212 Cirsium eatonii var. hesperium (Eastw.) Keil, comb. et stat. nov.—212 Cirsium eatonii var. peckii (L.F Hend.) eil, comb. et stat. nov.—212 Cirsium eatonii var. viperinum DJ. Keil, Cirsium edule var. macounii (Greene) J. Keil, comb. et stat. nov.—213 Cirsium edule var. wenatchense DJ. Keil, var. nov.—21 Cirsium horridulum var. megacanthum (Nutt.) DJ. Keil, comb. et stat. nov.—214 Cirsium in var. davisii (Cronquist) DJ weil comb. et nov.— Cirsium i inamoenum (Greene) DJ. Keil, comb. nov.— Cirsium occidentale var. lucianum DJ. il, var. nov.—214 Cirsium Gclvoce niin var. martinii (P. Barlow-Irick) DJ. Keil, comb. et Stat. nov.—21 Cirsium Jaevieatinena var. aridum R.D. Dorn) a Keil, comb. et stat. nov.—21 Cirsium scariosum var. americanum (A. Gray) DJ. Keil, comb. nov.—215 Cirsium scariosum vat. citrinum (Petrak) DJ. Keil, ed nov.— Cirsium eats ar. coloradense SS dip Dy. Keil, ab. et stat. ae Girciuan pero var. congdonii ore & Frankton) DJ. Keil, on et stat. nov.—215 Cirsium scariosum var. robustum D.]. ell, var. nov.—215 Cirsium scariosum var. toiyabense D.]. ell, var. nov.— Crataegus chrysocarpa Var. vernonensis J.B.Phipps & O’Kennon, var. nov.—73 Crataegus sheila-phippsiae J.B. oe J O’Ke nov.— — eg a Bal ae sp. —79 Dendrocalamus stocksii (Munro) M. Kumar, Remesh & Unnikrish- nar oe nov.— Eremogone capillaris var. americana (Maguire) R.L. Hartman & Rabeler, comb. nov.—239 Eremogone congesta var. cephaloidea b.) R.L. Hartman & Rabeler, mb. nov.—239 Eremogone congesta Var. charlestonensis (Maguire) R.L. Hartman & Rabeler, comb. nov.— Eremogone congesta var. crassula Maguire) R.L. Hartman & Rabeler, comb. nov.—239 Eremogone congesta var. glandulifera (Maguire) R.L. Hartman & Rabeler, comb. nov.—239 Eremogone congesta var. 2 ogee he) Renta Rabeler, comb. ane Eremogone congesta var. simulans Maguire) R.L. Hartman & Rabeler, comb. noy.—239 Eremogone congesta var. oe S. Watson) R.L. Hartman & Rabeler, comb. ope ee Eremogone congesta var. suffrutescens ( ra rtman @ Rabeler, comb. nov.—239 Eremogone congesta var. wheelerensis Maguire) R.L. Hartman & Rabeler, comb. nov.—240 aa 28 eastwoodiae var. adenophora a & Peebles) R.L. Hart- & Rabeler, comb. nov.—240 Ere sabeone eraay (Dougl. ex Hook.) man & Rabeler, comb. arene Eremogone franklinii var. thompsonii (M. Peck) R.L. Hartman & Rabeler, comb. nov. Eremogone hineii var. plateauensis (Maguire) R.L. Hartman & Rabeler, et nov.—240 enc kingii var. rosea (Maguire) R.L. Hartman & Rabeler, comb. et stat. nov.—240 Eremogone macradenia var. arcuifolia Maguire) RL. Rabeler, comb. nov.— Eremogone macradenia vat. ferrisiae (Abrams) R.L. Ha & Rabeler, comb. et stat. nov.—240 Eremogone macradenia var. kuschei (Eastw.) R.L. Hartman & Rabeler, comb. noy.—240 Erigeron clokeyi var. pinzliae Nesom, var. nov.—28 Erigeron davisii Sa Nesom, comb. et Erigeron bare a nom. nov.—28 Erigeron klamathensis (Nesom) ean. comb. et stat. nov.—20 Erigeron poliospermus var. disciformis oe onq.) Nesom, comb. et stat. —24 Peern porsildii Nesom & Murray, om. no Erigeron robustior eae Nesom, comb. et stat. nov.—2 Gundlachia diffusa (Benth.) Urbatseh & R.P Roberts, comb. no ee riskindii (B.L. Turner & gford) ee - R.P. Rob- ares comb. nov. Gawiiaen 4 te ‘ F Blake) Urbatsch & R.P. Roberts, comb. vi— Gundlachia truncata (G.L. Nesom) Urbatsch & R.P. Roberts, oak. nov. eo56 Krigia cespitosa var. gracilis (DC.) KL. Chambers, stat. nov.— Krigia x shinnersiana K.L. Chambers, brid nov.— Leucophyllum coahuilensis Henrickson, sp. nov.— Medranoa Urbatsch & Roberts, gen. nov.—25 Medranoa parrasana (S.F. Blake) Urbatsch & R.P. Roberts, comb. Microseris laciniata subsp. detlingii -Chambers, subsp. nov. —200 Microseris 1 K.L.Chambers, subsp. nov.— Neonesomia Urbatsch & RP. Roberts, en. nov.—252 aed johnstonii (G.L. Nesom) Urbatsch & R.P. Roberts, comb. nov. 253 Neonesomia palmeri (A. Gray) Urbatsch & RP. Roberts, comb. nov.—253 Packera streptanthifolia var. borealis (Torr. & A. Gray) D.K. Trock, comb. nov.—289 Packera subnuda var. moresbiensis Q.A. Calder & . L. Taylor) D.K. Trock, comb. noy.—289 Persicaria ere var. beyrichiana (Cha aa 1) Free ab no Prenanthes carrii JR Se RJ. non, & WC. Holmes, sp. nov_-187 Prestonia ts J.E Morales, sp. nov.—16 Prestonia Selina J.E Morales & A. uentes, sp. nov.—166 Pseudostellaria oxyphylla (B.L. Rob.) R.L. Hartman & Rabeler, comb. nov.—176 Solidago caesia var. zedia R.E. Cook & Semple, var. nov.— Solidago curtisii var. flaccidifolia (Small) R.E. Cook & as comb. et stat. nov.—223 Sicicuihus acostensis L.A. Gonzalez @& J.E Morales, sp. nov.—98 ee setifolia var. greggii (A. ay) Strother, comb. nov.—287 ees Urbatsch & RP. Roberts, gen. nov.—255 Xylovirgata pseudobaccharis (S.F. Blake) Urbatsch & R.P. Roberts, comb. nov.—256 ANEW SPECIES OP LEUCOPERYELUM (SCROPHULARIACEAE) AND COMMENTS ON RELATIONSHIPS OF THE GENUS James Henrickson epartment of Biolog California State University Los Angeles, California 90032, U.S.A. henrickson@mail.utexas.edu ABSTRACT Leucophyllum coahuilensis is described from a gypsum area in central Coahuila, Mexico. It is charac- terized by small, sessile leaves, a vestiture of dendritic hairs with long, slender radii, and blue to vio- let corollas with white on the throat floor with orange maculations. It is unique within the genus in having only 2 ovules per ovary. Recent cytological and molecular data regarding relationships be- tween the Scrophulariaceae and Myoporaceae are discussed. RESUMEN Se acscnle a Reucophy iis coaluilensis: como una eapecieh nueva de suelos yesosos del centro de Coahuila por presentatr | un pense: ee ba maculas anaranjadas. La nueva especie es la Unica del género gue fee dos se por ovario. Se ee datos eee recientes, y las relaciones del género con las con radios lees me tenues, y las corolas de ee Vv felet acon] blan ca familias Scrophulariaceae y Myoporaceae. INTRODUCTION Since the publication of the taxonomic monograph of Leucophyllum (Henrickson & Flyr 1985) two new taxa have been described (Nesom 1991, 1993) and a third new species is described Deen, eo iron op on chro- mosome numbers, anatomical features, and { g ps have also been published and are summarized herein. TAXONOMY While traveling in central Coahuila ona recently paved highway from San Pedro de las Colonias to Laguna del Rey, I saw extensive, previously unexplored, outcroppings of gypsum. I drove to the slopes, parked my vehicle and within 50 feet found this distinctive new species of Leucophyllum. Leucophyllum coahuilensis Henrickson, sp. nov. (Figs 1, 2). Typr: MEXICO. COAHUILA: N end of Valley of Acatita, 15.1 road mi N of Rancho Acatita where NE- SW running spur (Sierra del Cuchilla) extends to highway, the ridge containing extensive areas of gypsum; collections from the N side of this ridge; 26° 39'N, 103° SIDA 21(1): 1-9. 2004 BRIT.ORG/SIDA 21 Fic. 1.1L phy I huil A. Growth habit, the plant ab , the p m tall. B. Short shoot on young stem eis Pays Pgs | advitic lh Cc) 4 fal I n 1.4L lateral view, Flowe y vestitured calyx and Snreadiig corolla lobes. E. Corolla, face view, showing ae anthers and orange spots onthe tower une: ne la in lower throat and lower lobes. F. Corolla, face view, showing posterior stigma, i Bx 6;C x 1.3;D.x 3;E x 4;F x 4 (1 ) NOSXOTYNIH WATIAHdODNI1 Fic. 2. Leaf vestiture of Leucophyll huilensis. A. Small leaf 3 mm wide, adaxial surface, showing loose vestiture of moderately branched hairs. B. Transverse section of leaf showing thickness of the i i lati he leaf thick C | g leaf cl . 1. .£ igt pee Sr. eel mst AA q sees Il = 1.0mm. BRIT.ORG/SIDA 21(1) 10'W, 1070-1200 m, 20 Sep 1999, Henrickson 22601 with D. Riskind (HOLOTYPE: TEX; ISOTYPES: MEXU, NY, GH, US). bus li. Folia ovata ad elliptica, parva, Frutices globosi gent I Jum gyy sessilis, vestimento dendritico (cum radiis longis gracilibusque), sub pilis dendritici stipitato- glandulosa; axillae foliorum cum caespitibus trichomatum indutae; corollae profunde caeruleo- purpureae coloratae, cum coloribus luteis basi faucum, cum maculae aurantiaceae ad medium faucum, 10-13 mm longae. Fructus cum solis duobus seminibus instructus (non cum seminibus numerosae in fructo quoque, ut in caeteris speciebus generis) Divaricately branched, broadly globose, densely vestitured, whitish shrubs 3-6 dm tall, 4-8 dm wide, becoming rather thorny due to retention of old divergent stems; young stems with long-shoot internodes 3-13 mm long, the internodes densely white pannose with short dendritic hairs and an overstory of scattered, slender, longer dendritic hairs, with a shorter series of nearly sessile stipitate glands; understory dendritic hairs with 3-5 short nodes, each with 1-4 radii, 0.2-0.35 mm long, the longer scattered emergent hairs somewhat cylindrical with the central axis somewhat zig-zagged, 0.8-1.2 mm long, with 6-10 nodes, the internodes 0.1-0.15 mm long, the radii 1-2 per node, 0.2-0.3 mm long; with age the longer vestiture weathering away to forma closer pannose vestiture that eventually is replaced as older stems develop a light grayish, weathered bark. Leaves alternate on long shoots, and crowded on axillary fascicles; leaf blades ovate, ovate-elliptical, elliptical to obovate, (3.5-)5-9(-12) mm long, (2.2-)3-5 (-7.6) mm wide, obtuse to rounded, sometimes acute at the tip, broadly cuneate to rounded at sessile bases, entire, flat or somewhat saddle shaped, witha white, loose, uniformly dense vestiture on both surfaces, the hairs dendritic, 0.7-13 mm long, with zig-zagged rachies with 3-7 nodes, with internodes 0.1-0.16 mm long, the radii 1-2 per node, 0.15-0.4 mm long, + 0.03 mm thick, the terminal arm often longest, the dendritic vestiture rather open, with the green leaf sur- face visible through the vestiture, with a close understory of stipitate glands 0.05-0.2 mm long, the basal adaxial surface leaves witha tuft of non-branched, white hairs 0.8-1.7 mm long. Flowers solitary in axils of the long-shoot leaves, on pedicels 1-4.7 mm long, + 0.25 mm thick, with a dense dendritic vestiture as on the stems; calyces 6-9.5 mm long, the 5 sepals separate to within 0.5 mm of the base, the sepals lanceolate to linear-lanceolate, attenuate, 6-8.5 mm long, to 0.8-1 mm wide, broadest in the middle and narrowing below, strongly dendritically vestitured outside with hairs to 1.5 mm long and with an under- story of stipitate glands 0.05-0.2 mm long, the vestiture longer and more dense at the sepal base, the inner sepal surfaces green, with only scattered stipitate glands 0.1-0.3mm long or also with scattered unbranched hairs; corollas strong blue to violet, with white-yellow at the tube base, and white with scattered or- ange-brown dots on the throat floor, the orange spots + 0.5mm in diameter in several irregular series, the corollas 10-13 mm in total length (with lobes ex- tended), the tube 7-8 mm long, to 4-4.5 mm wide at the distal throat, the 5 HENRICKSON, A LEUCOPHYLLUM 5 corolla lobes + equal in size and shape and spread, + oblong, broadly based, rounded at the tips, the upper 2 lobes (3.7-)4-5.5 mm long, 3.2-4.5 mm wide, the lower 3 lobes 4.2-6 mm long, 3-5.5 mm wide, the lobes spreading, often concavo-convex, the corollas moderately stipitate glandular outside with erect hairs to 0.1-0.3 mm long, the lower throat villous within with loosely arranged unbranched, wavy, white hairs 1-1.5 mm long that somewhat block the orifice to the throat; stamens 4, included, epipetalous; anthers white, each 2 lobed, the lower (anterior) 2 anthers 1.2-1.5 mm long (after anthesis), borne on free fila- ments 1-2 mm long, the adnate filament bases 3-5 mm long, the upper (poste- rior) anthers 1.7-1.8 mm long, borne on free filaments to 2-3 mm long, the ad- nate filament bases 3.2-6 mm long; ovary densely vestitured at the tip with erect, weakly branched hairs to + 1 mm long, with the radii also erect; carpels and locules 2, ovules | per carpel produced in the distal fourth of the septa; styles 4.5-7.8 mm long, the stigmatic tip borne + at the distal corolla tube throat, beyond the anthers, sparsely pubescent throughout with scattered non- branched hairs 0.2-0.3 mm long. Mature fruit and seeds unknown but ovoid immature seeds seen to 2 mm long, 1 mm wide. Additional collections: MEXICO. Coahuila: ca. 31.5 rd mi S of Quimicas del Rey on paved road south to yes Pedro de las Colonias, area of steep gypsum on E side of rd near 26° 42'N, 103° 10'W, Henrickson 26 (TEX); N end of Valley of Acatita, 15.1 rd mi N of Rancho Acatita a NE-SW running spur 7 del Cuchilla) extends to highway, the ridge with extensive areas of gypsum, on solid gypsum one with limestone, + 26° 39'N, 103° 10'W, 20 Sep 1999, Henrickson 22604 with D. Riskind TEX); W end of the Sierra de los Organos, 16 mi N of San ae ro (= Rancho Acatita) on Finisterre- cum del Rey highway, in gypsum-limestone area, near 26° 39N, 103° 09'W, ua m, 24 ea See: 23016 (TEX); same location, date, Henrickson 23025(TEX). All w = distributec Leucophyllum coahuilensis is characterized by its small, sessile, ovate leaves, its dense, loose vestiture of dendritic hairs with moderately long rachis intern- odes (mostly 0.1-0.15 mm long) and few (1-2 per rachis node) thick radii mostly 0.2-0.3 mm long, 0.02-0.03 mm thick (Fig. 2). The sessile leaves also have a dis- tinctive tuft of straight, unbranched hairs at the axil with the stem (Fig. 2C)-a feature not found elsewhere in the genus. All vestitured portions also have an understory of sessile or short-stipitate glands. This same vestiture also occurs on the pedicel and outer sepal surface. While the stems have a dense low un- derstory vestiture that completely obscures the stem epidermis, the vestiture of the leaves is more open, with the green leaf surface clearly visible through the leaf vestiture, unlike some taxa, where the vestiture completely covers the leaf surface. The corollas are dark blue-violet with white marked with orange maculations in the lower throat; they are moderately short (10-13 mm in total length) and are stipitate glandular outside with a dense beard of wavy hairs on the basal throat within. The tip of the ovary has a dense beard of elongate branched hairs. Unlike all other species of Leucophyllum, ovules are only 2, one per carpel. 6 BRIT.ORG/SIDA 21(1) Within the genus L. coahuilensis is easily set apart by its two ovules per ovary, its distinctive vestiture, flower color, and sessile, ovate leaves with a basal tuft of unbranched hairs on the leaf axil. Itappears most similar to L.candidum LM Johnston and L. pruinosum LM Johnston, both of which also have alternate leaves, orange maculations in the lower corolla throat, and vestitured ovary tips. The dendritic vestiture of L. candidum, however, has very short rachis intern- odes, with moderately long lower radii, and highly reduced upper radii, often not much longer than the width of the rachis internodes. As the terminal por- tions of the hairs fall away, the vestiture becomes very close and uniform (pannose), completely covering the leaf surface. Leucophyllum candidum also has petiolate, more broadly ovate, sometimes opposite leaves, and often larger flowers. It occurs from southern Big Bend Texas, eastern Chihuahua, through west-central Coahuila to northern Zacatecas and northeast Durango; its distri- bution surrounds that of L. coahuilensis, but L. candidum is not known from gypseous soils. The new species is also somewhat similar to L. pruinosum, but the latter is distinguished by its vestiture that has longer, more slender radii (0.4-0.5 mm long, 0.01-0.015 mm in diameter); by petiolate, thinner, more orbicular leaves often with crisped margins; by more open, abruptly ampliate corolla throats; a preference for non-gypseous substrates; and a distribution around the confluence of the states of Tamaulipas, Nuevo Leén and San Luis Potosi (Henrickson & Flyr 1985). The type locality of L.coahuilensis lies about one half kilometer east of the paved highway between Quimica del Rey and the loop highway northwest of San Pedro de la Colonias, about 30.5 miles (51 km) south of Quimica del Rey and 52 miles (84 km) north of the turnoff from the loop road east of Finisterre. It is part of Fjido Rio Aguanaval. The type locality lies on the northern slopes of a northeast-southwest running spur from the range (locally called the Si- erra de Cuchilla) that extends from the main range towards the highway. The area contains areas of mixed gypsum and clay, and other areas of nearly pure gypsum where the new species is common. Associates in the area include both obligate gypsophiles: Fouquieria shrevei, Dyssodia gypsophila, Tiquilia gypso- phila, Petalonyx crenulata, Dicranocarpus parviflorus, Nerisyrenia sp., Nama constancia, Haploesthes greggii, Euphorbia crepitata var. crepitata, Tiquilia gossipina, Selinocarpus purpusianus, Mentzelia mexicana, Drymarid elata, etc. and non-gypsophils: Grusonia braditiana, Jatropha dioica, Euphorbia antisiphilitica, Cordia parviflora, Castilleja lanata, Agave lecheguilla, Acacia neovernicosa, Allionia incarnata, Tiquilia greggii, Machaeranthera pinnatifida Viguiera dentata, Tidestromia gemmata, Anulocaulis eriosolenus, Kallstromia grandiflora, and Larrea tridentata. On all trips to the site, the plants always produced a few flowers but no —_ HENRICKSON LEUCOPHYLLUM 7 mature fruit. | have concluded that the plants were idling, ie., just producing a few flowers at a time, but not maturing seeds. Perhaps after periods of strong rainfall, the plants will flower more strongly and have the resources to set seed. RELATIONSHIPS AND MORPHOLOGY OF THE GENUS Evidence of a close phylogenetic relationship between Leucophyllum and the Myoporaceae has been presented from a cpDNA-based molecular phylogenetic study by Olmstead et al. (2001). In this study the Scrophulariaceae is separated into three distinct clades. Leucophyllum remained in their Scroph I, or Scrophulariaceae senso stricto clade, showing a strongly supported relationship with Myoporaceae, with 100 percent bootstrap values and a high (44) decay value. Olmstead et al. (2001) suggested submersion of Myoporaceae into Scrophulariaceae s.s. along with the Loganiaceae. Robert Chinnock (pers. comm.), who has graphed Eremophila of the Myoporaceae, concurred and considered that Myoporaceae should be recognized as a subfamily of the Scrophulariaceae s:s. In the monograph by Henrickson and Flyr (1985) the single chromosome count for Leucophyllum listed in Flyr’s (1970) dissertation, obtained from pol- len mother cells, was reported as both n = 15 and n = 16. The former number was a typographic error and the latter number may have been erroneous as well as Flyr (1970) indicated that he had difficulty in obtaining a clear chro- mosome spread. Read and Simpson (1989) reported x = 17 for Leucophyllum based on multiple root-tip counts from three species: L. frutescens, 2n = 34; L. minus, 2n = 34, and L. candidum, 2n = 68, the first two diploid; the latter a tetra- ploid. Thus the base number for the genus appears to be x = 17. Within the Scrophulariaceae, x = 17 has been reported from some mem- bers of the Tribe Gratioleae, namely Bacopa Aubl. (2n = 68; Chandron & Bhavanandan 1981), Limnophila R. Br. (n = 17, 34; Chandrum & Bhavanandran 1986; Subramanian & Pondmudan 1987) and Picrorhiza Royle in Benth. (n = 17; Jee et al. 1987). A base number of x = 17 also has been reported for Veronica L. (tribe Digitaleae) along with x = 7, 8, 21, 32. Within the Scrophulariaceae base numbers range from x = 6-20 and some genera, as Veronica, show wide varia- tion in base numbers. Within the Myoporaceae, Barlow (1971) provided chromosome counts for 58 of the 214 species of the large genus Eremophila (Chinnock pers. comm.). All species were x = 18, with diploids, tetraploids, and hexaploids occurring, Only two other counts have been published for Myoporumare n = 54, presum- ably triploid numbers for M. laetum and for M. laetum var. decumbens (Hair & Beuzenberg 1959), and 2n = 68, a tetraploid based on x = 17, for M. boninense subsp. boninense by Ono and Masuda (1981). These are the only x = 17 counts recorded for the Myoporaceae thus far. Chinnock (pers. Comm..) considers the 8 BRIT.ORG/SIDA 21(1 = Ono and Masuda (1981) count to be in error. A single count for Bontia daphnoides (Chinnock, pers. comm.) was n= 18. No counts are known for other genera of the family. The cytological data give little information of position of the genus. Niezgoda and Tomb (1977) found that the Leucophylleae and Myoporaceae have tricolpate, diorate pollen grains of a distinct type not found elsewhere in the Sy aleae. Karrfalt and Tomb (1983) further noted that the Myoporaceae r are distinctive in having epithelium-lined secretory cavities in young stems and leaves. Their study showed that Leucophyllum has air cavities in the leaves that expand in a similar manner, although these lack epithelial linings and con- tents. They considered that the unlined air cavities of Leucophyllum may be homologous with the epithelial-lined secretory cavities in Bontia of the Myoporaceae. Lersten and Beaman (1998) found no support for the contention that air spaces in Leucophyllum were modified secretory cavities of the type found in Myoporaceae, however, they did find a single pair of epithelial-lined secretory cavities at the very leaf tips in three species of Leucophyllum. These did show a morphological similarity between pecan mand Myoporaceae. Similar cavities are known from the genus Capraria, but its familial relation- ships are also under investigation (Lerston & Curtis 2001). Carlquist and Hoekman (1986) studied wood anatomy of Myoporaceae anc noted that certain xylem differences of Leucophyllum would exclude the genus from the Myoporaceae, but that there was a overall similarity of the woods of Myoporaceae and Scrophulariaceae as well as Gesneriaceae. Commentsonother species: Since publication of Henrickson and Flyr (1985) Guy Nesom has published two additional species of Leucophyllum. His L. hintoniorum Nesom from gypsum in the southeastern corner of Nuevo Leon, southeast of Aramberri (Nesom 1991) is distinctive in having a vestiture of three types of hairs:a very short understory of stellate-dendritic hairs that form adense felty base layer;a larger series of stellate-dendritic hairs; and slender uniseriate gland-tipped hairs. With age the larger hairs fall alway leaving a closely pannose vestiture on both leaves and stems. The latter published L. alejandrae Nesom froma more arid gypsum area SW of Galeana, Nuevo Leon, about 68 km away, (Nesom 1993), has a similar vestiture, but narrower, more crowded leaves and smaller flowers. Subsequent collections of L. hintoniorum include specimens with much narrower, more crowded leaves, reducing the differences between the taxa. Unfortunately, L. alejandrae is known from only two collections, only one with flowers. More collections are needed of L.alejandrae in order to evalu- ate whether it should be considered separate from L. hintoniorum. eam ACKNOWLEDGMENTS Ithank Neil Harriman for the Latin translation, Thomas Wendt for the Spanish abstract translation, David Riskind for his keen observation while in the field, and the Plant Resources Center for use of facilities. HENRICKSON, LEUCOPHYLLUM 9 REFERENCES Bar.ow, B.A. 1971.Cytogeography of the genus Eremophila. Aust. J. Bot. 19:295-310. Cartouist, S.and D.A. Hoekman. 1986.Wood anatomy of Myoporaceae: ecological and sys- tematic considerations. Aliso 11:317-334. CHANDRAN, R.and K.V. BHAVANANDAN. 1981.In:Chromosome number reports LXXIl. Taxon 30:698. CHANDRAN, R. and K.V. BHavanannan. 1986. Cytological investigation of the family Scrophulariaceae 1.Limnophila R. Br. Cytologia 51:261-270. Fiyr, L.D. 1970. A systematic study of the tribe Leucophylleae (Scrophulariaceae). Ph.D. dissertation. University of Texas, Austin. Hair, J.B.and E.J. Beuzenserc. 1959. Contributions to a chromosome atlas of the New Zealand Flora—2 Miscellaneous families. New Zealand J. Sci. 2:148-156. HENRICKSON, J. and DLL. Fiyr. 1985. Systematics of Leucophyllum and Eremogeton (Scroph- ulariaceae). Sida 11:107-172. Jee, V., U. Dear, and P. KacHroo. 1987. Addition to the cytological conspectus of alpine subalpine flora of Kashmir Himalaya. CIS Chromosome Inform. Serv. 43:7—9. Karrralt, E.E. and A.S. Tome. 1983. Air spaces, secretory cavities, and the relationship be- tween Leucophylleae (Scrophulariaceae) and Myoporaceae. Syst. Bot. 8:29-32 Lersten, N.R.and J.M. Beaman. 1998. First report of oil cavities in Scrophulariaceae and reinves- tigation of air spaces in leaves of Leucophyllum frutescens. Amer. J. Bot. 85:1646-1649, Lersten, N.R. and J.D. Curtis. 2001. Idioblasts and other unusual internal foliar secretory structures in Scrophulariaceae. Plant Syst. Evol. 227:63-73. Nesom, G.L. 1991. A new species of Leucophyllum (Scrophulariaceae) from Nuevo Leén, México. Phytologia 71:337-339. Nesom, G.L. 1993. Leucophyllum alejandrae (Scrophulariaceae), a new gypsophilic species from Nuevo Leon, México. Phytologia 74: 296-304. Niezcopa, C.J. and A.S. Tome. 1975. Systematic palynology of the tribe Leucophylleae (Scrophulariaceae) and selected Myororactae. Pollen et Spores 17:495-516. OLMSTEAD, R.G., C.W. DEPAMPHILIS, A.D. Wotre, N.D. Youne, WJ. ELIsens, and PA. Reeves. 2001. Disin- tegration of the Scrophulariaceae. Amer. J. Bot. 88:348-361. Ono,M.and Y.Masupa. 1981.Chromosome numbers of some endemic species of the Bonin Islands Il. Ogasawara Res. 4:1—24. READ, J.C. and B.J. Simpson. 1989. Documented plant chromosome numbers 1989: 2. Chro- mosome counts of three species in the genus Leucophyllum. Sida. 13:369. SUBRAMANIAN, D. and R. Ponpmuol. 1987. Cytotaxonomical studies of south Indian Scrophulariaceae. Cytologia 52:529-541. BRIT.ORG/SIDA 21(1) BOOK REVIEW Gary PAUL NaBHANand ANA GUADALUPE VALENZUELA~ZAPATA. 2004. Tequila! A Natural and Cultural History. (ISBN 0-8165-1937-4, pbk.). The University of Ari- zona Press, 355 S. Euclid, Ste. 103, Tucson, AZ 85719, U.S.A. (Orders: 520- 621-1441, fax 520-621-8899, www.uapress.arizona.edu). $14.95 pbk., $29.95 hbk., 160 pp., 20 b/w, 51/2" x 91/4", It is always a pleasure to share in someone else's passion. From the very beginning of this book, it is obvious that Ms. Valenzuela-Zapata and Mr. Nabham enjoy a passion for the agave plant. They bring us along when they gaze over vast hillsides of blue agave and when they compare varieties from all areas of Mexico. They require us to mourn with them how the over planting of one species has led to serious Boe and infestation of the agave fields. They a us into > the ve as oe well trained h twill _or jimadores, chop leaves off 2g | Pr es roasted and distilled into tequila. Where the authors do not take us is out onto the patio to sip the finished product with them so we can learn about the different varieties and how to choose a good tequila. The Mexican government's descriptions of the four kinds of tequila are on page 52-3 and then a series of pictures shows the reader every stage of tequila production from planting propagules to the aging process of the fin isnee liquor. Otherwise there is no tasting guide, as one might expect from a book called “Wine!” or dka!” ry . } ¢ trea 1 t 15 ri A+] Good i ing froma Cea = terms to species fee iptions of agaves used in the tequila indlustiy. Theres is rialonth ral history of pulque, mescal and mae The main focus of the book to this reader remains the eine effects of relying on one species of agave. Whether or not it was the authors’ intention, the next time | take a sip of tequila I will be See of the “ paneer so aia in the fields, which isa bitter aftertaste to a favorite drink.—Karen Hall, Library | Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A. — SIDA 21(1): 10. 2004 AMARANTHUS ACANTHOBRACTEATUS (AMARANTHACEAE) James Henrickson Department of Biology California State University Los Angeles, California 90032, U.S.A. henrickson@mail.utexas.edu ABSTRACT A a hol faa ‘te sel f I ld Bill Viesca in southern Coahuila, Mexico. It is a sister species to the dioecious A. acanthochiton, that has been recognized as a distinct genus Acanthochiton. Both species are psammophytic, dioecious annuals with large chartaceous, sharp-pointed pistilate praets: The new taxon differs from A.acanthochiton in its longer, less cordate, distally recurved its larger seeds, larger pistillate inflores- cences, and in many minor sharicien es! The new species also is larger, with pistillate plants ma- turing as tumbleweeds RESUMEN Se describe Amaranthus hol i a la ciencia, de las dunas de arena de Bilbao, cerca de Viesca, en el sur de Coahuila, co eee una especie hermana de A acanthochiton, especie dioica que ha sido Seereeaca en un genero Cee Bee nthochiton, Ambas especies son anuales, dioicas y panes) i I g cartaceas y puntiagudas. El t A ih ree eee qe aes eee ain menos eordagas, y ec unvadas distalmente pee aS semillas mas eae por ie inflorescencias Adem ] af 4 ag | J +2 j B ba ft ss | of nueva son mas anes y, al madurar y secarse el eae: INTRODUCTION Dioecious Amaranthus species are native to North America. They initially were treated in the genera Acnida L. (with narrow pistillate bracts and 0-5 pistillate sepals) and Acanthochiton Torr. (with very large, conduplicate, chartaceous pis- tillate bracts) (Bentham & Hooker 1880; Standley 1917; Schinz 1934) until Sauer (1955) combined both genera with the monoecious Amaranthus species. Sauer (1957) noted that hybrids between the taxa previously in Acnida resulted in fertile offspring (Murray 1940), while hybrids between the Acnida group and monoecious Amaranthus species formed sterile hybrids, this implying that the Acnida taxa may represent a distinct phylad. However, Murray (1940) found that hybrids between species of monoecious Amaranthus often produced ster- ile hybrids as well. The relationship of Acanthochiton to the other dioecious taxa has not been specifically addressed. While some regional floras have con- tinued to recognize Acnida and Acanthochiton as a distinct genera (Correll & Johnston 1970; Martin & Hutchins 1980) their combination with Amaranthus SIDA 21(1): 11-17. 2004 12 BRIT.ORG/SIDA 21(1) has been recognized in the Flora North America series (Mosyakin & Robinson 2003). While Acanthochiton, initially appears distinct from Amaranthus, Sauer (1955) points out that the differences are only quantitative and that the stami- nate plants of the taxon are very similar to those of Acnida species. Thus Sauer (1955) placed Acanthochiton wrightii Torr.into Amaranthus. Due to the existence of a previously published Amaranthus wrightiiS. Wats., he made the combina- tion Amaranthus acanthochiton Torr.) Sauer. In this paper, I present a second species related to Amaranthus acanthochiton from the Bilbao Dunes near the town of Viesca in southern Coahuila. TAXONOMY AND DISCUSSION Amaranthus acanthobracteatus Henrickson, sp. nov. (Figs. 1, 2). Type: MEXICO. COAHUILA: ca. 37 air mi ESE of Torreon on sand dunesat Bilbao, (8 mi NW of Viesca): dioecious; female plant 8 x 18 dm tall-wide with the stems dense, male plant 7 x 10 dm, with stems more erect and open; 25° 25'33.6'N, 102° 53'33.3'W, 3650 ft. elev, 13 Nov 2002, Henrickson 23209 (HOLOTYPE: TEX; ISOTYPES: MEXU and to be distributed). Amaranthus acanthochiton (Torr) Sauer primo adspectu maxime simile, sed bracteis feminis ad maturitatem angustatis basin versus (non cordatis basi), spinis ad apicem bracteae recurvatis, non rectispinis, bracteis tenuibus, non incrassatis, sine danas ae pus rainies Biandion pus L7- fF. 2 (versus 1.2-1.4) mm longis, Dioecious, depressed-globose, coarse annuals of sand-dune ee Pistillate plants densely branched, to 4-9 dm tall, 9-18 dm wide; central stems erect, to 20 mm in diameter at the base; internodes 3-13 cm long, branching alternately throughout; young stems 1-1.5 mm in diameter, striate, mottled green between the + 10 yellowish ribs. Staminate plants 3.5-9 dm tall, 4-15 dm wide, with more vertical stems to 8 mm in diameter developed from basal decumbent stems. Leaves of pistillate and staminate plants similar; basal leaves linear-lan- ceolate, 5-8(-10) cm long, 5-8(-10) mm wide, ascending, obtuse to acute, apicu- late at the tip, cuneate at the base; petioles 10-20(-30) mm long; upper-inflo- rescence leaves more linear, 1-5 cm long, 1-5 mm wide, the margins of all leaves strongly crisped, whitish, the primary-secondary veins whitish beneath. Pis- tillate plants with flowers axillary to the linear mid- upper-stem leaves, in ir- regular, dichasial inflorescences that eventually develop into an elongated clus- ter of + overlapping bracts to 25 mm long, the inflorescences dichasial at the base, with the two axies usually not branching above, the large, ovate, strongly folded bracts obscuring the upper inflorescence arrangement, the lowest bracts smallest, to 5 mm long, 3mm wide, the mid bracts largest, 10-16.5 mm long, to 10 mm wide, the distal, youngest bracts + reduced, 8-12 mm long, the bracts thickened, V-shaped in transverse section, with the midvein straight or out- wardly curved, with the sharp tips reflexed, the bracts rounded, cordate only with age at the base, broadest in the lower half, and tapering to the tip, the adaxial bract surface white throughout, the abaxial (outer) bract surface white along the base, the mid and distal portions green, not conspicuously veined, HENRICKSON, A NEW SPECIES OF AMARANTHUS ties 14 dm wide; , 77 A. Pistillate plant. ca.9 dm tall Fic.1.A note density of branching. B. Lower C. Staminate plant, ca.8 dm Ad dhl 4 long petiole J stem leaf, sh ip. F, Post-mature pistillate plant, upside down—a J h tumbleweed. 14 BRIT.ORG/SIDA 21(1) Fig. 2. moniparisen of pistate a bracts, flowers, seeds of A th thobracteat 1 A. acanthochiton. A- ss A. I note narrow base, bract shape and decurved tip B.\ ciated perianth CM d (all f Henrickson 23209 \ D-F: A. acanthochiton. D. h | illate t note cor- date hase folded +e kh j ot pel . Lot footie J vers 4 th FE Mat A all ¢ (, WILCLI\ Tha eae 4 toathl \ } bracts (A, D); the 1 mm scale is for B- Cand E-F. the outer margins thick membranous, white, entire or variously undulate to coarsely toothed, (0.3-)0.5-1.0 mm thick, the midvein white, with age the lower portion of the bracts becoming + spongy and sometimes the mid portions and margins also becoming spongy at maturity. Pistillate flowers with 2-3 sepals, the two marginal sepals largest, 3-4.5 mm long, with one larger than the other, both swollen-spongy and truncate at the base, the midribs green, excurrent, the whitish margins broad along the claw, and much expanded in the ovate blade below the acute-obtuse tip, the third sepal (when present) oblong, 0.8-2.7 mm long, abaxial. Ovary compressed; styles 2-3, 1.5-2.2 mm long, stigmatic and pilose along their inner surfaces. Staminate plants producing dichasial glom- erules of 7-20+ staminate flowers at 2-15(-30) mm intervals along the upper stems, the glomerules subtended by narrow, crisped-margined leaves; bracts 1.2-2(-3.8) mm long, sepal like in structure, white scarious except along the green, excurrent midvein; sepals 5, oblong-elliptical, concavo-convex, 3.3-4.3 HENRICKSON AMARANTHUS 15 mim long, 1.2-1.4 mm wide, scarious except along the slender green midveins, rounded to 2-toothed at the tips, the midveins excurrent for 0.4-0.9 mm, the sepals strongly spreading at anthesis exposing the anthers; anthers 5, oblong, 2-2.2 mm long, + 0.7 mm wide (before anthesis), apiculate at the tip, yellow, post-anthesis anthers 1.5-1.7 mm long, the filaments initially 0.9 mm long, elongating to 2.0 mm at anthesis. Mature utricles compressed obovate, membra- nous, the body to 2.3 mm long, to 1.5 mm wide, witha visible circumscissile line of dehiscence in the distal third, the bottom portion smooth or rugose, the cap rugose or not, + beaked below the filiform styles. Seeds compressed obovoid, smooth, shiny, reddish-brown to reddish-black, L.7-2.2 mm long, 13-16 mm wide and 0.8-1.05 mm thick, the embryo radicle pointing downward. Figs. 1, 2. Additional collections: Mexico. Chihuahua: 36 air mi ENE of Escalon along trail to Esmaralda in Bolson de Mapimi region of CDR scrub; dioecious iene on eaney knoll, near 26° 57'N, 103° 52'W, 3500 = 19 Sep 1972, Henrickson 7755 (RSA); Coahuila. Sa Bilbao, 6 mi via Hwy 91, 5S of Mex Hwy 40, robust annual to 5 dm tall, in active dunes, 25° 25'N, 102° 55'W, 12 Sep 1984, |.E. Bowers 2930, a TL Burgess, R.M. Turner (ARIZ); = 37 air mi ESE of Torreon, on the sand dunes at Bilbao on Hwy 68, about 10 mi NW of Viesca, near 25° 25'N, 102° 53'W, + 1000 m, 17 Oct 2002, Henrickson 23171, with D. Riskind; same locality, 13 Nov 2002: Henrickson 23202, 23203, 23204, 23205, 23208: 23206, 23207, all TEX and to be distributed. There is no question that the new taxon is related to Amaranthus acanthochiton. Both are branched dioecious annuals of dune systems or sandy soils. In both the leaf blades are crispate; both have broadened and thickened pistillate bracts that fold over flowers and fruit. Both have two large pistillate sepals and 1-3 smaller ones. But Amaranthus acanthochiton differs from the new taxon in several fea- tures (Fig. 2). The bracts of A.acanthochiton are more reniform in two-dimen- sional outline, typically cordate at the base, the sharp tip is straight or only slightly curved (but not distinctly deflexed at the tip), the green, mid-bract becomes thickened, spongy and develops a conspicuous raised-reticulate ve- nation pattern on the outer base surface at maurity, the outer membranous margins are more strongly erose-crenate, typically much narrower, (0.4-0.6 mm wide), and the mature pistillate inflorescences are smaller, with fewer flowers. Also, in Amaranthus acanthochiton, the pistillate flowers have two larger mar- ginal, clawed sepals, and may have 1-3 smaller, oblong, membranous sepals, 1 on one side, and 0-2 on the other face, however, in the smallest bracts, the pistil- late flowers may just have 5 small, oblong sepals. Overall the sepals in A. acanthochiton are similar to those of the new species. But in some collections the blades of the larger sepals can become strongly expanded and develop a branching network of veins and have denticulate margins (Fig. 2 E). The larger sepals may also develop thickened, spongy bases at maturity as in the new taxon. Seeds in both are similar in orientation, color and shape, but are smaller (1.2- 1.4 mm long, 0.95-1.05 mm wide) in A. acanthochiton 16 BRIT.ORG/SIDA 21(1) Amaranthus acanthochiton is known from sandy flats and dunes from east- ern Arizona (Navajo, Greenlee cos.); western New Mexico (San Juan, Rio Arriba, McKinley, Valencia, Catron, Socorro, Otero, Dona Ana, Luna, and Hidalgo cos.); Texas (El Paso, Hudspeth and Jeff Davis cos.) and northern Mexico, Chi- huahua (Samalayuca Dunes south of Cd. Juarez) (Sauer 1955). Reports by Turner et al. (2003) of the species in Brewster, Presidio and Webb cos. have not been substantiated In Amaranthus acanthobracteatus, staminate plants have a much more open habit than the pistillate plants.ascan be seen in Fig, | A~B. Staminate plants branch at the base and the vertical lateral stems have fewer lateral branches creating an open growth form that would appear to allow for air flow between the stems and outward dispersal of pollen from the plants (Fig. 1C). The pistillate plants, in contrast, are more densely branched at the base and above; the plants forming a dense broadly globose growth form with the main lateral branches curving upward. That would presumably cause entering pollen to slow down, perhaps increasing the possibility of finding a suitable stigmatic surface. The plants are well known to the local public and in wet years are very common on the dunes where they occur nearly exclusively with Tidestromea lanuginosa (Nutt.) Standl. The plants, along with those of Tidestromia lanuginosa, are gathered for animal feed by local townfolk. The local name for the plant is “Torillo” or “Herba Voladora,” the latter (flying herb) implying that the plants become tumbleweeds at maturity. The overall architecture of the pis- tillate plants is very similar to that of Salsola, (Chenopodiaceae) the Russian thistle, and a later trip to the type locality showed that mature plants indeed become tumbleweeds (Fig. 1F). However, unlike Salsola, where a distinct abscis- sion layer forms at the stem-root junction, no distinct abscission layer forms between roots and stem. Rather, as the plants mature, their fleshy roots die, dry out, and eventually break away, leaving the broadly globose plants to tumble across the dunes dispersing the pistillate infructescences and seeds. ACKNOWLEDGMENTS I thank Neil Harriman for the Latin translation, Thomas Wendt and Fernando Chiang for the Spanish translation of the abstract, Bobbi Angell for Figure 2, ARIZ, and SRSC for loan of specimens, David Riskind for causing the initial visit the dunes at Bilbao, which I later hit at the peak of flower, and the Plant Resources Center of the University of Texas, Austin for use of facilities. REFERENCES BentHam, G.and J.D. Hooker. 1880. Amarantaceae. In: Genera plantarum. Vol 3.Reeves & Co., London. Pp. 20-43. Corrett, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner HENRICKSON AMARANTHUS 7 Maartin, W.C. and C.R. HutcHins. 1980. A flora of New Mexico. 2 Vols. J. Cramer, Vaduz. Mosyakin, S.L.and K.R. Rosinson. 2003. Amaranthus. In: Fl. N. Amer, 4:410-435. Murray, M.J.1940.The genetics of sex determination in the family Amaranthaceae. Genet- ics 25:409-431 SAuER, J. 1955. Revision of the dioecious Amaranths. Madrofio 13:5-46. Sauer, J. 1957.Recent migration and evolution of the dioecious Amaranths. Evolution 11: 11-31, ScHINZ, H. 1934. Amaranthaceae. In: A. Engler & K. Prantl. Die nattirlichen pflanznfamilien 16c:7-85. STANDLEY, PC. 1917. Amaranthaceae. In: N. Amer. fl. 21(2):95-169. Turner, B.L., H. NicHoLs, G. Denny, and O. Doron. 2003. Atlas of vascular plants of Texas, Vol. 1,, Sida, Bot. Misc. 24, Botanical Research Institute of Texas. BRIT.ORG/SIDA 21(1) BOOK REVIEW Benoit Dayrat. 2003. Les botanistes et la flore de France: trois si¢cles de découvertes. (ISBN 2-85653-548-8). Publications Scientifiques du Muséum national d’Histoire Naturelle, 57, rue Cuvier, 1-75005 Paris, FRANCE. (Orders: Tel. [33] (0) 1-40 79 3700; Fax [33] (0)1-40 79 3858; e-mail: diff.pub@mnhn fr). Price not given, 690 pp., Illus., color plates, portraits, 6 1/2" x 91/2" This massive work of scholarship is not, Dayrat declares, a history of botany in France but rather the history of the discovery of plants in France from reas to the early twentieth century. He details the id valid in current flora. The lives of the early explora- lives of those authors of at least doctors, pharmacists, members of the mailitary and of the clergy, essentially amateurs, the tions through the various regions of France, their are tr ‘amiphe a tragedies, the ney and y additional botanists are intellectual intrigues secon ee reading. By their associa ny cameoed in extensive footnotes, which also me bibioeerhleal aca of published works as well as references from many, often ohsciie resources, such as negions) botanical journals. Each bi- again listed under ne apHY conclides with the vali Every person cited is indexed in one of two indices, the other being comprehensive. Much botanical history is necessarily included in these biographies: the history of botanical investigations of such influential persons as Lamarck and Jussieu, of Candolle, of the eccentric Al- exis Jordan; the first microphotography by Brébisson, and the development of the academic disci- so considerable history of the many botanical and Linnaean societies pline of botany. There is a oe France and the establishment of her numerous botanical garden a final word Dayrat, a post-doctoral student at the California oan of sciences, reflect- 2 on Cea plants,urges the documentation of the history of discoveries of fauna and flora throughout t This is an important work and should be made accessible to all botanists and those in related sciences.—Joann Karges Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A. SIDA 21(1): 18. 2004 TAXONOMIC REEVALUATIONS IN NORTH AMERICAN ERIGERON (ASTERACEAE: ASTEREAE) Guy L.Nesom Botanical Research Institute of Texas 9 Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Four taxa of Erigeron are raised from varietal to specific rank: E. breweri var. klamathensis to E. klamathensis (Nesom) Nesom, comb. et stat. nov., E. decumbens var. robustior to E. robustior (Cronq.) Nesom, comb. et stat. nov, . C Rap ment vat. alin to E. ee (Cronq.) Nesom, comb. et stat. nov., he and E. utahensis var. s} as F. sf morphological distinctions of these taxa are ea to those of other accepted species os ae Erigeron ea aan var. disciformis (Cro ond; ) Nesom, com ifo Erigeron clokeyi inzliae N , Var. NOV, es ae the major portion of of the range of the species: the typical variety is pcacel to Clark Co., Nevada nom. nov, representing roe . Erigeron a species endemic to California, replaces the later fescue Erigeron Gr tracyi Greene is an earlier name for E. colomexicanus A. NES Texonoipe clarifications are provided for E.acris, E. uncialis and E. cavernensis, and fo catus, E. ochroleucus E. lackschewitzii, and E. parryi. A lectotype is peraee for E. ce ie vat. ae RESUMEN Cuatro taxa ac ene ot se elevan del rango varietal al especifico: E. breweri var. klamathensis a E. klamathensi ) Nesom, comb. et stat. nov, E. decumbens var. robustior a E. robustior (Cronq.) Nesom, comb. et stat. nov, E. engelmannii var. foes davisii (Crong.) Nesom, comb. et stat. nov. y iss édhonsts var. ae es tratado como F. sparsifolius Eastw. Las acs ey de estos taxa son equivalentes a las de otras especies aceptadas de Erigeron. Erigeron poliospermus vat. disciformis (Cronq.) Nesom, comb. et stat. nov, se eleva de rango desde E. poliospermus forma disciformis. Erigeron clokeyi var. pinzliae Nesom, var. nov. ocupa la mayor parte del area de la especie; la variedad tipica esta restringida a Clark Co., Nevada. Brigeron greenei Neo nom. nov., a aes una especie endémica de California, reemplaza a Greene. Erigeron tracyi Greene es un nombre mas antiguo para E. colomenientiis A. ‘Nels, Se wee ioe taxonomicas para E. acris, E. uncialis y E. cavernensis, y para las especies interrela- adas E. radicatus, E. ochroleucus, E. lackschewitzii, y E. parryi. Se designa un lectotipo para E. ci stenophyllus var. tetrapleuru Taxonomic modifications are required for North American Erigeron L., preced- ing a treatment of the genus for the Flora of North America project. A new va- riety is described, a replacement name is provided for a species currently rec- ognized by a later homonym, one taxon is raised from the rank of forma to variety, and four taxa previously recognized as varieties are treated at specific rank. Morphological distinctions of these former varieties are as significant as those separating many other species of Erigeron, and species rank is consistent SIDA 21(1): 19-39. 2004 20 BRIT.ORG/SIDA 21(1 with current taxonomy within the genus. Clarifications of the taxonomic status of other taxa also are provided. Fuller mor phological descriptions will be pro- vided in the FNA treatment. Erigeron breweri var. klamathensis at specific rank aay klamathensis (Nesom) Nesom, comb. et stat. nov. BASIONYM: Erigeron wert A. Gray var. klamathensis Nesom, Phytologia 72:175. 1992. Type: U.S.A, CALIFORNIA HUMBOLDT Co. Trinity Summit, 2 mi SE of Devil’s Hole, exposed rocky points in woods, westerly exposure, 26 Jul 1935, J.P Tracy 15515 (HOLOTYPE: UC! ISOTYPES: JEPS!, MO!, TEX!). —_ Flowering Jun-Sep. Outcrops, ridges, crevices, rocky slopes, over shale, granite, serpentine, peridotite, chaparral, oak-pine, fir-oak, mixed evergreen woodlands; (450-)700-2150 m; California, Oregon. Prior to the recognition of var. klamathensis, most previous identifications had referred these plants to the sympatric Erigeron foliosus Nutt. var. confinis (TJ. Howell Jeps. l originally treated var. klamathensis within E. breweri A.Gray (Nesom 1992a) primarily because of similarities in vestiture and habit but here hypoth- esize that a closer relationship of var. klamathensis is more likely with E. foliosus. Recognition of E. klamathensis at specific rank emphasizes its geographic and morphological distinctions and its ambiguous evolutionary affinity. Erigeron klamathensis occurs in the Klamath Ranges of Siskyou, Trinity, and Shasta counties, California, and adjacent Oregon (Nesom 1992a, Map 5). It is disjunct from E. breweri, which is distributed in the Sierra Nevada, south- western California, and parts of the Great Basin province (Nesom 1992a, Maps 3 and 4) and is more similar in range and ecology to E. foliosus, which is prima- rily a species of coastal ranges (Nesom 1992a, Maps 6 and 7). Erigeron klamathensis produces glandular phyllaries similar to those of E. breweri var. breweri and strongly lignescent bases like E. breweri var. porphy- reticus (as well as E. foliosus var. confinis). The spreading-deflexed orientation of the stem vestiture in E. klamathensis is similar to that commonly found in E. breweri, but the sparsely pilose-hirsute vestiture of long, stiff hairs differs from the hirsutulous vestiture of E. breweri. Cauline vestiture in Erigeron foliosus is consistently antrorsely strigose to nearly or completely absent, but the habit of EF. foliosus Nutt. var. confinis is nearly identical to that of E. klamathensis and the phyllaries are similarly glandular. Their similarity in overall appearance, geography, and ecology has led to mixed collections: e.g.,a collection from Josephine Co., Oregon (Denton 2409, HSC) has one plant of E. klamathensis and several stems of E. foliosus var. confinis, show- ing no intermediacy. Intermediates between the two are encountered but they are not common. The rayless Erigeron petrophilus var. viscidulus (A. Gray) Nesom also is simi- lar in habit to E. klamathensis, has similar involucral vestiture, and is partially NESOM, TAXONOMIC REEVALUATIONS IN ERIGERON 21 sympatric with it (Nesom 1992a, Map 8). Var. viscidulus should considered among possible close relatives of E. klamathensis. The radiate taxa under consideration can be identified by the following contrasts. Erigeron breweri is represented in the key by var. breweri because it is the only variety of E. breweri similar to E. klamathensis in its densely glan- dular phyllaries lacking non-glandular hairs. 1. Stems strigose to glabrate Erigeron foliosus 1, Stems hirsute to hispid-hirsute or pilose-hirsute. 2. Sterns arising from slender, woody basal offsets, these from a strongly devel- oped, woody root; hairs of stems 0.5-1 mm long; inner phyllaries with broad, white, thickened margins, lacking distinctly demarcated green apical areas____—s—s«sEErigeron klamathensis 2. Stems arising from slender fibrous-rooted rhizome-like bases, without a strongly developed woody root; hairs of stems 0.1-0.4 mm long; inner phyllaries with green apical areas Erigeron breweri var. breweri In addition to the difference in vestiture, stems of FE. klamathensis average con- siderably shorter than E. foliosus var. confinis [6-15(-20) cm tall vs. 10-)15-35 (-50) cm] and the root system usually is distinctly thicker and woodier than in var. confinis. Erigeron decumbens var. robustior at specific rank Erigeron robustior (Crong.) Nesom, comb. et stat. nov. BASIONYM: Frigeron decumbens Nutt. subsp. robustior Cronq., Brittonia 6:174. 1947. Erigeron decumbens Nutt. var. robustior (Cronq.) Cronq., Vase. Pl. Pacific Northwest 5:175. 1955. TyPE: U.S.A. CALIFORNIA. HUMBOLDT Co.: valley of South Yager Creek, 26 Jun 1932, J.P. Tracy 10252 (HOLOTYPE: UC). Erigeron robustior is known from Humboldt, Trinity, and Mendocino counties in northwestern California and is disjunct from E. decumbens, which is known from six counties of northwestern Oregon. The two taxa also are separated by consistent morphological differences (key below). It is possible tl and E. robustior have an evolutionary sister relationship, but the magnitude of difference between them is consistent with other accepted species of Erigeron, particularly within the “Erigeron eatonii A. Gray group,” of which they area part. E. decumbens — . Involucres (4.5-)5-6 mm high, 9-12 mm wide, phyllaries linear-lanceolate, apically linear-acuminate; disc corollas 3-3.5 mm long; cypselae 1.2-1.6 mm long; heavy soils in seasonally wet or dry upland prairie grasslands; 100-300 m elev. decumbens . Involucres 6-8.5 mm high, 12-18 mm wide, phyllaries narrowly oblanceolate to lanceolate, apically acute to acuminate; disc corollas 3.5-4.5 mm long; Lead (1.8-)2-3.2 mm long; rocky or gravelly slopes, sometimes over serpentine, glade and meadows, sagebrush; 700-1500 m elev. aon robustior — Comments regarding the biology of Erigeron decumbens sensu stricto on the Center For Plant Conservation website (CPC 2003) note the following: “This 22 BRIT.ORG/SIDA 21(1) rare species spreads vegetatively via rhizomes over very short distances [of] about 4 inches ( @0 E. engelmannii Calid kal £ WW . AIV .ndq DDIT Open circles r 1998; Albee et al. 1988) and from Cronquist (1947). Fic. 1. Distributi f Eri davisii and E. engelmannii g ican : .°'S F with observations of Cronquist (1947), who noted close similarities among these taxa. All three have ray corollas very narrow (0.8-1.1 mm wide vs. 1.2-1.8 mm wide) and reflexing at the tube-lamina junction (vs. broader and coiling at the tips), disc corollas distinctly inflated and indurate above the tube (vs. not in- flated or indurate), and achenes oblong (vs. narrowly obovate). Erigeron 24 BRIT.ORG/SIDA 21(1 = engelmannii shares with E. concinnus the unusual combination of densely short-hairy disc corollas and an outer pappus of narrow to broad scales. In contrast to E. pumilus and E. concinnus and their other close relatives, the stems and leaves of E. engelmannii do not have dense, minute glandularity and ray corollas appear not only to reflex (as in the E. pumilus group) but also to coil at the tips (as in the E. poliospermus group). This apparent combination of ray behaviors in a single species is rare in Erigeron. In sum, EF. engelmannii has features of both the E. pumilus group and the E. poliospermus group. 1. Involucres 3.5—5(—6) mm high;ray corollas 5-10 mm long, laminae 0.8—1.1 mm wide, apparently both coiling at the tips and reflexing at the tube-lamina junction; disc corollas inflated and indurate above the tube, puberulent; cypselae 1.4-1.8 mm long, oblong; outer pappus of narrow scales or setae Erigeron engelmannii _ Involucres 5-8 mm high; ray corollas 8-14 mm long, laminae 1.2-1.8 mm wide, coiling at the tips, not reflexing; disc corollas not inflated or indurate, glabrate; cypselae 2.1-2.5 mm long, obovate; outer pappus of setae Erigeron davisii Stems and involucres of E. davisii are more densely hairy than in E. engelmannii, easily seen with a collection of specimens of both; the difference is difficult to characterize ina key. Erigeron poliospermus forma disciformis at varietal rank Populations of discoid plants of Erigeron poliospermus from localities in central Oregon were originally treated by Cronquist at rank of forma. Geographically discrete discoid population systems, without other morphological differentia- tion, are generally recognized at varietal rank in other species of Erigeron. Erigeron poliospermus A. Gray var. disciformis (Cronq.) Nesom, comb. et stat. nov. Erigeron oo A. Gray forma disciformis Cronq., Brittonia 6:194. 1947. Type: U.S.A. OREGON. CROOK Co. near camp on Hay Creek, rocky hillsides, 840 m, 12 Jun 1894, J.B. Leiberg 212 (HOLOTYPE: NY; ISOTYPE: GH). 1. Stems branched at or below tae basal leaves and branches originating on elongate internodes from proximal 1-6 cm of primary stem; stems and involucres densely minutely glandular, without non-glandular hairs or sparsely ei Erigeron poliospermus var. cereus 1. Stems unbranched, basal leaves and stems originating from se nodes at the caudex apex; stems and involucres sparsely minutely glandular and densely hirsute 2. Heads discoid, ray florets absent Erigeron poliospermus var. disciformis 2. Heads radiate, ray florets present and conspicuous Erigeron poliospermus var. poliospermus Erigeron utahensis var. sparsifolius at specific rank Cronquist (1947, p. 273) observed that Erigeron utahensis A. Gray and E. sparsifolius Eastw.“intergrade so completely that specific recognition is scarcely possible” and treated them within a single species. He later noted (Cronquist 1994, p. 342) that “the species consists of two wholly intergradient varieties of NESOM, TAXONOMIC REEVALUATIONS IN ERIGERON 25 ff oa e @A A A @ E. utahensis ® A E. sparsifolius a a | 1. sparsifolius.R d f llecti NY and BRIT. as well as several I Fic? Distributi £ Lyi J citations from Cronquist (1947). strikingly different aspect.” A large set of collections (NY) confirms the “strik- ingly different aspect” of these taxa and, in contrast to Cronquist’s view, indi- cates that they are distinct in morphology as well as phenology. The two are sympatric over a significant area (Fig. 2) and are ecologically similar, but in Washington, Garfield, and western Kane counties, Utah, where E. utahensis is abundant, E. sparsifolius apparently does not occur. In San Juan Co., Utah, where both are common, apparent intermediacy in some collections may be evidence of hybridization, but more generally, E. sparsifolius has been identified consis- tently and it appears to be reproductively isolated from E. utahensis. The view that they are “wholly intergradient” is not corroborated by herbarium mate- 26 BRIT.ORG/SIDA 21(1) rial. And apparently from field observations, AH. Holmgren noted (label of Holmgren 16229, NY, San Juan Co.) that E. sparsifolius is “specifically distinct from E. utahensis.” The two species can be identified by the following contrasts. _ Cauline leaves linear, bracteate, relatively even-sized above midstem and continu- ing to immediately proximal to heads; heads (1-)3-10 from eee well above midstem;involucres 3-5 mm high, 5-8 mm wide; ray florets 10-14(-20), corollas 4— 8 mm long; disc corollas viscid-puberulent with blunt-tipped hairs; flowering Jun— Sep Erig = eron sparsifolius 1. Cauline leaves gradually smaller distally, absent proximal to heads; heads 1-3(-5) from branches from midstem or above; involucres 5-7 mm high, (7-)12-15 mm wide; ray florets 28-40, corollas 10-18(-20) mm long; disc corollas sparsely strigose- villous with needle-like hairs; flowering mid Apr—Jun(—Jul) Erigeron utahensis Erigeron sparsifolius Eastw., Proc. Calif. Acad. Sci. 2,6:297. 1896. Erigeron utahensis Gray var. sparsifolius (Eastw.) Cronq., Brittonia 6:273. 1947. TyPE: U.S.A. UTAH. SAN JUAN Co: Willow Creek, 14 Jul 1895, A. Eastwood 48 (HOLOTYPE: CAS; ISOTYPES: GH, US!). Wyomingia vivax A. Nels., Bot. Gaz. 56:70. 1913. TyPE: U.S.A. San Juan Co.: Geyser Canyon, least slope of La Sal Mountains, dry rocky hills, 9000 ft, 30 Jul 1912, EP. Walker 355 (HOLOTYPE: RM; ISOTYPE: GH!, US). Flowering Jun-Sep. Rocky or sandy soil, soil pockets and crevices in sandstone, canyon bottoms, stream terraces; 1100-1700 m; Arizona, Colorado, Utah. Erigeron utahensis A. Gray, Proc. Amer. Acad. Arts 16:89. 1881. Type: U.S.A. UTAH. [KANE Co.:] Kanab, Mrs. A.P. Thompson s.n. (HOLOTYPE: GH}; internet image!). Erigeron stenophyllus var. tetrapleurus A. Gray, Proc. Amer. Acad. Arts 8:650. 1873. Erigeron tetrapleurus (A. Gray) Heller, Bull. Torrey Bot. Club Ree 8.1898. Erigeron utahensis A. Gray var. tetrapleurus (A. Gray) Cronq,, Brittonia 6:272. 1947. LecToTyPe, designated here: U.S.A. Kane Co. Kanab, Mrs. A.P Thompson s.n. (GHI, internet Eiminee)) Gray also cited another Utah collection (FM. Bishop s.n., 1873)—this is mounted on the same sheet as the lectotype. Flowering mid Apr-Jun(-Jul). Rocky slopes, cliff bases, ledges, and crevices, sandstone outcrops and terraces, sandy soil, gravelly limestone, shale, cotton- wood floodplains, creosote bush, blackbrush, blackbrush-Joshua tree, warm desert shrub, salt desert shrub, mountain brush, pinyon-juniper, oak-maple- aspen; 800-2100(-2450) m; Arizona, California, Colorado, New Mexico, Utah. It seems likely that FE. utahensis eventually will be discovered in southern Nevada. A new variety within Erigeron clokeyi Erigeron clokeyi is distinct in a number of features: a low, relatively caespitose habit; stems erect to basally decumbent-ascending and mostly monocephalous; minutely glandular stems, leaves, and phyllaries; nonglandular cauline hairs spreading-deflexed; leaves narrowly oblanceolate; and ray corollas reflexing at the tube/lamina junction. The species has been treated as a single unit (Cronquist 1947, 1994; Nesom 1992b), but two expressions of leaf vestiture ex- ist within the species. Plants from the Charleston Mountains in Clark Co., Ne- NESOM, TAXONOMIC REEVALUATIONS IN ERIGERON 27 ———————— _O E. clokeyi var. clokeyi ae se ——_ e ° . @ E. clokeyi var. pinzliae ~~ \ of i) ‘ \ \ a Bee - +s 1! Fic. 3. Distributi f£Cyri a) as) J £ W 4 vata 1DDIT *aL 4 bse? £ CalElars /9NNA\ J ’ vada (the type locality), have hirsute-strigose leaves; those from other areas of the range (California, Nevada, west-central Utah), including most of the closest populations in southeastern Inyo Co., California, have hispidulous to hirsutu- lous leaves (Fig. 3). Some plants from Inyo Co. show a tendency toward strigose 28 BRIT.ORG/SIDA 21(1) foliar vestiture. Cypsela size is slightly but consistently different, and the Clark Co. plants appear to have narrower leaves and a greater tendency for the stems to be decumbent-ascending. The two varieties are ecologically similar. Erigeron clokeyi Cronq., Brittonia 6:214. 1947. Type: U.S.A. NEVADA. CLARK Co. Charleston Mountains, Lee Canyon, brushy meadow, yellow pine belt, 2700 m, 12 Jul 1937, 1. W. Clokey 7742 (HOLOTYPE: NY! ISOTYPES: LL!, MINN, MO!, NY!, PH, POM RY, SMU! TEX!, UC, US!, VDB!, WS, WTU). Erigeron clokeyi Cronq. var. pinzliae Nesom, var. nov. Type: U.S.A. NEVADA. MIN- L Co: Wassuk Range, road to Mt. Grant summit, 0.7 road mi below spring, T8N, R28E. NE 1/4 sect. 13, ca. 10,000 ft, 7 Sep 1995, A. Pinz! 11733 (HOLOTYPE: BRIT, ISOTYPE: NSMC). Differt a E. clokeyi sensu stricto vestimento foliorum hispidulo vel hirsutulo et cypselis minoribus. Flowering Jun-Sep. Dry, rocky habitats, dry meadows, sometimes with sage- brush or mountain mahogany, treeless areas and often with yellow, bristlecone, or limber pines; 2200-3450 m; California, Nevada, Utah. Differences between the two varieties are summarized here. 1. Leaves hirsute-strigose, hairs basally ascending, otherwise straight and ee antrorsely appressed; cypselae 2.2-2.5 mm; Charleston Mountains, Clark C Erigeron clokey var. clokeyi . Leaves uniformly hispidulous to hirsutulous, hairs stiffly spreading to spreading arching; cypselae 1.8-2 mm; east-central California, southern Nevada, west-central Utah Erigeron clokeyi var. pinzliae — New name fora cs aeeaes species Erigeron greenei Nesom, nom. nov. REPLACED SYNONYM: Erigeron angustatus Greene, Bull. Calif. Acad. Sef 1(3):88. 1885 (non Erigeron angustatus Fries ex Nym., Consp. FL Europ. 2:389. 1879). TYPE: U.S.A. CALIFORNIA. NAPA Co: dry hills on either side of Napa Valley, Jun-Oct,[Napa, 13 Aug 1874], E.L. Greene 339 (not located with certainty, see comments in Nesom 1992; probable type material GH). The name Erigeron angustatus Greene has been used (Nesom 1992) for a discoid species now known to occur in Lake, Napa, Sonoma, Tehama, Trinity, Shasta, and Siskyou counties, California. Because E. angustatus Greene is a later homonym, it is replaced here. The new epithet commemorates Edward Lee Greene (1843-1915), whose perception of supraspecific natural groups and generic boundaries, in many instances, has proved to be remarkably close to modern concepts. Status of Erigeron cavernensis Erigeron cavernensis has been treated as a synonym of E. uncialis (Cronquist 1994, Nesom 1992b) but E. uncialis var. conjugans, which closely approaches E. cavernensis in its geographical range (Fig. 4), is markedly different from the latter and perhaps more similar to E. cronquistii. Erigeron uncialis var. uncialis and E. uncialis var. conjugans have features in common between themselves and contrast as a unit with E. cavernensis. NESOM, TAXONOMIC REEVALUATIONS IN ERIGERON 29 co be, E. uncialis @ var. uncialis O var. conjugans © Fic. 4. Distributi f Eri jali 1 F. cavernensis. R d f llecti t NY, NSMC, and BRIT. 1. Phyllaries eglandular or sparsely glandular near the apices and along midregion; stems and leaves eglandular; leaves striqose to hirsute-villous, vestiture less dense on abaxial surfaces; cypselae 1.3-1.8 mm long __ Erigeron uncialis 30 BRIT.ORG/SIDA 21(1) 1. Phyllaries evenly densely glandular; stems and leaves glandular; leaves hirsute-ca- nescent, equally hairy on both surfaces; cypselae 1-1.2 mm long Erigeron cavernensis Erigeron uncialis S.F Blake, Proc. Biol. Soc. Wash. 47:173. 1934. Type: U.S.A. CALI- FORNIA. SAN BERNADINO Co. Clark Mountain, 7000 ft, Jun 1933, E.C. Jaeger s.n. (HOLOTYPE: POM; ISOTYPE: US). an + | ly VI 1, Stems 0.8-2.5 cm high, hirsute-villous; leaves 1-2 cm long, hirsute strigose Prigérah uncial var. uncialis . Stems 3-7 cm high, loosely villous-strigose;leaves 2-4 cm long, sparsely and clos strigos rigeron conn var. conjugans a. Erigeron uncialis S.F. Blake var. uncialis. Flowering May-Jul. Crevices, cliff bases, oot in limestone, pinyon-juniper, pine-fir; 1900-2600 m Californ b. Erigeron uncialis S.F Blake var. conjugans S.F. Blake, Proc. Biol. Soc. Wash. 47:174. 1934. Erigeron uncialis SF Blake subsp. conjugans (S.F Blake) Cronq., Brittonia 6:211. 1947. TYPE: U.S.A. NEVADA. CLARK Co. Charleston Mts., Kyle Can- yon, Big Falls, crevices of vertical rock faces, 9000 ft, 3 Sep 1927, C.L. Hitchcock s.n (HOLOTYPE: POM; ISOTYPE: US). Flowering May-Aug. Crevices in limestone cliffs and boulders, yellow pine or limber pine; 2200-2800 m; Nevada. Erigeron cavernensis Welsh @ Atwood, Great Basin Naturalist 48:495. 1988. Type: U.S.A. NEVADA. WHITE PINE Co:: Schell Creek ee 25 ue mi SE . Ely, ca. 2 km NE of summit of Cave Mountain, 3172-3233 m, li d rubble, Pinus flexilis-P longaeva community, 18 Jul 1981, B. Welsh, 5 eae and E. Neese 910 (HOLOTYPE: BRY; ISOTYPES: NY!, POM, RM, UNLY, US|, UT). Flowering Jun-Jul. Limestone ridges, outcrops, and cliffs, often with bristle- cone pine, limber pine, spruce; 2100-3400 m; Nevada, known only from the White Pine Range of White Pine County and adjacent Nye County. Status of Erigeron radicatus and E. ochroleucus var. scribneri Confusion has existed in the distinction between Erigeron radicatus and E. ochroleucus, but the hypothesis is advanced here that they are distinct species sympatric over a significant area. In this view, E. radicatus has a wider geo- graphic distribution (Fig. 5) than previously recognized and E. ochroleucus is more restricted in range (Fig. 6). Small plants of Erigeron ochroleucus, often identified as E. ochroleucus var. scribneri, approach E. radicatus in aspect and many plants of E. radicatus have been identified as E. ochroleucus var. scribneri. Erigeron radicatus is distinctive in its branched caudex, short-villous cauline vestiture, smaller leaves with more reduced vestiture, smaller heads, involucral hairs usually with colored crosswalls, and fewer pappus bristles (see key couplet below). A thick taproot and unbranched caudex usually are contrasting features of E. ochroleucus and the phyllaries of E.ochroleucus often are apically linear-acuminate and loose, a feature not found in E. radicatus. NESOM, TAXONOMIC REEVALUATIONS IN ERIGERON 31 Xf Cae, a. “Uy af I ab E. radicatus i | eS AS ( each es ea a ras Ai TT a fe . ie . vom Fic 5 iat aes ar g D 4, IDA c n fi ; fide Caleb Morse ifi For the most part, Erigeron radicatus seems consistently distinct from E. ochroleucus, but I have identified as E. radicatus a few plants with involucral vestiture lacking colored crosswalls (eg., Carbon Co., Wyo.: Dorn 3687, RM; Fre- mont Co., Id.: Moseley 835, RM) and some plants as E. ochroleucus with involu- cral vestiture with colored crosswalls (e.g., Big Horn Co., Wyo.: Hurd 208, RM; Sheridan Co., Wyo. Nelson 6149, RM; Johnson Co., Wyo.: Nelson 5984, RM). A few plants with an unbranched caudex are identified here as E. radicatus (e.g., Big Horn Co., Wyo. Williams 3221, RM; Gallatin Co., Mont.: Dorn 914, RM). Some of these plants of apparently intermediate morphology may represent hybrids or introgressants. The chromosome number is reported as 2n = 36 from Cheviot Mt., Alberta (Packer & Witkus 1982). A count of 2n =18 from Albany Co., Wyo- ming (Semple & Chmielewski 1987) was from Erigeron simplex Greene, the voucher misidentified by Nesom as E. radicatus. Erigeron radicatus often is scapiform at relatively high elevations (2750- 3350 m); more eastern populations in the Great Plains at lower elevations (1450- 32 BRIT.ORG/SIDA 21(1) 2550 m) tend to have leafier stems. Plants of the collection from Weld Co. Colo- rado (rocky ridge ca. 3.4 mi N of Rockport, 6000 ft, Dorn 8222, RM), have slightly narrower phyllaries and involucral trichomes essentially without colored crosswalls, but in other respects they are similar to typical (but smallest) E. radicatus. Some of the low-elevation populations in Albany and Carbon cos., Wyoming, produce atypically short rays, and | initially regarded the eastern, low-elevation plants as taxonomically distinct. Finally, however, I was unable to find significant difference in other features. Even so, a species over such a wide range of elevation is unusual, both in the U.S.A. and in Canada, where E. radicatus occurs from montane sites in Alberta to localities at lower elevation in Saskatchewan. On the other hand, E. ochroleucus, one of its closest relatives, occurs over an equally wide elevational range. Erigeron radicatus Hook., Fl. Bor-Amer. 2:17, t. 123.1834. Type: CANADA. ALBERTA. Mountains near Jasper’s Lake, Rocky Mountains, Drummond s.n. (SOTYPE, fide an- notation in 1945 by Cronquist: NY!). The NY sheet does not have collection infor- mation other than a label noting “Erigeron divaricatum Hook., very rare, Hoo Am.” Erigeron macounii Greene, Pittonia 3:162. 1897. Type: CANADA. ALBERTA: Sheep Mountain, Waterton Lake, 28-31 Jul 1895, J. Macoun 10858 (type material: NY!). Citec as asynonym of Erigeron ochroleucus var. scribneri; placed here asa synonym of E. radicatus by Cronquist (1947) because of its small (5-3 cm long) leaves and few (8) pappus bristles. The leaves, however, are linear-lanceolate and densely strigose and rays are bluish - features more consistently characteristic of E. ochroleucus. Sas huberi Welsh & Atwood, Rhodora 103:71. 2001. Type: U.S.A. UTAH. DUCHESNE CO. Uinta ntains, Lake Fork Mtn., TZN RSW S16, NW1/4 of NEI/4, Uinta Base Meridian, 2 ants growing along windswept ridge crest above limestone talus slopes, rocky soils, 10,900 Jul 1998, A. Huber and C. Wedig 3825 (HOLOTYPE: BRY: ISOTYPES: MO!, NY! US internet 0 a Flowering May-Aug. Rocky slopes, Hues and summits, ledges and crevices, outcrops and talus, usually limestone alpine tundra: (1450-)1600-2750(- 3350)m: British Columbia, Alberta; Saskatchew panels. Idaho, Montana, Nebraska, North Dakota, South Dakota, Utah, Wyoming. . Stems (1-)2-6(-12) cm high, usually arising from tips of short, thickened caudex branches; stems short-villous; leaves (0.5-)1—5(-8) cm long, sparsely loosely stri- gose adaxially, glabrous and shiny abaxially; involucres (3-)4-6(-8) mm high, hairs of involucre usually with colored crosswalls; pappus bristles (6-)7-11 Erigeron radicatus Stems (2-)8-18(-30) cm high, arising from a nearly common point near apex of thick taproot, caudex peer inbranched; stems loosely strigose; leaves (2—)4—9(- 12) cm long, usually strigose on both surfaces at least on proximal 1/3-3/4 of blade, glabrous distally; involucres 5.5—7 mm high, hairs of involucre usually without col- ored crosswalls; pappus bristles 11-15 Erigeron ochroleucus Sans Erigeron ochroleucus Nutt., Trans. Amer. Philos. Soc., 2, 7:309. 1840. Type: U.S.A. [probably central Wyoming, perhaps Natrona Co}. “Plains of the Oregon” [Trail], [ca. Jun, 1834], T. Nuttall s.n.(GH!, PH, UC-photo and fragment). NESOM, TAXONOMIC REEVALUATIONS IN ERIGERON 33 Erigeron ochroleucus Nutt. var. scribneri (Canby ex Rydb.) Cronq., Brittonia 6:189. 1947. ete scribneri Canby ex Rydb., Mem. New York Bot. Gard. 1:405. 1900. TyPE: U.S.A. MONTANA. [MEAGHER Co.! Little Belt Mountains, 12 Aug 1883, EL. Scribner 77 ee Erigeron oe Can t. Gaz. 15:150. 1890) was published as a “nomen provisorum Erigeron see Canby & Rose, Bot. Gaz. 15:05. . Erigeron montanus Rydb. [nom. nov, Bull. Torrey Bot. Club 24:296. es 7 Wyomingia ce na (Canby & Rose) A. Nels., Man. Rocky Mt. Bot. 531. 1909. TyPE: U.S.A. MONTANA. PARK Co. Jun 1889, F. Tweedy s.n. (NY- 2 sheets!). Erigeron laetevirens Rydb., Bull. Torrey Bot. Club 28:506. 1901. TyPE: U.S.A. MONTANA. SHERIDAN Co. Big Horn Mountains: Little Goose Creek, 8700 ft., Jul 1899, F Tweedy 2005 (NY!). Flowering Jun-Aug. Rocky or sandy slopes, limestone outcrops and ridges, ta- lus, sagebrush-grassland, juniper-mountain mahogany, ponderosa pine, lim- ber pine, limber pine-Douglas fir, alpine tundra; 1100-3000(-3300) m. Alberta, British Columbia; Montana, Nebraska, South Dakota, Wyoming. The record for British Columbia is added from a report by Roemer (1996, as E. ochroleucus var. scribneri). Nesom and Murray (2004) report E. ochroleucus in arctic and boreal Alaska and immediately adjacent Yukon, long disjunct from the primary range in the western U.S.A. and adjacent Canada. Plants of Erigeron ochroleucus are consistently relatively large in stature and white-rayed in northeastern and central Wyoming (Campbell, Converse, Crook, Fremont, Hot Springs, Natrona, Niobrara, and Weston cos.), where they occur at elevations of 1100-1900(-2400) and at similar elevations in the more montane areas of north-central Wyoming and adjacent Montana (Fig. 6). These plants match the type of the species. In the latter areas, however, at elevations characteristically about 2150-2750 m and ranging up to 3350 m, the plants are smaller and commonly blue-rayed, matching the type of E. ochroleucus var. scribneri, but such plants al mmonly extend downward to 1600 m in these montane areas, fas in the area of elevational overlap so much morphological variability exists that it seems impossible to distinguish var. scribneri. Smaller, blue-rayed plants also are occasionally encountered even in areas of predomi- nantly larger, white-rayed ones. Reported chromosome numbersare 2n =18 from southern Alberta (Chinnappa @ Chmielewski 1987) and Sheridan Co., Wyo- ming Jones & Smogor 1984). A count of 2n = 54 from Niobrara Co., Wyoming (Semple 1985) was from Erigeron caespitosus Nutt., the voucher misidentified by Nesom as E. ochroleucus Status of Erigeron lackschewitzii Erigeron lackschewitzii was compared in its original description with E. grandiflorus Hook., but it instead is very similar and closely related to E. ochroleucus. Nesom (1989) treated it asa synonym of E.ochroleucus, but exami- nation of additional collections confirms it as a distinct species. The distribu- tion record for Glacier Co. (Fig. 6) is based on the citation in Lesica (2002). The record for Alberta (Waterton Lakes National Park) is added fide Joyce Gould (Alberta Natural Heritage Information Centre). 34 BRIT.ORG/SIDA 21 =~ = ) : Involucres 5. 5-7 mm high; phyllaries i esas glandular, hairs of villous estiture without colored crosswalls;ray corollas white or blue; disc corollas 2.8-3.6 mm; pappus bristles 12-15 Erigeron ochroleucus . Involucres 6-8 mm high; phyllaries densely and conspicuously glandular, hairs o villous vestiture with dark purple ses ray corollas purple to lavender; dis corollas 3.5-4.3 mm; pappus bristles 1 rigeron penis Erigeron pared Nesom & W.A. Weber, Madronio 30:245. 1983. Type: U.S.A. MONTANA. TETON Co. Bob Marshall Wilderness Area, Flathead Range, summit of ee eae Pass, 2365 m, large colony in small, dry meadow on the narrow saddle, 29 Jul 1978, K. Lac kschewitz 8487 (HOL OTYPE: MONTUS ISOTYPES: COLO!, NY!). i Flowering Jul-Aug. Rocky slopes and ridges, terraces, talus, meadows, usually calcareous; 2250-2500 m; Alberta; Montana. Status of Erigeron parryi Cronquist (1947) maintained Erigeron parryi, noting (p. 190) that it probably is “merely an unusual form” of E.ochroleucus, but he later (1955) treated it (at least by implication) as a synonym of E. ochroleucus. Collections similar to the type, however, from the region of the type locality in southwestern Montana and ad- jacent Wyoming (Fig. 6) suggest that E. parryi is distinct. Leaves of E. parryi are equally hairy (hirsute to strigose-hirsute) on both surfaces, contrasted with the reduced vestiture (loosely strigose) on adaxial surfaces of E. ochroleucus leaves. Also, E. parryi tends to have smaller heads with fewer rays and the caudices sometimes are branched. The variation in orientation of vestiture is unusual, and as between E. ochroleucus and E. radicatus, the nature of the differentia- tion (or lack of differentiation) between E. ochroleucus and E. parryi is not clear. . Leaves 1-2.5 cm long, narrowly oblanceolate, equally hairy on both surfaces; caudices branched or not; involucres 4-6 mm high ie 10 mm wide; ray florets 22-30 ____ Erigeron parryi 1. Leaves 2-6 cm long, linear to narrowly ob late, striqose adaxially, less hairy to glabrous abaxially; caudices usually not berciee involucres 5.5-7 mm high, 10- 15(-18) mm wide; ray florets 30-62 Erigeron ochroleucus Erigeron parryi Canby & Rose, Bot. Gaz. 15:65. 1890. Type: U.S.A. MONTANA. BEAVERHEAD Co.: Grasshopper Creek, dry hills, 7000 ft, Jul 1888, F Tweedy 15(GH internet image!, NY!, US?). Canby and Rose cited only “Frank Tweedy 15” as the type. Plants perennial, taprooted, caudices with or without short, thickened branches. Stems 1.5-5 cm, erect, short-hirsute to loosely strigose-villous, eglandular. Leaves basal and cauline, basal narrowly oblanceolate, 1-2.5cm long, 0.8-2.5 mm wide, entire, cauline on proximal 1/2-2/3 of stem, gradually reduced distally, densely strigose to strigose-hirsute on both surfaces, eglandular, eciliate. Heads 1; in- volucres 4-6 mm high, 7-10 mm wide; phyllaries in (2-)3 equal to subequal series, filiform-attenuate and purplish at apex, sparsely to densely villous-stri- gose, basal cross-walls sometimes purple, sparsely minutely glandular. Ray flo- rets 22-30, corollas 6-8 mm, laminae white to pink or bluish, not coiling or re{lexing. Dise florets: corollas 2.4-3 mm, throat not indurate or inflated. NESOM, TAXONOMIC REEVALUATIONS IN ERIGERON 35 = i Fi ‘ rf ©, ochroleucus a a 4 E. lackschewitzij | _ Tf \ | [P] E. parryi - i ce oer I i Cl + = | ' oe ee 2 a) + Fic. 6. Distributi f Eri hrol E. lackschewitzii, and E. parryi t NY , parry and RM (see comments in text). Cypselae 2-2.3 mm, 2-nerved, densely strigose; pappus bristles 12-15, readily deciduous, outer setae or scales prominent. Flowering Jun-Aug. Open, rocky sites, limestone and quartzite; 1600- 2250(-2600) m; endemic to southwestern Montana and adjacent Wyoming. Additional collecti ined: U.S.A. MONTANA. Beaverhead Co.: crest of Red Butte, ca.8 mi NW of Lima, common, calcareous soil, with Eriogonum mancum and Oxytropis besseyi, T13S, ROW, Sec 10, 6200 ft, 6 Jul 1986, Lesica 3928 (NY); exposed ridge crest 2 mi S of Grasshopper Creek, common in gravelly limestone and quartzite-derived soil, 6200 ft, with Sphaeromeria capitata and Eriogonum mancum, 22 Jun 2003, Lesica 8657 (RM); Tendoy Mts., above Muddy Creek Rd, abundant on crest of ridge with Lesquerella alpina, Penstemon args ee opappus sp., 1135, RLOW, Sec 34, 7000 ft, 27 Jun 1987, Lackschewitz 11307 (NY). Carbon Co Lockheart Ranch, T8S, R28E, $13, Poa cover, 4400 ft, 9 Jun 1983, Lichvar 5913 oe Big oe Mountain, gravelly soil, S slope, grassland, 8500 ft, 12 Jul 1926, Williamson gs M).J erson Co.: slopes of small hill at the head of Negro Hol- low 6 mi NE of Caldwell limestone-derived soil, 5200 ft, with Hymenoxys acaulis and Eritrichium howardii, 17 Jun 2003, Lesica 8630 (RM). Madison Co.:S end a ae Mts., 3 miS and 3 mi W from Ruby or Dam, and aw of yng Springs occasional i g grassland, 5790 ft, with Opto outa ae oO 36 BRIT.ORG/SIDA 21(1) tragalus vexilliflexus, Chrysothamnus nauseosus, Stipa comata, Aster scopulorum, non 1996, Heidel and edad 1488 — pee hland Mountains, ow ridge ca. 1 mi S of Victoria Mine W of Silver Star, locally limestone-derived soil, 5250 ft, with Cercocarpus ledifol pues riogonum mancum, 18 Jun 2003, Lesica 8640 (RM). Wyomine. Park Co.: Absaro Shoshone River drainage, ridge E of Pagoda Creek, ca. 1.5-2.5 miS of US Hwy 14-16 & 20, open rocky areas with scattered Douglas fir and limber pine, 7200-8000 ft, 1] Jun 1986, Evert 9860 (RM). Leaves and stems of Lesica 3928, 8640, and 8657 and Williamson 28 are hirsute with stiffly spreading hairs, similar to those of the type collection; leaves of Lackschewitz 11307, Lesica 8630, Lichvar 5913, Heidel and Cooper 1488, and Evert 9860 are hirsute-strigose with loosely appressed hairs. Although the vestiture of the latter six collections approaches the orientation of that of E.ochroleucus the hairs are stiffer and slightly shorter than characteristic of E.ochroleucus Peter Lesica (submitted) has reached a similar conclusion regarding the distinction of Erigeron parryi, based on field experience, more numerous col- lections than recorded here, and a morphometric study. My conclusions were reached independently of his but were based on his collections, in significant part, at NY and RM. His concepts of E. ochroleucus and E. radicatus also are similar to those outlined here but not identical. us A on Inom. illeg.], New Man. Bot. Centr. Rocky Mts. 530. 1909. Erigeron lapiluteus isan illegiti- mate ¢ replacement name for E. yellowstonensis. ri elongatus Ledeb.[nom. inval.], Icon. Pl. Fl. Ross. 1:9, tab. 31.1829 (non E. elongatus Moench 1802) Erigeron acris var. ons vaapae se & Cajand., Suom. Kasv. 566. 1906. Eri - ot. (Lund.) 1843:120. 1843. Erigeron acrisL. subsp. politus soe iiab i jaacm: Pl. Fennos va Orient. 56.1901 (non Schinz & R. Keller 1909). ae asteroides Andrz. ex Besser ee Enum. PL. Volhyn. 33. 1822 (non Roxb. 1814). Erigeron acris L. var. asteroides (Andrz. ex Besser) DC., Prodr. 5:290. 1836. Trimorpha acris (L.) S.F Gray var. ast les (Andrz. ex Besser) Nesom, ie 67:64. 1989. Listed by Tzvelev ( a synonym of iene en E. podolicus Besse Erigeron ae Gaudin (misapplied), : oe 5:265. on Erigeron acris L. var. angulosus (Gaudin) Vacc., Cat. Pl. Vall. Aoste 1:350 Erigeron droebachiensis O. Mueller aren FI. Dan. 5, 15:4, tab. 874. 1782. Erigeron acris L. 38 BRIT.ORG/SIDA 21(1) var. droebachiensis (O. Mueller) Blytt, Norges Fl. 1:562. 1861. Erigeron acris L. subsp. droebachiensis (O. Mueller) Arcang., Comp. F1. Ital. 340. 1882. ACKNOWLEDGMENTS lam grateful to the staffs at GH, MO, and NY for help during visits there, HSC, NSMC, and RM for loans of specimens, Anita Cholewa (MIN) for observations regarding the type of Erigeron davisii, John L. Strother for observations on UC collections and editorial comments in review, Caleb Morse for observations on KANU collections, Peter Lesica for comments on E. parryi and relatives, Joyce Gould (Alberta Natural Heritage Information Centre) for information on E. radicatus and E. lackschewitzii in Canada, and John Semple for providing im- ages of vouchers for previously published chromosome counts, and Dave Boufford for advice on lectotype selection for E. stenophyllus var. tetrapleurus. REFERENCES Avsee, B.J., L.M. SHuttz, and S. Goopric. 1988. nue of the vascular plants of Utah. Utah Mu- seum of Natural History. Accessed tee 2003. CatFiora. 2004. Information on California plants for education, research and conserva- tion. [web application]. Albany, California: The CalFlora Database [a non-profit organi- zation]. Accessed Jan 2004. CENTER For PLANT CONSERVATION (CPC). 2003. CPC national collection plant profile. Accessed Dec 2003 Cuinnapra, C.C. and J.G. CHMiELEwsKi. 1987. Documented plant chromosome numbers 1987: 1. Miscellaneous counts from western North America. Sida 12:409-417. Cronquist, A. 1947.A revision of the North American species of Erigeron, north of Mexico. Brittonia 6:121-302. Cronauist, A. 1955. Vascular plants of the Pacific Northwest, Part 5: Compositae. Univ. of Washington Press, Seattle. Cronauist, A. 1994. Intermountain flora. Vol.5, Asterales. New York Botanical Garden, Bronx. C7eRePANOV, S.K.1995.Vascular plants of Russia and adjacent states (the former USSR).Cam- bridge Univ. Press, Cambridge, England GLeAson, H.A. and A. Cronouist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada (ed. 2). The New York Botanical Garden, Bronx, New York. Greene, E.L. 1902. A fascicle of new Compositae. Pittonia 5:55-64. Hara, H. 1939. Some notes on the botanical relationship between North America and Eastern Asia. Rhodora 41:385-392. Huwten, E. 1968a. Comments on the flora of Alaska and Yukon. Arkiv Bot. n.ser. 7, 1:1-147. Huten, E.1968b. Flora of Alaska and neighboring territories. Stanford Univ. Press, Stanford, California. Jones, A.G. and R.A. Smocor. 1984. In: Chromosome number reports LXXXIl. Taxon 33: = + al NESOM, TAXONOMIC REEVALUATIONS IN ERIGERON 39 Lesica, P, 2002. A flora of Glacier National Park, Montana. Oregon State University Press, Corvallis. Lesica, P. Submitted. The resurrection of Erigeron parryi (Asteraceae: Astereae). Brittonia Nesom, G.L. 1989. Infrageneric taxonomy of New World Erigeron (Compositae: Astereae). Phytologia 67:67-93. Nesom, G.L. 1992a. Revision of Erigeron sect. Linearifolii (Asteraceae: Astereae). Phytologia T2208: Nesom, G.L. 1992b. Erigeron and Trimorpha (Asteraceae: Astereae) in Nevada. Phytologia Nesom, G.L. 1993. Erigeron (50 species, 71 taxa). The Jepson manual: higher plants of Cali- fornia. J. Hickman (ed.). Univ. California Press, Berkeley. Nesom, G.L.and D.F. Murray. 2004. Notes on North American arctic and boreal species of Erigeron (Asteraceae: Astereae). Sida 21:41-57. Packer, J.G. and R. Wirkus. 1982. In: IOPB chromososome number reports LXXV. Taxon 31: 363-364. Rocky Mountain Hersarium. 1998. Atlas of the vascular plants of Wyoming. Accessed Dec 2003. Roemer, H. 1996. Two plants new to the flora of British Columbia. Bot. Electronic News No. 148, 2 Nov 1996. Accessed Dec 2003. Sempte, J.S.1985.Chromosome number determinations in fam.Compositae tribe Astereae. Rhodora 87:517-527. Sempte, J.S. and J.G. CHMieLewski. 1987. Chromosome number determinations in fam. Compositae tribe Astereae. Il. Additional counts. Rhodora 89:319-325. Sioa, O. 1998. Taxonomic problems in Erigeron sect. Trimorpha (Compositae) in Eurasia. Preslia 70: 259-269. STROTHER, J.L.and W.J. Ferarte. 1988. Review of Erigeron eatonii and allied taxa (Compositae: Astereae). Madrono 35:77-91. Tzvetev, N.N. (ed.). 2002. Flora of Russia: The European part and bordering regions. Vol. VII (English translation of 1994 original). A.A. Balkema, Rotterdam/Brookfield. BRIT.ORG/SIDA 21(1) BOOK REVIEW EsMonpD Harris, JEANETTE Harris, and N.D.G. JAmes. 2003. Oo a British History. (ISBN 0-9538630-8-5, pbk.). Windgather Press, Ltd., 29 Bi Road, Bollington Macclesfield, Cheshire SK10 5NX, UK. (Orders: Distributed by Central Books, 99 Wallis Road, London E95LN, UK). US $30.00, 208 p., 10 col., +9 b/w illus., 7 1/4" x 93/4", This is not a book about the historical significance of oaks in Britain. This is not a study of the many and varied uses that oaks have supplied over the centuries. This is not a guide to the cultivation of oaks and the restoration of aged and historic woodlands. This is not about the myths and symbology that have surrounded the oak since time immortal. The Oak: A British History is all of these and more. Written in a easy to read manner which belies the authority behind the text, this will be a classic for years to come The oak has always held pride of place amongst trees in Britain. For centuries, its durability, strength and attractiveness have made it the timber of choice. When the British state was forged in he seventeenth and eighteenth centuries, these qualities made it a metaphor for the virtues of the nation. This book tells the story of man’s use of ot wonderful nz — resource a that oak ide. The fond still has a rich future, both as material and asa key element i 5 authors are concerned with how people have managed and exploited oa ea over time and with the uses to which oak timber has been put, in ships, furniture and buildings. As practicing foresters, they revisit with an experts eye the ae anaes ee of the oats methods of a sca P : life acorn to standard. They reveal the skills nee de dto ore with oak timber, and tell the story of the great ne of iron- smelting and Aan which relied on the tree. They also explore the myths, symbols and cul- tural associations that have connected people in Britain with the oak over hundreds of years. An appendix lists over 700 p ene significant oak trees, with notes on their location, present con- dition and historical connections. This book is a cultural history not only of a tree, but also of a timber. It reclaims the disappearing forestry and carpentry skills of our ancestors and shows how, in an era of climate change, oak can enrich our future as well as our past. ing Ye using mane aging Oo. Esmond Harris has spent a oe a: 2 as a forester, and is a Past Director of the Royal Forestry Society. He wrote The Guinness Book of Trees 1981), Trees.986), and Arboles (1998). Jeanette Harris is a farmer and author. Together they wrote the best-selling Reader’s Digest Guide to the Trees and Shrubs of Britain (981) and Wildlife Conservation in Managed Woodlands and Forests (1997). They run a small farm in Cornwall, where their renovation of woodlands won the 2002 Duke of Cornwall’s Award for Forestry and Conservation. N. D. G. James, OBE, was President of the Royal Forestry Society and author of several tree books. He died in 1993, having laid the foundations for this book.—Gary Jennings, Botanical Research Institute of Texds, 509 Pecan Street, Fort Worth, TX 76102- 4000, U.S.A SIDA 21(1): 40. 2004 NOTES ON NORTH AMERICAN ARCTIC AND BOREAL SPECIESOR ERIGERON (ASTERACEAE: ASTEREAE) Guy L.Nesom David F. Murray Botanical Research Institute of Texas University of Alaska Museum treet 907 Yukon Dri Fort Worth, Texas 76102-4060, U.S.A. Fairbanks, Alaska 99775-6960, U.S.A. ABSTRACT Eri lifl icted to the western U.S.A. and southwestern Canada and is treated here to inelude reed cee poplton CE. pul ea as well as triploids (E. grandiflorus sensu stricto). The distinction between E 1 E. simplex has been made primarily on the ba- sis of differences in size and p a levee but identifications based on morphology often are arbi- trary because differences appear to be widely overlapping. Plants previously identified as F. grandiflorus var. arcticus Porsild are here treated as Erigeron porsildii Nesom & Murray, nom. nov. Distinctions are clarified among E. grandiflorus sensu stricto, E. porsildii, E. muirii, E. yukonensis, ane - oo i ROrOREneT lS E Bo eal nes not eae eoente popone a type collection 4 ] i unknown. E ‘igerondenalti 7] - ( includes E. erie var. dilatatus and Bo mexiae and is contrasted with EF. pallens (an endemic of southwestern Canada) and FE. purpuratus. Recent collections Sominieu the occurrence of E. ochroleucus in arctic and boreal Alaska and immediately aguaaey Yukon, disjunct more than 2800 kilometers from the closest localities in its main range to the south. RESUMEN Erigeron grandiflorus esta restringido al oeste de Estados Unidos y suroeste de Canada y es tratado aqui pa ra incluir las poblaciones diploides extensas (E. simplex) asi como a las triploides (E. stricto). La distincion entre E. Bee y E. simplex ha sido hecha en base a las diferencia de tamano y ae de ploidia, pero las pepe acienes pasadas en la morfologia son a Plantas identificadas rc E. gra anaifto orus vat. arctic Porsild se dee aqui como Erigeron porsildii Nesom & Murray, nom. nov. Se clarifican la entre E. grandiflorus sensu stricto, E. porsildii, E. muirii, E. yukonensis, y la asiatica E. apn — hulteniit no ha sido — a es después de la coleccion tipo Smala cuyas Erigeron denalii incluye E. lente var. Pie sig E. mexiae y se contrasta con E. vallens fam endemismo del suroeste de Canada) y E. purpuratus.R aexistencia de E.ochroleucusen Alaska artica y boreal y en Yukon ae og cae ee dB IRNN ‘2 desde las localidades mas proximas en su area principal en el sur. = Various taxonomic problems regarding alpine, arctic, and boreal Erigeron in North America have been brought into focus during preparation of a treatment of the genus for the Flora of North America (FNANM) volumes. Here we exam- ine some of those problems in detail and provide expl ion and documenta- tion for new taxonomic interpretations. SIDA 21(1): 41-57. 2004 42 BRIT.ORG/SIDA 21(1) Status of Erigeron grandiflorus Erigerongrandiflorus Hook. was described from southern Alberta, Canada (see typification below), and has since been recognized as a species distributed in alpine regions of the western U.S.A. and adjacent Canada and disjunct north- ward into arctic Canada and Alaska. Specimen citations by Cronquist (1947) for E. grandiflorus were mostly from arctic collections and his descriptive mea- surements reflected this inclusive view. The arctic plants were recently segre- gated as E. grandiflorus subsp. arcticus Porsild and are here treated as a sepa- rate species (see topic below). Erigeron simplex Greene, which has been recognized as a widely distrib- uted species of alpine habitats in the western U.S.A,, is very similar to E. grandiflorus. Cronquist found overlapping differences between these species in leaf shape and vestiture, number of pappus bristles, and outer pappus mor- phology. After separating the arctic plants from Cronquist’s concept of E. grandiflorus, we find that features defining E. grandiflorus and E. simplex are even more strongly overlapping. Since Cronquist’s monograph (1947), and apart rta- tion study of arctic and alpine species (1971), Erigeron grandiflorus has been rec- ognized in the conterminus U.S.A. only in Montana (Dorn 1984) and Colorado (Weber 1987, 1990; Weber and Wittman 1992). In these treat ts, contrasts of E. grandiflorus with E. simplex largely repeated measurements from Cronquist’s study. Both E. grandiflorus and E. simplex were included in the Alberta flora by Moss (1959), also using species descriptions essentially taken from Cronquist (1947). Scoggan (1979) included E. simplex as a questionable member of the Ca- nadian flora, based on the sole record from Moss, but Packer's revision of the Alberta flora (Moss 1983) treated only E. grandiflorus, with no mention of E. simplex. Eri- gerongrandiflorus also has been recognized in British Columbia by Douglas et al. (1998), whose descriptive measurements gave broader ranges than Cronquist’s. Spongberg (1971) regarded Erigeron grandiflorus strictly as an apomictic triploid (compared with strictly diploid E. simplex) and documented its occur- rence in Utah, Colorado, and Wyoming. At some localities, he found triploids “growing intermixed with plants of a larger population of Erigeron simplex” (e.g., Clear Creek /Grand Co. line, above Berthoud Pass, Spongberg 67-243, TEX). He did not provide a key, but from his comments and annotations, E. grandiflorus in southern Canada and the western United States differed only quantitatively in his concept from the more widespread E. simplex, with in- volucres and florets at the higher end of the ranges of size measurements. Spongberg (1971, p. 200) also noted that “because of the intergrading of mor- phological features of plants of Erigeron grandiflorus ... the single most impor- tant criterion indicative of this taxon is highly irregular [in shape] and greatly abortive pollen.” These pollen features result from meiotic anomalies associ- ated with the triploid condition. aa) ‘rom § | ‘SC L Oo Pee AND BOREAL SPECIES OF ERIGERON 43 We observe that triploids (including the largest-headed plants) appear to be consistently stipitate-glandular over the whole stem with glandular tri- chomes mostly 0.2-0.3 mm high, while the diploids (smaller-headed plants) commonly are essentially eglandular or stipitate-glandular with shorter tri- chomes only just beneath the heads. Many smaller-headed plants, however, are variably glandular, trichome sizes vary, and all collections of Erigeron simplex from Arizona and Utah appear to be stipitate-glandular. At least one collection of relatively small-headed plants was counted as triploid and identified by Spongberg as E. grandiflorus (Spongberg 69-30, TEX). Spongberg hypothesized that the triploid plants (= Erigeron grandiflorus sensu stricto in his view) incorporate a genomic element from an ancestor other than E. simplex, but we find that morphological distinctions between the ploidal races are too arbitrary to allow consistent identification. Until more convincing evidence is at hand regarding the evolu ony divergence of these taxa, and until some way might be found to dist hthem with more precision, E. grandiflorus is treated here as including the plants generally identified as E. simplex. Erigeron grandiflorus Hooker, Fl. Bor-Amer. 2:18, plate 123. 1834. Type: CANADA. [ALBERTA]. “Summits of the Rocky Mountains,” Drummond s.n. (GH!, KEW?). Non Nea lead non Hoppe ex DC. 1836; non Sessé & Mocino 1894. Drummond's col- lectio was made in June or July, 1826 (Drummond 1830), in the vicin- ity tape or between Jasper and “Lac-la-Pierre,” which is about 60 miles north of Jas asks a Greene, FL. Francisc. 387.1897. LECTOTYPE (Cronquist 1947): U.S.A. Colorado: no her data, 1875, E.L. Greene s.n. (ND-G). poets leucotrichus Rydb., Bull. Torrey Bot. Club 28:23. 1901. Type: U.S.A. WYOMING. Big Horn Mountains, 8000 ft, Jul 1899, F Tweedy 2003 (HOLOTYPE: NY)). Plants perennial, from short, horizontal or erect, fibrous-rooted rhizomes, cau- dex essentially unbranched or with short, thick branches. Stems 2-25 cm high, erect to basally decumbent-ascending, sparsely to moderately pilose to villous- hirsute, variably stipitate-glandular over whole or part of stem, sometimes es- sentially eglandular. Leaves basal and cauline, basal persistent, oblanceolate to obovate or spatulate, apically rounded, 1-6(-9) cm long, 2-6(-14) mm wide, entire, cauline quickly or gradually reduced upward, not subclasping, sparsely hirsutulous or villous to sparsely strigose or glabrate, sometimes sparsely glan- dular. Heads |; involucres 5-8(-10) mm high, 8-20 mm wide; phyllaries in 2-3 series, green or purplish, moderately to densely woolly-villous with flattened hairs, sometimes with reddish crosswalls, minutely glandular at least near the tips. Ray florets 50-130, corollas 7-11(-15) mm long, laminae blue to pink or purplish, rarely white, coiling. Disc corollas 2.4-4(-5) mm long, throat not in- durate or inflated. Cypselae ca. 1.8-2.4 mm long, 2-nerved, strigose; pappus of (7-)10-18(-22) bristles, with an outer series of narrow scales. 2n = 18, 27 (Spongberg 1971; numerous diploid counts in literature as Erigeron simplex). 44 BRIT.ORG/SIDA 21(1 Flowering Jul-Aug (-Sep). Rocky sites, meadows, alpine or near timber- line; 2900-4200 m. Canada (Alberta, British Columbia); U.S.A. (Arizona, Colo- rado, Idaho, Montana, Nevada, New Mexico, Oregon, Utah, Wyoming). Status of E. grandiflorus sensu stricto ane E. isi das subsp. arcticus The type collection of Erigeron grandiflorus and later collections from the same region (southwestern Alberta and adjacent British Columbia) are disjunct by more than 1500 kilometers from the more northern range of E. grandiflorus subsp. arcticus Porsild. The southern populations are located at the northern tip of the range of the species identified here as E. grandiflorus sensu stricto, including E. simplex. Porsild (1955) observed that E. grandiflorus comprised three “races”: Rocky Mountain, arctic, and Alaskan. The ‘Rocky Mountain’ race corresponds to E. grandiflorus sensu stricto, while we are unable to separate the “arctic” and “Alaskan” races and regard them both as composing Porsild’s subsp. arcticus. Our study of these two entities concludes that they are distinct by anumber of nonoverlapping features and that both are appropriately treated at specific rank. 1. Basal leaves oblanceolate to obovate or spatulate, 2—6(-14) mm wide, apically rounded, cauline oblanceolate to narrowly lanceolate, never subclasping; villous involucral vestiture ae hairs, often with reddish crosswalls; ray corollas 7-11( 15) mm long; 2n = Erigeron grandiflorus 1. Basal He a ne to narrowly obovate, (3-)5-14 mm wide, apically cute, cauline narrowly ovate to ovate-lanceolate or lanceolate, often subclasping; villous involucral vestiture of whitish hairs, without colored crosswalls; ray corollas 13-17 mm long; 2n = 36 Erigeron porsildii Erigeron porsildii Nesom & Murray, nom. nov. Erigeron grandiflorus Hooker subsp. rcticus Porsild, Mus. Natl. Canada, Publ. Bot. (Ottawa) 4:67. 1975. TYPE: CANADA. NORTHWEST TERRITORIES: Victoria Island, SW coast, vicinity of Holman Island trad- ing post, dry, gravelly slope, 8 Aug 1949, A.E. Porsild 17342 (HOLOTYPE: CAN, color image!, photos in Porsild 1955! IsoTYPE: ALA!). Non Erigeronarticus Rouy, Fl, France 8:160. 1903. Rouy’s epithet was a variant spelling applied to an “arctic” plant and precludes adoption at specific rank of the similar Porsild name, which denotes the same geography (ICBN, Art. 53, Ex. 9). In order to retain the original type des- ignation, we have chosen to rename the taxon at specific rank. The replacement A.E. Porsild (1901-1977), whose studies and publications over a 60 year period greatly advanced knowledge of the flora of arctic and boreal America. —— — Plants} ial, from a short, horizontal or erect, fibrous-rooted rhizome (some- times appearing more like a taproot), sometimes with short caudex branches. Stems (2-)10-20(-25) cm high, erect, sparsely to moderately villous with hairs 0.5-1.6 mm long, usually stipitate-glandular over whole stem with hairs 0.05- 0.4 mm long. Leaves basal and cauline, basal oblong-oblanceolate to narrowly obovate, 3-12 cm long, (3-)5-14 mm wide, entire, cauline oblong-lanceolate to lanceolate, often subclasping, gradually reduced upward or nearly equal-sized, densely hirsute to coarsely villous, sparsely stipitate-glandular to minutely AND BOREAL SPECIES OF ERIGERON 45 glandular. Heads 1; involucres 6-10 mm high, 12-20 mm wide; phyllaries in ca. 2 series of equal length, narrowly lanceolate, apically acuminate, purple at least at the tips, densely hirsute, hirsute-villous, or villous with whitish hairs with- out colored cross-walls, sparsely stipitate-glandular to minutely glandular. Ray florets 65-110, corollas 13-17 mm long, laminae 1.2-L.7 mm wide, blue to purple or lavender, less commonly white, weakly coiling. Disc corollas 3.8-4.5 mm, puberulent with glandular hairs, throat not indurate or inflated. Cypselae 2- 2.5mm, 2-nerved, sparsely strigose; pappus of 14-20(-25) bristles, with a promi- nent outer series of setae or narrow scales. 2n = 36 (Dawe & Murray 1981). Flowering mid Jun-Aug(-Sep). Alpine ridges and slopes, rock outcrops, cliffs and talus (often calcareous) slopes, shaly gravel and scree, bluffs, grassy ravines, tundra, meadows; (150-)600-1600(-2100) m. Canada (N.WT: Districts of Franklin, pee Yukon); U.S.A. (Alaska). Additional col d. Alaska. Wrangell-St. Elias ee Park and Preserve, eee Quad, oo Mts., vic. plateau W of Nizina River, above head of Nikolai Creek, SE-facing shaley scree near ridge crest, 1370 m, 24 Jul 1996, Batten and Barker ae (ALA); Mt. McK ne National Park, Cathedral Mountain, mile 36, 1-10 Jul 1964, ea s.n.(NY); Demarcation Point Quad, Kongakut river, 30 km S of Beaufort Sea coast, S-facing river bluff, dominant willow, 29 Jun 1984, McCarthy s.n. (ALA), Tetlin National Wildlife Refuge, N oo an Mentasta Mts., vic. ee River valley,‘4940 Hill’ site, alpine screes and seeps, 1506-1628 m, 25 Jul 1996, Moran 45 (ALA), Mt. McKinley Quad, mountain slope along Stony Creek, a 1941, Murie s.n. (ALA); Philip Smith Mountains Quad, Yukon River-Prudhoe Bay Haul Road, just E of Galbraith L, tundra slopes and conglomerate out- crops, 915-1220 m, 20 Jul 1976, Murray zi aie ie Jonnson (AL A Circle Ques sUKOE ee Up- lands, nouth Hous Birch Creek drainage, al | g Windy Creek, in mica-schist bedwodle ie dry, S- eae herbaceous slope ae ee 1100-1220 m, 8 Jul 1996, Parker et al. 6537 (ALA); Gates of the Arctic National Park and Preserve, Chandler Lake Quad, Arctic Foothills, Castle Mtn., northern ridge of summit area, 700-1070 m, al- pine re heath, cliffs, scree aah meadows along drainages, in lush herbaceous, N-facing meadow, 30 Jul 2002, Parker et al. 1 ALA); Baird Mountains Quad, Noatak National Preserve, Central ee River valley, ee aie vic. of “Wolf control” cabin, ca. 7 km upriver from Mukachiak Cr. mouth, S-facing acidic bedrock bluffs above floodplain, dry mesic shrub and open shrub-herba- ceous vegetation, growing in open, grassy slope, 140-200 m, 19 Jul 2003, Parker et al. 14959 (ALA, BRIT); Baird Mountains Quad, Noatak National Preserve, Central Noatak R. valley, Sekuiak Bluff, vic. of upstream end of bluff and VABM Windy, N bank of river, L1OO-150 n eee outcrops and scree, limestone-acidic contact zone, 27 Jul 2003, Parker et al. 15264 (ALA, BRIT nali National Park and Preserve, Healy Quad, wen Range, S-facing slopes of main ridge NW i ae, Creek, 1 km SSW of VABM Riley, confined to turfy, graminoid-forb meadow-tundra area on steep, S-facing slope, 3700 ft, 18 Jun ee Roland and ae 3821 (ALA). Yukon. Kluane National Park an Preserve, Dezadeash Quad, St. Elias Mts., King’s Throne, vic. Kathleen Lake and Haines Road, 24 Jul 2000, Caswell 359 (ALA); setae Mts., fine broken stones and turf in saddle and adjacent slopes, 2000 ft, 13 Jul 1982, Cody and Ginns 31788 (AL A): se of Herschel Island, Me ner Bay, Beaufort Sea, slope above stream, 20 Jul 1975, Cooper 75E (NY), 14 Jul 1978, Cooper 634D (NY), 19 Jul 1975 Cooper 54A (NY); Herschel Island, 14 Jul 1906, ae sn. (NY) Kluane Lake Quad, NW of Slims River, 4500-6300 ft, 9Jul 1967, Murray 933 (ALA) Erigeron porsildii contrasted with E. koraginensis A range of opinions regarding the definitions and distinctions (or lack of dis- 46 BRIT.ORG/SIDA 21(1) tinction) among Erigeron grandiflorus, E. porsildii (= E. grandiflorus subsp. arcticus), E.muirii,and the Asian species FE he raginensis is summarized in Elven et al. (2003, as related by Aiken et al. 2003). It has not been clear whether E. koraginensis occurs on the North America continent or whether it might be conspecific with an earlier-named American species, particularly E grandiflorus/porsildii. Erigeron koraginensis and E. porsildii are the most simi- lar among these species, but here, based on material housed at ALA, we con- clude that E. koraginensis and E. porsildii are distinct and note that we have seen no collections of FE. koraginensis from North America. 1. Leaves sparsely to moderately villous on both surfaces; villous Dist aae vestiture of whitish hairs, without colored crosswalls; ray florets 65-110, corollas m long; disc corollas 3.8-4.5 mm long; outer pappus a prominent series of ae seta or scales Erigeron porsildii |. Leaves glabrous or essentially so on adaxial surfaces, villous abaxially; villous in- alice vestiture of hairs with purple crosswalls; ray florets 45-73, corollas 9-13 mm long; disc corollas 3-3.9 mm long, outer pappus of a few minute setae a narrow scales rocisn koraginensis mat koraginensis (Komarov) Botschantzey, Not. Syst. Herb. Inst. Bot. Acad. i. URSS 16:391. 1954. Aster koraginensis Komarov, Fl. Penins. Kamtsch. 3:125. 1930, Erigeron komarovii Botschantzev es i is (Komarov) Voroshilov, Byull. Glavn. Bot. Sada (Moscow) 84:34. 1 Erigeron komarovii Botschantzev, Not. Syst. Herb. Inst. Bot. Acad. Sci. URSS 16:391. 1954. Aster consanguineus Ledeb., Fl. Ross. 2, 2:473. 1845. Erigeron consanguineus(Ledeb.) Novopokr, Bot. Mat. (Leningrad) 7:137. 1938 (non Cabrera 1937). Plants perennial from lignescent thickened but elongate rhizomes 1-4 cm long. Stems erect, 4-25 cm high, sparsely to moderately villous, densely villous be- neath the heads, hairs 0.5-1.5(-1.8) mm long, often with colored crosswalls, stipi- tate glands 0.1-0.3 mm high with colored cross walls. Leaves mostly basal or basal and cauline, basal oblanceolate to oblanceolate-spatulate with rounded to acute apices, 2-7 cm long, 2-10 mm wide, cauline oblanceolate to narrowly lanceolate, Gif present) quickly or gradually reduced on lower 1/3-2/3 of stem, not clasping, sparsely villous and minutely and short-stipitate glandular abaxially, glabrous adaxially, margins ciliate and stipitate-glandular. Heads 1; involucres 6-9 mm high, 11-18 mm wide; phyllaries linear-lanceolate, usually purple, in 2 series of equal length, moderately to densely villous, hairs usually with purple crosswalls (at least in proximal cells), stipitate glandular. Ray flo- rets 45-73, corollas 9-13 mm long, 1.2-2 mm wide, blue to white, coiling. Disc corollas 3-3.9 mm long, very sparsely villous, lobes narrowly triangular. Cypselae 2.2-2.6 mm long, narrowly oblong in outline, sparsely strigose-vil- ous, 2-nerved; pappus bristles 19-23, of uneven thickness and length, outer se- ries a few minute setae and narrow scales. Description drawn from nine ALA collections from northeastern Russia; it corresponds closely to the description — — AND BOREAL SPECIES OF ERIGERON 47 of E. komarovii provided by Botschantzev (1959). 2n = 18 (summary in Aiken et al. 2003). The names Erigeron koraginensis and E. komarovii were proposed in 1954 on the same page of publication. Erigeron komarovii was a replacement name for the later homonym E. consanguineus (1938), which was based on Aster consanguineus (1845); E. koraginensis was based on Aster koraginensis (1930). We follow the observations of Dr. Kanchi Gandhi (pers. comm.) in recognizing the priority of E. koraginensis if E. komarovii and E. koraginensis are combined: “Although the priority of the type of E. komarovii starts from 1845, the priority of the epithet itself starts from 1954. In contrast, the priority of the type and of the epithet in E. koraginensis start from 1930.” Erigeron koraginensis is distributed in arctic and eastern Siberia, Wrangel Island, and the Kamchatka region (Botschantzev 1959; Tzelev 2002). Yurtsev et al. 1989 said explicitly that E. komarovii occurs on Wrangel Is. and replaced the name EF. muirii erroneously used in prevous accounts. Czerepanov (1995) listed E. koragi isas a synonym of E. komarovii; Botschantzev (1959) maintained them as separate species but by widely overlapping differences. The type of E. koraginensis was collected in Kamchatka; the type of E. komarovii was collected from Karaginskii Island, in the Bering Sea just off the coast of northeast Kamchatka. Taxonomic status and distribution of Erigeron muirii Hultén (1968) treated Erigeron muirii Gray as a subspecies of E. grandiflorus Ga) 2 as identified here), noting that its densely lanate leaves and in- tinguish it from the typical expression of the latter. Elven et al. (2003) also note that ‘The hairs of E. muirii are very thin, curly, and intermingled, and they are totally different from the stiffer and straighter hairs of E. grandiflorus and E. koraginensis.” Erigeron muirii and E. porsildii occur sympatrically in northeastern Alaska, but E. muirii usually grows in exposed, rocky settings, whereas E. porsildii tends to favor meadows. Chromosome counts from Alaska show E. muirii to be diploid, E. porsildii tetraploid (Dawe & Murray 1981). Two unusual plants on a sheet with six others of typical E. muirii may be hybrids between EF. muirii and E. uniflorus var. eriocephalus J. Vahl) Boivin (Alaska: vicinity of Cape Thompson, 26 Jul 1976, Roseneau s.n., ALA), but we have not seen evidence of hybridization or intergradation between E. muirii and E. porsildii and agree with Cronquist (1947), Porsild and Cody (1980), Cody (2000), and others in observing that E. muirii is appropriately treated at specific rank. Erigeron muirii A. Gray, Proc. Amer. Acad. Arts 17:210. 1881. Erigeron grandiflorus Hook. subsp. muirii (A. Gray) Hultén, Ark. Bot. 7:132. 1968. TyPE: U.S.A. ALASKA. Cape Thompson, 1881, J. Muir s.n. HOLOTYPE: GH)). 1. Stems and leaves moderately lanate, involucres densely lanate, all eglandular;cauline leaves narrowly elliptic-lanceolate, not subclasping Erigeron muirii 48 BRIT.ORG/SIDA 21(1) 1. Stems and leaves sparsely villous, involucres villous, all glandular to stipitate-glan- dular; cauline leaves narrowly ovate to ovate-lanceolate or lanceolate, often subclasping Erigeron porsildii Erigeron muirii has been regarded as an endemic of northern Alaska (Murray & Lipkin 1987, Lipkin and Murray 1997), but a herbarium sheet of E. muirii at US() bears the label “Wrangel Is., Dr. Ross, Corwin 1881.” Hultén (1950) com- mented that personal communication from Soviet botanists led him to doubt the occurrence of E. muirii on Wrangel Island (in the Arctic Ocean, between the East Siberian Sea and the Chukchi Sea, off northeastern Russia) and to hy- pothesize that the specimen at US is incorrectly labeled. Nevertheless, he felt that this explanation needed confirmation. Knowing of the specimen at US and lacking authentic material of Erigeron muirii at LE, VV. Petrovsky compared the written descriptions and the photo- graph in Hultén (1950) of E. muirii with specimens of E. komarovii (= E. koraginensis, as recognized here, see section above) and concluded they were conspecific. Petrovsky (1987) therefore included E. muirii in the Arctic Flora USSR, under which he included E. komarovii Botsch. as a synonym (among many synonyms). Following exchanges of specimens between LE and ALA, it was possible for both sides to examine authentic E. muirii and E. komarovii and to determine that they are not the same. Yurtsev et al. (1989) removed E. muirii from their list of Wrangel Island plants. The voyage in which Muir participated had numerous ports-of-call, includ- ing Cape Thompson (the type locality of Erigeron muirii, in northwestern Alaska) and Wrangel Island. In accounts of this voyage published by Muir (1883, 1917), plant lists for Wrangel Island do not include any species of Erigeron. It is reasonable to presume that a mistake was made during processing of the speci- mens and that a Wrangel Island label was assigned to an Alaskan collection, just as Hultén had earlier surmised. Specimens from Herschel Island on the coast of the northwestern extreme of Canada very near the Alaskan border were included by Hultén (1968) in Eri- geron muirii(as E. grandiflorus subsp. muirii). We have determined these speci- mens as E. porsildii and note that other collections from Herschel Island are cited above for E. porsildii. Collections of E. muirii from which our discussion is drawn are mostly at ALA. The status of Erigeron hultenii Erigeron hultenii was noted by its author to be “closely allied to the polymor- phic arctic-alpine species E. grandiflorus WJ. Hooker” (Spongberg 1973, p. 116) and to have a “close morphological resemblance to some plants of E. grandiflorus from Alaska” [= E. porsildii in the present sense] (p. 119). Contrasts with E. porsildii, however (as in the couplet below), exclude it from that species. De- spite additional exploration at and near the type locality, plants corresponding — AND BOREAL SPECIES OF ERIGERON 49 to E. hultenii have not been recollected and we do not find that E. hultenii fits with any other known Alaskan species. We continue to regard it as an anoma- lous plant of unknown relationships. ]. Leaves 1-velr 1ed; ray florets 65-1 10; disc corollas 3.8-4.5 mm long; -achenes 2-nerve ed; outer pappus a prominent series of long setae or scales Bagerton porsildii . Leaves 3-veined; ray florets 20-45; disc corollas 2.5-3 mm long; achenes 4-nerve outer pappus barely if at all evident Erigeron hultenii Erigeron hultenii Spongberg, Rhodora 75:116. 1973. Type: U.S.A. ALASKA: Campbell Creek Valley, 11 mi N of Anchorage, 7 Aug 1965, L. Strutz s.n. (HOLOTYPE: S). Taxonomic status of Erigeron yukonensis Erigeron yukonensis was included within the “E. grandiflorus agg.” by Polunin (1959), but Hultén (1950) hypothesized it to be closely related to E. glabellus Nutt. Later, Hultén formalized this (1967) by reducing E. yukonensis to varietal rank within E. glabellus. We agree with a number of recent botanists who find E. yukonensis to be a distinct species, although its evolutionary relationships are not clear—its eglandular vestiture, elongate basal leaves, and 1-4 heads are un- characteristic of most of the arctic monocephalous Erigeron species. Plants of E. porsildii with elongate-lanceolate leaves sometimes approach single-headed E. yukonensis in aspect, but E. porsildii differs especially in its stipitate-glan- dular stems and its tendency for oblong-lanceolate to lanceolate, often subclasping cauline leaves. Erigeron yukonensis Rydb., Bull. New York Bot. Gard. 2:185. 1901. Type: CANADA. KON: Dawson, 23 Jul 1899, R.S. Williams s.n. (HOLOTYPE: NY). Erigeron glabellus Nutt. var. yukonensis (Rydb.) Hultén, Ark. Bot. 7(1):132. 1967. Plants perennial, fibrous-rooted rhizome, sometimes appearing like a taproot, multicipital. Stems 10-30(-40) cm high, erect to basally ascending, often pur- plish at base, sparsely to moderately hirsute to villous-hirsute, eglandular. Leaves basal and cauline, lower narrowly oblanceolate, rarely oblanceolate-obovate, entire, (1-)4-16 cm x 2-8 mm, middle and upper reduced and narrowly lan- ceolate, not clasping, sparsely hirsute to glabrous, eglandular. Heads |-2(-4) from branches at midstem; involucres 6-10 mm high, 12-19 mm wide; phyllaries in 2(-3) series, linear-lanceolate with linear-acuminate tips, sparsely to moder- ately woolly-villous with whitish, flattened hairs without colored cross-walls, eglandular to very sparsely minutely glandular. Ray florets 42-82, corollas 10- 15mm long, laminae white to pink or pink-purple, coiling. Disc corollas 3.8-5.2 mm long, throat not indurate or inflated. Cypselae 1.7-2 mm long, 2-nerved, strigose; pappus of 15-20 bristles, with an evident outer series of short bristles, squamellae, or scales. Flowering Jun-)Jul-Aug (-Sep). Rocky sites and meadows, near timber- line and above; 2850-3150 m. Canada (N.WT: District of Mackenzie; Yukon); U.S.A. (Alaska). 50 BRIT.ORG/SIDA 21(1) Additional ¢ i camined. Canada. Yukon: S end of Lake Kluane, alpine, 24 Jul 1944, Ander- son 9472B (NY); Moosehide ae sie W-lacing slope, open rocky slope, 20 Jun 1949, Calder 3266 (NY); Lower Klondike, last half of Jun 1919, Cockfield 64 (CAN-photo ALA!) near Collinson Head, E end of Herschel Island, van we Bay, Beaufort Sea, 16 Jul 1975, Cooper 38A (NY); Dawson, 30 Jun 1914, Eastwood 454 (CAN-photo ALA}; Red Mountain near Fort Selkirk, shady slopes, 17 Jul 1899, Gorman 1109 (NY); Ft. Selkirk, 1-9 Jul 1891, Hayes s.n. (NY); White Horse, Yukon River, 1 Sep 1902, Macoun 78996 (NY, CAN-photo ALA); Fort Selkirk, 18 Jul 1899, Tarleton 141 (NY). NWT: Inuvik, damp vella and Cysto rock ledges by Dolomite Lake in open sunlight, just above ledge with Woodsia gla meris fragilis, 7 Jul 19605, Swales 522 (CAN-photo ALAI). roe Cronquist (1947) cited a collection of Erigeron yukonensis from District of Mack- enzie (Richards Island, 1896, Stringer 16, no herbarium cited). Wiggins and Tho- mas (1962) cited an Alaskan collection in AMES from north of the Brooks Range (Utukok River, 5 Jun-15 Jul 1947, Thompson s.n.), but the a of this collec- tion needs to be verified. Cody (2000, p. 581) noted that E. yukonensis occurs in “the Yukon Territory west of latitude 135 W, in northern Disteiet of Mackenzie east to Coronation Gulf and in Nahanni National Park.” See also Map 1104 in Porsild and Cody (1980) and a map in McJannet et al. (1993, p. 38). The Erigeron purpuratus—E. pallens group Treatments of these species have varied considerably, but we recognize Erigeron purpuratus Greene, FE. denalii A. Nels., and E. pallens Cronq. as distinct species. Erigeron pallens is rare and endemic of the high mountains of southeastern British Columbia and adjacent Alberta, while the other two are more common and occur in Alaska, Yukon, District of Mackenzie, N.WT., and extreme north- western British Columbia, long disjunct from the range of E. pallens. The erect, inrolled laminae of the ray florets (giving them a “filiform” aspect) of all three species distinguishes them from other species of Erigeron sect. Tridactylia Nutt., of which they are members. The leaf Red: of E. pallens, however, is more simi- lar to that of E. vagus Payson and E. flabellifolius Rydb. (also sect. Tridactylia) of the northwestern US.A., which are geographically closer to E. pallens than E. denalii and E. purpuratus. Frigerondenalii has been treated asa synonym ora variety of E. purpuratus (e.g., Cronquist 1947, Hultén 1968, Boivin 1972). They are similar in many ways, including the characteristic purplish pappus bristles, and perhaps are most closely related to each other, but the two taxa appear to be partially sympatric and differences in vestiture and leaf morphology are constant. The habitat of E. purpuratus, sandy or gravelly alluvium, also is remarkably constant and dis- tinct from that of E. denalii. Probably correlated with the loose sands and grav- els of its habitat, plants of E. purpuratus almost always developa long and rela- tively slender but woody taproot, taproots of E. denalii are less well defined and often do not show on collections, and the plants usually develop slender rhizome-like caudex branches, well adapted to the downslope instability of the screes typically inhabited by the species. — ues AND BOREAL SPECIES OF ERIGERON 3 1. Leaves entire or shallowly 3-lobed at the obtuse to nearly flat apex; pappus tawny- white to yellowish Erigeron pallens ], Leaves entire or with 1-2 enalloyy lobes on aI en pappus usually sae 2. stems 1 —5 cm tall: te-spatulate, 2—4(- wide, minutely glandular and moderately vot, involuctes eeneey pigs: ______Erigeron denalii 2. Stems (2-)3-9(-14) ¢ | ly oblanceolate, 1-2.5(-3) mm wide, merely minutely Slane or sie Sey hirsute-villous; eee sparsely pilose Erigeron purpuratus Erigeron purpuratus Greene, Pittonia 4:155. 1900. Type: CANADA. YUKON: Fort Selkirk, sandy river banks, 28 Jun 1899, M.W. Gorman 1065 (ND-G)). Plants perennial, taprooted, from a diffuse caudex divided into system of slen- der rhizome-like branches. Stems (2-)3-9(-14) cm high, minutely glandular, villous. Leaves mostly basal, linear- to narrowly oblanceolate, 1.5-3(-4.5) cm long, 1-2.5(-3) mm wide, entire or (mostly on early-produced leaves) with | or 2 shallow lateral lobes, minutely glandular, sparsely villous or without nonglandular hairs. Heads 1; involucres 6-9(-10) mm high, 9-15 mm wide; phyl- laries in 2-3 series, minutely glandular, sparsely villous, hairs with colored cross- walls and up to 3 mm or without nonglandular hairs. Ray florets 40-90, 5-9 mm long, laminae filiform, 0.3-0.8 mm wide, erect, white to pink or purplish, not coiling or reflexing. Disc corollas 4-5 mm long, throat not indurate or in- flated. Cypselae 1.8-2.7 mm long, 2-nerved, sparsely strigillose to glabrate; pa- ppus of 28-40 pinkish to purplish bristles, without an evident outer series. 2n = 18 (Dawe & Murray 1979). Flowering Jun-Aug. Sandy and gravelly stream banks, gravel bars, sandy beaches, gravelly slopes, alpine tundra; 150-1350 m; Canada (Yukon); U.S.A. (Alaska). Erigeron denalii A. Nelson, Amer. J. Bot. 32:289. 1945. Type: U.S.A. ALASKA. Mt. McKinley National Park: Sable pass, at highest point of Park road, rocky slopes, ca. 4000 ft, 8 Aug 1939, A. Nelson and R.A. Nelson 4058 (HOLOTYPE: RM ISOTYPE: US)). ath i a Greene var. dilatatus Boivin, Phytologia 23:134. 1972. TypE: CANADA. Dis- F MACKENZIE: Redstone River Region, 4 July 1963, Kvale & Haggard 131 (HOLOTYPE: oo. Erigeron mexiae K. Becker, Brittonia 28:144. 1976. TYPE: CANADA. YUKON: Dempster Highway, N awson City, E-facing slope just W of Mile 41, in scree, 5000 ft, 8 Jul 1973, C.W. Greene 481 (HOLOTYPE: NY}; ISOTYPE: UAC). Plants perennial, taprooted, with a diffuse, slenderly branching caudex. Stems 1-5cm high, villous, granular-glandular. Leaves basal and cauline, spatulate to oblanceolate, entire or shallowly 3-lobed at the apex, 1-2 cm long, 2-4 mm wide, mid-cauline larger than the basal and lower cauline, villous, granular-glandu- lar. Heads 1; involucres (4-)5-7 mm high, 9-12 mm wide; phyllaries in 2(-3) series, densely lanate-villous, the hairs to 3 mm long, often with dark purple crosswalls, glandular. Ray florets 30-55, 4-8 mm long; laminae erect, rarely spreading, filiform to narrowly straplike, 0.3-1.2(-2) mm wide, white to laven- 52 BRIT.ORG/SIDA 21(1) der. Disc corollas 3.8-4.8 mm long, throat not indurate or inflated. Cypselae oblanceolate-oblong, 3-3.8 mm long, 2-nerved, strigillose to strigose-hirsute; pappus of 20-25 bristles, more or less purple-red tinged or sometimes whitish (variable within populations), without an evident outer series. Flowering Jun-Aug. Open alpine and subalpine habitats, tundra slopes, openings in spruce-fir woods, gravelly and shaley scree, cut banks; 900-2150 m: Canada (N.WT: District of Mackenzie; Yukon; extreme northwestern British Columbia); U.S.A. (Alaska). Plants of the type collection of Erigeron mexiae have ray corollas with rela- tively broad laminae (1.5-2 mm wide), but ray corolla width is variable in EF. denalii and E. mexiae is here regarded as an extreme variant within the spe- cies--in other characters there appears to be no difference. Erigeron pallens Cronquist, Brittonia 6:240. 1947. Type: CANADA. BRITISH COLUM- BIA: Glacier, Mount McDonald, 7500 ft, 1 Aug 1913, Butters and Hol way 799 (HOLO- TYPE: MINN). Plants perennial, taprooted, caudex divided into system of slender, rhizome- like branches. Stems 2-4 cm high, sparsely hirsute-villous, minutely glandu- lar. Leaves mostly basal, oblanceolate to spatulate, 1-2.5 cm long, 2-4 mm wide, entire or shallowly 3-lobed at the obtuse to nearly flat apex, sparsely villous, minutely glandular. Heads 1; involucres 6-8 mm high, 9-13 mm wide; phyllar- ies in 2-3 series, commonly purplish or purple at the tips, sparsely villous, the hairs often becoming tawny, without purple cross-walls, minutely glandular. Ray florets 50-65, corollas 4-5(-6) mm long, laminae white to pink or purplish, ca. 0.5-0.8 mm wide, erect. Disc corollas 3.5-4.4 mm long, throat not indurate or inflated. Cypselae length not observed, 2-nerved, sparsely strigose; pappus of 25-30 tawny to purplish bristles, with a few, short, outer setae. 2n = 18 (Packer & Witkus 1982). Flowering Jul-Aug. Alpine talus, shale slides, steep and sparsely vegetated slopes; ca. 2100-3200 m; Canada (Alberta, British Columbia). A nomenclatural combination attributed to G.W. Douglas (Erigeron purpuratus Greene subsp. pallens, such as by Moss 1983 and Packer & Witkus 1982), apparently was never validated and is a nomen nudum. Erigeron ochroleucus in Alaska Previously undetermined collections at ALA document the occurrence of Eri- geron ochroleucus Nutt. in arctic and boreal Alaska and immediately adjacent Yukon, disjunct more than 2800 kilometers from the closest localities in its main range, which runs from Nebraska and South Dakota through Wyoming and Montana into southern Alberta and British Columbia (see map in Nesom 2004). The arctic and boreal plants are unequivocally identified as E. ochroleucus, al- though they differ in some ways from those of the southern system: stem height ranges shorter (2-15 vs. (6-)8-18(-20, 30) cm), leaf length ranges shorter (2-7 ~ AND BOREAL SPECIES OF ERIGERON 53 vs. +-9(-12) mm), leaves are consistently linear to linear-oblanceolate (vs. lin- ear to narrowly oblanceolate, 1-4(-5) mm wide), hairs of the villous involucral vestiture often (vs. uncommonly) have red crosswalls, ray florets range fewer in number ((18-)30-50 vs. 30-62), and the disc florets range greater in length (3.5-4.5 mm vs. 2.8-3.6 mm). The Alaska and Yukon plants plants might be for- mally treated as a variety, emphasizing their geographic disjunction, but mor- phological differences are overlapping and plants from the western U.S.A. can be found that closely match the arctic plants. In the interpretation of Nesom (2004), no varieties are recognized in the southern population system of E. ochroleucus. Erigeron ochroleucus, although very restricted in Alaska and Yukon, is nevertheless common along a section of the Porcupine River from Rampart House in Yukon, but near the International Boundary, downstream below the Lower Ramparts, some 80 kilometers total. The climate is hypercontinental and habitat for this species is at the base of arid slopes with an eastern to southern exposure and adjacent terraces. The vegetation is discontinuous and steppic in character: Artemisia frigida, Bupleurum americanum, Calamagrostis purpurascens, Agropyron spicatum, Phlox hoodii, and Erigeron caespitosus ar prominent. Several species also restricted in Alaska and Yukon, and some are also disjunct from their main ranges: Festuca lenensis, Carex filifolia, Carex duriuscula, Eriogonum flavum var. aquilinum, Alyssum americanum, and Townsendia hookeri. The two arctic localities for E.ochroleucus limatically very different from the Porcupine River, but edaphically similar: dry sands and gravels (Atqasuk) and well-drained soils (Kadleroshilik Pingo). Collections examined. U.S.A. Alaska: Arctic Slope District, 70 air mi S of Point Barrow, near Eskimo village of Atkasook, 2.9 km SE of Meade River Camp, 15 m, 31 Jul 1975, Komarkova 45 (ALA). Black River Quad: Arctic NWR, steep southeast-facing steppe slope on Porcupine River, 366 msm, 66° 58' 56'N, 142° 49' 46" W, 4 June sie Murray, Emers, and Jorgenson 12,128 (ALA), Yukon Flats NWR, steep southeast-facing steppe slope on Porcupine River, 66° 59' 15" N, 142° 58' 20" W, 6 June 1996, rray, Emers, and Jorgenson 12,153 (ALA). Porcupine River, E of old archeology cam p, 66° 59'N, 142° 49'W S facing bluff, 20 m above river shore, 18 Jul 1995, Reitz s.n.(ALA and to be distributed). Porcu- pine River, 67° 20’ N, 141° 18’ W, 20 Jul 1980, Thorson 7-20-80-4 (ALA). First bluff inside Alaskan border [with Yukon, Canada], on Porcupine River, 67° 21'N, 141° 17'W, 275 msm, 17 July 1981, Viereck and Foote 4536 (ALA). Beechey Point Quad, Kadleroshilik Pingo (no. 41), Kadleroshilik study area, 70° OLN, 147° 41'W, gravelly areas, 26-28 Jul 1986, Walker 86-31 (ALA). CANADA. Yukon: Porcupine River, Rampart House, 11 July 1951, Loan 568b (DAO) Because Erigeron ochroleucus has not been included in floristic accounts of arc- tic areas, a description is provided here, drawn from 25 plants of the collections cited below. A detailed comparison can be made by comparing this description with that for “southern” E. ochroleucus in the Erigeron treatment for the devel- oping FNANM volumes (Nesom in prep.). Plants perennial from a thick, woody taproot, caudex multicipital, un- branched or rarely with short branches. Stems basally ascending, 2-15 cm, 54 BRIT.ORG/SIDA 21(1 loosely strigose to villous-strigose, eglandular or sparsely minutely glandular. Leaves mostly basal, linear to linear-oblanceolate, entire, 2-7 cm long, 1-1.8mm wide, cauline quickly reduced upwards, sparsely and loosely strigose adaxially, abaxially glabrous at least on distal 1/4-1/2 of blade and strigose proximally, eglandular. Heads 1; involucres 6-8 mm high, 12-16 mm wide; phyllaries in 3-4 series, densely hirsute-villous, hairs whitish or with red-purple crosswalls, densely and minutely glandular. Ray florets (18-)30-50, corollas 8-12 mm, lami- nae mostly white, less commonly bluish, 1-2(-2.5) mm wide, coiling. Disc co- rollas 3.5-4.5 mm long, not indurate or inflated. Cypselae 2-2.5 mm, 2-nerved, strigose-villosulous; pappus bristles 12-15, with a conspicuous outer series of thick setae, scales, or squamellae. The status of Erigeron pumilus in Yukon Porsild (1975) noted the occurrence in southwestern Yukon of Erigeron linearis (Hook.) Piper, as the single collection, H.M. and L.G. Raup 12918, had been iden- tified on the original label. Hultén (19607, 1968) referred to this collection as E. pumilus Nutt. as it has continued to be recognized (e.g, Cody 2000). With ca- veats as below, our identification of the three plants on this sheet is E. caespitosus Nutt. In E. pumilus (in contrast to E. caespitosus), the stems and involucres are hirsute to hispid-hirsute and glandular, the ray corollas are linear and ref lexing, and the disc corollas have distinctly indurate and inflated throats. CANADA. Yukon Territory: Alpine grassy-stony slope, rays lavender, mountain slope on east side of Big Arm, Kluane Lake, app. Lat. 61° 33'N, 138° 40'W, 20 Aug 1944, H.M. and L.G. Raup 12918 (ALA). Erigeron caespitosus has an extended geographic distribution, ranging from Arizona and New Mexico northward into Alaska; it is relatively common in southwestern Yukon (see map in Cody 2000). The Yukon plants in question have strigose cauline vestiture, which contrasts with the normally deflexed to spreading vestiture of E. caespitosus, but such variation exists sporadically in the species in other parts of its range, including Saskatchewan as well as vari- ous places in the western U.S.A, and the vestiture of Raup 12918 is more charac- teristic of E. caespitosus for 5-10 mm under the heads. The plants of 12918 also have linear to linear-oblanceolate basal leaves, narrower than normal for E. caespitosus, but narrow-leaved plants more clearly identified as E. caespitosus also occur in the Kluane Lake area (Murray, pers. observ.). The plants of 12918 are atypical, but we conclude that latent populational tendencies are sporadi- cally expressed in this species--at least a hypothesis of recent hybridization or genetic introgression is not supported, because hybridization with any other species in southwestern Yukon probably would have more conspicuously in- fluenced the morphological features of 12918. AND BOREAL SPECIES OF ERIGERON 55 ACKNOWLEDGMENTS We are grateful to Barney Lipscomb (BRIT) for help with locating bibliographic material, to NY staff for help on a recent visit, to Kanchi Gandhi for observa- tions and advice on nomenclatural problems, to Susan Aiken for her review comments and a color image of the holotype of Erigeron grandiflorus subsp. arcticus, to Robert Lipkin for review comments, and to ALA for a loan of speci- mens to Nesom at BRIT. We gratefully acknowledge the confidence gained by learning that Ron Hartman (RM) had earlier reached the same conclusion re- garding the identity of the Alaskan- Yukon populations of Frigeron ochroleucus. — REFERENCES Aiken, S.G., MJ. Datiwitz, L.L. Consaut, C.L. MCJANNeT, LJ. Gittespig, R.L. Botes, G.W. Arcus, J.M. GitvetT, PJ. Scort, R. Etven, M.C. LEBLANc, A.K. Brystinc, and H. Soistap. 2003 [1999 onwards]. Flora of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 29th April 2003. Accessed February 2004. Bown, B. 1972.Flora of the prairie provinces. Part Ill (continued). 113.Compositae.Phytologia 23:85-140. BOTSCHANTZEV, V.P. 1959. Erigeron. Flora U.R.S.S.25:191-288. Copy, W.J. 2000. Flora of the Yukon Territory (ed. 2). NRC.CNRC, NRC Research Press, Ottawa. Cronouist, A. 1947.A revision of the North American species of Erigeron, north of Mexico. Brittonia 6: 121-302. Dawe, J.G. and D.F. Murray. 1979. In IOPB chromososome number reports LXIll. Taxon 28: 265-268. Dawe, J.C.and D.F. Murray. 1981.Chromosome numbers of selected Alaskan vascular plants. Canad. J. Bot. 59: 1373-1381. Dorn, R.D. 1984. Vascular plants of Montana. Mountain West Publishing, Cheyenne, Wyoming. Doua.as, G.W., G.B. Stracey, D. Meipincer, and J. Pouar. 1998. Illustrated flora of British Colum- bia. Vol. 1, Gymnosperms and Dicotyledons (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks, Ministry of Forests. Victoria, British Columbia. Drummonb, T. 1830. Sketch of a journey to the Rocky Mountains and to the Columbia River in North America. Bot. Miscellany (WJ. Hooker) 1:178-219. Even, R., D.F. Murray, and B.A. Yurtsev. 2003. Checklist of the Panarctic Flora (PAF) Vascular Plants. Draft version, National Centre of Biosystematics, Natural History Museums and Botanic Garden. University of Oslo. Hutten, E. 1950. Flora of Alaska and Yukon. Vol. X.C.W.K. Gleerup, Lund.. Hutten, E. 1967. Comments on the flora of Alaska and Yukon. Ark. Bot., Bd. 7, 1:1-147. 56 BRIT.ORG/SIDA 21(1) Hutten, E. 1968. Flora of Alaska and neighboring territories. Stanford Univ. Press, Stanford, California. Lipkin, R.and D. Murray. 1997. Alaska rare plant field guide. U.S. Dept. Interior, Washington, D.C. (loose leaf, without pagination) McJANNet, C., G. Araus, S. Eptunp, and J. Cavouerte. 1993. Rare vascular plants in the Canadian Arctic. Syllogeus no. 72.Canadian Museum of Nature. Moss, E.H. 1959. Flora of Alberta. Univ. of Toronto Press, Toronto, Canada. Moss, E.H. 1983. Flora of Alberta (ed. 2, rev. by J.G. Packer). Univ. of Toronto Press, Toronto, Canada. Muir, J. 1883. Cruise of the Revenue-Steamer Corwin in Alaska and the N.W. Arctic Ocean in 1881.Notes and memoranda: medical and anthropological; botanical; ornithologi- cal. Govt. Print. Off, Washington. Muir, J. 1917.The Cruise of the Corwin. Journal of the arctic expedition of 1881 in search of DeLong and the Jeannette. Houghton Mifflin Co., N.Y. Murray, D.F. and R. Lipkin. 1987. Candidate threatened and endangered plants of Alaska with comments on other rare plants. Univ. Alaska Mus., Fairbanks. Nesom, G.L. 2004. Taxonomic reevaluations in North American Erigeron (Asteraceae: Astereae). Sida 00:000-000. Packer, J.G. and R. Witkus. 1982. ln IOPB chromososome number reports LXXV. Taxon 31: 363-364. Petrovsky, V.V. 1987. Erigeron. In: B.A. Yurtsevy, ed. Arctic Flora USSR. Vol. X. Nauka, Leningrad. (in Russian) Powunin, N.1959.Circumpolar arctic flora. Oxford Univ. Press, Oxford at the Clarendon Press. Porsitp, A.E. 1955. The vascular plants of the western Canadian Arctic Archipelago. Natl. Mus. Canad. Bull. 135. Porsito, A.E. 1974 [1975]. Materials for a flora of Central Yukon Territory. Natl. Mus. Canada, Publ. Botany 4:1—77. Porsito A.E. and W.J. Copy. 1980. Vascular plants of the Continental Northwest Territories, Canada. National Museum of Canada, Ottawa. Scoacan, HJ. 1979.The flora of Canada. Part 4—Dicotyledoneae (Loasaceae to Compositae). Natl. Mus. Nat. Sci. Publ. Bot. No. 7(4). SPONGBERG, S.A. 1971. A systematic and evolutionary study of North American arctic and alpine monocephalous species of Erigeron (Compositae). Ph.D. diss., Univ. of North Carolina, Chapel Hill. SPONGBERG, S.A. 1973. A new Alaskan species of Erigeron L. (Compositae). Rhodora 75: 116-119 Tavetev, N.N. (ed.). 2002. Flora of Russia: The European part and bordering regions. Vol. VII. A.A. Balkema, Rotterdam/Brookfield. Weser, W.A. 1987. Colorado flora: Western slope. Colorado Associated University Press, Boulder. Weeer, W.A. 1990. Colorado flora: Eastern slope. University Press of Colorado, Niwot. AND BOREAL SPECIES OF ERIGERON Weeer, W.A. and R.C. Wittman. 1992. Catalog of the Colorado flora: A biodiversity baseline Univ. Press of Colorado, Boulder Wicains, ILL. and J.H.THomas. 1962. A flora of the Alaskan arctic slope. Univ. of Toronto Press, Toronto, Canada. Yurtsey, B.A., T.G. Potozova, and N.A. Sexretareva. 1989. Additions and corrections to the list of vascular plants of Wrangel Island. Communication 2. Byull. Moskovsk. Obshch. Isp. Prir,, Otd. Biol. 94:116-126. (in Russian) BRIT.ORG/SIDA 21(1) BOOK NOTICE Ben-Erik VAN Wyk and MicHAcL WINK. 2004. Medicinal Plants of the World. (SBN 0-88192-602-7, hbk.) Timber Press Inc. 1335.W. Second Ave, oe 450, Portland, OR 97204-3527, US.A. (Orders: wwwtimberpress.com berpress.com 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $39.95, 480 pp., 793 color photos, 3 tables, 137 ae 6 sal x 9 a ll id he | known Medicinal Plants ofthe Worldis and most important medicinal slants, including those of special eonimercal or historical interest. This ee isa Be ro es oon their botany, main traditional uses, active ingre- dients, ] iden efficacy (if known) in a compact, colorful, reference text. Medicinal Pl lantsdoesa wonderful job of concept ing the more than 320 plants in the book. The regulatory status of each plant is also given—whether it is listed in one or more harmacopoeias, 5 io 5 cals the German Commission E monographs, the new ESCOP monographs of the European Community and the World Health Organization's series of monographs (all species treated in the ast-mentioned three works will be found in this book). In the back of the book, the authors do well to include “A Quick Guide to ee ized ecu Plants” and a “Glossary of Chemical, Medical °T geared toward the specialist and not the “av as itles. o. and Pharmaceutical Terms.” The | gardener.” The guide is ee alshabene i by genus species, and common names in os ome previous knowledge of botanical names, the user will be referencing the ae fre- quently. Nine hundred plants are covered in the book, but 320 have comprehensive detail including: plant description, geographic origin, therapeutic category, historical and modern uses, active ingre- dients and pharmacological effects —Asha McE ish, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 21(1): 58. 2004 NOTES ON TYPIFICATION IN PLUCHEA (ASTERACEAE: PLUCHEAE) Guy Nesom Botanical Research Institute of Texas 9 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Pong nowlenclature and Hypiicar ation are detailed for taxa of the Pluchea oe Pluchea I gnated for Pluchea glabrata DC, C ichx. Gynema viscide Ral, Pluchea petiolata Cass., and Conyza camphorata Pursii Pluchea iConyas) marilandica (Michx.) Cass., which is the type of the genus, is based on a Dillenius illustration that shows features anomalous for Pluchea. The identity of several one ponent taxa also remains ambiguous, but none of them appears to have a bearing on accepted nomenclature. RESUMEN Se detalla la nomenclatura formal y la tipificacion de taxa del grupo Pluchea odorata—Pluchea camphorata. Se designan lectotipos para Pluchea glabrata DC., Conyza marilandica Michx.,Gynema viscida Raf., Pluchea petiolata Cass., y Conyza camphorata Pursh. Pluchea (Conyza) marilandica (Michx.) Cass., que es el tipo del género, esta basada en una ilustracion de Dillenius que muestra caracteristicas andmalas para Pluchea. La identidad de otros varios taxa también queda ambigua, 7 7 ‘ y | Jat ] + weed 1] if 5 Parece (Cer p In connection with production of a taxonomic treatment of Pluchea for the Flora of North America (FNA) volumes of Asteraceae, an attempt is made here to clarify typifications for various names associated with two, relatively wide- spread species of Pluchea with petiolate, non-clasping leaves. Pluchea odorata (L.) Cass. The typical form of Pluchea odorata occurs from northern South America through the West Indies (the type is from Jamaica) and Mexico and is widely distributed through the southern United States, from California and Nevada to Texas and Oklahoma and eastward to North Carolina and Virginia. A large- headed variety of this species, primarily occurring in the northeastern United States, is P odorata var. succulenta (Fern.) Cronq. (Cronquist 1980; Khan & Jarvis 1989) Pluchea odorata (L.) Cass. [var. odorata]. Pluchea odorata (L.) Cass., Dict. Sci. Nat. 42:3. 1826. Placus odoratus (L.) M. Gomez, Anales Soc. Esp. Hist. Nat. 19:273. 1890. Pluchea odorata (L.) DC. var. normalis Kuntze, Revis. Gen. Pl. 1:357. 1891 (nom. in- valid; includes the type of the autonym). Conyza odorata L., Syst. Nat., ed. ee ee 1759. Linnaeus cited “Sloan. jam. t. 152. f. 1. Plum. ic. 97.” LECTOTYPE (Urb. e Godfrey 1952:241-242, 245): JAMAICA. Sloane, Voy. Jamaica 1: plate 152()). eo SIDA 21(1): 59 — 64. 2004 BRIT.ORG/SIDA 21 = 1) The plant illustrated is in the Sloane Herbarium—no data regarding collector, lo- cality, or cage eee Gillis 1977)! internet image !). Although Codirey’ s acceptance of an earlier oie of type by Urban formalized the lectotypification of P odorata, Godfrey misinterpreted the Sloane illustration as referring to the woody species now generally identified as Pluchea carolinensis Jacq.) D. Don. Application of this latter name was clarified by Khan and Jarvis (1989). The herbaceous species treated by Godfrey as P. purpurascens is correctly identified as P odorata (see Gillis 1977). — hea Ss (Sw.) DC, Prodr. 5:452. 1836. Placus purpurascens (Sw.) M. Gomez, Anales Hist. Nat. 19:273. 1890. Conyza purpurascens Sw., Prodr. 112. 1788. Type: JAMAICA: Swartz cel ‘Conyza I. Brown. jam. 318.” and “Conyza major odorata—Sloan. h. I. 258. t. 152. a ” Browne's citation (1756) was “Conyza major odorata, Xe. Slo. Cat. 124. & H.t. 152. f. reference to the same Sloane illustration, which is the same as the lectotype of omnes ove). LECTOTYPE (Howard 1989:589): JAMAICA. Sloane, Voy. Jamaica L: plate 152(1). 1707. = ene illustrated is in the Sloane Herbarium, as cited above for Pluchea odorata. Pluchea purpurascens (Sw.) DC. var. glabrata (DC.) Griseb., Cat. Pl. Cub. 150. 1866. Placus P purpurascens (Sw.) M. Gomez var. glabratus (DC.) M. Gomez, Anales Soc. Esp. Hist. Nat. 19:273. 1890. Pluchea glabrata DC., Prodr. 5:452. 1836. Type: GUADEL OUPE: de Candolle cited the following: “in Guadalupa legit olim cl. Badier et recentius Perrottet ad Pointe-a-Pitre.” Of speci- verbarium, one is labeled simply mens identified as Pluchea glabrata in the de Candolle “Guadeloupe” and may be the Badier collection: oe eeu more specific data—*Pte. a Pitre, 17 Juin 1824, Perrottet 1825.” This plant also lal ,and (Urban 1902) indicates that Perrottet was there at least for a period during 1824. The Pe rrottet collection is designated here as the LEC as DC fiche 905). luchea petiolata Cass., Dict. Sci. Nat. 42:2. 1826. TyPE: Details of the protologue are too abbrevi- ated to Se the ee “ses calathides sont petites, subglobuleuses, et disposées en corymbe terminal; ... ses squames sont ovales, aigués, presque membraneuses, parsemées de glandes...” The description of — heads with acute, membranous, sparsely glandular phyl- ‘ata, the heads ina ea corymb and laries suggest that it might be orata. a ae that “Nous avons observé cette ovate phyllaries are more like Pluc hea oc plante dans Vherbier de M. de Jussieu, ou elle étoit innomimeée,’ citing no further detail. One of two _ in the Jussieu herbarium might have been the one observed by Cassini: (1) Cat. No. 8646 (P-Jussieu, fiche 634!)—identified as “Conyza marylandica Michx.,” with no collection ane ciae Phe plant has short-petiolate leaves and a terminal capitulescence and is Pluchea ela (2) Cat. No. 8662 (P-Jussicu, fiche 635!)—identified only as “Conyza vulgari accedens [1. P purpurasc.,” ee co pais on es aes CSt. Domingo”); this slant also has short- petiolate leaves anda I and is a representative ol Pluchea odorata var. odorata. This specimen is here d aannied as the LECTOTYPE of Pluchea petiolata luchea camphorata (L.) DC. var. angustifolia Torr. & A. Gray, Fl. N. Amer. 2:261. ne Coa ita id Nutt., J. Acad. Nat. Sci. Philadelphia 7:109. suede Roxburgh 1832). Type: USA. “In South Carolina, from whence it was received by Mr. Durand” (specimen not located). Torrey - ae cited the Nuttall publication in full and apparently intended to recognize Nuttall’s Conyza angustifolia at reduced rank, not realizing that the basionym published by Nuttal was a later homonym. Following Article 58 of the [CBN (Saint Louis Code 2000), the name at varietal ee can be attr eee to Torrey and Gray asa nomen novum, Pluchea odorata var. succulenta (Fern.) Cronq, Pluchea purpurascens (Sw.) DC. var. aan ee Fern., Rhedoia: - 227. 1942. Pluchea odorata var. succulenta (Fern.) Cronq,., Fl. Southeast. U.S. 1:175. 1980. Type: UNITED STATES. MASSACHUSETTS: Back Bay, Boston, ditch in old marsh, 17 Sep 1910, EF Williams s.n. (HOLOTYPE: GH). NESOM, NOTES ON TYPIFICATION IN PLUCHEA 61 Pluch ilandica (Michx.) Cass., Dict. Sci. Nat. 42:2. 1826. ue marilandica Michx., Fl. Bor. Ane 2 126. 1803. TYPE: Michaux cited “DILL. Elth. t. 88. f. 104. nondum perfecte florida” and gave as possible Pes ee “Baccharis foetida? Walt.” and oe camphoratum? Linn noted that the habita “a Pensylvania ae Cana the oe PE, a designated Baie) must be the piesa ce the only elthamiensis |: t.88. f. 104.1732. This paatie identified in the Hortusas' ‘Conyza Marilandica, flore purpureo” and is shown with broadly lanceolate, sessile, serrate leaves and minutely pubescent stems (as illustrated), but the capitulescence is relatively elongate, broken into several subclusters borne on leafy branches and illustrated details show the heads as cylin- dric-campanulate with phyllaries in ca. 4-5 series and the bisexual florets as strongly gob- let-shaped, features not characteristic of Pluchea. The specimen in the Sherardian Herbari ium (OXF) upon which the drawing was based was identified as “Conyza marylandica” by Klinsmann (fide Druce and Vines 19 The voucher specimen at OXF notated “88 104 105,” bears two branches and was annotated by James Reveal in 1982 as Pluchea purpurascens (= P. odorata). A digital image made available by Stephen Harris, Curator of Oxford Univer- Herbaria, confirms that identity. An early annotation identifies it as “Pluchea marylandica.” According to Dr. Harris (pers. comm), “This is the speci- men that Druce placed into the Hortus sequence when he worked on the mate- rial with Vines in the early 1900s. However, the specimen appears to have been incorporated from the Sherardian Herbarium and bears a different polynomial from that ewer in the Hortus.” Clokie (1964, p. 89) observed that “very few of the that [Druce] did take out lof the Sherard Herbarium] can be iden- tified as originals of the plants in the Hortus Elthamiensis” and she noted spe- cifically that the voucher for t. 88, fig. 104 is ‘not the original specimen.” The original label on the OXF sheet identifies the collection first as “Baccharis mariana flora pulchra rubenta Acta Philos. No. 246. pag. 401. pl. 22 1690” (by Sherard), then as “Conyza mariana flora pulchra rubenta Mem. Ac. R. Se. 1719. p. 299 ns.” (by Dillenius). Thus, the identity of this specimen is clear, but its status as a voucher for the subject of the Dillenius illustration is not. Because Cassini typified his new genus Pluchea by Pluchea marilandica, typification of the genus is ambiguous to the same extent. In Cassini's establishment of the genus and transfer of Conyza marilandica to Pluchea, he noted “Cette plante habite !Amerique septentrionale” and de- scribed “l’enchantillon que nous avons observé dans lherbier de Michaux... .” Two plants in the Michaux herbarium (PI, P-Michaux fiche 100!) are identified as Conyza‘marylandica. One is labeled simply “Conyzal,] Maryland,” while the second plant is labeled “Conyza marylandical,] Virgin. et Carol."—both speci- mens are representatives of Pluchea camphorata, although the morphological details included by Cassini (Dict. Sci. Nat. 42:2) do not clearly indicate that he was describing P. camphorata rather than P. odorata. ms Pluchea camphorata (L.) DC. Based on a photograph of the type of Erigeron camphoratus L. received at Har- 62 BRIT.ORG/SIDA 21(1) vard from the British Museum, Fernald (1939) made this assessment: “That it is a fragment from the summit of a plant of P petiolata Cass. will be evident to those who know that plant” (p. 461, photo published on p. 459). Fernald’s judge- ment has been accepted by those who have dealt with this common species (e.g. Godfrey 1952, Cronquist 1980, Nesom 1989, and various other floristicians) and is confirmed here. A color image of the Clayton specimen is currently internet-posted as part of the John Clayton Herbarium (The Natural History Museum 2000) a camphorata (L.) DC, Prodr. 5:451. 1836. Erigeroncamphoratus L., Sp. Pl. 864. 1753. Linnaeus cited “Hort. ups. 259” and “Gron. virg. 97. Habitat in Virginia.” Gronovius (Flora Virginica, ed. 1, 1739, p. 96) provided a description and referred to “Clayt. n. 165.” The description in Linnaeus’s own Hortus Upsaliensis of 1748 was nearly identical to the later one of 1753 and also referred to “Gron. virg. 96.” As observed by Fernald (1939, p. 460), “Linnaeus composed his diagnosis by adding a portion of Clayton’s own description [on the specimen itself] of his plant to that of Gronovius.” LECTOTYPE (Fernald 1939:459-461): UNITED STATES. VIRGINIA: with- out other data, Clayton 165 (BM!, photo-GH, photo- Fernald (1939)! internet as ; accessed December 2000. Ursan, |. 1902.Notae biographicae peregrinatorum Indiae occidentalis botanicorum.Symb Antill. 3:14-146. A REVIEW OF CRATAEGUS SERIES ROTUNDIFOLIAE (ROSACEAE) IN WESTERN CANADA J.B.Phipps RJ.O’Kennon Dept. of Biology Botanical Research Institute of Texas The University of Western Ontario 09 Pecan St. London, Ontario, N6A 5B7, CANADA Fort Worth, a ae 4060, U.S.A. jphipps@uwo.ca okennon@brit.org ABSTRACT Par ate ay eek one | ee This paper discusses the circumscription of Crataegus ser. R materials (full descriptions, line illustrations and specimen citations her not previously avail- able) for the taxa of the series occurring in western Canada, defined for the purposes of this paper as British Columbia and Alberta. The variable, widespread and common species C. chrysocarpa is re- viewed for this area and a local, rather distinct new variety, var. vernonensis, is recognized. In addi- tion, one species new to science, C. sheila-phippsiae is described Key Worps: Crataegus, Rosaceae, ser. Rotundifoliae, western Canada, C. sheila-phippsiae sp. nov, C chrysocarpa var. vernonensis var. Nov. RESUMEN Cpdi rae es | Ee | “Cs d Cratdegus ser Rotundifoliae y (descripciones complies. ilustraciones y citas de ued que no estaban disponibles previamente) para los taxa de 1 oeste de Canada, definido a los efectos de este articulo como ae Bente y Alberta. La especie variable, muy difundida y comin C. GNI sOUarpEe es rev pease para esta area y se reconoce una nueva variedad local bastante diferente, var. vernonensis ibe una nueva especie para la ciencia, C. sheila-phippsiae. Since the first author's review (Phipps 1997) of red-fruited Crataegus in western North America was prepared, intensive collecting, particularly in the British Columbia Okanagan, has revealed a number of new species, especially in ser. Purpureofructi J.B. Phipps & O’Kennon and ser. Douglasianae (Rehder ex CK. Schneider) Rehder. Here, we deal with ser. Rotundifoliae (Eggl.) Rehder (1940) in western Canada, a very wide-ranging series of hawthorns. They occur from the Pacific Northwest of the USA and southern British Columbia in a broad band through southern Canada and the northern United States to Newfound- land and southern New England extending locally southward, in the Rocky Mountains to Colorado and in the Appalachians to northern North Carolina. The series is a re-ranking of Eggleston’s section Rotundifoliae (1908b) which in turn isa renaming of Sargent’s (L905) ‘natural group’ Coccineae which Eggleston (1908b) considered to be improperly typified. If one follows Eggleston (op. cit.) in which C. rotundifolia Moench is divided into two varieties, var. rotundifolia (now C. dodgei Ashe) and var. chrysocarpa (Ashe) Eggl. (now C. SIDA 21(1): 65-77, 2004 66 BRIT.ORG/SIDA 21(1) chrysocarpa Ashe), one might typify series Rotundifoliae with C. dodgei, which would seem plausible as C. dodgei possesses a round to broadly elliptic leaf and C. chrysocarpa normally possesses a + rhombic leaf. Unfortunately, Moench’s illustration (1785) shows a hawthorn with an elliptic leaf, pointed at both ends, nothing like the leaf-shape of C. dodgei and not much like that of C.chrysocarpa and, regrettably, completely at variance with the text of the protologue. The poor quality of Moench’s protologue for C. rotundifolia was further compounded by the fact that his name was invalid, C. rotundifolia Lam. (1783) being an ear- lier name for a different species. In this paper, we avoid the problems associated with typifying C. rotundifolia, by considering both C. chrysocarpa and C. dodgei (not in the area of this paper) to be members of series Rotundifoliae. Nevertheless, there is a serious argument for separating C. dodgei and immediate relatives from C. chrysocarpa and immediate relatives to create two series at which point the typi- fication problem would need to be addressed. In western Canada, all members of series Rotundifoliae fall into the C. chrysocarpa group and there is consider- able variation within it, some only recently recognized. Inasmuch as the C. dodgei group does not occur in western Canada and is perhaps best treated asa different series as suggested above, the following char- acteristics of ser. Rotundifoliae may be understood as only applying to the west- ern forms of the series, ie. the non-dodgei group of species. Series Rotundifoliae in this restricted sense are among the smaller hawthorns, typically multi- stemmed bushes 2-4 m tall and very thorny. The leaves are + rhombic to + ovate in shape, 2-4 lobed per side with LII (leaf incision index; IFI in latin) 5-20 %, rather glandular on the margins and petiole and typically very hairy adaxially. Anthesis is early, inflorescences 5-18 flowered, with usually pubescent branches bearing caducous, linear-oblong, gland-margined bracteoles. Flowers are mid- sized, 13-18 mm diam; hypanthium usually hairy; calyx-lobes quite strongly glandular but weakly serrate; stamens 10 or rarely 20; anthers ivory to cream or less commonly pink; styles 3-4. Fruit 8-12 mm diam, usually torbicular, hairy or not; calyx-lobes spreading else broken off; nutlets 3-4, sides smooth. At least superficially, the most similar series in western Canada are Macracanthae (Loud.) Rehder with eglandular petioles and laterally eroded nutlets and Purpureofructi, which are much larger plants, have stout, shorter thorns and plum-red to purple-black ripe fruit with sides of nutlets + eroded. Eggleston (1909) placed his C. williamsii in ser. Rotundifoliae but this species has laterally eroded nutlets, generally slightly larger leaves and is now placed in ser. Purpureofructi. The region explicitly covered by this paper is British Columbia and Alberta although the descriptive text should also cover C. chrysocarpa in Saskatchewan and Manitoba. PHIPPS AND O’KENNON, REVIEW OF CRATAEGUS SERIES ROTUNDIFOLIAE 67 TAXONOMIC TREATMENT We recognize here two species of ser. Rotundifoliae: C. chrysocarpaand C. sheila- phippsiae, the former with three varieties. The last-named species is new to sci- ence and brings to five the number of Crataegus species endemic to the remark- able hawthorn-rich region of the northern Okanagan. The detailed locations cited may be found, more or less precisely, by reference to our sketch-map (Fig. 1). KEY TO SPECIES 1. Stamens 20; inflorescences glabrous 1.C. sheila-phippsiae, sp. nov. 1. Stamens 10; inflorescences normally + hairy 2.C. chrysocarpa 1. Crataegus sheila-phippsiae J.B. Phipps @ O’Kennon, sp. nov. (Fig. 2) Type: CANADA: BRITISH COLUMBIA. Northern Okanagan, Spallumcheen Mun., McLeery Rd, 50° 20'59" N, 110° 08' 40" W, roadside, edge of thicket, 1314 ft asl, bush 4.5m tall, 18 pale pink anthers, 16 May 2002, J.B Phipps 8334 (HOLOTYPE: UWO; ISOTYPES: ALTA, CAN, DAO, TRT, UBC, UVIC, V). pinae 2.5-4 cm longae; + tenues vel validae, rectae vel leviter Frutices, usiter arborescentes, 3-5 malti: recurvatae, annotinae nitenter atrobrunneae, veteriores atrocinereae; ramuli extendentes glabri, virides; annotini valde nitento-brunnei, veteriores atrocinerei. Folia decidua; petioli 10-20 mm longi, graciles, pubescentes, glandulis secundum m paucis vel pluribus; laminae 4-5 cm longae, + rhombeae in forma g general, apex acuminata, ee cuneata; a 4 gente lobi per latus, max. IFI ca. bescentes, abaxiales 20%; oes pu pilosae solo in venis ruined et in axillis; feaane craspedodroma, 4-5 nervatae per latus. eee 8-18-florae; ramuli glabri, bracteolis caducis, linearibus, membranaceis, glandulo- marginatis. Flores 14-19 mm diam.; Geena glabrum extrinsecus; lobi calycis triangulares, glandulo-marginati sed vix serrati; petala alba, + circularia; stamina 18-20, antheris pallide roseis (raro moderate roseis - eee 8407); styli 4-5, exorientes a caespite parvo pilorum. Fruc- tus 10-12 mm diam., subglobosi, clare carmineo-scarlatini in Augusto exeunte, in maturitate fortiter rubro-purparei; lobi calycis patentes; pyrenae 4-5, dorsaliter sulcatae, lateribus planis. Shrub, usually somewhat tree-like, 3-5 m tall; thorns 2.5-4 cm long, moderately slender toa little stout, straight or slightly recurved, dark shiny blackish brown at 1 yr, older deep gray; extending twigs glabrous, green, at | yr rich shiny brown, older gray. Leaves deciduous; petioles 10-20 mm long, slender, with few to sev- eral glands along the sulcus, pubescent; blades 4-5 cm long, + rhombic in gen- eral shape, tip acuminate, base cuneate, 3-4 sharp lobes per side; max LIl ca. 20%; margins with very small sharp teeth; scabrous-pubescent adaxially, hairy in the axils and parts of the main veins abaxially; venation craspedodromous, ca. 5 veins/side. Inflorescences 8-18 flowered; branches glabrous bearing early caducous, linear, membranous, gland-margined bracteoles. Flowers 14-19 mm diam.; hypanthium externally glabrous; calyx-lobes triangular, gland-mar- gined, barely serrate; petals white, + circular; stamens 18-20, normally pale pink but occasionally medium-pink (Jackson for Phipps 8407) or puce (Phipps & O’Kennon 6917); styles 4-5, arising from a small tuft of hairs. Fruit 10-12 mm diam., subglobose, brilliant crimson-scarlet in late August deepening toa dark 68 BRIT.ORG/SIDA 21(1) burgundy at full ripeness, sey preading; nutlets 4-5, dorsally grooved, sides plane. Crataegus sheila-phippsiae is a relatively scarce species restricted to the northern Okanagan of British Columbia but in some localities rather common as, for instance, just north of the Indian Cemetery on hwy 97A about 2 km south of Enderby. It grows in relatively open and apparently rather dry sites and as such appears to be quite xeromorphic like other members of the same series. Crataegus sheila-phippsiae is a typical rhombic-leaved member of ser. Rotundifoliae. It is easily differentiated from C. chrysocarpa var. chrysocarpa, to which it is most similar, by its + glabrous nature, 20 stamens, usually pale pink anthers and greater single-trunk dominance. In the past difficulties have arisen differentiating C. sheila-phippsiae from C. shuswapensis (ser. Douglasianae) in flower in those rare cases where C. shuswapensis has 15-18 stamens but we be- lieve that they are now most easily resolved at the flowering stage by the better development of abaxial tufts of hair in the vein axils of C. sheila-phippsiae whereas C. shuswapensis is usually glabrous there. When in fruit, however, C. shuswapensis is quite different from C. sheila-phippsiae. The authors now sus- pect that the attribution of 18 stamens to C. shuswapensisin Phipps & O’Kennon (2002) may refer toa plant of C. sheila-phippsiae with deeply colored anthers. Of course, the critical eye will also notice differences in leaf color and slight differences in shape, together with subtle habit differences, all difficult to put into words. Also anther color in C. shuswapensis is much more variable than in C. sheila-phippsiae, ranging from pink to puce. Persons with the resources might investigate the possibility that C. sheila- phippsiae may be a diploid, sexual counterpart of C. chrysocarpa, like the C. suksdorfii/ douglasii pair so elegantly demonstrated by Dickinson et al. (1996). The species epithet recognizes the important contribution to the study of hawthorns by Sheila Phipps, wife of the first author, which include the typing over the years of hundreds of pages of manuscript, help with proofing of printed text, tending to young Crataegus seedlings and assisting at the scene the year this species was recognized in the field with certainty for the first time. Additional specimens examined: CANADA: British Columbia. Northern Okanagan. Mabel Lake Rd., 1.0 km E of centre of Shuswap R. bridge at Enderby, bush 3.5 m tall, fruit burgundy, 07 Sep 2001, J.B. Phipps 8279 & R,J.O’Kennon (BRIT, UWO), ca. 1.2 mi E of Enderby on road to Mabel Lake, bush 3 m tall, fruit burgundy-red, 20 Aug 1994, J.B. Phipps 7015a & RJ.O’Kennon (UWO), ca. Lmi S of Enderby, gravel driveway to NNE off hwy 97A at Indian Cemetery, bush 2.5 m tall, fruit deep blood red, 20 Aug 1994, J.B. Phipps & RJ. O’Kennon 7003 (BRIT, UWO); same loc., bush 2.5 m tall, pink anthers, 16 May 2002, J.B. Phipps 8335 (BRIT, UWO); same loc., bush 2.5 m tall, 20 pale pink anthers, 16 May 2002, J.B. Phipps 8336 (BRIT, UWO). Same loc., bush 4 m tall, 20 pale pink anthers, 16 May 2002, J.B. Phipps 8338 (BRIT, UWO); same loc., bush 2.5 tall, 20 pale pink anthers, 16 May 2002, J.B. Phipps 8339 (BRIT, UWO); same loc., bush 3 m tall, 20 pale pink anthers, 16 May 2002, J.B. Phipps 8340 (BRIT, UWO). same loc., bush 3 m tall, 20 pale pink anthers, 16 May 2002, J.B. Phipps 8342 (BRIT, UWO). Same loc., bush 3 m tall, anthers 18, light pink, 22 May 2002, R. Jackson for J.B. Phipps 8380 (BRIT, UWO); same PHIPPS AND O’KENNON, REVIEW OF CRATAEGUS SERIES ROTUNDIFOLIAE 69 Ps t [hwy 978 v 1 | Enderby". Ka «Deep ' * Creek ae ~ Fe tee fi eee rail Hi ty Met Bory Ra. n 7 FortuneCreek o “KH aa) . Mountain View Rd. rmgtrong WN -1] o = uae -~7 “Schubert Rd: (eas * Schubert A \ a Otter L. Cr x. Grandview 2 Re ~\Flats Rd.» — 4 eee 3 ‘ yy, Ger \ ge Silver Star ntain ah Lake ‘| a pleasant Valley Rd. +) Fic. 1. Sketch-map of the Northern Okanag I I g loc., bush 2 m tall, 20 light pink anthers, 22 May 2002, R. Jackson for J.B. Phipps 8381 (BRIT, UWO): same loc., bush 3 m tall, 20 light pink anthers, 22 May 2002, R. Jackson for J.B. Phipps 8383 (BRIT, WO): same loc., large bush, 20 light pink anthers, 22 May 2002, R. Jackson for J.B. ear WO); same loc. fruit deep red, 25 Aug 2002, R. Jackson for J.B. Phipps 8458 (BRIT, UWO); same loc.. fruit very dark red to deep purple, 22 Sep 2002, R. Jackson for J.B. Phipps 8475 (BRIT, UWO); 70 BRIT.ORG/SIDA 21(1) Spallumcheen Mun., Schubert Rd., bush 2 m tall, fruit reddish plum, 19 Aug 1995, J.B. Phipps 7168 BRIT, UWO); same loc., bush, 3.5 m tall, pale pink anthers, 18 May 2002, J.B. Phipps 8362 (BRIT, UWO), Spallumcheen Mun., McLeery Rd., bush 3 m tall, fruit crimson-scarlet, 23 Aug 2000, J.B. Phipps with S.R. Phipps 8174 (BRIT, UWO); same loc., large bush, anthers 20, light pink, 20 May 2002, R. Jackson for J.B. Phipps 8376 (BRIT, UWO), Spallumcheen Mun,, Stepney Cross Rd., bush 4.5 m tall, fruit red- ca ay 06 Sep 2001, J.B. Phipps with RJ. O’Kennon 8270 (BRIT, UWO); Spallumcheen Mun, ak ust N of Otter Lake Cross Rd., fruit very dark, 22 Sep 2002, R. Jackson for J.B. Phipps pera oe. same loc., 25 Aug 2002, R. Jackson for J.B. Phipps 8461 (BRIT, UWO); Spa allumcheen Mun., Otter Lake Cross Rd. between Hales Rd. and ee Rd., bush 2.5 m tall, fruit burgundy or darker, 20's stamens, 06 2001, J.B. Phipps & RJ.O’Kennon 8269 (BRIT, UWO); same loc., bush, 3.5m tall, pale pink anthers, 18 May 2002, J.B. Phipps 8362 (BRIT, UWO),; same loc., bush 1.5m tall, 10 May 2002, J.B. sence (BRIT. UWO); same loc., bush, 2 m tall, 20 stamens, pale pink anthers, 18 May 2002, J.B. Phipps 8361 (BRIT, UWO); same loc.,, fruit deep red, 25 Aug 2002, R. Jackson for J.B. Phipps 8462 (BRIT, UWO); some additional duplicates will be distributed = later. 2. Crataegus chrysocarpa Ashe, 1900 (FIREBERRY HAWTHORN). Crataegus rotundifolia Moench, 1785, sens. auctt., non Lam., 1783. Crataegus columbiana Howell, 1898, sens. auctt. Amer. Crataegus piperi Britton, 1901. Crataegus chrysocarpa var. piperi (Britton) Kruschke (1965). Crataegus columbiana var. chrysocarpa (Ashe) Dorn, 1988. Crataegus rotundifolia var. chrysocarpa (Ashe) Eggl., 1908a. Crataegus columbiana var. piperi (Britt.) Egg],]908a. TyPE: U.S.A. COLORADO. LARIMER Co. foothills of Cache le Poudre Mountains, 6000 ft, 20 Aug 1899, W.W. Ashe s.n. (NY). Bushes 2-3.5 m tall, sometimes colonial; extending twigs usually appressed- pubescent; l-year old branches usually dull yellowish-greenish brown or gray- brown to light tan in the east, rich tan but sometimes (as in MN and QU) shiny deep reddish- brown; thorns variable, 3-6 cm long, + slender to * stout, straight to slightly recurved, shiny, dark mahogany or blackish at 1 yr. Leaves decidu- ous; petioles 1-2.5 cm long, + pubescent, usually narrowly winged distally, + glandular at least initially; laminas 2-6 cm long, + rhombic to broad-elliptic, very occasionally broad-ovate in general shape, 2-4 lobed, the lobes sharp to blunt, LII 5-20%; margins with numerous small teeth, these caducous gland- tipped; venation craspedodromous, about 5-6(-7) main lateral veins per side; usually appressed-short pubescent above and pubescent mainly on the veins beneath, variably glabrate later, autumn color yellow, or purplish in very dry spots. Inflorescence 5-10 flowered; branches densely to sparsely villous, bear- ing caducous, linear, membranous, gland-margined bracteoles; anthesis early. Flowers ca. 15-20 mm diam.; hypanthium villous; calyx-lobes ca. 4 mm long, triangular, margins glandular-serrate, pubescent abaxially; petals + circular, white; stamens 10(-20), anthers only cream or ivory in the area covered by this paper, styles 3-4(-5). Fruit 8-10 mm diam., usually + spherical, usually scarlet to deep red or darkened to burgundy if dried and shriveled, + pubescent, calyx remnants present. Nutlets 3-4(-5), grooved dorsally, their faces plane. Crataegus chrysocarpa is the second most wide-ranging of North Ameri- can Crataegus species reaching from the landward side of the coastal ranges in —_— PHIPPS AND O’KENNON, REVIEW OF CRATAEGUS SERIES ROTUNDIFOLIAE 71 Fic. 2. Line drawing of C. sheila-phinnsiae from J.B. Phinns 8232 (UWQ) and] BR Phinn<@R OK, 810 (UWO), flower- g PIPE [ ing and flower details; R. Jackson for J.B. Phipps 8467 (UWO), fruiting and fruit detail. Scale bars = 1 cm. S. Laurie- Bourque del. 72 BRIT.ORG/SIDA 21(1) British Columbia, Washington and Oregon to the Atlantic. Its southern limit is Colorado, further north than the New Mexicoand Arizona southern limit for C. macracantha which otherwise has a broadly similar but rather more south- erly range. Crataegus chrysocarpa is abundant in the grasslands of the Cana- dian Prairie Provinces where it nearly always occurs in sunny areas in the bot- toms of draws, seepage zone edges on slopes or along shallow ditches. It may also be found occasionally under aspen, generally at the edge of a stand or where the stand is thin and is found in similar habitats in valleys of the inter-moun- tain areas. Crataegus chrysocarpa is the most northerly and cold-adapted spe- cies of Crataegus in North America nearly reaching Hudson Bay in Ontario. Crataegus chrysocarpa is a multi-stemmed bush, small for a hawthorn, usually 2-4 m high and, not surprisingly for so abundant a species, somewhat variable, although through most of its western range rather uniform. However a fairly well-marked regional variant, var. piperi, which is more thoroughly dis- cussed in Phipps (1997) and which might be worth raising to subspecific rank, occurs west of the Rocky Mountains. Crataegus piperi has also been synonymized with C. columbiana Howell but the first author showed that C. columbiana was based on two species and lectotypified it on the C. douglasii element (Phipps, 1995). The characteristic ripe fruit color of Crataegus chrysocarpa var. chrysocarpa in western Canada is bright red, in spite of the species epithet. However, about the third week of Au- gust it is a fairly bright orange, which ripens through orange-red to the final color. The fruit of a small percentage of specimens of C. chrysocarpa var. chrysocarpa observed in the Cypress Hills, Saskatchewan in late August 2001, however, were still golden-orange when most around were bright red. Also, in the same area and time, a lanate form of the type variety was found to have greenish-yellow fruit when congeners were bright red but its fully ripe color seems to occur later and is deep red. A few specimens from Saskatchewan, oth- erwise indistinguishable from the typical variety, have 20 stamens. See Phipps (1997) for citation of specimens, line illustration and distribution map. We also take the opportunity here to describe asa variety new variety of C. chrysocarpa, the Okanagan endemic taxon, var. vernonensis. The name Crataegus rotundifolia Moench has quite frequently been used for C. chrysocarpa Ashe, for instance by Packer (1983), though starting with Eggleston (1908a) as noted in the discussion of the series. However, as noted above, Eggleston's sense of C. rotundifolia var. rotundifolia was obviously C. dodgei, a different species. pa KEY TO VARIETIES OF C. CHRYSOCARPA IN WESTERN CANADA 1. Anthers ivory to cream; ripe fruit usually bright red. 2. Leaf-blades generally smaller (2-4 cm long) usually sharply but less deeply lobed (LIL 5—15%); leaves + glabrous abaxially 2a. var.chrysocarpa PHIPPS AND O’KENNON, REVIEW OF CRATAEGUS SERIES ROTUNDIFOLIAE 73 2. Leaf-blades larger (4-6 cm long), sharply and quite deeply lobed (LII < 20%), densely pubescent abaxially 2b. var. piperi . Anthers pale pink; ripe fruit usually deep claret-colored 2c. var. vernonensis, var. Nov. 2a. Crataegus chrysocarpa var. chrysocarpa. TYPE: as species. Characteristics as key. Variety chrysocarpa occurs practically throughout the range of the species and in western Canada in the valleys of southern inter- mountain British Columbia and the grassland biomes of Alberta, Saskatchewan and Manitoba where it is nearly the only hawthorn. The indumentum of the inflorescence is generally moderately pubescent but rare, densely lanate or + glabrous forms also occur. A broad-leaved form originally described as C. sheridana A. Nels. from Colorado might be varietally distinct and should be looked out for in the area of this paper. Its nearest proven occurrence is in North Dakota. Overall the variation of the typical variety in western Canada is con- siderable and warrants further study. 2b. Crataegus chrysocarpa var. piperi (Britton) Kr a 1965. Type: U.S.A.. WASH- INGTON: Pullman, Sep 1897, C.V. Piper 1535 (N Characteristics as key. Widespread in the intermountain zone from Oregon to British Columbia and extending eastwards to Montana, var. piperi is the only variety at the western edge of the species’ range. In western Montana interme- diate forms occur while in the Okanagan when varieties piperi and chrysocarpa occur together they maintain their distinctness. The indumentum of the inflo- rescence is + lanate. Ze. gees chrysocarpa var. vernonensis J.B.Phipps & O’Kennon, var. nov. e E: CANADA: BRITISH COLUMBIA: Okanagan Valley, Okanagan LR. no. err Rd., 2 mi SW jet. Hwy 97, natural hedges, bush 3 m tall, habit very erect, fruit plum colored, flesh succulent-mushy golden, 26 Aug 2000, J.B. Phipps & S.R. Phipps 8178 (HOLOTYPE: UWO; ISOTYPES: BRIT, CAN, TRT, V). Frutices, saepe multicaules; ramificatione pauca pro ratione, range -3 m altae; spinae 3-4.5 cm longae, nitenter brunnescentes secundo anno exeunte, cinerascentes in maturitate; ramuli tendentes cri pilosi et rubi in juventute, glabrescentes; annotini + nitenter brunnei, veteriores atrocinerei. Bolla ene petioli 10-20 mm longi, primum + dense-pubescentes, in maturitate ay a aliquot glandulis secundum sulcum; laminae 3-6 cm longae, ovatae vel ovato-rhombeae in forma generali; apex obtusa vel subacuta; basis cuneata; lobi laterales 3-4/latus, acuti vel + obtusi apice vix cus an max ae us. ca. 15%; mS gone bue HURST supncute Dieter basales; 1c] venatio craspedodroma, 3-4 } scabro-pubescentes, persistentes; saperier: abaxiales fere Biabiae sed pilosae cecum venas, + coriaceae in maturitate et saepe leviter convectae, olivaceae; color autumnalis flavus purpura saepe = superposita. Inflorescentiae 5-14 -florae; rami lanati, ferentes caducas, anguste-oblongas membranaceas, ao uli-marginatas, aliquantum brunneo-rubras bracteolas. Flores 18-20 mm ee hypanthiut sla infra, glabrum supra; lobi ee minute glandulo- serrati; stamina 8-10, aye red pallide roseis; styli 4-5, exorientes a br it pilorum. Fructus 12 min diam. subglobosi, + pilosi, rubro-purpurei vel fere rae tend: caro aurea: lol ycis semi-persistentes, erecto-patentes; pyrenae 4-5, sulcatae dorsaliter; lateribus planis. 74 BRIT.ORG/SIDA 21(1) Shrubs, often multistemmed with relatively little branching, us. 2-3 m tall; thorns 3-4.5 cm long, shiny brownish black at 2 yrs, graying with age, some- what stout, straight to slightly recurved; extending twigs crisped hairy young, green tinged reddish young, glabrescent; at one year + shiny brown; older dull gray. Leaves deciduous; petioles 10-20 mm long, + dense-pubescent at first, largely persisting, with several glands along the margins of the sulcus; blades 3-6 cm long, ovate to ovate-rhombic in general shape, tip obtuse to + acute, base cuneate; lateral lobes 3-4/side, acute (more likely in smaller or vegetative leaves) to somewhat rounded with a barely cuspidate tip, max. LI] us. ca. 15%, margins with very small subacute teeth except near the entire base; venation craspedo- dromous with 3-4 principal lateral veins/side; adaxially appressed scabrous- pubescent, persisting; abaxially pilose on the veins, surfaces glabrous, rather coriaceous at maturity and often slightly convex, dull olive green; autumnal color yellow often overlayed with dull purple. Inflorescences 5-14-flowered; branches lanate, bearing caducous, narrow-oblong, membranous, somewhat brownish-red, gland-bordered bracteoles. Flowers 18-20 mm diam.; hypan- thium externally lanate below, glabrous above; calyx-lobes triangular, minutely glandular-serrate; stamens 8-10, anthers generally pale pink; styles 4-5, arising froma very short tuft of hairs. Fruit 10-12 mm diam. subglobose, + hairy, bur- gundy or lake to nearly black when over-ripe; flesh golden; calyx-lobes semi- persistent, erecto-patent; nutlets 4-5, dorsally grooved, laterally smooth. Crataegus chrysocarpa var. vernonensis is locally common in the northern Okanagan valley between the Vernon area and Salmon Arm. It occurs in the more xeric habitats in which hawthorns of the region are found, for instance, sometimes tolerating sandy soil. For this reason and its early ripening the fruits are apt to shrivel during September when the fruits of other hawthorns in the area are normally still plump. Perhaps for this reason it is under-collected. The particularly xeromorphic nature of var. vernonensis is such that it is seldom found growing among the larger, more mesomorphic hawthorns such as mem- bers of ser. Purpureofructi or C. okennonii. It is among the earliest to flower of the hawthorns of the area, typical for all ser. Rotundifoliae. Crataegus chrysocarpa var. vernonensis is known from about 17 collection numbers and in diagnostic characteristics it is remarkably uniform. The multi- stemmed, relatively little-branching, erect habit is conspicuous in well-devel- oped plants, as are the lanate inflorescences, pale pink anthers and fruit already ripe and burgundy-colored in late August. These characters help to distinguish the new species from other members of ser. Rotundifoliae while the tendency to blunt lobes in the leaf helps to distinguish it in midsummer from rare lanate forms of sympatric var. chrysocarpa. a. Additional specimens examined. CANADA: British Columbia: Okanagan Valley, Okanagan Indian Reserve no. 1. Hawhola Rd., erect bush, 8 ft. tall, 1] May 2002, ].B. Phipps & R,J.O’Kennon 8313 (UWO). Okanagan Indian Reserve no. 1, Westside Rd, N end, bush 4 ft. tall, fruit purple, 18 Aug 1994, J.B. PHIPPS AND O’KENNON, REVIEW OF CRATAEGUS SERIES ROTUNDIFOLIAE 75 Fic. 3. Line drawing of C. chrvsocarpa var. vernonensis from J.B. Phi 889 (LIWO) 1/B Phi 883q (UWO). flower- J 7 P PrP \ U rP i dfl tails; J.B. Phipps and R.J. O’Ke 989 (UWO) and R. Jackson for J.B. Phi PI PP g 8460 (UWO), fruiting and fruit details. Scale bars = 1 cm. S. Laurie-Bourque del. 76 BRIT.ORG/SIDA 21(1) Phipps & RJ. O’Kennon 6985 (UWO); Spallumcheen Mun,, top of ravine above Old Kamloops Rd., opposite Swan Lake, bush 3 ft. tall (= JBP 6889 ), 19 Aug 1994, J.B. Phipps & RJ. O’Kennon 6987 (BRIT UWO): same loc., 3 May 1994, J.B. Phipps 6889 (= JBP 6987) (UWO); same loc., both 24 September 1993, J.B. Phipps & RJ. O’Kennon 6795 (UWO), 6796 (UWO); Spallumcheen Mun., Pleasant Valley Rd., N of Vernon, bush 1.5 m tall, 26 September 1993, J.B. Phipps and RJ. O’Kennon 6816 (UWO); same loc., 26 September 1993, J.B. ame and RJ. O’Kennon 6816a (UWO); same loc., 26 September 1993, J.B. Phipps & RJ. O’Kennon 6817 (UWO),; same loc., bush, 5 ft. tall, 2 May 1994, J.B. Phipps 6883a (UWO. Spallumcheen Mun,, hwy 97, 1/2 mi W jet. Westside Rd, bush 2 m tall, 19 Aug 1994, J.B. Phipps & RJ. O’Kennon 6994 (BRIT, UWO): Spallumcheen Mun,, eastern end L. & A. Cross Rd. (bank below hwy 97 A), bush 1.3m Hau, 18 Aug 1995, J.B. Phipps 7160 (UWO), same loc., bush 1.5 m tall, 12 May 2002, J.B. Phipps & RJ.O’Kennon 8321(BRIT, beeen Spallumcheen Mun., Otter Lake Cross Rd., bush 2.5 m tall, 16 May 2002, J.B. cs 8331 (UWO); Spallumcheen Mun., McLeery Rd., ca. 2 mi S Enderby, 25 Aug 2002, R.Jackson for J.B. Phipps ete UWO). cay Indian Reserve no, 2, bush 3m tall, 16 May 2002, J.B. Phipps 8337 (BRIT, UWO); Salmon Arm, oldt icipal di site, hawthorn thickets, bush 3.5 m tall, fruit red-burgundy, 20 Aug 1994, is PI hipps & RY. oO Kennon 7028 (UWO). Additional du- plicates may be distributed later. ACKNOWLEDGMENTS Thanks are due to my colleague, Antony Littlewood, Dept. of Classical Stud- ies, University of Western Ontario for critically checking my Latin diagnoses; to the Natural Sciences & Engineering Research Council of Canada for discov- ery award Al726 which supported the work; to Sheila Phipps who provided invaluable help as mentioned under the eponymous species and to Susan Lau- rie-Bourque who continues to provide exquisite illustrations. REFERENCES AsHe, W.W. 1900. Crataegus chrysocarpa.|n: Some new species of the genus Crataegus and notes on some dichotomous Panicums. North Carolina Coll. Agric.Mech. Arts 175:110. Bovin, B. 1967. Crataegus. In: Flora of the Prairie Provinces. Provancheria 2:51—52. Britton, N.L.1901.A new Crataegus from Washington. Torreya 1:55. Dickinson, T.A.,S. BeLaousorr, R.M. Love and M.Muniyamma. 1996. North American black-fruited hawthorns. |. Folia Geobot. Phytotax. 31:355-371. Dorn, R.D. 1988. Vascular plants of Wyoming. Mountain West Publishing, Cheyenne, WY. Ecateston, W.W.1908a.The Crataegi of the northeastern United States and adjacent Canada. Rhodora 10:73-84. Eaceston, W.W. 1908b. Crataegus sect. Rotundifoliae. In: Robinson, B.L.& M.L. Fernald. Gray’s New Manual of Botany, ed. 7. 467-469. American Book Co., New York, Cincinnati and Chicago. Ecateston, W.W. 1909. New North American Crataegi. Bull. Torrey Bot. Club 36:639-642. Harms, V.L. 2003. Checklist of the vascular plants of Saskatchewan. Univ. Saskatchewan Extension Press, Saskatoon, SK. Howett, TJ. 1898. Crataegus columbiana.|n: Flora of northwestern America 1:163.Published by author, Portland, OR. PHIPPS AND O’KENNON, REVIEW OF CRATAEGUS SERIES ROTUNDIFOLIAE 77 KruscHe, E.L. 1965. Contributions to the taxonomy of Crataegus. Milwaukee Public Mus. Public. Bot. 3:1-273. Lamarck, J.B.P. De M. 1783. Crataegus rotundifolia. In: Encyclopédie Méthodique 1:84. Pantouche, Paris and Plonteaux, Liege. Moench, C. 1785. Crataegus rotundifolia. In: Baume und Stauden. J.G. Fleischerisch, Frank- furt & Leipzig. Packer, J.G. 1983. Flora of Alberta, 2nd ed. Univ. Toronto Press, Toronto. Pripps, J.B. 1995. The identity of Crataegus columbiana and its relationship to C. piperi (Ro- saceae), Taxon 44:405—408. Puipes, J.B. 1997. Introduction to the red-fruited hawthorns (Crataegus, Rosaceae) of west- ern North America. Canad. J. Bot. 76:1863-1899. Puipps, J.B. and RJ. O’Kennon. 2002. New taxa of Crataegus (Rosaceae) from the northern Okanagan-southwestern Shuswap diversity centre. Sida 20:115-144. ReHper, A. 1940. Manual of cultivated trees and shrubs hardy in North America. 2nd ed. Macmillan, New York. SARGENT, C.S. 1905. Manual of the trees of North America. Houghton Mifflin, Boston and New York. 78 BRIT.ORG/SIDA 21(1) BOOK NOTICES New York BOTANICAL GARDEN ANbREW HENDERSON. 2002. Evolution and Ecology of Palms. (ISBN 0-89327-444- 5, pbk.). The New York Botanical Garden Press, 200th Street & Kazimiroff Boulevard, Bronx, NY 10458-5126, U.S.A. (Orders: 718-817-8721, 718-817-5126 fax, http://www.nybg.org, nybgpress@nybg.org). $28.00, 198 pp., 38 b/w photos, 37 line drawings, 37 tables, glossary, bibliography, index, 7" = 10" pally Pu families of plants found in tt | regions of the world. Within the tropics, espe- I blisher Comments: “Palms are hee me most sige diverse, and economically important cially in lowland rnoist forests, palms are extremely abundant in terms of both numbers of species and individual, and are clearly an important part of the ecosystem. in addition, palms have a fasci- nating life history, much of it related to their morphology: they are not trees, yet they have woody stems and are not classified as herbaceous plants.” “Henderson brings together and analyzes the relevant literature and data in an attempt to un- derstand something of the evolution and ecology of the palm family, and integrates this desparate i oD knowledge into a cohesive whole.” Henry A. GLEASON AND ARTHUR CRONQUIST. 2004. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, Second Edition, corrected. (ISBN 0-89327-365-1, hbk.). The New York Botanical Garden Press, 200th Street & Kazimiroff Boulevard, Bronx, NY 10458-5126, U.S.A. (Orders: 718- 817-8721, 718-817-5126 fax, http://wwwnybg.org, nybgpress@nybg.org). $69.00, 862 pp., 6" x 9". This is the 7 printing of the second edition; the first printing was in 1991. This new printing comes with corrections (suggested by users) to the good keys and descriptions, new information includes photographs and short ears of Gleason and Cronquist, author abbreviations standardized new and much 1 sive indexes, and a new looking cover design. SIDA 21(1): 78. 2004 CROTON BIGBENDENSIS (EUPHORBIACEAE), A NEW SPECIES FROM TRANS-PECOS, TEXAS B.L. Turner Plant Resources Center University of Texas Austin, Texas 78712-0471, U.S.A. ABSTRACT Croton bigbendensis B.L. Turner, sp. nov. is is described from Trans-Pecos, Texas. It is closely related to C. dioicus Cav. and has passed for tl treatments of Croton for the Southwestern United States and Mexico. Croton bigpenile nsis is largely confined to the lower elevations of the Big Bend Region in Brewster, Presidio, and Hudspeth counties where it dominates sandy flats along the Rio Grande. It is readily distinguished in the field by its bushy habit, and elongate upper internodes, the latter having linear-lanceolate leaves. So far as known, it does not co-occur with C. dioicus, nor does it appear to intergrade with the latter in regions of near contact. Key Worps: Croton, C. dioicus, Euphorbiaceae, Texas RESUMEN Se describe Croton bigbendensis B.L. Turner, P nov. de oan: Pecos, Hee Ee a enlaeele con C. dioicus Cav. y ha pasado por tal especie en la mayoria de de los Estados Unidos y México. Croton bighe ndensis esta epee: a ba ee ejacone de la region de Big Bend en los condados de Brewster, Presidio y H a lo largo de las Haniras arenosas s del Bie nies Se distingue bole en d campo por su habito nee e ultimos con hojas linear-lanceoladas. Por lo conocido hasta ahora dina poco | grad él en las regiones de contacto cercano. Field work in the Trans-Pecos region over a several year period has convinced me that there exists an undescribed species of Croton, this long hidden within the taxonomic fabric of C. dioicus. Its description follows: Croton bigbendensis B.L. Turner, sp. nov. (Figs. 1,2,3). Type: U.S.A. TEXAS: PRESIDIO :ca.20 mi W of Hwy. 90 along county road 2017 (dirt road to the Rio Grande), sandy flats along old flood plain of Rio Grande, 22 Aug 2002, B.L. Turner 22-204A (pistillate) and 22-204b (staminate) (HOLOTYPE: TEX; ISOTYPES: MEXU, SRSC). Similis Croton dioico Cy sed potas su acca aut aan estad 0.5m altus, ramosissimus e basi p g 1.5-3.0 plo longioribus. Suffruticose perennial herbs or ans nee 40-60 cm high, the stems arising from lignescent tap roots. Stems densely pubescent with sessile peltate scales, the latter ca. 0.4 mm across. Lower leaves lanceolate; uppermost intern- odes elongate, bearing linear-lanceolate leaves 5-7 times as long as wide, pu- bescent like the stems. Staminate flowers with mostly (8-)9-12(-15) stamens; SIDA 21(1): 79-85. 2004 80 BRIT.ORG/SIDA 21(1) filaments pilose, 1.5-2.0 mm long. Pistillate flowers with style branches ca. | mm long, fruits globose, ca. 55mm long, 5 mm wide. Chromosome number, 2n = 28. Representative Specimens: UNITED STATES. TEXAS: Brewster Co.: Castalon Ranger Station, 17 Aug 1972, Bacon & Hartman 1438 (LL); Big Bend Natl. Park, Chisos Mts., 26 Jun 1937, Marsh 41 (TEX); flats N of Chisos Mts., 8 Jul 1932, Muller s.n. (TEX); near upper Tornillo Creek Bridge, “subshrub to 0.5 m high,” 28 Apr 1984, Powell & Powell 4329 (TEX); 24.9 mi S of Marathon, 16 Jun 2003, Turner 23-152 (TEX); 25.7 mi along La Linda highway from highway 385, 16 Jun 2003, Turner 23-155 (TEX), 4 mi of highway 2627 along dirt road to Brushy Creek Canyon,17 Jun 2003, Turner & Dodson 23-163 (TEX); between Todd Hill and Burro Mesa, yellow clay soils, 3200-3500 ft, 1 Aug 1955, Warnock 12789 (LL, SRSC):; Nine Point Mesa Ranch, 22 Apr 1998, Webster & Westlund 32586 (TEX): near Rio Grande, creek bottoms, Compton Place, 30 Aug 1910, Youngs.n. (TEX). Hudspeth Co.: ca. 5 mi Sol Finley RR Station, along dirt road to IH 10, 3 Sep 2002, Turner 22-226 (SRSC, TEX); 6 mi W of Fort Hancock, 7 Sep 1955, Warnock 13767 (SRSC). Presidio Co.: Big Bend Ranch along Rio Grande, mouth of Canyon Colorado, 5 Oct 1975, Butterwick & Lamb 1771 (TEX); 4.5 mi N of Ruidosa, 29 May 1941, Hinckley 1541 (TEX); 3 mi N of Candelaria, 3 Sep 2000, Billie & Matt lurner 20-508 (MEXU, NY, SRSC, TEX). MEXICO: CHIHUAHUA. Mpio. Manuel Benavides. ca. 7 mi W of Providencia, silty desert plains, locally abundant, 10 Aug 1940, 1. M. Johnston & Muller 106 (LL). — The species is named for the Big Bend region of Texas and closely adjacent Mexico, to which it is largely restricted. Altogether, +4 collections of C. bigbendensis were examined in this study (LL, SRSC, TEX): 22 from Brewster Co. 18 from Presidio Co., and two from Hudspeth Co. Only a single collection was examined from Mexico, this cited in the above. Johnston (1959) provided a detailed treatment of the Texas species of Cro- ton. In this he noted that C. dioicus is typified by material collected in the state of Hidalgo, Mexico. He further commented that it is a “widespread” (distrib- uted from Texas to southern Mexico) and “somewhat weedy” taxon, occurring “in disturbed ground along roads and railroads.” In spite of the fact that he knew the Trans-Pecos region quite well, he did not discern the species proposed herein. Finally, it should be emphasized that, to my knowledge, the two species do not occur together, nor do they appear to intergrade in regions of near contact (cf. figs. 4 and 5), hence my recognition of C. bigbendensis as a good biological species. Such “cryptic species” are becoming increasingly common in Texas, even in regions that have been well studied: for example, the two new species of Liatris created out of the fabric of Liatris punctata Hook. (Nesom & O’Kennon 2000), and that of Nemophila sayersensis carved out of the previously well-studied N. phacelioides Nutt., Simpson et al. 2001). — Lectotypification Johnston (1959) placed C. neomexicanus Muell. Arg. in synonymy with C. dioicus, having not seen type material, although he was aware that several early workers had taken up the name C. neomexicanus for what should have been called C. dioicus. Indeed, early on | thought the former name might apply to what I describe herein as C. bigbendensis. To resolve this issue | borrowed type TURNER, A NEW SPECIES UF FPEQUS ACUIUN UF IEARS NOLOTYPE OF: Th TEX: Croton n gence B. L. Turner > BLL. Turner, 2002 PLANTS OF TEXAS Creton diceus Cav. ESIDIO COUNTY: northwestermmost part of county ca, 20 miles west By highway 90 long county mod 2017 which nuns into a dirt road poing Jo the Rio Grande, plants collected in Mat area nest io large metallic shed ca, 200 yards from Rio Grande g SN, 104 SW B.L. Turner 22. 204A 22 Aug 2002 nan Fic. 1. Croton bigbendensis; holotype (B.L. Turner 22-2044, pistillate, TEX) 82 BRIT.ORG/SIDA 21(1) UNIVERSITY OF EXAS HERBARIUM (OL i Cet ——— ATH OT EL WHEY ann I " a bi. if sir Aebharaalh a PLANTS OF TEXAS Det: BLL Turner, 2002 ade ei d tolaus Cw. aA Det BL Turner PRESIDIO COUNTY ca 2 miles west & H : e iH if Ff é i : i q & ow sy B.L. Turner 22-2048 22 Avg 2002 iain 00398032 A Fic. 2. Croton bigbendensis; holotype (B.L. Turner 22-204b, staminate, TEX) TURNER AENIVIN VE TLAAD L | as ae eae oe ae ee eee shea! | lee ie ae oe ee ie es ae ee a eee cy L oa ee Re re pepe tt ty Pk te Hie Vie ie ete ae de gee ae ee ee ee ee a a a a e*eiei@ eee, kK © De ee re ee 5 Bao ie Te Ss Loree pee i | OY O F e j Sh, | e e rexel Bec a Pe. * 7 ra = be ae Sa eo ey Va Bae Ne OGG We teg ies Ce rT Tee. " ¢ ere Tel Se ye se we, ) paket ee ae | wtp N Rs in Orb os Bo | : * em, a ry mae A on a oe { i \ —i, Ob Oo | ae ae NL we ~ Xe G00 en a A a OO “ O se! e % L vane Se iad o, Oe hy we 4 ae \, i 7 pop hs [ iSiet 75 7 = ae 3 ony Z CROTON ee | le © bigbendensis i | | § @ dioicus 7 sg oe eet / Fic. 3. Distributi f Croton bigt 1 C. dioicus in Texas materials of C.neomexicanus from several institutions, the taxon itself typified by diverse collections, some of these said to have been collected by Charles Wright in the Trans-Pecos region of Texas. Clearly the name needed lectotypifi- cation, which follows. In the protologue of C. neomexicanus, its author cited three collections: Charles Wright 1800, w/o locality, 1851-52; Charles Wright 642 “Expedition from western Texas to El Paso, New Mexico, May-October, 1849,” and Berlandier 3211, Mexico: Nuevo Leon “in campsis,” Jun 1844. Specimens of these several collec- tions were borrowed from GH and NY. From among these Wright 1800 (GH) was selected asa suitable lectotype; isolectotypes were also recorded at GH and NY. The lectotype has both male and female plants mounted on the same sheet, as does the GH isolectotype, the latter mounted on the same sheet with Wright 642 (a paratype). All of the specimens concerned are quite similar, possessing the habit, upper nodes, and leaf shape of rather typical C. dioicus. BRIT.ORG/SIDA 21(1) J a \ ical habit of C di oF .Typ Fic.4 15 mi. NW of Presidio, Texas; note the bushy a \ IP father of the sheriff of Brewster Co., Pv rounded habit of th Texas). TURNER, A NEW SPECIES OF CROTON FROM THE TRANS-PECOS REGION OF TEXAS 85 Chromosome numbers Urbatsch et al. (1975) reported chromosome counts for 11 collections of C. dioicus (nine of these were diploid with 2n = 28, and two tetraploid with 2n = 56 pairs). Examination of the vouchers concerned (LL, TEX) revealed that only one of these (Brewster Co: Bacon & Hartman 1438) belonged to what is here described as C. bigbendensis. Since chromosome numbers of the C. dioicus collections included both diploids and tetraploids, chromosome number alone is not useful in dis- tinguishing between the two taxa. ACKNOWLEDGMENTS My wile Gayle provided the Latin diagnosis. Grady Webster and Mike Powell reviewed a preliminary draft of the paper. The following herbaria provided loans of type materials: GH, NY, and US. Thanks to all for the kindness rendered. REFERENCES JOHNSTON, M.C. 1959. The Texas species of Croton (Euphorbiaceae). SouthW. Naturalist 3: 175-203. Nesom, G.L.and R.J.O’Kennon. 2001.Two new species of Liatris series Punctatae (Asteraceae: Eupatorieae) centered in north central Texas. Sida 19: 767-788. Simpson, B.B., D.M. Hetrcort, and J.L. Nerr. 2001. A new cryptic species of Nemophila (Hydrophyllaceae) from Texas and the lectotypification of N. phacelioides Nuttall. Lundellia 4:30-36. Urparscn, L.E.et al. 1975.Chromosome numbers for North American Euphorbiaceae. Amer. J.Bot.62:494-500. BRIT.ORG/SIDA 21(1) BOOK NOTICE DesoraH P. DeLMER, Hans J. BOHNERT, and SABFEHA MERCHANG (eds). 2003. Annual Review of Plant Biology: Volume 54, 2003. (ISBN 0-8243-0054-6, hbk; ISSN 1040-2519). Annual Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews.org, 800-523- 8635, 650-493-4400, 650-424-0910 fax). $165.00 (USA), $170.00 Unt'L), 751 pp. 6" x 9" Contents of Volume 54 of Annual Review of Plant Biology: Conjectures, Refutations, and Extrapolations Understanding the Functions of Plant Disease Resistance Proteins Protein cea ii in Plants Plant Peroxiredoxin Nitric Oxide: The fecitiy of an Extensive Signal Molecule Biosynthesis and Metabolism of Brassinosteroids The COPY Signalosome: Regulating Plant Development Through the Control of Proteolysis — lron Transport and Signaling in Plants From Bacterial Glycogen to Starch: Understanding the Biogenesis of the Plant Starch Granule The Plant Cell Cycle Phospholipid-Based a ing in Plant Gibberellins and Flowering of Grasses aa Cereals: Prizing Open the Lid of the “Florigen” Photosynthesis of Overwintering amie Plants Structure of Linkage Disequilibrium in Plan Single-Nucleotide Mutations for Plant see Genomi How Do Cells Know What They Want To Be When They oe Up? Lessons from Epidermal Patterning in Arabidopsis Transfer Cells: Cells Specialized for a Special Purpose Chloroplast Movement Cryptochrome Structure and Signal Transduction Membrane-Bound Diiron Carboxylate Proteins Lignin Biosynthesis Apomixis: A Developmental Perspective Molecular Mechanisms and Regulation of K+ Transport in Higher Plants Perception and Signal Transduction of Cytokinins Functional Genomics of P450s Metabolomics in Systems Biology Remodeling the Cytoskelton for Growth and Form: An Overview with Some New Views SIDA 21(1): 86. 2004 A NEW SPECIES OF CENTAURIUM (GENTIANACEAE) FROM TRANS-PECOS TEXAS B.L. Turner Plant Resources Center The University of Texas Austin, Texas 78712-0471, U.S.A. ABSTRACT Centaurium blumbergianum B.L. Turner, sp. nov. is described from near Ruidosa, Presidio Co., Texas. It is a perennial species with broad open, cymes having no obvious close relatives among the North American taxa described to date. It occurs only along saline streams and in seeps at the base of lime- stone walls in dead-end canyons where it grows with or near a number of other rare taxa such as the recently described localized endemic, Arida mattturneri and the equally rare A. gypsitherma (cf. Sida 20 [4]). Photographs of type material are presented, along with an account of its possible rela- tionchj t tax [ RESUMEN Se describe Centaurium blumbergianum B.L. Turner, sp. nov, de cerca de Ruidosa, Presidio Co, Texas. Es una especie perenne con cimas abiertas anchas, GUE no Baie parents pions entre los taxa eg | norteamericanos descritos hasta ahora. yen filtrados de la base de paredones calcareos de los finales ciegos de canones s donde crece junto o cerca de ciert numero de otros taxa raros tales como el endemismo descrito Arida mattturneri y la igualmente rara A th f. Sida 20 ua Se presentan fotografias del material tipo, junto con una relacion de su porites eae blumbergianum B.L. Turner, sp. nov. (Figs. 1, 2 ). Type: U.S.A. TEXAS: Presidio Co., ca. 2.2 mi NNW of Ruidosa in “Blumberg Canyon,” growing along sulphur and about tinajas, 26 Jul 2003, Matt W. Turner 101 (HOLOTYPE: TEX; ISOTYPES: MO, SRSC). Simile Centaurio tenuifolio (Mart. & Gal.) B.L. Rob. sed habens petala parviora pallidaque et inflorescentiam expansiorum cum flores pauciores. Perennial tufted herbs to 40 cm high. Stems 4-sided, 1-2 mm across, their edges adorned with minute, somewhat scabrous ridges, otherwise glabrous or nearly so. Leaves opposite throughout, linear, glabrous, those at mid-stem mostly 15- 25 mm long, 0.4-0.6 mm wide. Flowers 3-15 per stem, forming an open cyme up to 20cm across, the pedicels mostly 1.5-3.0 cm long. Sepals 5, linear-lanceolate, 7-10 mm long, united at their base for ca. | mm; apices slender and narrowly apiculate. Corollas pink; tubes 7-9 mm long, becoming transparent at matu- rity, constricted just below the 5 lobes, the latter 8-9 mm long, 3.5-4.5 mm wide. Stamens 5, exserted from tube for 2-3 mm, the anthers ca. 3 mm long at first, but 2-3 mm long and markedly helical at full anthesis. Style exserted from the tube for ca. 4 mm, the stigma somewhat bilobed, ca. 0.5 mm across. Capsules SIDA 21(1): 87-91. 2004 88 BRIT.ORG/SIDA 21(1) UNIVERSITY OF HERBARIUM PLANTS OF TEXAS entisnacese Centaurium blumbergianum B. L. Turner. sp. neve Det BLL. Turner PRESIDE e \TY: of Rui “Bhanberg Canyon,” growing along percanial sulphur streams and abwet tinajas, FIRST RECORD FOR TEXAS 30 00.45 NL Ons Ww Maw W, Turner yoy 26 Jul 2003 hy , ce Au Fic.1 Fee at ee eee holotype (M.W. Turner 107, TEX). J at TURNER TLARD 89 “er g o Be ’ 5 git ; oa Py t; Lion a 4. Meaty Bt th 4 te “ee See we a Fic. 2G H h] } - + on \e-l £ lo /] ) 90 BRIT.ORG/SIDA 21(1) Fic. 3. James and Olivia Blumberg. bivalvate, narrowly oval, 8-10 mm long, ca. 3 mm across; ovules numerous to each carpel. Seeds ovoid, brown, ca. 0.6 mm long, 0.4 mm wide, at maturity sculpted with a peanut-like raised reticulum. Among the Texas species, C. blumbergianum has no close relatives, although an anonymous reviewer suggests that it might be compared favorably with C. TURNER, A NEW SPFEXILS VE ICARS 91 multicaule B.L. Rob. a rather small annual with broadly lanceolate to oblan- ceolate basal leaves. In contrast, C. blumbergianum is a relatively robust peren- nial with persistent linear basal leaves. In her treatment of the Mexican and Central American species of Centaurium by Broome (1974), C. blumbergianum will key to or near C. brachycalyx Standl. & L.O. Williams, a species of southern Mexico and Central America. It differs from the latter in numerous characters, including habit, in- florescence, and leaf shape. Except for its small flowers and expanded inflores- cence, C. blumbergianum would appear to be closest to C. tenuifolium (Mart. & Gal.) B.L. Rob.,a taxon confined to the mountains of western Mexico (Nayarit to Guerrero). Among the species of Centaurium in the U.S.A. it most closely approaches Centaurium namophilum Reveal, Broome & Beatley, a localized species of Nye Co., Nevada and closely adjacent California; C. blumbergianum differs in hav- ing a more robust perennial habit and much more diffuse inflorescence. Centaurium namophilum and the closely related C. tricanthum (Greisb.) B.L. Rob. (according to Reveal et al. 1974), are said to “occur near or along stream drainages, marshy places or on alkaline soil.” Such habitats also house C. blumbergianum. Etymology.—It is a pleasure to name this remarkable new species for Mr. James Blumberg and his wife Olivia (Fig. 3), residents of Ruidosa, Texas. They own the property concerned and Mr. Blumberg eagerly accompanied Matt Turner and myself in our efforts to collect type material. ACKNOWLEDGMENTS lam grateful to James Blumberg for permission to visit the site concerned and to my wile Gayle for the Latin diagnosis. A special thanks to Robert Harms for helping “upgrade” the picture of the Blumbergs. James Reveal and an anony- mous reviewer made helpful suggestions. REFERENCES Broome, C.R. 1973. Systematics of Centaurium (Gentianaceae) of Mexico and Central America. Ph.D. Dissertation, Duke Univ., Durham, North Carolina, University Microfilms, Ann Arbor, Michigan. ReveAt, J.L., C.R. Broome, and J.C. Beatiey. 1974. A new Centaurium (Gentianaceae) from the Death Valley region of Nevada and California. Bull. Torrey Bot. Club 100:353-356. Turner, B.L. 1993. The Texas species of Centaurium (Gentianaceae).Phytologia 75:259-275, BRIT.ORG/SIDA 21(1) BOOK NOTICE DEBORAH P. DELMER and SABEEHA MERCHANG (eds). 2004. Annual Review of Plant Biology: Volume 55, 2004. (ISBN 0-8243-0655-4, hbk; ISSN 1543-5008). Annual Reviews Inc., 4139 E] Camino Way, PO. Box 10139, Palo Alto, CA 94303- 0139,US.A (Ones www.A g, 800-523-8635, 650-493-4400, 650-424-0910 fax). $173.00 (USA), $178.00 (intl .),659 pp., 6" x 9". Contents of Volume 55 of Annual Review of Plant Biology: Alternative NAD(P)H Dehydrogenases of Plant Mitochondria An Untoreseen Voyage to the World of Phytochromes Sterols Biosynthesis and Accumulation o Decoding Ca2+ Signals Through Plant Protein Kinases DNA Methylation and Epigenetics Genetical Regulation of Time to Flower in Arabidopsis thaliana How Do Crop Plants Tolerate Acid Soils? Mechanisms of Aluminum Tolerance and Phosphorous Efficiency Metabolic Channeling in Plants Molecular Mechanism of Gibberellin Signaling in Plants Naturally Occurring Genetic Variation in Arabidopsis thaliana Phosphoenolpyruvate Carboxylase: A New Era of Structural Biology Phytoestrogens Plastid Transformation in Higher Plar Reactive Oxygen Species: Metabolism, oe Stress, and Signal Transduction Rhamnogalacturonan II: Structure and Function of a Borate Cross-Linked Cell Wall Pectic Polysaccharide Rising Atmospheric Carbon Dioxide: Plants FACE the Future Single-Cell C4 Photosynthesis Versus the Dual-Cell (Kranz) Paradigm Symbioses of Grasses with Seedborne Endophytes The Generation of Ca2+ see in Plan The Ubiquitin 26S Proteasome Proteol “ie Pat ul bale Mechanisms for Organic Focus of Carbon om Nitrogen Between Source and Sink S Vectors for Gene Sliencing: Many Targets, Many Tools V neers Chromosome Structure/Organization way SIDA 21(1): 92. 2004 A NEW COMBINATION IN DENDROCALAMUS (POACEAE: BAMBUSOIDEAE) Muktesh Kumar & Remesh M. N. Unnikrishnan otany Division Department of Bota Kerala Forest Research Institute SVRNSS College, TP Puram, a Peechi,680 653, Thrissur, Kerala, INDIA Kottayam, INDIA muktesh@kfri.org ABSTRACT A new combination, Dendrocalamus stocksii Munro) M. Kumar, Remesh & Unnikrishnan, is pro- posed to accommodate a widely occurring economically important bamboo in South and central India which was formerly oe ee an Oxytenanthera Munro and Pseudox ytenanthera Soderstr. & RP. Ellis. RESUMEN Se propone una nueva cunuaneae cpsuupeann us hana: tune) M. an ar, > Remiesh = Unnikrishnan, para acomoda sur y centro re la India que fue descrito anteriormente en Oxyte nanthera Munro y Pseudo- xytenanthera Soderstr. &@ RPE INTRODUCTION During revisionary studies on some Indian bamboos, the authors undertook a detailed study on the delimitation of Dendrocalamus Nees, Oxytenanthera Munro, and Pseudoxytenanthera Soderstr. &@ R-P. Ellis. All of the species de- scribed in the above genera were critically examined using the specimens de- posited in various herbaria including the type specimens. Field examination of the Indian species belonging to these genera was carried out. We found that O. stocksii Munro showed more affinity to the Genus Dendrocalamus in the veg- etative and floral characters. Oxytenanthera stocksii was first described by Munro (1868) based on the specimen collected by Stocks from the Concan area. This species is distributed in South India, along the Konkan coast up to Karwar. Subsequently, Talbot also collected it from the Coompta River in 1884 and Karwar in 1889. Munro's spe- cies was recognized by many later workers, e.g., Beddome (1873), Gamble (1896), Camus (1913), etc. It was, however, Holttum (1956) who asserted that the genus Oxytenanthera was truly represented by the African species O. abyssinica due to the peculiar nature of its ovary ie. the ovary attenuate upwards very gradu- ally into a more or less three angled hollow structure which bears the stigmas at its apex and the cavity of the style appears not to be continuous with the cavity which contains the ovule. He concluded that the rest of species described SIDA 21(1): 93-96, 2004 94 BRIT.ORG/SIDA 21(1) within this genus should be transferred to some other genera. Based on this analysis, Majumdar (1989) transferred this species to a new genus, Pseudotenanthera Majumdar. Unfortunately, Pseudotenanthera is nom. superfl. and illegitimate for Pseudox ytenanthera Soderstr. & Ellis 1988). Nguyen (1990) treated this taxon under the genus Gigantochloa but it could not be included under this genus due to the presence of stamens with free filaments. Therefore, Naithani (1991) transferred this species to Pseudoxytenanthera as P. stocksii (Munro) Naithani. Pseudoxytenanthera is characterized by the presence of straggling culms, inflorescence with spikelets of semiverticillate clusters, mona- delphous stamens, and three plumose stigmas. Pseudoxytenanthera stocksii possesses erect culms, free stamens, and a single plumose style. Although the re thors described this species with fused filaments, during the present ay it was observed that the anther filaments are short and do not show true monadelphous condition. A close ination from the young stamens to ma- ture stamens revealed that the filaments are totally free. Owing to these char- acteristic features, it cannot be accommodated within the genus Oxytenanthera Munro either. Oxytenanthera stocksii differs from O. abyssinica, the type spe- cies, by having free stamens and monostigmatic ovary. In O. abyssinica the sta- mens are monadelphous and the style is divided into three stigmas. The culms and branching patterns of Oxytenanthera stocksii also resemble those of species of Dendrocalamus Nees. As in the case of the type species (ie, D. strictus), O. stocksii shows erect culms with short internodes, which have narrow lumen (solid). The inflorescence of O. stocksii is a large panicle of spi- cate heads. Even in Dendrocalamus, the inflorescence is composed of round congested globose heads. In both species, the spikelets are few-flowered and there are no lodicules. In O. stocksii and in D. strictus, the palea is keeled and ciliate on the keels and the paleas of the upper flowers are not keeled. The sta- mens have short-apiculate anthers and free filaments. Another important char- acter is the vestiture of style and stigma. In both species, the style is sparsely ciliate and ends in a single feathery stigma. In O. stocksii and other species of the genus Dendrocalamus the basal nodes bear aerial roots. Oxytenanthera stocksii can be easily separated from Dendrocalamus strictus by the eons auricles and bristles in the culm sheath, comparatively large leaves (15-22 x 15—2.5 cm), slender spikelets, slightly apiculate anthers with short filaments, elongated ovary, and oblong caryopsis. The above observations eee the aes of Oxytenanthera stocksii from the genera Oxytenanthera, Pseudoxytenantheraand Gigantochloa, which justify its inclusion within the genus Dendrocalamus. Morphological charac- teristics of D. stocksii are compared with those of the type species of Dendrocala- mus, Oxytenanthera, Pseudoxytenanthera, and Gigantochloa in Table 1. While describing Oxytenanthera stocksii, Munro had also noticed its simi- KUMAR ET AL., A NEW COMBINATION IN DENDROCALAMUS Taste 1. Comparison of Dendrocalamus ae _ Dendrocalamus strictus, Oxytenanthera abyssinica, en monadelpha an 1atter (The type species of genera). Characters D. strictus D. stocksii 0. abyssinica P. monadelpha Gigantochoa atter Culm surface = matt, wax glossy, wax glossy, wax glossy, wax glossy, wax thin furry thin thin thin Culm wall thick walled thick walled thick walled moderately moderately thickness thick walled thick walled Culm sheath small and well-developed small and well-developed well-developed auricle rudimentary with oral setae rudimentary — with oralsetae with oral setae Spikelets 2-3 flowered 2-3 flowered 1-3 flowered 1-3 flowered 3-4 flowered Lemma sparsely hairy — glabrous sparsely hairy — glabrous glabrous Apex of anthers sea slightly perfectly perfectly perfectly iculate apiculate apiculate apiculate apiculate Filaments free free United united united Stigma single single Three three single larity to Dendrocalamus strictus. The type specimen of O. stocksii, housed at Kew (K), is annotated as D. stocksii Munro, and therefore it is evident that Munro had previously considered this species within Dendrocalamus. However, he treated the species within Oxytenanthera due to the presence of slightly api- culate anthers and striated membranous lower palea. The similarities between this species and Dendrocalamus were also mentioned by Gamble (1896) who pointed out that the narrow leaves, long petioles and culm sheath were similar to that of D. strictus. These similarities also led to the misidentification of this species by subsequent authors. All the previous studies including that of Munro and Gamble were merely based on herbarium specimens, and some of the field characters and floral characters such as the nature of filaments, ovary, palea etc., were omitted. Therefore, they did not observe the correct and distinct char- acteristics of the species. The field observation on the floral and vegetative char- acters confirm the present concept of the species and hence a new combina- tion within Dendrocalamus is proposed. This bamboo species, which is economically important, is widely culti- vated throughout South India and is utilized for house construction, basket making, ladders, poles, and for several other purposes. SYSTEMATIC TREATMENT Dendrocalamus stocksii (Munro) M. Kumar, Remesh & Unnikrishnan, comb. nov. Oxytenanthera stocksit Munro, Trans. Linn. Soc. London 26:130. 1868. Pseudotenanthera stocksii (Munro) R.B. Majumdar, Fl. Ind. Enumerat.-Monocot. 280.1989. Gigantochloa stocksii Munro) Nguyen, Bot. Zhurn. Akad. NAUK. 75:224. 96 BRIT.ORG/SIDA 21(1) 1990. Psuedoxytenanthera stocksit (Munro) Naithani, J. Bombay Nat. Hist. Soc. 87:440. 1991. Pseudoxytenanthera stocksii (Munro) T.Q. Nguyen, Bot. Zhurn. NAUK 76:993. 1991. TYPE: INDIA. SOUTH INDIA. Concan: Stocks s.n. (LECTOTYPE, here selected: K). Vernacular names.—Uyi, Mula (Malayalam), Konda (Karnataka). Distribution—Endemic to Northern Western Ghats. South India; north- ern Kerala and Karnataka pane Concan coast, Goa, Maharashtra. Sel 1 Speci INDIA. K I Dist.: North Kasargod, Bamboo Products Exports 140317 (DD); Kenenatead. 29 Oct 1999 Riveendvan 20637 (KFRD. Thrissur Dist: eel ee 2000 M. Remesh 20646 (KFRD; 26 Feb 2001 Unnikrishnan 74039 (CALL. Goa: South G Ary Mar 1985, H.B. Naithani 1189 (DD). Karnataka: North Kanara Dist.: Coompta, ce 7 ei 269 (BSD); Flora of North Kanara W.A. Talbot 549974 (CAL). ACKNOWLEDGMENTS The authors wish to acknowledge J.K. Sharma, Director, Kerala Forest Research Institute, Peechi for his keen interest and encouragement and for providing the research facilities. Help rendered by S.A. Renvoize, Herbarium, Royal Botanic Garden, Kew for sending the cibachrome sheets of the type specimen depos- ited in K and the authorities of Indian Herbaria (CAL, CALI, DD, BSI, MH, and BLAT) for permission to consult specimens are gratefully acknowledged. The authors are also thankful to the Department of Science & Technology, Govt. of India, New Delhi, for the financial support to conduct this study. REFERENCES Beopome, R.H. 1873. The flora sylvatica for southern India. Madras 3:232-233. Camus, E.G. 1913.Les Bambusees. Lechevalier, Paris. Pp.149. Game _e, J.S. 1896. The Bambuseae of British India. Annals of the Royal Botanic Garden, Calcutta 7:75-76. Houttum, R.E. 1956.The classification of bamboos. Phytomorphology 6:73-90. Munro, W. 1868. A monograph of Bambuseae. Trans. Linn. Soc. London 26:126-127. Magumpar, R.B. 1989. In: Karthikeyan, S. et al. Flora Indicae, ser. 4, 2(Enumeratio Mono- cotyledonae). Botanical Survey of India, Howrah, Calcutta. Pp. 274-283. NartHani, H.B. 1991. Nomenclature of Indian species of Oxytenanthera Munro. J. Bombay Nat. Hist. Soc. 87:439-440 Nouyen, T.Q. 1990. New taxa of bamboos (Poaceae, Bambusoideae) from Vietnam. Bot. Zhurn. Akad. Nauk. 75(2):221-225. UNA NUEVA ESPECIE DE STRUTHANTHUS (LORANTHACEAE) PARA COSTA RICA J.Francisco Morales Instituto Nacional de eas ) Apto 2 22-3 Sant Di d ae COSTA RICA RESUMEN ithus (Loranthaceae) para Costa Rica. Ademas se adjunta Se describe una nueva especie de una clave para las especies del genero hes tee en el pais, incluyendo taxas no reportados atin, pero presentes en zonas cercanas de Nicaragua y Panama. ABSTRACT A new species of Struthanthus from Costa Rica is described. A key to th » known for the country is given, including two species recorded from ee ada Nicaragua, but not re- ported from Costa Rica yet. La familia Loranthaceae esta ampliamente representada a lo largo del tropico, conteniendo alrededor de unos 60 géneros y 1000 especies (Kuijt 2001). En Costa Rica se han reportado hasta la fecha 6 géneros y unas 28 especies (Morales, en prep.). Este grupo de plantas esta ampliamente distribuido y se encuentra en una amplia gama de habitads y ecosistemas, encontrandose desde el nivel del mar hasta los 3200 metros sobre el nivel del mar, tanto en bosques secos a muy huamedos 0 areas o paramos. Su forma de crecimiento les ubica en una posici6n muy particular desde el punto de vista antrdpico, ya que en cultivos frutales representa un problema que se refleja en la disminucion de la produccion al parasitar dichos cultivos, mientras que en el manejo de los bosques naturales son una fuente de alimento para varias especies silvestres, principalmente aves. Struthanthus es un género neotropical constituido por unas 70 especies (Kuijt 2001), de las cuales se conocen para Costa Rica 13 especies. Dentro de la familia se puede confundir con Panamanthus, pero este Ultimo se distingue por tener inflorescencias con las flores solitarias, mientras que en Struthanthus las flores estan dispuestas en diadas o triadas. También se puede confundir con Phthirusa, cuyo caracter diferenciante radica en la presencia de flores usualmente bisexuales, en contraste con Struthanthus que posee flores funcionalmente unisexuales. En los ultimos 15 anos, la exploracion de regiones montanosas y practica- mente inexploradas del Valle de Candelaria en Acosta, San José y de la Vertiente Atlantica de la Cordillera de Talamanca, Limon, han producido la descripcién de varios taxa (Hammel & Zamora 1990, 1993; Morales 1997, 1999), algunos de SIDA 21(1): 97-102. 2004 98 BRIT.ORG/SIDA 21(1) ellos localmente endémicos y restringidos a estas zonas geogralicas. Durante la preparacion del t de Loranthaceae para el Manual de la Flora de Costa Rica, se encontro una nueva especie de Struthanthus. Conocida hasta el momento de las areas geogralicas antes expuestas, se describe a continuacién Sag acostensis L.A. Gonzalez & J.F Morales, sp. nov. (Fig. 1, 2). Tipo. RICA. SAN JOSE: Acosta, Valle del Candelaria, Fila Zoncuano, ie 1050 mM, ol 1995 C1), Morales 4582 (HOLOTIPO: INB; ISOTIPO: MO). S. burgeri Kuijt cui affinis, corollae sessilis differt Epifita escandente, hemiparasita. Entrenudos de las hojas de 1.5-6.5 « 1.5-4.0 mm. Tallos teretes a subteretes, esparcidamente ferrugineo-lenticelados. Hojas opuestas a subopuestas, peciolos de 3-8 mm de largo y 1.5-2.0 mm de ancho, con sus bordes continuos a los margenes de la lamina: lamina de 8.0-12.5 cm de largo y 3.5-8.2 cm de ancho, ovada a eliptica, estrechandose gradualmente hacia el apice, aguda, atenuada a cuneada basalmente, borde entero y ligeramente revoluto cuando seco, esencialmente glabra, venacién pinnada, venas secundarias de 3-9 pares conspicuas, las terciarias conspicuas y a veces formando reticulos. Inflorescencias 1 a 6 por axila, 0.7-4 cm, con 2-4 triadas, subtendidas por bracteas de 0.7-2 mm, una por cada flor: pounce 03-25 cm de largo, raquis de las triadas 1.5-7.0 mm de largo; flores amarillentas, sésiles unisexuales; las masculinas desconocidas, las pistiladas 4-5 mm de diametro, corola ca. 3mm, estaminodios fusionados con los pétalos, estilo apical, estigma capitado. Fruto 4.0 x 3.0 mm de diametro, anaranjado al madurar, subgloboso, obtuso en la base, la superficie cubierta con numerosas y diminutas ranuras 0 cavidades. Distribucion. —esta especie se encuentra en bosques muy humedos en la Fila Zoncuaco (Acosta) en el Valle del Candelaria, y en la Cordillera de Talamanca (Alto Urén), a elevaciones entre 700-1100 m. Floracién y fructificacion entre Julio y Agosto. Struthanthus acostensis se reconoce con facilidad por sus inflorescencias con 2-4 triadas pediceladas, flores y frutos sésiles y tallos teretes a subteretes. Anteriormente esta especie fue identificada como Panamanthus panamensis (Rizzini) Kuijt, pero este taxa se caracteriza por sus flores solitarias, contrastando con Struthanthus acostensis que posee flores en triadas. También ha sido identificada como S. burgeri Kuijt; sin embargo, esta ultima especie se caracteriza por poseer inflorescencias con mas de 4 triadas y flores pediceladas, mientras que S. dcostensis tiene flores sésiles dispuestas en inflorescencias de 2 a 4 triadas. Por otro lado, se puede separar con facilidad de S. verstedii (Oliv) Standl. por sus hojas muchisimo mas grandes, peciolos mas largos y desarrollados, asi como por sus flores mas grandes y distribucion geografica y preferencias ecoldgicas distintas. La etimologia de la especie hace referencia a la localidad tipo, zona de cuyas BIODEVERSTDAD DE PUTO: NACTONATL “SAF quer COSTA RICA LORANTHACEAE dup.s 4 Struthnanthus Det. rae Cantén de Talamanca Alt rén. Siguiendo el camino entre ins jGuebrada hang, y Crorina. “59'00"H 700 oe "ple or amarillo canes s. ae mi oreae Frutos maduros anaranjado: Mdaly 194 Gerardo Herrera Hated ya te peated tye COSTA RICA (CR) is EOTAWECAL GARDEN (¥ ASTETS 4 Herrera 3326, NB) nO} A. Gonzalez & J.F.M 100 BRIT.ORG/SIDA 21(1) exploraciones botanicas en los ultimos 10 anos, han revelado una serie de novedades botanicas y nuevos report os y que por ree tiempo han pasado inadvertidos, como consecuencia de la extensa deforestacion prevaleciente desde hace mas de 150 anos Paratipos. Costa Rica. Limon: Talamanca, Alto Urén, 23 Jul 1989 (f1, fr), Herrera 3326 (INB, MO) A continuacion se presenta una clave para las especies del género Struthanthus conocidas hasta el momento para Costa Rica. Dos especies adicionales, reportadas en zonas limitrofes de Panama y Nicaragua que podrian encontrarse eventualmente en el pais son incluidas Triad Asi bsésil la antesis,el pedunculo a i i det 1.5mm largo, oe veces algunos ee en reennceeien ate eens r s los frutos mas de 9m | ar. fniloreseenidaxe con numerosas bracteas 10.01 [4] 6. Ratio of Mean corolla lobe length: = Continuous Mean sepal length Adaxial corolla pubescence is much more frequent among members of the MCCG (Fig. 3). In fact, two thirds of the examined individuals in this group were pubescent (41 of 62). The MCCG appears to be restricted to a more nar- rowly defined southeastern range east of the Mississippi (Fig. D. Except for a few outliers (e.g.,GA2, NC20, VAI, VA23), the geographic clus- tering is supported by the midpoint-rooted neighbor-joining tree (Fig. 4). Al- though bootstrapping yielded little support for any one branch of the tree (due to the small number of characters employed), the tree remains informative. Members of the MCCG and UCCG are grouped together—indicating greater within-group similarity across the five morphological characters than between groups. Individuals of the UCCG exhibit longer mean corolla lobes (mean=7.68) and higher mean corolla lobe length to mean sepal length ratios (mean=2.48) than individuals of the MCCG (Table 2). Results of analyses of variation (ANOVA) indicate the differences in both mean corolla lobe length and corolla:sepal length ratio to be highly statistically significant between the two groups (Table 3). It is interesting to note that the rare pubescent individuals of the predomi- nantly glabrous UCCG appear to exhibit a similar frequency distribution of corolla lobe lengths and corolla:sepal ratios as pul t members of the MCCG. ANOVA tests show no significant difference between pubescent UCCG and pubescent MCCG in both mean corolla lobe lengths (F=0.042 < Forit=3.99) and the ratio (F=0.604 < Fe;jp=3.99)—although this may be a factor of the small sam- pling size of pubescent UCCG members (n=3). Glabrous members of the MCCG appear to exhibit corolla lobe lengths much larger than the more frequent (in terms of specimens examined) pubescent members of the MCCG (Fig. 5B). How- ever, the difference in mean corolla lengths between glabrous and pubescent MCGG is not quite significant (F=3.559 < Fcrir=3.96). The difference in the 103 BRIT.ORG/SIDA 21(1) a a a Oo awed { a ~{* fe) _—— : ce me . oh i ‘eae } s ~ O 7 a Ba ‘ MILES % t a 1D0 2 3100 aga 6 \ l aoe | 1 1 Ll } o » ~ wee Io} Tautatle ( \ L fel 1 \ be 4 am ied ul a ‘ t \ 1 J nrollac ith UTUlas CHCl a corolla:sepal ratio between glabrous and pubescent MCCG members is also not quite significant (F=3.746 < Fcrir=3.96). Although the frequency peak for corolla:sepal ratios of glabrous MCCG members appears to graphically coin- cide with the peak of glabrous UCCG members (Fig. 5C & D), ANOVA results show a strong statistical difference between the two (F=9.467 > Feyit=3.93). DISCUSSION The issue of whether more than one Gonolobus species should be recognized for the southeastern United States has long confronted taxonomists and has been additionally complicated by the nomenclatural confusion surrounding Gonolobus suberosus (L.) R.Br. (see Drapalik 1969; Reveal & Barrie 1992). Small (1933), and later Perry (1938), recognized two species of Gonolobus in the South- east based on corolla pubescence and the ratio of corolla lobe length to sepal length. Small (1933) referred the taxon with glabrous corollas and lobes more than twice as long as the sepals to G. gonocarpos (Vincetoxicum gonocarpos sensu Small), whereas the taxon with pubescent corollas and lobes twice as long or less than the sepals was referred to G. suberosus (V. suberosum sensu Small). KRINGS AND QIU- 109 12.00 10.00 , , E 8.00. SRal om ‘S roy 5 "ee 2 6.00 Nat Sains ns & » = $ | ao o 4.00 Oo i” => @ Green, pubescent =” wh > Green, glabrous 2.00 mg Multi-color, pubescent Multi-color, glabrous 0.00 0 2 4 6 Sepal length (mm) £ 1 ial k ha, £1 | aes pee ee | I 7 £ IAk er Caney | th Fi t eastern United States. Fernald (1950) added some less than distinct characters to the mix, including flower bud (abruptly acuminate” vs. “gradually acute or acuminate”), calyx pubescence (“practically glabrous” vs. “glabrous or ciliolate apically”), and co- rolla lobe shape (“broadly lanceolate” vs. “linear-lanceolate”), while basically maintaining Small’s (1933) pubescence and ratio characters. Gleason (1952) treated only G. gonocarpos, stating that G. suberosus was “erroneously” accred- ited to the range of Britton and Brown’s flora. However, in a footnote, Gleason (1952) maintained the distinction between the taxa based on adaxial corolla pubescence, forwarded by previous authors (Small 1933; Perry 1938; Fernald 1950). In light of the present analyses, these concepts of specific delimitation are untenable as stated. Drapalik’s (1969) findings of overlapping combinations of character states among Gonolobus taxa in the southeast are upheld by our study. However, over- lapping character presence/absence combinations are insufficient argument against recognition of multiple taxa. Especially at the infraspecific level and in hybrid zones, some level of character overlap can be expected between indi- viduals sharing some range continuity. Our data support the notion of previ- ous workers (e.g., Small 1933; Perry 1938; Fernald 1950; Gleason 1952), that at least two Gonolobus entities occur in the Southeast that could be accorded for- 110 BRIT.ORG/SIDA 21(1) 100 | ; = _ _ [ — ——s | | . : = {1 Pubescent = Glabrous MCCG Ly Wl ‘allaa al J 4] Ie | A W {RACY } PA } mal rank. However, contrary to previous workers, we propose that the two groups respectively be defined by uniformly-green versus multi-colored corolla lobes, rather than by corolla/sepal length and adaxial corolla lobe pubescence. Al- though there are significant differences in mean corolla lobe lengths and the ratio of corolla lobe length to sepal length (Table 3), these character states can overlap at the individual level and cannot consistently separate the taxa. Simi- larly, adaxial corolla pubescence cannot consistently separate the taxa, being present in both members of the UCCG and the MCCG. However, distinct differ- ences are apparent in the frequencies of the pubescence trait (Figs. 3 & 5). In addition, with respect to their geographic distribution, the frequency of gla- brous, multi-colored flower collections increases conspicuously in the zones where the UCCG meets the MCCG (e.g., in Alabama). Similarly, the rare indi- viduals bearing adaxially pubescent, uniformly green corollas occur well within the range of the MCCG. Thus, the respective changes in pubescence frequen- cies in the MCCG or occurrence of rare character states in the UCCG outside its primary range may be cautiously hypothesized to be an effect of genetic inter- change between two taxa intergrading in distribution. Unfortunately, frequency histograms of corolla lobe length and ratio of corolla lobe length:sepal length size classes within the UCCG and MCCG shed weak light on the matter of intergradation (Fig. 5). Although our analyses sup- port the recognition of two Gonolobus entities in the Southeast, the question remains whether these should be recognized at the species level or below. Our KRINGS AND QIU- NJ Fic. Miata) Midpomns TouIsG Heighborse 6 TX18 | Thad MS18 MS17 MSs8 MS6 MS5 MS3 ree Wwe No2o To Part B TIT J + L . cad Unitenmny greet corollas —eithe g ored corollas VICM LVIVITGS 112 BRIT.ORG/SIDA 21(1) ty~ RI Co WG Won ~ as 4 > oe OER Gt Gist DS ONT na pei al red Sas cad week al pel sa nt ugh sa bho Con Fic. 4 (Part B). data can be interpreted either way—two morphologically divergent species potentially hybridizing in overlapping zones, resulting in some morphological intergradation, or two diverging subspecies with morphological intergradation in zones of range overlap. The former interpretation could result from applica- tion of a quantitative or phenetic species concept, in which species are distin- guished based on differences in means of variables. Although commonly KRINGS AND QIU- 113 ABLE 2. Descriptive summary statistics for continuous floral characters of the uniformly colored corolla group (UCCG) and the multi-colored corolla group (MCCG). Mean sepal length Mean corolla Mean ratio lobe length (corolla lobe length: sepal length) Uniformly colored — 3.21 (SD=0.76) 7.68 (SD=1.47) 2.48 (SD=0.61) Multi-colored 3.08 (SD=0.74) 5.59 (SD=1.73) 1.83 (SD=0.44) | +|, if, | | { {| Taste 3. ANOVA results for floral characte and the multi-colored corolla group. in both cases, the null hypothesis of no eon dene is rejected. Mean corolla lobe length Source of Variation S$ df MS F P-value F crit Between Groups 153.1516 ] 153.1516 61.08722 1.15E-12 3.908255 Within Groups 353.5008 141 2.507098 Total 506.6524 142 Ratio of mean corolla lobe length: mean sepal length Source of Variation SS df MS F P-value F crit Between Groups 14.78613 1 14.78613 50.19068 6.1E-11 3.908255 Within Groups 41,53849 141 0.294599 Total 56.32462 142 applied, the underlying theory of this concept remains unclear (see Luckow 1995). In contrast, application of a phylogenetic species concept suggests two infraspe- cific taxa—populations exhibiting high frequencies of unique traits that may become ‘fixed’ in the future (Nixon & Wheeler 1990). However, a cladistic analy- sis that includes other congenerics is necessary to test this hypothesis (Nixon & Wheeler 1990). In the absence of additional data, we cautiously choose to follow (Drapalik 1969) in recognizing a single species until additional evidence is available. Interestingly, the overall biogeographic pattern exhibited by the two Gonolobus entities has been observed for other taxa (including fish!) in the Southeast (see Avise 1994) and may be the result of past environmental shifts such as the oceanic incursions and retreats that define today’s southeastern Coastal Plain (Sorrie & Weakley 2001). To further elucidate both the question of rank and evolutionary history of the complex, we are conducting genetic analyses of southeastern Gonolobus populations using molecular markers and integrating phylogenetic data from Caribbean Gonolobus species—some of which have been suggested to be nearest relatives of the southeastern taxa (Scheele 1848). 114 BRIT.ORG/SIDA 21(1) UCCG MCCG 40 20 35 1) Pubescent 11 Pubescent 30 |mGlabrous | 15 ; Oo Glabrous 25 ee 20 » 10 15 10 5 | 5 as | me h eka | EE IE ex A 201-4 401-6 6.01-8 8.01-10 >10.01 B 201-4 401-6 601-8 8.01-10 >10.01 UCCG MCCG 26 20 24 18 | 22 11 Pubescer | (i Pubescet 20 Glabrous 16 "® Glabrous 18 1 : 14 | 16 14 im . e 12 1 10 | 10 Ae ah is 8 1 é 6 : : | 4 4 = : 4 | 5 i z = 2? { mu 0 ma l| al ie a_ 8 a lt : 1.001- 1.501- 2.001- 2.501- 3,001- >3501 © ll - - - - - > 3 ; ; ; ; 1.001- 1.501-2 2.001- 2.501-3 3.001- >3.501 Cc 3 3.5 D 1.5 2.5 3.5 . < L: rq £] Fig. 5. PIeQuenicy MotUgranls tur W {RASC RE 2 f Ila InAho } +} my Rte Nn. AA ,Yy L Ct ye eg 1 ry \ character state ranges. Should workers choose to formally recognize the two entities, we provide a brief discussion regarding available names. For more detail regarding the par- ticularly complicated history and nomenclature involved, readers are urged to see Reveal & Barrie (1992) and Krings (2001). In choosing a name for the taxon with uniformly colored petals, past names published for southeastern Gonolobus taxa were examined. Walter’s type of Gonolobus gonocarpos (Walter 687, BM!) contains only leaves and roots. In addition, no mention regarding the pubes- cence of the corolla is made in his protologue (Walter 1788). Thus we cannot be certain to which species the plants that Walter based his description on be- long. However, due to the rare occurrence of uniformly colored species in the Carolinas, we propose that his type be considered to belong to the multi-color group. Types for Gonolobus macrophyllus Michx. and Gonolobus laevis A. Gray, non Michx. are relegated to the multi-color group for the same reason. The KRINGS AND QIU- } 115 protologue of Gonolol latus Scheele (1848) is based on the only eligible type specimen collected west of the Mississippi (Lindheimers.n.) and notes gla- brous adaxial corolla lobes. Thus, the correct name for the uniformly green- flowered taxon should be based on Gonolobus granulatus Scheele and the name for the multi-color flowered taxon should be Gonolobus suberosus (L.) R.Br. Al- ternatively, if the two entities are recognized at the infraspecific level, the cor- rect species name is Gonolobus suberosus (L.) R.Br. ACKNOWLEDGMENTS We thank the following institutions and their curators for assistance with speci- men inquiries, providing photographs of material, and/or providing specimen loans: BM, BRIT, DUKE, FLAS, GA, KY, LL, MISS, MO, NCU, NO, OK, P TENN, TEX, UNA, USCH, USF WILLI. Helpful methodological and nomenclatural com- ments were provided by Tom Wentworth and Paul Fantz, respectively. David Thomas provided PAUP support. Eric Sundell and an reviewer pro- vided thoughtful comments on a previous version of the manuscript. REFERENCES Avise, J.C.1994, Molecular markers, natural history,and evolution.Chapman and Hall, New York. Drapaik, D.J.1969.A biosystematic study of the genus Matelea in the southeastern United States. Ph.D. Dissertation, University of North Carolina, Chapel Hill. Ewan, J. 1939. Bibliographical miscellany. Ill. Publication dates of Asa Gray’s “Botanical Con- tributions.” Amer. Midl. Naturalist 22:218-222. Fernato, M.L. 1950. Gray’s Manual of Botany. American Book Company, Boston Forster, P.l. 1991. The correct publication dates for some genera and species of Asclepiadaceae described by Robert Brown. Asklepios 52:78-79. Gieason, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United States and adjacent Canada. New York Botanical Garden, Bronx. Houmaren, P.K., N.H. Houmaren, and L.C. Barnett. 1990. Index Herbariorum.New York Botanical Garden, Bronx. Krinas, A. 2001. Neotypification of Enslenia albida and a new name in Ampelamus for Cynanchum laeve (Apocynaceae, Asclepiadoideae). Sida 19:925-929. Liede, S. 1996. Sarcostemma (Asclepiadaceae) — a controversial generic circumscription reconsidered: Morphological evidence. Syst. Bot. 21:31-44. Lieve, S. 1997. American Cynanchum (Asclepiadaceae)—A preliminary infrageneric classi- fication. Novon 7:172-181. Liebe, S.and A. TAuser. 2000. Sarcostemma R.Br. (Apocynaceae—Asclepiadoideae)—a con- troversial generic circumscription reconsidered: evidence from trnL-F spacers. Plant Syst. Evol. 225:133-140. Luckow, M. 1995. Species concepts: assumptions, methods, and applications. Syst. Bot. 20:589-605. 116 BRIT.ORG/SIDA 21(1) Masperiey, J. 1985. Jupiter Botanicus—Robert Brown of the British Museum. J. Cramer, Braunschweig. Nixon, K.C. and Q.D. WHeeLer. 1990. An amplification of the phylogenetic species concept. Cladistics 6:211-223. Perry, M.L. 1938. Gonolobus within the Gray’s manual range. Rhodora 40:281-287. Raprorb, A.E., H.E. AHtes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. University of North Carolina Press, Chapel Hill. Reveat,J.L.and FR. Barrie. 1992. Matelea suberosa (L.) Shinners (Asclepiadaceae)—once again. Bartonia 57:36-38. Rosarti, TJ. 1989.The genera of suborder Apocynineae (Apocynaceae and Asclepiadaceae) in the southeastern United States. J. Arnold Arbor. 70:443-514. ScHeeLe, A. 1848. Beitrage zur flor von Texas. Linnaea 21:747-768. SHinners, LH. 1950. The species of Matelea (including Gonolobus) in North Central Texas (Asclepiadaceae). Field & Lab. 18:73-78. SHinneRS, L.H. 1964. Texas Asclepiadaceae other than Asclepias. Sida 1:358-367. Smait, J.K. 1933. Manual of the southeastern flora. University of North Carolina Press, Chapel Hill, Sorrie, B.A.and A.S. Weaktey. 2001.Coastal Plain vascular plant endemics: Phytogeographic patterns. Castanea 66:50-82. Starteu, FA. and R.S. Cowan. 1976. Taxonomic literature:a selective guide to botanical pub- lications and collections with dates, commentaries and types, Vol. |: A-G. Bohn, Scheltema, and Holkema, Utrecht. Stearn, W.T, 1960. An introduction to Robert Brown's “Prodromus florae novae hollandiae.” Pp.V-LII in fascimile edition, Weinheim. Stevens, W.D. 2001. Asclepiadaceae. Monogr. Syst. Bot. Missouri Bot. Gard. 85:234-270. SuNDELL, E.1981.The New World species of Cynanchum L.subgenus Mellichampia (A. Gray ex S.Wats.) Woods. (Asclepiadaceae). Evolutionary Monogr. 5:1-62. Sworrorb, D.L. 2002. PAUP 4.0510. Sinauer Associates, Sunderland. Water, T. 1788. Flora Caroliniana. Murray Printing Co. (Photolithographers), Cambridge. Woopson, R.E. 1941. The North American Asclepiadaceae. Ann. Missouri Bot. Gard. 28: 193-244. POLYPHYLY OF THE GENUS ECHITES (APOCYNACEAE: APOCYNOIDEAE: ECHITEAE): EVIDENCE BASED ON A MORPHOLOGICAL CLADISTIC ANALYSIS Justin K.Williams i tant oloigical Sciences Sam n State University Huntsvil Hs ae 77341-2116, USA. ABSTRACT A cladistic analysis was penlonmied: to test bane a of Echites (Apocynaceae: Apocynoideae: Echiteae). For the analysis 40 | characters were coded for 421 ingroup taxa (22 genera) and three outgroup species (two nee TI 1€ circumscribed is polyphyletic. However, sees sel ad within the original descriptions of the two subgenera de- scribed in Echites form Key Worps: Echites, Prestonia, Thenardia, and Thevetia Apocynaceae, cladistics, morphology, Neo-tropics studies (Struwe et a aoe Bade et al. 1996; pROLRISICE and Als bert, 2001). Allof the genera of the Apocy in Nerium), except Adenium. Colleters (4-6 & 11). Thomas and Dave (1991) provided a discussion of the systematic pce tions of colleters in the Apocynaceae that will not be repeated here. Endress et al. (199 et al. (1998) utilized calycine colleters in their studies (character 12 here). 1 have expanded the use of colleters by including the presence or absence of colleters on other parts of the plant. Character 4, colleters around the stem; Character 5, colleters present on the apex of the leaf petiole of the upper leat surface, a ee ars obec conve —o as it is shared by members of the ‘ ‘Mandevilla” clade and the y species of Mandevilla studied te This character is a ene which unites M. subsagittata and M. hirsuta Domatia (7). Domatia are only present in two of the genera studied here ee id and Forsteronia. ae character appears spikes oa haven arisen in & 9). Distinctness of the secondary venation of leaves a tertiary venation of leaves nas not oa utili ae ina mo Ee al analysis ORE ation in the field coupled with herbarium venation. The lack of secondary veins is a character uniting Thevetia ahouai with C pena The lack of tertiary veins is a character that unites shenayats and Echites subg. Pseudechites. C ze (10). The majority of taxa in the Apocynoideae have sepals 1-3 mm long, a few have sepals as ie 5-15 mm. Overall this character is es variable with ae sepals occurring Ho f, ft LO unify ALLOW OL t eS randomly ial the t 5 I PP cies of Presto eran (12). With the exception of Parsonsia (valvate), dextrorse aestivation is present in all of the taxa in the Apocynoideae included in this study. Aestivation type is one of the few syna- pomorphies that distinguishes the Apocynoideae from the Rauvolfioideae (sinistrorse aestivation). Corolla shape ,color and size(13, 14, 18). Corolla shape was utilized by Endress et al. (1996), color and tube size are new characters. Most of the taxa presented in this work have yellow or white corollas. However, some have maroon corollas. Color was used because Laubertia and Prestonia portobellensis possess maroon corollas. Consequently the character was utilized to test if the species Sees ae testing the seas ee) of Prestonia. In addition, the two subgenera of Echites have rolla colors. Int | as utilized to test the monophyly of Echites. Corolla tube size is a as an augment to the as ity that exists in corolla shape. Utilization of tube size ph 7 hat although both subg. of Echites have salverform corollas, there exists a con- id be rice in Ane ee of the corollas Corolla witl d corona | I si corolla with infrastaminal appendages (15, 16, 17) These ie ieee were utilized a discussed by Endress et al. (1996). ‘ilament length (19). An examination of the filaments of the taxa in this analysis indicates that filament length appears to be Seas y corre aes with generic Sanus Short filaments are typical of the taxa in the subg. E 1), while medium length fi with the pene clade (Fig. D lg one filaments a are only] present in two noe ane 1s oc of Ty. : L * ‘ 1 nthers from ribs, stamen exposure, and anther dehiscence (20, 21, 23) Anther ribs are only present in Thevetia and a bera and are used mainly to establish the monophyly of the outgroup. Anther exy and dehiscence were utilized and discussed in Endress et al. (1996). WILLIAMS, EVIDENCE FOR A NEW NEOTROPICAL GENUS OF APOCYNACEAE 129 pical appendages on anthers (22). It was presumed before the analysis was conducted that this character was highly convergent. However, it was included as a reference for testing the mono- phyly of Tinti oe which has two species with and one species without elongate apical an- ther appendag Anther | es anther- Ca head Sac ine pistil head type (24, 25, 26). At least five different types of anthers and pistil heads hibited in the Apocynaceae. The different anther and pistil head types have been discussed in beara (1930) and Fallen (1986). The important traits that characterize the different anther types are the connective, theca positioning and fertility and the base of the anther body. The union of the anthers and the pistil head is a synapomorphy that unifies the Apocynoideae. Nectary (27). povera types of nectaries are ies by the Apocynaceae. Three different types I lintoa cup, a ane nectaries fused into a cup type two. Type oe refers to the nectaries of Thevetia and Cerbera, that are twice as large and twice as mace? as Biss found in ae pogynl eae hology, infl I hing (28, 29, 30). The struc- g tare of the inflorescence hasnot been used in hological cladisti lysis of the Apocynaceae 1 account of the inflorescence ee inthe A Apocynacede that will not be repeated eke Within the taxa examined only the W Apocyneae (Apocynum) have terminal inflorescences, with all of the Echiteae poses: an axillary a lorescence. Inflores- cence branching is one of the oe separating subg. Echites (not branched) from subg. Pseudechites (branched). Fruit type, follicle orientation, follicles moniliform, follicle color, fruit dehiscence, fruit texture (31-36). Of the above characters, only fruit dehiscence has been utilized in a cladistic study (Endress et al. 1996). Fruits have been an underutilized resource in the systematics of the Apocynaceae. This is mainly due to the paucity of fruiting herbarium specimens. Collecting trips were made by the au- thor specifically in the latter part of the flowering season, for the hes of collecting fruits. From these observations, a pattern cee Many of the taxa with presumed relationships had similar frui es. Characters observed were the union of the follicles, f ee at apex, follicles spreading, or fused eee The fusion of ve follicles, is a useful character in distinguishing species within genera (e.g. Echites), but overall the cladistic analysis indicated that follicle union is a convergent character, with spreading and fused follicles OeCUrEINE ErOuE TOU the eDOCS meLlene. In addition, follicle texture was noted. Some follicles were membranous while others were firm and woody. This character was useful in ees Echites as polyp g. Echites with woody follicles and y Sup bg. Pseudechites with herbaceous follicles). Lastly it it was noticed that some taxa had straight fol- ly in the “Prestonia” clade. uit color was used to test the mon opny ly ot The vetia. ass character isasy ane uniting T. show and Cerbera, indicating that T cumscribed is paraphylet s with coma (37). This character was utilized by aos et al. (1996) and by ace and fies Cones subsequently discussed by them Pollen apertures, pollen exine a pollen shape, pollen size (39-42). Pollen ano and e pattern were utilized in Endress et ‘i “99 6). a Rauvolfioideae and the Apocynoideae are ioe by the apertures of ie pollen, with tri-porate pollen asa aporphy uniting the Apocynoideae. In addition, the pollen of taxa in the Apocynoideae is consistently smooth vs. the Rauvolnoidede which has various exine patterning. Pollen shape, also : Ips to ec the f e Apocynoideae. In general, the Rauvolfioideae have triangular-rounded pol- len vs. the merits that are consistently spherical. Pollen size was useful for determining in- tergeneric and intrageneric relationships. For instance, pollen size supports Echites as polyphyletic (subg. Echites, 45-60 um vs. subg. Pseudechites, 23-35 um). In addition within Mandevilla, pollen size oe M. hirsuta and 7 subsagittata to be closely related. 130 BRIT.ORG/SIDA 21(1) ACKNOWLEDGMENTS [thank the curators of the following herbaria for allowing me to study and bor- row their specimens: BM, BRIT, CHAPA, F FLAS, G,GH, K, MA, METPEC, MEXU, MO, NY, P, SHST, TAMU, TEX, US, WIS. l also thank B.B. Simpson for providing comments on the running and interpretation of the cladistic analysis, and most especially, I thank Mary Endress for her patience, support and superbly accurate and critical review of the article as well as contribution to the results section. REFERENCES Enpress, M.E., B. SENNBLAD, S. NILSSON, L. CiveyReL, M. CHASE, S. HuYSMANS, E. GRaAFsTROM, and B. Bremer, 1996. A phylogenetic analysis of Apocynaceae s. str. and some related taxa in Gentianales: a multidisciplinary approach. Opera Bot. Belg. 7:59-102. Enoress, M.E. and P. Bruyns. 2000. A revised classification of the Apocynaceae s.|. Bot. Rev. 66:1-56. FALLEN, M. 1986. Floral structure in the Apocyna Morphological, functional and evo- lutionary aspects. Bot. Jahrb. Syst. 106:245-286. Gentry, A. 1983. A new combination for a problematic Central American Apocynaceae. Ann. Missouri Bot. Gard. 70:205-206. Heaos, M. J. and PJ. be Lance. 1998. Parsonsia praeruptis (Apocynaceae): a new threatened, ultramafic endemic from North Cape, New Zealand. New Zealand J. Bot. 37:1-6. KITCHING, I.J., PL. Forey, C.J. Humpuries, and D.M. Wittiams. 1998. Cladistics. Oxford University Press, Maboison, W.P. and D.R. Maooison, 1992. MacClade: Analysis of phylogeny and character evolution. Version 3.0. Sinauer Associates, Sunderland, Massachusetts Monacuino, J. 1959. A new Echites from Mexico. Bull. Torrey Bot. Club 86:245, Morates, J.F. 1996. Una nueva especie de Tintinnabularia (Apocynaceae). Novon 6: 392-394 Morates, J.F. 1997. A reevaluation of Echites and Prestonia sect. Coalitae (Apocynaceae). Brittonia 49:328-336. Morates, J.F. and J.K. Wituams. 2004. Allotoonia, a new neotropical genus of Apocynaceae based on a subgeneric segregate of Echites. Sida: 21:133-158. PoraieTer, K. and V. Acsert. 2001. Phylogenetic relationships within Apocynaceae s.. based on trmL intron and trnL-F spacer sequences and propagule characters. Ann. Missouri Bot. Gard. 88:523-549. SennBLAD, B.and B. Bremer. 1996. The familial and subfamilial relationships of Apocynaceae and Asclepiadaceae evaluated with rbcL data. Pl. Syst. Evol. 202:153-175. SENNBLAD, B., M.E. Enoress, and B. Bremer. 1998. Morphology and molecular data in phyloge- netic fraternity: the tribe Wrightieae (Apocynaceae) revisited. Amer. J. Bot. 85: 1143-1158. Struwe, L., V.A. ALBerT, and B. Bremer. 1994. Cladistics and family level classification of the Gentianales. Cladistics 10:175-206. WILLIAMS, EVIDENCE FOR A NEW NEOTROPICAL GENUS OF APOCYNACEAE 131 Sworrorb, D.L.1993.PAUP:Phylogenetic Analysis Using Parsimony.Version 3.1.1.Computer program distributed by the Illinois Natural History Survey. Champaign, Illinois THomas, V.and V. Dave. 1991.Comparative and phylogenetic significance of the colleters in the family Apocynaceae. Feddes Repertorium 102:177-182. WiLuIAMS, J.B. 1996. Parsonsia.In:Flora of Australia. Volume 28, Gentianales. Melbourne:CSIRO Australia. Pp. 154-189. WILLIAMS, J.K. 1996. The Mexican genera of the Apocynaceae (sensu A.DC.), with key and additional taxonomic notes. Sida 17:197-214. Wituiams, J.K. 1998. A revision of Thenardia H.B.K. (Apocynaceae, Apocynoideae). Lundellia 1:78-94. WILLIAMS, J.K. 2002a. A further evaluation of Echites sect. Yucatanense (Apocynaceae) with additional notes on the genus. Brittonia 54:310-317. Wituiams, J.K.2002b. Thoreauea (Apocynaceae: Apocynoideae), a new genus from Oaxaca, Mexico. Lundellia 5:47-58. Woooson, R.E., Jk. 1930. Studies in the Apocynaceae.I.A critical study of the Apocynoideae (with special references to the genus Apocynum). Ann. Missouri Bot. Gard. 17:1-212. Woopson, R.E., Jr. 1931. New or otherwise noteworthy Apocynaceae of tropical America. Ann. Missouri Bot. Gard. 18:555. Woopson, R.E., Jk. 1935. Observations on the inflorescence of Apocynaceae with special reference to the American genera of Echitoideae. Ann. Missouri Bot. Gard. 22:1-40. Woooson, R.E., Jk. 1936. Studies in the Apocynaceae IV.The American genera of Echitoideae (concluded). Ann. Missouri Bot. Gard. 23:169-438. Woooson, R.E., Jr. 1938. Apocynaceae. In: N.L. Britton et al. North Amer. FI. 29:103-192. Woooson, R.E., Jk. 1960. Loganiaceae, Apocynaceae, and Asclepiadaceae. In miscellanea taxonomica. Il. Ann. Missouri Bot. Gard. 47:79. BRIT.ORG/SIDA 21(1) BOOK NOTICE Douc.as J. Futuyma, H. BRADLEY SHAFFER, and DANIEL SIMBERLOFF (eds). 2003. An- nual Review of Ecology, Evolution, and Systematics: Volume 34, 2003. ISBN 0-8243-1434-4, hbk; ISSN 1543-592). Annual Reviews Inc., 4139 El Camino Way, P.O. Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www. AnnualReviews.org, 800-523-8635, 650-493-4400, 650-424-0910 fax). $160.00 (USA), $165.00 (Int’L), 716 pp., 6" x 9". Volume 34 of Annual Review of Ecology, Evolution, and Syst ic i 24 papers: Effects of introduced bees on Avian sexual dichromatism in relation to Be ee oa and Hts Paleobiogeography: the relevance of fossils to biogeography The ecology of bird introductions The effects of genetic and geographic structure on neutral variation Data, models, and decisions in U.S. Marine fisheries management lessons for ecologists Partitioning of time as an ecological resou Performance comparisons of co-occurring native and alien invasive pl ants: implications for conservation and restoration enetic variation in rare and common plants The ecology and evolution of insect baculoviruses rn, process, scale, and synthesis Latitudinal gradients of biodiversity: patte Recent advances in the (molecular) phylogeny of vertebrates The role of reinforcement in speciation: theory and ae Extra-pair paternity in birds: causes, cone and conflict Species-level paraphyly and polyphy y, causes, and consequences, with insights f animal mitochondrial DNA Protective ant as interactions as model systems in ecological a | y research Functional matrix: a -eptual framework for predicting are e ane offects on ecosystem processes Effects of habitat fragmentation on biodiversity Social organization and parasite risk in mammals: integrating theory and empirical studies he community-level consequences of seed dispersal patterns The ecology and ev aaah seed as aca a theoretical perspective Analysis of rates of Development i the g sedation of evolutionary change within insect species Flexibility and specificity in coral-algal symbiosis: diversity. ecology, and biogeography of Symbiodinium SIDA 21(1): 132. 2004 ALLOTOONIA, A NEW NEOTROPICAL GENUS OF APOCYNACEAE BASED ON A SUBGENERIC SEGREGALE OF EGRITES J.Francisco Morales Justin K.Williams Instituto Nacional de Bi panera NTE) gees Poaceae Sciences Apto 22-3100, Santo Doming Sam Houston State Univ COSTA RICA Huntsville, ss 77341-21 i is ABSTRACT Alloto ia (Ay ) ne otropl¢ i 1f Echitesisd ibed. In addition oO (ead ©) a ae to species descriptions, distribution maps, ill | ined aa Pe pi OVIadcag for all species. Key Worps. Apocynaceae, Apocynoideae, Allotoonia, Echiteae, Echites, Neotropics, Mesoamerica, Mexico RESUMEN Allotoonia (Apocynaceae), un nuevo género neotropical segregado de ites es descrito. Se brinda una clave para todos los taxa, asi como descripciones, mapas de dicmibaciéa ilustraciones y especimenes examinados para todas las especies. In his monumental revision of Echites P Browne, Woodson (1936) established two sub : Echites (Euechites) and subg. Pseudechites Woodson. Subgenus Pse udeenaies included the species E. turbinata Woodson and E. tuxtlensis Stand and was distinguished from subgenus Echites by its smaller corollas (1.25-2.5 cm long vs. 5-8 cm long), lobes narrowly oblong and reflexed (vs. obliquely obovate and spreading), and its many flowered lax inflorescence (vs. compact and few flowered). A third species, E. parviflora Sessé & Moc. was subsequently added to subg. Pseudechites by Woodson (1938). Echites went relatively unnoticed for close to 60 years, until Morales (1997) provided a synopsis of the genus. In his work, Morales described an additional species in subg. Pseudechites (E. puntarenensisJ. F Morales) as well as presented an argument for maintaining Prestonia agglutinata Jacq.) Woodson in Echites, resurrecting the older name Echites agglutinata. The morphological cladistic analysis of Echitess.l.and other genera of sub- family Apocynoideae presented in Williams (2004) indicates that Echites sensu Woodson is polyphyletic. Subgenus Pseudechites shows a stronger affinity to other genera of Echiteae sensu Endress and Bruyns (2000) (e.g., Laubertia, Prestonia, Thenardia) than it does to subg. Echites. Consequently, the authors have chosen to recognize the species in subg. Pseudechites as a distinct genus (Allotoonia described below) separate from Echites s.s. Table | documents the SIDA 21(1): 133-158. 2004 134 BRIT.ORG/SIDA 21(1) differences between Allotoonia, Echites, and other selected genera of New World Echiteae. MORPHOLOGICAL CHARACTERS Leaves.—The leaves are opposite and entire. Four of the five species have leaves with obscure secondary venation making them readily discernible from Echites One species, however, A. agglutinata, has conspicuous secondary veins. Allotoonia also shares with others neotropical Apocynoideae, intra-petiolar colleters. All species have glabrous leaves. The leaves of A. turbinata are often two to three times longer and broader than those of the other four species. Field studies have shown that these characters are uniform and consistent within a population. Inflorescence.—Allotoonia is readily distinguished from Echites by the in- florescence structure. The inflorescence of Allotoonia is a helicoid cyme once, twice to occasionally thrice branched, while in Echites the inflorescence is a modified dichasium, sometimes appearing racemose, and occasionally reduced to just one flower. The bracts are always inconspicuous and scarious. Flowers.—The calyx is regular, pentamerous, and with a solitary opposite colleter on the adaxial surface of each sepal, as found in many others members of the Echiteae (e.g., Prestonia, Thenardia). The sepals are narrowly ovate and 1-3.5 mm long. The shape and dimensions of the sepals for each species corre- sponds to the shape and dimensions of the bracts subtending the pedicel. The corolla is salverform, the tube glabrous without. The aestivation is dextrorse as are all other neotropical members of the Apocynoideae. The co- rolla lobes are very narrowly elliptic to almost filiform, acuminate to long- acuminate at the apex and conspicuously wavy and twisted distally. Allotoonia isthe only neotropical Apocynaceae genus with wavy and twisted filiform lobes at the anthesis (Figs. 1-2). In Echites, the corolla lobes are obovate. The stamens are included and the anthers are strongly attached at two lev- els to the spool shaped style head (Fig. 3). The anther shape differs between Allotoonia and Echites (Fig. 4). In Allotoonia an anther is more or less continu- ous (except for a slight dorsal protuberance), without a marginal rib. In Echites the anther has a conspicuous marginal rib that is extended from the basal pro- jections along the entire length up to the apex (Fig. 1 B). Fruit.—The fruits of Allotoonia are composed of two herbaceous, pendu- lous, follicles (Fig. 5) developing from two carpels postgenitally united at the apex, sometimes free at maturity. The follicles are continuous to obscurely moniliform. The fruits of Echites are woody and never moniliform. Seeds.—The seeds of Allotoonia are glabrous, 6-20 per follicle, and rostrate, with the beak slender, narrowly and conspicuously elongate (Fig. 6), while in Echites the seeds are inconspicuous to moderately rostrate, with the beak short ALLOTOONIA GENUS OF APOCYNACEAE 135 (sometimes somewhat elongate in E. yucatanensis) and gradually acuminate toward the micropylar end (Fig. 7). Allotoonia J.F Morales = . naa gen. nov. Type. Allotoonia agglutinata Jacq.) J.-E Morales & J.K. Will Hetba volubilis, BORSIeFenieae, Laubertiae, Parsoniae, Prestonieae, Thenardiae et Thoreauiae affinis: ms | ek - ia . ] 12 7 ] 5 OR ty | membranacea, eg] rt I lobis filiformis, revolutis. tub PI licul 1; +3] } < iC cole regulatis 5-partita inserta Lactescent lianas, usually herbaceous, more rarely suffruticose, not woody. Stems terete to subterete, somewhat flattened at the nodes, glabrous, not lenticellate, with inconspicuous intrapetiolar colleters, the latex usually wa- tery. Leaves opposite, entire, membranaceous, glabrous or glabrate, the second- ary and tertiary veins usually inconspicuous or obscure, petiole with many, minute and filiform colleters in the axils Inflorescence a helicoid cyme, axil- lary, glabrous, many-flowered, usually long-pedunculate, bracts scarious, in- conspicuous. Flowers pentamerous, the sepals free or very slightly imbricate basally, scarious, bearing a solitary, episepalous entire to variously laciniate colleter within; corolla salverform, glabrous to very minutely and inconspicu- ously puberulent without, without corona lobes or annular corona, the tube usually longitudinally grooved, with an infrastaminal ribs present below each filament, the limb 5-parted, actinomorphic, dextrorsely convolute, the lobes very narrowly elliptic to almost filiform, acuminate, twisted and wavy distally; stamens 5, included, inserted in the upper part of the corolla tube near the mouth, anthers connivent and agglutinated to the style head, attached in two points to the style head, glabrous to glabrate, thecae with the base sagittate, 2- auriculate, auricles short, but conspicuous, short-acuminate, filaments short, puberulent to pilose, carpels 2, united at the apex by a common stylar shaft surmounted by the fusiform, spool-shaped style head, ovary glabrous, ovules numerous, multi-seriate, nectarines five, separated and distinct, rarely slightly connate basally. Follicles continuous and terete to subterete, or obscurely moniliform, slender and smooth, usually glabrous; seeds comose at the micropilar end, conspicuously rostrate, minutely rugose, numerous. This genus comprises five species distributed from southern Mexico to northern Panama, and the West Indies. Allotoonia is named after Dr. Anthony Leeuwenberg, from the Wageningen University (WAG), The Netherlands, in recognition of his work in the Apocynaceae. The genus Allotoonia is in the subfamily Apocynoideae, tribe Echiteae (En- dress & Bruyns 2000), as evidenced by the following characters: anthers with the thecae connivent, agglutinated and strongly attached at two levels to the spool shaped style head. Taste 1. Morphological characters in Aliotoonia and seiected genera of the Echiteae S Genera Allotoonia Echites Forsteronia Laubertia Prestonia Thenardia Thoreauea Morphological Characters Latex Clear White White Clear White Clear White Secondary veins Usually Conspicuous Conspicuous, Conspicuous Conspicuous Usually Conspicuous INCONSpiCuOUs rarely inconspicuous inconspicuous Inflorescence Helicoid cyme Modified Thyrsiform Scorpioid cyme, Racemose, Subumbellate Trichotomously dichasium sometimes subracemose, cyme reduced and umbellate, Subumbellate appearing simple subumbellate, cyme orumbelliform = corymbose or subcorymbose Bracts Scarious Scarious Scarious Scarious Foliaceous to Scarious Scarious scarious Sepals Scarious Scarious Scarious Scarious Foliaceous to Scarious Scarious scarious Sepals Solitary and Solitary and Laterally or evenly Lacking Solitary and Solitary and Solitary and colleters opposite opposite disposed, rarely opposite opposite opposite lackin Corolla Salverform Infundibuliform Salverform to rotate Salverform Salverform to Rotate Urceolate rarely to campanulate infundibuliform 2 Corona Absent Absent Absent Present, Present, Absent Present, a continuous continuous dissected interior & around around to corolla mout S the corolla the corolla = mouth mouth = Genera Allotoonia Echites Forsteronia Laubertia Prestonia Thenardia Thoreauea Morphological Characters Lobes Very narrowly Obovate Oblong to Obovate Obovate to Ovate Ovate elliptic to almost lanceolate ovate filiform, acuminate, twisted and wavy distally Stamens Included, Included, Included or Included or Included or Exserted Apically exserted insertion inserted in the inserted about variously variously apically upper part,near mid exserted exserted exserted the mouth corolla tube Filaments Free Free Free orconnate Free Free or Partially Partially connate connate Disk or 5, usually 5, but usually Usually (3-) 5- 5,separated to Annular, entire to 5, separated 5, separated nectaries separated, rarely connate lobed irregularly 5-lobed or slightly connate _ basally basally connate in 5 divided basally nectaries Follicles Slender, Slender, der, Slender Divaricate, Slender, Unknown continuous and — continuous divaricate or, moniliform, rarely fusiform, moniliform terete, or and terete arallel, continuous _ continuous or obscurely continuous rarely moniliform moniliform to moniliform Coma seeds Conspicuously Inconspicuously Truncate, sessile Truncate,sessile Truncate Sessile Unknown rostrate to moderately rostrat VINOQOLOTIV IVIDVNADOdY 40 SNNID ZEL BRIT.ORG/SIDA 21(1) Fic. 1 Aliant . data A. Hahi er | AA } ( 8242, INB). B. Close-up of the flowers, showing twisted and elongate corolla lobes (Morales 8242, INB). KEY TO THE SPECIES OF ALLOTOONIA . Corolla lobes (17-)20-27 mm long; corolla tube constricted below the insertion of the stamens; montane rain forest and related disturbed areas between 1500- 2350 m pie A.turbinata |. Corolla lobes 4-14(-16) mm long; corolla tube straight or slightly constricted be- low the insertion of the stamens; tropical deciduo orest, premontane moist for- est, gallery forest, and related disturbed areas, bewteen 0-1 200(-1400) m. 2. Secondary venation of leaves raised and conspicuous, tertiary venation of leaves obscure; corolla lobes glabrous; plants of the West Indies A. agglutinata 2. Secondary venation inconspicuous, not raised, tertiary venation usually incon- spicuous; corolla lobes pubescent or glabrous; plants of Mexico and Central America 3. Corolla lobes shorter than tube, glabrous; bud pointed; tube 7-9 mm long, ca. 1 mm diam., constricted between middle to base of lobes A. tuxtlensis 3. Corolla lobes longer than tube, hirsute or glabrous; bud round; tube 4-6(-7) mm long, ca.2 mm diam., straight, not constricted. 4. Corolla orange to yellow-orange, the lobes hirsute; pedicels 2-9 mm; an- thers 3.5-4m A. parviflora 4, Corolla white, creamish white to pinkish white, the lobes glabrous to glabrate; pedicels 11-24 mm; anthers (5-)6-8 mm A. caudata ALLOTOONIA GENUS OF APOCYNACEAE 139 Fic.2. All j iflora. A. Habit and infl (Morales & Abarca 8810, INB). B. Close-up of the flowers, show- ( rS scetan A | + ll lanl ThA iy PAL OoOoT1n , INB) 1. Allotoonia ied ae Jacq.) J.F Morales &J.K. Williams, comb. nov. (Figs. 8, . BASIONYM: Echites oe inata Jacq., Enum. Syst. Pl. 13. 1760. Anechites ee eh Miers, Apocyn. S. Amer. 236. 1878. Prestonia agglutinata Jacq.) oodson, Ann. Missouri Bot. Gard. 18:552. 1931. Type. HAITI: Cap. Francais, Select Stirp. Am. Hist. 1763. (LECTOTYPE, here designated). Echites circinalis Sw., Prodr. 52.1788 Se Don, Gen. Hist. 4:83. 1837. Anechites circinalis (Sw.) Miers, Apocyn. S. Amer. 236. 1878. TyPE: data lacking (fl), Rohr 93 (LECTOTYPE, here designated: C; ISOLECTOTYPE: P- oo Echites sanguinolenta Tussac, Fl. Antill. 95, t. Ll. Haemadictyon nutans (Anders.) A. DC. var. sanguinolenta (Tussac) A. DC., Prodr. 8:426. 1844. Type. HISPANIOLA: not located. Echites circinalis Sw. var. thomasiana A. DC.,, Prodr. 466. 1844. Anechites thomasiana (A. DC.) Miers, Apocyn. S. Amer. 237. 1878. TyPE: ST. THOMAS: exact locality lacking, 1841 (fD, Siedrichsthat 1 240 (HOLOTYPE: W). Echites eties Sessé & Moc., Naturaleza (Mexico City), ser. 2, 2, app. 45. 1893. TypE: PUERTO RICO r Castellum del Morro, Oct., year lacking, Sessé & Mocino 5075 (LECTOTYPE, here eee MA). Suffruticose to herbaceous liana, stem terete to subterete, glabrous to glabrate. Leaves: petioles 5-20 mm long, with minute pectinate colleters in the axils or sometimes eglandular, glabrous, blade 3-13.5 x 2-8.5 cm, membranaceous, el- 140 BRIT.ORG/SIDA 21 — = fs oat3 ae MARRS 28 ’ 2 . cocbeatice: wet, fase? Re wets GIVE TRE e 3 t t 1.5mm 3mm Fic. 3. All ia parviflora style head and anthers, show- | Fic. 4. Allotoonia and Echites anthers, dorsal view. A. ingtl levels of I he spool shaped styl Allotoonia parviflora. (Morales 3139, INB). B. Echites head. (Morales & Abarca 8810, NB umbellata. (Whiteford 8209, INB). aS liptic, narrowly elliptic to narrowly ovate, apex caudate to obtuse, base obtuse to somewhat attenuate basally, glabrous, midvein impressed on both surfaces, the secondary venation conspicuous on both sides. Inflorescence usually longer than the subtending leaves, lax, peduncle 1.5-7 cm, pedicels 2-5 mm, bracts l- mm long, ovate, acute, scarious; sepals 1-1.5 x 0.5-1 mm, ovate, acuminate, gla- brous, glabrate, or minutely and inconspicuously puberulent, colleter some- what laciniate to deeply so; corolla salverform, white, creamish white to cream, glabrous without, tube 5-6 mm long, I-15 mm diam, somewhat inflated at the attachment of the stamens, pubescent within near the mouth, lobes 4-6 « 1-2 mm, glabrous; stamens: filaments ca. 4 mm, ee to psig) pubescent, anthers 4-5 mm, glabrous, auricles ca. 0.2 mm, ovary ca. 2mm, siaprous, style head 1.5-2 mm long, nectaries I-1.5 mm, entire and conspicuously separate. Follicles 11-31(-46) x 0.4-0.8 cm, continuous, smooth and glabrous, usually straight; seeds 6.5-10 x 1.5-2 mm, rugose, the tannish coma 2-4 cm. — ALLOTOONIA GENUS OF APOCYNACEAE 141 Fic. 5. Allotoonia parviflora. Follicles BRIT.ORG/SIDA 21(1) Fic. 6. Allotoonia seeds. A. A util (Axelrod & Axelrod 3091, INB). B. A. turbinata (Gomez-L. et al., 14142, |INB).C.A. tuxtlensis (Arvigo et al. 536, INB).D. A. parviflora (Morales 3161, INB) J ————— A | Faerun eee | lcm Fic. 7. Echites seeds. A. E. bellata (Trejo et al 530, INB). B. £ y ALLOTOO NIA GENUS OF APOCYNACEAE Fic. 8. Allot ta A. Fertile shoot 1mm ter. C. Corolla. D. Disk and ovary (Zano niet al. 47 057, |NB). 144 BRIT.ORG/SIDA 21(1) aw, ae e =A. agglutinata ~ 4 m =A. caudata Fic. 9. Distribution of A. agglutinata and A. caudata. Habitat and distribution.—Dry forest, gallery forest, shrubby vegetation, thickets, and disturbed areas in the West Indies (Hispaniola, Cuba, Puerto Rico, Jamaica, part of Greater Antilles), at elevations of 0-350 m. (Fig. 9). Associated species include Abutilon and Plumeria. Phenology.—Flowering specimens have been collected in January, May-June and September-October. Fruiting collections have been made in January-Feb- ruary and September-October. Local names.—abrazapalo (Dominican Republic); azufaifo, babeiro (Puerto Rico). Allotoonia agglutinata is the only species of the genus with conspicuously raised secondary veins. In addition, it has glabrous corolla lobes that are longer than the tube and it is distributed throughout the West Indies. The corolla bud is contorted, similar to the bud of A. tuxtlensis (Fig. 10). A complete list of synonyms for this species as presented by Woodson (1936) is provided here in order to indicate that an earlier generic name does not exist for any of the described species of Allotoonia. Although the types collection of Echites sanguinolenta Tussac and E. circinalis Sw. var. thomasiana A. DC. were not located, they are included in the synonymy based on their original descrip- tions, which matches the Allotoonia agglutinata concept used here. Within Allotoonia, A. agglutinata has the highest number of synonyms, and is the only — ALLOTOONIA GENUS OF APOCYNACEAE 145 a. 1G. 10. Fl buds of the species of Allotoonia. A.A. parviflora (Williams 96-88, SHST).B. A. turbinata (Breedlove 25058, TEX). C. A. tuxtlensis (Herndndez 488, TEX). species that has been described in multiple genera. None of the genera listed above are legitimate alternatives to Allotoonia. Observations of the Sessé & Mocino collections by the second author indi- cates that the specimen number 5075 is labeled Echites obtusifolia. The species name is written on the back of the herbarium sheet, which perhaps accounts for the specimen being overlooked. Consequently, specimen number 5075 of the Sessé and Mocino herbarium is here designated as the lectotype for Echites obtusifolia. Selected specimens examined. JAMAICA: Data ms Swartz 265 (BM). PUERTO RICO: Patillas, Guardarraya, Cerro Mala Pascua, 13 Oct 1991, Axelrod & Axelrod 3091 (INB, UPRRP); Isla de Desecho, W of Puerto de los Botes, Breckon 4994 (MAPR), csi 5019 (MAPR): Sak G le, Susua forest, 7 Nov 1990, oes . plc 3287 (MAPR, NY); Salinas, Camp Santiago, section Lima, date lack- ing, Ross& N 53 (MO); near Cayey, Pedro Avila, 22 Sep 1895, Sintenis 2457 (S); Quebradillas, 24 Jun 1991, oe & Gereau 10493 (MO); Guanica, 25 Jan 1886, Urban 3565 (BM, P [2 sheets]); Punta Tuna, S of Yabacoa, 11 Oct 1968, Wagner 1690 (INB). DOMINICAN REPUBLIC: Cabo Francés Viejo to Cabrera, 28 Sep 1969, eeu 16152 (INB, NY); La Romana: Rio Cumayasa, road San Pedro de Macori- La Romana, 29 Oct 1992, Zanoni et al. 47051 (INB, JBSD). HAITI: La Hotte, Bras Gauche river, 8 Nov 1924, Ekman ceed Totue Island, Basse-Terre, 28 Oct 1925, Ekman 5131(S, US). TORTOLA: Belmont Beach, 30 Oct 1965, D’Arcy 292" (MO). VIRGIN ISLANDS: Francis Bay area, 6 Jun 1985, Acevedo et al. 902 (BM, NY); ses Bay Quarter, road to Bordeaux, ies 1991, Acevedo & Siaca 3819 (INB, US); Reef Bay, Dittlif point, 20 Jan 1991, Acevedo & Siaca 3974 (s 2. Allotoonia caudata (Woodson) J.F. oe cons nov. (Figs. 1, 11). BASIONYM: Echites puntarenensis J.F Morales, Brittonia 49:332. 1997. nom. nov. Prestonia caudata Woodson, Ann. Missouri Bot. Gard. 47:79. 1960, non Blanco (1837). TYPE: 146 BRIT.ORG/SIDA 21(1) COSTA RICA: Puntarenas: vicinity of Cascajal, 25 km ESE of Puntarenas, 3 Jul 1949 (£D, Holm & Iltis 243 (HOLOTYPE: MO; ISOTYPES: A, CR, G, GH, P) Suffruticose lianas, stem terete to subterete, glabrous. Leaves: petioles 6-19 mm long, with minute pectinate colleters in the axils, glabrous; blade 6-14(-16) x 15-5 cm, membranaceous, elliptic to narrowly elliptic, more rarely narrowly obovate-elliptic, apex acuminate, base obtuse to cuneate, glabrous, midvein irapressed on both surfaces, the ey venation n scarcely impressed or in- conspicuous on both sides, veinlets usually Inflorescence longer than the subtending leaves, lax, peduncle 4.5-l1 cm, n, pedicels 11-24 mm, bracts ca. 1mm long, ovate, acuminate, scarious; sepals 1 x 0.5-1 mm, ovate, acute, gla- brous, the colleter entire to subentire; corolla salverform, white, creamish white, or pinkish white, glabrous without, tube 6-8 mm long, 2-2.5 mm diam, straight, glabrate around the stamens within, lobes 8-15 x 1.5-2 mm, glabrous to glabrate; stamens: filaments less than 0.5 mm, inconspicuous, anthers (5-)6-8 mm, gla- brous, auricles ca. 0.8 mm; ovary 1.5-2 mm, glabrous, style head 1-2.5 mm long; nectaries 0.5-1 mm, entire. Follicles unknown. Distribution.—Dry forest, savannas, and disturbed areas in northern Costa Rica, southern Nicaragua and El Salvador, 0-600(-1000) m. (Fig. 9). Associated species include Curatella, Lonchocarpus, Plumeria, and Stemmadenia. Phenology.—Flowering specimens have been collected in February, April, June-July, September, and November-December. Local names.—bejuco de Veneno (Costa Rica, Guanacaste, Nicoya). Allotoonia caudata is very similar to A. parviflora and usually misidentified in herbarium specimens. However, the former taxon is easily dis- tinguished by its white, creamish white, or pinkish white flowers (vs. orange to yellow-orange), with glabrous or glabrate corolla lobes (vs. densely hirsute). Morales (1997) recognized that Prestonia caudata Woodson should be treated in Echites but a new name was necessary, because the epithet was already oc- cupied by several homonyms (E. caudata L., E.caudata Burmanf.,and E. caudata Blanco), thus, the name FE. puntarenensis was proposed. With the transfer of this taxon to the new genus, the original basionym can be used according to article 58.3 of The Code. Therefore, the respective combination is made and E. aN is reduced to synonymy. d. EL SALVADOR. Santa Ana: Ser ee 3 Sep 1994, Villacorta & Méndez 2181 (B LAGU, MO). NICARAGUA. Chontales: 3 km N of St. Tomas, 6 Apr 1982, Moreno 16081 (MO). Rivas: Isla Omotepe, Balgue, 28 Nov 1982, Moreno ae (MO); Isla Ometepe, Balgue, 18 Jan 1983, Moreno 196564 (BM, MO); Isla de Ometepe, 15 Sep 1983, eek 22179 (MO); Isla Omotepe, Mérida, 11 Feb 1984, Robleto 199 (MO); Omotepe, Volcan Maderas, 18 Jun 1984, Robleto 912 (MO), 16 Jan 1985, Robleto ee ees COSTA RICA. Guanacaste: Pilas de Bejuco, 18 Nov 1994, Estrada & Rodriguez 283 (CR, O): Canas, Rio — 26 Jun 1942, Leén 894 (F); Nicoya, 25 Jan 1985, Poveda & Castro 3926 ne MO); scrub near Nicoya, May 1900, Tonduz 13940 (MO). Puntarenas: Paquera a Playa Organos, 25 Dec 1993, Hammel & Garita 19306 (CR, INB, MO). San José: San Juan de Mata, cabeceras Quebradas Yeguas y Paso Agres, 11 Nov 2001, Morales 8242 (INB, MO). ALLOTOONIA GENUS OF APOCYNACEAE 147 Fic. 11. All j 1 A. Flowering shoot. B. Sepals. colleters, nectaries, and ovary. C. Corolla (Morales 8242. |INB) g I 1 y \ ' ) 148 BRIT.ORG/SIDA 21(1) 3. Allotoonia parviflora (Sessé & Moc.) J.F Morales & J.K. Williams, comb. nov. (Figs. 2, 12, 13) BASIONYM: Echites ea Sessé & Moc., Naturaleza (Mexico City), ser. 2, 1, app. 28. 1888, non Roxb (1832). TypE: MEXICO. MICHOACAN: Apatzingan, date lacking (f1), Sessé G Mocino ee ECTOTYPE: fide Williams 2002, MA; ISOLECTOTYPE: F; photo F neg. 41238, INB ex MA). Echites woodsoniana Monac., Bull. Torrey Bot. Club. 86:245-247, f. 1. 1959. Prestonia woodsoniana (Monac.) A.H. Gentry, Ann. Missouri Bot. Gard. 70:205. 1983. TYPE: MEXICO. MICHOACAN: Apatzingan, 13 Oct 1939 (f1), Hinton 15325 (HOLOTYPE: NY; ISOTYPES: GH, MO, NY, TEX, US [2 sheets}) Suffruticose liana, stem subterete, glabrous. Leaves: petioles 3-15 mm long, with minute pectinate colleters in the axils, glabrous; blade 3.2-9.2(-11.4) x 0.9-3.6 cm, membranaceous to somewhat subchartaceous, elliptic, narrowly elliptic to narrowly ovate, apex acuminate, acute to acute-mucronate, base obtuse to cuneate, glabrous, midvein impressed on both surfaces, the secondary vena- tion scarcely impressed or inconspicuous. Inflorescence longer than the sub- tending leaves, lax, peduncle 1.3-7 cm, pedicels 2-9 mm, bracts 1-1.5 mm long, almost linear, acuminate, scarious; sepals 1-15 x 0.5-1 mm, ovate to narrowly ovate, acute to acuminate, glabrous to glabrate, more rarely minutely and in- conspicuously papillate, the colleter very slightly erose; corolla salverform, or- ange to yellow-orange, glabrous without, tube 3.5-6 mm long, ca. 1.5 mm diam, straight, gaat within around the stamens, lobes 8-14(-16) « 15-2 mm, densely hirsut ially; stamens: filaments less than 0.5 mm, inconspicuous, anthers 3.5-4 mm, glabrous, auricles ca.0.8 mm, ovary 1.5-2 mm, glabrous, style head 2-3 mm long, nectaries 0.5-1.5 mm, distinct to somewhat connate, entire. Follicles 30-40 x 0.4-0.5 cm, continuous to obscurely moniliform, smooth and glabrous, somewhat twisted distally; seeds 21-27 x 1-15 mm, rugose, the creamish coma 3.8-4.2 cm. Distribution.—Dry forest, premontane moist forest, disturbed areas, savan- nas, semideciduos forest, rocky forested outcrops, and gallery forest in Mexico, Guatemala, Honduras, Nicaragua, and Costa Rica, at elevations of 0-1100(-1400) m. (Fig. 13). Associated species include Anacardium, Astronium, Bursera, Enterolobium, Lonchocarpus, Ficus, and Plumeria. Phenology.—Flowering January, April-May, and July-December. Fruitng collections have been collected in January-April and November Local names.—Bejuco de Sapo (Mexico, Guerrero, Zirandaro). This species has a complicated taxonomic history having been treated re- peatedly in Prestonia. The history of this species is well documented in Will- iams (2004). Allotoonia parviflora is immediately distinguished by its rounded corolla bud (Fig. 10) and pubescent corolla lobes (Fig. 2 B). It is often confused with A. tuxtlensis, which has a pointed corolla bud (Fig. 10) and glabrous corolla lobes. Also, Allotoonia parviflora consistently has corolla lobes longer than the tube, while the corolla lobes in A. tuxtlensis are consistently shorter than the tube. ALLOTOONIA GENUS OF APOCYNACEAE 149 A—f) fram A, v [, [+ Nisin es A ; if (A C. Corolla. D. N 8241, INB; E from Morales 3739, INB). A. Flowering shoot. B y E. Fruits 150 BRIT.ORG/SIDA 21(1) The holotype of E. woodsoniana (=Allotoonia parviflora) at NY has corolla lobes 15-16 mm long, making it appear to be a specimen of A. turbinata. Allotoonia turbinata differs from A. parviflora by its flowering bud, which has the corolla lobes extended upwards rather than folded in (Fig. 10). Although Echites parviflora Sessé & Mog. is a later homonym of E. parviflora Roxb., its use in the new combination Allotoonia parviflora (Sessé &t Moc.) J.E Morales & J.K. Williams is legitimate according to article 58.3 of The Code. Monachino (1959) recognized that Echites parviflora Sessé & Mog. isa hom- onym of E. parviflora Roxb., and thus regarded its use by Woodson (1936) as illegitimate. Unfortunately, Monachino did not have access to the type of E. parviflora Sesse & Mog. and was therefore uncertain as to its true identity. Rather than proposing a new name for the species he decided to re-describe it as a new species, giving it the name E. woodsoniana Monach.,,as was exposed previously by Gentry (1983). Observations of the lectotype of E. parviflora at MA by the second author confirm that the specimen is conspecific with E. woodsoniana. Echites woodsoniana is here treated as a synonym of the new combination Allotoonia ee Specimens examined. MEXICO. Guerrero: Tario, Coyuca, 5 Apr 1935, Hinton et al. 7589 (P, US); Tario, Coyuca, 4 Feb 1934, Hinton een El Talamo, road Zirandaro-Guayamo, 6 Sep 1982, Soto & ne 4355 (INB, MEXU). Jalisco: Jilotlan de Los pes Rio Tepalcatepec, 21 Sep 1983, Martinez et al. 436 CINB, MEXU). Michoacan: + kme of Apatzingan along river, 3 Jan 1997, Williams 96-88 (SHST, a Arteaga, road to Playa Azul, 11 Nov 1977, Koch etal. 77462(US). Arteaga, road Arteaga to Nueva Italia, 27 Sep 1983, Martinez & Stevens 4597 (INB, MEXU); along road to oo ria 1983, Miller et al. 456 (MO [2 sheets). Oaxaca: Mpio. San Miguel Chimalapa, Rio Escondido (Arroyo Baul), W of the union with Rio Portamonedas and Benito Juarez, ca. 38 km in a straight line to the N of San Pedro Tapanatepec, 8 Oct 1985, Maya 2336 (TEX). Quintana Roo: N de Estero Franco, 30 Jul 1984, Cabrera & Cabrera 6885 ce U, MO). GUATEMALA. Chiquimula: between Ramirez and Cumbre de Sie 15 Oct 1940, Standley 74562 (F). HONDURAS. Comayagua: Comayagua Valley, 1] Sep 1974, Hazlett s.n. (MO); Quebrada Chicuas, cerca de El Agua Salada, 21 Jul 1962, Molina 10996 (EAP, F, NY). NICARAGUA. Chontales: near Cuapa, 14 Jul 1976, Neill 619 (DUKE, MO); N of Cuapa, ee Neill 2488 (MO), N of Cuapa, 21 Jan adie Stevens 6106 (BM, MO), ca. 2.8 km about Cuapa c 1983, Stevens 22695 (DUKE, MO, NY). Rivas: Isla Omotepe, 27 Apr 1984, Robleto 396 (MO); eee Robleto 537 (MO), 14 Dec 1984, Robleto 1611 (MO), Volcan sae 21 Sep 1984, Robleto 1218 ae COSTA RICA. Alajuela: road to Upala, Hacienda Carbor v 1985, Gomez etal. 2 (CR, F MO). Guanacaste: Finca Tenorio, Las Canas, 18 Jul ee Creel ae (MO); La Pacifica, 26 Jan ae Gentry 307 (MO), Finca La Pacifica, Canas, 21 Sep 1975, Janzen 10148 (MO); Rio Tenorio, near La Pacifica, 20 Nov 1972, Opler 1567 (COL, CR, E MO, NY). Puntarenas: Monteverde Biological Reserve, 27 Aug 1993, Bello G Cruz 5278(CR, INB, MO). San José: Acosta, Valle del Rio Candelaria, near Puente, 15 Apr 1995, Hammel 19760 (INB, MO); Acosta, Rio Candelaria, 19 Nov 1994, Morales 3139 (CR, INB [2 sheets], MO), Morales 3161 NB, MO), Cerros de Caraigres, Rio La Me a, Ceiba Este, Morales & Corrales 6033 (INB, MO); Rio Jorco, Acosta, between Bajo Badilla and Bajo Cardenas, 22 Aug 1998, Morales 6481 (NB, MO): Rio Candelaria, W of Resbalon, 3 Dec 2001, Morales & Quirés 8241 oi INB, MO). Acosta, between Monterrey and Las Ceibas, | Dec 2002, Morales & Abarca 8810 (INB 4. aay turbinata (Woodson) J.-F Morales & J.K. Williams, ates nov. (Figs. 14). BASIONYM: Echites turbinata Woodson, Ann. Missouri Bot. Gard. 21:615. ALLOTOONIA GENUS OF APOCYNACEAE 151 400 km } * apres : ‘ ) ¢ = Allotoonia tuxtlensis > re 4= Allotoonia parviflora ae i er a «= Allotoonia turbinata sf i a ots Pb S a fv oa 5. x Pit: a Fic. 13. Distribution of A. parviflora, A. turbinata and A. tuxtlensis. 1934. TYPE: COSTA RICA: Alajuela: Rancho Flores, 22 Feb 1890 (fl), Tonduz 2147 (HOLOTYPE: B [destroyed]; LECTOTYPE, here designated: US; ISOLECTOTYPES: BR [2 sheets]). Suffruticose liana, stem terete to subterete, glabrous. Leaves: petioles 7-21 mm long, without colleters in the axils, glabrous; blade 4.5-14.5(-17) x 2.1-8.7 cm, membranaceous, rarely subchartaceous, elliptic to narrowly elliptic, apex cau- date-acuminate, rarely acuminate, base obtuse to cuneate, sometimes oblique, glabrous, midvein impressed on both surfaces, the secondary venation slightly impressed or inconspicuous. Inflorescence shorter than the subtending leaves, lax to somewhat agglomerate, peduncle 1.5-5.1(-5.4) cm, pedicels 4-15 mm, bracts 1-2.5 mm long, very narrowly elliptic to linear, acuminate, scarious; se- pals 2-3.5 x 1-15 mm, ovate, long-acuminate, glabrous to glabrate, minutely rugose, the colleter irregularly fimbriate; corolla salverform, greenish yellow, glabrate to inconspicuously and sparsely puberulent without, tube 4-6 mm long, ca. 1.5 mm diam, inflated at the attachment of the stamens, puberulent to glabrate within around the stamens, lobes (17-)20-27 x 1 mm, glabrous to glabrate; stamens: filaments ca. 1 mm, glabrate, anthers 3-3.5 mm, glabrous, auricles 0.5-0.8 mm, ovary 1.5-2 mm, glabrate, style head 1-1.5 mm long, necta- ries ca. 1.5 mm, entire. Follicles 20-31 x 0.4-0.6 cm, obscurely moniliform, smooth and glabrous; seeds 27-30 x 2-3 mm, rugose, the tannish coma 3-5.5 cm. 152 BRIT.ORG/SIDA 21(1) 2.5mm Fic. 14. Allotoonia turbinata (A—C from Kupper 937, M; D from Gomez-L. et al., 12142, INB). A. Flowering shoot. B. Sepals, colleters, nectaries, and ovary. C. Fruits. ALLOTOONIA GENUS OF APOCYNACEAE 153 Habitat and distribution.—Montane Rain forest and related disturbed ar- eas in Mexico (Chiapas), Honduras, Costa Rica, and Panama, between 1500-2350 m. (Fig. 13). Associated species include Clethra, Eugenia, Liquidamber, Pinus Quercus, and Randia Phenology—Flowering specimens have been collected in January-May, June, and September. Fruiting collections are reported in January-February, and June. Allotoonia turbinata is immediately distinguished by its corolla lobes 1.2- 2.7 cm long. This species is similar to A. parviflora but differs in its corolla lobes which are longer and extended upwards in bud (vs. rounded in bud, Fig. 10). The elevation range of this species (between 1500 m and 2350 m) is unique, considering that the other species in Allotoonia are found mostly at low eleva- tions and the fact that in Mesoamerica very few apocynaceous lianas grow at 1500 m altitude. This species was originally considered endemic to Costa Rica (Rancho Flores, Cordillera Volcanica Central), but further collections have broadened of its known range. 9 Specimens examined. MEXICO. Chiapas: Mpio. Villa Corzo, E base of Cerro Tres Picos near Cerro Bola along a logging road SW of Colonia Agronomos Mexicanos, 4 May 1972, Breedlove 25058 (MO [2 sheets], TEX). HONDURAS. Cortéz: Montane Idalfonso, N de Cofradia, 17 Apr 1957, Molina 6203(EA F). COSTA RICA. Alajuela: near Carrizal, 31 Jan 1992, Gomez-L. et al. 12142 (INB, USJ); Pods, 27 Mar 1932, Kupper 937 (M [2 sheets]). Heredia: Monte Barva, 1845-1848, Oersted 15544 (C); Las Lajas de San Isidro, Sep 1900, Pittier 14035 (US). PANAMA. Chiriqui: vicinity of Cerro Punta, 21 Jan 1939, Allen 1524 (MO); Rio Chiriqui Viejo Valley, between El Volcan and Cerro Punta, 15 Mar 1938, White 6 (MO, US); vicinity of Casita Alta, Volcan de Chiriqui, 28 Jun 1938, Woodson et al. 988 (GH, MO, NY) 5. Allotoonia tuxtlensis (Standl.) J.-F Morales & J.K. Williams, comb. nov. (Figs. 4, 15). BASIONYM: Echites tuxtlensis StandL., Contr. U.S. Natl. Herb. 23.1164. 1924. Type: MEXICO. CHIAPAS: near Tuxtla, 1 Sep 1895 ({D, Nelson 3080 (HOLOTYPE: US). Sutfruticose liana, stems terete to subterete, glabrous. Leaves: petioles 3-11 mm long, with minute pectinate colleters in the axils, rarely eglandular, glabrous; blade 4.5-11 x 1.4-3.7(-4.8) cm, membranaceous, narrowly elliptic to narrowly ovate, apex acuminate, base obtuse to cuneate, midvein impressed on both sur- faces, the secondary venation slightly impressed or inconspicuous. Inflorescence longer than the subtending leaves, lax, peduncle 2.6-8 cm, pedicels 8-19 mm, bracts 0.8-1.5 mm long, ovate, acuminate, scarious; sepals 1.5-L8 x 1mm, ovate, acuminate, glabrous, the colleter entire or somewhat laciniate; corolla salver- form, yellow to creamish yellow, glabrous without, tube 8-10 mm long, 1.5-2 mim diam, inflated at the attachment of the stamens, puberulent within around the stamens, lobes 6-8 x 1-1.5 mm, glabrous; stamens: filaments inconspicuous, anthers ca. 4 mm, glabrous, auricles ca. 0.5mm, ovary l-1.5 mm, glabrous, style head 1.5-2 mm long, nectaries 0.5-1 mm, distinct to somewhat connate, entire. Follicles 13-15 x 0.3 cm, continuous, smooth and glabrous, sometimes some- what twisted distally, seeds 11-13 x 1.5 mm, minutely and incot Ispic uously rug- ose, the creamish coma (2.3-)2.5-3.2 cm. _— BRIT.ORG/SIDA 21(1) 154 Fic. 15. Allotoonia tuxtlensis (A—C from Arvigo et al. 536, INB; D from Renderos 378, INB) A. Fertile shoot, showing the D. Corolla ls B. Sepa I ALLOTOONIA GENUS OF APOCYNACEAE 155 Habitat and distribution.—Tropical deciduous forest and disturbed areas of Mexico (Chiapas, Campeche, Oaxaca, Quintana Roo, and Yucatan), Belize, Guatemala, Honduras, and disjunct into El Salvador, 100-1200 m. (Fig. 13). As- sociated species include Aspidosperma, Bursera, Ficus, Pinus, Quercus, and Spondias. This species is occasionally cultivated and it is suspected that the populations in Northern Mexico (San Luis Potosi, Querétaro, and Veracruz) are introduced rather than native. Phenology.—Flowering all the year. Fruiting collections have been made in January and May. Local names.—Tasnk’ub ts’aah (México, Huasteca): centipede vine (Mexico); bejuco lengua de culebra (El Salvador, Ahuacapan, San Benito); loroco de Culebra (El Salvador, Cuscatlan). The name Loroco is used for others Apocynaceous lianas in northern Mesoamerica [e.g,, Fernaldia pandurata (A. DC.) Woodson. Allotoonia tuxtlensis is readily distinguished from the other species of Allotoonia by its contorted and pointed corolla buds (Fig. 10) and its corolla lobes shorter than the tube. In both these respects, A. tuxtlensis resembles A. agglutinata. The main differences between the two taxa is the conspicuous sec- ondary veins in the leaves of A. agglutinata and its restriction to the West In- dies. At present no specimens with visible secondary veins in the Mexican and Central American collections of A. tuxtlensis have been observed. This species was considered to be restricted to the Caribbean slopes in northern Mesoamerica (Morales 1997), but it was recently collected in the Cuscatlan Department, El Salvador Alcorn (1984) reported that this species is cultivated and protected in San Luis Potosi, Mexico. She reports that the leaves are used as a medicine in the cure of respiratory problems, including wheezing and difficulty in breathing. The leaves are boiled in a tea and drunk. Specimens examined. MEXICO. Campeche: Calakmul, road to Dos Caminos, 14 Oct 1997, Alvarez 401 (MEXU, MO), Santa Maria Xacabacab, 7 Mar 1982, Cabrera et al. 2062 (INB, MEXU); Champoton, road to Escarcega, 23 Oct 1997, C vali & May 4628 (CICY, MO); Calakmul, road Xpujil-Campeche road, 22 Nov 1997, Lira et. al. 360" (MEXU, MO); Calakmul, 24 Nov 1997, Martinez et. al. 29893 (MEXU, MO), Hopelchén, SE of Xpujil, 23 Jan 1996, Pascual 150 (MO). Chiapas: between Pinola (Las Rosas) and Soyatitan, 27 nee 1981, Breedlove 52441 (MO [2 sheets); See 14 Sep 1981, Breedlove 52710 (MO); Ococingo, Rio Usumacinta, + Dec 1984, Martinez 8987 (IN U); Cerro Brujo, Ocozocuautla, 25 Oct 1985, ie ne AEXU, a NW of Boca Lacantum, 3 a 85, Martinez 14468(MEXU, MO); Ococingo, oe to Palenque, + Nov 1985, Martinez 14883 (MEXU, MO); Ococingo, 10 Jan 1986, Martinez 16205 (MEXU, MO); Ocosingo, road to Chapul, S of Boca nee 18 Apr 1986, Martinez 18362 (MEXU, os Comitan, road to Tzimol, 20 Sep 1988, Martinez & Stevens 23940 (INB, MEXU, MO), Tonala, Sep 1913, Purpus 6825 (BM, GH); La nae ates 21 Feb 1984, Shilom 7306 (INB, MEXU);E of Cintalapa, 19 Nov 1984, Téllez et al. 8126 (IN nee Oaxaca: Santa Maria Chimalapa, Paso La Cueva, Rio Corte, 9 Oct 1984, Her eee pee ere MO, TEX). ee Mpio. Lanad de Matamoros, W of Tilaco, road to Santa Ines, 6 Aug 1985, Fe a 3121 (IEB). cea Roo: N of Estero Franco, road to Tomas Garrido, 30 Jul 1984, Cabrera & Cabrera ae (MEXU, MO). San Luis Potosi: Mpio. San Antonio, San Pedro, 26 Oct 1978, Alcorn 2102 (TEX). Veracruz: Mpio. Chicontepec, in the town of Tlacolula, 26 Feb 1980, Calzada 5866 (F). Yucatan: Progreso, 1932, Flores s.n. (F). GUA- TEMALA. Huchuetenango: between Democracia and Santa Ana, Sierra de los Cuchumatanes, 25 Aug 1942, Steyermark 51307 (F, MO). HONDURAS. Morazan: along Jicarito creek, 13 Aug 1947, Molina 485 (EAP. F). Ocotepeque: road to Esquipulas, 29 Aug 1968, Molina 22423 (F NY). BELIZE. Belize: Belize Camp, 15 May 1991, Arvigo et al. 536 INB, NY); Savanna, 2 Jun 1974, Dwyer 12613 (MO). paeaes High Ridge, 1931-1932, Gentle 439 (BM, MO), Northern River, Dec 1933, Gentle 1022 (MO, NY). T lumbia road, Dec 1946, Gentle 6116 (F MO, NY, eee EL SALVADOR. Abuachapan: sound eee El BRIT.ORG/SIDA 21(1 z 1953 (MO): San F Santa Lina, S of Ahuachapan, 5 Dec 994, | Corozo, 13 Apr 2000, Rosales 544 (B, L AGU, MO), u May 2 2000, Rosales 702 (LAGU Rosales 785 (LAGU, MO): San Benito, 15 Jun 1995, Sandoval & Sandoval 27 (LAGU, MO). Cuscatlan: . 0: Co- ), MO), 19 Aug 2000, artificial lake near Cerron Grande, 9 Dec 1997, Renderos 378 (B, BM, BONN, INB, LAGU, MQ). Santa Ana: S of Metapan, San Diego mountain, May 1995, Linares & Martinez 2728 (MO). APPENDIX Li NUMERICAL LIST OF ACCEPTED TAXA Allotoonia turbinata (Woodson) J.F Morales & eS lotoonia agglutinata Jacq.) J.K Morales & J.K. Williams Allotoonia caudat i Weedson) |B Morales Allotoonia parvifl S & Moc.) J. Morales & J.K. Williams .. Williams A Williams APPENDIX 2: INDEX TO NAMES IN SYSTEMATIC TREATMENT Allotoonia agglutinata Jacq.) J.E Morales & J.K. Williams caudata (Woodson) J.E Morales parviflora (Sessé & Moc.) J.Z Morales & J.K. Wil- es Woodson) J.F Morales & J.K. Williams ee (Standl.) J. F Morales & J... Williams Anechites avglutinata Jacq.) Miers (= A. agglutinata) circinalis (Sw.) Miers (= A. agglutinata) circinalis Sw. var. thomasiana A. DC. (= A. utinata) thomasiana (A. DC.) Miers Echites agglutinata Jacq. (= A. agglutinata) circinadlis Sw. (= A. agglutinata) obtusifolia Sessé & Mog. (= A. agglutinata) puntarenensis J.F Morales (= A. caudata) sanguinolenta Tussac (= A. agglutinata) turbinata Woodson (= A. turbinata) ixtlensis Standl. (= A. tuxtlensis) woodsoniana Monac. (= A. parviflora) Haemadictyon circinalis (Sw.) G. Don (= A. agglutinata) nutans (Anders.) A. DC. var. sanguinolent (Tussac) (= A. agglutinata) Prestonia agelutindta Jacq.) Woodson (= A. agglutinata) caudata Woodson (= A. caudata) woodsoniana (Monac.) A.H. Gentry (= A parviflora) APPENDIX 3: INDEX TO EXSICCATAE Alcorn, J., 2102 (5). Allen, P, 1524 (4). Alvarez, D., 401 (5). Arvigo, R. et al., 536 (5). Axelrod, F & B. Axelrod, 3091 (1). Bello, E. & FE. Cruz, 5278 (3) Breckon, G., 4994 (1); 5019 (D. Breedlove, D., 25058 (4); 52441 (5); 52710 (5). Cabrera, E. & H. Cabrera, 6885 (5). Cabrera, E. et al., 2062 (5). Calzada, J., 5866 (5). Carnevali, G. & F. May, +628 (5). Croat, T., 268 (3). DArcy, W, _ (1). Dwyer, J., 12613 (5). Ekman, ‘i 3 an 5131 CD). Estrada, A. & A. Se ee 283 (2). eee R. 31215). Flores, R., (5). Friedri¢ = sthal , 240 (). itoonta tuxtlensis (Standl,) J.E Morales & J.K. d ALLOTOONIA GENUS OF APOCYNACEAE 157 Garcia, R. & G. Caminero, 3287 (1). Nelson, E., 3080 (5). Gentle, P, 439 (5); 1022 (5): 6116 (5). Oersted, A., 15544 (4). Gentry, A, 307 (52). Opler, P, 1567 (3). Gomez-L., J. et al., 12142 (4). Pascual, A., 150 (5). Gomez, L. et al., 23879 (3). Pittier, H., 14035 (4). Hammel, B., 19760 (3). Poveda, L. & V. Castro, 3926 (2). Hammel, B. & M. Garita, 19306 (2), Purpus, C., 6825 (5) Hazlett, D., s.n. Lat Renderos, M., 378 (5) Hernandez, Robleto, W, 199 (2): 306 (3); 537 (3); 912 (2): 1218 Hinton, G., iN cre (3). ae 1701 (2). Hinton, G. et al., 7589 (3). Rohr, J.,9 Holm, R. & H. Iltis, 243 (2). oe 2 (5); 702 (5): 785 (5). Janzen, D., 10148 (3). Ross, R. & D. Meletische, 253 (1). Koch, et al., 77462 (3). le E. & M. Sandoval, 27 (5). Kupper, W,, 937 (2) essé y Pa M. & J. Mocino. 5075 (1); 5068 (3). Linares, J. & C. Meine 1953 (5); 2728 (5). ane 306 (5). Liogier, B., 1615 as ae P, 2457 (1). Lira, E. et al., 360° (5). Soto J. & G. Silva, 4355 (3). Martinez, E. 7) 14303 (5); 14883 (5); 16205 sane P, 74562 (3). (5); 183 Stevens, D., 6106 (3): eae Martinez, : - = Stevens, 4369 (3); 4597 (3): Seek P, 51307 23940 (5). Swartz, O., 265 (1) Martinez, E. et al., 29893 (5). Taylor, C. & R. Gereau, 10493 (1). Maya, S., 2336 (3). Téllez, O., 8126 (5). Miller J. et al., +56 (3). Tonduz, A., aa (4); 13940 (2). Molina, A., 485 (5); 6203 (4); 10996 (3); 22423(5). Urban, L., 3565 Morales, J. F, 3139 (3); 3161 (3); 6481 (3); 8242 (2). — Villacorta. R. a Méndez, 2181 (2). a) ee Morales, J. & R. Abarea, 8810 (3). Vere . — OCD. Morales, J.F & J.F Corrales, 6033 (3). White Morales, J.F & T. Quirés, 8241 (3) ae 4 06- 88 (3). Moreno, P, 16081 (2); 18888 (2): 19664 (2): 22179 ~~ Woodson, R. et al., 988 (4). (2). Zanoni, T. et al., 47051 (1). Neil, D., 619 (3); 2488 (3). ACKNOWLEDGMENTS We wish to thank the curators of A, B, BM, BR, BRIT, C, CAS, CGE, CICY, CIIDIR. CIMI, CM, COL, CR, DUKE, EAP ENCB, F FI, FI-W, G, GH, HAL, HBG, HUA, IEB, JAUM, JEPS, K, LAGU, LD, LE, M, MA, MEDEL, MEXU, MICH, MO, NY, O,P PMA, TRIN, TEX, U, UC, UPS, US, USE USJ, UVAL, W, WAG, WIS, XAL for providing specimens for examination. The senior author thanks Jorge Leon, for giving access to original references in his personal library. The second author thanks Mary Endress and the researches at the Plant Resources Center at the Univer- sity of Texas in Austin. This research was financed in part by the Netherlands Development Agency (NEDA) through the project Development of Biodiversity Knowledge and Sustainable Use in Costa Rica conducted by the National Biodiversity Institute of Costa Rica (INBio). This research was also possible thanks to the Cooperation Agreement between the Ministry of Environment 158 BRIT.ORG/SIDA 21(1) and Energy (MINAE) and INBio in order to accomplish the National Biodiversity Inventory. REFERENCES Atcorn, J.B. 1984. Huasteca Mayan ethnobtoany. University of Texas press, Austin. Eworess, M.E.and Bauyns P.2000.A revised classification of the Apocynaceae s.|. The Botani- cal Rev. 66:1—56. Gentry, A.H. 1983. A new combination for a problematic Central American Apocynaceae. Ann. Missouri Bot. Gard. 70:205-206. Monacino, J. 1959. A new Echites from Mexico. Bull. Torrey Bot. Club 86:245-247. Morates, J.F. 1997.A reevaluation of Echites and Prestonia section Coalitae (Apocynaceae). Brittonia 49:328-336. Wituiams, J.K. 2002. A further evaluation of Echites sect. Yucatanense (Apocynaceae) with additional notes on the genus. Brittonia 54:310-317. Wituiams, J.K. 2004. Polyphyly of Echites (Apocynaceae: Echiteae) based on a morphologi- cal cladistic analysis. Sida 21:117-131. Woopson, R.E. 1936. Studies in the Apocynaceae IV. The American Genera of Echitoideae. Ann. Missouri Bot. Gard. 23:169-438. Woopson,R.E.1938.(Asclepiadales) Apocynaceae.|n:J.H. Barnhart,ed.N. Amer. FI. 29(2):179- 185. New York. ESTUDIOS EN LAS APOCYNACEAE NEOTROPICALES IV: NOTAS TAXONOMICAS EN PRESTONIA (APOCYNOIDEAE, ECHITEAE) CON UNA NUEVA ESPECIE DE ECUADOR J. Francisco Morales Instituto Nacional de Biodiversidad (INBio) partado Postal 22-3100 Santo Domingo, Heredia, COSTA RICA RESUMEN Se discute la utilidad taxonomica de los sépalos para delimitacion de secciones dentro del género 7 ae Po Prestonia. Asimismo, se d lor, P. amabilis, muy inusual por su caliz campanulado. ABSTRACT The taxonomic use of the sepals features for intergeneric delimitation in Prestonia is discussed. Prestonia amabilis (Apocynaceae, Apocynoideae), a new species from Ecuador, is described and il- lustrated Durante la preparacion de una monografia de Prestonia (Apocynaceae, Apocynoideae), un total de 55 especies han sido reconocidas, distribuidas desde México al N de Argentina y las Islas del Caribe (Morales, en prep.). Dado el alto numero de especies de Prestonia originalmente descritas basadas en una sola coleccién (Woodson 1936), y la alta sinonimia prevaleciente en otros géneros, donde se han presentado situaciones similares (Morales 1999), ha sido necesaria la evaluacion de varios caracteres tradicionalmente usados para separar especies (e.g., grado de laceracion de los coléteres de los sépalos y de los nectarios 0 disco nectarifero), lo cual ha comprobado que algunos de ellos son muy inestables y no deben de ser usados como el caracter principal para el reconocimiento a nivel especifico, tal y como fue demostrado por Morales (1997 b) para las especies mesoamericanas de la secci6n Tomentosae. Esto ha conllevado a la sinonimizacion de varios nombres. Las medidas aqui mencionadas estan basadas en los resultados obtenidos durante la preparacion de una nueva monografia del género. La forma y consistencia de los sépalos fue un caracter clave usado por Woodson para separar secciones dentro de Prestonia. Sin embargo, dado que la utilidad de estos caracteres es cuestionable (para separar secciones) y junto con el descubrimiento de una nueva especie de Ecuador, con un céliz bastante inusual, una explicacion se brinda a continuacion. Los sépalos en Prestonia son siempre libres longitudinalmente, estando tnicamente ligados en forma basal al hipantio. Los sépalos pueden ser conspicuamente foliaceos 0 subfoliaceos (e.g., SIDA 21(1): 159 — 163. 2004 160 BRIT.ORG/SIDA 21(1) P. portobellensis (Beurl.) Woodson) a escariosos (e.g., P. quinquangularis Jacq.) Spreng.). Woodson (1936) distinguié tres secciones basado en la consistencia de las sépalos. De esta manera, él separ la seccion Acutifoliae de las secciones Annulares y Iomentosde por sus sépalos pequenos e inconspicuos, escariosos 0 levemente foliaceos y usualmente reflexos apicalmente (vs. grandes y conspicuos, foliaceos a subfoliaceos y no reflexos apicalmente). Sin embargo, estos caracteres no son totalmente confiables para la separacion de estas secciones y la busqueda de otros mas consistentes debe prevalecer, debido a que son subjetivos y estan supeditados al criterio de cada tax6nomo. Primero, en las especies incluidas en la seccion Acutifoliae, los sépalos no son consistentemente reflexos en el apice, pudiéndose encontrar apices reflexos o no reflexos. Luego, los terminos foliaceo, subfoliaceo, o levemente foliaceo son totalmente subjetivos, si no son acompanados de medidas y otros caracteres adicionales, que definan claramente el concepto del monografiador. Por otro lado, algunas especies incluidas en las seccion Annulares (e.g., P. longifolia (Sessé & Moc.) J. F Morales) tienen sépalos escariosos a muy levemente subfoliaceos, de 3-6(-8) x 2-3 mm, no reflexos apicalmente, los cuales no difieren en gran medida de aquellos de otras especies incluidas en la seccion Acutifoliae (e.g., P. lagoensis (Mull. Arg.) Woodson), con sépalos de 3-6 x 1.5-2 mm, reflexos 0 no reflexos apicalmente. Por lo tanto, se sugiere utilizar otros caracteres mas consistentes, como el tipo y densidad del indumento de flores y frutos, presencia o ausencia de los lobulos coronales en el interior de la corola, nivel de insercidn de los lobulos coronales (apicalmente exsertos 0 levemente incluidos, con los apices casi al mismo nivel de la corona anular vs. profundamente incluidos, con los apices por debajo de los apices de las anteras) y textura de sépalos y hojas. Sin embargo, la definicion definitiva de las secciones seran comentadas con detalle en la proxima monogratia del género. La ultima monografia del género Prestonia (Apocynaceae, Apocynoideae) fue hecha por Woodson (1936), quien acept6 61 especies. Cinco especies fueron reportadas en ese entonces para Ecuador, P. mollis Kunth, P. parvifolia K.Schum. ex Woodson, P. peregrina Woodson, P. rotundifolia kK. Schum. ex Woodson y P. schumanniana Woodson, cuatro de ellas endémicas y conocidas por menos de dos colecciones, con solo P. mollis reportada también en Pert. Desde 1936, el numero de expediciones en Ecuador se han incrementado en forma notable, sobre todo en los ultimos 30 anos, lo que producido un aumento considerable en el numero de colecciones de Apocynaceae disponibles hoy en dia. Estos esfuerzos resultaron en la descripcidn de nuevas especies, algunas de ellas muy raras y endémicas (Morales 1997 a). El proceso de revision de especimenes para una nueva monogratfia de Prestonia ha llevador a la necesidad de sinonimizar algunos taxa y elevado el ntmero de especies conocidas para Ecuador a 15 (dos de ellas atin sin describir) y llevado a la necesidad de sinonimizar algunos taxa. Una de estas especies inéditas es muy notable por tener los s¢épalos connados MORALES, UNA NUEVA ESPECIE DE PRESTONIA EN ECUADOR 161 basalmente, conformando una base campanulada, algo bastante inusual dentro del género y que no fue reportado por Woodson (1936). Especies con sépalos connados a lo largo del tercio o mitad basal son muy raras y solamente otros dos taxas tienen la misma caracteristica, P. haughtii Woodson (incluyendo P. macrophylla Woodson) y P. rotundifolia K. Schum.ex Woodson. Estas tres especies estan restringidas a lo largo de los Andes desde la Cordillera Occidental en Colombia, a través de Ecuador hasta el N de Pert y en forma general son conocidas por muy pocas colecciones. Por lo tanto, esta nueva especie es descrita a continuacion. Prestonia amabilis J.F Morales, sp. nov. (Fig. 1). Tipo. ECUADOR. PASTAZA: Haci- San Antonio de Baron von Humboldt, 2 km NE de Mera, 27 Feb-19 Mar 1985 (fl, fr), Neill et al. 5975 (HOLOTIPO: INB; IsoTIPOS: MO, NY, USF fotocopia en INB). Suffruticosa volubilis, ramulis minutis puberulis maturitate sparseque puberulis vel glabratis; folia elliptica, 13-23 _ 6.5-13.5cm, glabra, glabrata vel sparseque puberula, apice breviter acuminata, basi eae obtusa vel rotundata petiohas l later alibus Be acteis bs - mm longis, calycis laciniis ovalis vel ov ale el ili (10- Mee Enna tubo extus glabro, 14-15 mm vel is; folliculis 18-27.5 cm, glabratis. Liana, ramitas diminutamente ferrugineo-puberulentas cuando jovenes, muy esparcidamente puberulentas a glabradas en la madurez, con secrecion lechosa, inconspicuamente lenticeladas, coléteres interpeciolares inconspicuos, 0.5-1 mm de largo. Hojas: lamina 13-23 x 6.5-13.5cm, eliptica, obtusa y abruptamente corta-acuminada o apiculada apicalmente, obtusa a redondeada basalmente, coriacea a subcoriacea, algunas veces algo revoluta marginalmente, glabra y usualmente brillante adaxialmente, inconspicuamente papilada-puberulenta a glabrada abaxialmente, venacion secundaria y terciaria conspicuamente impresas, peciolo 1-2 cm. Inflorescencia corimbosa, axilar, usualmente mas larga que las hojas adyacentes, con muchas flores, diminutamente y densa a esparcidamente ferrugineo-puberulenta, pedtnculo 6.5-21 cm, pedicelos (1-) 1.7-2.8 cm, bracteas 1-2 x 0.5-1 mm, escariosas; base del caliz conspicuamente campanulada, sépalos (0-)12-19 x (3-)4-6 mm, fusionados basalmente a a largode1/3-1/2desu longitud, coridceosa subcoriaceos, a angostamente ovado-elipticos, acuminados a cortamente acuminados, no reflexos apicalmente, diminuta y densamente ferrugineo-puberulentos, raramente glabrados, foliaceos, coléteres 1-1.5 mm de largo, enteros, subenteros, o diminutamente erosos; corola hipocrateriforme, amarilla y moteada con rojo, glabra exteriormente, tubo 14-15 x 3-4 mm, recto, lobulos coronales 1.5-2 mm, incluidos o apicalmente exsertos, corona anular entera, conspicua, lobulos 9- ll x 7-9 mm, obovados; anteras 5 mm, glabras, los apices conspicuamente exsertos, ovario 1-1.5 mm, glabro, cabeza estigmatica ca. | mm, disco sobrepasando el ovario, usualmente profundamente y irregularmente penta- lobado, eroso a algo lacerado. Foliculos 18-27.5 x 0.8-1.2 cm, continuos a levemente moniliformes, esparcidamente lenticelados, glabrados; semillas 16- 19 mm, glabras, la coma 3.5-4.8 cm, café oscuro. BRIT.ORG/SIDA 21(1) Fig. 1. Prestonia amabilis (A-E de Palacios 10102, USF; F-G de Brehm s.n. (INB)). A. Ramita florifera. B. Caliz, pedicelos, y bracteas.C. Caliz abierto lo lat lif | énalos fusi D. Tubo del la abierto, mostrando 4 P las anteras y lobul | pi | E. Disco. F. Foliculos. G. Semilla MORALES, UNA NUEVA ESPECIE DE PRESTONIA EN ECUADOR 163 Distribucion, habitat, y ecologia.—Restringida por el momento a Ecuador, donde crece en bosques muy humedos y areas de vegetacion perturbada, en elevaciones de 300-1100 m. Especimenes con flores fueron colectados de Febrero hasta Junio, y de Agosto hasta Diciembre. Especimenes con frutos fueron colectados en Febrero, Marzo, y Noviembre. Paratipos. ECUADOR. Morona -Santiago: |.im6n Indanza, cuenca del rio Coangos, rio Tsuirin, Oct 1999 (fD, Ronquillo et al. 1018 (MO). Napo: carretera Coca-Loreto, cerca a rio Pinguillo, 20 Oct 1988 (fl), Cerén et al. 5331(MO, USF); Shinguipino, entre rios Napo y Tena, 30 Sep 1960 (fl), Grubb et al. 1688 (K, NY); Tena, carretera Campococha-Chontapunta, 23 Ago 1997 (f1), eee bomen O), NNO de Coca, rio Huashito, 27 Oct 1982 (£1), Pennington 10637 (K, QCA). Pastaza: Arajuno, campamentos 11- 12, 15-20 Sep 1998 (f1), Freire et al. 3433 (MO); Puyo, Santa Cecilia, a ine 1992 (£1), as 10102 (MO, QCNE, USF); Puyo, Los Vencedores, carretera Puyo-Macas, 8 Jun 1996 (£1), Soejarto et al. 9651 (F); Pozo Villano, 3 Dic 1991 (£1), Tipaz et al. 426 (MO, QCNE). Zamora-Chinchipe: Parque Nacio- nal Podocarpus, 4 km al S de Zamora, 30 Nov 2000 (f], fr), Brehm s.n. (INB); Nangaritza, Miazi, rio Nangaritza, 28 Jul 1993 (st), Gentry 80568 (MO). Prestonia amabilises similar a P. haughtii Gincluyendo P. macrophylla Woodson) por tener sus sépalos connados basalmente y formando una conspicua base campanulada, pero difiere por su inflorescencia repetidamente ramificada, sépalos diminutamente ferrugineo-puberulentos (vs. glabros o glabrados), anteras apicalmente exsertas (vs. incluidas) y l6bulos coronales levemente exsertos 0 al menos al nivel de la corona anular (vs. profundamente incluidos). Esta especie esta ademas algo relacionada a P. annularis, pero difiere por sus inflorescencias mas grandes, sépalos ferrugineo-puberulentos, base del caliz campanulada, y frutos mas gruesos y algo lenosos. AGRADECIMIENTOS Quiero agradecer a los herbarios K, MO, NY, QCA, US, USF por el envio de mate- rial en préstamo. Asimismo, se agradece al personal del Herbario de la Univer- sidad de Antioquia (HUA), por las facilidades brindadas durante mis visitas a Medellin, Colombia. También deseo agradecer a Gunnar Brehm y Juergen Homeier, quienes suministraron fotografias, material preservado y duplicados de Prestonia amabilis. REFERENCIAS Morates, J.F. 1997 a. A synopsis of the genus Allomarkgrafia (Apocynaceae). Brittonia 49: 337-345. Morates,J.F. 1997 b.A synopsis of the genus Prestonia section Tomentosae in Mesoamerica. Novon 7:59-66. Mora es, J.F. 1999. A synopsis of the genus Odontadenia (Apocynaceae), series of revisions of Apocynaceae XLV. Bull. Jard. Bot. Belg. 67:381-477. Woodson,R.E. 1936. Studies in the Apocynaceae.IV.The American genera of Echitoideae. Ann. Missouri Bot. Gard. 23:276-436. 164 BRIT.ORG/SIDA 21(1) BOOK NOTICE PaMELA A. MATSON, ASHOK GADGIL, and DANieL M.KAMMEN (eds). 2003. Annual Review of Environment and Resources: Volume 28, 2003. ISBN 0-8243-2328-9, hbk: ISSN 1543-5938). Annual Reviews Inc., 4139 El Camino Way, P.O. Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews.org, 800-523-8635, 650-493-4400, 650-424-0910 fax). $185.00 (USA), $190.00 (Int'l), 648 pp., 6" x 9". Contents of Volume 28 of Annual Review of Environment and Resources: L. Earth’s Life Support Systems Climate Change, Climate Modes, and Climate Impacts the Atmosphere Evaluating Uncertainties in Regional Photochemical Air Quality Modeling = he Cleansing Capacity o Transport of Energy, Information, and Material Through the Biosphere lobal State of Biodiversity and Loss Patterns and Mechanisms of the Forest Carbon Cycle Il. Human Use of Environment and Resources Dynamics of Land-Use and Land-Cover Change in Tropical Regions Urban Centers: An Assessment of Sustainability Water Us ae Cereal Demand While Protecting Natural Resources and Improving Environmental Qua State of a World’s Fisheries Green Chemistry and Engineering: Drivers. Metrics, and Reduction to Practice Ill. Management and Human Dimensions International E l Agreements: A Survey of Their Features, Formation, and Effects Tracking Multiple Pathways of Human Exposure to Persistent Multimedia Pollutants: Regional, Continental, and Global Scale Models Geographic Information Science and Systems for Environmental Management The Role of Carbon Cycle Observations and Knowledge in Carbon Management Characterizing and Measuring Sustainable Developmen Just Oil? The Distribution of Environmental and Social pe of Oil Production and Consumption SIDA 21(1): 164. 2004 ESTUDIOS EN LAS APOCYNACEAF NEOTROPICALES V. UNA NUEVA ESPECIE NUEVOS REPORTES. Y NUEVA SINONIMIA EN LAS APOCYNACEAE DE BOLIVIA J.Francisco Morales Alfredo Fuentes Instituto Nacional de pia idad Heeane del eke ene ee) A artado 22- M i Gir Mercia Santo Domingo, Heredia, ee C Universidad Auto M | R Cre MICTETIO Casilla 2489, canta Gis BOLIV RESUMEN Se describe e ilustra una nueva especie del Benero ms stontia, y taxas relacionados son discutidas. Se bri lde13 Bolivia y se proponen nuevas sinonimias en los géneros Himatanthus, Prestonia, y Tbe r dened ABSLRACT A new species of Prestonia is described and illustrated, and its relationships with related taxa are discussed. In addition, 13 new records of Apocynaceae (subfamily Apocynoideae) are reported for Bolivia and new synonymy on Himatanthus, Prestonia, and Tabernaemontana are proposed. Bolivia es uno de los paises menos explorados desde el punto de vista botanico en América del Sur. Hasta la fecha, no se ha realizado ningtin tratamiento inte- gral para su flora y practicamente, la unica referencia disponible es el trabajo de Foster (1958), el cual consiste en un catalogo basado en la informacion de trabajos monograficos previos. Al igual que casi todo el resto de la flora, no existen tratamientos para la familia Apocynaceae (subfamilia Apocynoideae) y las Unicas referencias parciales son las monografias de varios géneros (e.g, Monachino 1943, 1945a; Morales 1999, 2002; Woodson 1933, 1936). Elincremento de la actividad botanica en los tltimos 20 anos en el pais, han provocado un notable aumento de nuevos reportes y novedades en la flora en general (e.g., Al- Shehbaz 1999; Fuentes 1998; Justiniano y Toledo 2001; Moraes 1996; Navarro 1997; Zuloaga et al. 1993). Como parte del proceso de elaboraci6n de las Apocynaceae para el Catalogo de las Plantas Vasculares de Bolivia, una serie de novedades taxonomicas han sido encontradas. Estas incluyen géneros y especies anteriormente no citados (e.g., Lacmellea, Stipecoma), nuevas sinonimias (e.g., Himatanthus, Prestonia) y una nueva especie de Prestonia. Para el caso de nuevos reportes, solo se cita un especimen representativo por Departamento, y cuando el caso loamerita, se brindan ios adicionales para cada especie o género. Asimismo, solo se citan géneros 0 especies que no hayan sido reportados nteriormente en el checklist de Foster (1958) 0 en los trabajos de Chavez de Michel (1993) Ezcurra (1981, 1984), Fallen (1983), Gensel (1969), Hansen (1985), SIDA 21(1): 165-174. 2004 166 BRIT.ORG/SIDA 21(1) Killeen et al. (1998), Leeuwenberg (1994), Marcondes-Ferreira (1988), Meyer (1955), Monachino (1943, 1945a, 1945 b), Morales (1997, 1999, 2002, 2003), Plumel (1991), Rao (1956), Sakane y Sheperd (1987) y Woodson (1933, 1935, 1936, 1951). Para la determinacion de la nueva sinonimia en Prestonia, todas las colecciones tipo fueron examinadas (a menos que se indique lo contrario) como parte del trabajo monografico realizado por el primer autor para la serie Flora Neotropica. Asimismo, las medidas de P. lagoensis (Mull. Arg.) Woodson y Mandevilla scabra (Hoffmans. ex Roem. & Schult.) K. Schum. utilizadas como referencia de comparacion en la descripcion de P. boliviana y M. symphitocarpa, fueron extraidas de las monografias de Prestonia y Mandevilla (Morales, en prep.), las cuales pueden diferir en alguna forma de los trabajos de Woodson (1933, 1936). NUEVA ESPECIE Prestonia boliviana J.-F Morales & A. Fuentes, sp. nov. (Fig. 1). Tipo: BOLIVIA. CHUQUISACA: valle del Rio Limon, entre Padilla y Monteagudo, 13 Feb 1994 (£1), Wood 7970 (HOLOTIPO: LPB; ISOTIPOS: INB, Kk). Suffruticosa volubilis, ramulis glabris. Folia late ovalia, 8-9 x 4-5 cm, glabra, apice acuminata, basi obtusa vel rotundata, petiolus 1.5-2.5 cm. Inflorescentia racemosa, lateralibus, bracteis 15-2 mm longis, scariaceis, calycis laciniis anguste ovatis, 2.5-3 mm longis, glabris, corola salverformis, tubo extus glabro, 8-9 mm, appendicibus lobis coronae paulo exsertis. Folliculis ignotis. Liana, ramitas glabras, con secrecion acuosa, coléteres interpeciolares inconspicuos, menos de 0.5 mm de largo. Hojas: lamina 8-9 x 4-5 cm, anchamente ovada, acuminada apicalmente, obtusa a redondeada 0 cordada basalmente, delicadamente membranacea, no revoluta marginalmente, glabra, venas secundarias levemente impresas, venaciOn terciaria Casi inconspicua o inconspicua, peciolo 1.5-2.5 cm. Inflorescencia racemosa 0 corimbosa y dicotomicamente ramificada, pero ramificandose solo una vez cerca de la base y pareciendo subracemosa, axilar, usualmente mas larga que las hojas adyacentes, laxa, con muchas flores, glabra a glabrada, pedunculo 5-6.5 cm, pedicelos 2-3 cm, bracteas 1.5-2 x 0.5-1 mm, escariosas; base del caliz corta e inconspicua, no campanulada, sépalos 2.5-3 x 1mm, libresa lo largo de suentera longitud, delicadamente membranaceos, muy angostamente ovados, acuminados, los apices algo reflexos apicalmente, glabros a glabrados, escariosos, coléteres 0.5-1 mm de largo, muy diminutae inconspicuamente lacerados; corola hipocrateriforme, crema, glabra exteriormente, tubo 8-9 x 2.5mm, recto, lobulos coronales 2.5-3 mm, apicalmente exsertos, corona anular entera, conspicua, lobulos 8-10 x 5mm, obovados; anteras 4 mm, glabras, los apices algo exsertos, ovario ca. 5 mm, glabro, cabeza estigmatica ca. 2 mm, nectarios ca. | mm de largo, enteros a subenteros. Foliculos desconocidos. Distribuci6n, habitat y ecologia. Endémica al departamento de Chuquisaca, Bolivia, donde crece en bosques estacionalmente humedos y areas de vegetacion disturbada asociada de la formacion Boliviano-Tucumana, en elevaciones Fic. 1 Prestonia ee he LPB). A. parma florifera. B. pe pedicelos, ai Ne C.Sé y coléteres, vista adaxial. D. C lobul 7 Uvall¥, 168 BRIT.ORG/SIDA 21(1) entre 1300-1500 m. Especimenes con flores han sido recolectados en Febrero y Mayo. Paratipos. BOLIVIA. CuuQuisaca: entre Padil 9326 (LPB). a y Monteagudo, rio Marcani, 1 Jun 2003 (£1), Morales Prestonia boliviana pertenece a la seccion Acutifoliae por sus flores glabras exteriormente y sépalos pequenos e inconspicuos (Woodson 1936). Esta especie esta cercanamente relacionada con P. lagoensis (Mull. Arg.) Woodson, ya que vegetativamente son muy similares (e.g, textura y forma de las hojas, estructura de la inflorescencia) y ambas tienen flores con los l6bulos coronales conspicuamente exsertos apicalmente. Sin embargo, P. boliviana es facilmente separada por sus [lores con los sépalos 2.5-3 x 1 mim (vs. 3-6 x 1.5-2 mm), tubo de la corola mas pequeno, 8-9 mm de largo, (vs. 13-20 mm) y pedicelos mas largos, con 20-30 mm de longitud [vs. 8-14(-18) mm]. NUEVOS REPORTES 1. Couma macrocarpa Barb. Rodr, Vellozia (ed. 2) 1:32, pl. 1, f. b. 1891. Solo hemos visto un especimen estéril, pero que sin duda pertenece a esta especie. Debidoa la escasez de colecciones suramericanas en la primera mitad del siglo XX, no fue reportada para Bolivia por Monachino (1943). El género Couma se reconoce facilmente por sus hojas verticiladas, peciolos con coléteres basales y frutos bayas carnosas, usualmente comestibles. El género esta ampliamente distribuido en toda la cuenca amazonica, siendo C. macrocarpa la especie mas comun. Especimenes representativos examinados: BOLIVIA. Pando: Federico Roman, Rio Negro, tributario del Abuna, Vargas et al. 988 (LPB, USZ). 2. Forsteronia affinis Mull. Arg., Fl. Bras. 6(1):100, pl. 30. 1860. Fosteronia es un género mayormente Suramericano, con unas pocas especies presentes en las Islas del Caribe, México y Mesoamérica. Ampliamente distribuida en Colombia, Ven- -zuela, Pert y Brasil, esta especie era desconocida en Bolivia (Hansen 1985), pero recientes colecciones en el Departamento de Santa Cruz, han confirmado su presencia en este pais. Especimenes representativos examinados: BOLIVIA. Santa Cruz: oe Nacional Noel Kemplf Mercardo, Campamento Las Gamas, Foster et al. 476 (MO, USE, L 3. Forsteronia australis Mull. Arg., Fl. Bras. 6(1):103. 18600. Esta especie es bastante rara y anteriormente era conocida solo en Brasil, donde se reportaba de localidades aisladas en los estados de Bahia, Ceara, Goias, Minas Gerais, Rio de o (Hansen 1985). En Bolivia se conoce solamente en el —_— Janeiro y Sao Pau departamento de Santa Cruz, de los bosques semideciduos chiquitanos. Especimenes representativos examin: a BOLIVIA. Santa Cruz: Florida, Quebrada La Coca, al SO de Bermejo, Nee 44409 (MO, NY, USF, 4. Forsteronia graciloides Woodson, Ann. Missouri Bot. Gard. 22:163-165. 1935. Aunque anteriormente fue conocida solo de Pert y una coleccion disyunta de Colombia (Hansen 1985), especimenes adicionales han ampliado su rango de extension a Ecuador y Bolivia, donde es reportada por primera vez. Esta especie se puede reconocer con facilidad por el relativamente pequeno tamano de sus hojas, con las laminas obovadas y la presencia de domacios pubescentes a lo Bre del nervio central abaxialmente. los: BOLIVIA. La Paz: Abel Iturralde, Parque Nacional Maididi, rio nai arroyo Rudidi, Paniagua et al. 5099 (LPB, MO). 5. Galactophora calycina (Huber ex Ducke) Woodson, Ann. Missouri Bot. Gard. 19:50. 1932. Galactophora es un género de hierbas erectas tipico de zonas abiertas y sabanas, restringido a Suramérica, donde ocurre desde la Amazonia colombiana hasta Brasil y Bolivia. Anteriormente era conocida solo en Colom- bia, Venezuela y Brasil. Aunque historicamente G. calycina se ha considerado como una especie distinta de G. crassifolia (Mull. Arg.) Woodson (Woodson 1936), es probable que representen una misma entidad, dado que la unica diferencia permisible entre ambas es la presencia 0 ausencia de la pubescencia en hojas e inflorescencias y existen especimenes con un amplio grado de variacion en relacion a la densidad del indumento, situaci6n que se repite en otros géneros de la tribus Apocyneae (e.g., Odontadenia) y Mesechiteae (e.g,, Mandevilla). En todo caso, ambas especies son exactamente iguales en el resto de caracteres ene ee los: BOLIVIA. La Paz: Iturralde, Luisita, Haase 827 (PB). Santa én et al. 2545 (INB, MO, SCZ). — Gia Velasco, entre Florida y Bella Vista, Guil 6. Lacmellea aculeata (Ducke) Monachino, Lloydia 7:292. 1944 [1945]. El género Lacmellea esta confinado mayormente a Sur América, donde la mayor cantidad de especies estan restringidas a la cuenca amazonica, aunque si bien, unas pocas especies estan presentes en Mesoamérica (Morales 1998). La Ultima monografia fue hecha por Monachino (1944), quién entonces no reporté ninguna especie para Bolivia. Lacmellea aculeata esta ampliamente distribuida en la cuenca baja amazonica, desde Colombia y las Guyanas hasta Brasil y Bolivia. Una especie adicional, probablemente no descrita y cercanamente relacionada con este taxon, con hojas densamente pubescentes abaxialmente y conocida solo por ecciones con frutos del Pee de Pando, se encuentra en Bolivia. E t los: BOLIVIA. Pando: Rio Madeira, cerca de Abuna, Prance et al. 8358 (NY Us) 7. Lacmellea arborescens (Mull. Arg.) Markgr. Notizbl. Bot. Gart. Berlin-Dahlem 15:626. 1941. La revision de Lacmellea de Monachino (1944) reporto esta especie como endémica a Brasil, pero el proceso de elaboracion de una nueva monogratia del género (Morales, en prep.), ha extendido su rango hasta Pert y Bolivia. Esta especie esta cercanamente relacionadaa I icipesisstind ja (Mull. oo ) Markgr. pero puede ser separada por sus hojas con nervi I mente 170 BRIT.ORG/SIDA 21(1) por 2a 4mm, ldbulos de la corola de 1-1.5 mm de longitud y frutos globosos a subglobosos, de 9 a 12 mm de diametro. Otras especies relacionados a estos taxones por Monachino (1944), (e.g., L. peruviana) seran sinonimizadas en la — revision del género (Morales, en prep.). IA. Pando: Federico Roman, Rio Abuna, Fortaleza, back naval Abus: senda hacia la cachuela, Vargas et al. 1066 (F LPB, MO, USZ). 8. Mandevilla symphi pa(G. Meyer) Woodson, Ann. Missouri Bot. Gard. 19:70. 1932. Aunque originalmente fue conocida como restringida a las Guyanas y Trinidad (Woodson 1933), en el proceso de elaboracion de una nueva monografia del género, se ha encontrado que Mandevilla symphitocarpa es una rara especie, distribuida ampliamente en toda la cuenca baja amazonica (Morales, en prep.). Este taxa puede ser confundida con la comtn M. scabra (Hoffmansegge ex Roemer & Schultes) K. Schumann, pero M. symphitocarpa se puede reconocer facilmente por sus hojas, con la lamina foliar usualmente glabra (vs. variadamente pubescente a glabrada), flores con la parte inferior del tubo de la corola 4-5 mm de ancho (vs. 2-3 mm), foliculos 25-32 cm de largo (vs. 9-23 cm) y semillas 13-16 mm de longitud (vs. 10-12 mm), con la coma 33-37 mm de largo (vs. 14-24 mm). Especimenes representativos seg BOLIVIA. Santa Cruz: Serrania de Huanchaca, estacion los Fierros, Foster et al. 13733 (LPB, USZ). 9. Odontadenia anomala (Van Heurck & Mill. Arg.) J.F Macbr, Field Mus. Nat. Hist., Bot. Ser. 11:35. 1931. Anteriormente reportada como endémica para Pert (Morales 1999), varias colecciones de esta especie fueron localizadas en varios herbarios bolivianos, lo que ahora amplia el rango de distribucion hasta Bo- livia. Una de las especies mas distintivas en el género por sus hojas irregularmente verticiladas. Especimenes representativos examinados: BOLIVIA. Beni: Vaca Diez, Rio Beni, puerto de Florida, Moraes 580 (LPB, USF). Pando: Puerto Candelaria, Rio Madre de Dios, Moraes 528 (LPB, MO, P. SD. Santa Cruz: Velasco, Campamento La Toledo, camino hacia Florida, Guillén & Roca 2474 (USZ). 10. Prestonia lagoensis (Mtl. Arg.) Woodson, Ann. Missouri Bot. Gard. 23:296. 1936. Esta especie fue conocida por mucho tiempo solo por la colecci6n tipo, hecha por Warming en Lagoa Santa, Minas Gerais, Brasil. Sin embargo, con la futura sinonimizacion de P. lindmanii (Malme) Hoehne (Morales, en prep.), dentro de esta especie, su rango se extendera notablemente. A pesar de ser relati comun en Brasil y Paraguay, es conocida en Bolivia por menos de rres colecciones tres : Especimenes representativos examinados: BOLIVIA. Santa Cruz: Velasco, Cerro Pelao, Guillén & Surubi 1251 (MO, USZ). 11. Rauvolfia mollis S. Moore, Trans. Linn. Soc. London, Bot. Ser. 2, 4:393. 1895. Rauvolfia mollis, no reportada anteriormente para Bolivia (Rao 1956), es una especie cercanamente relacionada con R. ligustrina Willd. ex Roem. & Schult. y ambas especies pueden confundirse con facilidad, ya que los caracteres que las separan citados por Rao (1956) algunas veces tienden a traslaparse. En términos generales, R. mollis se reconoce de R. ligustrina por sus inflorescencias largamente pedunculadas y el indumento usualmente hirsuto de tallos e inflorescencias. los: BOLIVIA. Beni: Mamoré, 104 km al N de Trinidad, camino a con jeaaul n, Mowaes et al. 1521 (LPB). Santa Cruz: Velasco, San Juancito, 27 km al N de San Ignacio, Seidel & Beck 379 (LPB, MO, SI). 12. Stipecoma peltigera (Stadelm.) Mull. Arg., Fl. Bras. 6(1):176. 1860. En la monografia de Woodson (1936), Stipecoma era conocido unicamente por menos de seis colecciones, de los estados de Bahia, Goids, Minas Gerais en Brasil. Sin embargo, el incremento de la actividad botanica en dicho pais en el ultimo siglo ha incrementado notablemente el numero de especimenes de esta especie. Anteriot te endémico para Brasil, es reportado por primera vez para Bolivia. Especimenes representativos examinados: BOLIVIA. Santa Cruz: Chiquitos, La Mina, Motacuzal, Chochi, Wood & Landivar 17548 (K, LPB) 13. Vinea major L., Sp. Pl. 1:209. 1753. Esta hierba de origen europeo ha sido introducida ampliamente en los trdpicos y se puede encontrar en algunos sitios cercanos a la Paz creciendo escapada de cultivacion. Especimenes representativos examinados: BOLIVIA. La Paz: Murillo, Jardion Botanico Municipal, Solomon 15602 (LPB, MO). NUEVA SINONIMIA 1. Himatanthus sucuuba (Spruce ex Mull. Arg.) Woodson, Ann. Missouri Bot. Gard. 25:198. 1938 [1937]. Plumeria succuba Spruce ex Mull. Arg,, Fl. Bras. 6(1):40. 1860. Tipo: BRASIL. AMAZONAS: Manaus, Oct 1851 (EL), Spruce 1848 (HOLOTIPO: K, ISOTIPOS: FP Himatanthus segelloue (K. Schum. ex Markgr.) Plumel, ee ae 1991. Plumeria tarapotensis K. Schum. ex Margr., Notizbl. Bot. Gart. Berlin-Dahlem 11: 339. 1932. Tipo: ren Remate dos Males, Ducke 21607 (HOLOTIPO: - ISOTIPO: P) El basidnimo de Himatanthus tarapotensis fue incluido anteriormente en la sinonimia de H. succuba por Woodson (1938), pero fue elevado al rango de especie por Plumel (1991). Sin embargo, las diferencias mencionadas por Plumel para distinguir ambas especies (hojas oblongas en H. succuba vs. hojas elipticas en H. tarapotensis) no son consistentes ni relevantes para mantener ambos taxas como entidades diferentes, pues corresponden a leves variantes morfologicas, sin ninguna discontinuidad evidente o aislada. Ademas, las colecciones tipo no difieren en el resto de caracteres morfoldgicos (e.g., inflorescencia, flores). 2. Prestonia tomentosa R. Br, Mem. Wern. Nat. Hist. Soc. 1:70. 1811. Tiro: BRASIL. Rio DE JANEIRO: Rio de Janeiro, s.d. (fl), Banks 684 (HOLOTIPO: BM, foto en BM, INB, NY). 172 BRIT.ORG/SIDA 21(1) Prestonia cornutisepala Rusby, Mem. New York Bot. Gard. 7:329. 1927, syn. nov. Prestonia cephalantha Rusby, Mem. New York Bot. Gard. 7:330, 1927, syn. noy. T1pO: BOLIVIA. LA PAZ: Ixiamas, 17 Dic 1921 (1D, Cardenas 1928 (HOLOTIPO: NY; ISOTIPOS: K, NY, n.v.). Prestonia cornutisepala y P. cephalanta fueron descritas basadas en la misma colecci6n tipo. Prestonia cornutisepala, descrita originalmente para Bolivia, no fue incluida en la monografia de Woodson (1936). La coleccion tipo no difiere en ninguna forma del tipo de P. tomentosa, y por lo tanto debe ser sinonimizada. 3. Tabernaemontana vanheurckii Mull. Arg., Obsery. Bot. 168. 1871. Tipo: PERU, RTIN: cerca de Tarapoto, 1855-1856 (1D), Spruce 4209 (HOLOTIPO: AWH; ISOTIPOS: BM, BP, BR, C, CGE, E, G, GH, GOET, K, L, MO, NY). Tal t unguiculata Rusby Mem. New York Bot. Gard. 7:324. 1927, syn. nov. TIPO. BOLIVIA. LA PAZ: Huachi, cabeceras del Rio Beni, 21 Ago 1921 CD, White 461 (HOLOTIPO: NY; ISOTIPOS: GH, US). ~ Tabernaemontana unguiculata fue incluida en la sinonimia de T. heterophylla por Leeuwenberg (1994). Sin embargo, desptes del estudio de las colecciones tipos de ambas especies, es claro que T. unguiculata es coespecifica con T. vanheurchii y por lo tanto, debe ser removida de la sinonimia de T. heterophylla, tal y como fue previamente propuesto por Killeen et al. (1993). AGRADECIMIENTOS Queremos agradecer a los curadores y directores de los siguientes herbarios B, BM, BOLY, C, CGE, E, F G,G-DC, GH, K, M, MO, NY, LPB, P. US, USZ por el préstamo y uso de sus colecciones. El primer autor agradece al Missouri Botanical Garden (MO) por el generoso envio de material por regalo de identificacion, asi como a Stephan Beck (LPB) por la ayuda brindada en su estadia en Bolivia. También queremos agradecer a Mary Endress(Z), por ayudar a clarificar la identificacion de las especies de Malouetia presentes en Bolivia. REFERENCIAS At-SHEHBAZ, |. 1999, Lepidium beckii (Brassicaceae), a new species from Bolivia. Novon 9:5—7. CHAvez DE MicHeL, R. 1993. Apocynaceae. En: T. Killeen, E. Garcia, y $.G. Beck, eds. Guia de arboles de Bolivia. Herbario Nacional de Bolivia, La Paz, Bolivia y Missouri Botanical Garden, St Louis, USA. EzcurrA, C. 1981. Revision de las Apocinaceas de Argentina. Darwiniana 23:367-484 Ezcurra, C. 1984. Apocynaceae. En: A.L. Cabrera, dir. Flora de la Provincia de Jujuy (Rep. Argentina) 8:84-117.Col. Cient. INTA, Buenos Aires. FALLEN, M.E. 1983. A taxonomic revision of Condylocarpon (Apocynaceae). Ann. Missouri Bot. Gard. 70:149-169. Foster, R.B. 1958.A catalogue of the ferns and flowering plants of Bolivia.Contr. Gray Herb. 184:1-223 Fuentes, A. 1998. Contribucién a la flora lenosa de los bosques de tierras bajas del este de Santa Cruz; adiciones a la guia de arboles de Bolivia. Revista Soc. Boliv. Bot. 2:46-59. Gensel, W.H. 1969.A revision of the genus Thevetia (Apocynaceae). Masters thesis. Univer- sity of Connecticut [University Microfilms International, Ann. Arbor.] Gentry, A.H. 1974. Notes on Panamanian Apocynaceae. Ann. Missouri Bot. Gard. 61: 891-900. Gentry, A.H. 1984. New species and combinations in Apocynaceae from Peru and adja- cent Amazonia. Ann. Missouri Bot. Gard. 71:1075-1081. Hansen. B. 1985.A monographic revision of Forsteronia (Apocynaceae). Ph.D. dissertation. University of South Florida. U.S.A. [University Microfilms International, Ann. Arbor] Justiniano, J. & M. Toledo. 2001. Contribucién de nuevos registros a la guia de arboles de Bolivia. Revista Soc. Boliv. Bot. 3:304-316 KiLLEEN et al. 1998. Apéndice 1: Listado de plantas vasculares del Parque Nacional Noel Kempff Mercado y sus alrededores. En:T. Killeen y J. Schulenberg. A biological assess- ment of Parque Nacional Noel Kempff Mercado. RAP Working Papers 10.Washington = LeeuwenserG, A.M. 1994. A revision of Tabernaemontana. The New World species and Stemmadenia. The Royal Botanic Gardens, Kew, Richmond, England Marconbes-FerreiRA, W. 1988. Aspidosperma Mart., nom. cons. (Apocynaceae): estudos taxondmicos. Ph D. thesis, Universidade Estadual de Campinas, Brasil Meyer, E.1955.The genus Vallesia,a systematic study. Masters's Thesis, Washington Univer- sity, St. Louis, Missouri. 1-46. [University Microfilms International, Ann. Arbor.] MonacHino, J. 1943. A revision of Couma and Parahancornia (Apocynaceae). Lloydia 6: MonacuHino, J. 1944. A revision of Lacmellea and the transfer of Zschokkea (Apocynaceae), Lloydia 7:275-302. MOoNAcHINo, J. 1945 a.A revision of Hancornia (Apocynaceae). Lilloa 11:19-48. MonacHino, J. 1945 b. A revision of Macoubea and the American species of Landolphia (Apocynaceae). Lloydia 8:291-317. Moraes, M. 1996. Novelties of the genera Parajubaea and Syagrus (Palmae) from interandean valleys of Bolivia. Novon 6:85-92. Morates, J.F. 1997 [1998]. A synopsis of the genus Macropharynx (Apocynaceae). Rhodora 99:252-262. Morates, J.F. 1998. Synopsis del género Lacmellea (Apocynaceae) en Mesoamérica, con una nueva especie de Costa Rica. Novon 8:259-262. Morates, J.F. 1999. A synopsis of the genus Odontadenia (Apocynaceae) in A.J.M. Leeuwenberg (editor), Series of revisions of Apocynaceae XLV. Bull. Jard. Bot. Nat. Belg. 67:381-477. Morates, J.F. 2002. Studies in Neotropical Apocynaceae |:A revision of the genus Laubertia (Apocynaceae). Rhodora 104:170-185 Morates, J.F. 2003. Studies in Neotropical Apocynaceae Ill: A revision of the genus Secondatia, with discussion of generic classification. Candollea 58:305-319. Navarro, G. 1997. Izozogia nellii (Zygophyllaceae), nuevo género y especie del Gran Chaco de Santa Cruz. Novon 7:1—5. 174 BRIT.ORG/SIDA 21(1) PiumeL, M. 1991. Le genre Himatanthus (Apocynaceae) Révision taxonomique. Bradea 5 (supl.):1-118. Rao, A.S. 1956. A revision of Rauvolfia with particular reference to the American species. Ann. Missouri Bot. Gard. 43:253-355 SakaNe, M. y G.J. SHEPERD, 1987. Uma revisao do genero Allamanda L. (Apocynaceae). Rev. Brasil. Bot. 9:125-149. Woooson, R.E. 1933. Studies in the Apocynaceae. IV. The American genera of Echitoideae XXVI. Ann. Missouri Bot. Gard. 20:605—790, Woopson, R.E. 1935. Studies in the Apocynaceae. IV. The American genera of Echitoideae XXVI. Ann. Missouri Bot. Gard. 22:153-306. Woopson, R.E. 1936. Studies in the Apocynaceae. IV. The American genera of Echitoideae XXVI. Ann. Missouri Bot. Gard. 23:169-438. Woopson, R.E. 1938. Studies in the Apocynaceae. VII. An evaluation of the genera Plumeria L.and Himatanthus Willd. Ann. Missouri Bot. Gard. 25:189-224. Woopson, R.E. 1951. Studies in the Apocynaceae. VIII. An interim revision of the genus Aspidosperma Mart. & Zucc. Ann. Missouri Bot. Gard. 38:119-206. ZuLOAGA, F., O. Morrone, y T. Kitten. 1993. Gerritea, a new genus of Paniceae (Poaceae: Panicoideae) from South America. Novon 3:213-219. PSEUDOSTELLARIA OX YPHYLLA (CARYOPHYLLACEAE), A LONG OVERLOOKED SPECIES FROM NORTHERN IDAHO Ronald L.Hartman Richard K. Rabeler Rocky Mountain Herbarium University ea es Herbarium Department of Botany 3600 Varsity Driv University of Wyoming Ann Arbor, oe. igan a 08- ae U.S.A. Laramie, Wyoming 82071-3165, U.S.A. rabeler@umich.edu rhartman@uwyo.edu ABSTRACT cy ll ‘ p! Tl] RT Rob | } a i ss 1 5 cs fee | ] Bi eee zi 1; ture since its rie even in 1898. Our study indicates that Stellari phylla is congeneric wit : sthe new sompmatons is Sai d. The three western North American mem- 4 toll] bers of P forming a shallow dish, thus os the see oe @R.L. Hartman and Pjamesiana (Torr) WA. Weber & R.L - Hartman) in ae: internodal lines of hairs (0.5- : mm wid | 1 branches of the inf] , the stems sof F hat Cs 2) ne pase ee aoe 2 or . prONONS ] D 1 1] : £ ai > Lf . “Ot the congener widely distributed in ve western United States. Pseudostellaria oxyphylla ane ies Gena share a reduction i in stamen pubes from 10 to 5 and seeds with elongate, tuned kuperces ae with or 1 conical projec tions respectively, ll found along stream bane ofan under and near conifers. RESUMEN Se ha redescubierto Stellaria oxyphylla B.L. Rob. en nae norte de poate un taxon raramente aaa en la bibliografia desde q describio en 1898. t inGiGaale Stellanwoxvnlat congenérica con Pseudostellaria ooas 2 que se hace | una nueva eae Los tres miembros del oeste de Estados Unido a hacia atras 2 63 aes formando un plato ee eevemenee de este modo las semillas. Pseudostellaria ee difiere de P sierrae Rabeler & R.L. Hartman y P jamesiana (Torr.) WA Weber & R.L. Hartman por tener lineas de pelos internodales (0.5-5 mm de ancho) en los tallos : ramas de las floes pero como | eae peak IS tallos oe P. ae son cuadrados e seccion transversa t esiana, el congénere lao rey en di oeste de los Estados Unidos. Pecullostell arid oxyphylla y P. sierrae mparten una reduccion en el numero de estambres de 10 5, y semillas con tubérculos alargados y eae cada uno con 5 a 12 o mas glandulas estipitadas diminutas 0 proyecciones cénicas, respectivamente. Pseudostellaria oxyphylla se encuentra a lo largo de los bancos de torrentes, a menudo debajo y cerca de coniferas. Stellaria oxyphylla B.L. Rob. has rarely appeared in the literature since it was described in 1898. Most recently, the taxon was relegated to synonymy under Stellaria calycantha (Ledeb.) Bong. as CL. Hitchcock considered it “the robust extreme’ of that taxon (Hitchcock et al. 1964). Rabeler 1986) considered Stellaria oxyphylla distinct from the S. calycantha complex and placed it tentatively SIDA 21(1): 175-179. 2004 176 BRIT.ORG/SIDA 21(1) under Pseudostellaria jamesiana (Torr.) W.A. Weber & R.L. Hartman. Our re- cent collaboration on the Caryophyllaceae for Flora of North America (Hart- man & Rabeler in prep.), necessitates that we resolve the taxonomic status of this plant. Recent field work has provided new material and information on the geographic and ecological distribution of this taxon. Populations of Poxyphylla appear restricted to stream margins in the St. Joe Mountains, Kootenai and Shoshone counties, Idaho, an area where P jamesiana is absent. It likely repre- sents a taxon of conservation concern but intensive field studies are warranted. After studying the available material of Stellaria oxyphylla again, espe- cially in light of our recent description of Pseudostellaria sierrae Rabeler & R.L. Hartman and its relationship to P. jamesiana (Rabeler @ Hartman 2002), we conclude that these species are congeneric. We here make the following new combination, Pseudostellaria oxyphylla (B.L. Rob.) R.L. Hartman & Rabeler. The description, illustration (Fig. 1), and geographic and ecological notes supple- ment the only other published account of the species—the type description. We investigated the relationship between the North American and Asian species of Pseudostellaria when P. sierrae was described (Rabeler @ Hartman 2002), an overview of the genus can also be found in that article. At least one synapomorphy unites the western United States members of Pseudostellaria: six capsular valves that dehisce by rolling back tightly 2 or 3 revolutions, forming a shallow dish exposing the basal placentae with | or 2 seeds that may persist, at least briefly, prior to dispersal. In general appearance, P oxyphylla is most simi- lar to P jamesiana. Pseudostellaria oxyphylla and P sierrae share a reduction in stamen number from 10 to 5 and have seeds with elongate, rounded tubercles each having 5 to 12 minute stipitate glands or conical projections, respectively. All three taxa have rhizomes with occasional axillary buds, but unlike P. oxyphylla, P jamesiana has tuberous thickenings of the rhizomes, while P sierrae has tuberous, cigar-like thickened roots. Considerable excavation of the rhizomes of several plants of Poxyphylla did not reveal additional perennating structures. oe Sy few fully mature seeds are preserved for these taxa on her- barium spe Ihy true for Pjamesiana. Unlike Psierrae (northern C alifornia) and P jamesiana a (widespread), with geographical ranges that overlap in part, P oxyphylla appears to be disjunct from the latter taxon. Pseudostellaria oxyphylla (B.L. Rob.) R.L. Hartman & Rabeler, comb. nov. (Fi 1). BASIONY™: Stellaria oxyphylla B.L. Rob., Bot. Gaz. (Crawfordsville) 25:165, pl. B. fig. 5, 1898. Alsine oxyphylla (B.L. Rob.) A. Heller, Cat. N. Amer. pl. ed. 2, p. 4. 1900. Type: U.S.A. IDAHO. KOOTENAI Co.: on St. Joseph’s River [?], Wiesls]ner’s Peak, Sandberg, MacDougal, and Heller 608, 8 Jul 1892 (£D, 1,800 m (HOLOTYPE: GHI; ISOTYPES: CAS! NY! PH!, POMI, US). Perennial, mostly glabrous herb, erect to sprawling, 20-25 cm tall, with stems arising singly, at varying intervals, along rhizomes. Roots vertical to spreading, filiform, 1-8 cm long or more; rhizomes often extensive, branched, whitish to 177 Fic.1.P | lari hylla.A.and B., habit, the stem inA.d hed f het ith rhi to the rigdhtin B.C YPM) 7 ' 7 g ct. . a - hed. F. Capsule, dehisced, with sut li lyx. G. Upper intern- g Caly Pp Flower. D. Sepal E oP | > L + ee eH ‘4 1 rr r tan, square in cross section to rounded with age, 0.5-1 mm in diameter, shiny, internodes 0.1-2 cm long or more, when internodes contracted achlorophyllous bracts often dense, overlapping, axillary buds 2-3 mm long. Stems mostly simple, square in cross section, internodes 1-5 cm long or more below, 1-2.5cm long in inflorescence, glabrous except for a dense, internodal line of pubescence, 0.5-5 mm wide, trichomes straight to curly, most recurved, uniseriate of 5-9 178 BRIT.ORG/SIDA 21(1) elongate cells (possibly somewhat glutinous as evidenced by debris adhering to older material). Leaves opposite, sessile, the primary leaves 8-11 pairs on main l late to lance-elliptic, ascending to spreading, 6-12 x 0.7-1.2 cm, little reduced above, exstipulate, bases connate with transverse margin 0.3-0.6 mm wide, leaf surface flat, often pustulate (30x), margins often revolute, smooth to granular or sometimes papillate, sparsely ciliate proximally, midrib often sparsely pubescent adaxially, smooth, shiny, and often prominent abaxially. Inflorescence consisting of paired flowers, or one abortive, on pedicels often terminating secondary branches in the distal 3-7 axils, progressively reduced upward from pairs of normal leaves to scarious bracts, lanceolate to linear-lan- ceolate, 2-10 mm long; pedicels 1.5-4.5 mm long. Flowers perfect, chasmo- gamous, regular; sepals, inner lanceolate, outer narrowly ovate, 6.5-7 x 1.5-2.5 mim, glabrous, margins narrowly scarious, ciliolate in proximal half, midrib and lateral pair of nerves obscure, becoming evident in fruit, acute to acuminate, in fruit proximal 1/5 of sepals often with a thickened, elliptical patch; petals 5, white, strap-shaped, expanding to twice the width distally, 8-9 x 15-2 mm, broadly notched, notch 0.8-1 mm deep, the lobes rounded; stamens 5, each with a circular, thickened, basal gland 2-2.5 times the filament width, alternating with the petals, filaments 4-4.5 mm long, anthers yellow; ovary 3-carpellate, with 6-12 ovules; styles 3, 3.5-4 mm long, stigma terminal and adaxial, linear, minutely papillate (0x). Capsule ovoid, 4-4.5 mm long; valves 6, dehiscing by rolling back tightly 2-3 revolutions, forming a shallow dish exposing the basal placentae and plump funiculi. Seeds 1-2, reddish-brown, 2.5-2.6 mm long, cir- cular, plump, radicle prominent, mature seeds with most tubercles broadly coni- cal to elongate, rounded, interdigitating, each with 8-12 or more stipitate glands ca. 0.015 mm long or less (50x). During the August visit to Pine Creek, most of the inflorescences were brown; most of the fruit aborted. This is likely a consequence of the dry summer. Similarly, it is very difficult to find fruiting material and seeds of Pseudostellaria jamesiana, in part because the fruits and seeds frequently abort or mature late in the fall and thus are seldom collected. While this species has been known for over a century, to our knowledge it had been collected only twice, and never in fruit. The only collection subse- quent to the type was gathered by J.H. Christ in 1940 (Christ 10949, NY). In Au- gust of 2000, Hartman relocated it and returned in June of 2001 to obtain addi- tional material. The known geographic range appears restricted to the St. Joe Mountains and vicinity: extreme west-central Shoshone and southeastern Kootenai counties, Idaho. stem, Specimens examined: U.S.A. Idaho. Kootenai Co.: St. Joseph River [?], Wiesls|ner’s . ak, 1,800 m., 8 Jul 1892, Sandberg, ecco & Heller 608 (HoLoTyPE: GH; isoTyPes: CAS, NY, PH, POM, US); Pine Creek, se before Middle Fork Pine Creek, on jeep trail to Mount Wiessner, T47N RIE S15, ie 800 ft., 25 Aug 000, Hartman oe (MICH, MONT, RM); Pine Creek, 0.5 air mi NE of Middle Fork Pine Creek, T47N RIE $12, 2,700 ft., 25 Aug 2000, Hartman 70410 (MICH, RM), 12 Jun 2001, Hartman 71073 (ID, MICH, NY, RM, UC, WTU). Shoshone Co.: cliffs along St. Joe River, 7 mi N of Calder, 19 May 1940, Christ 10949 (NY). The only floristic work that treats Stellaria oxyphylla as a recognized taxon is the unpublished “Manual of the Flora of Northern Idaho” by Carl Epling and Joseph Ewan dated 1941 (page 258 of 1121 page manuscript, original at MO; cop- ies at RM, UC, provided by Alan Whittemore). Several locations were visited along the St. Joseph River, including the seg- ment in the vicinity of Christ’s collecting site, but Pseudostellaria oxyphylla was not relocated. The habitat is given as “cliffs,” perhaps at their bases. Ac- cording to the late Douglass Henderson, “Christ's labels are to be viewed as “prob- ably inaccurate” (letter to Rabeler, Sep 1987). After Hartman discovered populations of the taxon along Pine Creek, south of Pinehurst, he hiked the switchbacks from the southwest to the saddle, ca. 6,000 feet elevation, between Latour Peak (6,408 feet) and Mount Wiessner (6,185 feet). Neither suitable habitat for nor populations of Pseudostellaria oxyphylla was encountered on the slopes and drainages. ECOLOGY Pseudostellaria oxyphylla appears restricted to banks along perennial streams and rivers and adjacent moist sites; often under conifers or at the edge of conif- erous forests. Due in part to an often extensive rhizome system, 20 to 100 or more flowering stems were found scattered in these sites that often dry out over the summer. On Pine Creek, the scattered populations were found over a 2.5 mile stretch of the creek valley. ACKNOWLEDGMENTS We wish tothank Carolyn Crawford for the illustration and the curators of CAS, GH, NY, PH, POM, and US for loans to Rabeler, ID, MONT, MONTU, WS for loans to Hartman, and CAS, MO, and UC for visits by Hartman and Rabeler. REFERENCES HitcHcock, C.L., A. CRONQuisT, M.Ownsey, and J.W. THompson. 1964. Vascular plants of the Pacific Northwest. Part 2. Univ.Wash. Publ. Bot. 17(2):1-597. RageLer, R.K. 1986.Revision of the Stellaria calycantha (Caryophyllaceae) complex and taxo- nomic notes on the genus. Ph.D diss., Michigan State Univ., East Lansing. Rasecer, R.K.and R.L. HARTMAN. 2002. Pseudostellaria sierrae (Caryophyllaceae),a new species from California. Novon 12:82-86. Rosinson, B.L. 1898. New species and extended ranges of North American Caryophyllaceae. Bot. Gaz. (Crawfordsville) 25:165-171. 180 BRIT.ORG/SIDA 21(1) BOOK NOTICE Rosert K. WEBSTER, GEORGE BRUENING, WILLIAM O. Dawson, and NEAL K. van ALFEN (eds). 2003. Annual Review of Phytopathology: Volume 41, 2003. (ISBN 0- 8243-1341-0, hbk; ISSN 0066-4286). Annual Reviews Inc., 4139 El Camino Way, P.O. Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www. AnnualReviews.org, 800-523-8635, 650-493-4400, 650-424-0910 fax). $165.00 (USA), $170.00 (Int'l), 704 pp., 6" x 9" Contents of Volume 41 of Annual Review of Phytopathology: Perspectives on Plant and Soil Nematology James Gordon Horsfall: Nonconformist and Founding Father Carl Freiherr von Tubeutf: Pioneer in Biological COntrol of Plant Diseases Epidemiology and Management of Tomato Spotted Wilt in Peanut Brome Mosaic Virus RNA Replication: Revealing the Role of the Host in RNA Virus Replication Cultural and Genetic Approaches to Managing Mycotoxins in Maize Regulation of Antibiotic Production in Pseudomonas spp. and Implications for Biological Control of, Plant Disease The Threat of Plant Pathogens as Weapons against U.S. Crops Gibberella from A(venacea) to Z(eae) Evolution of Wheat Streak Mosaic Virus: Dynamics of Population Growth Within Plants May Explain Limited Variation Molecular Basis of Pto-Mediated Resist to Bacterial Speck Disease in Tomato Parasitic Nematode Interactions with Venamels ata Blanks Ecology of Mycorrhizae: A Conceptual Framework for Complex Interactions Among Plants and Fungi Advances in Molecular-Based Diagnostics in Meeting Crop Biosecurity and Phytosanitary Issues Development of Alternative Strategies for Management of Soilborne Plant Pathogens Currently, ontrolled Through Methyl Bromide Patterns of Pesticide Use in California and the Implications for Strategies for Reduction of ou Pesticides Innovations In Plant Pathology Teaching Of Smuts, Blasts, Mildews, and Blights: cAMP Signaling in Phytopathogenic Fungi The Ecological Significance of Biofilm Formation by Plant-Associated Bacteria oe Sensing | in Plant-Pathogenic Bact ity — Plant Pathogenic Mollicute Relationships with its Two Hosts, the Plant and the he, Leafhopper Vector Sp Pathogen Self-Defense: Mechanisms to Counteract Microbial Antagonism Luteovirus-Aphid Interaction Ecology and Epidemiology of ee and Plasmodiophorid Vectors The Potential of Optical Canopy Measurement for Targeted Control of Field Crop Diseases Engineering of Transgenic Plant Nematode Resistance Establishment of Biotrophy by Parasitic Fungi and Reprogramming of Host Cells for Disease, Resistance SIDA 21(1): 180. 2004 THE GENUS PRENANTHES (ASTERACEAE: LACTUCEAE) IN TEXAS Jason R. Singhurst Robert J.O’Kennon Wildlife Diversity Program Botanical Research Institute of Texas Texas Parks and Wildlife Department 09 Pecan Street Austin, Texas 78704, U.S.A. Fort Worth, Texas 76102-4060, U.S.A. Walter C. Holmes Department of Biology Baylor University Waco, Texas 76798-7388, U.S.A. = ABSTRACT Tae pECe of Prenantnes occur in Texas pignantne scare seen O’Kennon, & Holmes is de- Plateau vegetational region. Prenanthesaltissimais known only sinners and Newton counties in extreme southeastern Texas. Prenanthes barbata is more widespread but rare in rich hardwood and pine-oak forests in the Piney woods of eastern Texas, with one record in the Post Oak Savannah of Lamar County. A key to species, descriptions, distribu- tion map, and lists of exsiccatae are included. RESUMEN En Texas hay tres especies de Prenanthes. Se describe Prenanthes carrii Singhurst, O’Kennon, & Homes como especie nueva endémica de la region de vegetacion de Edwards Plateau. Prenanthes ce tnicamente de los condados de Jasper y Newton en el extremo sureste de Texas. Prenanthes barbata es mas frecuente pero raro en los bosques de madera dura y de pino-roble en los de con una cita en la sabana de Post Oak del condado de Lamar. Se incluye una clave de especies, descripciones, mapas de distribucion, y listas de exsicatas. INTRODUCTION Correll and Johnston (1970) recognized two species of Prenanthes L. in Texas. One, P. altissima L. was identified from three specimens all collected the same year from the same location in Newton County. Prenanthes barbata (T. & G.) Milstead ex Cronquist was identified from two specimens, one each from Jas- per and Nacogdoches counties. By 1993 knowledge of the genus in Texas was based upon seven additional collections of P. barbata and two additional col- lections of P. altissima. In 1993 and 1994 Singhurst (1996) conducted a status survey of P. barbata in Texas that increased the understanding of its distribu- tion and habitat restrictions. That survey prompted the present expanded study. The purpose of the present paper is to (1) describe a new species of Prenanthes from the Edwards Plateau vegetational region of Texas, (2) report the additional information regarding the Texas distribution of P. altissima and P. barbata, (3) provide a key to distinguish the three species in Texas and (4) SIDA 21(1): 181 - 191. 2004 182 BRIT.ORG/SIDA 21(1) remark on the rarity and abundance of all species treated. Field research and collections made between 1993 and 2003 form the primary basis of this study. The majority of specimens collected are deposited in the Baylor University Her- barium (BAYLU). Other specimens from ASTC, BRIT, LAMU, Rice University Herbarium, Houston Texas (hereafter cited as RICE), SBSC, TAES, TAMU, TEX- LL, US, and the personal herbarium of Steve Orzell and Edwin Bridges were examined and annotated. Cultivation studies of both P. barbata (1993-1994) and P. altissima (1998) were conducted at the Stephen F Austin Arboretum and Tucker Estate, Nacogdoches, Texas. The new species was cultivated in 2003 at the residence of O’Kennon in Fort Worth. Though limited, these cultivation studies provided growth form information that was otherwise unavailable. TAXONOMIC TREATMENT Prenanthes L., Sp. Pl. 797-798. 1753. Nabalus Cass., Dict. Sc. Nat. 34:94, 1825. Perennial herbs with milky juice and tuberous roots. Leaves alternate, simple and entire to toothed to few-lobed or deeply cleft, or lower ones sometimes sev- eral-foliate. Capitula mostly numerous in an elongate, erect and often nodding paniculiform capitulescence; flowers ligulate, perfect, 5-15 in number, white, yellow-white, to greenish-white; involucres cylindric, of 4-8 principal phyllar- ies, the outer phyllaries (bracts) much reduced; receptacles epaleate. Cypselae cylindric or nearly so, glabrous, mostly reddish-brown, multicostate; pappus of numerous deciduous capillary bristles. Prenanthes is a genus of approximately 40 species (Rao & Datt 1996) with a predominantly north a epee” except for one species of south central Africa (Milstead 1964). are recognized. Subgenus Nabalus includes all species of North America and northeast Asia, wile the European and African species are in subgenus Prenanthes. Classification of the genus is complicated by frequent hybridization and by extreme morphological varia- tion (Cronquist 1980). Plants of the genus are com y known as rattlesnake root, cankerweed, or gall of the earth. The name aa is from Greek prenes (drooping) and anthe (flower). This name refers to the nodding habit of the capitula. KEY TO THE SPECIES OF PRENANTHES IN TEXAS 1. Capitula 5-6 flowered; involucres with 5-6 principal phyllaries, glabrous P.altissima . Capitula 10- 15 baila involucre with 6-8 principal phyllaries, pubescent with long coarse h 2. Lower and oe m leaves shallowly or deeply pinnately lobed, the upper lan- ceolate to oblong; petioles usually shorter than the blades; capitulescence race- mose to slightly paniculate P.barbata SINGHURST ET AL., A NEW SPECIES OF PRENANTHES FROM TEXAS 183 2. Lower and midstem leaves sagittiform, the upper ovate to triangular-deltate; petioles usually longer than the blades, capitulescense paniculate P. carrii Prenanthes altissima L., Sp. Pl. 797.1753. Nabalus altissimus (L.) Hook., FI. Bor. Amer. 1:294. 1833. TyPE: LINN; IDC microfiche 177.536.HL1. Prenanthes altissima L. var. cinnamomea Fern., on 10:95. 1908. Type: U.S.A. MISSOURL Monteer, 5 Oct 1905, Bush 3534 (HOLOTYPE: GH Perennial herbs from a thick and knotty corm-like taproot. Stems erect to 200 cm tall, glabrous, striate, the upper portions not branched or rarely so, leafy towards the base and sparse to leafless in the immediate vicinity of the capitulescence. Leaves ovate to triangular, 3-5 lobed, 2-14 x 15-12 cm, the lower ones sometimes withered before anthesis but can also be present; venation pin- nate, with 4-6 pairs of prominent secondary nerves separating from the midvein at an angle of ca. 45’, the nerves arcuate, tertiary and quartenary nerves reticu- late, surfaces glabrous to villous; bases widely cordate to an obtuse insertion at the petioles, margins dentate, the teeth 0.5-1 cm distant, apices acute to acumi- nate; petioles 0.5-8.5 cm long, wingless throughout its length; upper leaves re- duced in size; deltoid to reniform, shortly petiolate, margins slightly dentate. Capitulescene paniculate, heads disposed in loose axillary clusters in upper axils, 20-28 cm tall. Capitula slender, 12-15 mm long; peduncles to 2-3 mm long, puberulent; bracts (outer phyllaries) lanceolate to ovate, 1-2 mm long, apices acuminate, margins puberulent; primary phyllaries 5, 10-12 mm long, lanceolate, green to lilac, apices acuminate, glabrous. Florets white, 5-6 in num- ber, 10-11 mm long, tubes ca. 6 mm long, ligules 5 mm long, 5-nerved, teeth 5, linear, sparingly glandular, surfaces glabrous, anthers + mm long; stigmatic surfaces densely hispid to setose. Cypselae obscurely 5-angled, 5 mm long, golden-brown. Pappus bristles ca. 40-50, yellow-brown, ca. 8-9 mm long, mar- gins finely scabrid. Distribution.—Known only from Jasper and Newton counties in extreme southeast Texas (Fig. 1); also Quebec to Maine, S to Georgia, Tennessee, Kentucky, Arkansas, and Louisiana. Phenology.—Flowering from late August through October. ene pone TEXAS. Jasper Co.: Sally Withers Lake area, Autumn 1974, Watson s.n. (RICE) Newton Co. 1414 about 3 mi SE of Burkeville, 23 Mar 1969, Correll 36864 (TEX-LL); FR 1414 about 3 mi SE - Burkeville, 15 Oct 1969, Correll 38172 (TEX-LL); FR 1414, about 3 mi SE of Burkeville, 29 Oct 1969, Correll & Correll 38217 (TEX-LL); Scrappin Valley, 25 Oct 1974, Watson s.n.; Scrappin Valley Preserve, Temple Inland Property, 12 mi NNE of Burkeville, 2 Apr 1998, Singhurst G Watson 6548 (BAYLU); jct. FM 645 and Hickman Creek, W on Hickman Creek 0.4 mi, 3 Apr 1998, Singhurst & Watson 6559 (BAYLU); Canyon Rim Trail, Tex. Hwy 87, 0. 5 s N of Mayflower Community, 30 Sep 2001, Holmes 11997 & Singhurst (BAYLU); Louisiana Pacific Corp. Nature Trail, 4 mi N of jet. of Tex. Hwy 87 and R255 on Hwy 87, 23 Sep 2003, Singhurst ve (BAYLU); Tex. Hwy 87, 2.1 mi S of jet. with Tex. Hwy 63, SE on Co. Rd. 2.7 mi to Simms Cemetery, Yellow Bayou bluffs and banks, 23 Sep 2003, Singhurst 12494 (BAYLU). 184 BRIT.ORG/SIDA 21(1) A a a» C ry A rN a £4 ry e a @ Sle Fic. 1. Distributi f th p, hes inT p thes altissima WH, P barbata A, and P. carrii @. — Prenanthes altissima is known from seven locations in the northern portion of Jasper and Newton counties. These locations are the southwestern limit of P. altissima’s distribution in Texas. Outside Texas, the nearest documented occur- rences are approximately 120 km distant in Evangeline and Rapides parishes, Louisiana (Thomas & Allen 1996). The nearest Arkansas population (Smith 1979) is about 320 km distant. In Texas the species occurs on primary and sec- ondary terraces along stream banks of small perennial spring-fed creeks and banks of larger streams with flood plains that daa h ardwood forests. Prenanthes altissima is associated with Fagus grandifc alia grandiflora, Acer barbatum, Quercus alba, Hamamelis virginiana, iH. ve rnalis, ee ben- zoin, Lilium michauxii, lipularia discolor, Sanguinaria canadensis, Polygonatum biflorum, Polystichum acrostichoides, Thelypteris hexagonoptera, Trillium grac- ile, Spigelia marilandica, Solidago auriculata, and S. caesia. SINGHURST ET AL., A NEW SPECIES OF PRENANTHES FROM TEXAS 185 Cronquist (1980) recognized two varieties of Prenanthes altissima, var. altissima with whitish to pale brown pappus, occupying the greatest part of the distribution of the species, and var. cinnamomeaof Missouri, Arkansas, Loui- siana, and, though not cited by Cronquist, presumably Texas. This variety is characterized by bright yellow-brown or almost orange pappus. Gandhi and Thomas (1989) used this varietal name to refer to the Louisiana forms of this taxon. However, Thomas and Allen (1996) merged var. cinnamomea with var. altissima in the Atlas of the Vascular Flora of Louisiana. Color variation of the pappus does not appear to justify recognition at the varietal level. Prenanthes barbata (Torr. & A. Gray) WL. Milstead ex A. Cronquist. Brittonia 9223. 1977. deena eee DC. var. barbatus Torr. & A. Gray, Fl. N. Amer. 2:481. 1843. P aria Pursh var. barbata (Torr. & A. Gray) A. Gray, Syn. Fl. N. ichey 12): 434, 1886. Nabalus barbatus (Torr. &@ A. Gray) Heller, Muhlenbergia 1:8. 1900. Nabalus serpentaria Pursh var. barbatus (Torr. & A. Gray) Mohr, Contr. US. Natl. Herb. 6:755. 1901. TyPE: U.S.A. ALABAMA: Buckley s.n. (SYNTYPE: GH; ISOSYNTYPE: NY). The GH specimen is here designated as lectotype. The other syntype cited, Macbride s.n. from the Saluda Mountains of [Polk Co., North Caro- lina], was not located but from location seems to refer to P. serpentaria Pursh. Perennial herbs from thick corm-like taproots, occasionally connected by slen- der rhizomes. Stems erect, 50-150 cm tall, simple below, glabrous, the upper portions often branched, glabrate to pilose to hispid to semiarachnose to vil- lous, leafy except in the immediate vicinity of the capitulescence. Leaves oblan- ceolate to spatulate, 5-20 x 1.5-5 cm, the lower ones usually withered before anthesis, venation pinnate, with 4-6 pairs of prominent secondary nerves sepa- rating from the midvein at an angle ofc Cana eearers the nerves straight to arcuate, tertiary and quartenary ner late, surf labrate t ingly puberulent-pilose; bases attenuate, the lower to long winged petioles, margins entire to denticulate to dentate to lacerate-parted (especially lower leaves), the teeth 1-2 cm distant, apices acute to rounded; petioles 0.5-6.5 cm long, winged for most of its length; upper leaves reduced in size; elliptic to oblanceolate, sessile to shortly petiolate, margins mostly entire to denticulate to less ly den- tate to lacerate-lobed at the bases. Capitulescene racemose to paniculate, 5-36 cm tall. Capitula cylindric to semicampanulate; peduncles to ca. 1 cm long, glabrate to tomentose to villous; bracts (outer phyllaries) lanceolate to narrowly ovate, 2-5 mm long, apices acuminate, margins subentire, midribs with few to numerous coarse bristles; primary phyllaries 6-8, 13-16 mm long, linear-ob- long to narrowly lanceolate, green to purplish, apices acute to rounded to an acute point, finely puberulent, occasionally with glandular hairs, margins en- tire, midribs with few to numerous coarse bristles. Florets white, 10-12 in num- ber, 12-13 mm long, tubes ca. 5-6 mm long, ligules 5-6 mm long, 5-nerved, teeth 5, linear, sparingly glandular, surfaces glabrous, anthers 5-5.5 mm long; stig- matic surfaces setose-hispid. Cypselae obscurely 5-angled, 5-5.5 mm long, 186 BRIT.ORG/SIDA 21(1) golden-brown. Pappus bristles ca. 50, white to yellow-brown, ca. 8 mm long, margins scabri Distribution.—Eastern Texas from Hardin County north to Lamar County (Fig. 1); also Alabama, Arkansas, Georgia, Kentucky, Louisiana, and Tennessee. Phenology.—Flowering from late August through November. Specimens examined. TEXAS. Angelina Co.: Comp. 94, Angelina National Forest, ca. 0.7 mi S of junc- tion of FS Rd 303 and FS Rd 302 on FS Rd 303, jet. of FS Rd 303 and Big Creek, 11 Nov 1994, Singhurst 3609 (BAYLU). Cass Co.: ca. 0.4 air mi SW of jet. of Tex. Hwy ll and Tex. Hwy 8 at Linden, 24 Oct 1994, Singhurst 3404 (BAYLU). Cherokee Co.: ca. 0.5 air mi SE of jet. of U.S. Hwy 69 and FM 241, 17 Nov 1994, Singhurst 3610 (BAYLU). Hardin Co.: W side of Silsbee near Mill Creek, Rd 327, 22 Sep 1971, Watson 766, 767, & 768 (RICE); W of Silsbee on Tex. Hwy 327 near Mill Creek, 29 Sep 1971, Amerson 806 (SMU). Jasper Co.: U.S. Hwy 96, ca. 2.0 mi S of Jasper, 10 Oct 1946, Lundell & Lundell 14674 (TEX), ca. 3.5 mi SW of Jasper on FM 777, 4 Nov 1982, Agilvsgi 8270 (TAMU); S of Walnut Run Creek on U.S. Hwy 96, 1.8 mi S of jet. U.S. Hwy 190 and 96 in Jasper, 29 Oct 1993, Singhurst 3401 (BAYLU); roadside park ca. 1.6 mi S of U.S. Hwy 190 and 96 in Jasper, 13 Oct 1982, Cheatham s.n. (TEX). Lamar Co.: FM 906, 1.4 mi E of jet. of FM 906 and US. Hwy 271 at Mid City, 18 Oct 2002, Singhurst & Harris 11345 (BAYLU). Nacogdoches Co.: ca. 8.0 mi NE of Nacogdoches, 4 Oct 1941, Parks 1373 (TEX), 1372 (SMU); Tex. Hwy 21, 100 yards W of Loco Bayou, ca. 0.4 mi E of Co. Rd. 829, 21 Oct 1993, Carr 13246 (TEX-LL); Little Loco Bayou, ca. 9.0 mi W of Nacogdoches on Tex. Hwy 21, ca 2.1 air mi ESE of Winter Hill, 1994, 14 Oct 1993, Singhurst 3406 (BAYLU); ca. 1.95 mi W of jet. of FM 95 and FM 1878, 5 Sep 1994, Singhurst 3407 (BAYLU), ca. 9.2 mi NNE of Nacogdoches on U.S. Hwy 59, 4.5 mi SSW of Garrison on U.S. Hwy 59, 1 Nov 1993, Singhurst s.n. (BAYLU); ca. 4.2 mi NE of jet. of Loop 224 and FM 1878 in ies on FM 1878, Carrizo Estates, 5 Sep 1994, Singhurst 3408 (BAY LU); ca. 6.9 NW from jet. of U.S. Hwy 59 and FM 343 on FM 343, ca. 0.9 air mi ESE of Winter Hill, 5 Sep 1994, Singhust 3601 a v a ees W end of fe Valley Drive, 5 Sep 1994, Singhurst 3602 (BAYLI Lake, ca. 1.5 SW of Barnes Lake Dam, 5 Sep 1994, Singhurst eee U). Newton oe jet. of ee Sear i y 190,7 Mar 2002, Singhurst 11202 (BAYLU) Co.: Morgan Creek, FM 1988, 2.6 mi SW of jet. Tex. Hwy 146 and FM 1988, 27 ae 1994, we ea Peers .U). Rusk Co.: ca. 2.5 air mi NNE of jet. of Tex. Hwy 322 and U.S. Hwy 2 24 Oct 1994, Singhurst 3402 (BAYLU). Sabine Co.: Comp. 66, Sabine National ee ia Hills ca. 4.4 mi NE of jet. of Tex. Hwy 21 and FM 3153.14 Aug 1994, Si ae es (BAYLU); Comp. 69, Sabine National Forest, ca. 1.3 mi E of jet. of FS Rd 131 and FS Rd 131-A on FS Rd 131-A, ca. 2.1. air miS of jet. of U.S. Hwy 87 and FS Rd 131, 14 Aug 1994, Singhurst 3607 (BAYLU); a 72, Sabine National Forest, ca. 3.4 mi from jet. of Tex. Hwy 2land U.S. Hwy 87, ca. 0.5 air mi SW of Red Hills Lake, 14 Aug 1994, Singhurst 3608 (BAYLU), Tex. Hwy 21, 5.9 road mi E of jet. FM 225 at pla 12 Oct 1993, Carr 13246 (TEX-LL). San Augustine Co.: Comp 65, Sabine National Forest, ca. 0.8 mi on FR that enters Comp. 65 from W, 14 Aug 1994, Singhurst 3604 (BAYLU), ca. 3.5 air mi NNE of jet. “ol Attoyac oe and Tex. ou 21, 14 Aug 1994, Singhurst 3605 (BAYLU); Spring Ridge on Arenosa Creek, ca. 12 mi WNW of San Augustine, 29 Sep 2001, Holmes & Singhurst 11970 (BAYLU). Shelby Co.: sin Na- tional Forest, ee RD, Compartment 51, ca. 1.4 air mi NNW of jet. of FM 1279 and Tex. Hwy 147, 1+ Aug 1994, Singhurst 3400 (BAYLU); Comp. 51, Sabine National Forest, ca. 2.2 mi aa jet. of FM 1279 and FM 147 on FM 147, 22 Apr 1987, Orzell & Bridges 5126 (pers. herb.). The distinctness of Prenanthes barbata was first recognized but never formal- ized by Milstead (1964). The name was subsequently adopted by Correll and Johnston (1970) and gained general acceptance; it was formalized by Cronquist in 1980. SINGHURST ET Al OF PRENANTHES FROM TEXAS 187 The species was considered rare by Correll and Johnston (1970) because it was known only from Jasper and Nacogdoches counties. Turner et al. (2003) presented an expanded distribution in east central Texas that included six coun- ties. Turner et al. also used the name to refer to plants of the Edwards Plateau which, in this study, are considered a distinct species. Prenanthes barbata oc- curs in mesic ravine slope forests that are dominated by Fagus grandifolia- Quercus alba series (Diamond et al. 1987) and on mesic hardwood sites with Quercus shumardii, 9. muhlenbergii, Q. alba, Q. michauxii, Pinus taeda, Carya ovata, and C. myristicaeformis. Prenanthes carrii J.R. Singhurst, RJ. O’Kennon, & WC. Holmes, sp. nov. (Fig.2). YP TEXAS. BANDERA Co. Lost Maples State Natural Area, above and be- low Upper Forks of Mystic Canyon Trail, 1 Sep 2003, Singhurst & Singhurst 12496 (HOLOTYPE: BAYLU). AP. } Bak a a [a Dich : Le ty $31, a FE aa y > Perennial herbs from tuberous tap roots with similar side roots. Stems simple, erect, occasionally rather long branched in area of inflorescence, 80-150 cm long, strigose to tomentose in upper half, scattered strigose to glabrous basally. Lower leaves sagittiform (ovate, ovate-deltate to broadly elliptic in general con- tour), 13-25 x 7-12 cm, light green, chartaceous, venation pinnate, with 3-5 pairs of prominent secondary nerves separating from the midvein at an angle of about 45 degrees, nerves straight to slightly antrorsely curved, tertiary and quartenary veins reticulate; surfaces glabrate to lightly puberulent-setose especially on nerves, bases attenuate, truncate to widely cordate, occasionally deeply pin- nately divided at base near petioles, margins coarsely and irregularly dentate, mostly 1-2 cm distant, teeth subspinose-mucronate, apices acute to rounded: petioles 2.2-13 cm long, broadly winged for half or more of its length by the decurrent leaf bases; upper leaves reduced in size, elliptic, occasionally nearly sessile or with winged-decurrent petioles to about 1 cm long, otherwise similar to lower leaves; bracteal leaves reduced in size, less prominently dentate to oc- casionally subentire. Capitulescence paniculate, 20-50 cm tall; capitula cylin- dric to semicampanulate; peduncles, 4-9 mm long, tomentulose, with 5-12 or more linear-subulate to narrowly lanceolate hispid bracts (outer phyllaries) 2- 4 mm long; primary phyllaries 8, 9-11 mm long, mostly lime green to pinkish- rose to lavender, linear-subulate to lanceolate, apices acuminate, outer surfaces glabrate except for the sparingly hispid midrib, apical margins minutely cili- ate. Florets 9-11 per capitulum; 11.5-13.5 mm long, tubes +-7 mm long, ligules ca. 7 mm long, 5-nerved, teeth 5, linear-oblong, 0.07-0.1 mm long, apical sur- faces papillate; anthers ca. 5.5 mm long; stigmatic surfaces densely hispid-se- tose. Cypselae irregularly angled to more often terete, ca. 6.3 mm long, golden yellow to tan, prominently 12-15 costate. Pappus bristles white to tan to yellow, 7-8 mm long, 30-40, margins scabrid. 1) BRIT.ORG/SIDA 21( A ~~ Se . = SS a t i ee Z Pee en tea | B. upper bracteal leaves, C. middle leaf, D. b f. E. florets. F pappus, G Fic. 2. Prenanthes carrii: A. habit (BAYLU) J} SINGHURST ET AL., A NEW SPECIES OF PRENANTHES FROM TEXAS 189 Distribution.—Southwest Edwards Plateau (Bandera, Gillespie, Kerr, and Real counties) of Texas (Fig. 1.) Phenology.—Flowering late August to November. Paratypes: TEXAS. Bandera Co.: Lost Maples State Natural Area, Upper Can Creek, 17 Nov a Singhurst 8533 (BAYLU); Love Creek Preserve, The Nature Conservancy of Texas, 2 Nov 2002, ls een cee otyre: BAYLU). Gillespie Co.: 7 mi N of Harper inrich creek canyon o sa One 993, O’Kennon 11914 (TEX-LL). Kerr Co.: along rocky spring branch, Lacey’s Ranch, 3 Oct 1916, iene ot specific peas 13 Oct 1940, Parks s.n. (TAES); 12 mi S of Kerrville along Lamb Creek, Oct 1993, O’Kennon 11898 (TEX-LL); 5.3 mi N of jet. of Kerr/Bandera County line on Tex. Hwy 16, Upper Lamb Creek, 17 Nov 1 a Singhurst 8534 (BAYLU), cultivated [propagules from Upper Lambs Creek], 18 Oct 2002, O’Kennon s.n. (BAYLU). Real Co.: Lost Maples State Natural Area, 27 Oct 2001, Singhurst 11526 (BAYLU); upper reaches of Mill Creek, Grey Wolfe Ranch, 2 Nov 2002, Singhurst 11554 (BAYLU). Prenanthes carrii is morphologically similar to P. crepidinea Michx.,, P. alata (Hook.) D. Dietr, P. sagittata, (A. Gray) A. Nels., P. bootii (DC.) D. Dietr, and P. barbata. This group was proposed as a new ease ction in Milstead’s 1964) un- published (thus never formalized) dissertation. The group is characterized by paniculate to racemose capitulescences, leaves at least short petiolate, flowers white to creamy and usually numbering 7-38 per capitulum, inner phyllaries 6-15, and outer phyllaries (bracts) mostly 7-13 per capitulum. Leaf character- istics of P. carrii resemble the sagittiform shaped leaves of P. alata, P. crepidinea, and P. sagittata. The new species appears to be most closely similar to P. barbata, particularly in the paniculate nature of the capitulescence. The two species may be distinguished by the characters referenced in the key. Additional traits dis- tinguishing P. carrii from P. bar bata include its taller height of 80-150 cm, stri- gose to tomentose vestiture, and primary phyllaries 9-11 mm long. Prenanthes barbata is usually 55-125 cm tall, has tomentulose to subarachnose vestiture, and has primary phyllaries 11-14 mm long. Prenanthes carrii occurs primarily in rich soils in woodlands at the upper reaches of canyons where springs flow due to geologic contacts. Where these contacts occur, there is a vegetation transition between the species of Quercus laceyi, Q. muhlenbergti, Q. texana, and Acer grandidentatum and the creekside seepage shelves dominated by Platanus occidentalis, Cephalanthus occidentalis Adiantum capillus-veneris, Thelypteris kunthii,and Cladium mariscoides. The species is normally associated with Aristolochia serpentaria, Carex edwardsiana, C. planostachys, Lindera benzoin, Senecio obovatus, and Verbesina virginica. Other central Texas endemics occurring with or near P. carrii include Chaetopappa effusa, Clematis texensis, Matelea edwardsensis, Philadelphus texensis, Tragia nigricans, Tridens buckleyanus, and Styrax platanifolius var. stellatus. Etymology.—The species is named in honor of William F. Carr of The Na- ture Conservancy of Texas. Bill is deeply committed to preserving the botani- 190 BRIT.ORG/SIDA 21(1) cal heritage of Texas and is currently one of the most active plant collectors in the state. ACKNOWLEDGMENTS We wish to thank the curators of the herbaria cited for access to the specimens used in this study. Special recognition is given to Geraldine Watson who ac- companied the senior author to the sites where she had previously collected Prenanthes. Amber Swanson, Curatorial Assistant of the Harvard University Herbaria, provided a digital photograph of the designated lectotype of Prenanthes barbata. We thank Guy L. Nesom for his Latin translation and for reviewing the manuscript. Appreciation is also expressed to the landowners who graciously permitted access to their properties and to Bill Carr for his field assistance and access to Love Creek Preserve. We also give special thanks to David Riskind at Texas Parks and Wildlife Department who granted us access to Lost Maples State Natural Area and reviewed the preliminary manuscript. We also want to thank Larry Brown of Spring Branch Science Center Herbarium and Sandi Elsik and Warrens Pruess of Rice Herbarium for their assistance with specimen records. We also extend our deepest gratitude to Linny Heagy. Her amazing artistry exemplifies the details of the newly-described Prenanthes carrii. —_— REFERENCES Corrett, D.S. and M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner Cronauist, A. 1980. Vascular flora of the southeast United States. Vol. 1. Asteraceae. Univ. North Carolina Press, Chapel Hill. Diamono, D.D., D.H. Riskino, and S.L.Orzett. 1987. A framework for plant community classifi- cation and conservation in Texas. Texas J. Sci. 39:203-221. GANDHI, K.N. and R.D. THomas. 1989. Asteraceae of Louisiana. Sida, Bot. Misc. 4. Botanical Re- search Institute of Texas, Fort Worth. Musteap, W.L. 1964.A revision of the North American species of Prenanthes. Unpub.disser- tation, Purdue University. Rao, R.R. and B. Darr. 1996. Diversity and phytogeography of Indian Compositae. In: DJ.N. Hind, ed.-in-chief, Composite: systematics. Proceeding of the international Compositae conference, Kew, 1994. Royal Botanic Gardens, Kew. Pp.445-461. SinGHURST, J.R. 1996. The status of nine endangered plants of East Texas: historical, ecologi- cal and phytogeographical notes. Unpub. thesis, Stephen F. Austin University, Nacogdoches, Texas. Smith, E.B. 1979. An atlas and annotated list of the vascular plants of Arkansas. Published by the author. Fayetteville. THomas, R.D. and CA. Aten. 1996. Atlas of the vascular flora of Louisiana. Vol. II. Dicotyle- SINGHURST ET AL., A NEW SPECIES OF PRENANTHES FROM TEXAS 191 dons. Acanthaceae-Euphorbiaceae. Louisiana Department of Wildlife and Fisheries Natural Heritage Program, Baton Rouge. Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida, Bot. Misc. 24. Botanical Research Institute of Texas, Fort Worth. 192 BRIT.ORG/SIDA 21(1 = BOOKS RECEIVED SLavoko Matic (Editor in Chief). 2003. Obiéna Bukva (Fagus sylvatica L.) U Hrvatskoj (common beech, Fagus sylvatica L. in Croatia). (ISBN 953098571-1-2, hbk.). Akademija Sumarskih Znanosti and co-published with Hrvatske Sume d.o.0 Grad Zabreb, Gradski ured za Poljoprivredu I Sumarstvo, Zagreb Croatia. Price not given, 855 pp., b/w photos, color photos, maps, drawings, graphs, tables, 61/2" x 91/2", JoserH H. KIRKBRIDE, JR., CHARLES R.GUNN, and ANNA L. WEITZMAN. 2003. Fruits and Seeds of Genera in the Subfamily Faboideae (Fabaceae). Volume 1 and volume 2.US.Department of Agriculture, Technical Bulletin No. 1890.U.S. Department of Agriculture, National Technical Information Service, 5285 Port Royal Road, Springfield, VA 22161, U.S.A. (Orders: 703-605-6000, 306-966-5567 fax). Price not given. 1212 pp., b/w photos, drawings, keys, descriptions, 81/2" x RICHARD “THE BUGMAN” FAGERLUND AND JOHNNA STRANGE. 2004. The Bugman on Bugs: Understanding Household Pests and the Environment. (ISBN 0-8263-3363- X, pbk.). University of New Mexico Press, 1720 Lomas Blvd. NE, | Univer- sity of New Mexico, Albuquerque, NM 87131-0001, U.S.A. (Orders: 800-249- 7737, 505-277-9270 fax, unmpress.com). $14.95, 131 pp., 17 halftones and 30 b/w illus. 5 1/2" x 9" Howarp Scott Gentry. 2004. Agaves of Continental North America. (ISBN 0-8165- 2395-9, pbk.). The University of Arizona Press, 355 S. Euclid, Ste. 103, Tucson, AZ85719. (Orders: 520-621-1441, 520-621-8899 fax, www.uapress.arizonaedu) $49.95, 670 pp., 422 illustrations, 7" x 10". TRACEY PARKER. 2003. Manual of Dendrology Jamaica. (ISBN 976-610-504-9 pbk.). Forestry Department, Ministry of Agriculture, 173 constatn Spring Road, Kingston 8, Jamaica (West Indies) (Orders: 876- ek 2667, 876-924-2626 fax, www.orestry.gov.jm, email: forestrydepat y.govjm ). +494 pp., 21/2" x 8" —_ SIDA 21(1): 192. 2004 TWO NEW SUBSPECIES OF MICROSERIS LACINIATA (ASTERACEAE) FROM THE SISKIYOU MOUNTAINS Kenton L. Chambers Dept. of Botany & Plant Pathology Oregon State University 2082 Cordley Hal Corvallis, Oregon 97331, U.S.A. chamberk@science.oregonstate.edu eo ABSTRACT Ee Tee | sos 1 . . . | } re J sho af 1 f the f ‘loristi- cally “central” region of the ae Mountains a Orton and adjacent California. These taxa have novel morphological features differentiating them from the related Microseris laciniata subsp. laciniata and subsp. leptose pala also found in this region. The populations are kept separate by allo- patry and by adaptation to different edaphic conditions, and thus have different vegetation associa- tions. Intermediate populations occur where pairs of taxa meet and the ecological differences are modified by habitat disturbance. RESUMEN RAS oe | I _ 4 : j ] Se describen siskiyouensis y subsp floristica “central” as las Montanas Siskiyou de pOtenen y la adyacente Came Estos taxa fenea caracteristica | laciniata y subsp. leptosepala nie abe se encuentran en esta region. Las poblaciones se manbenes eee por alopatria y por adaptacion a condiciones edaficas diferentes, y asi tienen diferentes asociaciones de vegetacion. Aparecen poblaciones intermedias donde contactan pares de taxa y las diferencias ecoldgicas se modifican por alteraciones del habitat INTRODUCTION The perennial species of Microseris subgen. Scorzonella were last reviewed taxo- nomically by Chambers (1957) in preparation for the generic treatment in Abrams and Ferris’ Illustrated Flora of the Pacific States (Chambers 1960). An attempt was made to compartmentalize the extensive variation of M. laciniata into two subspecies, subsp. laciniata and subsp. leptose pala, based principally on the shape, width, and pubescence of the involucral phyllaries and on differ- ences in distribution. The species ranges from western Washington south through Oregon to just north of San Francisco in California, principally west of the Cascade Range but extending east to Klamath and Lake Counties, Oregon, and Siskiyou, Lassen, and Modoc Counties, California. The subsp. leptosepala occurs in the more coastal parts of the Coast Range, especially in the Klamath Mountains region, but is also in northwestern Oregon including the lower Wil- lamette Valley. The subsp. laciniata is more interior, being the common form SIDA 21(1): 193 - 205. 2004 194 BRIT.ORG/SIDA 21(1) from Pierce County, Washington south to Douglas County, Oregon and from southern Humboldt County to Sonoma County, California, as well as in the Great Basin part of the range. Intergradent forms are frequent, especially in the Wil- lamette Valley and Curry County, Oregon, the northern California Coast Range, and in populations east of the Cascade Range. In this earlier study, additional puzzling variation was found in the Siskiyou Mountains of southwestern Oregon and adjacent California, but it could not be resolved using the few available herbarium specimens from this rugged and poorly explored region. This variation was lumped into Microseris howellii, which we now know to be a narrowly endemic species which is limited to ex- posed serpentine sites in the Illinois River Valley of Oregon. Field research by the present author suggested that two additional forms of M. laciniata should be recognized taxonomically from the Siskiyou region, and their ranges have recently been mapped through the collecting efforts of a number of interested colleagues (see Acknowledgments section). These entities have been presented informally (Chambers 1993) but have not yet been validly published. Some diagnostic features of these new entities were presented by Mauthe et al. (1982) and compared with 9 other samples of Microseris laciniata repre- senting much of the species’ range. Capitulum and fruit morphology was the principal object of this investigation. It was suggested that there was “a rather small number of unlinked genes allowing for a rather free recombination of character states” and that “the characters that distinguish these two groups of populations (subsp. laciniata and subsp. leptose pala) recombine freely” in some Sedat Crossing experiments (Pires 1980) show that hybrids between taxa n the M. laciniata complex are no less pollen-fertile than intrataxon crosses. i important result of these and previous studies of M. laciniata (Bachmann and Price 1979) was the validation of pappus part number as a highly consis- tent morphological feature of these two named subspecies. In both, the number varies among cypselae of single heads but is always constrained from 5 to 10. Frequencies of average pappi numbers follow patterns of rather simple Poisson distributions between 5 and 10,as would result from the interaction of a mini- mum of two pairs of genes (Bachmann & Chambers 1978 Both of the newly recognized subspecies have average pappi numbers that are above 10-up to 16.6 in subsp. detlingii and 20.3 in subsp. siskiyouensis. In other respects, these two new subspecies are very different in morphology and in edaphic and vegetation associations. The newly available suite of collections of these taxa shows that each has an area of intergradation with an adjacent subspecies of M. laciniata—subsp. detlingii with subsp. laciniata, and subsp. siskiyouensis with subsp. leptosepala. Hence they are parts of a single genetic complex and are best placed as subspecies rather than as separate species. Their addition to the flora of the Siskiyou Range means that this region is an unusu- ally rich center of diversity for perennial Microseris, containing 5 recognized Nar CHAMBERS, NEW SUBSPCUICS UF 195 taxa. These taxa have defined distribution patterns and characteristic habitat preferences, which allow them to persist as separate populations in this lim- ited geographic region. Microseris es (Hook.) Sch. Bip. subsp. siskiyouensis K.L. Chambers, subsp . 1, 2, 3). Type: US.A. CALIFORNIA. DEL Norte Co.: Hwy. 199, 5.0 mi by oa the Smith R. from Patricks Ck. Guard Station, on a rocky diabase knob overlooking the river, in shallow soil covered by mosses, lichens, and Selaginella sp., with Melica harfordii, Luina hypoleuca, Sedum sp., 10 Jul 1964, K.L. Chambers 2242 (HOLOTYPE: OSC; ISOTYPES: MO, NY, UC). "1: Microseride laciniato subsp. leptose palo (Nutt.) K.L.Cham| i is pappi 10-24 varians setis minute barbellatis differt; chromosomatum numerus 2n = -18, Perennial herbs with 1-3 fleshy biennial taproots; stems to 65 cm high, well- branched above, except on depauperate plants, leafy, the floral peduncles aris- ing terminally and from leaf axils; leaves linear or narrowly lanceolate, attenu- ate, sometimes sparsely toothed or pinnatifid with narrow lobes; involucres 10-17 mm high, witha series of lanceolate inner phyllaries and a variable num- ber of shorter outer ones ranging from linear to lanceolate-attenuate or del- toid-attenuate (Fig. 1B shows the narrowest type of outer phyllaries), both outer and inner series usually densely furfuraceous (that is, covered with the minute scaly trichomes that are a generic feature of Microseris), inner phyllaries usu- ally lightly black-villous dorsally and minutely white-strigulose ventrally; flo- rets 13-48+ in number, with yellow ligules 15-22 mm long; cypselae pale to dark brown, 3.5-5.5 mm long, 10-ribbed, the ribs usually smooth or lightly scabrous, often lightly hispid near the apex on outer fruits; pappi scales 10-24, 0.5-2.0 mm long, lanceolate to linear-lanceolate, white or brownish, bristles minutely barbellate, white. In the region under discussion, subsp. leptosepala and subsp. siskiyouensis are practically indistinguishable in habit. Therefore, Figure 1 can represent ei- ther taxon. Cypsela and pappi are illustrated in Figure 2, with enlargements of the pappi to show the bristle differences from the scabrous-awned subsp. leptosepala, left, and the plumose-awned species Microseris nutans (Hook.) Sch. Bip. on the right. The bristle difference with subsp. leptosepala breaks down where the taxa are sympatric in western Curry Co., Oregon and Del Norte Co., California. Here both subspecies have barbellate bristles, and they are distin- guished only by pappi number—5-10 per cypsela in subsp. leptosepala, 10-24 per cypsela in subsp. siskiyouensis. Distribution. —Microseris laciniata subsp. siskiyouensis is limited to the Siskiyou Mountains in Del Norte and Siskiyou counties, California and Josephine and Jackson counties, Oregon. The most common habitat is in grassy openings of second growth woodlands, in non-serpentine soil or well-developed forest soil over serpentine bedrock. It also occurs on non-serpentine rock outcrops, as at the type locality. Frequently associated woodland species are Pseudotsuga BRIT.ORG/SIDA 21(1) 196 siskivouensis. A. Habi I Fic. 1. Mi CHAMBERS ZaaTITIT OE LLL Ls eNOS SANNA 3 a , : — eins aeeced LLL LLL SO tk 5-9 ESSAY 3mm —<—— aa! SS ————— oy 7 Fig. 2. Cypselae and pappi of various Microseris taxa. A. Cypsela of M. laciniata subsp. siski, is bearing 15 pappus parts. B. Pappus part of M. laciniata subsp. lacpataiel or subsp. /eptosepala, with scab bristle. C. P D. P. sae subsp. siskiyouensis, with minutely t sela of M. laciniata subsp. detlingii bearing 12 pappus parts. F. Pappus part of M. laciniata subsp. detlingii, with mi- t of M. nutans, with plumose bristle. E. ae barbellate bristle. BRIT.ORG/SIDA 21(1) 198 124°30' 124° 123°30' 123° 122730" 4s | | | CU RRY fy a COOS DOUGLAS i | | : es ple --r x be 1 7 = pore a j por ur | ie “OSEAN JACKSON / arnt oma \ ome 30 1 R. GP Rogue Gfp oo é Le X 00 1 O°. X ! gp fe) My GS ' M ed & e J ce) ay ° ed ay Ps ° 7 : ° 0 “fy | x/e e “e R wy ae a U ‘o 0 o 9° 4° B \e/ £(* , e 7 Oregon e A ce lg — 30’ Fic. 3. Distribution of Mi Si Black circles = M. laciniata subsp. siskiyouensis; open circles = M. /aciniata subsp M. laciniata subsp. laciniata. Cities: B= Brookings; cc Microseris howellii is omitted. , 2 GaldaR h- Cp ¥ tc D AA RMaodf, nd NVI D MN MICQIOTU. J t C +t Clty: CR yr Aatil, UE lS Fidos, CHAMBERS 199 menziesii, Pinus lambertiana, P. jeffreyi, Calocedrus decurrens, Quercus garryana, Q. kelloggii, Arbutus menziesti, Arctostaphylos canescens, Rhododen- dron macrophyllum, and Ceanothus integerrimus. Elevations are 30-2100 m. Flowering.—May-Jul, depending on habitat and elevation. Figure 3 maps the known populations of subsp. siskiyouensis, represented by black circles, while the known localities for subsp. leptosepala in the Siskiyous are shown as open circles. The pattern of parapatry and the region of contact in western Josephine County are evident. Two sites have been identified, marked by small Xs, where intermediate populations are known, containing many plants with 10 or fewer pappus parts and others with average pappi numbers in the range of subsp. siskiyouensis. One such population, at Gasquet Flat, Del Norte County (see Chambers 5168, below) was well sampled by Mauthe et al. (1982), who reported the average numbers of pappi in a sample of 149 plants. The fre- quencies of plant averages, grouped by whole numbers, are as follows: 9-10: 56; 10.1-11: 55; 11.1-12: 15; 12.1-13: 8 13.1-14: 8 14.1-15: 6, 15.1-16: 1. Although the ge- netics of pappi numbers are not known, one can speculate that this is a popula- tion of subsp. leptosepala showing a history of hybridization with subsp. siskiyouensis. The second site with an intermediate population, West Side Road in Josephine County (Chambers 5522, below), though not as well sampled, shows a similar range of pappi numbers between the two sub select plants in 10 populations of subsp. siskiyouensis more remote from subsp. leptosepala had the following average pappi numbers: 13.2, 15.0, 15.2, 15.5, 15.6, 16.8, 17.0, 17.3, 17.6, and 20.3. The large Xs in Figure 3 mark populations assign- able to subsp. laciniata; these are discussed below under subsp. detlingii. For comparison Collections of Microseris laciniata subsp. siskiyouensis mapped in Figure 3. U.S.A. CALIFORNIA. De qb Co.: Old Gasquet Toll Road by Middle Fork of Smith R., 1 Jun 1935, Tracy 11223 (UC); Li of Monumental, 13 Jun 1936, nee 15220 (UC); French Hill, 2 mi S of Gasquet, 20 Jun 19 ae ae (UC); intersection of Hayne’s Flat Road, Coon Mtn., ae 1950, Tracy 18991 (UC); head i- Blue Ck. 1 mi E of lnm Ree 19fu 1950, Tracy 19136 (UC); Smith R. on Hwy. 199, 9.7 mi NE of Hwy. LO1, 5 Jul 1961 DS); Pine Flat, 26 Jun 1938, iio see QGEPS); Old Gasquet Toll Road near eighiceauniie Ck. 26 6 n es VanDeventer 167 JEPS); Gordon Mtn. N of Big Flat, 24 Jun 1952, Munz 17729 (NY); Crazy Peak area on Road 49906-053, 1 Jun 1997, Paetzel & Bell s.n. (OSC), Rd. 40503 S of Waldo, Six Rivers Natl. For. 1 mi S of border of Siskiyou Natl. Forest, 30 Jun 1973, Denton 2916 (OSC, WTU); Old Gasquet Toll Road 2.7 mi up from Smith R. bridge at Gasquet, 11 Jul 1964, Chambers 2246 (OSC); Old Gasquet Toll Road 4.7 mi up from Smith R. bridge at Gasquet, 1] Jul 1964, Chambers 2247 (OSC). Siskiyou Co.: Klamath R. 2 mi N of Swillup Ck. Ranger Station, | Jun 1942. Stebbins & Beetle 3273 (UC). OREGON. Jackson Co.: Collins Mtn. near Steamboat, 13 Jul 1950, Whittaker 184 (WS): summit slopes of Big Red Mtn., 15 Jul 1950, Whittaker s.n. (WS). Josephine Co.: Old Gasquet-O Brien Toll Road 10.1 mi NE of Patrick Ck. Road, 8 Jun 1962, Breedlove 3193 (DS); north- ern ae nts Sppeilkc hg tion, 9 Jun 1962, Breedlove 3241 (DS), Waldo Junction, 18 May 1930, Kildale Takilma, 24 Jun 1918, Peck 7955 (GH, WILLU); Murphy Ck. near Murphy, 12 Jul 1950, ‘Whittaker 155, 159 (WS); Illinois R. Valley SW of Cave Junction, to E of West Side Rd., 11 May 1989, Brock 242 (OSC), valley of East Fork of Illinois R. by French Flat, 1 Jun 1988, Kagan 6018801 (OSC); Illinois R. Valley, West Side Road S of Cave Junction, 4 Jun 1991, Chambers 5609 (OSC); Illinois R. Valley, Rockydale Rd. 2.0 mi N of Waldo Rd. E of O’Brien, 15 Jun 1998, Chambers 6113 (OSC), Waldo 3 BRIT.ORG/SIDA 21(1) Hill Lookout Rd. L6 mi S of Waldo, 15 Jun 1998, Chambers 6123 (OSC); junction of Waldo Hill spur road with road to Sanger Pk., 3 Jul 1965, Chambers 2364 (OSC). Collections of Microseris laciniata subsp. leptosepala mapped in Figure 3. U.S.A. CALIFORNIA. Del orte Co.: Gasquet Flat, alluvial flat in mixed evergreen woodland by the Smith R., TI7N, R2E, $20, 24 Jun 1984, Chambers 5168 (OSC). OREGON. Curry Co.: 10 mi N of Carpenterville, 7 Jul 1939, Peck 20450 (WILLU); Brookings, 11 Jul 1919, Peck 8790 (WILLU), Rogue River 5 mi below Mule Ck., 21 Jun 1917, Peck 3502 (WILLU), Snow Camp Meadows, 3 Jul 1929, Leach 2244 (ORE); Mine Cabin, Collier Trail, 28 Jun 1929, Leach 2277 (ORE); near Agness, 25 Jun 1933, Leach 4428 (ORE); Waldeens, 29 Jun 1934, eee sae vgs Rock, 30 Jun 1934, Leach 4701 (ORE); Vulcan Peak, T39S, RIT W, S15, 23 Jun 1980, Hess s.n.(O gness road, 2 mi W of Illinois R. junction with Rogue R., 18 Jun 1984, Stansell s.n. (OSC); on he E ie Gold Beach, 23 Jun 1982, Stansell s.n. (OSC): Pine Point Forest T375, RI3W, SI8, 27 Jun 1974, Hawk s.n. (OSC); above Wren Cabin, 1375, R14W, S12, 28 Jun oo Ritieniouse 1480 (OSC); 10 km S of Gold Beach, T37S, RI4W, S24, 16 Jun 1980, Sundberg 1098 (OSC); Fairview Meadow, T37S, R12W, 518, 11 Jul 1981, Chambers 4872 (OSC); Gold Beach to Agness road 1.6 mi W of Illinois R. bridge, 23 Jun 1984, Chambers 5162 (OSC). Josephine Co. Eagle Gap, || mi W of Selma, 23 Jun 1930, Leach 2897 (ORE, WILLU):7 mi W of O'Brien, T40S, RLOW, $26, 14 Jun 1990, Kagan s.n. (OSC); Hlinois R. road, 1375, ROW, $32, 31 May 1988, Kagan 5318801 (OSC): Illinois R. v ae W of Selma, T37S, ROW, $23, 19 Jun 1969, White/Lillico 266 (OSC); old road to Buckskin Pk RIOW, 524, HH Jul 1989, Rolle 256 (OSC); Bolt Mtn. 9 mi SW of Grants Pass near Applegate R. 9 i 1996, Mazzu s.n. (OSC). — _— Poe (Hook.) Sch. Bip. subsp. detlingii K.L. Chambers, subsp. nov. (Figs. 2, 4, 5). Type: U.S.A. OREGON. JACKSON Co: Siskiyou Pass, S side on the old ae where it joins Hwy. 5, 2.1 mi N of Hilt exit, grassy openings in Quercus breweri/Amelanchier pallida brushland, in heavy clay soil on slope above road, 22 Jun 1967, K.L. Chambers 2868 (HOLOTYPE: OSC; ISOTYPES: BRIF-SMU, CAS, MO, NY, RSA, UC, US, WS, WTU). Microseride | sIm111 lis ac ua marginibus foliorum plerumque integris caule non ramoso radice fodsies maine segmentis pappi numeris 9-19 varians squamis +-9 mm setis minute barbellatis differt; chromosomatum numerus 2n = 18. Perennial herbs with 1-2 much elongated fleshy biennial taproots; stem erect, to 55 cm high, usually simple, leafy near the base; leaves lanceolate or oblan- ceolate, acute to attenuate, tapering below to a clasping, winged petiole, gla- brous, entire or rarely sparingly dentate or pinnatifid, margins often undulate; head single ona terminal, naked or bracteate peduncle, a second peduncle some- times arising from the axil of a lower leaf; involucres 13-25 mm high, the inner series of phyllaries equal, lanceolate, often black-villous dorsally, the outer phyl- laries imbricate in several series, broadly lanceolate to elliptic or round, cuspi- date to acuminate, glabrous, sometimes purple-spotted, the outermost 3-7 mm wide; florets 18-85+, with yellow ligules 18-22 mm long, often purple-striped dorsally; cypselae 5-9 mim long, gray to brown, 10-ribbed, ribs smooth or hispid on outer fruits; pappi scales 9-19, 4-9 mm long, lanceolate, silvery (brownish on herbarium specimens), bristles minutely barbellate. Distribution.—Microseris laciniata subsp. detlingii isendemic toa limited area east and south of Medford and Ashland, Jackson County, Oregon, extend- ing north to near Butte Falls and south over Siskiyou Pass to the California state CHAMBERS, NEW SUBSPECUIES UF Fic. 4. Mi is laciniata subst detlingii A. Hat is. B. Pressed head showing phyllaries. Tcm 202 BRIT.ORG/SIDA 21(1) 123° 422° 30' | | t me | 42° | 30! | | | | | | | | | | | I i | | | | 2 ls : i 15 2 { 1s =] , i Oregon we S| aye sisKiYou California \ : 20 kan a | Fic. 5. Distributi f Mi j I Jack OR k Co., CA. Black circles = M. laciniata subsp. detlingii; Xs = M. laciniata subsp. laciniata. Cities: A = Ashland; B= Butte Falls; E= a Point; G = Gold Hill; H = Hornbrook; M = Medford; R = Ruch. Dashed line = Interstate Highway 5. line (Fig. 5). One population has been found in adjacent Siskiyou County, Cali- fornia. The subspecies occurs only in montmorillonite clay soil, sticky when wet and hard and cracked when dry, on grassy slopes and openings in shrublands and forest edges. The geology of the type area, south of the summit of Siskiyou Pass, was included in the thesis of Richard Carlton (1972), who iden- tified the underlying rocks at the type locality as fossil-bearing claystones and siltstones of the early Eocene Colestin Formation, lacustrine in origin and pos- sessing clay minerals of the montmorillonite-mica type. Farther north, near Ashland and Medford, similar clay soil develops in younger Eocene deposits of volcanic-derived sedimentary rocks (McKnight 1971). The complex geology of this region also includes sandstones and volcanic lahar and ash-flow deposits. Adaptations of subsp. detlingii to this unusual substrate include an exception- ally deep, slender taproot (Fig. 4) and the ability to reproduce clonally by ad- ventitious buds on the lateral rootlets. Colonies are often limited to patches of CHAMBERS 203 loose soil turned up by gophers or squirrels. Associated species are Pinus jeffreyi, Quercus breweri, Q. garryana, Amelanchier pallida, Ceanothus cuneatus, Arc- tostaphylos viscida, Toxicodendron diversifolium, Festuca idahoensis, Achnatherum lemmonii. Elevations are 600-1450 m. Flowering occurs May- Jun. Etymology.—The name is in honor of Prof. LeRoy Detling, longtime curator of the University of Oregon herbarium, whose 1950 collection first alerted the author to the peculiar features of this plant. Figure 5 shows the limited distribution of subsp. detlingii, as well as the nearby occurrence of populations, marked by Xs, which the author places in subsp. laciniata. The latter specimens, listed below, occur on substrates other than the “heavy clay soil” or “rocky clay soil” consistently mentioned on the labels of subsp. detlingii specimens. North of Medford, subsp. laciniata is found on rocky alluvium, as at the Agate Desert (Chambers 3080), and differs from subsp. detlingii in having pinnate leaves, multiple arched-ascending stems from the base, consistently LO pappi, and lacking an unusually elongate taproot. However, the pappi scales are up to 4 mm long, nearly twice the usual range of subsp. laciniata, and are barbellulate. On Kanutchan Creek just north of Little Butte Creek, collections by Greenleaf (1418, 1435) contain both subsp. detlingii and plants with highly pinnatifid leaves and basal branching like that of subsp. laciniata at Agate Desert. The habitat is described as Quercus garryana/Pinus ponderosa/ Arbutus menziesii woodland. We expect that more intergradation will be found between the two subspecies in this area, depending on soil type and disturbance, for example by erosion and cattle grazing. A pappus member of subsp. detlingii is shown in Figure 2B, where it is compared with both subsp. ceuuaaie and with the common ie in subspp. leptosepala and laciniata. Snes of A aseris lacini 1 in Figure 5. U.S.A. CALIFORNIA. Siskiyou d. W ik Fen Gate Reservoir 12 mi N of Klamath R. bridge at Iron Gate Dam, 16 Jun 1998, hae ber rs a 32 sade we OREGON. Jackson Co.: High hills opposite Ashland, Jun 1889, Howell s.n. (MIN, MSC, ORE, UC, US); slope of Grizzly Peak, 17 Jul 1913, Peck 7667 (WILLU); Klamath Hwy. 7 mi SE of Ashland, a 1927, Peck 15000 (WILLU); S slope of Siskiyou Mtns. near California a ee 1930, Henderson 13256 (ORE); mae Pass, T41S, R2E, $8, 11 Jun 1950, Detling 6635 (DS, ORE, UC, WTU); Dead Indian Memorial Rd. E of Ashland, T39S., R2E, $5, 23 May 1995, Straw 3274 es ante Top RNA, T355S, RIE, S15, 25 Jun i Wineteer s.n. (SOC); Kanutchan Ck., T35S., RLE, $27, 31 May 1983, Greenleaf 1435 (OSC); Dead Indian Memorial Rd. E of Ashland, T385, R2E, 524, 9 Jul 1982, Kagan s.n. (OSC); S of Butler Ck., hills N of Ashland, 9 May 1993, Brock 486 (OSC); W slope of Roxy Ann Pk. E of Medford, 30 May 1993, Brock 496 (OSC); Lick Ck., T36S, RIE, Sl, Brock 797 (OSC); Heppsie Mtn., T37S, R2E, S2, Brock 807b (OSC); Lick Ck., T35S, R2E, 529, 14 Jun 2001, Knapp 614001 (OSC); Old Siskiyou Pass Rd., 3.5 miS of summit at junction with Hwy. 5, 29 May 1965, Chambers 2348 (OSC); Old Siskiyou Pass Rd. 0.7 mi N of junction with Hwy. 5, 22 Jun 1967, Chambers 2869 (OSC); Old Siskiyou Pass Rd. 2.7 mi N of junction with Hwy. 5, 22 Jun 1967, Cham 7 rs Se (OSC); Siskiyou Pass summit on the old road, T40S, R2E, $32-33, 13 Jul 1978, Chambers 4524 ); Siskiyou Pass, S side, on aban- doned stretch of old road ca. 0.25 mi N of California state line, 16 Se 1998, Chambers 6131 (OSC). Collections of Micro: | ta sub lin Figures 3and 5.U.S.A. OREGON. Jack- by rer 204 BRIT.ORG/SIDA 21(1) son Co.: Sam's Valley N of Medford, 4 May 1930, Henderson 12388, 13253 (ORE); 5 mi W of Fish Lake, T36S, R3E, $35, 27 Jul 1989, Rolle 280 (OSC); Kanutchan Ck. ca. 4 mi E of Eagle Point, 26 May 1983, Greenleaf 1418 (OSC); Sam’s Valley, T35S, R2 W, $32,6 May 1961, Cham bers 1601 (OSC), 2.5 mi E of Hwy. 62, White City N of Medford, 16 May 1971, Chambers 3044 (OSC); Agate Desert N of Medford, Kirtland Rd. 1 mi W of Table Rock Rd., 16 May 1971, Chambers 3080 (OSC). Josephine Co.: Fish Hatchery Rd. 0.7 mi W of New Hope Rd. S$ of Grants Pass, 14 Jun 1998, Chambers 6105 (OSC). DISCUSSION The Klamath Region, including the Siskiyou Mountains, has long been recog- nized as an area of high endemism and as a center of floristic diversity (Whittaker 1961). Factors favoring this diversity, mentioned by Whittaker, in- clude a steep climatic gradient from the coast inland, high rainfall and moder- ate temperatures, much-dissected topography, and diversity of bedrock, and hence of soil types. The complex geological history and origin of the varying substrates are described by Coleman and Kruckeberg (1999). Types of endemism and an analysis of the endemic flora were presented by Smith and Sawyer (1988). These authors list the two taxa newly described here, whose names were avail- able on herbarium annotations, and also Microseris howellii,a previously named endemic of serpentine barrens in the Illinois River valley of Josephine County, Oregon. This species, and the other Microseris taxa mentioned above, illustrate very well the importance of edaphic and climatic factors in keeping separate the parapatric members of this complex. Proceeding from west to east, Microseris laciniata subsp. leptose pala occu- pies the more coastal region of Curry County and is found on serpentine bar- rens as well as non-serpentine meadows and forest edges. The peculiar vegeta- tion and characteristic flora on serpentine barrens in the Siskiyous are discussed in Coleman and Kruckeberg (1999). Examples of serpentine sites among the specimens of subsp. leptosepala cited above are Pine Point (Hawk s.n.), Gold Beach to Agness road (Chambers 5162, Stansell s.n.), Signal Buttes (Stansell s.n.), S of Gold Beach (Sundberg 1098), Buckskin Peak (Rolle 256), and 7 miles W of O’Brien (Kagan s.n.). Endemic to serpentine barrens farther east in the Illinois River valley is M. howellii, a close relative of M. laciniata, having 5-10 pappi but differing in its pappi scales 3-5 mm long. Parapatric in Oregon with M. howellii is M. laciniata subsp. siskiyouensis, which avoids open, rocky serpen- tine barrens but occurs in adjacent forested sites in loam soil. No hybrid popu- lations have been noted between these two taxa. To the east, M. laciniata subsp. laciniata is on alluvial and deeper loam soils in grasslands and mixed oak wood- lands near the Rogue River. Finally, M. laciniata subsp. detlingii is endemic to montmorillonite clay soils from 600-1450 m elevation in the Medford-Siskiyou Pass area. The differentiation of Microseris taxa in the Siskiyous, and their mainte- nance as genetically separate populations, has involved both an adaptation to different substrates and a geographical separation into different climatic zones. CHAMBERS 205 This has led to an unusual richness of species and subspecies in this limited region of southwestern Oregon and adjacent California, which is in line with the frequently mentioned floristic diversity of the Klamath-Siskiyou Mountains in general. ACKNOWLEDGMENTS The author gratefully acknowledges the help of the following colleagues in vari- ous aspects of this study: Konrad Bachmann, Richard Brock, Richard Carlton, John Dilles, Jimmy Kagan, Linda Mazzu, John Megahan, Bruce Rittenhouse, Wayne Rolle, Veva Stansell, and Richard Straw. REFERENCES BACHMANN, K. and K.L. CHampers. 1978. Pappus part number in annual species of Microseris (Compositae, Cichoriaceae). Pl. Syst. Evol. 129:119-134. BacHMANN, K. and H.J. Price. 1979. Variability of the inflorescence of Microseris laciniata (Compositae: Cichorieae). Pl. Syst. Evol. 131:17-34 CarLTon, R.W. 1972. Stratigraphy, petrology, and mineralogy of the Colestin Formation in southwest Oregon and northern California. Ph.D. Thesis, Oregon State Univ., Corvallis. CHameers, K.L. 1957. Taxonomic notes on some Compositae of the Western United States. Contr. Dudley Herb. 5:56-68. CHamBers, K.L. 1960. Microseris. In: Abrams, L.R. and R:S. Ferris, eds. Illustrated flora of the Pacific States. Vol. IV. Stanford Univ. Press, Stanford, CA. Pp. 554-562. CHameers, K.L. 1993. Microseris. In: Hickman, J.,ed. The Jepson manual: higher plants of Cali- fornia. Univ. California Press, Berkeley. Pp. 316-319. Coteman, R.G. and A.R. Kruckeserc. 1999, Geology and plant life of the Klamath — Siskiyou Mountain region. Natural Areas J. 19:320-340. McKnict, B.K. 1971. Petrology and sedimentation of Cretaceous and Eocene rocks in the Medford—Ashland region, south-eastern (sic!) Oregon. Ph.D. Thesis, Oregon State Univ.,, Corvallis. Mautue, S., K. BACHMANN, K.L. CHampers, and HJ. Price. 1982. Variability of the inflorescence among populations of Microseris laciniata (Asteraceae, Lactuceae). Beitr. Biol. Pflanzen i) a er Pires, MJP. 1980. Morphogenetic studies of intraspecific hybrids of Microseris laciniata (Hook.) Sch. Bip. M.S. Thesis, Oregon State Univ., Corvallis. Smith, J.P. Jr.and J.O. Sawyer. 1988. Endemic vascular plants of northwestern California and southwestern Oregon. Madrofo 35:54-69. Wuirtaker, R.H. 1961.Vegetation history of the Pacific Coast states and the “central” signifi- cance of the Klamath region. Madrono 16:5-23. BRIT.ORG/SIDA 21(1) BOOKS RECEIVED WiiiaM A. Weer, ELS, (Eb). 2004. The Valley of the Second Sons (Letters of Theodore Dru Alison Cockerell, a Young English Naturalist, Writing to His Sweetheart and Her Brother about His Life in West Cliff, Wet Mountain Valley, Colorado 1887-1890). ISBN 0-9710609-9-1, pbk.). Pilgrims Process, Inc., 4066 Niblick Drive, Longmont,CO80503.(Orders: 303-554-6641, wwwPilgrimsProcess.com). $39.95, 358 pp., 7 1/2" x 9". GLAFIRO J. ALANIS a °S§ AND ae ‘(GO GONZALEZ ALANIS ee lana Nativa Ornamen- tal DP. Ao A de las Especies sare el Paisaje Urbano. (ISBN ‘970- 649-]22-3 ori ). Univer- sidad Autonoma de Nuevo Leon R. Ayuntamiento de Monterrey, 2000-2003. (Orders: Email: galaris@ccrdsi.uani.mx). Price not given, 127 pp.,color photos, 81/4" x 81/4 JOsEPH FELSENSTEIN. 2004. Inferring Phylogenies. (ISBN 0-8893-728-5, pbk.). Sinauer Associates, Inc. Publishers, 23 Plumtree Road, Sunderland, MA 01375, U.S.A. (Orders: fax 413-549-1118, orders@sinauer.com, wwwsinauercom). $59.95, 664 pp., b/w photos, graphs, 7" x 9". The book is divided into 35 chapters: 1) Parsimony methods. 2) Counting evolutionary changes. 3) ow many trees are there? 4) Finding the best tree by heuristic search. 5) Find the best tree by branch and bound. 6) Ancestral states and branch lengths. 7) Variants of parsimony. 8) Compatibility. 9) Statistical properties of parsimony. 10) A digression on history and philosophy. 11) Distance matrix methods. 12) Quartets of species. 13) Models of DNA evolution. 14) Models of protein evolution. 15) Restriction sites, RAPDs, — and microsatellites. 16) Likelihood methods. 17) Hadamard meth- an ods. 18) B f phylogenies. 19) ae models, trees, and clocks. 20) Bootstrap, jack- knife, and permutation tests. 21) Paired-sites tests. 22) Invariants. 23) Brownian motion and . frequencies. 24) Quantitative characters. 25) Comparative methods. 26) Coalescent trees. 27) Likeli- hood calculations on coalescents. 28) Coalescents and species trees. 29) Alignment, gene ae and genomics. 30) Consensus trees and distances between trees. 31) Biogeography, hosts, and para- sites. 32) Phylogenies and paleontology. 33) Tests based on tree shape. 34) Drawing trees. 35) Phylog- eny software. SIDA21(1 ): 206. 2004 NEW TAXA AND NEW COMBINATIONS IN NORTH AMERICAN CIRSIUM (ASTERACEAE: CARDUEAE) David J. Keil Bi jologic al Sciences Department California P. olytechnic State ( University San Luis Obispo, California 93407, U.S.A. dkeil@calpoly.edu ABSTRACT Six new varieties are proposed in North American Cirsium ees cla ): C. arizonicum (A. Gray) Petrak var. tenuisectum DJ. Keil var. nov, C. eatonii (A. Gray) BL. Rob. var. viperinum DJ. Keil, var. nov, C. edule Nutt. var. wenatchense D.J. Keil, var. nov, C. occidentale ae var. lucianum DJ. Keil, var. nov. C. scariosum Nutt. var. robustum DJ. Keil, var. nov, C. scariosum Nutt. var. toiyabense DJ. Keil, var. nov. Additionally, twenty new combinations are presented: C. arizonicum (A. Gray) Petrak var. bipinnatum (Eastw.) DJ. Keil, comb. nov. C. arizonicum (A. Gray) Petrak var. chellyense (RJ. Moore & Frankton) DJ. Keil, comb. et stat. nov, C. arizonicum (A. Gray) Petrak var. rothrockii (A. Gray) DJ. Keil, comb. et stat. nov, C. clavatum (M.E. Jones) Petrak var. americanum (A. Gray) DJ. Keil, comb. nov, »C. clavatum (MEF. Jones) Petrak var. osterhoutii (Rydb.) DJ. Keil, comb. et stat. nov, C. ) J.T. Howell var. ieee eae Keil, comb. et stat. nov. C. eatonii (A. Gray) BL. Rob. var. eri phal (A. Nelso DJ. Keil, comb. nov, C. eatonii (A. Gray) B.L. Rob. var. hesperium (Eastw.) DJ. Keil, comb. et stat. nov, C. eatonii (A. Gray) B.L. Rob. var. peckii (L.F Hend.) DJ. Keil, comb. et stat. nov. C. eatonii (A. Gray) B.L. Rob. var. var. clokeyi (S.F Blake) DJ. Keil, comb. et stat. nov. —_ C. edule Nutt. var. macounii (Greene) DJ. Keil, comb. et stat. nov, C. horridulum Michx. var. mepacanthum (Nutt.) DJ. Keil, comb. et stat. nov., C. inamoenum (Greene) DJ. Keil, comb. nov, C. ) D. i} . Keil var. davisii (C Hae J. Keil, comb. et stat. nov, C. ochrocentrum Gray var. martinii (P. i Trick) D.J. Keil, comb. et stat. nov. C. pulcherrimum (Rydb. ) K.Schum. var. aridum (R.D. Dorn) DJ. Keil, comb. et stat. nov., C. seariosum Nutt. var. americanum (A. Gray) DJ. Keil, comb. nov, C. seariosum Nutt. var. citrinum (Petrak) DJ. Keil, comb. nov, C. seariosum Nutt. var. coloradense (Rydb.) DJ. Keil, comb. et stat. nov. C. scariosum Nutt. var. congdonii (RJ. Moore & Frankton) DJ. Keil, comb. et stat. nov. Lectotypes are designated for several of these taxa. RESUMEN S i iedad Ci (A d \deN arizonicum (A. Gay iParak var. tenuisectum DJ. Keil var. nov. C. eatonii (A. Gray) B.L. Rob. var. viperi DJ. Keil var. nov., C. edule Nutt. var. I DJ. Keil, var. nov, C. occidentale (Nutt. oe var. lucianum D). Keil, var. nov, C. scariosum Nutt. var. robustum DJ. Keil, var. nov, C. scariosum Nutt. var. toiyabense DJ. ae var. nov. Se presentan ademas veinte combinaciones nuevas: C. arizonicum (A. Gray) Petrak var. Eastw. eil, comb. nov, C. arizonicum (A. Gray) Petrak var. chellyense (RJ. Moore & Frankton) DJ. Keil, — et stat. nov., C. arizonicum (A. Gray) Petrak var. rothrockii (A. Gray) DJ. Keil, et stat. nov. C. clavatum (MLE. Jones) bee var. americanum (A. Gray) DJ. Keil, comb. nov, C. clavatum (M.E. Jones) Petrak var. db.) DJ. Keil, comb. et stat. nov, C. cymosum (Greene) J.T. Howell var. canovirens (Rydb.) DJ. Keil, eee et stat. nov. C. eatonii (A. Gray) B.L. Rob. var. eriocephalum (A. Nelson) DJ. Keil, comb. nov, C. eatonii (A. Gray) B.L. Rob. var. hesperium (Eastw.) DJ. Keil, comb. et stat. nov. C. eatonii (A. Gray) B.L. Rob. var. peckii (L.F Hend.) DJ. Keil, comb. et stat. nov. C. eatonii (A. Gray) B.L. Rob. var. var. clokeyi (S.F Blake) DJ. Keil, comb. et stat. nov, C. edule Nutt. var. = SIDA 21(1): 207-219. 2004 208 BRIT.ORG/SIDA 21(1) macounii (Greene) DJ. Keil, comb. et stat. nov, C. horridulum Michx. var. megacanthum (Nutt.) DJ. Keil, comb. et stat. nov, C. inamoenum (Greene) DJ. Keil, comb. nov. C. inamoenum (Greene) DJ. Keil var. davisii (Cronquist) DJ. Keil, comb. et stat. nov, C. ochrocentrum A. Gray var. martinii (P. Barlow-lrick) DJ. Keil, comb. et stat. nov, C. pulcherrimum (Rydb.) K. Schum. var. aridum (R.D. Dorn) DJ. Keil, comb. et stat. nov, C. scariosum Nutt. var. americanum (A. Gray) DJ. Keil, comb. nov, C. scariosum Nutt. var. citrinum (Petrak) DJ. Keil, comb. nov, Cc. scariosum Nutt. vat. coloradense (Rydb.) DJ. Keil, comb. et stat. nov. C. scariosum Nut J. Moore & Frankton) DJ Keil, comb. et stat. nov,, También se designan lectétipos de varios ae estos taxa. Preparation of a taxonomic treatment of Cirsium Mill. (Asteraceae: Cardueae) for the upcoming Asteraceae volumes of the Flora of North America has en- tailed a reevaluation of many of the published taxa. The only comprehensive treatment of North American Cirsium species (Petrak 1912, 1917) is long out of date. Revisions or cytotaxonomic studies have been published for several spe- cies groups (Frankton & Moore 1961, 1963; Moore & Frankton 1962, 1964, 1965, 1966, 1969, 1974, Ownbey 1952; Ownbey & Hsi 1963, 1969; Ownbey & Olson 1969; Ownbey et al. 1975), and many of the species have been included in state or regional floristic treatments (e.g., Cronquist 1955, 1994; Dorn 1992; Gardner 1974: Howell 1960; Keil & Turner 1993; Munz & Keck 1959; Munz 1974; Weber 1996; Weber & Wittmann 1987; Welsh 1983; Welsh et al. 1993), but these treat- ments have been uneven and limited in scope. Political boundaries often mark the application of differing taxonomic constructs. In my investigations | have examined North American Cirsium as a whole and across the breadth of its geographic range. Cirsium well deserves its reputation as a taxonomically difficult genus. One of the most challenging aspects for a taxonomist studying New World Cirsium is the presence of species complexes that are apparently evolutionary works in progress. Some of the thistles, especially in the mountainous western part of North America, are frustratingly polymorphic with mosaic and overlapping patterns of variation and intergradation of characters. Early taxonomists, bas- ing their work on a limited sampling of the morphological diversity, named many of the forms as species, and the literature is robust with species names. The infilling that results from more collectors visiting more localities within the ranges of these complexes has blurred the boundaries between many of the proposed species and often added forms that do not “fit” the characters of named species. As I faced the challenges of preparing the FNA treatment I recognized that maintaining some of the named entities as species would for consistency require a further proliferation of species names. I have chosen to go the other way. Instead continuing the proliferation of ill-defined microspecies | treat the plants in question as rapidly evolving, only partially differentiated assemblages of races that have not reached the level of stability that is usually associated with the concept of species. In a molecular phylogenetic investigation Kelch and Baldwin (2003) found unusually low rates — KEIL, NEW TAXA 209 of molecular divergence relative to the ecological diversity of New World Cirsium and suggested that the genus has undergone a rapid ecological diver- sification ona continental scale. Certainly there is much variation within these groups that deserves a level of taxonomic recognition, or that at least should be mentioned, but I think it much more prudent to recognize these as infraspe- cific taxa—entities that may be expected to freely intergrade—than species. I view species as entities that should be consistently recognizable. Within a large genus such as Cirsium the amount of infraspecific variation and the degree of geographical separation among races varies from species to species and from geographical race to geographical race. Races occupying contiguous habitats often intergrade freely in zones of contact whereas races currently isolated by geographic or habitat barriers often have more discrete variation patterns. Both situations may occur in a single species. Infraspecific taxa in Cirsium have been recognized in various taxa at the rank of form, variety, and subspecies. In the Flora of North America (Flora of North America Editorial Committee 1996) the trivial rank of form, represent- ing sporadic variants without a geographic range, is not used. Some Cirsium variants recognized in the past as varieties are the equivalent of forms. In Cirsium both variety and subspecies have been used in the context of morpho- logically disti hable geographical races. The rank of variety has been used more widely. The Flora of North America Guide for Contributors (Flora of North America Editorial Committee 1996) stipulates that only one infraspecific rank (subspecies or variety) be recognized within a genus. As used by most workers in Cirsium taxonomy the choice of rank has been a matter of preference, and I view them as equivalent. I have chosen to use variety rather than subspecies because only one change from subspecies to variety has to be made to follow the Flora of North America Guidelines whereas the use of subspecies as the infraspecific rank for formally recognized geographical races would require many more nomenclatural innovations. The overall product of my studies will be detailed in the FNA treatment (Keil, in press). Presented below are six newly proposed varieties and valida- tion of twenty new combinations for North American Cirsium. Cirsium arizonicum (A. Gray) Petrak var. bipinnatum (Eastw.) DJ. Keil, comb. nov. BASIONYM: Cnicus drummondii Torr. & A. Gray var. bipinnatus Eastw., Zoe 4:8. 1893. TYPE: COLORADO: Johnston Canon near where it joins Mancos River, Sep 1892, Eastwood s.n. (HOLOTYPE: CAS; ISOTYPE: MIN) Cirsium sa eeere (A. Gray) Petrak var. chellyense (RJ. Moore & Frankton) eil, comb. et stat. nov. BASIONYM: Cirsium chellyense RJ. Moore & Frankton, i J. Bot. 52:547. 1974. Type: ARIZONA. Apache Co: Canyon de Chelly, 1 Jul on Norman 680 (HOLOTYPE: DAO). 210 BRIT.ORG/SIDA 21 = 1) Cirsium arizonicum (A. Gray) Petrak var. rothrockii (A. Gray) DJ. Keil, comb. et stat. nov. BASIONYM: Cnicus rothrockii A. Gray, Proc. Amer. Acad. Arts 17:220. 1882. TYPE: ARIZONA: Rocky Canyon, 1874, Rothrock 289 (lectotype, here chosen from syntypes, GH). Gray (1882) described Cnicus rothrockii based upon two collec- tions (Rothrock 289 and Lemmon 2794) Moore and Frankton (1974) selected Rothrock 289 as the lectotype collection of Cnicus rothrockii. However, they cited as lectotype specimens of this gathering from both GH and US. According to Article 9.2 of the International Code of Botanical Nomenclature (Greuter et al. 2000) “a lectotype is a specimen or il- lustration designated from the original material as the nomenclatural type.” Moore and Frankton’s designation of two specimens at different institutions as lectotype fails the requirement that a single specimen serve in that role. lagree with their designation of Rothrock 289 as the lectotype collection and here choose the GH specimen as lectotype for the taxon. Cirsium arizonicum (A. Gray) Petrak var. tenuisectum DJ. Keil, var. nov. TyPe: CALIFORNIA. SAN BERNARDINO Co.: New York Mountain pBeystons Canyon, 2.1 road mi from Ivanpah Road, ca. 1670 m, Pi Juniperus osteosperma woodland; scattered in rocky wash channel; associates: Fallugia paradoxa, Prunus fasciculata, Gutierrezia sarothrae, Atriplex canescens, Eriodictyon angustifolium, Yucca brevifolia, Opuntia echinocarpa, Menodora scabra, Ephedra nevadensis, Pen- stemon sp., Machaeranthera canescens, Purshia mexicana, Muhlenbergia porteri, Ericameria linearifolia, Salvia dorrii, Lycium cooperi, Yucca baccata, Berberis sp., Rhus trilobata, Sphaeralcea sp., lpomopsis agregata, Datura wrightii, Artemisia dracunculus, and Achnatherum hymenoides, 5 Nov 1999, Keil 2840] (HOLOTYPE— flowering stem of plant D, OBI; soTyPes [the letter in parentheses represents the individual], ASU (A), BRY (C), CAS (B), CDA (C), DAO (D), GH (D), MIN (D), MO (C), NY (C), OBI (A, C), RSA (A), TEX (D), UC (B), US (D)). ibus et costis abaxialibus glabris ad tomentosis trichomatibus tenuibus non-septatis, spinis eerie oliorum 5-30 mm longis, et corollis rubellopurpureis distinguatur. Stems thinly arachnoid tomentose, + glabrate, without septate trichomes. Leaves deeply divided, often nearly to the midvein, abaxially arachnoid tomen- tose or sometimes glabrate, without septate trichomes, adaxially thinly arach- noid or glabrate; principal marginal spines 5-30 mm, often stout; cauline leaves narrowed at base to truncate or + clasping, but not or only slightly decurrent. Involucres cylindric to campanulate; phyllary spines 5-25 mm, stout. Corollas reddish purple, 25-35 mm, the tube 10-13, throat 5-8 mm, lobes 10-13.5 mm; style tips 1-2 mm. ParatyPes: CALIFORNIA. San Bernardino Co.: New York Mountains. Carruthers Canyon, 1615-1645 m, 6 Nov 1999, Keil 28402 (NMC, OBI, OSC); New York Mountains, Keystone Spring, 1585 m, 13 Oct 1935, Munz 13862 (UC). Nevada. Clark Co.: SW slope of NW Spring Mts., upper Clark Canyon, yellow pine vegetation, 7000-7600 ft, 9 Jul 1970, Beatley 11355 (UNLV), Charleston Mts., Charleston Peak, gravelly wash, with Pinus scopulorum, 2270 m, 18 Jul 1938, Clokey 8182 (PH, UC, UTCQ), WTU()), Charleston Mts., Kyle Canyon at bridge, wash with aspens, 2300 m, 3 Aug 1935, Clokey 5609 (UC), Charleston Mts., Charleston Park, gravelly wash, yellow pine belt, 2300 m, 25 Jul 1936, Clokey 5609 = (UC, UTC, WTU); Charleston Mts., Lee Canon, in limestone, 8000 ft, 4 Aug 1913, Heller 11057 (PH, UC); Charleston Mts., Pine Cr, 6700 ft, 4 Sep 1927, Jaeger 2202 (UC); Charleston Mts., near Charleston park resort, plentiful, 8000 ft, 12 Sep 1925, Jaeger s.n. UC); Spring Range, Toiyabe National Forest, Kyle Canyon Rd. (Nevada Rte. 157) just W of junction with Nevada Rte. 157, Ponderosa pine forest, 6930 {t, 17 Aug 2002, Keil 29112 (OBL); a noes Toiyabe National Forest, Las Vegas Ski and 5.69 Snow board area in Upper Lee Canyon, 36. 6° W, subalpine forest with limber pine, bristle- ne pine, white fir, quaking aspen, wax currant, oe eee ae 9200 ft, 17 Aug 2002, Keil 291 Ridge Park- | way, North Carolina, oie an allopolyploid of the cross Krigia bi Krigia montana (Michx.) N RESUMEN Se propone la nueva combinacion Krigia cespitosa (Ral.) K.L. Chambers ilis, basada en Apogon gracilis DC., y se hace una lectotipificacion del basionimo. Se crea un neoupe para Serinia cespitosa Raf. El hibrido hexaploide Krigia x shinnersiana se describe del Blue Ridge Parkway, North Caro- lina, y representa un alopoliploide del cruce Krigia biflora (Walter) SF. Blake x nee montana (Michx.) Nutt. THE VARIETIES OF KRIGIA CESPITOSA Krigia Schreb., Nomen Conservandum|, has a large synonymy, considering its rather modest number of species. Shinners (1947) recognized only seven spe- cies but listed eight synonymous generic names. This excess of names may have resulted from the various authors’ overemphasis on pappi differences and the contrast between annual and perennial habit within the genus. One species, as recognized here, consistently lacks pappi and, on this basis, was segregated as Serinia Raf. (Rafinesque 1817) and Apogon Elliott (EHiott 1823). DeCandolle (1838:261) placed Serinia in his Compositae Incertae Sedis but recognized Apogon humilis Elliott and added a second species, Apogon gracilis DC. These two spe- cies were again recognized by Gray (1884), and a third species, A. wrightii A. Gray, was described, differing from the others by the possession of “an obscure vestige of pappus.” The final union of Apogon and Serinia with Krigia was done by Shinners (1947), who included Krigia gracilis DC.) Shinners and Krigia oppo- sitifolia Raf. (=Krigia cespitosa (Raf.) K.L. Chambers, formerly Apogon humilis) but placed Apogon wrightii Gray in synonymy under the latter species. ‘As explained by Kim and Turner (1992), the International Code of Botanical Nomenclature (Greuter et al. 1988) erroneous stated that the basionym of the conserved type species, Krigia virginica (L.) Willd, is Tragopogon virginicus L., whereas it should be Hyoseris virginica L. This mistake is perpetuated in the current Saint Louis Code (Greuter et al. 2000).| second the proposal by Kim and Turner that this be corrected in the next edition of the Code. SIDA 21(1): 225-236. 2004 226 BRIT.ORG/SIDA 21(1) We now know much more about this cluster of taxa, from chromosome studies and the detailed molecular analyses by Kim and co-workers (Kim & Mabry 1991; Kim et al. 1992; Kim & Jansen 1994). Krigia gracilis and K. cespitosa both are diploid annuals with n = 4 and have similar epappose, fusiform cypselae. They are practically indistinguishable in the chloroplast cDNA and nuclear rDNA traits studied (summarized in Kim & Turner 1992). Krigia wrightii (A. Gray) K.L. Chambers ex K.-J. Kim is annual and tetraploid, with n = 9, and has more barrel-shaped cypselae often with a pappus of vestigial scales (figured in Kim & Turner 1992:180). Ina cladistic analysis of cDNA and rDNA data, a 100% bootstrap value provided by 51 synapomorphies separated it from the cluster that includes K. cespitosa and the related K. occidentalis Nutt. It is probably an ancient allopolyploid between a K. cespitosa-like species and one of the pap- pus-bearing annual species of the genus (Kim & Turner 1992). Prior to the work of Kim and Turner, systematists in Texas recognized Krigia gracilis and K. cespitosa as different species (Shinners 1958; Correll & Johnston 1970). Their emphasis was on morphological traits, especially the difference in size of heads and florets (corollas 5-10 mm long, involucres 5.3-8.5 mm high in flower, 6.2-8.5 mm high in fruit in K. gracilis, versus corollas 2-4 mm long, in- volucres 3-4.3 mm high in flower, 3.3-5.3 mm high in fruit in K. cespitosa). To these authors, K. gracilis was endemic to central and south-central Texas in the Blackland Prairie, occasionally west to the Llano region, mostly in clay-loam soil but also in adjacent regions of sandy soil, even becoming weedy in fields and disturbed ground (Shinners 1947). Krigia cespitosa is much more wide- spread, extending {rom central Texas east to Florida, the Carolinas, and Vir- ginia, and north to Kansas, Missouri, and Tennessee. In herbarium studies done much earlier by the present author, the area of overlap between the varieties in Texas formed a north-south band, from Denton and Tarrant Cos. east to Hunt Co. in the north, and from Travis and Hays Cos. east to Harris Co. in the south, with an extension east to Polk and Angelina Cos. Kim and Turner (1992) took a different view of these taxa. Based on their field and herbarium studies, they extended the range of K. gracilis from eastern Texas to “adjacent Oklahoma, Arkansas, and Louisiana,” and they noted the presence of populations interme- diate in head and flower size. Due to this pattern of intergradation and to the high similarity revealed by their DNA analysis, they proposed that the large- flowered types be reduced to the rank of K. cespitosa forma gracilis (DC) K-J. Kim (Kim & Turner 1992:196; Diggs et al. 1999). My purpose here is to make a different proposal for these taxa, based on several considerations. The first is the geographical pattern, in which a wide- spread small-flowered type intergrades at the western edge of its range witha morphologically distinctive and geographically limited large-flowered type. Geographical races marked by interbreeding in the area of overlap are more CHAMBERS, TAXONOMIC NOTES ON KRIGIA 227 often accorded the rank of variety than of forma. Weight must also be given to the nature of the morphological differences, as they represent a developmen- tally coordinated syndrome of variation in the reproductive organs. The large- flowered race may differ in breeding system, through pollinator attraction, an increased frequency of outcrossing, and genetic mixing in the otherwise self- pollinating reproductive mode exhibited by the two varieties (Kim & Turner 1992). Intraspecific differences in reproduction associated with conspicuous floral-size differences have been noted in other genera of Cichorieae and have been accorded varietal status. Examples known to the author are Agoseris heterophylla (Nutt.) Raf. var. californica (Nutt.) Jeps. (Chambers 1963), and Glyptopleura marginata D.C. Eaton var. setulosa (A. Gray) Jeps. Jepson 1925), both of which are large-flowered races in typically smaller-flowered species. To allow the recognition of this morphologically distinctive geographic race at a higher taxonomic level than forma, the following combination is proposed: Krigia ae (Raf.) K.L. Chambers var. se (DC.) K.L. Chambers, stat. nov. 1). Apogon gracilis DC., Prod. 7:79. 1838. Krigia gracilis (DC.) Shinners, Ste 1:205. 1947. Krigia cespitosa (R if ) K.L. Chambers forma gracilis (DC.) K.- J. Kim, Brittonia 44:196. 1992. Type: [TEXAS] “de Bejar a Austin, Avril 1828, Berlandier No. 1637” (LECTOTYPE: G-DC, the larger left-hand plant of 3 on the sheet; ISOLECTOTYPES: G, GH, US). The need for lectotypification of the basionym was first suggested by the word- ing of DeCandolle’s description, which includes the phrase “achaeniis scabris brevissime papposis.” Since Krigia gracilis, as the name has been used by Ameri- can botanists, totally lacks pappus, | became concerned that DeCandolle was dealing witha mixed collection, perhaps including K. wrightii or K. occidentalis. With the kind assistance of Dr. F Jacquemoud and Dr. L. Gautier of the Her- barium, Conservatoire et Jardin botaniques de la Ville de Geneve, it has been determined that the Berlandier type sheet is a mixed collection, the left-hand plant possessing epappose cypselae and the right-hand plant having vestigial pappus resembling the illustration of K. wrightii published by Kim and Turner (1992:180). To maintain the present usage of the epithet gracilis, | have desig- nated the left-hand plant as the lectotype. The isolectotype at GH studied by Shinners and Kim consists only of gracilis-type plants that lack pappi. Another duplicate in the general herbarium at Geneva, ex Herbier Moricand, clearly shows two gracilis-type plants and two of the pappus-bearing species. My notes from 1965 on the isolectotype at US, “ex Herb. Musei Britannici,” state that it has 2 plants of the gracilis-type and 4 plants with muticous pappi. Whether the second species in Berlandier’s collection is K. wrightii or a muticous form of K. occidentalis is not clear, although a photograph of the Herbier Moricand sheet strongly suggests the latter. BRIT.ORG/SIDA 21(1) NAVE NOE Fic. 1. Type sheet of Apogon gracilis, G-DC; lectotype plant is the tall specimen at the left. CHAMBERS, TAXONOMIC NOTES ON KRIGIA 229 Krigia cespitosa (Raf.) K.L.Chambers var. pias Serinia cespitosa Raf. Fl. ludov 149.1817. Krigia? oppositifolia Raf, op. cit. 57, nom. invalid. Krigia cespitosa (Raf.) K.L. Chambers, J. Arnold Arbor. 54:52- 3 1973. Krigia cespitosa (Raf). K.L. Cham- bers forma cespitosa in K.-J. Kim, Brittonia 44:196. 1992. TYPE: LOUISIANA: Natchitoches, open ground, 15 Apr 1915. EJ. Palmer 7220 (NEOTYPE: MO! ISONETOTYPE: NY!) Asis well known, Rafinesque’s Florula Ludoviciana (1817) was a translation of part of the French traveler C.C. Robin’s publication describing his voyages to Louisiana, Florida, and the West Indies, in which Rafinesque proposed many new species and genera. Collections by Robin have never been located. The above species was described first on page 57 as Krigia? oppositifolia, but that name was rejected by the author on page 149 of the “Additions,” and Serinia cespitosa was substituted (Chambers 1973). American authors have been in agreement as to what species Rafinesque was referring, but its name has not before been typified. A HYBRID KRIGIA FROM THE MOUNTAINS OF NORTH CAROLINA It was long ago reported that a hexaploid hybrid population of Krigia occurs on the Blue Ridge Parkway northeast of Ashville, North Carolina (Chambers 1965; Vuilleumier 1973), but its formal taxonomic recognition has been delayed until the present. The parental taxa are K. montana (Michx.) Nutt. (n = 10) and K. biflora (Walter) SF Blake (n = 5), whose area of sympatry is limited to the southern Appalachian Mountains. Krigia montana is endemic to this region (Shinners 1947; Wiser 1994), occurring mainly on granitic balds, cliffs, talus, and roadbanks, principally in North Carolina but extending into South Caro- lina, Georgia, and Tennessee in similar habitats. It is consistently tetraploid (Tomb et al. 1978). Krigia biflora is much more widespread, occurring in wood- lands and low prairies throughout the northeastern United States and parts of southern Canada, ranging south to Georgia, Alabama, and Arkansas, with dis- junct populations in Colorado, New Mexico, and Arizona (Kim & Turner 1992). In the Blue Ridge of North Carolina it is found principally in the shrub bald community (Ramseur 1960), and both diploid (n = 5) and tetraploid (n = 10) populations are known. At Craggy Gardens on the Blue Ridge Parkway, ca. 12 air line km northeast of Asheville, Buncombe County, hexaploid plants were found that were inter- mediate in various respects between the two above species. Their population will be described below. Samples from this site, along with samples of K. montana and of diploid and tetraploid K. biflora were contributed to the re- search of K.-J. Kim, University of Texas, and are mentioned in his various pub- lications. His biochemical studies involved the whole genus and included both chloroplast DNA, examined through restriction site analysis, and nuclear rDNA, for which restriction sites and ITS sequences were utilized. Because the chloro- plast genome is inherited maternally, it was expected that the hybrid’s chloro- 230 BRIT.ORG/SIDA 21(1) plasts would resemble one parent more than the other. In the cladistic analysis, the maternal parent proved to be diploid K. biflora (Kim et al. 1992). In the ITS sequence study, the hybrid did not associate closely with either parent but in- stead, in the strict consensus tree, formed an unresolved branch at the node basal to the two parents (Kim & Jansen 1994). However, a relationship of the hybrid with K. montana was shown in the analysis based on rDNA restriction sites (Kim and Mabry 1991). The hexaploid “exhibited combined length polymor- phism patterns of the diploid K. biflora and the tetraploid K. montana.” These studies leave no doubt that the Craggy Gardens plants are indeed the allopoly- ploid hybrid of K. biflora x K. montana. In order to call attention to this hybrid and to provide a name for use by those interested in studying it further, the following taxon is here described: Krigia < shinnersiana kK... Chambers, hybrid nov. (Fig. 2). Typr: US.A. NORTH CAROLINA. BUNCOMBE Co. Craggy Gardens, Blue Ridge Park way 18.6 mi N of junc- tion with Hwy. 70 E of Asheville, at the observation hut on grassy bald above the parking lot, elev. 5340 ft, grassy turf over granite substrate, 2] Jul 1968, K.L. Cham- bers 2880 (HOLOTYPE: OSC; ISOTYPES: GH, MO, NY, OSC, US). 2 Hybrida inter Krigiam eee amet ~ montanam intermedia caule inferne et superne ramoso foliato ramis superne folia redacta ferens longi-pedunculatis 1-3 simul terminatis, cypselis 3.5 mm longis, floribus aurantiaco- ice eenecoin atum numerus 2n = 60. Perennial herb with short rhizome, propagating by adventitious buds from the rootlets, stems decumbent or erect, to 65cm tall, leaves of the basal rosette 8-30 cm long, oblanceolate, tapering to a narrowly winged petiole, acute, entire or dentate to pinnatifid with remote, straight or arcuate, acute lobes, lower cauline leaves similar, 14-25 cm long, upper leaves reduced, lanceolate with broad, clasp- ing petiole, peduncles 14-28(-30) cm long, gue villous Is apex, in- volucre 7-10 mm high in flower, 8-13 mm high in fruit, phyllaries 11-13, equal, lanceolate, attenuate, green, glabrous or lightly glandular-villous, florets 20- 23mm long. The name honors Lloyd H. Shinners, whose valuable monograph (1947) forms the basis for studies of the genus. Additional collections: U.S.A. NORTH CAROLINA. Buncombe Co.: Blue Ridge Parkway, mile 364.4 Craggy Flats, in bald near trail shelter on Craggy Gardens nature trail, 14 Jul 1977, I. Govus 231 and Dan Pittillo(WCUH). Buncombe Co.: Scattered in grassy bald, top of Craggie Mt., 15 Jul 1935, H). Oosting 35453 (DUKE). A comparison of habit and flower color in Krigia x shinnersiana and its two parents is given in Table |. Differences in the cypselae also were noted (Fig. 3). Fruits of the hybrid were consistently longer, at 3.5mm, than in either K. biflora (3.0 mm) or K. montana (2.5-3.0 mm). In K. biflora the ribs of the cypselae are low and of equal prominence nearly to the base, whereas in K. montana, 3 ribs are usually more prominent than the others in adaxial view, some other ribs being suppressed. In K. x shinnersiana the ribs in adaxial view arise at the base in groups of 3, with the central rib being more prominent to near the cypsela CHAMBERS, TAXONOMIC NOTES ON KRIGIA KRIGIA X SHINNERSIANA K. Annotated: K. L. Chambers 2/4/2004 Hybrid behoeen giles x B:Plorn VRIVDESIT? OF SSRSARIUE (hb. Sel) Krigia montana (Michx.) Nu gs Lie Ls] vet PLANTS of NORTH CAROLINA KRIGIA BUNCOMBE EG} keke af the Blue Ridge et 18.6 m c. 70 east of Asheville, bald above the asters , on 2S Hwy. KERSAPLUM ervation hut on grassy pea lot. Elev. 0 fee 4myQy: over granite substrate. 4 PIDRS transplants flowered and coll ORESOM STATE 1968 in garden at Kensington, K. L. Chambers (2880-3) UNIVERSITY Fic. 2. Holotype of Krigia x shinnersiana, OSC 232 BRIT.ORG/SIDA 21(1) nee ee SS — FS Imm (Chambers 1404), B. K. x shinnersiana Fic. 3. Cypselae of Krigia = shinnersiana and its parental taxa. A. Krigi (Chambers 1362), C.K. montana (Chambers 1360). Pappi not shown. apex. The number of bristles per cypsela was compared, using mature heads collected in glass vials to avoid shattering the delicate pappi (Table 2). Bristle numbers in K. biflora are distinctly greater than in K. montana, while the hy- brid numbers vary from those of K. montana up to an approximately interme- diate number. lf bristle number is assumed to have a genetic component, the range of numbers in the hybrid, sampled at different times from the type local- ity, suggests that two or more different genotypes are present. The type locality is reached by a trail heading south 0.5 km from the Visi- tors Center at the Craggy Gardens parking area, toa hikers’ shelter constructed of weathered chestnut beams. The grass bald south of the shelter is being in- vaded by small shrubs of Rhododendron catawbiense from the surrounding heath bald community (Ramseur 1960:90). Krigia montana is common on Craggy Pinnacle north of the parking area but was not seen in the heath or grass bald communities along the south trail. Krigia biflora was never noted in this vicinity during my field studies. Observations made in 1995 found the hy- brid to be abundant in two areas of the bald, forming colonies near small shrubs of Rhododendron and Vaccinium. The absence of variation in transplants from CHAMBERS, TAXONOMIC NOTES ON KRIGIA 233 Taste 1. Comparison of habit and flower color of Krigia biflora, K. montana, and K. x shinnersiana. *Measured by means of Nickerson Color Fan (Munsell Color Co., Baltimore, MD). Floral pigments of Krigia spp. identified as carotenoids by Harborne (1997). Krigia biflora Krigia < shinnersiana Krigia montana Stems naked below or with one large leaf low-down; modified clasping leaf higher up. Main stem ending in umbellate cluster of 2-6 naked floral peduncles. Terminal cluster of peduncles subtended by 1-3 bracts up to 3(—5) cm long. Later inflorescences may arise in axils of clasping upper leaf and lower stem leaf. First floral peduncle not arising singly from basal leaf rosette. Flower color Munsell Hue 10YR 8/10 moderate orange yellow.* Same as K. montana Main stem ending in cluster of 1- naked floral peduncles. Terminal peduncles subtended by 1-3 bract- like leaves up to 9(-11) cm long. Same as K. montana Same as K. biflora Flower color Munsell Hue 5Y 8/12, vivid yellow. Cauline leaves well-developed, with axillary branches above basal leaf rosette. Main stem usually terminated by a single naked floral peduncle. Terminal peduncle subtended by 1-2 leaves up to 18 cm long. Later inflorescence branches arise in axils of stem leaves and leaves subtending terminal peduncle First floral peduncle may arise singly near basal leaf rosette. Flower color Munsell Hue 7.5Y 8/12, vivid greenish yellow. Taste 2. Mean number of pappi bristles per cypsela in plants of Krigia biflora, K. montana, and K. x shinnersiana. Collections of the latter taxon were made at Craggy Gardens at various times. The standard deviation, range, and number of cypselae sampled are given. See Appendix for other locality information. Collection Mean S.D. Range Number 1207 biflora 2x 36.86 2.97 30-42 28 1404 biflora 4x 31.88 3.15 26-39 24 2894 montana 15.38 1.36 13-18 16 2881 montana 18.67 1.36 16-22 27 2887 montana 21.13 1.46 20-23 8 2889 montana 21.88 1.45 19-25 16 2888-1 shinnersiana 18.49 1.64 14-22 38 2880 shinnersiana 19.48 1.35 16-22 29 2888-2 shinnersiana 20.74 1.95 18-23 50 2879-2 shinnersiana 21.16 241 17-26 76 1362 shinnersiana 24.17 2.17 21-28 12 2879-5 shinnersiana 24.80 2.07 20-28 20 2879-3 shinnersiana 27.70 1.52 25-30 20 234 BRIT.ORG/SIDA 21(1 = each colony suggests that reproduction is largely clonal. One such colony was located 31 m southeast of the shelter, just east of an incised walking trail, while aS was found 28 m southwest of the shelter, west of a second trail. The tw nies differ in leaf shape (nearly entire vs. sharply pinnately lobed) and a represent different penoty pee. Fruit-set is high when hybrid plants are cross-pollinated experimentally, but seedling a may be infre- quent in the dense stand of Carex and grasses of the As discussed by Wiser and White (1999), grass bald of the Southern Ap- palachians were used as summer grazing pastures until around 1930, but they are not maintained by current natural processes and are subject to woody plant invasion. The bald at the type locality, under the name Craggy Flats, was de- scribed by Pittillo and Govus (1978), who included a brief list of the herbaceous flora. Krigia x shinnersiana was mentioned in this list under the name K. biflora (voucher checked by the author). It is tempting to speculate that the hybrid originated over 70 years ago, in the period of cattle grazing, when habitat dis- turbance was greater than now and open ground was more available for hybrid seedling establishment. The two parental species must have been in genetic con- tact over a long period of time in this mountainous region, and we can expect to find the products of their hybridization at other sites as well. Evidence of this is the discovery of pentaploid plants referable to Krigia biflora at two sites along the trail up Mt. Pisgah from the Blue Ridge Parkway, Haywood County, NC (Chambers 2891, 2893). The habitat was trail-side in a shrub bald about 50 m below the summit communications tower, in very different conditions than the grass bald at Craggy Flat. Morphology of the plants suggests a contribution of K. montana to their origin. It is hoped that this report will stimulate interest and further study by persons to whom these sites on the Blue Ridge Parkway are readily accessible. — APPENDIX! COLLECTIONS CITED Krigia biflora —NEW JERSEY. Atlantic Co.: Nesco, 21 May 1957, Chambers 1207. WEST VIRGINIA. Nicholas Co.: 7.7 mi N of Mt. Nebo, 30 Aug 1958, Chambers 1404. NORTH CAROLINA. Heywood Co.: trail up Mt. Pisgah, where vegetation changes to Kalmia scrub with oaks, 21 Jun 1968, Chambers 2891; same, 50 es down soe from summit tower, Chambers 2893 Krigia montana.—NORTH CAROLINA. Rutherford Co.: Chimney oe Park, oa te Chambers 907, re 1360. Buncombe Co.: Blue ae Parkway, Craggy Gina 20 Oct Chamber ame, natu trail by Craggy Gardens ey: lot, 20 Jun 1968, Chambers oe mbe Co.: Blue ae nn vay, Balsam Gap, 6.5 mi of Craggy Gardens, 20 Jun 1968, C ae 2889. SOUTH CAROLIN Greeneville Co.: Caesar's a 22 Jun 1968, Chambers 2894. Krigia X shinnersiana —NORTH CAROLINA. Buncombe Co.: grassy bald at observation hut on trail S of Craggy Gardens parking lot, 8 Jun 1958, Chambers 1362; same, 20 Oct 1967, Chambers 2879, 2880; same, 20 Jun 1968, Chambers 2888. CHAMBERS, TAXONOMIC NOTES ON KRIGIA 235 ACKNOWLEDGMENTS lam pleased to acknowledge the help of the following persons during the course of these studies: David Boufford, Henrietta Chambers, Kanchi Gandhi, Laurent Gautier, Bonnie Hall, F Jacquemoud, Aaron Liston, John Nelson, J. Dan Pittillo, Beryl Simpson, Rena Schlachter, Chris Tanner, and Carroll Wood. | appreciate Guy Nesom andan for carefully reviewing the manuscript. REFERENCES CHamBERS, K.L. 1963. Amphitropical species pairs in Microseris and Agoseris (Compositae: Cichorieae). Quart. Rev. Biol. 38:124-140. CHameers, K.L. 1973. Krigia cespitosa. In: B.S. Vuilleumier, The genera of Lactuceae (Compositae) in the Southeastern United States. J. Arnold Arbor. 54:52-53. CHameers, K.L. 1965. An allopolyploid Krigia from the southern Appalachians. Amer. J. Bot. (Abstr.) 52:658. Corrett, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner DeCanvoite, A.P. 1838. Prodromus systematis naturalis regni vegetabilis. Pars 7. Treuttel & Wurtz, Paris. Dicas Jr. G.M., B.L. Liescome, and RJ. O’Kennon. 1999. Shinners & Mahler's illustrated flora of north central Texas. Botanical Research Institute of Texas, Fort Worth. ELuiott, S. 1824.A. sketch of the botany of South-Carolina and Georgia. Facsimile ed. 1971, Hafner Publ. Co., New York. Gray, A. 1884. Synoptical flora of North America. Vol.|, Part Il. lvison, Blakeman, Taylor & Co., New York. Greuter, W. et al. 1988. International code of botanical nomenclature. Regnum Veg. 118. Koeltz Scientific Books, Koenigstein Greuter, W. et al. 2000. International code of botanical nomenclature. Regnum Veg. 138. Koeltz Scientific Books, Koenigstein Harsorne, J.B. 1977. Variations in pigment patterns in Pyrrhopappus and related taxa of the Cichorieae. Phytochem. 16:927-928. Jepson, W.L. 1925. A manual of the flowering plants of California. Univ. of California, Berkeley. Kim, K.-J.and TJ.Masry. 1991.Phylogenetic and evolutionary implications of nuclear ribo- somal DNA variation in dwarf dandelions (Krigia, Lactuceae, Asteraceae). Pl. Syst. Evol. 177:53-69. Kim, K.-J. and B.L. Turner. 1992. Systematic overview of Krigia (Asteraceae—Lactuceae). Brittonia 44:1 73-198. Kim, K.-J., B.L. Turner, and R.K. JANSEN. 1992, Phylogenetic and evolutionary implications of interspecific chloroplast DNA variation in Krigia (Asteraceae—Lactuceae). Syst. Bot. 17:449-469 Kim, K.-J.and R.K. JANsen. 1994. Comparisons of phylogenetic hypotheses among different BRIT.ORG/SIDA 21(1) data sets in dwarf dandelions (Krigia, Asteraceae): additional information from internal transcribed spacer sequences of nuclear ribosomal DNA. PI. Syst. Evol. 190:157-185. Pittito, J.D.and T.E.Govus. 1978.|mportant plant habitats of the Blue Ridge Parkway from the Great Smoky Mountain National Park to Roanoke, Virginia. Southeast Regional Of- fice, National Park Service. Rarinesque, C.S. 1817. Florula ludoviciana. C. Wiley & Co., New York. Ramseur, G.S. 1960. The vascular flora of high mountain communities of the southern Ap- palachians. J. Elisha Mitchell Sci. Soc. 76:82-112. SHinners, L.H. 1958. Spring flora of the Dallas-Fort Worth area, Texas. Lloyd H. Shinners, Dallas. SHINNERS, L.H. 1947. Revision of the genus Krigia Schreber.Wrightia 1:187-206. Tome, A.S., K.L.CHameers, D.W. KyHos, A.M. Powett, and PH. Raven. 1978.Chromosome numbers in the Compositae. XIV. Lactuceae. Amer. J. Bot.65:717-721. VuiLLEUMIeR, B.S. 1973. The genera of Lactuceae (Compositae) in the southeastern United States. J. Arnold Arbor. 54:42-93. Wiser, S.K. 1994. High-elevation cliffs and outcrops of the southern Appalachians: vascular plants and biogeography. Castanea 59:85-116. Wiser, S.K.and PS.Wuite. 1999. High-elevation outcrops and barrens of the southern Appa- lachian Mountains. Pp. 119-132. In: R.C. Anderson, J.S. Fralish, and J.M. Baskin, eds. Sa- vannas, barrens, and rock outcrop plant communities of North America. Cambridge Univ. Press. NEW COMBINATIONS IN NORTH AMERICA EREMOGONE (CARYOPHYLLACEAE) Ronald L. Hartman Richard K. Rabeler Rocky Mountain Herbarium niversity of Michigan Herbariu Department of Botany 3600 Varsity Drive University of fe Ann Arbor, Michigan 48108-2287, U.S.A. Laramie, Wyoming 82071-3165, U.S.A. rabeler@umich.edu ice eeraulent: ABSTRACT The following 18 n bi 1 in _— 1g I ie 15 ill is Var. americana, E. congesta var. caphalbides E. congesta var. charlestonensis, E. congesta var. ercsule E. congestd var. glandulifera, E.congesta var. prolifera, E. congesta var. ane E. congesta var. subcongesta, E. congesta var. Sul Jailescems, E. congesta var. wheelerensis, E. eastwoodiae var. adenophora, E. franklinii, E. franklinii ve kingii var. plateauensis, E. kingii var. rosea, E. macradenia var. arcuifolia, E. macradenia vanjeraae ane E. macradenia var. kuschei. RESUMEN Se proponen dieciocl | E E illaris var. americana, E. ere var. cephaloidea, E. congesta var. chavlesipenct B, aieda var. masala E, congesta var. glandul ee i sce a var. prolifera, E. congesta var. simulans, E. ee var. subcongesta, E. congesta var. su , E.congesta var. wheelerensis, E. eastwoodiae iden or E ee Efi dnblan var. the mpsonii, E. kingii var. plateauensis, E. kingit var. rosea, E. iene nia var. arcuifolia, E macradenia var. ferrisiae, y E. macradenia var. kuschei. During preparation of the treatment of the Caryophyllaceae for Flora North America (Rabeler and Hartman, editors, in prep), the decision was made to rec- ognize the genus Eremogone Fenzl ~ Arenaria subgenera Eremogone (Fenzl) Fenzl and Eremogoneastrum F. Williams sensu McNeill 1962). Nepokroeff and Wagner (pers. comm.; see also Nepokroeff et al. 2001) con- ducted a DNA survey of the Caryophyllaceae, using sequence data from matK and the rpsl6 intron. Over 79 species constituting 38 genera were included, with emphasis on the subfamily Alsinoideae. The phylogeny illustrated Arenaria (sensu McNeill (1962), excl. Minuartia and Moehringia) to be polyphyletic, with two major, distinct clades. While one clade includes members of four of McNeill’s subgenera, including Arenaria serpyllifolia L., the type of Arenaria, and the genus Moehringia, the second clade includes seven species (five from subgenus Eremogone, three North American and two Asian, and two Asian members of subgenus Eremogoneastrum). Within that clade, Arenaria bryophylla Fern. (subg. Eremogoneastrum from Asia) clusters closely with A. aculeata S. Watson and A. kingii (S. Watson) MLE. Jones (subg. Eremogone from North America), suggesting that the relationships between the subgenera may also need exami- SIDA 21(1): 237-241. 2004 238 BRIT.ORG/SIDA 21(1) nation. The results indicated that the Eremogone group, including both sub- genera, is strongly supported as monophyletic. Eremogone was treated by McNeill 962) as a subgenus of Arenaria con- taining eight sections (64 species, our compilation; ca. 70 species (Zhengyiet al. 2001)) closely allied with subgenus Eremogoneastrum (ca. 22 species). Mem- bers of subgenus Eremogone are found from western Europe through Asia, east tosouthern Alaska and northwestern Canada, disjunct to southwestern Canada and the western United States. The composition of Eremogone sensu Ikonnikov (1990) excludes McNeill’s monotypic section Monogone Maxim. (the Tien Shan Mountains and the Altai Region), two of the species from his section Scariosae (northern Iran and Turkish Armenia), and his sect. Pungentes (two species, Spain and North Africa). As outlined by McNeill 1962), Arenaria subgenus Fremogoneastrum is confined to three centers with two species (Arenaria hookeri Nutt. and A. franklinii Dougl. ex Hook. a third, A. pinetorum A. Nelson is a variety of the former) in western North America, the type, A. festucoides Benth., known from India, Pakistan, and China, and the remainder (ca. 6 species) Sino-Himalayan in distribution. Seventeen species are now known from China (Zhengyi et al. 2001); most are Sino-Himalayan, nine were published since 1979. Ikonnikov makes no mention of Arenaria subgenus Eremogoneastrum in either of his syn- opses of Eremogone (1973, 1990). In North America, both Maguire (1947, 1951; treated in Maguire (1951) as Arenaria sect. Pentadenaria Williams) and Hickman (1971) recognized a broader concept of Arenaria section Eremogone, including the two western American members of subgenus Eremogoneastrum (A. hookeri and A. franklinii). Maguire treated those species as related to but of uncertain place- ment, while Hickman considered them to be intermediate between sections Eremogone and Alsine (= Minuartia). Eremogone as presented here is currently accepted in floras of Colorado (Weber & Wittmann 1992 and subsequent up- dates at http://cumuseum.colorado.edu/) and Wyoming (Dorn 2001). Morphologically, Eremogone consists of woody based perennials with a cespitose or matted habitat, filiform to subulate leaves with scarious bases, stiffly erect or ascending flowering stems, and cymes that are open to congested or umbellate. This is contrasted with the remaining North American (north of Mexico) members of Arenaria (ca. 9 species), that are annuals or perennials with either a caespitose habit or ascending to trailing stems, broader (ovate to lanceolate, sometimes narrowly so) leaves, and flowers often solitary or in few- flowered, open cymes. Ikonnikov (1973) and Holub (1974) both noted that a base chromosome number of x = 11 also distinguished Eremogone from Arenaria (x = 10). Of the approximately 30 species of Eremogone, as here defined, which have been counted, only three counts (including two for E. fendleri that appear not to have been vouchered; that taxon was counted by Hartman (1971) as 2n = 44) are not based on x = 1]. Contrary to their notion, Arenariass. is cytologically more di- verse; while many counts are based on x = 10, counts based on x = 11 are known in two subgenera as well as five sections of subg. Arenaria (Rabeler, pers. com- pilation from standard chromosome atlases). Most of the nomenclatural combinations for the 13 North American spe- cies of Arenaria to be included in Eremogone are available Ikonnikov 1973, 1974, Weber et al. 1981); by contrast, only a few of the infraspecific taxa have the ap- propriate combinations (Weber et al. 1981; Dorn 2001) The following combinations are needed for the genus Eremogone in North America in anticipation of the Flora North America treatment (Hartman, Rabeler and Utech, in prep). Eremogone capillaris (Poir.) Fenzl var. americana (Maguire) R.L. Hartman & er, comb. nov. BASIONYM: Arenaria capillaris Poir. subsp. americana Magu- ire, Bull. Torrey Bot. Club 74:41. 1947. Arenaria capillaris var. americana (Magu- ire) RJ. Davis, Madrono 11:144. 1951. Eremogone americana (Maguire) Ikonn., Novosti Sist. Vyssh. Rast. 11:174. 1974. Eremogone congesta (Nutt.) Ikonn. var. cephaloidea (Rydb.) R.L. Hartman & comb. nov. BASIONYM: Arenaria cephaloidea Rydb., Bull. Torrey Bot. Club 39:316. 1912. Arenaria congesta Nutt. var. cephaloidea (Rydb.) Maguire, Bull. Torrey Bot. Club 74:46. 1947. Eremogone congesta (Nutt.) Ikonn. var. charlestonensis (Maguire) R.L. Hartman & Rabeler,comb. nov. BASIONYM: Arenaria congesta Nutt. var. charlestonensis Maguire, Bull Torrey Bot. Club 73:326. 1946. Eremogone congesta (Nutt.) [konn. var. crassula (Maguire) R.L. Hartman & Rabeler, comb. nov. BASIONYM: Arenaria congesta Nutt. var. crassula Maguire, Bull. Torrey Bot. Club 74:45. 1947. Eremogone congesta (Nutt.) [konn. var. glandulifera (Maguire) R.L. Hartman & abeler, comb. nov. BASIONYM: Arenaria congesta Nutt. var. glandulifera Magu- ire, Amer. Midl. Naturalist 46:501. 1951. Eremogone congesta (Nutt.) [konn. var. prolifera (Maguire) R.L. Hartman & beler, comb. nov. BASIONYM: Arenaria congesta Nutt. var. prolifera Maguire, Bull. Torrey Bot. Club 74:47. 1947. Eremogone congesta (Nutt.) [konn. var. simulans (Maguire) R.L. Hartman @ er, comb. nov. BASIONYM: Arenaria congesta Nutt. var. simulans Maguire, Bull. Torrey Bot. Club 74:48. 1947. Eremogone congesta (Nutt.) Ikonn. var. subcongesta (S. Watson) R.L. Hartman & eler, comb. nov. BASIONYM: Arenaria fendleri A. Gray var. subcongesta S. Watson, Botany [Fortieth Parallel]. 5:40. 1871. Arenaria congesta Nutt. var. subcongesta (S. Watson) S. Watson, Bot. Calif. 1:69. 1885. Arenaria subcongesta (S.Watson) Rydb., Bull. Torrey Bot. Club 24:244. 1897. Arenaria burkei Howell, FI. N.W. Amer. 1:85. 1897. Eremogone congesta (Nutt.) [konn. var. suffrutescens (A. Gray) R.L. Hartman & Rabeler, comb. nov. BASIONYM: Bre werina suffrutescens A. Gray, Proc. Amer. Acad 240 BRIT.ORG/SIDA 21(1) 8:620.1873.A a Nutt. var. suffrutescens (A. Gray) B.L. Rob., Proc. Amer. Acad. Arts 9: 295, 1804. Doenaae suffrutescens (A. Gray) A. Heller, Muhlenbergia 6:96. 1910. Eremogone congesta (Nutt.) Ikonn. var. wheelerensis (Maguire) R.L. Hartman & er, comb. nov. BASIONYM: Arenaria congesta Nutt. var. wheelerensis Magu- ire, Bull. Torrey Bot. Club 74:48. 1947. at aaa (Rydb.) Ikonn. var. adenophora (Kearney & Peebles) R.L. Rabeler, comb. nov. BASIONYM: Arenaria eastwoodiae Rydb. var. aa oe sa & Peebles, J. Wash. Acad. Sci. 29:475. 1939. Eremogone franklinii (Dougl. ex Hook.) R.L. Hartman & Rabeler, comb. nov, BASIONYM: Arenaria franklinti Dougl. ex Hook., Fl. Bor. Amer. 1:101. 1831. ies franklinii (Dougl. ex Hook.) R.L. Hartman & Rabeler var. thompsonii (M. Peck) R.L. Hartman & Rabeler, comb. nov. BASIONYM: Arenaria franklinii Danek ex Hook. var. thompsonii M. Peck, Torreya 32:149. 1932 (Thompsoni). ai oi kingii (S. Watson) Ikonn. var. plateauensis (Maguire) R.L. Hartman Rabeler, comb. nov. BASIONYM: Arenaria kingii (S. Watson) M.E. Jones subsp. Sees Maguire, Bull. Torrey Bot. Club 74:54. 1947. Arenaria kingii var. plateauensis (Maguire) Reveal, Great Basin Naturalist 35:344. 1975. Eremogone kingii (S. Watson) Ikonn. var. rosea (Maguire) R.L. Hartman & Rabeler, comb. et stat. nov. BASIONYM: Arenaria kingii (S. Watson) M_E. Jones subsp. rosea Maguire, Bull. Torrey Bot. Club 73:326. 1946. Eremogone macradenia (S. Watson) Ikonn. var. arcuifolia (Maguire) R.L. Hart- & Rabeler, comb. nov. BASIONYM: Arenaria macradenia S. Watson vat. arcuifolia Maguire, Bull. Torrey Bot. Club 74:51. 1947. Eremogone macradenia (S. Watson) Ikonn. var. ferrisiae (Abrams) R.L. Hartman Rabeler, comb. et stat. nov. BASIONYM: Arenaria macradenia S. Watson subsp. ferrisiae Abrams, Ill. Fl. Pacif. States 2:151. 1944 Eremogone macradenia (S. Watson) Ikonn. var. kuschei (Eastw.) R.L. Hartman abeler, comb. nov. BASIONYM: Arendrid kuschei Eastw, Proc. Calif. Acad. IV. 20:140. 1931. Arenaria macradenia S. Watson var. kuschei (Eastw.) Magu- ire, Bull. Torrey Bot. Club 74:51. 1947 ACKNOWLEDGMENTS We wish to thank Kanchi Gandhi (GH) for nomenclatural advice and Molly Nepokroeff (SDU) and Warren L. Wagner (US) for sharing with us results of their molecular work on the Car yophy llaceae. — REFERENCES Dorn, R.D. 2001. Vascular plants of Wyoming. Third Ed. Mountain West Publishing. Hartman, R.L. 1971. Chromosome numbers in Caryophyllaceae from Wyoming and adja- cent states. Bull. Torrey Bot. Club 98:276-280. Hickman, J.C. 1971. Arenaria, Section Eremogone, (Caryophyllaceae in the Pacific North- west:a key and discussion. Madrono 21:201-207. Ho.us, J. 1974. New names in Phanerogamae 3.Folia Geobot. Phytotax. 9:261-275. Hotus, J. 1977. New names in Phanerogamae 6. Folia Geobot. Phytotax. 12:41 7-432. IKONNiKov, S.S.1973.Notae de Caryophyllaceis, 1. Novosti Sist. Vyssh. Rast. 10:136—142. IKONNIKOV, S.S. 1974. Notula de speciebus Americanis generis Eremogone Fenzl (fam. Caryophyllaceae). Novosti Sist. Vyssh. Rast. 11:174-175. IKONNIKOV, S.S. 1990. Notae de Caryophyllaceis, 9. Novosti Sist. Vyssh. Rast. 27:63-65. Macuire, B. 1947. Studies in the Caryophyllaceae—lll. A synopsis of the North American species of Arenaria, sect. Eremogone Fenzl. Bull. Torrey Bot. Club 74:38-56. Macuire, B. 1951. Studies in the Caryophyllaceae—V.Arenaria in America North of Mexico. A Conspectus. Amer. Midland Nat. 46:493-511. McNett, J. 1962.Taxonomic studies in the Alsincideae:|.Generic and infra-generic groups. Notes Roy. Bot. Gard. Edinburgh 24:79-155. Nepoxroerr, M., W.L. Wacner, E.A. Zimmer, S.G. Wetter, A.K. Sakai, and R.K. RaBeter. 2001. Origin of the Hawaiian subfam. Alsinoideae and preliminary relationships in Caryophyllaceae inferred from matkK and trnL C-F sequence data. Botany 2001 Abstracts: 130. Bot. Soc. of America, Columbus, Ohio. (Also posted at http://www.botany2001.org) Weser, W.A., B.C. JOHNSTON, and R. Wittmann. 1981. Additions to the flora of Colorado-VIl. Brittonia 33:325-331. Weeer, W.A and R.C. Wittmann. 1992. Catalog of the Colorado flora: a Biodiversity Baseline. Univ. Press of Colorado, Boulder ZHENGYI, W., Z. Linua, and W.L. Wacner. 2001. 19. Arenaria. In: Zhengyi, W., RH. Raven, and H. Deyuan, eds. Flora of China 6:40-66. 242 BRIT.ORG/SIDA 21(1) BOOK NOTICES FLORAS AND CHECKLISTS Puinip A. Munz (Edited by Dianne Lake and Phyllis M. Faber). 2003. Introduction To Shore Wildflowers of California, Oregon, and Washington. Revised Edition. (ISBN 0-520-23639-4, pbk.). University of California Press, California/Prin- ceton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, US.A. (Orders: 609-883-1759, 609-883-7413 fax). $16.95, 234 pp., color photos, 5"

s, gymnosperms, dicots and mono- cots. There is a brief description et the e family followed bya key to the various taxa in the family with 14 -] : f ] a f } ] A g f the book is the color plate part in which almost one quarter of the taxa are alinsrented with a nice color photograph.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 761202-4060, U.S.A. VERNON L. Harms. 2003. Checklist of the Vascular Plants of Saskatchewan and the Provincially and Nationally Rare Native Plants in Saskatchewan. (ISBN 0-88880-471-7, pbk.). University of Saskatchewan, Extension Division, Kirk Hall Room 125, 117 Science Place, Saskatoon, SK 57N 5C8 CANADA. (Or- ders: 306-966-5565, 306-966-5567 fax). $39.95ed, 328 pp., 6" x 9" “The Checklist of the Saskatchewan Vascular Plants is a compilation of all native and naturalized vascular plants that occur in Saskatchewan.” The checklist provides users with relatively compre- es : ; eee ee ae ern sk o index. SIDA 21(1): 242. 2004 NEW COMBINATIONS IN THE GENUS GUNDLACHIA AND FOUR NEW GENERA OF ASTEREAE (ASTERACEAE) FROM NORTHERN MEXICO AND THE SOUTHERN UNITED STATES Lowell E. Urbatsch Roland P. Roberts Department of Biological Sciences Department of Biological Sciences Louisiana State University Towson University Baton Rouge, Louisiana 70803, U.S.A. Towson, Maryland 21252-0001, U.S.A. ABSTRACT New binati | a for species in Xyloth i ided, reflective of recent phylo- genetic data indicating the naomi tic nature of that genus ana the need to revise its taxonomy. Four species of Xylothamia are arenes under the genus Gundlacl ssitating the following new combinations: G. diffusa, indii, G. triantha (the type for lng: and G. truncata. Four Ww genera are recognized = fee without available eemeye? names. Neonesomia includes N. aia and N. palmeri. Each of the other t1 Chihuahuana, Medranoa, and Xylovirgata, and = accommodate C. purpusii, M. parrasana, and X. pseudobaccharis, respectively. Key Woros: Asteraceae, Astereae, Xylothamia, Chihuahuana, Gundlachia, Medranoa, Neonesomia, and Xylovirgata, Chihuahuan Desert, Sonoran Desert, Caribbean plants RESUMEN : as nuevas Soi inacioneS ye les nuevos géneros aus se ofrecen para las especies en oo mid son reflejo de | indican la naturaleza polifilética de ese gén oa de revisar su taxonomia. Chane especies de Xylothamia se incluyen en ele género Gundlachia que hace necesarias las nuevas combinaciones siguientes: G. dupes G. riskindii, G. triantha, (la tipo para Xylothamia), y G. truncata. Se r los linajes sin nombres genéricos disponibles. Neonesomia incluye N. johnstonii y N. paliaert Cae uno de los otros tres géneros es monotipico, Chihuahuana, Medranoa, y Xylovirgata, e incluyen a C. purpusii, M. parrasana, y X. pseudobaccharis, respectivamente. INTRODUCTION Xylothamia,a genus of nine shrubby species from northern Mexico, Baja Cali- fornia, and southern Texas, was proposed by Nesom et al. (1990) to accommo- date certain species of Haplopappus (sensu Hall 1928) and certain Ericameria (sensu Urbatsch 1978), plus similar and more recently described taxa. Studies designed to evaluate relationships among North American Astereae using par- simony analysis of external (ETS) and internal transcribed (ITS) DNA sequence data has shown Xylothamia to be polyphyletic (Urbatsch et al. 2003). Four of the nine species in Xylothamia and Gundlachia formed a robustly supported clade in this sequence-based study. The other five species were, in general, un- resolved in a clade that also contained species in the genera Amphiachyris, Bigelowia, Euthamia, Gutierrezia, Gymnosperma,and Thurovia (Urbatsch et al. SIDA 21(1): 243 — 257.2004 244 BRIT.ORG/SIDA 21(1) 2003). Figure | summarizes the phylogenetic relationships among taxa treated in the present study and those related. The tree is derived from figure 4 pub- lished in Urbatsch et al. (2003) where additional, explanatory information may be found. The purpose of this paper is to provide names for species of Xylothamia reflective of their phylogenetic placement in this sequence based study with consideration for their morphological and cytological variation as well. Gundlachia, according to Lane (1996), who last considered its taxonomy, is restricted to the Caribbean region and consists of two species, G. domingensis and G. corymbosa. The former is known from the Bahamas, Cuba, and the Do- minican Republic, while the latter consists of six varieties, and ranges from coastal Venezuela northward through the Greater and Lesser Antilles to the Bahamas and Cuba (Lane 1996). Although Gundlachia had not been included in their cpDNA-based study, Lane et al. (1996) and Nesom (1991, 1993) were in general agreement on its placement within the Gutierrezia lineage sensu Nesom (1993). Gundlachia is expanded herein to accommodate X. diffusa, X. riskindii, X.triantha, and X. truncata. Gundlachia now encompasses six species. Its geo- graphic range is expanded from the Caribbean and nothern South America to include parts of mainland Mexico, the Baja California peninsula, and south- western Texas. Formal nomenclatural combinations for these Xylothamia are made in keeping with the botanical code (Greuter et al. 2000). Generic syn- onymy for Gundlachia as treated herein is also updated because it encompases X.triantha, the type for Xylothamia. With regard to the other five species of Xylothamia, X. johnstonii, and X. palmeri constitute a robustly supported clade (Urbatsch et al. 2003) that is herein proposed as the new genus Neonesomia. The three remaining species of Xylothamia are each treated as monotypic genera because they are not unam- biguously supported as monophyletic or placed within existing genera based on DNA sequence data (Urbatsch et al. 2003), and they are each morphologi- cally unique. Chihuahuana, Medranoa, and Xylovirgatd are the generic names proposed for each of the three taxa. Gundlachia as reconstituted in this study and its sister clade containing the five former species of Xylothamia plus Amphiachyris, Bigelowia, Euthamia, Gutierrezia, Gymnosperma, and Thurovia are supported as a monophyletic lin- eage based on ITS/ETS sequences (Urbatsch et al. 2003) and approximates what Nesom (1991) called the Gutierrezia group. Nesom (1991) credited the cpDNA enzyme re a of Suh C982). and Suh and Simpson (1990) for initially helping tod tic. Although Gundlachia had not been part of the oreabone | DNA investigations, Nesom (1991) aligned it with this group because of its sharing several anatomical and morphological features dis- cussed in part by Anderson and Creech (1975). Chrysoma and Sericocarpus were subsequently aligned with the former ten genera of the Gutierrezia lineage (Nesom 1993) which Nesom subdivided groups. Amphiachyris, Gutierrezia, ASTEREAL 245 Gymnosperma, and Thurovia were assigned to the redefined Gutierrezia group. Features shared by most taxa in this groupare reduced pappus, short disc corolla lobes, annual life cycle in certain species, and base chromosome numbers of x = 4,5(Nesom 1993). The remaining six genera were designated the Euthamia group (Nesom 1993) that are typically perennial herbs or shrubs with pappus of capil- lary bristles and a base chromosome number of x = 9. Chrysoma and Sericocarpus in the ETS + ITS sequence based phylogenies were placed more closely to Solidago and allies (Urbatsch et al. 2003) and not part of the Gutierrezia lineage sensu Nesom (1993). Results by Noyes and Rieseberg (1999) for Sericocarpus were similar while Chrysoma was not included in their study. Lane et al. 1996), based on cpDNA restriction site data, provided support for a clade containing six of the ten genera of the Gutierrezia lineage. Chrysoma, Gundlachia, and Sericocarpus were not investigated and X ylothamia constituted part of their Ericameria lineage (Lane et al. 1996). Branch support and decay index scores for Xylothamia’s inclusion in the Ericameria lineage ranged from one to zero (Lane et al. 1996). Sequence support for the Gutierrezia group within the Gutierrezia lin- eage (sensu Nesom 1993) is equivocal based investigations of Urbatsch et al. (2003), depending on optimality criteria and databases analyzed. Amphiachyris, Gutierrezia, and Gymnosperma constitute a robustly supported lineage in the Bayesian ITS + ETS tree. However, Thurovia appears as an unstable member of the Gutierrezia group sensu Nesom (1993). Only when this data set is analyzed with PAUP* ratchet are Thurovia and Amphiachyris sisters (Urbatsch et al. 2003). Otherwise Thurovia was not affiliated with the Gutierrezia group. It was sister to Bigelowia in PAUP® ratchet of the combined ITS + ETS + indels and to Neonesomia in the Bayesian tree resulting from the combined ITS + ETS. Sequence data offered no support for the Euthamia group of the Gutierrezia lineage sensu Nesom (1993). Characters that support and diagnose the Gutierrezia lineage as delineated in Urbatsch et al. (2003) i.e, without Chrysoma and Sericocarpus, include leaves showing xerophytic adaptations such as often being filiform or otherwise re- duced in size, somewhat coriaceous in some taxa, usually punctate and resin coated, and often supporting a variety and abundance of trichomes; capitula generally small, borne in sessile clusters or on short peduncles; phyllaries ba- sally indurate with a thickened apical patch; stamen insertion below the apex of disk corolla tube; cypselae small, cylindrical to turbinate, and often sericeous NOMENCLATURAL TREATMENT 1. Chihuahuana Urbatsch & R-P. Roberts, gen. nov. Type: Ericameria purpusii Brandegee, Univ. Calif. Publ. Bot. 4:191. 1911. = Chihuahuana purpusii (Brandegee) Urbatsch & R.P. Roberts, combination made herein]. Fruticulus ramosus, folia lineari-lanceolata ca. 2 mm longa acuminata in axillis filiorum 3-10 mm 246 BRIT.ORG/SIDA 21(1) longorum linearium acuminatorum decurrenti demum cinereorum spinescentium fasciculatis; capitula ramos terminantibus 6- cae ted ere Seecclh , dicceidee corollae 8-10, luteola in lobos ovato- lanceolata inaequaliter sectis. Densely branching shrubs to 30 cm tall; stems mostly concealed by the persis- tent closely spaced leaves with small axillary clusters; leaves sessile, acicular, narrowly triangular with a thickened midrib composed mainly of a large bundle of fibers, erect, 3-10 mm long, ca. 1 mm broad at base, hirtellous due toa dense covering of uniseriate conic trichomes, bases somewhat decurrent, api- ces sharply acute; capitula discoid, solitary, campanulate, 4-5 mm wide; phyl- laries graduated, the inner 5-6 mm long; eradiate; disk flowers 8-10, corollas hairy 4.2-5.0 mm long; cypselae densely sericeous; x = 9. Prominent features, distribution, and relationships. —Chihuahuana’s needle- like, non-resinous, hirtellous leaves with a large midvein consisting of a bundle of fibers is unique in the Gutierrezia clade where its relationships are not fully resolved (Urbatsch 1975; Urbatsch et al. 2003). Etymology.—The generic name was selected because it is a member of the Chihuahuan Desert flora. — la. Chihuahuana purpusii (Brandegee) Urbatsch & RP. Roberts, comb. nov. BASIONYM: Ericameria purpusii Brandegee, Univ. Calif. Publ. Bot. 4:19]. 1911. Haplopappus [Aplopappus] purpusii (Brandegee) S.F Blake. Contr. U.S. Natl. Herb. 23:1491.1926. Xylothamia purpusii BeandegesG L. Nesom, Sida 14:112 1990. TYPE: MEXICO. COAHUILA: Cerro de Macho, Jun 1910, Purpus 4479 (HOLOTYPE: UC)). Distribution, ecology, and relationship.—Chihuahuan Desert region in south- western Coahuila and adjacent locales in Chihuahua and Durango, rocky hills in gypseous or limestone soils (Nesom et al. 1990). Detailed descriptive infor- mation and distributional data for this species (as X. purpusii) are provided in Nesom et al. 1990). Chihuahuana purpusii was originally described in Ericameria then trans- ferred to Haplopappus by Blake (1926). Hall (1928) placed the species in Haplopappus section Asiris with five other species mostly of Great Basin dis- tribution. This species was then reinstated in Ericameria (Urbatsch 1978) and subsequently placed in the newly created genus X ylothamia (Nesom et al. 1990). Foliar anatomy, morphology, and flavonoid profiles for C. purpusii are unique (Urbatsch 1978; Urbatsch et al. 2003). ETS/ITS sequence-based phylogenies in- dicate the affinities of C. purpusii are with the Gutierrezia clade rather than with species in section Asiris (Urbatsch et al. 2003). 2. Gundlachia A. Gray, Proc. Amer. Acad. Arts 16:100. 1880. TyPE: Solidago domingensis Spreng., Syst. Veg. 3:539. 1826 Xylothamia G.L. Nesom, Y.B. Suh, D.R. Morgan & B.B. Simpson ae 14: es mise! o- PE: Aplopappus aie en Blake, ]. Wash. Acad. Sci. 28:485. 1938. Gund! t Blake) Urbatsch P. Roberts. AND NEW GE AIITCACAC 247 Evergreen shrubs to 2.0 m tall, stems unbranched to abundantly branching, angular to minutely ridged; leaves usually evenly spaced along the stems, spreading to occasionally appressed, sessile to short petiolate, narrowly to broadly linear to obovate, flat to involute-terete, punctate, resinous; capitula usually clustered at branch tip, clusters few and sometimes hidden by the sub- tending leaves or numerous and organized into racemes or paniculate to corymbiform capitulescences; involucres cylindric, turbinate, or narrowly obconic, phyllaries 2-5 seriate, linear-lanceolate to ovate with an apical resin pocket, basally chartaceous, indurate, margins translucent; ray flowers 0-13, corollas 1-6.5 mm long, white to yellow; disk flowers 3-50, corollas 4.0-5.5 mm long, white to yellow, lobes L.0-2.5 mm long, slightly irregular, usually laxly recurved, style branches 1.0-2.2 mm long, linear lanceolate to ovate, ratio of appendage/stigmatic length variable with different species; cypselae turbinate to cylindrical, LO-2.5 mm long sparsely to densely pilose to sericeous: pappus of ca. 40, slender, flattened, barbellate bristles, 3.5-5.0 mm long; x =9 Prominent features, distribution, and relationships—Gundlachiaas treated in this study consists of six species known from the Caribbean, northern South America, northern Mexico, Baja California, and southwestern Texas. Its mono- phyly was robustly supported by DNA sequence data asa lineage sister to one consisting of several other taxa primarily from western North America. All taxa in this genus are shrubs usually with resinous, punctate leaves that are linear lanceolate to spatulate in three species, G. corymbosa, G. domingensis, and G. riskindii and filiform in the other three. Capitules« ences are often large, corym- bose to paniculate in G. corymbosa, G. domingensis, and G. diffusa; they are more reduced and cymose in X. triantha. Gundlachia riskindii has solitary capitula whereas G. truncata has two or three capitula clustered at their twig apices. Phyllaries in Gundlachia are resinous to glutinous and basally indurate. All species are xeropytes. Gundlachia corymbosa and G. domingensis in- habit mainly sandy or stony soils in or near coastal habitats of various Carib- bean islands, while G. diffusa occupies similar habitats of the Gulf of Califor- nia region of Sonora and Baja California. Gundlachia riskindii appears to be restricted to exposed limestone areas in pine-oak woodland at around 2100 m in Coahuila and adjacent Nuevo Leon, Mexico. The other two species occur in the Chihuahuan Desert region associated with elements more typical of that flora such as Atriplex, Larrea, Prosopis, Suaeda, etc. Specific descriptions, dis- tributional data, and supporting documentation for the species of Xylothamia here considered as Gundlachia are provided in Nesom et al. (1990). Similar data for Gundlachia truncata asa species of Xylothamiacan be found Nesom (1992). Lane (1996) provided detailed species descriptions, distributional data, other pertinent information, and keys to varieties of G. corymbosa. There is no indication whether the Gundlachia clade first evolved in the Caribbean, in Mexico, or elsewh ince species relationships within it are not 248 BRIT.ORG/SIDA 21(1 = well-resolved (Urbatsch et al. 2003). The clade sister to Gundlachia consists of North American taxa. These two clades taken together also appear to be related to North American species, although few Central and South American species have been investigated (Urbatsch et al. 2003). Regardless of where the Gundlachia clade first appeared, dispersal rather than plate tectonics must have been a factor in its evolution since the approximate present position of the Car- ibbean islands relative to North and South America predates the estimated age of the Asteraceae (Graham et al. 2000; Bohm & Stuessy 2001). KEY 10 SPECIES OF GUNDLACHIA _Leaves linear-lanceolate to spatulate, laminar, more than 2 mm wide. 2. . typically solitary at branch tips; plants of Coahuila and Nuevo Leon, xicO G. riskindii 2. ae generally in large, dense, corymbose or paniculate clusters several cm broad; plants of the Caribbean 3. Capitula in corymbose clusters; involucres cylindric; ray corollas <4 mm long; Bahamas, Greater and Lesser Antilles G. corymbosa 3, Capitula in paniculate clusters; involucres obconic or turbinate; ray corollas >4 mm long; plants of northern Bahamas, Cuba, and Hispaniola 1. Leaves filiform, less than 2 mm wide, nearly as thick as broad 4, Leaves densely clustered along the terminal 2-3 cm of the branches, surfaces glutinous, not evidently punctate; capitula eradiate, in clusters of 2-3, mostly concealed by foliage; disk flowers 3-5; Coahuila, Mexico, Cuatro Cienegas basin G. domingensis G. truncata 4, Leaves more widely spaced on stems, internodes generally much >2 mm long, surfaces resinous, Somewhat punctate; capitula iat or rays 1-3, rays mostly concealed by the involucre; disk flower 5. Stems glabrous; ray flowers 0-3; coat ee of the Gulf of California re gion, Baja California and Sonora, Mexic G. diffusa 5. Stems papillate, scabrous; rays absent; an flowers 3(-7); widespread in the Chihuahuan Desert of southwestern Texas, Chihuahua, Coahuila, Durango, and Nuevo Leon, Mexico G. triantha New specific combinations in Gundlachia 2a. Gundlachia triantha (S.F Blake) Urbatsch & R-P. Roberts, comb. nov. BASIONYM: Aplopappus{Haplopappus] trianthus SF Blake, J. Wash. Acad. Sci. 28:485. 1938. Ericameria triantha (S.F Blake) Shinners, Field & Lab. 19:133. 1951. Xylothamia triantha (S.F Blake) G.L. Nesom, Sida 14:113. 1990. Type: UNITED STATES. TEXAS. BREWSTER Co.: Chisos Mountains area, along road on Study Butte to Terlingua, 31 Aug 1937, B.H. Warnock 1126, (HOLOTYPE: USI ISOTYPE: LL). = Distribution, ecology,and relationships.—This species is widespread in the Chihua- huan Desert region ranging from Brewster County, Texas, to eastern Chihuahua, Coahuila, northeastern Durango, and west central Nuevo Leén, Mexico. It grows on gyp-seous, calcareous, igneous, or saline soil substrates on slopes or desert flats generally associated with Atriplex, Prosopis, Larrea, Suaeda,and various desert spe- cies at elevations 700-1500 m. Flowering normally occurs from July to October A F ASTEREAE 249 which undoubtedly depends on the timing and abundance of rainfall. Its resinous, d of ten three-flowered capitula are diagnostic. Based on sequence data G. triantha and G. truncata are strongly supported as sisters (Urbatsch et al. 2003) (see Fig. 1). Nesom (1992) indicated that the two are similar in having linear, involute leaves, eradiate, few-flowered capitula, and similar style appendages. Gundlachia diffusa receives moderate support as the sister to this clade in most analyses except for parsimony analysis of the ETS + ITS data sets where it is sister to G. domingensis (Urbatsch et al. 2003). When describing G. triantha Blake, (1938) discussed its morphological similarity to G. diffusa. Urbatsch (1978) noted the similarity in flavonoid profiles between these two taxa. At the time these studies were made G. truncata was unknown. Nesom et al. (1990) commented on the Euthamia-like inflorescences of G. triantha and G. diffusa. Sequence data show that Euthamia is but distantly re- lated to these taxa, indicating possible convergence in this feature (Fig. 1). 2b. Gundlachia diffusa (Benth.) Urbatsch & R.P. Roberts, comb. nov. BASIONYM: Ericameria diffusa Benth., Bot. Voyage H.M.S. Sulphur 2:23. 1844. non Aplopappus diffusus DC. 1836. Solidago diffusa (Benth.) A. Gray, Proc. Amer. Acad. Arts 5:159. 1861. Bigelovia diffusa (Benth.) A. Gray, Proc. Amer. Acad. Arts 8:640. 1873. Chrysoma diffusa (Benth.) E. Greene, Erythea 3:10. 1895. Xylothamia diffusa (Benth.) GL. Nesom, Sida 14:109. 1990. Type: MEXICO. BAJA CALIFORNIA SuR: Magdalena Bay, 1839, R. Barclay & B. Hinds s.n. (HOLOTYPE: BM?). adi-axially caniculate, filiform leaves, Linosyris sonoriensis A. Gray, Proc. Amer. Acad. Arts 8:291. 1870. Basionym: Haplopappus [Aplopappus] sonoriensis (A. Gray) S.F. Blake, Contr. U.S. Natl. Herb. 23:1490. 1926. Aster sonoriensis (A. Gray) Kuntze, Rev. 1:317. 1891. Type: MEXICO. Sonora: District of the Yaqui River, 1869, FE. Palmer s.n. (HOLOTYPE: GH)). Distribution, ecology, and relationships.—Gundlachia diffusa occurs in Baja Cali- fornia Sur and coastal regions of Sonora, Mexico, where it inhabits various soil types including coastal sand dunes, gravel plains, and salt flats from near sea level to around 100 m. Its habitat preferences appear to be similar to the two species of Gundlachia from the Caribbean. Based on leaf morphology the spe- cies resembles the two Chihuahuan Desert species in this genus. When well- developed, its inflorescences can be large and paniculate and appear similar to those in G. corymbosa. Its species relationships are somewhat equivocal but appear to be closest to G. triantha and G. truncata and are discussed under to the former. 2c. Gundlachia riskindii (B.L. Turner & Langford) Urbatsch & R.P. Roberts, comb. nov. BASIONYM: Ericameria riskindii B.L. Turner and Langford, Madronio 29:234. 1982. Xylothamia riskindii (B.L. Turner and Langford) G.L. Nesom, Sida 14:113. 1990. Type: MEXICO. COAHUILA: ca. 24 km E of Saltillo, S side of Sierra La Viga, ca. 6.5 km E of Jamé along wood cutter’s road, 10,00 ft, 15 May 1977, Henrickson et al 16156b (HOLOTYPE: TEX! ISOTYPE: MEXU, RSA). Distribution, ecology, and relationships.—This apparently rare species is known from southeastern Coahuila and adjacent Nuevo Le6n where it occurs on ex- 250 BRIT.ORG/SIDA 21(1) posed limestone areas and in gypseous soils in pine-fir-oak woodlands at 2100- 3000 m elevation. It is distinctive in having small spatulate leaves and radiate capitula with a large number of disk and ray florets. Originally, G. riskindti was described as Ericameria but then transferred to Xylothamia. Concerning its sister taxon relationships, results differ depending optimality criteria used in DNA sequence analysis. In the PAUP ratchet analysis of the combined ITS/ ETS sequences support is provided for its basal position in the Gundlachia clade. In the parsimony derived trees that included indels, the Caribbean and Mexi- can species are resolved as sister lineages, where it is basal to the latter (Urbatsch et al. 2003). When indels were excluded, the species is basal in the Gundlachia clade (Fig. 1). Flagelliform trichomes having a subterminal appendage attach- ment characterize the Caribbean species and similar trichomes are seen in G. riskindii. This unusual trichome type, along with spatulate leaves and certain DNA evidence suggest that G. riskindit may represent the ancestral state for Gundlachia or may bea link connecting the Caribbean and the Mexican species 2d. Gundlachia truncata (G.L. Nesom) Urbatsch & R.P. Roberts, comb. nov. BASIONYM: Xylothamia truncata G.L. Nesom, Phytologia 73:318. 1992. TyPE: MEXICO. COAHUILA: Mpio. Cuatro Cienegas, ca. 2 km W of town of Cuatro Cienegas, along dirt road paralleling railroad, hard packed gypseous sand, 18 Oct 1985, Nesom 5254 (HOLOTYPE: TEX). This rare species is know only from the Cuatro Cienegas basin in Coahuila, Mexico, where it was collected on alluvial, gypseous sands. It is readily recog- nized by its involute linear leaves crowded near the branch apices that largely concealing its flowering capitula (Nesom 1992). The nearly identical ITS/ETS sequences of G. truncata and G. triantha robustly support their sister relation- ship despite their distinctive morphologies. Nesom (1992) had suggested the possibility of G. truncata being an abnormal growth form of the later, but field observations offered no evidence supporting this hypothesis. Distribution, ecology, and relationships—Both Gundlachia corymbosa and G. domingensis grow in the Caribbean region. The former occurs in eastern Cuba, the Bahamas, the Greater and Lesser Antilles, and northern Venezuela. It is dis- tinguished from its counterpart by its corymbose capitulescences, cylindric involucres, ovate phyllaries, and ray corollas shorter than 4 mm. Six varieties are recognized within the species. Three of these, G. corymobsa varieties apiculata, cubana, and foliosa, have corymbs reduced toa few capitula concealed by the leaves, which is a feature seen in two species in Mexico, G. riskindii and G. truncata. The leaves of G. corymbosa var. apiculata, a taxon from the moun- tains of eastern Cuba, are similar in form to those of G. riskindii but larger. Gundlachia domingensis is known from the northern Bahamas, Cuba, and Hispaniola and is distinguished by its paniculate capitulescences, turbinate involucres, linear-lanceolate phyllaries, and ray corollas about 5.0 mm long, Both species appear to grow on well-drained sandy and stony substrates from near 251 ATCCLU AMIN DADcCoOTC CAC F ASTERCAL G 198 Te — Haan johnstonii Xj] Palme | emeoma) X X. palmeri 100/11 X. parrasana 1 ae ccharis 100/15 60/2 ap f Fealdeccharis 2 (Xylovirgata) [1] “5 ai Bua 2 19 (Chihuahuana) ooig F ig ia ia ia 4 61/3 apne SH gpa sada a Canadensis 5 fistutosa | S. fauics S. Meee Chrysothamnus depr essus 52itt C. viscidiflorus ee Sericocarpus Ericameria c i Ericameria ericoaides Ericameria t Ericameria nau Traydna { Batopilasia; Boltonia; Chloracantha H aplopappus; H azardia; | socoma; L Machaeranthera; Saas ay eee Xanthisma; Xanthocephalum; iza { Chaetopappa; Croptilon; Erigeron | onactis Majority rule Fig. 1. 50% “ 1 iL DALID¥ vate nt ly ’ +h hi AcT¢ r We Ast + ah cutindels and based onFig.4p blished in Urbatsch et al. (2003). F | bers desi k h support/branch | in bold highligt Jitionally t d as Xylothamia. Dash line | hes highlight taxa tradi- tionally treated as Gund hia.G i proj 1 in the j y indi 1 hetically. Taxa of the | t clad i fi | | | | ly li 1 fi | j lineage. 252 BRIT.ORG/SIDA 21(1) sea level to 2500 m elevation. Greater details for these two species are provided in Lane (1996). 3. Neonesomia Urbatsch & RP. Roberts, gen. nov. Type: Aster palmeri A. Gray, Proc Amer. Acad. Arts 17:209. 1882. = Neonesomia palmeri (A. Gray) Urbatsch @ RP Roberts. Frutices ad 3 m alta; caules ramosissimi aliquantum aromatici, ramunculis dune va porcatis; folia 1-4 cm longa 1-2(-5) mm lata linearia elliptica ad anguste oblanceolat cum costis elevatis abaxialibus; capitula radiata solitaria vel in cymis oie ie apices ramulorum, paniculis laxis; involucra plerumque turbinata 4-6 mm puekee oe ia valde gradata linaria; flosculi radii 5-15, corollae ca. 2-5 mm longa alba ad flava; 8-20, corollae 3.5-5 mm longa alba ad flava, limba aliquantum asymmetrice 5-lobata; pappi setosi; cy se ca. L.5 mm longae dense strigosae ad sericeas. Shrubs to 3 m tall; much-branched, somewhat aromatic, leafy twigs strongly ridged, internodes 2-10 mm long; leaves 1-4 cm long, l-2(-5) mm broad, linear, elliptic to narrowly oblanceolate, blades flat with conspicuously raised midvein abaxially, pubescent with flagelliform trichomes, margins entire to minutely ciliate; capitula solitary or in cymose clusters at branch tips these arranged in loose panicles; involucres usually turbinate, 4-6 mm long; phyllaries strongly graduated, linear, blunt to somewhat acute, thick, firm, resinous, mostly stramineous but with an apical glandular patch occupying much of the tip region; rays flowers 5-15 corollas ca. 2-5 mm long, white, pale yellow, or yellow; disk flowers 8-20, corollas 35-5 mm long, with a tube |-1.8mm long and poorly differentiated from the asymmetrical 5-lobed limb, shorter lobes 0.9-1.5 mm long, longest up to 3.3 mm long; pappus of setose bristles; cypselae ca. 1.5 mm long, densely strigose to sericeous; x = Prominent features, distribution, and relationships.—Neonesomia contains two species, N. palmeri from Nuevo Leon, Tamaulipas, and southern Texas and N. johnstonii from San Luis Potosi, which grow on rocky hillsides, brushy shrublands, and coastal dunes Johnston 1970; Nesom et al. 1990). Species in this genus are characterized by their shrubby habit, ridged twigs, flat leaf blades with prominent midveins, small radiate capitula, and white to yellow corollas with somewhat zygomorphic disc corollas. They resemble the herbaceous pe- rennials in Euthamia in leaf and growth form, but differ in being non-rhizoma- tous shrubs with deeply lobed, somewhat zygomorphic disc corollas. Etymology.—N muy Nesom, Botanical Research In- stitute of Texas, zealous pendent of the Astereae and other Asteraceae who has sig- nificantly contributed t ling their systematics. Additional information for these taxa is found in Nesom’s treatment of Xylothamia (Nesom et al. 1990). The genus Neonesomia is strongly supported as a clade in the gene-based phylogenies of Urbatsch et al. (2003). Based on parsimony analyses of ITS +ETS + INDEL data it is basal to a clade consisting of several xerophytic, mainly west- ern North American taxa, including Amphiachyris, Bigelowia, Chihuahuana, ARID. NICIAY CEN EDA AC ACTECDEAL ASTERCAC 253 Euthamia, Gutierrezia, Gymnosperma, Medranoa, Thurovia, and Xylovirgata (Urbatsch et al. 2003). Weak support for its sister relationship to Thurvoia was seen in the ITS/ETS Bayesian tree derived in that study. Otherwise its sister re- lationship was unresolved (Fig. 1) and kinship among the above cited genera is uncertain (Urbatsch et al. 2003). KEY TO SPECIES OF NEONESOMIA = . Ray and disk flowers 12-15 and 15-20, respectively; corollas yellow; flowering May to June; San Luis Potosi, Mexico johnstonii . Ray and disk flowers 5-11 and 9-14, respectively; corollas white to very pale yellow; mainly flowering August to October; southern Texas and Nuevo Leon and Tamaulipas N. palmeri — New combinations in Neosomia 3a. Neonesomia palmeri (A. Gray) Urbatsch & RP. Roberts, comb. nov. BASIONYM: Aster palmeri A. Gray, Proc. Amer. Acad. Arts 17:209. 1882. TYPE: UNITED STATES. TEXAS. MAVERICK Co. Eagle Pass on the Rio Grande, Sep-Oct, 1879. E. Pa Imer 516 (LECTOTYPE: GHI: ISOLECTOTYPES: PH, US; Johnston 1967). Isocoma palmeri (A. Gray) Shinners, Field & Lab. 18:2. 1950. Ericameria austrotexana M.C. Johnston, nom. nov, SouthW. Naturalist 12:106. 1967, non E. palmeri (A. Gray) H.M. Hall. Xylothamia palmeri (A. Gray) G.L. Nesom, Sida 14:110. 1990, non X. palmeri var. pachylepis (H.M. Hall) G.L. Nesom ex M.A. Lane & R.L. Hartman, Amer. J. Bot. 83:364. 1996 Distribution, ecology, and relationships.—This species is known from most coun- ties in southern Texas, ie., Atascosa, LaSalle, and San Patricio cos. southward into the states of Nuevo Leon and Tamaulipas, Mexico, mostly of open, brushy habitats on or near the Gulf Coast in sandy or saline soils from near sea level to 600 m. Neonesomia palmeri is distinguished from its sister species in having fewer flowers per capitula and paler yellow corollas. Its ITS/ETS sequences showed little differentiation from N. johnstonii in Urbatsch et al. 2003). Neonesomia palmeri was first described asa species of Aster and subsequently slaced in various other genera including Isocoma (Shinners 1950), Ericameria Johnston 1967), and Xylothamia (Nesom et al. 1990). Nesom et al. (1990) pro- posed a close relationship of N. palmeri to Medranoa (Xylothamia) parrasana and Xylovirgata(X ylothamia) pseudobaccharis, a hypothesis not robustly sup- ported by analyses of DNA sequences. 3b. Neonesomia johnstonii (G.L. Nesom) Urbatsch & RP. Roberts, comb. nov. BASIONYM: Xylothamia johnstonii G.L. Nesom, Sida 14:110. 1990. TYPE: MEXICO. SAN Luis Potosi: Bagre, Minas de San Rafael, May 1911, C.A. Purpus 5021 (HOLO- TYPE: GH! ISOTYPE: US). Distribution, ecology, and relationships.—This taxon is known from central San Luis Potosi, Mexico where it apparently grows on steep slopes at elevations of 120-1500 m. In addition to the diagnostic features presented under N. palmeri, flowering times also differ with N. johnstonii blooming in the spring and N. 254 BRIT.ORG/SIDA 21(1) palmer‘ in the fall. Habitat preferences for the two taxa differ: N. johnstonii grows on steep hillsides at higher elevations than N. palmeri, which inhabits the Gulf coastal plain often on sand dunes near the water. Based on sequence data N. johnstonii and N. palmeri form a robustly supported clade, but the two taxa differ little from one another in sequence data indicating their close affinity (Urbatsch et al. 2003). 4. Medranoa Urbatsch & Roberts, gen. nov. Typr: Ericameria parrasana S.F Blake, Contr. Gray Herb. 52:26. 1917. = Medranoa parrasana (S.F Blake) Urbatsch & RP. Roberts. ] ‘ ti ‘ lak 1 ‘Et Caan pF a i lloso- al andulosis viscosis non puberulis dae foliosis; folia linearia mucronulata ut ramuli punctata et viscose complanata supra suplana vel paullum concave sutus subconvexa; involucri 3-seriati Sc aies gradati = 5- i mm alti pe a supra glandulari-viscosa ceterum subglabra infra valde = subherbacea lanceolata munita. Shrubs to 2 m tall, ie ee ee ascending, mostly terete or remotely ribbed, bark smooth, becoming slightly fissured, tan becoming dark gray; twigs numerous, to 8 cm long, internodes smooth to obscurely sulcate, 1-4 mm long, green resinous, scabridulous; leaves evergreen, sessile, crowded, 5-12 mm long =< 1.5mm wide, narrowly elliptic-oblanceolate, flat to canaliculate, decurrent on stem, surfaces resinous, punctate; capitulescence somewhat corymbiform, capitula solitary or cymose at branch tips; involucres 2-3 seriate, cam panulate, 3-5 mm high, 4-7 mm wide; phyllaries imbricate, moderately graduated, mostly chartaceous, 2.5-4 mm long; capitula radiate, with ca 30 flowers, ray flowers 5- ll, pappus bristles ca. 80, setose, subequal, 2.5-4 mm long, silvery tan, corollas 5-8 mm long, ligules elliptic, 3-7 mm long, 1-2 mm wide; disk flowers ca 20, corollas pale yellow, 4-5 mm long, lobes spreading to recurved, unequal in length, shortest 0.8-12 mm long, longest 1.5-2.2 mm long, style branches 2-3 mm long, appendages narrowly linear, apices acute, 15-18 mm long, pappus same as on ray flowers; cypselae turbinate, ca. 2 mm long, pilose; x = 9. Prominent features, distribution, and relationships—Medranoa is unispecific with its only species occurring in the Sierra de Parras region of Coahuila and Zacatecas. Shrubby habit, deeply pitted, resin coated leaves, relatively large ca- pitula, and thickened style branches are features diagnostic for this taxon. Se- quence based data robustly support its place inthe Amphiachyris/Gutierrezia clade (Urbatsch et al. 2003). Weak support is provided in that study for a sister relationship with Chihuahuana (X. purpusii) or with a clade composed of Chihuahuana plus Xylovirgata (Xylothamia pseudobaccharis). Etymology—The generic name Medranoa is in honor of Francisco Gonzlez Medrano, MEXU. He has worked for many years on desert and dryland floras of Mexico—mostly Tamaulipas and Tehuacan, and has trained several young bota- nists. Additional information about this taxon can be found in the treatment of Xylothamia by Nesom et al. (1990). ASTCREAL 255 4a. Medranoa parrasana (S.F. Blake) Urbatsch @ R-P. Roberts, comb. nov. [BASIONYM: Ericameria parrasana S.F. Blake, Contr. Gray Herb.52:26. 1917]. Haplopappus parrasanus (S.F Blake) S.F Blake, Contr. U.S. Natl. Herb. 23:1490. 1926. Xylothamia parrasana (S.F Blake) G.L. Nesom, Sida 14:111. 1990. TyPE: MEXICO. COAHUILA: Sierra de Parras, rocky slopes, Mar 1905, Purpus 1005 (HOLOTYPE: GHI). Distribution, ecology, and relationships.—The single species in this genus grows on rocky slopes in the Sierra de Parras region of southern Coahuila and northern Zacatecas. Originally it was described in Ericameria by Blake (1917) who later transferred it to Haplopappus (Blake, 1926) Subsequently, it was placed in Xylo- thamia (Nesom et al. 1990). In the DNA Sequence: based trees, it is placed in the clade composed of Amphiachyris, Bigelowia, Chi Euthamia, GutierreZia, Gymnosperma, Neonesomia, and Thurovia (Urbatsch et al. 2003). Medranoa is basal toa clade of X. pseudobaccharis and Chihuahuana purpusii in parsimony analyses of the ETS/ITS/INDEL data set place. It is a weakly supported sister of Chihuahua purpusii in the Bayesian analysis (Urbatsch et al. 2003). Medranoa parrasana is distinguishable from other taxa in the Amphiachyris/Gutierrezia clade by the combination of its shrubby habit, nar- rowly elliptic-oblanceolate, flat-canaliculate, resin-covered leaves with numer- ous, well-organized depressions, and its campanulate capitula bearing 5-11 ray and 15-22 disk flowers. The thickened style branches wherein the vascular trace bifurcates or expands in size distally in each branch is unique among taxa in this clade. Additional information about this taxon can be found in the treat- ment of Xylothamia by Nesom et al. (1990). 5. Xylovirgata Urbatsch & R-P. Roberts, gen. nov. Type: Haplopapus pseudobaccharis SF Blake, J. Wash. Acad. Sci. 40:47.1950. = Xylovirgata pseudobaccharis (S.F Blake) Urbatsch & R-P. Roberts Frutex scoparium metralis glaberrimus modice resinosus, ramis et ramulis multis erectis pallide Scena : ] nga a 1; sas A ] 1 : . Oo o oO ie = vel solum supra obscurissime punctata usque ad 1.4 cm longa 1 mm lata. Intricately branched, broomlike shrubs to 1 m tall; stems slender, bark becom- ing whitish with age; branches and twigs strongly ridged and angled; leaves present mainly on present years growth, widely spaced, somewhat erect, blades 2-15 mm long, <1 mm wide; capitula radiate solitary at branch apices arranged in loose racemes; involucres campanulate to turbinate, 3-4 mm wide, phyllar- ies graduated, the inner 3-5 mm long, thickened subapical glandular structure present; ray flowers 3-6, ligules apically 2-3 denticulate, 2.3-3.0 mm long, ca. | mm wide; disk flowers 7-14, corollas 4.0-4.5 mm long unequally 5 lobed, shorter lobes ca. 1.3 mm long, longer lobes ca. 2.3 mm long; cypselae ca. 1.3 mm long, sericeous; pappus similar on disk and ray cypselae, ca. 30 subequal setose bristles, x= 9. Prominent features, distribution, and relationships.—X ylovirgata is unispe- cific and known only from western Coahuila, Mexico. It is recognized by its 256 BRIT.ORG/SIDA 21(1) broom-like appearance due to its woody, erect, intricately branched habit, as referenced by its generic name, and its conspicuously ridged stems, reduced, widely spaced, inconspicuous leaves. This entity is clearly related to taxa in the Amphiachyris/Gutierrezia clade, where there is weak support for its sister re- lationship to Chihuahuana in trees resulting from parsimony analysis of ETS/ ITS sequence data (Urbatsch et al. 2003). When indel data were added to the data set support for this relationship is supported more robustly. Bayesian and bootstrap analyses resulted in its placement as one of several unresolved basal elements in the Amphiachyris/Gutierrezia clade (Urbatsch et al. 2003). Addi- tional descriptive data for this taxon can be found in Nesom et al. (1990). 5a. Xylovirgata pseudobaccharis (S.F Blake) Urbatsch & R.P. Roberts, comb. nov. BASIONYM: Haplopapus pseudobaccharis S.F. Blake. J. Wash. Acad. Sci. 40:47. 1950. Ericameria pseudobaccharis (S.F. Blake) Urbatsch, Sida 7:299. 1978. Xylothamia pseudobaccharis(S.F Blake) G.L. Nesom, Sida 14:112. 1990. TypE: MEXICO. COAHUILA: arid limestone hills of Sierra Paila, Valle Seco, General Cepeda, 1700 m, 4 Jul 1944, J.C. Hinton (G.B. Hinton et al.) 16546 (HOLOTYPE: USk ISOTYPES: GH!, NY). Distribution, ecology, and relationships.—X ylovirgata pseudobaccharis is known only from western Coahuila, Mexico, where it grows on limestone or gypsum slopes. Its suggested relationships to Neonesomia (Xylothamia palmeri, X. johnstonii), Medranoa (X. parrasana), and Gundlachia riskindii CX. riskindii) (Nesom et al. 1990) are not supported by sequence-derived phylogenies (Urbatsch et al. 2003). Sequence data provides some support for its sister rela- tionship with Chihuahuana purpusii. Additional descriptive and distributional data for this taxon as Xylothamia pseudobaccharis are presented in Nesom et al. (1990). ACKNOWLEDGMENTS We thank Kurt Neubig for providing the Latin diagnoses, Tom Wendt (LL, TEX) for his assistance in providing materials for DNA extraction and other her- barium matters, and to herbarium curators at BRIT, GH, MICH, MO, NY, UC. and US for their assistance. We appreciate Guy Nesom reviewing the manuscript. REFERENCES ANbeRSON, L.C. and J.B. CreecH. 1975. Comparative leaf anatomy of Solidago and related Asteraceae. Amer. J. Bot. 62:486-493. Buake, S.F.1917.New and noteworthy Compositae, chiefly Mexican. Contr. Gray Herb. 52: 16-59, Biake, S.F. 1926. In: PC. Standley. Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23: 1313-1721. Buaxe, S.F. 1938. Eleven new American Asteraceae. J.Wash. Acad. Sci. 28:478-492. Boum, B.B and TF. Stuessy. 2001. Flavonoids of the sunflower family (Asteraceae). Springer- Verlag, Vienna, Austria. ASITCREAC 257 Granam, A., D. Cozano, A. Areces-MALLes, and N.O. Freberiksen. 2000. Studies in Neotropical paleobotany. XIV. A palynoflora from the middle Eocene Saramaguacan Formation of Cuba. Amer. J. Bot. 87:1526-1539. GreuTer, W., CHAIRMAN OF THE EDITORIAL COMMITTEE, ET AL. 2000. International code of botanical nomenclature (St. Louis Code). Regnum Veg. 138. Koeltz Scientific Books, Konigstein Hatt, H.M. 1928. The genus Haplopappus: a phylogenetic study in the Compositae. Carn- egie Inst. Wash. Publ. 326:i-viii, 1-391. JOHNSTON, M.C. 1967. Ericameria austrotexana M.C. Johnston (Compositae), nomen novum. SouthW. Naturalist 12:106-109. JouNsToN, M.C. 1970. Ericameria. In: Correll, D.S.and M.C. Johnston. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Texas. Pp. 1577-1 D1 ox Lane, M.A. 1996. Taxonomy of Gundlachia (Compositae: Astereae). Brittonia 48:532-541. Lane, M.A.,D.R.Morcan, Y. SUH, B.B. Simpson, and R.K. Jansen. 1996.Relationships of North Ameri- can genera of Astereae, based on chloroplast DNA restriction site data. In: D.J.N. Hind and H.J. Beentje, eds. Compositae:systematics—Proc. Int. Compositae Conf, Kew, 1994. Royal Botanic Gardens, Kew, UK. Vol. 1:49-77 Nesom, G.L.1991.Morphological definition of the Gutierrezia group (Asteraceae: Astereae). Phytologia 71:252-262. Nesom, G.L. 1992. A new gypsophilic species of Xylothamia (Asteraceae: Astereae) from the Cuatro Cienegas area of Coahuila, Mexico. Phytologia 73:318-320. Nesom, G.L. 1993. Taxonomic infrastructure of Solidago and Oligoneuron (Asteraceae: Astereae) and observations on their phylogenetic position. Phytologia 75:1-44. Nesom, G.L., Y. Sux, D.R. Morcan, and B.B. Simpson. 1990. Xylothamia (Asteraceae: Astereae), a new genus related to Euthamia. Sida 14:101-116. Noves, R.D. and L.H. Rigseserc. 1999. ITS sequence data support a single origin for North American Astereae (Asteraceae) and reflect deep geographic divisions in Aster s./. Amer. J. Bot. 86:398-41 2. Suinners, L.H. 1950. Notes on Texas Compositae-lV. Field & Lab. 18:25-32. Sun, Y. 1989. Phylogenetic analysis of chloroplast DNA in North American Astereae (Asteraceae) based on chloroplast DNA. Ph.D. dissertation, University of Texas, Austin, Texas, USA. Sux, Y and B.B. Simpson. 1990. Phylogenetic analysis of chloroplast DNA in North American Gutierrezia and related genera (Asteraceae: Astereae). Syst. Bot. 15:660-670. Urearscu, L.E. 1975. First chromosome number reports for some Compositae. SouthW. Naturalist 20:271-287. Ureatscu, LE. 1978. The Chihuahuan Desert species of Ericameria (Compositae: Astereae). Sida 7:298-303. Urearscn, L.E., R.P. Rogerts, and V. Karaman. 2003. Phylogenetic evaluation of Xylothamia, Gundlachia, and related genera (Asteraceae, Astereae) based on ETS and ITS nrDNA sequence data. Amer. J. Bot. 90:634-649. 258 BRIT.ORG/SIDA 21(1 BOOK NOTICES FLORAS AND CHECKLISTS NaTHAN SMITH, Scott A. Mort, ANDREW HENDERSON, DENNIS WM. STEVENSON and Scott a ALI p(eds,) 2004. Flowering Plants of the Neotropics. (ISBN 0-691-11694- Tee Princeton University Press, 41 William Street, Princeton, NJ 08540 S.A. (Orders: 609-258-5714, 609-258- 7. fax). $75.00, 616 pp., 307 eolet ane 258 line illus., 6 tables, 81/2" x 11", The American tropics is one of the great hotspots of ie diversity, harboring 35% of the world’s flowering plants. This magnificent new book from Princeton University Press provides standard treat- ments for more than 280 families of flowering plants that occur in this region, and is the culmina- tion of the work of 150 botanists from around the world. The goal of the editors was to provide an authoritative reference to plant families that are known to occur in tro apc! America. T ne h lave ac- complished their goal, as this book will be an essential ref earn- ing about the rich alice As the neotropical flora. a text a ccoriees features of each family, the diversity ol g classification geogray | natural history. and import int uses. Students learning troy sicalfl {lowering plant famili d their cl ill fine fea tures and descriptions especii ay) useful. The excellent identification key to the families treated in Isto Id i ppendix V will be of specie) interest toanyone who wishes to learn to identify ian owering plant families. The editors al other useful refer- the book and the Ai ence oe suena asa pens botanical glossary, and comparisons - Cais concepts across the vz ncluding Cronquist, Dahlgren, and Judd et al. More than 300 color the photographs and 250 botanical i drayanes provide expert visual aids for identifying and learn- ing about the families —John Janovec, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth. TX 76102-4060, U.S.A RJ. PETHERAM and B. Kox. Photography by E. Bartlett-Torr, 2003. Plants of the Kimberley Region of Western Australia. Revised Edition. (ISBN 1-920694- 04-8, pbk.) University of Western Australia Press. (Orders: International See Book Services, 920 NE 58" Avenue, Ste. 300, Portland, OR 97213- , 303-287-3093, 503-280-8832 fax, www.isbs.com, email: a ens $40.50, 553 pp., color photos, 6" 8 1/2". Publisher Comments: “For many years Plants of the Kimber ley Region of Western Australia has been an important resource for pastoral managers and rangeland advisors in managing vegetation and and-use issues. This revised edition includes changes to 50 on names, and also updates the intro- luctory sections about the Kimberley region and the principles of ‘rangeland management. The 240 ys species covered in the book are organised in three sections: grasses and herbs (110), shrubs (40) and trees (90), and constitute a unique flora not dealt with in any other single volume.” Wi hee ] ith its stre aightforward text and excellent photographs this book will also be a valuable ref- erence lor students of ecology and range science, as well as appealing to nature-lovers, conservation- ists and travellers in the Kimberley region.” CHARLES M. ALLEN, DAWN ALLEN Newman, and Harry WINTERS. 2004. Grasses of Louisiana. Third Edition. (ISBN 0-9718625-1-6, pbk.). Allen’s Native Ven- tures, LLC, 5070 Hwy 399, Pitkin, LA 70656, U.S.A. (Orders: 337-328-2252, WwWww.nativeventures.net, email: native@c Lnet). $20.00, 374 pp.,3 maps, b/w line drawings, 7" x 10". SIDA 21(1): 258. 2004 A NEW SPECIES OF CALYCADENIA (ASTERACEAE) FROM NORTH CENTRAL CALIFORNIA Robert L. Carr Gerald D. Carr Department of Biolo Department of a Eastern Washington University University of Hi Cheney, Washington 99004, U.S.A. 31 rcarr@ewu.edu Honolulu, Hawaii 96822, U.S.A. gerry@hawaii.edu ABSTRACT An unusual form of Calycadenia from north central California was studied in greenhouse culture and confirmed to be self- oe a eoueison rare in Caco aed, aces on the distinctive mor- phology and breeding system of this species, Calycadenia micrantha. Although ae ee to Ci truncata and apparently included in C. truncata DC. subsp. microcephala H.M. Hall ex D.D. Keck, this name was rejected as a pease for the new species be- cause of ambiguities sere type, the original description | 1 and/or anno- tated by Keck. RESUMEN c lia ¢ : 1 Pe Perea | fonortedeCalit : ie ae ] y se confirm6 que es autocompatible, una caracteristica rara en Calycadenia. Se describe e ilustra como una nueva especie pasate en la diferente mre y sistema reproductor de este taxon, ie truncata y aparentemente incluida en C. truncata DC. subsp. microcephala H.M. Halle ex Re Be este nombre fue rechazado como basionimo oo la nueva espec cie ae: el tipo, la descripcion . I ] original, y | I ) por Keck. INTRODUCTION For some years we have been aware of a unique population of plants related to Calycadenia truncata and growing on Elk Mountain in Lake County, Califor- nia. In 1990 we received a similar specimen from David W Isle, Forest Botanist for the Mendocino National Forest in California. The plant had been collected near the Wilson Camp area of southern Colusa County. Morphologically, the plant was clearly related to Calycadenia truncata. However, like the Elk Moun- tain specimens, Isle’s plant was shorter and more slender than most and had very tiny heads with only 1 or 2 obscure, tiny ray flowers and 1 or 2 disk flowers. Subsequently, several additional populations of this taxon were located in adjacent Lake County. The heads and rays suggested the possibility of self-com- patibility, a condition rare in Calycadenia. In conjunction with ongoing bio- systematic research on Calycadenia,a number of populations of this taxon were studied in greenhouse culture. All individuals were self-compatible, a condi- tion known to exist in only one other species of Calycadenia, C. hooveri G.D. SIDA 21(1): 259 — 265. 2004 260 BRIT.ORG/SIDA 21(1) Carr. In addition, although dozens of reciprocal crosses of these populations with self-incompatible forms of Calycadenia truncata have been made, cypselae with embryos were generated only when the self-compatible form served as the female parent. Carr (1975) noted the same phenomenon in C. hooveri and suggested that it may be the result of unilateral interspecific incompatibility, as discussed in Lewis and Crowe (1958). The investigation of these unique plants led to re-evaluation of a previ- ously described taxon, Calycadenia truncata DC. subsp. microcephala H.M. Hall ex D.D. Keck. According to Keck (1946), “This subspecies is separated from Calycadenia truncata subsp. scabrella (Drew) Keck, to which it is most nearly related, by the reduced number of disk-florets (3 or 4 instead of the usual 8 to 15) and the smaller heads.” Additionally, he recognized the existence of intergra- dation between these subspecies. It appears from Keck’s description and dis- cussion that his small-headed taxon clearly includes the tiny-rayed self-com- patible taxon described here but also includes other small-headed C. truncata populations. Keck (1946) listed a number of specimens, and subsequently annotated numerous additional specimens, representing C. truncata subsp. microcephala, including locations from southern Trinity County to Lake County and in the Santa Lucia mountains of Monterey County. Some of these specimens are rep- resentative of a small-headed, self-compatible taxon but others represent popu- lations of self-incompatible C. truncata plants with somewhat larger heads and rays. Depauperate individuals are common in populations of annual plants, es- pecially in harsh years. Specimens of C. truncata prepared from such depau- perate plants, especially those poorly pressed or without flowering heads at an- thesis, could easily be confused with the new, small-headed, self-compatible taxon proposed herein. To better understand his concept of C. truncata subsp. microcephala, at- tempts were made to field-verify all collections referred to this taxon by Keck in his original publication (1946) and subsequent specimen annotations. This was difficult as most collections cited, including the type locality, have not been re-located. Additionally, the type locality cited (H.M. Hall 9602, Mill Creek Can- yon about 8 miles eastward from Ukiah, Mendocino Co., CA.) has no habitat that would support Calycadenia within ca. 3-4 miles. Hall may have acciden- tally written down the wrong mileage or possibly transposed a 3 with an 8. Regardless, there is very little likelihood that the type specimen was collected near the suggested location. There are sites 2-4 miles east of Ukiah with habi- tat that may have supported Calycadenia in the past but these sites are now largely occupied by agriculture. In any case, there appears to be no way to re- late the type specimen to an extant population in the field. After careful consideration, and for a variety of reasons, we believe the cir- cumscription of Keck’s C. truncata subsp. microcephala is not the same as the CARR AND CARR, OF CALYCADENIA FROM CALIFORNIA 261 new species proposed herein. The questionable nature of the type, the ambigu- ity in the description and cited specimens, and the likelihood that Keck had no knowledge of the derived breeding system of the new taxon described below, lead us toconclude that use of the epithet “microcephala’ for this new species is untenable. RESULTS AND DISCUSSION Calycadenia micrantha R.L. Carr & G.D. Carr, sp. nov. (Fig. 1). Type: U.S.A. CALI- FORNIA. TRINITY Co.: ca. 1 mi N of Mad River Rock on rd. to Mad River Rock from Low Gap, S of the Mad River Ranger Station, 40°23110'N, 123°29'01"W, 1340 m, 22 Aug 2003, R.L. Carr 3801 (HOLOTYPE: UC; ISOTYPES: OSC, US). Herba annua. Caule graciles l- Seuiramonun gl b | Folia £ | basalia 2-5 cm x 2-3 mm proxima aliter istali ae linearia. Bracteae pedunculares 2-4 mm hispidulae plus minusve pectinato-fimbriatae glande una oo capitata terminali. Ca- pitula L(-3) in quoque nodo; phyllariis 1-2, 4-5 mm; paleis 2-3, 5-6 mm hispidulis villosulis ad marginem superum interdum cum glande una parva capitata erminal flosculis radialibus 1-3 lamina 2-2.5(-3.5) mm sinu 0.5(-1) mm; flosculis disci 1-3; li | rugosis glabris epapposis; cypselis disci plerumque abortivis glabris epapposis. Annual herbs. Stems 1-5 dm, slender, generally less than 2(-3) mm diameter at the base, branches often many, generally beginning near mid-stem, arcuate to ascending; glabrous, often purplish, especially distally. Leaves in basal rosette 2-5 cm long, 2-3 mm wide, sessile by a widened base, becoming more remote and reduced distally, linear, hispidulous adaxially and along the margins, of- ten with longer, hispid hairs adaxially; leaves of the inflorescence 5-20 mm long, linear with a widened base, hispidulous, sometimes with a few awn-like bristles along the margin. Heads |(-3) per node, sessile or nearly so. Peduncular bracts 2-4 mm, these and associated reduced leaves terete to strongly flattened, glabrous to hispidulous, commonly also with 1-8 long, pectinate bristles on the margins; apex bearing | large tack-shaped gland. Phyllaries |-3(-6), 4-5mm, each partly enfolding a ray cypsela, the abaxial surface glabrous to more or less hispidulous, especially toward the tip, sometimes bearing a few scattered, stout bristles; distal margins with shaggy hairs; apex occasionally with a single small tack-shaped gland. Receptacle paleae 2-3; 4-6 mm, each associated with a disk cypsela, the abaxial surface glabrous to more or less hispidulous, especially to- ward the tip, sometimes bearing a few scattered, stout bristles, distal margins with shaggy hairs; apex occasionally with a single, small, tack-shaped gland. Ray florets 1-3(-6); fertile, corolla bright yellow, laminae 2-2.5(-3.5) mm long by 2-4 mm wide, 3(-4)-lobed, sinuses ca. 0.5 mm, the middle lobe(s) smallest, symmetric, oblong to narrowly triangular, the outer lobes asymmetric, basi- cally oblong to obovate but excursion of outer margin from midline greater than that of the inner margin, the tube 1-1.5 mm. Disk florets 1-3, 3-4 mm, yel- lowish. Ray cypselae ca. 3mm long, ca. 2mm wide, more or less triangular, rough- wrinkled, glabrous, pappi none. Disk cypselae ca. 3 mm long, mostly abortive, 262 BRIT.ORG/SIDA 21(1 Fic. 1. Cal 4 : : n Habit. B. Capitulum, lateral view. C. Capitulum from above. D. Peduncular bract (ap- pressed to capitulum). E. Ray cypselae. F, Peduncular bract tip with tack-shaped gland. Photos of Holotype, R.L. Carr 3801 (UC). terete when developed and tapered toward the base, smooth to very slightly ridged, glabrous, pappi none. Self-compatible. 2n = 14 (Carr 1977). Flowering Jun-)Jul-fall. Paratyres: U.S.A. CALIFORNIA. Colusa Co.: Wilson Camp SW of Stonyford, near the Colusa/Lake Co. line, ca. 3 air mi SE of Goat Mtn., 1400 m, 23 Sep 1994, R.L. Carr 2656 (UC). Lake Co.: 0.4 mi S of Elk aaa along Elk Mtn. Rd. 1140 m, 10 Aug 1974, G.D. Carr 771 (UC); S side of Elk Mtn. along Elk Mtn. Rd., 0.45 mi N of 14 mi marker, N of Upper Lake, 1150 m, 30 Sep 1986, R.L. Carr 2258 (UC); 0.7 mi NE of ae Rock Rd. on Bartlett Springs Rd., 2.1 mi NE ok L — iew Campground, 6.8 mi from with Hwy. 20, 1150 m, 23 Sep 1994, R.L. Carr 2658 (UC); S facing slope on Elk Mtn., . Mtn. Rd. to Upper Lake at jet. with forest rd. (17N28), ca. mile post 16.6 on rd., 1100 m, 23 Sep 1994, R.L. Carr 2660 (UC); S facing slope of Elk Mtn., 3.3 mi N of the entrance to Middle Creek C me on rd. from Upper Lake to Elk Mtn., ca. 1100 m, 23 Sep 1994, R.L. Carr 2661 (UC); E of Elk Mtn. Rd. on Deer Valley Campground Rd. (ONOL), 1.3 mi W of Deer Valley Campground, 1005 m, 18 Jul 1997, R.L. ae 3353 (UC); rd. to Bear Creek (LONOI) ca. 2.1 mi E of Deer Valley Campground, 0.8 mi W of Dry Oak Camp- ground, 1130 m, 18 Jul 1997, R.L. Carr 3354 (UC); ca. 200 yds SW of Little Pinnacle Rock Peak, on forest oS CARR AND CARR, A NEW SPECIES OF CALYCADENIA FROM CALIFORNIA 263 Fic. 2. Capitulae from greenhouse-grown plants. A. Calycadenia t ta, R.L. Carr 2224. B. Calycadenia micrantha, G.D Carr 771 rd. lONO1 ca. 1.9 mi W of its jet. with rd. to Bartlett Springs, 1350 m, 07 Oct 1997, R.L. Carr 3395 (UC): above Old Rd. to Witter Bie ca. 0.2 mi FE of Witter Springs site N of Lakeport, 39°11'29'N, 122°59'30"W, 500 m, 23 Aug 2003, R.L. Carr a 2(UC): along Hwy. 175, W of Lakeport, ca. 2.5 mi W of jet. with Hwy. 29, 38°59'43'N, 122°55) 48"W, 500 m, 23 Aug 2003, R.L. Carr 3803 (UC). Monterey Co.: jet. with =H Fort Hunter Liggett boundary, ridge crest near South Coast Ridge Rd., ca. 12.9-13 rd. mi S of Nacimiento-Fergusson Rd. and ca. 0.3 mi jet. with Burma Rd., 1000 m, 15 Jun 1998, E. Painter & E. Neese (s.n.) (SBBG); Fort Hunter Liggett (Training Area 17), near Burro Rd., ca. 2.0 km N of Three Peaks, ca. 3.25 km W of Burro Mtn., 700 m, 15 Jun 1995, E. Neese & E. Painter ge (SBBG); Fort Hunter Liggett (Training Area 17), near Burro Rd., ca. 2.0 km N of Three Peaks 5 km W of Burro ., 700 m, E. Painter, E. Neese & A. Hazebrook HL3005 (SBBG). Trinity Co.: sere of the Mad ee ca. 0.2 mi E of end of county rd. across Mad River, 40°09'48'N, 123°13'21"W, 899 m, 21 Aug 2003, R.L. Carr 3800 (UC) pa Distribution and ecology.—Dry, open, rocky ridges, hillsides and talus; 500-1,500 m; Colusa, Lake, Trinity and Monterey counties, California. This species grows only in areas of low plant density, in or closely associated with exposed rocky areas or areas of packed mineral materials. For this reason and because survi- vorship is low, populations are generally small with few individuals. The spe- cies is surviving in an extremely limited and very fragile habitat. Etymology.—Name Gr, micr, small; anth, a flower. Referring to the reduced ray flowers compared to most other species of Calycadenia. Compared to some other taxa of Calycadenia, C. micrantha is remarkably uniform morphologically. The variation that is seen between populations is 264 BRIT.ORG/SIDA 21(1) Tase 1. Comparison of selected features of Calycadenia micrantha and C. truncata. Character Calycadenia micrantha Calycadenia truncata Stem height 1-5 dm 2-12 dm Leaf length 2-5cm 2-10cm Peduncular bract length 2-4mm 1-12 mm Phyllary length 4-5mm 5-10 mm Ray floret number 1-3(-6) 3-6 Ray corolla lamina length 2-2.5(-3.5) mm (4-)5-12 mm Disk floret number - 3-25 Disk floret length -4mm 4-6 mm Breeding system Self-compatible Self-incompatible about that seen within populations. The overall size, branching and coloration of the plants, the overall size and position of the heads, the number, size, and position of the peduncular bracts, involucral bracts, ray ligules, ray achenes, receptacular bracts, disk flowers, and disk cypselae is very constant. The major variation seen is in the vestiture of the basal/proximal cauline leaves and that of the leaves associated with heads and the peduncular, involucral, and receptacular bracts. The basal and proximal cauline leaves are nearly always hispidulous but the presence of longer, stiff, bristly hairs ranges from sparse to rather dense. The leaves and bracts of the inflorescence are nearly always more or less hispidulous but the presence of pectinate hairs and other bristles varies considerably, as does the presence of hairs on or near the tips of the involucral and receptacular bracts. Additionally, the presence of the smaller tack-shaped gland on the tips of the involucral or receptacular bracts is variable, although uncommon overall. As discussed above, the populations treated here as Calycadenia micrantha have previously been considered conspecific with C. truncata. Salient features that help distinguish the two species as circumscribed here are presented in Table 1. Some of the most striking differences relate to the reduced capitulum associ- ated with the self-compatible breeding system found in C. micrantha (Fig. 2). In greenhouse cultivation of C. micrantha, we have noted that the middle lobe of ray flower laminae is commonly subdivided, yielding a 4-lobed lamina with two small symmetric central lobes and two larger, asymmetric outer lobes. Heads may contain a mixture of ray flowers with ‘normal’ and ‘aberrant lami- nae. It isnot known to what extent this phenomenon occurs in natural popula- tions. We have noted the same phenomenon at very low frequencies in other species or hybrids of Calycadenia in cultivation. CARR AND CARR, A NEW SPECIES OF CALYCADENIA FROM CALIFORNIA 265 ACKNOWLEDGMENTS We are grateful to Kenton L. Chambers, emeritus professor of botany, Oregon State University, for preparation of the Latin diagnosis. We also thank the cura- tors at UC and SBBG for making specimens available to the first author. We thank Bruce G. Baldwin (UC) and an anonymous reviewer for reviewing the manu- script. REFERENCES Carr, G.D. 1975. Calycadenia hooveri (Asteraceae), a new tarweed from California. Brittonia 27:136-141. Carr, G.D. 1977. A cytological conspectus of the genus Calycadenia (Asteraceae): an ex- ample of contrasting modes of evolution. Amer. J. Bot.64: 694-703. Lewis, D. and L.K. Crowe. 1958. Unilateral interspecific incompatibility in flowering plants. Heredity 12:233-256. Keck, D.D. 1946. A new subspecies of Calycadenia truncata from the Coast Ranges of Cali- fornia. Leafl.W. Bot. 4:259-260. 266 BRIT.ORG/SIDA 21(1) BOOK NOTICES Oxford University Press Trp J. Case, Martin L. Copy and Exrquirt Ezcurra. (eds.). 2002. A New Island Bio- geography of the Sea of Cortés. (ISBN 0-19-513346-3, hbk.). Oxford Uni- versity Press. (Orders: Oxford University Press, 2001 Evans Road, Cary, NC 27513, U.S.A. 800-451-7556, 919-677-1303 fax, www.oup.com). $95.00, 652 pp., figures, maps, graphs, tables, 6 !/2" x 91/2" Publisher Comments: “This updated and Casali A New Island Heed ly of the Sea of Cortés, first published nearly 20 years ago, int new and broader studies enco ssing more taxa and more complete island coverage. The paca synthesis provides a basis for furthe research and ex- porainen in upcoming years of the biologically fascinating Sea of Cortés region.” “The > new Biogeography includes a section on the conservation issues in the Sea of Cortés, past accomplishments, and conservation needs as yet outstanding,” DonaLb G. Crossy. 2004. The Poisoned Weed, Plants Toxic to Skin (ISBN 0-19-515548- , hbk.) Oxford University Press. (Orders: Oxford University Press, 2001 Evans Road, Cary, NC 27513, US.A.,800-451-7556, 919-677-1303 fax, www.oup.com). $59.95, 214 pp., figures, graphs, 24 color figures, 61/2" x 91/2". Publisher Comments: “Toxic plants affect the skin of almost everyone on earth. Many such “dermatotoxic” species were eae oe to the saan ane? ater generated centuries of medical research, but The Poisoned W oy nt I bout this topic. Cros covers a wide range of these plants.” Poison ivy and relatives (Anacardiaceae) are not the only culprits included in the book. JAMES F. HANCocK. 2004. Plant Evolution and the Origin of Crop Species. Second Edition. (ISBN 0-85199-685-X, hbk.) CABI Publishing, Wallingford, Oxon OX10 8DE, UK and 8756 Massachusetts Avenue, 7'* Floor, Cambridge, MA 02139, U.S.A. (Orders: www.cabi-publishing.org, Oxford University Press, 2001 Evans Road, Cary, NC 27513, U.S.A., 800-451-7556, 919-677-1303 fax, www.oup.com). $100.00, 313 pp., graphs, b/w photos, line drawings, 6 !/4" x 91/4" ublisher Comments: “This book i bably the only currer combine discussion of plant evolution with that of crop origins. ihe fist edition was published in 1992 by Prentice-Hall, and has now been fully revised to reflect recent advances. The key features include a description of the pro- cess of ev ee in native ane snauasee oe of plants, a review of when and where ae I f the subsequent development of crops over time. Ther is ae greater integration of the information on evolution and crop origins compared with the i edition. SIDA 21(1): 266. 2004 TWO NEW VARIETIES OF AGOSERIS (ASTERACEAE: LACTUCEAE) Gary |. Baird Department of Biology Brigham Young University-ldaho 525 South Center St. Rexbura, Idaho, U.S.A. 83460-1100, U.S.A. bairdg@byui.edu ABSTRACT — andiflora var. leptophylla occurs predominately west of the Cascade Mountains om EG diesen British Columbia to western Oregon and is morphologically and geographically dis- tinct from A. grandiflora var. grandiflora. ane two varieties are parapatric and intermediate forms are found where they occur together. A hylla var. quentinti is known from Sonoran Desert regions of Arizona and New Mesice a is sceuinalite ally isolated and morphologically distinct. RESUMEN A : If] ] } ayl] ‘ ] . ] 1,] wi = a | pee Agoserisgrandifl ir. leptophy ascadas lesde el de la Columbia Britanica | loested de Otepon yicsmonly Feice yigecereuceients ie) distinta de A. grandiflora var. grandiflora. Las dos intermedias alli donde conviven. Agoseris heterophylla var. quentinil es conueiea de las regiones del desierto de Sonora en Arizona y Nuevo México, esta aislada geograficamente y es diferente morfologicamente. INTRODUCTION Agoseris Raf. isa genus of perennial, lactucoid herbs that are found throughout western North America. One species is known from temperate regions of south- ern South America. Some Agoseris are considered taxonomically challenging. This is due, in part, to similar or overlapping morphologies between certain species and/or the formation of occasional hybrids between sympatric species. In addition, most members of the genus are very widespread and often contain local forms or regional phases that can appear quite distinct. Past attempts to formally recognize these forms or phases have largely failed as they either oc- cur as sporadic populations or they form broad clines, thus their separation becomes arbitrary. Despite these challenges, in a recent review of Agoseris (Baird 1996) two variants were discovered that were geographically and morphologi- cally distinct enough to merit nomenclatural recognition. aa prendre (Nutt.) Greene var. leptophylla G.1. Baird, var. nov. Type: U.S.A. HINGTON: PIERCE or LEWIS CO.: “dry oper n ground, upper valley of the Ye an River,” 19 Jul 1896, Allen 225 (HOLOTYPE: GH; ISOTYPES: CASI, DSI, K, NY/[2 sheets], UCI, WS). SIDA 21(1): 267-274. 2004 268 BRIT.ORG/SIDA 21(1) Varietas haec ab var. grandiflora differt capitulis minoribus et flosculis paucioribus, plerumque 40- 60+, phyllaciisiavelucriedpe cum maculis purpureis, foliis et lobis oe plerumaque ca. 2-4 mim latis, magis saepe in habitationibus humidis vel sylvaticis vel umbrios Perennial herbs, + acaulescent: leaves linear-filiform to —, oblanceolate, 10-25(- 36) cm x 1-4(-8) mm (excluding lobes), toothed (rarely) to laciniately lobed or pinnatifid, sub-glabrous to villous; lobes mostly in 3-5 opposite to sub- opposite pairs, linear to filiform, spreading to antrorse, often with a reduced secondary tooth or lobe on distal side of base of each primary lobe; heads borne singly, erect, scapiform; peduncles 15-40 cm tall at anthesis, 25-75(-96) cm tall at maturity, mostly 2-4 leaf lengths when mature, 3-4 mm in diameter, proxi- mally glabrate, distally villous to tomentose, non-glandular; involucres cam- panulate, 2-4 cm tall at maturity; phyllaries + ovate-attenuate, in 4-5 series, subequal at anthesis, unequal at maturity, usually entire, rarely dentate, herba- ceous, often purple spotted and/or with a rosy-purple medial stripe, adaxially sub-glabrous tot tose, abaxially glabrous or villous, margins ~ ciliate; hairs whitish-opaque or translucent, non-glandular; outel phyllat ies apically spread- ing to squarrose, not overtopping the inner series at anthesis; inner phyllaries erect, elongating, exceeding the outer at maturity; receptacles naked; florets +0- 60+; corollas equal to or just exceeding phyllaries at anthesis, yellow, often with an abaxial purplish stripe on the outermost; tubes 4-5 mm; ligules 3-5 = 1 mm: anthers ca. 1 mm; eypselae pale brown to whitish, 15-24 mm long, 10-ribbed, beaked, glabrous or slightly scabrous, homogenous or outermost slightly dif- ferent; bodies fusiform, 3-6 mm, abruptly contracted to their beaks: ribs ridged to sub-alate, straight, beaks 11-18 mm ong, filiform, mostly 3-4 lengths of cypsela bodies; pappi of capillary bristles in 2-3 series, 7-15 mm long, white; n =9(Tombet al. 1978, voucher: Chambers 2238 (OSC-143201), reported as Agoseris apargioides subsp. maritima). Common name.—Puget Sound agoseris. Agoseris grandiflora var. leptophylla occurs primarily west of the Cascade Mountains from Vancouver Island and southwestern British Columbia, south throughout the Puget Sound trough and Willamette Valley to southwestern Oregon and northwestern California (see Fig. 1). Italso occurs eastward through the Columbia River gorge and sporadically on the eastern slopes of the Cas- cade Mountains in Washington and Oregon. Some specimens from the moist, western slopes of the Rocky Mountains of British Columbia and northern Idaho (panhandle region) are assignable to var. leptophylla. Ecological notes on speci- men labels suggest that var. leptophylla occurs most commonly in lowland prai- ries or open forest habitats within the region outlined. As a whole, A. grandiflora manifests two or three geographic phases that grade together and cannot be satisfactorily or consistently separated. Con- versely, var. leptophylla represents what is arguably the most distinct phase o the species, with the specimens of var. leptophylla from the Puget Sound region BAIRD, TWO NEW VARIETIES OF AGOSERIS 269 Agoseris grandiflora @ var. grandiflora O var. leptophylla (Dashed lines indicate regions of sympatry where intermediate forms occur) 1 A . Jt aeth ig? fA . eT . NI tha * Tl + n Fic, re J J r primarily morphologically the most distinct from var. grandiflora. Within this region var. leptophylla appears to completely supplant var. grandiflora. Jones (1954) felt this was only the “expressions of different environmental regimens” and that the morphological overlap and geographical transition between this variant 270 BRIT.ORG/SIDA 21(1) and the remainder of the species great to justify recognition of intraspe- cific taxa. The separation and transition between the two varieties of A. grandi- flora, however, are no different than that found between other well-accepted varietal pairs within Agoseris (e.g., A. aurantiaca var. aurantiaca and var. purpurea, or A. glauca var. glauca and var. dasycephala). In the Columbia River gorge and southern Willamette Valley, extending into northern California, and in the panhandle region of northern Idaho, the two varieties do intermingle and intergrade such that the distinction between them falters and not all speci- mens are clearly assignable to one or the other variety. In general, var. leptophylla differs from var. grandiflora by its more diminutive size, slender, laciniately lobed leaves, slender peduncles, smaller heads with fewer florets, and outer phyl- laries subequal to (rather than longer than) inner phyllaries at anthesis. The following couplet will assist in separating the two varieties: 1. Leaves + entire, toothed, or pinnatifid, mostly 10-35 mm wide (excluding lobes), rarely less, lobes lanceolate to oblanceolate; Haare 3.0-5.5 cm tall at maturity, florets mostly 150-500-+, rarely fewer; outer p ries often with a purplish medial stripe, usually not spotted var. grandiflora . Leaves mostly laciniately pinnatifid, mostly 2-4 mm wide (excluding lobes), rarely wider, lobes filiform to narrowly lanceolate; involucres 2.0-4.0 cm tall at maturity, florets mostly 40-60, rarely more; outer phyllaries occasionally with a purplish medial Stripe but often purple spotted var. leptophylla — ne Specimens assigned here to var. leptophylla have often been identified as Agoseris laciniata (Nuttall) Greene or A. grandiflora var. laciniata (Nuttall) Jepson, names based on Stylopappus laciniatus Nuttall. In 1834-1835, Thomas Nuttall collected the types of four Agoseris taxa from the Willamette Valley of western Oregon, all of which he placed in Stylopappus. Three (Stylopappus elatus, S. laciniatus, and S. laciniatus var. longifolius) were collected at the mouth of the Willamette River and belong to the enigmatic Agoseris xelata (Nuttall) Greene Jones 1954; Baird 1996). The fourth (Stylopappus grandiflorus) was col- lected on the “ high plains” or “ hills” of the Willamette (the exact location is not known but possibly in present-day Marion or Polk Counties; see Ewan, 1971) and isthe type of Agoseris grandiflora; the specimen is teratological but is clearly assignable to A. grandiflora. The type description of S. laciniatus seems to de- scribe A. grandiflora var. leptophylla and applying the name “laciniatus’ to this variant of A. grandiflora seems an appropriate course of action. However, the lectotype (Nuttall’s original gathering at BM) of S. laciniatus appears to be part of the A. xelata hybrid complex and is therefore excluded from A. grandiflora Jones 1954; Baird 1996). The lectotype of S. laciniatus var. longifolius (also tera- tological) is more clearly of hybrid origin, with A. grandiflora var. leptophylla as one of the putative parents. Representative collections of Agoseris grandiflora var. leptophylla: CANADA. BRITISH COLUMBIA: Alberni-Clayquot R.D.: Alberni region, 26 Jun 1907, Rosendahl! 1969 (GH, MO, NY, UC). Capital R.D.: BAIRD, TWO NEW VARIETIES OF AGOSERIS 271 Near Victoria, 21 Apr 1885, Fletcher s.n. (GH, US); vicinity of Victoria, 31 May 1893, J. Macoun 573(GH, MO): Goldstream, Vancouver Island, 12 Jun 1939, Eastham s.n.(UBC); Maxwell Mt., Salt Spring Island, 7 Aug 1955, Ashlee s.n. (UBC). Central Kootenay R.D.: Longbeach, Nelson, 12 Jul 1937, Eastham s.n. (UBC); Gray Creek, Kootenay Lake, 12 Jul 1941, one n. — Cowichan Valley R.D.: Cowichan Lake, a ie Buckland 44 (UBC). Greater Vancouver R.D.: South Face of Black Mt., 15 Jun 1912, Davidson UBC). Nanaimo R.D.: Cameron Lake, 14 Jul 1917, fone s.n. (GH); First Lake, Nanaimo River ais, aes ees Dombois 61-3 (UBC); Parksville, — 1961, Taylor 3098 (UBC). U.S CALIFORNIA: Del Norte Co.: On road to — Basin, 1 mi. W of Doe Flat, 2 Aug 1955, Van Deventer 1003 JEPS). Humboldt Co.: Mackay Prairie, Tr it, 25Jul] ee Tracy 14234 (UBC); Sceaine at Nend of town, 18 Jul 1942, Trac 17275 (UBC) Guvely place at summit of ridge near “Clea 31 Jul 1950, Tracy 19233 (UC). Mendocino Co.: South Fork of Eel River, near the Mendocino as a 6 Jul 1918, Tracy 5074 (UC). Trinity Co.: Two mi W of Hayfork, 23 Jun 1943, Pitelka 256 (UC); Trinity Centre, 27 Jun 1982, Straley 2341 (UBC). IDAHO: Clearwater Co.: One mile S of Weipe, 26 Jun 1941, Davis 3592 (UC). Idaho Co.: Above Little Granite Creek, 13-30 Jun 1937, ais rd 265 (UC). Kootenai Co.: Coeur d’Alene, EF slope of Tubb’s Hill, 28 Jun 1913, Rust 316 (US). Shoshone Co.: Coeur d’Alene Mountains, between Old Mission and Wardner, 30 Jul 1895, Leiberg 1413 ae US); pen cae Station, on the road to Avery, + Aug 1941, Wilson 488 (GH, UC). OREGON: Benton Co.: Corvallis Jun 1916, Gilbert 26 (OSC). Clackamas Co.: Boring, 18 Aug 1918, Diehm and Gorman 1339 mee Curry Co.: Brookings, 11 Jul 1919, Peck 8784 (WILLU); aay above Agness, Rogue River, 8 Jul 1929, Henderson 11386 (UC). Douglas Co.: Roseburg, 21 Jun 1916, Peck se LLU); Umpqua eae Forest, Bear Creek Road No. 2735, 21 Sep 1975, ee n. (ORE). Hood River Co.: Bonneville, 6 1895, Canby s.n.(US); Hood River, 23 Jun 1911, Peck 2453 (WILLU). ne Co.: Ashland, 19 Jun 2 Peck 14999 (WILLU); 4 mi E of Central Point, 22 May 1948, Peck 24856 (WILLU). canines Co.: Bank of Suttle Lake, 19 Jul 1925, Peck 14426 (WILLU). Josephine Co.: Grants Pass, 28 Jul 1913, Peck ae (WILLU); Takilma, 24 Jun 1918, Peck 7953 (WILLU). Klamath Co.: Klamath Falls, 28 Mas Se Peck 2446 (WILLU). Lane Co.: Amazon Slough, W of Eugene, 31 May 1925, Constance s.n. (UC); Spencer's Butte, 1] Jul 1933, Brown 229 (ORE). Lincoln Co.: Yachats, bluff above the sea, 25 Aug 1921, Peck 10612 (WILLU). Linn Co.: Cascade Mtns., vicinity of Tombstone Pass, Iron Mtn., Cone Peak, Tombstone rairie, by Hwy. 20, 16 Aug 1983, Chambers and oe 78 (OSC). Multnomah Co.: Dry hills in Port- land, 25 Jun 1886, oe 584 (US); St. Johns, 28 Jul 1902, Sheldon 1102] (US). Tillamook Co.: Neahkahnie, 3 Jul 1924, Peck 13313 (WILLU). Wasco Co.: The Dalles, 7 Jun 1869, Kellogg and Harford 604 (US). WASHINGTON: Clallam Co.: Mt. Angleles, 21 Jul 1931, Howell 7429 (CAS). Clark Co.: East Mill Plain, 27 Jun 1925, English 452 (US). Chelan Co.: Nason Creek, 30 Jul 1893, Sand berg and Leiberg 612 (GH, UC, US). Grays Harbor Co.: near Montesano, 27 Jun 1898, Heller and Heller 3964 (MO, NY, PH, US). Island Co.: Whidby Island, Deception Pass Park, Goose Rock, 8 Jul 1937, Smith 2113 (DS). Jefferson Co.: Evergreen, 13 Jul 1902, Conrad 326 (PH, US). King Co.: Seattle, 19 Jun 1889, Smith s.n. (US). Klickitat Co.: Bingen Mountain, 16 Jul 1907, Suksdorf 6007 (GH). Mason Co.: Olympic National Park, Lincoln Ranger Station, road shoulder, 10 Jul 1941, Rogers 860 (UC). Pierce Co.: Tacoma, edge of forest prairies, Boe 1908, Flett 3390 (UC). San Juan Co.: Friday Harbor, 25 Jun-1 Aug 1917, Zeller and Zeller 857 (GH, U 1g Agoseris spel deta (Nutt.) Greene var. quentinii G.I. Baird, var. nov. Type: U.S.A. ARIZONA: Pima Co. “On gravelly slopes in scrub oak - mesquite eee alti- ae 4000 ft., Sawmill Canyon, near upper well, Santa Rita Mountains,” 26 Mar 1945, Gould and Haskell 3045 (HOLOTYPE: LL; IsoTYPES: ARIZ!, CASI, Ds GH], NY}, UC!) A aa A ] 1 <" : ee | - } Sonn | Varietas haec aliis differt forma valde acauli slabris sed adaxialiter pubescentibus, lebis nORUnOeDS iS obtusatis raro foliis dentatis vel integris, scapis ongioribus sed foliis scapum Speen bus. acicuas costatis vel porcatis, sine variatione vanietatutn ceterarum. 272 BRIT.ORG/SIDA 21(1) Annual (winter annual?) herbs, acaulescent: leaves oblanceolate to spathulate, 2-12 cm x 3-9(-12) mm, spreading to prostrate, adaxially pubescent, abaxially glabrous, mostly lobed, rarely toothed; lobes in 2-3 pairs, rounded to blunt, lack- ing secondary denticulations; heads borne singly, erect, + sessile to scapiform:; peduncles to 26 mm tall at maturity, mostly less than 0.5 leaf lengths at anthe- sis (rarely longer), 0.5- 3 leaf lengths at maturity, villous to tomentose, proxi- mally + glabrate, distally tomentose; involueres campanulate to hemispheric, |-2cm tall at maturity, sometimes proximally pubescent, hairs yellowish-trans- lucent, glandular, phyllaries lanceolate, in 2-3 series, subequal at anthesis, un- equal at maturity, entire, herbaceous, often with a purplish medial stripe, adaxially lanate, hairs whitish-opaque, abaxially + glandular-villous, hairs purple-septate, translucent (intermixed with whitish-opaque hairs), margin- ally + ciliate to lanate; outer phyllaries erect to squarrose, not elongating at maturity; inner phyllaries erect, + elongating at maturity; receptacles naked; corollas + equal to phyllaries at anthesis, yellow, outermost often with an abaxial purplish stripe; tubes 2-3 mm; ligules 2-3 x 0.8-1.5 mm; anthers 1 mm or less; cypselae pale brown to whitish, 9-10 mm, ca. 10-ribbed, beaked, + glabrous, + homogenous; bodies fusiform, 3-4 mm, gradually tapering to abruptly narrow- ing to their beaks; ribs ridged, straight, not diminishing proximally; beaks 5.0- 6.5mm long, 1.5-2.5 lengths of cypsela bodies; pappi of capillary bristles in 2-3 series, 4-9 mm, whitish. Common name.—Arizona agoseris. Agoseris heterophylla var. quentinii is known from Arizona and New Mexico (see Fig. 2). It most commonly occurs in desert grasslands, scrublands, and open woodlands between 1200 and 2000 m. It is found on various moun- tain ranges from the vicinity of the San Francisco Peaks southeast ward to south- eastern Arizona and southwestern New Mexico. It has not yet been reported from Mexico, although it has been found not far from the border in the Baboquivari and Huachuca mountains of Arizona and the Peloncillo Moun- tains of New Mexico. The principle features that distinguish var. quentinii from the other two varieties of A. heterophylla are its strongly acaulescent form, leaves adaxially pubescent and abaxially glabrous, peduncles apically tomentose and typically much shorter than the leaves at anthesis, and homogenous cypselae that lack the morphological variation and heterogeneity typical of this species. The three varieties of A. heterophylla accepted here may be separated using the following key: 1. Corolla ligules 2-4 mm long, = equaling phyllaries; anthers less than 1.5 mm lon leaves entire, toothed, or lobed, the lobes mostly 2-3 paired; found in California and/or elsewhere. 2. Peduncles mostly 1.5-4.5 leaf lengths at anthesis, proximally gla! glabrate, ihe day f TdUIOUs OF |: distally glabrous or + pubescent; leaves glabrous or uniformly pubescent; wide- BAIRD, TWO NEW VARIETIES OF AGOSERIS e Avar. Seis spread in western North America but not known from Arizona or New Mexico var. heterophylla 2. Peduncles 0-1 leaf length at anthesis, proximally glabrous or glabrate, distally tomentose; leaves adaxially oo abaxially glabrous; desert regions of southern Arizona and New Mex var. quentinii Corolla ligules 10-15 mm long, ace exceeding phyllaries; anthers 2-4 mm long; leaves toothed to lobed, the lobes mostly 3-5 paired; coast ranges and foothills of central California var. cryptopleura — 274 BRIT.ORG/SIDA 21(1) Agoseris heterophylla var. quentinii is named in honor of Quentin Jones, Ph.D., who monographed Agoseris for his doctoral thesis jones 1954). His work was instrumental in stabilizing and establishing much of the current nomencla- ture within the genus. He was the first to recognize that specimens of A. heterophylla from Arizona and New Mexico are distinct from the remainder of the species. His manuscript name was never published. Observed collections of seer EEECOR AYE a var. feel U.S.A. ARIZONA: Cochise Co.: Three mi N of Mescal, 17 Mar 1945, Pultz and Philli { ins, Bass Canyon on Muleshoe Ranch, ca. 25 mi NW of Willcox, T12S, R2IF, 14 May 1983, Daniel and Butterick 2863 (ASU, NY) Coconino edona, 25 May 1941, Stitt and McLellan s.n. (ASU). Gila Co.: Pinal Mountains, Six aes 2 May 1968, Pase 1797 (ASU); Pinal Mountains, Russell Gulch, 2.7 mi below intersec- tion of forest roads 55 and 55A, 24 Apr 1970, Keil, McLeod, Lamb, and Lehto 16792 (ASU). Navajo Co.: White Mountain Indian Reservation, on grassy flats around Kinishba Ruin, 31 Apr 1947, Lane 1946 (ASU). Pima Co.: Plains near Arivaca, 6 Apr 1884, Pringle s.n. (PH-2, NY-2); Santa Rita Mountains, Stone Cabin Canyon, 17 Apr 1903, Thornber 374 (ARIZ-2, NY); Santa Rita Mountains, Stone Cabin Canyon, 5 May 1905, Thornber s.n.(ARIZ-2); Santa Rita Mountains, “ Rozemont” (Rosemont?), 12 Apr 1907, Thornber s.n. (ARIZ); Santa Catalina Mountains, 16.1 mi S of Oracle on road to Mt. Lemmon and 0.4 mi E on ranch road, 27 Apr 1973, Lehto, Hansel, and Pinkava 10848 (ASU). Santa Cruz Co.: Santa Rita Mountains, McCleary’s, base of Old Baldy, Apr 1901, Griffiths 2671 (NY). Yavapai Co.: 1.6 mi Nol Skull Valley, 8 May 1967, Keil, Pinkava, and Lehto 8147 (ASU); Weaver Mountains, Arrastre Creek, 18 May 1980, Butterwick and Hillyard 6439 (ARIZ, ASU); Finch Wash, E of Skull Valley, near National Forest boundary, 2 Jun 1980, Butterwick and ne 6697 (ASU); Woodchute Wilderness Area, NW base a Woodchute Mountain, ca. 9 W of os 7 meas 1992, Baker as (ASU). NEW MEXICO: Catron Co.: Base of Mogollon Mountains, $ h trail ca. 6 mi SE of Glenwood, 4 mi ae Hwy. 180, 21 May 1983, Soreng and Ward 2130C (NMC). Grant Co.: Mangas Spring, 27 May 1941, without collector (NMC). Hidalgo Co.: Peloncillo Mountains, Coronado National Forest, about one mile NW of Pendleton Ranch House along Cloverdale Creek, T33S, R21W, $5, 20 Apr 1986 Worthington 14026. 5(NMC, NY). Location Uncertain: Mexican Boundary Survey (without location, date, or collector; NY); Griffiths 2671 (without location or date; ARIZ); Arizona, “ Toros Canyon,” 28 Mar 1927, Peebles, Harrison, and Kearney 3769 (ARIZ q =) ACKNOWLEDGMENTS [thank the curators and staff members at ARIZ, ASU, CAS, DS, GH, JEPS, K, LL, MO, NMC, NY, ORE, OSC, UBC, UC, US, WILLU, WS, and WTU for their assis- tance in locating information and for the loan of specimens. REFERENCES Baird, Gl. 1996. The systematics of Agoseris (Asteraceae: Lactuceae). Ph.D. dissertation, University of Texas, Austin. EWAN, J. 1971.Classica Botanica Americana, Vol. 7: Thomas Nuttall, Genera of North Ameri- can Plants. New York. Jones, Q.1954.Monograph of Agoseris, tribe Cichorieae. Ph.D. dissertation, Harvard Univer- sity, Cambridge. Tome, A.S.,K.L.CHameers, D.W. KyHos, A.M. PoweLt, and PH. Raven. 1978.Chromosome numbers in the Compositae. XIV. Lactuceae. Amer. J. Bot.65: 717-721. LECTOTYPIFICATION OF FASSIPEORA APFINIS (PASSIFLORACEAE) AND DISCUSSION OF ITS GEOGRAPHIC RANGE WITHIN THE UNITED STATES Douglas H.Goldman tesa Epa . Harvard University | FCECOSUATICL oom myTivit ty A 7 Cambri dep M 02138, U.S.A. dgoldman@fas.harvard.edu ABSTRACT George Engelmann, in his des ion of Passiflora affinis, never designated a holotype for the spe- J re cies, nor has a lectotype been oe ated in a ocaall iiecrarure addressing Engelmann’s work or North American Passiflora. A lectoype is designated here and a discussion and clarification of the geographic range of this species in the United States is provide Key Worps: Passiflora affinis, lectotypification, geographic distribution, Ferdinand Lindheimer, George Engelmann, Charles Wright, Charles Parry, Benno Matthes RESUMEN | d ipcion de Passif ffinis lesi holotipo para la especie ni epee se ha designado un lectotipo en la litera subsieuients que trata el trabajo de Engel- mann o las Pass ifl ra lectotip 0, é ademas se hace una discusion y cl Gomene la distribucién geagrilied de la scpccie en los Estados Unidos. INTRODUCTION Passiflora affinis Engelm. isa warm-temperate and subtropical species of oak- juniper savannas and mesic woodlands from central Texas and northeastern Mexico (Schulz 1922; Killip 1938; Correll & Johnston 1970; Diggs et al. 1999). The type collection was made by Ferdinand Lindheimer in August to Septem- ber 1849 at Comanche Spring, Bexar County, Texas. The type series was distrib- uted in 1907 by the Missouri Botanical Garden asa part of a long-forgotten por- tion of the Flora Texana Exsiccata (Blankinship 1907), although it was originally described several decades earlier (Engelmann 1850a). The labels distributed with the type series in 1907 (Fig. 1) are potentially misleading, presenting the type locality as “Comanche Spring; New Braunfels, etc.,” Comanche Spring and New Braunfels occurring perhaps 30 miles apart. However, Engelmann (1850b), Blankinship (1907), Killip 1938) and Lindheimer correspondence with George Engelmann, found in the archives of the Missouri Botanical Garden, make clear that it was collected at Comanche Spring alone. Lindheimer spent the majority of the growing season of 1849 at that site, re- turning to New Braunfels only at the end of that year (Engelmann 1850b; SIDA 21(1): 275 — 285. 2004 276 BRIT.ORG/SIDA 21(1) LINDHEIMER; FLORA TEXANA. (SUPPLEMENTARY TO “FLORA TEXANA EXSICCATA.”) DISTRIBUTED BY THE MISSOURI! BOTANICAL GARDEN. PASSIFLORA AFFINIS, ENGELM, Type Collection. Collected by Lindheimer from 1849 to 1851. Aug. -Sept. aS 849. Comanche Spring; New Braunfels, etc. No. GL, Fic. 1. The collection label found on speci f the ty, ies of Passiflora affinis distributed in 1907 (see Blankinship 1907), this label from a duplicate at GH. Blankinship 1907). Comanche Spring was on the property of a friend of Lindheimer’s, Baron Ottfried Hans von (‘John O.”) Meusebach, an early settler and founder of several communities in central Texas (King 1967). The spring was probably very close to Meusebach’s house near the headwaters of Salado Creek, in what is now the Camp Bullis Military Reservation (Ivey 1979). Although Englemann (1850a) did clearly state the type locality in the protologue, he did not designate a holotype, and neither Blankinship (1907) nor Killip (1938) later designated a lectotype from the P affinis type series. Passiflora affinis Engelm., Bost. J. Nat. Hist. 6:233. 1850. (Fig. 2). Type: TEXAS: [BEXAR Co.:] Comanche Spring, Aug-Sep 1849, F Lindheimer 174 (LECTOTYPE, here selected: GHI: ISOLECTOTYPES: ARIZ!, BM!, BR!, BRIT-SMUI, Cl, CANI, G! (2), GHI, Kl, MEXU, MOL NY! Pl PH! TEX! UC), US! WI). Liana, often suckering from roots, stems terete, glabrous, minutely puberulent to scabrous when young; tendrils glabrous; stipules linear-setaceous, erect, gla- brous, 0.1-0.2 cm; petioles glabrous, glandless, 0.4-4.5 cm; leaves glabrous, en- tire, shallowly to deeply 3(-5)-lobed, broadly ovate in general outline, basally cordate, truncate, to cuneate, 1-8(-10) cm long by 1.5-10(-14) cm wide, lobes ovate to obovate, rounded to truncate, entire to apiculate, often basally nar- rowed, laminar nectaries round, often in two obscure lines, each running from base of leaf near petiole apex toward the larger sinuses; inflorescence of two or rarely one flower per node, flowers fragrant, erect, stipe + pedicel 1.0-3.5 cm, slightly elongating in fruit; floral bracts glabrous, 3, occasionally deciduous, green to purple, linear-subulate to rarely ovate, 0.1-0.3 cm; flowers 3-5 cm in diameter; sepals moderately reflexed, lanceolate to linear-lanceolate, rounded aes GOLDMAN, LECTOTYPIFICATION OF PASSIFLORA AFFINIS 277 Sayojeg jONUOD 10109 YYGON H, Flora Texana exsicata. - No. C200 hon 6 ee es po cote a> King a : eS hs ne thy ; : 2 2 bd Kart fe F, Langham pe las Tai - : . oo 4 , e Fic. 2. The lectotype of Passiflora affinis at GH. 278 BRIT.ORG/SIDA 21(1) to acute, pale green to white, 1.0-1.6 cm long, 0.2-0.4 cm wide; petals moder- ately reflexed, linear-lanceolate, subulate, acute, pale green to white, 0.6-1.3cm long, 0.1-0.2 cm wide; corona of 2 series of filaments, outer series linear-fili- form, sinuous, weakly reflexed or not, tapering distally and terminating in a clavate apex, basally purple, apically green, white in between, 0.9-1.8 cm long, inner series filiform, erect, clavate, pale purple with green basally, 0.15-0.3 cm long; floral tube shallowly convex to nearly flat, operculum erect, incurved, plicate, pale purple to white, 0.1-0.2 cm long, nectar ring absent, limen erect, incurved, white, 0.1 cm wide; androgynophore 0.7-1.0 cm long, terete, pale purple; stamen filaments green with purple, 0.4-0.6 cm long’ anthers green with purple margins, 0.25-0.4 cm long; ovary globose to subovoid, glabrous, styles purple, filiform, glabrous, 0.5-0.7 cm long, stigmas broadly expanded, round, 0.1-0.2 cm wide; berry, purple-black, ovoid, globose, to dorsiventrally com- pressed, 1.0-1.5 cm long, 1.0 cm wide; seeds dark brown to black, flattened, obo- void, acute, transversely sulcate, 0.25-0.35 cm long, 0.2-0.3cm wide, with white arillate swelling to one side; germination epigeal. It would be anticipated that a specimen chosen as the lectotype of an En- gelmann name at the rank of species or below would be at the Missouri Botani- cal Garden where many of his collections reside, St. Louis, Missouri, being Engelmann’s home from the early 1830s until his death in 1884 (White 1896; Yatskievych 1999). However, the protologue describes both flowering and fruit- ing material and none of the examined duplicates of this collection have flow- ers and fruit except for the specimen chosen as the lectotype at GH (Fig. 2). Most duplicates seen are sterile or have few fruit, the only other flowering speci- men being at BRITSMU. All of the duplicates distributed in 1907 have mass- produced labels numbered 817, whereas one of the two GH specimens, the one with flowers, has a much older, mostly hand-written label, numbered 174. Lindheimer had his own field numbers, whereas Englemann used separate numbers based on the distribution order of the Flora Texana Exsiccata follow- ing the Bentham and Hooker sequence (Blankinship 1907). Blankinship does note (p. 170) that 174 is the Lindheimer number, which corresponds to the En- gelmann number 817, therefore these numbers refer to the same collection. The handwriting on the label of the now-lectotype is almost entirely Engelmann’s, identifying the plant as Passiflora triloba. Elsewhere on the label and in the lower right-hand corner of the specimen is written “affinis” in what is prob- ably Asa Gray’s handwriting. In addition, this is the only specimen of the type series that indicates habitat information as it is cited in the protologue, stating that the plant was growing in “shady places” and “climbing high over trees.” None of the series distributed in 1907 gives habitat details. The only detail in the label of the lectotype that differs from the protologue and the 1907 labels is the date, the older label stating “Sept 1849” whereas the protologue and newer labels state “Aug.-Sept. 1849” (Fig. 1). However, this does not make the older la- — os GOLDMAN, LECTOTYPIFICATION OF PASSIFLORA AFFINIS 279 bel incongruent with the protologue, and because of the other information on itand the presence of flowers on this specimen alone it is the best choice for the lectotype. GEOGRAPHIC DISTRIBUTION In Texas P affinis is typicallyfound growing over Cretaceous limestone or much less commonly over Precambrian igneous rock or late Tertiary sandstone and clay (Sellards et al. 1932; Spearing 1991). In northeastern Mexico (see specimens examined) it grows over Cretaceous limestone, shale and sandstone (Direccion General de Geografia del Territorio Nacional 198la, b; Padilla y Sanchez & Aceves-Quesada 1992). The currently known distribution of Passiflora affinis in Texas is shown in Fig. 3, based on herbarium and literature surveys, indicat- ing that it is nearly restricted to the southern Edwards Plateau and Lampasas Cut Plains (Diggs et al. 1999; Turner et al. 2003). A population was once collected in Fayette County, Texas, by Benno Matthes (see specimens examined), representing the eastern range-limit of the species in the United States and in an area with different geology than where this spe- cies occurs elsewhere in Texas. Matthes, a German settler and naturalist in Fayette County, lived for a short period in the town of Round Top, where he collected several plant specimens from late 1853 until mid-1854 (Geiser 1941). In this area he apparently collected P affinis (-Matthes 274), with specimens deposited at the Muséum National d’Histoire Naturelle, Paris and the Naturhistorisches Museum, Vienna (Killip 1938). This specimen still exists at E but it was not found at W ina recent search, although W does have a fragmen- tary Matthes specimen of P lutea labeled “bei No. 274,” suggesting that the two species were probably growing together. Matthes initially outlined his botani- cal discoveries in this area, although without mentioning Passiflora (Matthes 1855a). He later provided a more detailed field account (Matthes 1861) where he mentioned finding at the upper margin of a riparian forest “Passiflora triloba” with “Clematis coccinea” and Cornus florida. This may have been to the south- west of Round Top near Cummins (“Cummings”) Creek, along which he had been known to collect (Matthes 1855b, 1861). Although mesic forest edges are appropriate habitat for both P affinis and P lutea, such a habitat at the inner edge of the coastal plain may seem more appropriate for the latter species (e.g. Cooperrider 1995; Alford 2000) versus those of P. affinis on the Edwards Pla- teau and associated uplift. However, much of Fayette County, including the Round Top area, is underlain by sandstone and clay of the Catahoula and Oakville formations, both which can contain calcareous portions, particularly the latter (Sellards et al. 1932; Proctor et al. 1974). Furthermore, the springs in Fayette County are also known to be alkaline (Brune 1981). This suggests that much of Fayette County can provide habitats consistent with the primarily calciphilic preferences of P affinis. =x 280 BRIT.ORG/SIDA 21(1) + 35° }H~ 30° }— 25° 105° 100° 95° Fic. 3.G maries (Diggs et al. 1999; Turner et al. 2003). £ Shia tS a IIn? tc rn J L Killip 1938) reported the presence of P.affinis in New Mexico, citing a Parry specimen at US from Dona Ana county, labeled “chiefly in the valley of the Rio Grande, below Donana.” This specimen was collected by the botanical survey party attached to the United States and Mexican Boundary Survey, consisting of John Bigelow, Charles Parry, Arthur Schott and Charles Wright (listed on the collection label as “CC. Parry, J.M. Bigelow, C. Wright, & A. Schott”; see speci- mens examined). However, the only P affinis collection cited in the botanical report of the Boundary Survey is by Bigelow in October (but with no year given) along the Devils River (Torrey 1859) in what is now Val Verde County, Texas, perhaps 350 miles from Dona Ana County, New Mexico. The Boundary Survey travelled within close proximity to the Rio Grande for several hundred miles south and west of Dona Ana County after the Treaty of Guadalupe Hidalgo GOLDMAN, LECTOTYPIFICATION OF PASSIFLORA AFFINIS 281 was signed with Mexico in 1848, following the end of the Mexican-American War (Emory 1857). Therefore it should not have been assumed that this speci- men was collected within the present boundaries of New Mexico. In contrast, Killip (1938) did not similarly interpret the vague labels of P affinis specimens collected by Charles Wright to suggest that this species oc- curred in New Mexico. Three collections could have been interpreted as such, one at GH (Wright 217) labeled “from Western Texas to El Paso, New Mexico, May to October 1849,” another at GH and US with the same information but numbered 218, and another with no number at NY and PH labeled “N. Mexico, 1851’ (see specimens examined). It is surprising that they were not, with El Paso attributed to New Mexico in two of these three collections and “N. Mexico” also potentially interpreted in the same way, or as “northern Mexico.” Wright’s number 217 is actually a distribution number given by Asa Gray for his Plantae Wrightianae part 111852), whereas Wright’s field number corre- sponding to 217 was 404, collected on the 9° of June, 1849. His travel route (Geiser 1935) and his field notes, summarized in Shaw (1987) and available in the Gray Herbarium archives at Harvard University, indicate that 217/404 was collected “on the banks of the Leona” close to present-day town of Uvalde, Texas. Wright 218 was the distribution number for Wright's field number 734, col- lected on the 25" of July 1849, among the “hills of the Devil’s River” in present- day Val Verde County, Texas, although Gray (1852) consistently referred to the Devils River as the “San Pedro River”, following the geographic names used by French (1850; Shaw 1987). Wright's “N. Mexico, 1851” may be the same as that referred to by Gray (1853), collected “on the Sabinal, western Texas, July,” in what is now Uvalde County. However, Gray (1853) gives no distribution number for this specimen. In Wright's 1851 field notes in the GH archives Passiflora is en- tered only twice, once having been found on Chicon Creek (in modern Zavala County, Texas) in May 1851 and another time found in the “mountains near Santa Cruz, climbing among rocks,” in Sonora, Mexico, July 1851, with no mention of the genus in his 1852 notes. The Chicon Creek Passiflora matches the entry in Gray (1853) for P tenuiloba and the Santa Cruz specimen matches the entry (and protologue) for P. inamoena (=P. bryonioides Kunth). In July 1851 Wright was hundreds of miles away from the Sabinal River, in the vicinity of El Paso and southwestern New Mexico (Shaw 1987). It is possible that he collected this P.affinis specimen in this area, but without entering it into his field book, al- though that seems unlikely because he was a fastidious note-taker during his explorations of the border region. The only time he appeared to relax his note- taking was at the end of his travels in 1852, on his return to San Antonio, when on July 2" he records his final collection in what is now western Crockett County, Texas (Shaw 1987), over 100 miles to the northwest of the Sabinal River in Uvalde County. It seems probable that Wright collected this specimen not in 1851 but in 1852, east of Crockett County on his return to San Antonio, which 282 BRIT.ORG/SIDA 21(1) would have taken him through present-day Uvalde County. Shaw (1987) men- tions that these post-July 2, 1852 notes tend to be folded-up in the fragment folders of the specimens themselves. Such notes tend to be only with GH speci- mens because of Wright's professional association with Gray. No duplicate of this specimen exists at GH so such information is probably lost. Furthermore, Gray often cut up notes and correspondence to make into fragment folders (W. Kittredge, pers. comm.). Thus if these notes still exist at GH they may form a packet on one of the over five million specimens in the herbarium. These three Wright collections and the single Parry collection were ob- tained within what is currently west-central Texas, some of this area having been included in a southeastward extension of the Mexican province of New Mexico prior to the signing of the Treaty of Guadalupe Hidalgo (Commons 1990). Thereafter, all that is presently part of modern Texas was considered to be within the United States, Texas having claimed most of what is now modern New Mexico, northward through parts of Colorado and Kansas, into Wyoming, this land later purchased from Texas by the United States in the Compromise of 1850 (Stephens & Holmes 1988; Beck & Haase 1989). With these Parry and Wright collections made after Guadalupe Hidalgo in an area that has subse- quently been considered to be part of Texas only, any interpretations in her- barium labels or in the literature of these collections having come from New Mexico or northern Mexico are in error. Despite the potentially misleading labels on these specimens, P affinis has never been reported from modern New Mexico in any floras of that state, which is fortuitous because a similar situation with vague herbarium labels may have led to the potential misreporting of P tenuiloba for New Mexico (Martin & Hutchins 1981). Therefore, it can be assumed that the geographic range of P. affinis is restricted only to central Texas, with scattered populations in north- eastern Mexico. The potential confusion created by old, vague labels, can lead to floristic uncertainty and this alone should be an inspiration to all botanists to be extremely thorough in making their collection labels. ted examined: MEXICO. Nuevo Leon. Mcpo. Lampazas de Naranjo: Rancho Resendez, Lampazas, 22 Jun 1937, M.T. Edwards 298 (GH). Mcpo. Monterrey: Lan Sierra Meee ienté a 23 Jul 1933, CH. Mueller & M.T. Mueller 100 (GH). Mepo. Santiago: trail bety Las Ajuntas, 24 Aug 1939, C.H. Muller 2955 (GH). Tamaulipas. Mcpo. ealie Vic sof El ae Cerro Zamora, 22 Aug 1930, H.H. Bartlett 11094 (GH). Mepo. Victoria: |2mi W of Victoria, 28 Aug 1948, Kenoyer & Crum 3366 (GH). U.S.A. TEXAS: Bexar Co.: Comanche Spring, New Braunfels, etc., Aug-Sep, 1849, F Lindheimer 817 [=174] (ARIZ, BM, ms BRIT-SMU, C, CAN, G [2], GH [2], K, MO, NY, P. PH, TEX, UC, US, W); San Antonio, 17 Oct 1920, E.D. Schulz 401 (US). Edwards Co.: Devil’s Sink Hole, 29 Sep 1940, V.L. Cory 35652 (GH); 39mi S. of Sonora, 16 Aug 1941, VL. Cory 38039 (GH); Devil’s Sink Hole, LOmi E. of Rock Springs, 24 Jul 1946, D.S. Correll 13412 (BRIT, NY); lOmi SE of Carta Valley, 7 Aug 1905, D.S. Correll 31537 (GH). Fayette Co.: no locality, no date, B. Matthes 274 (P). Gillespie Co.: 3mi E. of farm road, |mi S. of county line off rt. 16, 20 Jul 1967, D.S. Correll & HB. Correll 34292 (GH). Hays Co.: San Marcos GOLDMAN, LECTOTYPIFICATION OF PASSIFLORA AFFINIS 283 and vicinity, Jul 1897, S.W. Stanfield s.n. (NY); lake bank, San Marcos, 25 Jul 1941, B.C. Tharp s.n. (BRIT, GH, NY, PH, US). Kerr Co.: Hunt, shores of Guadalupe River, 4 Aug 1969, P. Fryxell 1099 (NY). Kimble Co.: Telegraph, 8 Oct 1916, E.J. Palmer 10942 (US); along Llano River at Junction, 6 Sep 1965, D.S. Correll 31527 (GH). Kinney Co.: Ft. Clark, 10 May 1893, E.A. Mearns 1439 (US); 10 3/4mi W of Laguna, 24 Sep 1939, V.L. Cory 33461(GH). Llano Co.: Enchanted Mt., Sian 26 Jun 1932, C.C. Alberss.n. nee Enchanted Rock, 1 Aug 1938, B.C. Tharp s.n. 3 on Co.: 3/4mi SW of Ft. Terrett, 20 Aug 19 VL. Cory 24090 (GH). Travis Co.: above Zilke aie ustin, 10 Oct 1945, B.C. Tharp 45-49 (GH, os Hamilton Pool, 8 Jul 1966, [.R. Crutchfield a (NY); Austin, 27 Aug 1978, J.M. MacDougal 448 (US). Uvalde Co.: from Western Texas to El Paso, New Mexico, May-Oct 1849, C. Wright 217 (GH [2]); N. Mexico, 1851, C. Wright s.n. (NY, PH). Val Verde Co.: chiefly in the valley of the Rio Grande, below Donana, no date, C.C. Parry, J.M. Bigelow, C. Wright, & A. Schott s.n. (US); from Western Texas to El Paso, New Mexico, May-Oct 1849, C. Wright 218 (GH [2], US). Undetermined locality: Nueces, 1880s, V. Havard s.n. (US). CULTIVATED: Cornell University, originally from U. Texas, San Antonio, 17 Jun 2001, D. Goldman 1769 (BH); Cornell University, originally from U. Texas, San Antonio, 10 Aug 2001, D. Goldman 1819 (BH); Cornell University, originally from U. Texas, San Antonio, 9 Sep 2001, D. Gold- man 2126 (BH); Cornell University, originally from Edwards Co., TX, 9 Sep 2001, D. Goldman 2127 (BH); Cornell University, originally from U. Texas, San Antonio, 28 Sep 2001, D. Goldman 2160 (BH): Cornell University, originally from U. Texas, San Antonio, 2 Nov 2001, D. Goldman 2161 (BH); Cornell University, originally from U. Texas, San Antonio, 2 Nov 2001, D. Goldman 2162 (BH). — ACKNOWLEDGMENTS Thanks is given to Lisa DeCesare, Gretchen Wade and Judy Warnement for li- brary and archives assist ; Ken Cameron, Kanchi Gandhi, Walter Kittredge, Scott “M” LaGreca, Barney Lipscomb and Tanya Livshultz for valuable discus- sions or assistance; John MacDougal and Kristen Porter-Utley for reviewing this manuscript and other related discussions; the staff at ARIZ, BM, BR, BRIFSMU, C, CAN, G, GH, K, MO, NY, P, PH, TEX, UC and US for sending loans and/or making digital images of specimens available; Alain Changy (P), Jochen Heinrichs (GOET) and Bruno Wallnéfer (W) for assistance with finding Matthes specimens; Bob Dirig, Kevin Nixon and Sherry Vance (BH), and Richard Zander (BUF) for receiving loans on my behalf; Marshall Enquist and Larry Gilbert (TEX) for sending live material for cultivation; Carol Bader, Paul Cooper, Jerry David, Kim Goodwin, Andy Leed and Melissa Lucknow for the facilitation of space and cultivation assistance at Cornell University; Andrew Colligan for as- sistance with the archives of the Missouri Botanical Garden; Renate Hellmiss and Daniel Schott for translations of Matthes manuscripts, Dan Fulop for the Spanish translation of the abstract and Hobbes Goldman for general support throughout. REFERENCES Atrorb, M.H. 2000. The vascular flora of Amite County, Mississippi. Sida 19:645-699. Beck, W.A., and Y.D. Haase. 1989. Historical atlas of the American West. University of Okla- homa Press, Norman, Oklahoma. BLANKINSHIP, J.W. 1907. Plantae Lindheimerianae part Ill. 18th Ann. Rep. Missouri Bot. Gard. Pp. 123-223. 284 BRIT.ORG/SIDA 21(1) Brune, G. 1981. Springs of Texas. Vol. |. Branch-Smith, Ft. Worth, Texas. Commons, A. 1990. Divisiones territoriales, historia de las divisiones politico-adminstrativas. 1.5.2.(1524-1776), 11.5.3 (1776-1821) ,11.5.4 (1810-1990). In: Atlas Nacional de México, Vol |, Universidad Nacional Autonoma de México, D.F., Mexico. Cooperribder, 1.S. The Dicotyledonae of Ohio. Part 2,Linaceae through Campanulaceae. Ohio State University Press, Columbus. Correll, D.S., and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner, Texas. Dicas, G.M., B.L. Liescome, and RJ. O'KENNON. 1999. Shinner and Mahler's illustrated flora of north central Texas. Sida, Bot. Misc. 16. Botanical Research Institute of Texas, Fort Worth, Texas. DIRECCION GENERAL DE GEOGRAFIA DEL TERRITORIO NACIONAL. 1981a. Carta Geoldgica, 1:1,000,000. Monterrey. Estados Unidos Mexicanos, Secretaria de Programacién y Presupuesto. DIRECCION GENERAL DE GEOGRAFIA DEL TeRRITORIO Nacional. 1981b. Carta Geoldgica, 1:1,000,000. México. Estados Unidos Mexicanos, Secretaria de Programacion y Presupuesto. Emory, W.H. 1857. Report on the United States and Mexican Boundary Survey, Vol. 1.A.O.P. Nicholson, Washington, D.C ENGELMANN, G. 1850a. Passiflora affinis. P. 233. In A. Gray, ed., Plantae Lindheimeianae Part Il. Boston J. Nat. Hist. 6. ENGELMANN, G. 1850b."Subjoined preface.” Pp. 234-240. In A.Gray, ed. Plantae Lindheimeianae Part Il. Boston J. Nat. Hist. 6. FRENCH, 5.G.1850.A report in relation to the route over which the government train moved from San Antonio to El Paso del Norte. In: Reports of the Secretary of War: with recon- naissance of routes from San Antonio to El Paso. Senate, 31°‘ Congress, 1°' Session, Ex. Doc. No. 64. Union Office, Washington, D.C. Pp. 40-54. Geiser, S.W. 1935. Charles Wright's 1849 botanical collecting trip from San Antonio to El Paso; with type localities for new species. Field & Lab. 4:23-32 Geiser, S.W. 1941. Dr. Benno Matthes: an early Texas herpetologist. Field & Lab. 9:37-45. Gray, A. 1852. Plantae Wrightianae Texano-Neomexicanae Part |: An account of a collection of plants made by Charles Wright, A.M.,in an expedition from Texas to New Mexico in the summer and autumn of 1849, with critical notices and characters of other new or interesting plants from adjacent regions, etc. Smithsonian Institution, Washington, D.C Gray, A. 1853. Plantae Wrightianae Texano-Neomexicanae Part Il: An account of a collec- tion of plants made by Charles Wright, A.M.,in western Texas, New Mexico and Sonora in the years 1851 and 1852. Smithsonian Institution, Washington, D.C Ivey, J.E. 1979. The Comanche Spring site.La Tierra 6:16-21. Kiup, E.P. 1938.The American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19:1-613. Kina, LM. 1967. John O. Meusebach, German colonizer in Texas. University of Texas Press, Austin, Texas. Maartin, W.C.,and C.R. Hutcins. 1981.A flora of New Mexico. J.Cramer, Vaduz, Liechtenstein. Mattes, B.1855a. Excursion von New-Orleans nach dem Urwald am Rio Colorado in Texas. GOLDMAN, LECTOTYPIFICATION OF PASSIFLORA AFFINIS 285 Allgemeine deutsche N burg, Germany. Mattes, B.1855b.Die Hemibatrachier im ean und die Hem parecnish Yon Nord- America im Speciellen. Allgemeine deutsche he Zeitung, Band |, No. 7:249-280. R. Kuntze, Hamburg, Germany. Mattes, B. 1861. Reise-Bilder. Bilder aus Texas. HJ. Zeh, Dresden, Germany. PADILLA Y SANCHEZ, R.J., and J.F. Aceves-Quesaba. 1992. Geoldgica, IV.1.1.In Atlas Nacional de México, Vol Il, Universidad Nacional Autonéma de México, D.F., Mexico Proctor, C.V., T.E. Brown, J.H. McGowen, and N.B. Waecuter. 1974. Geological atlas of Texas. Austin sheet. Bureau of Economic Geology, University of Texas, Austin. ScHutz, E.D. 1922. 500 wild flowers of San Antonio and vicinity. Published by the author, San Antonio, Texas. Seccaros, E.H., W.S. Aokins, and F.B. Plummer. 1932. The geology of Texas. Volume 1, stratigra- phy. University of Texas Bull. 3232. Austin. SHaw, E.A.1987.Charles Wright on the boundary, 1849-1852, or Plantae Wrightianae revis- ited. Meckler Publishing Corp., Westport, Connecticut. SpeARING, D. 1991. Roadside geology of Texas. Mountain Press, Missoula, Montana STEPHENS, A.R., and W.M. Hoimes. 1988. Historical atlas of Texas. University of Oklahoma Press, Norman. Torrey, J. 1859. Botany of the Boundary. |n:W.H.Emory,ed. Report on the United States and Mexican Boundary Survey, Vol. 2, part 1.A.0.P. Nicholson, Washington, D.C. Pp. 27-270. Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Introduction, dicots. Sida, Bot. Misc. 24. Botanical Research Institute of Texas, Fort Worth, Texas. Waite, CA. 1896. Memoir of George Engelmann, M.D., 1809-1884. Extract from Biog.Mem. Nat. Acad. Sci. 4. Judd & Detweiler, Washington, D.C YATskievYCH, G.A. 1999, Steyermark’s flora of Missouri. Missouri Department of Conserva- tion, with Missouri Botanical Garden Press, Jefferson City. he Zeitung, Band I, No.4:152-162.R. Kuntze, Ham- 286 BRIT.ORG/SIDA 21(1) BOOK NOTICES Timber Press Mark EAarGeand Joun E. BraGcins. 2004. Tree Ferns. ISBN 0-88192-630-2, hbk.). Timber Press, Inc. 133 S.W. Second Ave, pu 450, Portland, OR 97204-3527, U.S.A.(Orders: www.tim berpress.com, imberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $39.95, a ne 131 color photos, 3 tables, 12 line drawings and 15 maps, 7 3/8" x 10 3 This book i is ake erns that look like trees. Tree ferns are true ferns and reproduce as other ferns by produc spores. Some of the species are ator displayed in the many color photographs oa in : book. The dust jacket says, “ volume is the source of information on the living tree ferns. It surveys all the families, genera, and Ba including those of are a for oe 1] = Ii as ltl home ge irden. It offers up-to y ve everything On tree ae use to conservation. = recognition of the horticultural Importance op tree ncluding } ferns pests.’ —Barney L cau Botanical Research Institute of Texas, 509 ean Street , Fort Worth, Texas 761202-4060, U.S.A. Davip E. ALLEN and GABRIELLE HATFIELD. 2004. Medicinal Plants in Folk Tradition, an Ethnobotany of Britain and Ireland. (ISBN 0-88192-638-8, hbk.). Tim- ber Press, Inc. 133S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 303-227-2878, l- 800-327-5680, 503-227-3070 fax). $29.95, 432 pp., 31 color photos, 57 b/w illustrations and | map, 6" x 9". anne Comments: This book provides the first comprehensive account of medicinal uses of wild plants by the country folk of Britain and Ireland. Two of Britain’s foremost ethnobotanical scholars ormation gathered by the Irish sree i 300 published and unpublished sources, including in Folklore Commission in more aa han 1000 manuscript volumes, to chronicle the fascinating uses of more 400 plant species ong the many lands of plants recorded here are a seaweed used in Scotland against ‘all ne — ee except the Black Death’, a mushroom stewed in milk in Norfolk to soothe cancer of t throat, a fem identified in Ireland as a ‘herb of the seven gifts’ for its reputed ability to cure diseases, and the remarkable range of benefits attributed to nettles. Such information, in addition to being interesting in and of itself, offers a starting point for ay eas and pharmacological investiga- tions of plants whose utility may ie n overlool An appendix of veterinary uses is provided. nd 31 colour photographs by Deni Bown.” Illustrations include 57 carefully SIDA 21(1): 286. 2004 THYMOPHYLLA SETIFOLIA VAR. GREGGII (COMPOSITAE) John L. Strother UGHUliT University 1001 Valley Life Sciences Bldg. University of California Berkeley, California 94720-2465, U.S.A. strother@uclink4.berkeley.edu — The types of the names Thymophylla greggii and T.greggii var. radiata have been treated as convarietal within Dyssodia setifolia (Lagasca) Robinson or Thymophylla setifolia Lagasca; (e.g., M.C. Johnston, Field & Lab. 24:67. 1956; J.L. Strother, Univ. Calif. Publ. Bot. 48:65-66. 1969; J.L. Strother, Sida 11:378. 1986). I have used the name Thymophylla setifolia Lagasca var. radiata (A. Gray) Strother. That name is illegitimate under Article 11.6 of the International Code of Botanical Nomenclature (W. Greuter et al., 2000. Regnum Veg. 138:i-xviii, 1- 474). For treatment of the variety in forthcoming Flora of North America north of Mexico, a new combination is required: Thymophylla setifolia Lagasca var. greggii (A. Gray) Strother, comb. nov. BASIONYM: Thymophylla greggii A. Gray var. greggii, established by Thymophylla greggii A. Gray Oo [oe} var. radiata A. Gray, Smithsonian Contr. Knowl. 3(5):119. 1852. SIDA 21(1): 287. 2004 288 BRIT.ORG/SIDA 21(1) BOOK NOTICES Timber Press JENNIFER TREHANE. 2004. Royal Horticultural Society Plant Collector Guide Blue- berries, Cranberries and Other Vacciniums. (ISBN 0-88192-615-9, hbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, l- 800-327-5680, 503-227-3070 fax). $29.95, 272 pp. 66 color photos, 7" x 9" Publisher Comments: “This book clarifies the Vaccinium story for any reader, whether an enthusias- tic gardener or someone who is simply curious about blueberries and cranberries and would like to le ab l inform and inspire those who have either never grown Vaccinium plants before or have put just a > eee widen the picture and maybe even learn a litt out some of the lesser known vacciniums. It wi tentative toe in the water and would now like to go ina little deeper. The book contains a wealll 1 of information to get a reader started on growing blueberries, cranberries and even lingonberri Lee Reicu (lustrations by Vicki Herzfeld Arlein). 2004. Uncommon Fruits for Every Garden. (ISBN 0-88192-602-7, hbk.). Timber Press, Inc. 133 S.W. Sec- ond Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.tim berpress.com, mail@timberpress.com, 503-227-2878, 1-800-327- 5680, 503-227-3070 fax). $24.95, 308 pp., 51 color photos, 29 line drawings, 1 map, 6" x 9" Publisher Comments: “Lee Reicl ides a valuable guide to fruits and berries that add an adventur- L ous flavor to any garden. as pence names like jujube, Eee ed pop, and shipova may seem ex- otic at first glance, these fruits offer delectable willit 1g to goo ] y slightly oll the beaten path at local nurseries. Reliable in the fat hest siden situations, cold- eee and pest- resistant, they are as enticing to the beginning as to the advanced gardener. This expanded sequel to the author's celebrated Uncommon Fruits Worthy of Attention offers new fruits, new varieties, and new photos and illustrations to entice the reader into an exciting world of garden pleasure.” SIDA 21(1): 288. 2004 TWO NEW COMBINATIONS IN THE GENUS PACKERA (ASTERACEAE) Debra K. Trock Herbarium, Department of Plant Biology Michigan State University East Lansing, Michigan 48824-1312, U.S.A. ABSTRACT vo new varietal combinations in Packera (Asteraceae: Senecioneae) are made based on work com- pleted for the Flora of North America north of Mexico project. RESUMEN Seh d binaci varietales en Packera (Asteraceae: Senecioneae) en base al trabajo mpletado para el proyecto Flora of North America north of Mexico. Packerda isa relatively recent North American segregate of the much larger cos- mopolitan genus Senecio. Askell and Doris Love first proposed the recognition of Packera as a distinct genus based on chromosome numbers and morpho- logical trends (Love & Léve 1975). They initially transferred eight species of arctic plants to this genus. Subsequently other authors have transferred most of the remaining taxa (Weber & Léve 1981; Jeffrey 1992; Trock & Barkley 1998; Bain 1999; Kartesz 1999). Preparation of a treatment of Packera for the forthcoming Flora of North America north of Mexico has brought to light the need for two additional no- menclatural combinations Packera pe ah (Greene) W.A. Weber & A.Léve var. borealis (Torr. & A. .K. Trock, comb. nov. BASIONYM: Senecio aureus L. var. borealis Torr. & A. Cray FI. N. Amer. 2:442. 1843. Senecio cymbalarioides Nutt. var. borealis (Torr. & A. Gray) Greenm., Ann. Missouri Bot. Gard. 3:117. 1916. Senecio streptanthifolius Greene var. borealis (Torr. & A. Gray) J.F Bain, Rhodora 90:293. 1988. Packera subnuda (DC.) Trock & T.M. Barkley var. moresbiensis (J.A. Calder & R.L. Taylor) D.K. Trock, comb. nov. BASIONYM: Senecio cymbalarioides Nutt. subsp. moresbiensis J.A. Calder & Taylor, Canad. J. Bot. 43:1399. 1965. Senecio moresbiensis J.A. Calder & R_L. Taylor) G.W. Douglas & G. Ruyle-Douglas, Canad. J. Bot. 56:1710. 1978. Packera moresbiensis J.A. Calder & Taylor) J.E Bain, Novon 9:457, 1999. ACKNOWLEDGMENTS lam grateful to Luc Brouillet and Guy Nesom for reviewing this manuscript, SIDA 21(1): 289 — 290. 2004 290 BRIT.ORG/SIDA 21(1) and to Alan Prather and Ted Barkley for providing helpful comments. | would also like to thank the curators of the following herbaria for their loan of speci- mens: ALA, CAN, COLO, DAO, MICH, MONT, MSC, NY, OSC, V, WTU. REFERENCES Bain, J.F. 1999. New combinations in Packera (Asteraceae) and lectotypification of Packera ovina. Novon 9:457-459. JerFrey,C. 1992. The tribe Senecioneae (Compositae) in the Mascarene Islands with an an- notated world check-list of the genera of the tribe. Notes on Compositae: VI. Kew Bull. 47:49-109. Kartesz,J.1.1999.A synonymized checklist and atlas with biological attributes for the vas- cular flora of the United States, Canada, and Greenland. First Edition. In: Kartesz, J.7.and CA. Meacham. Synthesis of the North American flora, Version 1.0, North Carolina Bo- tanical Garden, Chapel Hill, NC. Love ,A.and D. Love. 1975. Nomenclatural notes on arctic plants. Bot. Not. 128:497-523. Trock, D.K.and TM. Barkley. 1998. Seven new nomenclatural combinations and anew name in Packera (Asteraceae: Senecioneae). Sida 18:385~—387. Weser, W.A. and A. Love. 1981. New combinations in the genus Packera (Asteraceae). Phytologia 49:44-50. A NEW COMBINATION IN PERSICARIA (POLYGONACEAE) Craig C.Freeman i . er Herbarium, Division Oy PNUCUECAT AlEL CAG EU UT Uv ‘Y OOO rch Univers ity Oftonsa 2045 uncon ventc Lawrence, Kansas 66047-3729 cfree@ku.edu ABSTRACT The following new combination in | icari i iana (Cham. & Schltdl.) M Gomez var. beyrichiana (Cham. Schl ‘al Grae Ereenian RESUMEN | nueva en Persicaria: Persicaria meisneriana (Cham. & Schltdl.) M. omnes var. heynichiana (Cham. & Schltdl.) CC. Freeman. — Persicaria Mill. sect. Echinocaulon (Meisn.) H. Gross is represented by five spe- cies in North America north of Mexico. Park (1988) monographed the section as part of Polygonum L. Following evidence from morphological (Haraldson 1978; Ronse Decraene & Akeroyd 1988; Ronse Decraene et al. 2000) and mo- lecular studies (Lamb Frye & Kron 2003), sect. Echinocaulon will be included in Persicaria Mill. in the forthcoming treatment of Polygonaceae in the Flora of North America. Consequently, the following new combination is required. Persicaria meisneriana (Cham. & Schltdl.) M. Gomez var. beyrichiana (Cham. & Schltdl.) C.C. Freeman, comb. nov. BASIONYM: Polygonum beyrichianum Cham. & Schltdl. Linnaea 3:42. 1828. Polygonum meisnerianum Cham. & Schltdl. var. beyrichianum (Cham. & Schltdl.) Meisn., Fl. Bras. 5:19. 1855. Tracaulon beyrichianum (Cham. & Schltdl.) Small, FL. S.E. USS., 380, 1330. 1903. TYPE: n.v, see Park (1986) ACKNOWLEDGMENTS I thank Kanchi Gandhi for his assistance verifying the nomenclature. Jim Zarucchi, Caleb Morse, Guy Nesom, and Mihai Costea provided valuable com- ments. REFERENCES Haratoson, K. 1978.Anatomy and taxonomy in Polygonaceae subfam.Polygonoideae Meisn. emend. Jaretzky. Symb. Bot. Upsal. 22:1- Lame Frye,A.S.and K.A. Kron. 2003. rbcl phylogeny and character evolution in Polygonaceae. Syst. Bot. 28:326-332. SIDA 21(1): 291 — 292. 2004 292 BRIT.ORG/SIDA 21(1) Park, C.-W. 1986. Nomenclatural typifications in Polygonum section Echinocaulon (Polygonaceae). Brittonia 38:394-406. Park, C.-W. 1988. Taxonomy of Polygonum section Echinocaulon (Polygonaceae). Mem. New York Bot. Gard. 47:1-82. Ronse Decraene, L.P.and J.R. Akeroyb. 1988. Generic limits in Polygonum and related genera (Polygonaceae) on the basis of flora characters. Bot. J. Linn. Soc. 98:321-371. Ronse Decraene, L.P, S.-P. Hone, and E. Smets. 2000. Systematic significance of fruit morphol- ogy and anatomy in tribes Persicarieae and Polygoneae (Polygonaceae). Bot. J. Linn. Soc. 134:301-337. AN EVALUATION OF ANTHENANTIA (POACEAE) Robert Kral Botanical Research Institute of Texas and Vanderbilt University (VDB) 509 Pecan Stree Fort Worth, Texas, 76102-4060, U.S.A. ABSTRACT Anthenantia, a small Panicoid grass genus of the southeastern U.S.A. is evaluated gua) The two previously known taxa, A. rufa (Elliott) Schultes, and A. villosa (Mich get witha proposed new species, A. texanda, are described, compared and illustrated ber sa a ships are discussed. RESUMEN Se evaltia morfologicamente Anthenantia, un pequeno género de graminea panicoide del sureste de Estados Unidos. re dos taxa conocides septs A. rufa (Elliott) Schultes, y A. villosa (Michx.) Beauv, junto con | xana, se describen, comparan,e ilustran, y se discuten rE sus relaciones. Anthenantia Beauvois is treated in most recent floras (Small 1903 1933; Hitch- cock 1951; Radford et al. 1968; Correll & Johnston 1970; Gould 1975; Hatch et al. 2001; Barkworth et al. 2003), asa panicoid grass genus of two species, these con- fined to the Coastal Plain Physiographic Province and adjacent physiography within the southeastern United States from eastern North Carolina to Florida, westward (exclusive of the Mississippi Embayment) to Arkansas and eastern Texas. The genus was named by Palisot de Beauvois (Ess. Agrost. 48, 151, pl. 10, f.7, 1812) and based upon Phalaris villosa Michx. (FL Bor. Amer. 1:43. 1803). A second species, Aulaxanthus rufus Elliott (Bot. S.C. & Ga. 1:103. 1821), was in- corporated by Schultes (Mantiss. 2:258. 1824), thus becoming A. rufa (Elliott) Schultes and forming a bitypic genus. So far as southeastern U.S.A. canes are concerned, the only supplemental descriptive information appears to have from G.V. Nash, who noted some extra-typical A. rufa in eee of that species from South Carolina westward, these mentioned in J.K. Small’s Flora (1903, p. 79), as “A form, A. rufa scabra Nash.” Lately I have had some questions on the genus, the provocation being first from trying to fit the two previously known species into my “Guide to the Flora of Alabama and Middle Tennessee” (in prep.), second from trying to understand the patterns of morphological variation over the known range of the genus. Trips to and from Fort Worth, Texas across the Gulf South in recent years, to- gether with fieldwork over much of the southeastern U'S.A. over the past four decades and study of collections in the herbaria BRIT/SMU, DUKE, MISSA, NO, SIDA 21(1): 293 — 310. 2004 294 BRIT.ORG/SIDA 21(1) TAES, TENN, TEX/LL, and USF have allowed me to form concepts regarding these taxa. Asa result, a third taxon has been discovered. In this study the sequence will be L) detailed discussion of Anthenantia as a genus, presented in standard descriptive sequence; 2) a descriptive key, in- cluding the new species; 3) technical descriptions of the three, each followed by some discussion of habitat and distribution; 4) illustrations, the first three be- ing figures of the species, the fourth a plate with additional morphological de- tail. — Anthenantia Beauv. Ess. Agrost. 48, 151, pl. 10, f. 7. 1812. Type Species: A. villosa Michx.) Beauv. [= Phalaris villosa Michx.]. Habit loosely caespitose, slender perennials, perennating by shallow, concavel y- arching, scaly rhizomes spreading as axillary branches from older shoot bases. Roots fibrous, shallow; adventitious roots not observed. Culms wandlike, mostly 70-120 cm tall, terete, finely multicostate, smooth, leafiest at and toward base. Leaves lowermost transitional to rhizomal scales, almost entirely multicostate, firm, scale, just upculm transitional to short, then longer, bladed members, then to uppermost ones, most distant and with long, tubular-conduplicate sheaths and short, erect blades; lower (basal) sheaths often open, either angled- or rounded-concave; upper sheaths progressively more rounded-conduplicate, tubular: leaf blades linear or lance-linear to linear-spathulate or linear-gladi- ate, flat to strongly involute or flattened-conduplicate, the margins variously hirsute or pilose-ciliate, scabro-ciliate, scaberulous, or entire, the apices mostly asymmetrically broadly acute, scabro-serrulate, often with midrib exsert asa mucronula; surfaces strongly multicostate, abaxially smooth, adaxially with strongest costae, these and their intervals smooth to variously scaberulous or papillate, sometimes with rounded microhairs, and in one species with erect, pilose intercostal hairs to 2 mm. Ligule present as a narrow, transverse, pur- plish or brownish, often sinuous zone, this elevated as a low, erect to antrorse ridge or scale crested with minute to elongate papillae, ciliae, tubercles or scales, or erose. Note! Measures and character of ligules are taken from lower culm leaves; ligules of median and upper leaves can sometimes have longer hairs or scales than given here! Inflorescence paniculate, the upper scape a slender, na- ked peduncle above the short, erect blade of the elongate-sheathed upper leaf, mostly narrowly oblong to elliptic, lanceolate or ovate in outline, compact or loose, mostly 10-20 cm, the lowest nodes with primary branches whorled and usually longest, the internodes progressively closer, with progressively shorter primary branches, ultimate branching and rebranching sinuous, bending up- ward, terminating in cupuliform receptacles. Spikelets erect on cuplike recep- tacle, at maturity broadly obovoid or ellipsoid, slightly compressed dorsiven- trally, 3-4 mm long, exposed surfaces greenish and/or reddish, and at maturity KRAL, AN EVALUATION OF ANTHAENANTIA 295 densely cloaked with narrow, longitudinal bands of reddish, pinkish or pale, sharp, straight trichomes 1-2 mm long, lending a fuzzy look to the whole inflo- rescence; first glume lacking; second glume and lower lemma broadly obovate, obovate or elliptic, cupuliform, slightly longer than all other spikelet parts, abruptly acuminate, 5-nerved with broad, hyaline, entire borders, the oppos- ing lemma slightly narrower, less convex, both with strong bands of trichomes alternating with the median nerves, but the outermost (lateral) nerves narrow- est, surmounted by waxy-papillose zones with protruding trichomes, the sec- ond glume with its thin border lapping over the edges of the opposing lemma. Lower (first) lemma and its palea sterile or male, rarely with a bisexual floret, the palea hyaline, oblong- oblong-ovate or oblong, mostly apically two-toothed, shallowly two-keeled, two-nerved, or nerves lacking, entire ‘with broad invo- lute borders. Second lemma and its palea enclosing a perfect flower, cartilagi- nous, narrowly ovoid or lance-ovoid, plano-convex, at maturity brown to deep red-brown or appearing nearly black, with hyaline borders broadening toward scale apex, those of lemma finely ciliate, both with acuminate, incurved, crisped-bordered tips, at anthesis looking like a slightly opened bird’s beak, the lemma back strongly rounded proximally, the palea slightly rounded apically, medially and distally two-nerved, shallowly two-keeled, plane or slightly con- cave between the nerves. Florets those of the lowest (first) lemma either male or sterile; those of the upper (second) lemma bisexual. Perianth of two, asym- metrically flabellate, bilobed lodicules. Stamens 3, anthers oblong-linear to elliptic-linear, extrorse, 1.5-2.5 mm long, the very accrescent filaments attach- ing to anther adaxially at a very short connective, shoving the anthers out of the floret apex as they reach lengths of 4 mm or more. Ovary at anthesis with uneven dorsiventral symmetry, looking much like an ovoid-bodied, small- headed insect, the arched abaxial and convex side with two short, broad, lat- eral grooves, the more level, adaxial side with a broad, shallow, concave surface, the small “head” with two lateral, narrowly subulate-terete branches, these bent outward, proximally, then upward, narrowing distally, ca. 2.5 mm long, each abruptly thickening to become the axis of a dark, dense stigmatic “brush” ca. 2.5 mm, these shoved out laterally as the floret opens. Caryopses broadly and tumidly obovoid, 1-1.8 mm long, yellow-brown, the bulbous apex apiculate, the oblique base dorsiventrally narrowed and oblique, the hilum an oblong depres- sion on the palea side, the embryo under a large, rimmed disc on the lemma side. Distribution.—Three species in North America, Atlantic and Gulf Coastal Plain and contiguous physiography exclusive of the Mississippi Embayment, North Carolina south to peninsular Florida, west to southern Arkansas and east- ern Texas. Phenology.—Typically flowering from mid-July through October. BRIT.ORG/SIDA 21(1) KEY TO THE SPECIES . Adaxial surface of leaf blade with erect or variously directed, strumos Plioelty hairs arising from intercostal sulcae; principal leaves with blades shall auricled, slightly bent outward from sheath; longer primary panicle branches 1/3-1/2 as long as whole panicle, these usually naked-based and sae ascending (to 45° or slightly more ) giving panicle an ovate or broadly ellip gl and low (outer) lemma surfaces often with longitudinal aaa reddish bands Seen tri- chomes commonly reddish, or pale with red or pink tips; ligule base reddish, its edge ascending-ciliate, the pale hairs to 1(-1. . Adaxial surface of leaf blade esa eens el any sort ous 1 occasional pa- illae or microhairs) > 1.A.texana strongly auricled and bent outward from their sheathed: or eeeking auricles and erect to gradually bowed outward with no geniculation;longer primary branches of panicle either less than 1/3 panicle length or branching near base, seldom (save in a few extremes of no. 3) widely ascending, and generally with a narrower outline and a denser look; Sine ane ial ul with or lacking red pigm ted longitudinal bands; ligule bas ao te-tuberculate to nent ciliolate or ciliate or with a line of narrowly etanea 2. Principal (lower) leaves eed auriculate at blades of lower culm lea breaking’ a junction of blade and sheath, thus from sheaths at narrow to wide angles, these same eee at ee Soil wih spreading to ascending stru- mose-hirsute cilia; pigmentation of leaves, spikelets and their trichomes usually with little orany red; fertile lemma and palea ethOuls brown; anthers at anthesis brown, ligule rim of lower principal leaves ly erose or lacerate-ciliolate or with a line of irregular, flat-based cilia mostly under 1 mm 2. Principal (lower, bladed) leaves weakly, if at all, auriculate at junction of blade and sheath, thus be zone, if viewed from side, usually ee but a slight pro- jection and a few e hairs, sometimes ascending 2 2.A. villosa pilose-ciliate at blade base, or lacking hairs ane blades not perceptibly’ breaking away (geniculate) but flowing in ir sheaths. suberect erleney excurved; pigmentation of leaves, smieiee ind their trichomes variously reddish or purplish, thus the foliage, scapes aaa spikelets darker; fertile lemma and palea similarly ar. but a darker, red- brown to near black;anthers at anthesis dark brown; ligule rim of lower principal leaves nearly perpendicular, its edge commonly papillose- tubercular, minutely ciliolate, finely erose, or with a line of short narrowly-triangular-based pale cilia th 3.A.rufa 1. Anthenantia texana R. Kral, sp. nov. (Figs. 1, 4). U.S.A. TEXAS. HOUSTON Co: 2.5 mi W of Kennard city limit by TX Hwy 7, in Sam Houston National Forest; sandy clay loam of clearings in and edges of pine (Pinus taeda, P. echinata) and hard- je 30 Sep 2002, R. Kral 92270 (HOLOTYPE: VDB; ISOTYPES: AUA, BM, BAYLU, DOV, DUKE, FLAS, FSU, GH, ILLS, JSU, K, KANU, M, MICH, MO, MU, NCSC, ate me P TEX, U, VAM, UNA, USCH, US, VPI, VSC, WILLD. Planta perennis, gracilis, laxe caespitose, squami-rhizomatosa, (40-)50-100(-120) cm alta. Folia principalia vulgo suberecta vel leviter excurvata, 15-50 cm longa, leviter auriculata, leviter geniculata ad basim culmorum approximate, sursum remota; ligula albociliata, ciliis (0.5-)l(-1.5) mm longis; ] } Cals ] laminae loliorm ada ialiter strumos o-pilosae, pilis erectes vel ascenden tibus, ad 2 mm longis. Squa- mae exteriores spicularum alterne longitudine rubrae et virides. Lemma et palea secunda coriacea atrocastanea. Antherae maturae atrolerrugineae, ca. 2mm longae. KRAL, AN EVALUATION OF ANTHAENANTIA 297 Perennial (45-)60-100(-120 cm high, loosely caespitose, the shoot bases con- nected by short (-7 cm) concavely arching, shallow scaly rhizomes 2-4 mm thick. Culms slender, wand-like, leafiest at and toward base, the lowermost leaves mere ribbed yellowish scales 5-15 mm, soon grading to bladed members. Principal leaves (15-)20-40(-60) cm, ascending, the longest with blades sev- eral times longer than their open but somewhat “V”-shaped or rounded sheaths., grading to the uppermost, this with its erect blade much shorter than its con- volute-tubular slender sheath; sheath summit with two low but evident, usu- ally cartilaginous, pilose-edged auricles, the transverse narrow, usually purple, ligular scale projecting forward at an angle, its edge a band of pale ciliae (5-)(- 15 mm; blade lance-linear to linear, mostly 4-7(-10) mm wide, base thickened at rounded auricle area, there often ascending-pilose, here breaking from the sheath, distally narrowing, often flattened or “v”-troughed to plane or variably deeply concave or almost conduplicate, sree narrowed to a broadly and obliquely angled apiculate tip, abaxial surfaces of sheath pale to green tinged with red, those of blades mostly deep green; adaxial surface of blades pilose with pale, erect or ascending, pustular-based trichomes to 2 mm arising from deep intercostal sulcae. Panicle outline ovate to elliptic or broadly lanceolate, 7-15(-20) cm, the lowest group most distant, the primary branches ascending, sometimes to 45° or slightly more, the longest of a whorl also naked-based and mostly 1/3-1/2 or more the total length of the panicle, the whole with second- ary branching progressively more and shorter toward branch ends, giving the wide panicle base a much more open look. Spikelets turgidly obovoid or ellip- , 3-4 mm, at flowering time with longitudinal smooth zones of second en and outer palea reddish alternating with green, the alternating rows of pustular-based trichomes red to pale pink or purple. First palea hyaline, 2-keeled, bifid, hairless, slightly shorter than the opposing lemma, tristaminate or sta- mens lacking. Second lemma and palea about equaling subtending second glume, a deep, lustrous red-brown or near black, the hyaline margins gradually widening, ciliolate, toa crisped, ciliate apical border; flower usually perfect sim- ply pistillate. Caryopses 1.5-1.8 mm long. Distribution.—Sands, sandy clay loam, sandy peat or silts of pine flatwoods, pine-oak barrens, bog edges, ditchbanks, clearings, Gulf Coastal Plain and con- tiguous physiography west of the Mississippi delta, southern Arkansas, Louisi- ana and eastern Texas. Phenology.—Flowering mid-July through October. Paratypes. U.S.A. LOUISIANA: Beauregard Parish: by US 171, 2.8 mi N of Ragley & jet. US 190E (with A. villosa) 14 Oct 2001, R. Kral 91997 (TROY, VDB, VSC); ca. 17 mi W of DeRidder by US 190, 30 Sep 2002, R. Kral 93287 (BM, BAYLU, CONN, CTB, EKY, FSU, GH, JSU, KANU, KNK, MICH, MISSA, MO, MSC, eae NCSC, NY, mae aa RM, TENN, U, UNA, USCH,US, VDB, VPI, VSC, WAT, WILLI. ‘ited by county, collector and collector’s number): ARKANSAS: Bradley Co.: andele McIntyre 2788 (V DB). Calhoun Co.: Sundell, Amason & Etheridge 7876 (BRIT); Orzell 298 BRIT.ORG/SIDA 21(1) 2 ne 5 ali. pa f re iS SS KP y, | ty \ > Fic. 1. Anthenantia texana (from the type, Kral 92270). A. Habit sketch. B. Leaf apex. C. Leaf sheath/blade junction, adaxial side (left), sector of leaf blade, adaxial side (right). D. Leaf sheath/blade junction, oblique view. E. Idealized cross- section of involute leaf blade. F. Spikelet at anthesis. G. Abaxial side of glume. H. Abaxial side of first (lower) lemma (above); idealized cross section of same (below). J. Fruit, apy i ity, two abcissed sti brush (al ): abaxial side of fertile ( My yy | (t low). K. Side view of fruit. yi KRAL, AN EVALUATION OF ANTHAENANTIA 299 & Bridges 3058 (SMU). Ouachita Co.: Thomas & Doffitt 171881 & 669 (BRIT, NLU). LOUISIANA: Allen Parish: Shinners 22102 (SMU). Beauregard Parish: Shinners 21554 (NCSC, SMU). Grant Parish: Tho- mas, Barrett, Jones 1105 (VDB); Thomas et al. 12231, 12565 (VDB). Jefferson Davis Parish: Shinners 21469 (SMU). Natchitoches Parish: B.R. @ H.M. MacRoberts 1202, 1550, MacRoberts 1198 (VDB); Tho- mas et al. 171,947 (BRIT). Winn Parish: Shinners 21940 (SMU). TEXAS: Angelina Co.: 13 Oct 1979, Fritz sn. (SMU). Aransas Co.: Blakey 45140 (TEX). Austin Co.: 15 Oct 1939, Tharp s.n. (TEX). Calhoun Co.: | Dec 1928, Tharp, s.n. (TAES, TEX). Freestone Co.: Kral 154 (SMU). Hardin Co.: Gould 11030 (SMU, VDB); Parks and Cory 19901 (TAES). Harris Co.: Fisher 10178. Houston Co.: Kral 93270 (VDB-Ty pel). Jackson Co.: Silveus 371 (BRIT, TEX). Jasper Co.: Correll 38165 (SMU); Silveus 840 (TEX). Liberty Co.: Gould 5419 (SMU, TAES). Newton Co.: Mahler 5199 with Weaver (SMU). Nueces Co.: Tharp 7918 (TEX). Robertson Co.: Lonard 2460 et al. (SMU); 2 Oct 1948, Parks s.n. (TAES); Trew, Jr. 97 (TAES). Tyler Co.: Brown 3425 (TEX); Cory 49972 (SMU) This species has in the past been identified as A. villosa, particularly the broader and more distinctively auriculate-leaved, paler-haired examples (ie. Texas speci- mens, Calhoun Co., 1 Dec 1928, B.C. Tharp [TEX], Nueces Co., 25 Oct 1931, B.C. Tharp [TEX]). Many, because of their darker green or reddish pigmentation of foliage, darker spikelet pubescence, narrow, more erect leaves, have been identi- fied as A. rufa (a large majority of east Texas examples, all Arkansas examples, most examples from western Louisiana. The “villosa” morphs of the species from southeast Texas may suggest a genetic influence of that species, since A. villosa is the only other Texas native, it and A. texana often mingling where ecotones are obliterated by disturbance. But, in such instances, the two appear to retain most, if not all, distinguishing features. In Beauregard Parish, western Louisi- ana, both west and east of DeRidder, | observed many hundreds of the two in recently logged areas of pine savanna. Ineach site A. villosa (Kral 91997B, 93286) occupied sandier small rises but often would be within a few feet of clumps of A. texana Kral 91997A, 93287), which would be on slightly moister substrate. In those sites I had no trouble distinguishing the two, the former with broader leaf blades distinctly “breaking” away and with paler spikelets in narrower, denser inflorescences, the latter with narrower, geniculate, but less evidently so, and with darker spikelets in broader, more diffuse inflorescences. I plan fur- ther fieldwork in western Louisiana so as to see if there are examples there of mixed populations of A. texana and A. rufa and will prepare a report on that situation. Should there be such mixtures, from what information I have so far, all the A. rufa would be predicted to be forma scabra, while A. texana should show no scabrosity and a consistently geniculate leaf blade, together with a longer ligule. So, while there are distinct overlaps in regard to given characters for the three taxa, the pale, intercostal pilosity of strumose-based hairs is unique to A. texand. 2. Anthenantia villosa (Michx.) Beauv, Ess. Agrost. 48, 151, t. 10, £.7. 1812. ga 4). BASIONYM: Phalaris villosa Michx., Fl. Bor. Amer. 1:43. 1803. TYPE: U.S.A. “ sabulosis Carolinae,” Michx. s.n. (HOLOTYPE: Pl). BRIT.ORG/SIDA 21(1 ’ a Pi ah ey eee 4 of oe 2 dear la villosa (Mict ‘.) Beauv (from Kral 90571) as ‘ie eheth: . Leaf apex, side view. C. Sector of lower eaf blade, adaxial viewed. Adaxial f leaf/st view of spikelet. F. An anomalous “extra” floret fo anther ( right). G. Spikelet at anthesis. H. Thieey views of fertile ae a abaxial (palea side) view, stigma brushes protruding (above), abaxial view (below, left); side view (below, right). 1. Mature caryopsis (Hilum side). KRAL, AN EVALUATION OF ANTHAENANTIA 301 Aulaxanthus ciliatus Elliott, Sketch Bot. S. Carolina 1:102. 1816. Aulaxia ciliata (Elliott) Nutt., Gen. PL. L47. 1818. Type: “in dry pine barrens, flowers September - October,” (HOLOTYPE: CHAR “BY-3965"1). Elliott se Ste possibility a his A.ciliatus might | ing “Phalaris villosa? Mich. 1. p. 43.” To be sure of Elliott’s concepts, I visited the Charles- ton arene in December 2003 to examine the type material. Thanks to the consideration of Albert Sanders, curator of the Elliott Herbarium, | was qu oe enowa the Elliott types of inthus ciliatus and aie rufus, both mounted on a single sheet and in 2008 ce The right hand specimen is a small but ay es specimen (CHARL “BY- ius ciliatus El- | Phalaris villo us Mich 5 3065”) of Anthenantia villosa, marked t by Weatherby as the type of Aulaxant liott by Weatherby in 1941. To be absolutely sure of the identity of Michaux’s Phalaris vil At my request, Barney Lipscomb (BRIT) made photocopies of my plates of A. villosa and A. rufa and sent them unlabelled to Porter Lowry (P). The Michaux type clearly matched my losa. osd, unlabelled plate of Anthenantia vi Panicum erianthum Poir, Encycl. Sup. 4:284. 1816. Panicum hirticalycinum Bosc. ex Roemer & Schultes, Syst. Veg. 2:468. 1817. Perennial (50-)60-130 cm high, loosely caespitose, perennating by short, scaly rhizomes from older shoot bases, Culms slender, wand-like, leafiest at or to- ward base. Lowest leaves imbricate, yellowish, multicostate scales, grading to long-bladed principal leaves 15-40 cm, the sheaths rounded-conduplicate, open- ing distally, there with a narrow, wavy-transverse ligule, each edge terminating at a prominent thickened auricle., there with a tuft of pilose hairs, the ligule base ascending at a wide angle from the leaf surface, or even an erect, low wall of tissue, its edge variously short-ciliate (mostly not over 0.5 mm), or irregu- larly papillate-erose or with short, narrowly triangular squamellae; blades mostly lance-linear, lower ones distinctly breaking away from sheaths at vari- ous angles sometimes nearly 90°, 4-9(-15) mm wide, plane or inrolled, margins strumose-hirsute- ciliate, often also strongly papillose at least proximally, apex flat, triquetrous to rounded-conduplicate, or compressed-conduplicate, usually obliquely acute, scabro-ciliolate. Upper culm exserted 1-2 dm above the-erect, short-linear upper leaf blade as a slender peduncle terminating at a mostly narrowly elliptic to cylindric or lanceoloid panicle, this (5-)10-20(-25) cm, the whorls of primary branches ascending to erect, mostly closely rebranching to produce a generally dense, rarely interrupted inflorescence of yellowish to sil- very-green spikelets. Spikelets obovoid to ellipsoid, 3-4 mm long, the rows of pustular-based trichomes typically silvery or pale, the outer scales with mostly green or pale green surfaces, the hyaline first palea slightly shorter than its lemma, often with a line of ascending clear trichomes lateral to each keel, the floret either sterile or staminate: second lemma and palea coriaceous with dis- tal edges hyaline-ciliate, surface brown to dark brown, this floret typically per- fect. Anthers at maturity narrowly oblong-linear, ca. 2.5mm, yellowish-brown. Fruit ca. 2mm long, yellow-brown. Distribution.—Sands, sandy clay, sandy loams, moist to rather dry sites, mainly pinelands, particularly the longleaf pine-turkey oak system, oak-pine 302 BRIT.ORG/SIDA 21(1) barrens and flatwoods, upper edges of bogs, ditchbanks, sandy clearings, At- lantic and Gulf Coastal Plain and contiguous physiography from North Caro- lina south to South Florida and west, except for the Mississippi Embayment, into eastern Texas. Phenology.—Flowering mid-July through October (or November in south- ern range). — |: ALABAMA. Autauga Co.: Kral 33566 (V DB); Harper 4464 (UNA, VDB): McDaniel 7002 (MISSA. VDB). Baldwin Co.: Kral 29795 (V DB): Kral 29847 (SMU. V DB); Pennell 4551 (DUKE); Shinners 28901(SMU); Tracy 8025 (TAES). Barbour Co.: Kral 28004 (SMU, VDB). Bibb Co.: Kral 52265 & 69548, Choctaw Co.: Kral 67840. Conecuh Co.: Kral 40972 (BRIT, VDB). Covington Co.: Kral 33668, 38107, 36804, 33688, sda mes Crenshaw Co.: Kral 21993, 33722 (VDB). Escambia Co.: Kral 32477, 33873 (VDB). G aponals 12225 (VDB). Houston Co.: MacDonald 3030 (VDB). Mobile Co.: Kral 29701] cn 25 i. (SMU, VDB); Silveus 1021 (BRIT, TEX). Monroe Co.: Kral 69707, 85370 (VDB). Russell Co.: Kral 44210 (VDB). Washington Co.: Kral 37263 (VDB); McDaniel 9913 (VDB). FLORIDA. Calhoun Co.: Godfrey 55581 (FSU, NCU, VDB), Godfrey & Kral 54160 (DUKE). Duval Co.: Curtiss 6258 (NCU). Escambia Co.: Silveus 5622 (TEX). Franklin Co.: Godfrey (FSU, VDB). Gadsden Co.: Godfrey 53585 (DUKE, FSU). Hernando Co.: Ray 9484 (FSU, USE VDB). Hillsboro Co.: Lakela 23376 (USF), 25374 (SMU, USF, VDB). Jackson Go.: Godfrey 54264 (FSU), 54383 (DUKE, FSU, USF VDB); Tacy 3850 (TAES). Leon Co.: Cle well 793 (V DB): Godfrey 56111 FSU, USE, VDB): Kral 1789 (FSU, SMU, NCSG). Reese 640 (NCU). Levy Co.: Kral 15269 (V DB). Liberty Co.: Thorne & Davidson nee possible hybrid! (FSU). Madison Co.: ‘Kral 6178 (FSU, VDB). Okaloosa Co.: Godfrey 57669 (FSU), 27 Sep 1950, West s.n. Santa Rosa Co.: Ford 5375 (NCU), Godfrey & Houk 62551 (SMU, VDB). in Co.: Kurz 169 (FSU). Walton Co.: Gals 57630 (FSU); Tyson 358 (USF). Washington Co.: Godfrey 55238 (FSU, NCSC). GEORGIA r Co.: Thorne 6416 (Emor y U.). sa Co.: E vles 6814 (Emot y U,). Charlton Co.: Kral aa (V cm ae 7290 with Carter (VDB, VSC). Early Co.: Kral 90344 (VDB); ae ne 6628 (Emory U.). Emanuel Co.: Kral 85477 (VDB): Wilbur a (FSU, NCSC, SMU). Taylor . Correll 8401 (DUKE); Kral 85477 (VDB). Ware Co.: Silveus 5370A (TEX). LOUISIANA. oe Parish: Shinners 22218 (NCSC, SMU, TEX); Kral 91997B (V Be) Kral 92286 (VDB). Grant Parish: Thomas et al. 3062 (VDB), 12565 (BRIT). LaSalle pate : aird 1069 (BRIT). Natchitoches Par- ish: Kral 16207 (VDB). Rapides Parish: Duncan 56031 (SMU, TAES, TEX), and 56041 (SMU, TAES). St. many Parish: Bro. Arsene et al. 11251 (SMU). Vernon Parish: oe & DePoe 273 (VDB); Kral - 319 (VDB). Washington Parish: Allen 8684 (VDB). MISSISSIPPL. Forrest Co.: KE. & L. Rogers 42011 (SMU): Weddle s.n.9/10/40 (MISSA). Greene Co.: Rogers 1747-A (SMU. Harrison Co.: [racy 3848 (NCU, TAES). Jackson Co.: A.B. & A.C. Seymour 178 (DUKE, NCU); 3 Aug 1889, Earle s.n. (DUKE); Weaver & Rushing 0186 (V DB). Jones Co.: Morgan 1439(V DB). Lauderdale Co.: McDaniel & Clarke 14601(VDB). Pearl River Co.: Jones & Sargent 13694 (V DB), 8432 (SMU): Reed 53 (FSU). Stone Co.: Shinners 28826 (SMU). NORTH CAROLINA. Bladen Co.: Blomquist 13622 (DUKE). Brunswick Co.: Blomquist 439 (FSU, NCSC). Cumberland Co.: Carter 2974 (VDB); Ahles & Leisner 33466 (NCU). Columbus Co.: Blomquist 14785 (DUKE). Duplin Co.: Ahles 33183 with Leisner (NCU, VDB). Harnett Co.: God rey 5685 (DUKE) Radford 8758 (NCU). Hoke Co.: Kral et al. 82990 (VDB); Ahles 36383 (NCU). Johnston Co.: Radford 29292 (NCU, SMU). Moore Co.: 10 Apr 1931, Blomquist sn. (DUKE). Richmond Co.: Correll 7139 (DUKE); ae 19232 (NCU). Scotland Co.: Ahles with Leisner 32863 (NCU). SOUTH CAROLINA. Allendale .« Ahles with Bell 18423 (NCU). Bamberg Co.: Ahles 37657 (NCU). Barnwell Co.: Batson, s.n. 27 Oct 1953 (NCU, USCH) Calhoun Co.: Ahles 35363 (NCU, VDB). Chesterfield Co.: Godfrey 8086 (DUKE): Radford 18646 (NCU). Colleton Co.: Ahles with Bell 15431(DUKE, NCU). Darlington Co.: 22 Aug 1908, Coker s.n. (NCU). Edgefield Co.: Radford 30195 (NCU). Georgetown Co.: Godfrey 8109 (DUKE). Ker- shaw Co.: Radford 29984 (NCU). Lee Co.: Radford 29340 (NCU). Lexington Co.: Radford 29856 (NCU). Marion Co.: Bell 11012 (NCU). Orangeburg Co.: Ailes 35131(NCU). Richland Co.: Godfrey 50758 (DUKE, , KRAL, AN EVALUATION OF ANTHAENANTIA 303 NCSC). Sumter Co.: Freeman 57880 (NCU). TEXAS. Hardin Co.: Parks & Cory 11145, 19900, 19902, 19903 (TAES). Jasper Co.: Correll 38164 (SMU); 22 Aug 1941, Tharp s.n. (TAES). Newton Co.: 16 Sep 1947, Lay s.n. (TEX); Cory 49807 (SMU). I have not yet seen the three Anthenantia in one place, although this might be possible in western Louisiana or even eastern Texas. But were such to happen, A. villosa would be distinguishable at a glance. It is the most robust of the three, has the least rusty pigmentation in leaf and culm (save sometimes for pinkish internodes and spikelet hairs), has generally paler vesture, the plants therefore giving the overall effect of pale green. The lower culm leaves are more visibly auriculate and tend to spread more, and the blade margins are more coarsely and conspicuously strumose-hirsute-ciliate. Pigmentation of anthers and of the coriaceous lemma and palea is paler. The hyaline palea of the lower (first) flo- ret is unique, since it usually has rows of ascending, long, stiff, pale trichomes parallelling and external to each of the two keels (see fig. 4).On the other hand, in character of culm, in pigmentation, character and orientation of leaves and in character, shape, and indumentum of inflorescence, it is easy to see why the other two have traveled under the same epithet for such a long time. This will be discussed under A. rufa. Anthenantia villosa occupies the driest habitats of the three. It extends fur- ther up sides of sandhills, is in higher zones within ecotones between uplands and low, and is in the higher spots in flatwoods and savannas. It is the most frequent associate of Aristida stricta in Longleaf pine sandhills. Future studies probably will show that there is genetic exchange between itand A. rufaand A. texana, particularly where ecotones have been broadened through disturbance. 3. Anthenantia rufa (Elliott) Schultes, Mant. 2:258. 1824. (Figs. 3, 4). BASIONYM: inthus rufus Elliott, Sketch bot. S. Carolina 1:103. 1816. Aulaxia She ee Nutt., yo Pl. 1:47. 1818. Panicum rufum (Elliott) Kunth, Revis. Gramin. 1:35. Monachne rufa (Elliott) Bertolini, Mem. Reale Accad. Sci. Inst. Bologna > 506, t. 41, f. 1.1850. Type: [SOUTH CAROLINA] “in savannas, and damp soils in the pine bar- rens, midway between Saltcatcher bridge and Murphy’s on the Edisto,” Elliott 523 (number — = Muhlenberg) (HOLOTYPE: CHARL!). Leptoco lrumn i Mull. Berol., Bot. Zeitung (Berlin) 19:314. 1861. TyPE: U.S.A. LOUIsI- ANA: ‘1831 eo T. Drummond. Perennial (50-)60-125 cm high, loosely caespitose, perennating by short, scaly rhizomes from older shoot bases. Culms slender, wand-like, leafiest at or to- ward base. Lowest leaves mere scales, transitional to principal leaves 12-60 cm, with blades much longer than sheath; sheaths often red-brown or purple tinged, variously folded conduplicately, narrowing gradually to similarly folded blade, the connecting auricle minute or not evident save as a few pilose spreading hairs at ligule ends or even these lacking; ligule wavy-transverse, usually a narrow, erect ridge 0.2-0.4 mm high, typically purplish, its upper edge papillate, mi- nutely lacerate, rarely ciliolate with cilia at most 0.5 mm (uppermost leaves 304 BRIT.ORG/SIDA 21(1 SS leer eaeanttitiaans Ly me ae A ae = os Fic.3. Antt ia rufa (Ell.) Schult. (from Kra/ 90579). A. Habit sketch. B leaf apex side view (left), fblad hj f blade/ st | view, ligul D. Spikelet at anthesis. E. ‘Cleared’ f rtile lemma with y developing fruit. F. Ada Galvien of anther. G. Adaxial view of first Howe palea I is Ct low left); side view of upper's peune: lemma aang palea (above ant), H. Perfect floret, (left) 1.T views of ripe fruit, side view (left), bile side (middle), embryo side (right). KRAL, AN EVALUATION OF ANTHAENANTIA 305 often with distinctly longer cilia!); blades mostly erect to ascending-excurved, continuous with sheath apex, not at all geniculate, linear to linear-gladiate or linear- spathulate, (3-)4-8(-10) mm wide, flat to rounded-involute or variously folded- conduplicate, margins entire,- or sparsely pilose-ciliate proximally, or scabrid, the surfaces pale to deep green, or purple-tinged, smooth to papillate or scaberulous on the costae, apex mostly conduplicately sharply folded, or open at a wide angle, obliquely broadly acute, apiculate. Panicles ovoid to lanceoloid, ellipsoid, or cylindric, (8-)10-20(-25) cm, branching mostly strongly ascend- ing to nearly erect, the lowest whorl often with some primary branches 1/3 the total panicle length or more, but mostly floriferous to near base, thus the inflo- rescence mostly dense. Spikelets mostly broadly ellipsoid to obovoid, 3-4 mm, with exposed outer surfaces alternating with longitudinal bands of green and red, the bands of trichomes reddish to deep purple, rarely pinkish, very rarely pale. Lower floret usually staminate, often sterile, sometimes perfect; upper flo- ret with coriaceous lemma and palea deep red-brown or castaneous, appearing nearly black, the hyaline borders broadening distally, there ciliate, often crisped and finely ciliate apically, their flower usually perfect. Anthers at anthesis nar- rowly oblong, ca. 2-2.5 mm, deep purple-brown, appearing black. Ripe cary- opses 1.5-2 mm, yellowish-brown. Distribution.—Sands, sandy peats, silts and sandy clay of pine flatwoods and bogs, edges of bogs, acidic seeps and seep slopes, and pine savanna, Coastal Plain and contiguous geology, North Carolina south to peninsular Florida, west into Louisiana, possibly eastern Texas. Specimens examined.—Note! In Small’s Flora of the Southeastern United States 1903, p. 97) appears the citation., under Anthenantia rufa “A form, A. rufa scabra Nash, differing from the above in having the sheaths and blades scabrous, occurs in similar situations in South Carolina, but mainly from Ala- bama to Louisiana. Fall.” This information appears in some present-day refer- ence sources, but inconsistently, sometimes the taxon being given as a variety, in other instances as a “form” (as per Small). Since Nash and Small were col- leagues at the New York Botanical Garden, and since Small passed along the characteristics in 1903 for what he referred to as a form, it must be assumed that the name received no further published attention. | have been unable to find any actual formal presentation, and Small appears to have let the matter lapse as of 1903; certainly it did not carry forward to his 1933 “Manual.” What- ever the case, Nash should be credited for his observations. It is true that there are populations of scabrid A. rufa from the Carolinas west to Louisiana, and there are associated characters such as the presence (usually) of a tuft of pilose hairs at a small triangular projection where ligule meets margin, this often ac- companied by a short line of slender pilose cilia above and below along con- tiguous margin. However, these latter characters tend to vary independently, as does the degree of scabrosity of leaf blades and sheaths. My own conclusion is BRIT.ORG/SIDA 21(1) 306 KRAL, AN EVALUATION OF ANTHAENANTIA 307 to leave the situation as Small had it. “Scabra” morphs show a gradation west- ward, particularly in the increase of scabrosity and hairs at and around the ligular edge, mostly from Alabama and panhandle Florida west into ‘Louisi- ana. Regrettably, large loans from DUKE and NCU were annotated A. rufa with- out checking “scabra” characters. They are entered with an asterisk so as to pro- vide an idea of distribution of A. rufa in the Carolinas, anda more careful check for the few, if any, “scabra” morphs will be made. Anthenantia rufa (Elliott) Schultes forma rufa ALABAMA. Baldwin Co.: soe (VDB). Butler Co.: Kral 62998 (VDB). Conecuh Co.: Kral 83306 (VDB); Kral 52367 (V : McDaniel 7958 (VDB); Kral 52367 (VDB). Houston Co.: Mac Donald 11801 (V DB), Brae eae) ae (VDB). Mobile Co.: John & Connie Taylor 15260 (BRIT). FLORIDA. no county but suspect Duval, 1883, A.H. Curtiss (TAES). Bay Co.: Kral 52190 (VDB); Godfr & Houk 61532 (SMU). Gadsden Co.: Kral 64456B (VDB). Liberty Co.: Godfrey 84413 (BRI aly - FSU) Wakulla Co.: Henderson 63-1379 (SMU). Walton Co.: Ward 7417 with Hunter (VDB). Washington Co.: Kral with Godfrey 5976 (DUKE, FSU, VDB). GEORGIA. Baker Co.: Kral 56732 (VDB). Coffee Co.: Kral 83766 (VDB). Long Co.: Kral 18864 (VDB). Pierce Co.: Kral 79975 (V DB). Turner Co.: hove with Carter 84177 (VDB). Wayne Co.: Duncan 7670 (SMU). Worth Co.: Kral 81802 (VDB). LOU NA. St. Tammany Parish: 17 Nov 1936, Penfound s.n. (NO). MISSISSIPPI. Hancock Co.: Clarke se aT Harrison Co.: Tracy 3819 & 8590 (TAES). Jackson Co.: Tracy 82 (TAES), Earle 239A. NORTH CA LINA. Bladen Co.: Ahles with Leisner 33368; Ahles nee (NCU) Brunswick Co.: Blomquist me (NCSC, SMU). Craven Co.: Brown 2332 (TEX). Duplin Co.: Ahles 35796A*. Pender Co.: Blomquist 10075, 10076 (TEX), Ahles 36235 (NCU, 5-MU). SOUTH CAROLINA. Bamberg Co.: Ahles 37755 (NCU). Ches- terfield Co.: Radford 18760* (NCU). Georgetown Co.: ee 31389* (NCU). f. jeg FE euck Anthenantia ALABAMA. Baldwin Co.: Wilhelm 1188 (VDB); ee on 89038 (V DB). Covington Co.: Duncan et al. 14181 ae Kral 41655, 80081D, 86880 (V DB). Escambia Co.: Kral 33829, 44787, 44885 (V DB). Geneva Co.: Kral 90294 (V DB). Mobile Co.: Kral 26949, 93368 (VDB). M e Co.: Kral 44380, 69624, 90519, 90572 (VDB). Washington Co.: LeLong 6818 (VDB); Kral 25901, 90526, 90527 (this last one first iden- o tified as A. villosa ts of pale spices a es set to be distributed, (V oe FLORIDA. Alachua Co.: Silveus 6742 ‘). Baker Co.: Godfrey 74696 (VDB). Bay Co.: 24 Oct 1980, Athey s.n. (VDB) GEORGIA. Tift Co.: eae 237 (TAES). Ware Co.: ge s Ge) Worth ( Co.: Kral 51569 (VDB). LOUISIANA. St. Tammany Parish: 17 Nov 1938, Se ane s.n. (NO). Vernon Parish: Thomas & Allen Fic. 4. Idealized sketct f floral and ive Anthenantia. A.A £-| Fiat (elas poe . L eo at blade base and on auride. B. Three ligular Bee idealized for A. villosa, with three cross- s-sections (top); side view of Jest neatly lunction, A. ee oe C. A. rufa, over feat, pcuoblaule ane cross- eeco at Peas (above)-, side fa (t D. A. t glume abaxial side (top), cross- section ie; uot for ae nS ot hairs); adaxial view (bottom). E. en papa abaxial side (top); cross iddle) (k ). F. Side view of A. ue shirelet lowe! (staminate) flower atanthedk: G, Side view of A. rufa“upper” floret at anthesis tig 1 (lef g section cane cu auones attish Mower) palea, A. villosa eiperow) deog f tion of I. Adaxial w,A 1 1 (upper) fl f A. rufa opened We degrees, the perfect a prior to anthesis, K. Reale view of young gj ium, abaxial side. L. Three A ct floret, Anthenantia.N ( i )fl A. villosa, just prior to is. 0. Side vi f lodicule, A I g the typical tion (left); ideog f position of flower parts, tha th + 4th + + SIS. the two lodicules, y 308 BRIT.ORG/SIDA 21(1) 98029 (NLU, VDB); Thomas 5568 (VDB). Washington Parish: Kral 83043 (VDB). MISSISSIPPI. Han- cock Co.: Jones 20345 (VDB); Sargent 9007 (SMU); Clarke 5887 (BRIT). Harrison Co.: Tracy 3819 (TAES). Stone Co.: Kral 93351 (VDB). NORTH CAROLINA. Hitchcock 290 (LL). CONCLUSIONS My study of Anthenantia has been based on field observation along with care- ful artwork and morphologic evaluation. Such foci allow the following: 1, Anthenantia, so far as the North American flora is considered, is a dis- tinct genus of panicoid grasses. It is the only genus in our area to combine (1) a scaly-rhizomatous, loosely caespitose habit (2) a paniculate inflorescence, wit (3) spikelets similar to those of Panicum but lacking a first glume, (4) the sec- ond glume and first lemma 5-nerved, with at least the median three flanked by narrow longitudinal rows of elongate, often papillose-base (strumose) tri- chomes, and (5) two florets, the lower “glumelike” lemma with its hyaline, bicarinate palea enclosing a male floret or sterile, the second (or upper) lemma and palea coriaceous, enclosing a (usually) perfect floret. 2. A third “species-level” taxon, Anthenantia texana, exists west of the Mis- sissippi Embayment, where it may share area, if not exact habitat, with A. villosa and possibly also A. rufa in eastern Texas and western Louisiana. Anthenantia texana may actually be the only species to be found in Arkansas. 3. Anexamination of the synonymy and taxonomic dispositions in Anthen- antia and morphologically adjacent genera seems (to me) to show a definitely confused set of concepts. Since the whole tribe, for that matter the whole family, is having a vigorous and very controversial “shakedown,” it is useful to point out two genera with strong resemblance to Anthenantia, namely Leptocory- phium Nees and Melinis P. Beauv. Leptocoryphium has two species: L. lanatum (Kunth) Nees ranges from Argentina north into Mexico and the Anti'lles, in the north of its range frequenting pine savanna, oak-pine land, open savanna, and pasturage within these systems. In short, it has an “Anthenantia-type” habitat. My first encounter with the species was in Nicaragua (“Zeylaya, burnt savanna by road to Limbaika, ca. 1 km e of jet. rd to Limbaika, 10 July 1982, R. Kral 64344"). I did not identify the plant but did note its strong resemblance to Anthenantia. From my recent examination of herbarium material and from excellent de- scriptions by Pohl for Flora Meso-america (1994) and Flora Nicaragua (2001), | am further intrigued. The species is described as having a cormose, fibrillose base but otherwise seems to differ vegetatively from Anthenantia in no signifi- cant way. The inflorescence is paniculate, sinuously branched, the spikelets are similar in design, the lower glume is lacking, the second glume and lower palea have the same nervation, with trichomes elongate and in longitudinal rows. The only significantly different character state in the spikelet seems to be in the lower floret, which has no palea and which is sterile. The upper floret in character and dimensions of lemma and palea and in perianth, stamens, and an a KRAL, AN EVALUATION OF ANTHAENANTIA 309 gynoecium is very similar. The fruits are similar. | suggest that if a tight de- scription of Leptocoryphium were spaced so that a similar one of Anthenantia were laid in on alternate lines, there would be a strong agreement. It is signifi- cant that one of the most excellent of observers, George Bentham, treated L. lanatumas Anthenantia lanata (Kunth) Benth. Melinis P. Beauv., an African genus of 22 species, also is similar to Anthenantia but appears to be a more distant relative and is in fact placed in subtribe Melinidinae by current authors, while Anthenantia and Leptocory- phium are morphologically aligned with subtribe Digitariinae. The two spe- cies of Melinis in the Americas are the weedy invasives M. repens (Willd.) Zizka ssp. repens (Rhyncheletrum repens [Willd] CE. Hubb. = R. roseum (Nees) Stapf & CE. Hubb.) and M. minutiflora P. Beauv, which are rapidly occupying dis- turbed sandy or gravelly areas (overfarmed situations, fields, railroad rights- of-way, etc.). The former has gotten into the range (if not the habitat) of Anthenantia in northern Florida, south Georgia, and Texas. The latter is so far confined to Florida. A quick scan of a living M. repens makes one note the strong resemblance to A. villosa. The general dimensions of culm, leaf (shape, surfaces, ciliate blade margins, the geniculate “bend,” the character of ligule and auricle) and the feathery panicle of pinkish to silvery-white hairy spikelets, all are de- ceptively similar. However, a closer look reveals a different plant base, namely caespitose but not rhizomatous, the culms often short-decumbent, themselves geniculate at base and with adventitious roots. In the panicle the spikelets, at first appressed-silk y-pubescent, later “fuzzy” with elongate spreading trichomes, are superficially similar to Anthenantia, but a closer inspection reveals (a) lat- eral compression rather than dorsiventral, (b) two glumes, both keeled apically and aristate, and (c) second glume and lower lemma about equal in length with lower half gibbous, the rounded backs promincety pustular-papillose, the hairs liberally i 1, not in longitudinal rows, the upper half abruptly narrowed toa strongly-later ally come keeled, aristate (in the glume) beak. Finally, while the two florets are similar in composition to Anthenantia (lower floret with a hyaline, 2-keeled palea and tristaminate flower, the upper one typically perfect and navicular) the lemma and palea are chartaceous rather than coria- ceous and show a slight lateral compression. The caryopses, while slightly similar, also show bilateral symmetry and an eccentric style base. Thus, I have the impression that Leptocoryphium, having so many charac- ters in common, could indeed be merged with Anthenantia, an opinion already given by George Bentham. For Melinis, on the other hand, the symmetry of spikelet, the presence of two awned glumes, the disposition of hairs and papil- lae on the laterally (rather than dorsiventrally) compressed scales or their tips, and the chartaceous, rather than coriaceous, laterally compressed upper floret constitute a significant set of differences and suggest a different evolutionary alliance 310 BRIT.ORG/SIDA 21(1) ACKNOWLEDGMENTS The help of curatorial and management staffs of all is hereby gratefully ac- knowledged as well as for careful editorial suggestions by agrostologist Victo- ria Hollowell (MO). The invaluable help of Barney Lipscomb and that of Porter Lowry (P) is greatly appreciated. Barney made digital images of my drawings of A. rufa and of A. villosa and sent them to Porter, who compared them with Michaux’s type collection of Anthenantia villosa. The kindness of these two gentlemen is gratefully acknowledged! — REFERENCES BarkworTH, M.E., K.M. Carpets, S.LonG,and M.B. Piep (eds.). 2003. Magnoliphyta: Commelinidae: Poaceae part 2. Flora of North America: North America north of Mexico. Oxford Uni- versity Press. New York. Vol. 25. PaLisor DE Beauvois, A.M.F.J.1812.Essai d'une nouvell agrostographie. ..de Fain, Paris, France. Corrett, D.S. and M.C. JoHnston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner E.uort,S.1821.A sketch of the botany of South Carolina and Georgia /(Facsimile of 18 1824 edition) Hafner Publishing Company, New York. Vol. 1. Goutp, F.W.1975. The grasses of Texas. Texas A&M University Press, College Station. Harcn, S.L., KN. Ganoui, and L.E. Brown. 2001.Checklist of the vascular plants of Texas. Agric. Exp. Sta. Misc. Publ. No. 1655. HitcHcock, A.S.1951.Manual of the grasses of the United States. Second Edition edited by A. Chase. Dover Publications, Inc. New York. Micaux, A. 1803. Flora Boreali-Americana...Parisii & Argentorati, France. Vol. 1:43. Naso, G.V. 1903. Anthenantia. In: Small, J.K., Flora of the southeastern United States. (Anthenantia rufa forma scabra Nash). P. 79. Pout, R.W. 1994. Leptocoryphium. In: G. Davidse, M. Sousa S., A.O. Chater, eds. Flora Me- soamerica: Alistmataceae a Cyperaceae. Universidad Nacional Autonoma de México, Missouri Botanical Garden, and Natural History Museum (London). Vol. 6: 371. Pout, R.W. 2001 Leptocoryphium. In: W.D. Stevens, C. Ulloa Ulloa, A. Pool, O.M. Montiel, eds. Flora Nicaragua: Angiosperms (Pandanaceae—Zygophyllaceae). Monogr. Syst. Bot. Mis- souri Bot. Gard. 85(3):2071. Ravrorb, A.E., H.E. AHLes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. The Univ. of North Carolina Press, Chapel Hill. Sma, J.K. 1903. Manual of the southeastern flora. The Science Press, Lancaster, Pennsylva- nia. SMALL, J.K. 1933. Manual of the southeastern flora. John Kunkel Small, New York. CALAMAGROSTIS COAHUILENSIS AND C. DIVARICATA (POACEAE: POOIDEAE: AGROSTIDINAE), TWO NEW SPECIES FROM MEXICO Paul M. Peterson and Robert J.Soreng Jesus Valdés-Reyna Department of Botany Pane anNG de Botanica Nati aie all ee peel) Univerisida noma Agraria ‘Antonio Narro” nian Inst apna a Washington, District ee umbia 2001 e pea Neder Coahuila MEXICO peterson.paul@ eon uaaan.mx ABSTRACT Calamagrosti huilensis P.M. Peterson, Soreng & See ome na, ASP: nov. es Cease paelosains P.M. Peterson & Soreng, sp. nov, from } occurs on calcareous slopes in the mountains of southeastern Coahuila ee Souiniwesien Nuevo ae anes S aan occurs on moist ee and mossy cliffs 2: scuiawesters anes heat eee 1.2mm long), nearly spice ia es, scaberul hises, gla pees ee scabrous sheath summits, and somewhat dense but not spikelike, areanieh panicles. Calamagrostis divaricata seems morphologically allied to C. pringlei but differs by having wider panicles (4-1 cm) with branches reflexed spreading and divaricate, shorter spikelets (3.4-4.3 mm long), shorter glumes (3.4-4.3 mm long) that are both l-veined, and shorter lemmas (3.8-4.3 mm long) with an awn inserted on upper 1/4 to 1/3. io) RESUMEN describen y se puseen ae Mexico Coie cagtllensts PM. Peterson, sane _ eas Reyna, sp. nov, y Cal g , Sp. nov. C se encuentra en laderas calca las montanas del sureste del estado de ernie y suroeste del tado Nuevo Leon; y C. divaricata se encuentra en laderas humedas y riscos musgosos del suroeste ‘el estado de sea Calamagrostis coahuilensis es See ae simil: ara e purpurascens pee difiere en posect anteras mas ou 1.2mm largo), ligulas casi lisas loso, apices de las y penicnias densas pero no espigadas, de color eee Calamagrostis divaricata | fologi doaC. pringlei pero difiere en poseer paniculas mas amplias (4- 10 cm) con Tomercone: reflexas, extendidas y divaricadas, espiguillas mas cortas (3.4-4.3 mm largo), glumas mas cortas (3.4-4.3 mm largo) ambas uninervadas, y lemas mas cortas (3.8-4.3 mm largo) con una arista insertada desde 1/4 hasta 1/3 arriba de la mitad. Calamagrostis is characterized as having single-flowered spikelets, one or three- veined glumes as long or exceeding the floret in length (rarely slightly shorter), non-keeled lemmas that are membranous or cartilaginous (infrequently hya- line), usually witha single dorsally attached awn (rarely awnless),a callus with a crown of hairs, caryopses with short hilums and without apical hairs, and lodicules that are apically membranous (Clayton & Renvoize 1986; Watson & SIDA 21(1): 311 - 320. 2004 312 BRIT.ORG/SIDA 21(1 Dallwitz 1992). Worldwide, Calamagrostis includes between 230 (Watson & Dallwitz 1992) and 263 (reported by D. Clayton & B. Simon, pers. comm. 2004) species. In North, Central, and South America 131 species are reported in Soreng et al. (2003). In México the following 11 species of Calamagrostis are recognized: C. erectifol id Hitche., C. eriantha (Kunth) Steud., C. guatemalensis Fitehe. 2G, intermedia (J. Presl) Steud., C. oridzabae (Rupr. ex E. Fourn.) Beal, C. pringlei Scribn. ex Beal, C. purpurascens R. Br, C. rigescens QJ. Presl) Scribn., C. tolucensis (Kunth) Trin. ex Steud., C. valida Sohns [syn=C. mcvaughei Sohns,; McVaugh (1983) was the first to select this name asasynonym, see Article 11.5 for equally published species in the Code of Botanical Nomenclature (Greuter et al. 2000), and C. vulcanica Swallen (Chimal 1987; Espejo-Serna et al. 2000). While collecting grasses and specifically searching for unique forms of Trisetum in Durango, México (Finot et al. 2004) in the fall of 2003 the first au- thor gathered a specimen that seemed to be an undescribed species. At first glance this specimen appeared to be two or more flowered but upon closer in- spection and under magnification all spikelets contained a single floret. After studying morphologically similar material from México collected by the first author, an additional undescribed species was found in Coahuila. The two new species are clearly members of subfamily Pooideae, tribe Poeae, subtribe Agrostidinae (Soreng et al. 2003, 2004). We describe these two specimens as new species of Calamagrostis — Calamagrostis coahuilensis P.M. Peterson, Soreng & Valdes-Reyna, sp. nov. (Figs. , 2). Type: MEXICO. COAHUILA. Sierra Madre Oriental, 32 mi SE of Saltillo and 8 mi SE of Jame on road to Sierra La Viga, 3240 m, 29 Sep 1990, P.M. Peterson, CR. Annable & J. Valdés-Reyna 10051b (HOL OTYPE: USI). A Calamagrosti purpurascenti R.Br. antheris 1-12 mm longis, ligulis lere laevigatis, rhachibus scaberulis, vagina apicali glabra vel scabrella antrorse, paniculis aliquantum densis sed non spicae L similibus, viridis, recedit. Caespitose perennial with intravaginal and extravaginal shoot initiation. Culms (22-)40-LOO cm tall, erect, glabrous; internodes glabrous. Sheaths 6-28 cm long, shorter than the internodes, glabrous, often fibrous below with age; margins smooth; cataphylls and lower sheaths glabrous; summit glabrous or minutely antrorsely scabrous. Ligules 5-8 mm long, membranous often lacerate, firmer below, nearly smooth, apex obtuse to acute. Blades 8-20 cm long, 2-4 mm wide, flat, apically acuminate, glabrous, scabrous above. Panicles (7-)9-20 cm long, 0.5-1.2 cm wide, contracted, somewhat dense but not spike-like, greenish; ra- chis scaberulous; inflorescence branches mostly 2-10 cm long below, the branches, ascending and tightly appressed, mostly [loriferous near base, one or two per node. Spikelets 5.8-7.2 mm long, |-flowered, tightly appressed to the branches, dark greenish; pedicels 0.6-4 mm long, ascending, scaberulous; rachilla 2.4-3mm long, covered with stiff hairs, the hairs 0.5-1 mm long. Glumes 5.8-7.2 mm long, lanceolate, about equal or subequal in length, longer than the PETERSON ET AL., 3133 Fic. 1. Calamagrosti huilensis (P Annable & Valdés-Reyna 100516). A. Habit. B. Inflorescence.C. Sheath, ligule, and blade. D. Spikelet. E. Lower glume, dorsal view. F. Upper glume, dorsal view. G. Floret, lateral view. H. Lemma, lateral view. I. Lemma, dorsal view. J. Palea with rachilla, dorsal view. K. Stamens. L. Pistil. M. Lodicules. N. Caryopsis, dorsal view. 314 BRIT.ORG/SIDA 21(1) \ = \ L \ Baja / ) California _ | : | ( ° ~ \ | Sonora \ \ \ \ Chihuahua N\ \ \ \ ae \ \ , ) \ { j ) ce >? \ J \ Coahuila \ J wS Baja \ >) \ / = a \. California\, | \. Sur x \ | \ ; ( J as aN \ ¢ > \ ) er a \ ? a a x ‘4 “Nuevo \ \ is \ ( Sinaloa / Durango ) — Leon \ \, i \ Rac 7 / { \e Monterrey ) J N \ = \ \ ? . ay ae \ \ \ { — \ s \ xf A i : tS Durango” A \ ‘ m - C. divaricata \ >. | / \ 200 km \ . / e. i, & a-C. coahuilensis , »>~ (San Luis) Tamaulipas, 200 miles \.{ 1. |Zacatecas\. Potosi \_, ? Fic ? i etethertt ae eee | Hy , a7 - i ae bY ?. , oe 4 RAAu floret, membranous, |-veined, scaberulous along vein; margins hyaline; apex acute. Lemmas 5-5.7 mm long, lanceolate, membranous, yellowish-green, con- spicuously 5-veined near base, scaberulous, awned; apex acute, entire or bifid; awn 8-10 mm long, golden below and purplish above, borne near the base on the lower 1/6, inserted in a groove below and twisted and once-geniculate above; callus obtuse with white hairs, the hairs 0.8-1.8 mm long. Paleas 5-5.6 mm long, about as long as the lemma, hyaline, 2-veined, the veins scaberulous, green; apex acute, entire. Lodicules ca. 1 mm long, lanceolate, lobed, glabrous. Stamens 3; an- thers 1-1.2 mm long, purple. Ovary ca. 0.2 mm long, glabrous; styles 2, separate; stigmas 2, feathery, whitish. Caryopses 2-2.5 mm long, fusiform, light greenish. Phenology.—Flowering in late August, September, and early October. Distribution —Calamagrostis coahuilensis is known from the mountains of Coahuila and Nuevo Leon growing on calcareous slopes with Abies mexicana Martinez, Pinus culminicola Andresen & Beaman, P. hartwegii Lindl. Hesperostipa spartea (Trin.) Barkworth, Poa ruprechtii Peyr, Holodiscus discolor (Pursh) Maxim., Quercus greggii (A. DC.) Trel., and scattered Populus tremuloides Michx. Addit | ined yPES): MEXICO. Coahuila: municipio de Arteaga, near summit of Coalon SE of San his de Las Alazanas and SE of Saltillo, 3120 m, 17 Oct 1989, P.M. Peter son, J. Valdés-Reynd & J.A. \ ‘a1 8399 (ANSM, US): 29 Aug 1986, J.A. McDonald (TEX). Nuevo Leon: municipio de Galeana, Cerro el Potosi, 3600 m, 3 Aug 1989, A. Garcia-Arévalo 66 (ANSM, CIIDIR): cumbre de Cerro Potosi, ca. Galeana, 29 Aug 1986, J.A. McDonald (TEX); Ladera S, 26 Jul 1985, J.A. McDonald (TEX): J. Ochoa-Guillemard 983 (COCA). PETERSON ET AL., 315 acre e te divaricata PM. Peterson & Soreng, sp. nov. (Figs. 2, 3). Type: CO. DURANGO. Sierra Madre Occidental, ca. 3 mi S of Hwy 40 at Mexiquillo, ee Paso Resbaloso (23°42'46.2"N-105°39'45.1"W), 2520 m, 14 Sep 2003, P.M. Peterson, M.S. Gonzdlez-Elizondo & G. Tefia-Gonzdlez 17774 (HOLOTYPE: US! ISOTYPES: ANSMI, CIEDIR!, MO! RSA! US). A Calamagrosti pringlei Scribn. ex Beal paniculis +-10 cm latis, ramis reflexis, effusis et divaricatis, spiculis 3.4-4.3 mm longis, glumis 3.4-4.3 mm longis, univenis, lemmatibus 3.8-4.3 mm longis, arista inserta supera 1/4-1/3, recedit. Loosely caespitose perennial with short rhizomes and extravaginal shoot ini- tiation. Culms 52-96 cm tall, erect to slightly decumbent near base, glabrous; internodes glabrous. Sheaths 8-20 cm long, shorter or longer than the intern- odes, glabrous; margins smooth; cataphylls and lower sheaths glabrous; sum- mit glabrous. Ligules 2-3.6 mm long, membranous often lacerate, decurrent below; apex truncate, ciliolate. Blades 15-60 cm long, 1-3 mm wide, flat or invo- lute, apically acuminate, glabrous, scabrous above. Panicles 9-16 cm long, 4-10 cm wide, ovate, open, lax, sparsely flowered, greenish; rachis scaberulous; in- {lorescence branches mostly 2-8 cm long below, the branches delicate, ascend- ing and spreading, naked near base, whorled; lower inflorescence nodes with 3-6 branches. Spikelets 3.4-4.3 mm long, l-flowered, spreading from the branches, yellowish-green; pedicels 1-5 mm long, ascending to reflexed and spreading, scaberulous; rachilla 1.6-1.9 mm long, covered with stiff hairs, the hairs up to 1.5 mm long. Glumes 3.4-4.3 mm long, lanceolate, about equal in length, shorter or longer than the floret, membranous, I-veined, scaberulous along vein; margins hvaline: apex acute to ac inate. Lemmas 3.8-4.3 mm long, lanceolate, membranous, yellowish-green, 5-veined, scaberulous, awned; apex acute, entire,awn 4-6 mm long, yellowish, borne on the upper 1/3 or 1/4, straight or twisted and once-geniculate; callus obtuse with short white hairs, the hairs 0.2-1 mm long. Paleas 3.8-4.4 mm long, as long or slightly longer than the lemma, hyaline, 2-veined, the veins scaberulous, green; apex acute, entire. Lodi- cules 0.7-0.8 mm long, lanceolate, lobed, glabrous. Stamens 3; anthers 1.8-2.2 mm long, yellow. Ovary 0.2-0.4 mm long, glabrous; styles 2, separate; stigmas 2, feathery, whitish. Caryopses 1.8-2.5 mm long, fusiform, light brownish. Phenology.—Flowering in September. Distribution —Calamagrostis divaricata is known only from the type lo- cality growing on moist slopes and mossy cliffs with Pinus cooperi CE. Blanco, P durangensis Martinez, P. thuite C. Ehrenb. ex Schltdl., Quercus sideroxyla Bonpl., Arbutus occidentalis McVaugh & Rosatti, Cupressus arizonica Greene, and A M U hle nl ergid dalamosde Vasey. DISCUSSION Calamagrostis coahuilensis remained undetected by biologists for almost 19 years. The first collections of this species were apparently made by McDonald ‘ih ih j r | i i Wi BRIT.ORG/SIDA 21(1) Fic. 3. ae divaricata nies Gonzdlez-Elizondo & Tena-Gonzdlez 17774). A. Habit.B. Inflorescence.C. Sheath, ligule, and blade. ventral view, I. aria oe es dorsal view. J. Palea, dorsal view. K. S les. Lodicu elet.E wer glume, dorsal view. F. Uppe rGlume, dorsal view. c Lema, dorsal view.H. Lemma LL Gyn noecium, mature. M ! PETERSON ET Al 317 in 1985 from the Cumbre de Cerro Potosi. All specimens that we have seen from Coahuila and Nuevo Leon that were previously determined as C. purpurascens or C. pringlei are C. coahuilensis. Therefore, C. purpurascens is not found in México. Calamagrostis coahuilensis is morphologically similar to C. purpurascens, but can be separated from the latter by having anthers that are shorter (1-1.2 mm long), ligules that are nearly smooth, a scaberulous rachis, sheath summits that are glabrous or minutely antrorsely scabrous, and green- ish panicles that are somewhat dense but not spikelike (Table 1). Populations of C. coahuilensis are separated from the closest known locality of C. purpurascens in the Sangre de Cristo Mountains, Taos County, New México (Allred 1993) by over 800 mi. McDonald (1990) and Garcia-Arevalo and Gonzalez-Elizondo (1991) first reported C. purpurascens from Coahuila and Nuevo Leon, México. Apparently, Manuel Gonzalez-Ledesma, John Reeder, and Charlotte Reeder iden- tified the grasses for McDonald (1990) since they appear in the acknowledg- ments for having reviewed that family. This is not surprising since in most char- acteristics the range of variation in C. coahuilensis overlaps that of C. purpurascens. An adequate illustration (Ochoa 983) of the new species appears in Beetle’s (1987) treatment of the grasses of México, although he referred to it as C. pringlei. It is surprising that C. divaricata is known only from a single, recent col- lection since it occurs near a rather heavily used tourist destination. Calamagrostis divaricata seems morphologically allied to C. pringlei and can be differentiated from the latter by having wider panicles (4-10 cm) with branches reflexed spreading and divaricate, shorter spikelets (3.4-4.3 mm long), shorter glumes (3.4-4.3 mm long) that are both 1-veined, and shorter lemmas (3.8-4.3 mm long) with an awn inserted on upper 1/4 to 1/3 (Table L.). The infrageneric classification of Calamagrostis has not been critically tested, and it is not usually discussed for the New World, other than the accep- tance of the segregate genus Deyeuxia by some South American taxonomists (Rugolo de Agrasar 1978, 1998; Villavicencio 1995). The genus (including Deyeuxia) is divided into at least four subgenera with many sections, subsec- tions, and series (Rozhevits & Shishkin 1963; Tzvelev 1976; Wasiljew 1960). All of these Russian agrostologists placed C. purpurascens in Calamagrostis sect. Deyeuxia (Clarion ex P. Beauv) Rchb., and Wasiljew (whose classification was world-wide in scope) placed C. pringlei in the same section. Several infrageneric taxa were accepted for the New World (Soreng et al. 2003). Based on callus length (1/2 or less the lemma length), lemmas membranous, and presence of a well-developed rachilla extention, both of the new species appear to belong in Calamagrostis subgen. Deyeuxia sect. Deyeuxia. A key for separating the two new species from other species of Calamagrostis in northern México (Chihua- hua, Coahuila, Durango, Nuevo Leon, Sinaloa, and Tamaulipas) follows. _ — Nair Tasce 1. Salient features comparing Calamagrostis hy yi is, C. divaricata, C. pringlei, C. purpurascens, and C. valida. Characters C. coahuilensis C. divaricata C. pringle C. purpurascens C. valida Cataphylls & lower leaf glabrous glabrous densely hirtellous glabrous or glabrous sheath abaxial surface scaberulous Leaf sheath summit glabrous or minutely glabrous glabrous glabrous or minutely Densely hirtellous abaxial surface antrorsely scabrous retrorse-Strigulose Leaf blade & ligule glabrous or nearly glabrous glabrous scabrous sparsely hirtellous adaxial surface smooth Inflorescence width 0.5-1.2 cm 4-10 cm - (1.5-)4-10 cm Inflorescence ascendi branches, arrangement & spikelet placement Spikelet length Spikelet color rachilla length rachilla hair length Glumes length Upper glume, number of veins Lemma length Lemma surface Lemma veins, aspect Lemmatal awn insertion Lemma apex ng and closely appressed; *+floriferous to base 5.8-7.2 mm greenish 24-3 mm 0.5-1 mm 5.8-7.2 mm one 5-5.7 mm smooth conspicuous base, lower 1/6 entire or bifid scending, reflexed spreading to divaricate; not floriferous to base yellowish-green 1.6-1.9mm 0.8-1.5 mm 3.4-4.3 mm one 3.8-4.3 mm scaberulous inconspicuous upper 1/4 to 1/3 entire ascending, loosel floriferous to base pale green to purplish 1.5-2 1-2 mm (4.5-)5-5.8 mm three 45-5 mm scaberulous INCONSpPICUOUS lower 1/3 to middle entire ascending and closely appressed, spike-like; +floriferous to base yellowish-purple 1.8-3 mm 1-2 mm 5-9 mm lower 1/6 to 1/3 bifid ascending, appressed to base spreading; + floriferous to base yellowish-purple 1.5-2mm 2-3 mm 3.4-6.5 mm (one) three 3.4-5 mm smooth inconsp./conspicuous near middle two or four setae or with age teeth (prolongation of veins) Callus hair length 0.8-1.8 mm 0.2-1 mm 0.2-1 mm 0.7-2 mm, 1.3-2.8(-3.5) mm occasionally absent Anther length 1-1.2 mm 1.8-2.2 mm 2-2: 1.5-3.5 mm 1.4-2.2 mm Anther color purple yellow yellowish-purple purple or yellow yellow or purple BLE (L)LZ VaIS/9¥O'LINS PETERSON ET Al 319 KEY TO THE SPECIES OF CALAMAGROSTIS IN NORTHERN MEXICO 1. Panicles 0.5-1.5 cm wide, contracted, de eee flowered;lemma awn inserted at bas oron sont 1/6 to near lower 1/4; anthers 1-1.2 mm long Calamagrostis | oeiuilend: — . Panicles 1.5-10 cm wide, narrow to open and loosely flowered; lemma awn in serted a lower 1/3, middle, and upper 1/4; anthers 1.4-2.4 mm long 2. Lemma apex with two or four setae or teeth (prolongation of veins); callus hairs 1.3-2.8 mm long; rachilla hairs 2-3 mm long; leaf blade, ligule, and summit of sheath sparsely hirtellous Calamagrostis valida 2. ile apex entire and without setae or teeth; callus hairs 0.2—1 mm long;rachilla hairs 0.8-2 mm long; leaf blade, ligule, and summit of sheath glabrous. 3. Te 3-veined; spikelets (4.5) 5—5.8 mm long;lemmas 4.5-5 mm Bee awn inserted on lower 1/3 to middle; panicles 2-4 cm wide, the branche loosely appressed or spreading but not divaricate ee. pringlei 3. Upper glume 1-veined; spikelets 3.4—4.3 mm long;lemmas 3.8-4.3 mm long, awn inserted on upper 1/3 to 1/4; panicles 4-10 cm wide, the branches re- flexed spreading, divaricate Calamagrostis divaricata ACKNOWLEDGMENTS We wish to thank the Smithsonian Institutions, National Museum of Natural History, Biodiversity and Inventory Program, and Research Opportunities Fund for supporting the fieldwork. Appreciation is extended to Alice R. Tangerini for providing the illustrations; Susan J. Pennington for preparing the distribution map; Alain Touwaide and Dan Nicolson for correcting the Latin diagnoses; Kelly Allred and Rich Spellenberg for checking the distribution records in New Mexico; and Teresa Mejia-Saulés for finding inconsistencies in the manuscript. REFERENCES Autrep, K.W.1993.A field guide to the grasses of New Mexico. Department of Agricultural Communications, College of Agriculture and Home Economics, New Mexico State University, Las Cruces. Beetve, A.A., E. Manrique Forceck, C.V. JARAMILLO Luque, M.P. GUERRERO SANCHEZ, J.A. MIRANDA SANCHEZ, C.l. Nunez Tancreoi, y C.A. CHIMAL HERNANDEZ. 1987.Las Gramineas de México, Tomo Il. 344 p. COTECOCA, S.A.R.H., México, D.F., México CHIMAL HERNANDEZ, C.A. 1987. Calamagrostis Adans. In: A.A. Beetle, E. Manrique Forcek., CV. Jaramillo Luque., M.P. Guerrero Sanchez., J.A. Miranda Sanchez., C.l. Nunez Tancredi, y C.A. Chimal Hernandez. 1987. Las Gramineas de México. Tomo Il. COTECOCA, S.A.R.H. México, D.F. Pp. 161-172. Crayton, W.D.and S.A. Renvoize. 1986.Genera graminum: grasses of the World. Her Majesty's Stationary Office, London. EspeJo-Serna, A., A.R. Lopez-Ferrari, and J. VALDES-Reyna. 2000. Poaceae. In: A. Espejo Serna and A.R. LOpez-Ferrari, eds. Las Monocotylidéneas Mexicanas: una sinopsis floristica, Partes IX-XI. Consejo Nacional de la Flora de México, A.C., Universidad Aut6noma Metropolitana-lzapalapa, y Comision Nacional para el conocimiento y uso de la Biodiversidad, México, D.F. Pp. 10:7-236 320 BRIT.ORG/SIDA 21(1) Finor, V.L., PM. Peterson, R.J. Sorenc, and F.O. ZuLoaca. 2004. A revision of Trisetum, Peyritschia, and Sphenopholis (Poaceae: Pooideae: Aveninae) in México and Central America. Ann. Missouri Bot. Gard. 91:1-30. GarciA-ArevaLo, A. and S. GONZALEZ-ELIzonbo. 1991. Flora y vegetacion de la cima del Cerro Potosi, Nuevo Leon, México. Acta Bot. Mex. 13:53-74. GreuTer, W.J. McNelt, F.R. Barrie, HM. Burpet, V. DEMOULIN, T.S. Ficcueiras, D.H. NicoLson, P.-C. Sitva, J.E. Skoa, P. TReHANE, NJ. TURLAND, and D.L. Hawkswortu. 2000. International code of botani- cal nomenclature (Saint Luis Code). Koeltz Scientific Books, KOnigstein, Germany. McDonato, J.A. 1990. The alpine-subalpine flora of northeastern México. Sida 14:21-28. McVaush, R. 1983. Flora Novo-Galiciana. A descriptive account of the vascular plants of western México. Gramineae. 14: 1-436. University of Michigan Press, Ann Arbor. Rozuevirs, R.Yu.and B.K. SHISHKIN. 1963 (1934). Flora of the U.S.S.R. vol. Il, XXIV.Gramineae Juss. National Science Foundation, Washington, D.C. Pp. 152-184. RUGOLO DE Acrasar, Z.E. 1978, Las especies patagdnicas del genero Deyeuxia Clar. (Gramineae) de la Argentina y de Chile. Darwiniana 21:417-453. RUGOLO DE Acrasar, Z.E. 1998. Deyeuxia. In: S.A. Renvoize, Gramineas de Bolivia. The Royal Botanic Gardens, Kew. Pp. 181-235. SorenG, R.J., PM. Peterson, G. Daviose, EJ. Juoziewicz, F.0. Zutoaca, T.S. FILGuEIRAS, and O. MorRONE. 2003. Catalogue of New World grasses (Poaceae): IV: subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1—730. SorenG, R.J., G. Davipse, P.M. Peterson, F.O. ZuLoaea, E. J. Jupziewicz, T.S. FItGueiRAS, and O. MORRONE. 2004.Catalogue of New World grasses (Poaceae). http://mobot.mobot.orgw3T/Search/ nwgc.html and Classification of New World Grasses. http://mobot.mobot.org/W3T/ Search/nwgclass.html Tzvetev, N.N. 1983 (1976). Grasses of the Soviet Union (Zlaki SSSR), part Il. Oxonian Press, New Delhi, India. VILLAVICENCIO, X, 1995, Revision der gattung Deyeuxia in Bolivien. 1-304. Ph.D. Thesis, Freien Universitat Berlin, Berlin. Wasivew, W.N. 1960. Das system der gattung Calamagrostis Roth. Feddes Repert. 63(31): 229-251. Watson, L. and M.J. Datuwitz. 1992. The grass genera of the World. C.A.B. International, Wallingford, U.K. ETHNOBOTANY OF RHODIOLA ROSEA (CRASSULACEAE) IN NORWAY Torbjorn Alm Department of Botany Troms@ Museum University of Troms@ N-9037 Tromse@, NORWAY ABSTRACT Rhodiola rosea is a widespread species in Norway. It is well known in Norwegian folk tradition, with a variety of vernacular names, of which many reflect its traditional uses. Past use asa cure for scurvy in cattle may explain names with the prefix kalv- (“calf”). Its widespread use as a hair wash is also reflected in vernacular names. In the past, Rhodiola was planted on turf roofs to protect them from fire, i.e. as an apotropaic (supposedly averting evil forces); this tradition is documented as early as the 13th century. SAMMENDRAG Rosenrot Rhodiola rosea er en vanlig og vidt utbredt art i Norge. Den er vel kjent i folketradisjonen, med en lang rekke folkelige navn. En god del av disse gjenspeiler artens tradisjonelle bruksomrader. Bruk som et botemiddel mot skjerbuk hos kyr kan forklare de mange navnene pa nal ly. Den flittige bruken til harvask gar likeens igjen i mange lokale navn. Tidligere ble rosenrot pa tovtak [£ for A beskytte dem mot brann, dvs. som er vernerad; denne tradisjonen er dokumentert allerede pa 1200-tallet. INTRODUCTION Rhodiola rosea L. (Crassulaceae) (syn. Sedum rosea (L.) Scop,, S. rhodiola DC.) is a common species in the mountains and coastal districts of Norway, occurring abundantly both on coastal cliffs and in alpine habitats, from sea level to 2280 m a.s.l.in the mountains of Central Norway (Elven 1994). The lowland plants belong to subsp. rosea, whereas those of high mountain habitats are similar to the arctic subsp. arctica (Boriss.) A. & D. Love (Elven 1994). The species as such is easily recognizable The genus Rhodiola, formerly often included in Sedum, comprises about 50 species, centered on the mountains of East Asia (Springate 1995). Whereas Sedum has hermaphroditic flowers, many Rhodiola species, including R. rosea, are dioecious (Lippert 1995). Rhodiola rosea is a hemicryptophyte with thick rhizomes. The annual shoots are unbranched, and densely clad with flat, fleshy leaves. Rhodiola rosea has a circumpolar-montane/alpine distribution (Hultén & Fries 1986; Lippert 1995). Hultén (1958) considered R. rosea “A collective spe- cies consisting of numerous races differing chiefly in size, form and dentation of the leaves and in the colour of the flowers.” a SIDA 21(1): 321 — 344, 2004 322 BRIT.ORG/SIDA 21(1) Rhodiola rosea has received increased attention during the last few years, not least due to its alleged medical properties, e.g. as an adaptogen, supposedly enhancing memory, stress mastering, etc.; some of these effects have been con- firmed by recent studies (Boon-Niermeijer et al. 2000; Spasov et al. 2000). Germano & Ramazanov (1999) provide extensive references to Russian litera- ture on its medical properties. For a review of current knowledge, see Brown et al. (2002). Folk tradition claiming positive health effects derives largely from Asia (eastern Russia, Mongolia, China). Norwegian tradition is less inclined to claim “wondrous” properties, but may provide valuable information of potential use Both the rhizomes and flowering stems of R. rosea have found a number of uses in folk tradition in Norway. The present paper aims at a comprehensive review of the ethnobotany of R. rosea in Norway, including vernacular names, medici- nal uses and other traditions. Unless otherwise stated, all citations have been Wn translated from Norwegian. Sources Hoeg (1974) assembled a vast body of information on plant names and uses in Norway, including a three-page chapter on Rhodiola rosea. It is largely based on his own data, and fails to incorporate more than a fraction of existing litera- ture, e.g. the interesting note of Soreide (1952). Nordhagen (1934) studied the old tradition of planting R. rosea on house roofs in Norway. Alm (1996a) dis- cussed its past use as a cure for scurvy. Recently, Dragland (2001) reviewed data on R. rosed as part of a project aimed at commercial cultivation in Norway. His report includes some ethnobotanical data, but these are largely culled from Hoeg (1974) and some secondary sources. Scattered notes on R. rosea and its uses in Norway occur in numerous other publications on folk medicine and other traditions. In addition to these, I have incorporated some data from archival sources, mainly NFS (Norsk olkeminnesamling/Norwegian folklore collection) and NOS (Norsk ordbok, seddelarkivet/Norwegian dictionary, card archive). Furthermore, some data have been excerpted from my own ethnobotanical field work in North Nor- way; these are referred to as “interview + year.” Informants are not identified here; transcripts and some recordings of the original interviews are stored at the Department of Botany, Tromse Museum (TROM). re Vernacular Names In Norwegian folk tradition, widespread and well-known plants are usually known either under a single, ubiquitous vernacular name (e.g. bldbaer, “blue berries” for Vaccinium myrtillus L.), or display a substantial array of widely dif- ferent names (e.g. more than 100 names for Dactylorhiza maculata (L.) S06, see Alm 2000). With more than a half-hundred Norwegian vernacular names re- corded so far, Rhodiola rosea belongs to the latter group. No record of its Norse — ALM, ETHNOBOTANY OF RHODIOLA ROSEA 323 name (or names) seems to have survived (Heizmann 1993): the oldest names included in the present paper date back to 1599. A compilation of Norwegian and Sami names is found in Table 1. Some are of local use, and may be confined toa single village; others are widespread. A couple of name-groups (see Fig. 1) are of particular interest: a) In Western Norway, R. rosea has a set of names with the prefix syste—or syster, systre, soster and similar terms (Hoeg 1974; Lagerberg et al. 1955; Lid 1941; Nordhagen 1934; Soreide 1952; Strom 1762), e.g. systegras (“-gras”), systelykjil C- key”), systeroter (“roots”). Nordhagen (1934) noted only a deviant form, systergras (“-grass”), and suggested that syster (interpreted as meaning “sister”) might refer to the occurrence of separate male and female plants—perhaps a rather too botanical explanation, though Heeg (1974) noted that people at least locally had noticed that there were two different kinds of plants. The latter au- thor recorded a number of similar names, partly with syste- and partly with syster-. He suggested the latter to be correct, perhaps influenced by the linguist Ivar Aasen’s record of sostregras (‘sisters grass”) at Sunnmere in the 1830s (see Lid 1941). Evidently, Hoeg was not aware of the discussion in Sereide (1952), who argued convincingly that the prefix was related to the verb syste or syfte, “clean” or “purify.” Syftesok is an old calendar term for July 2, at which date apotropaic plants (e.g. twigs of Alnus sp.and Juniperus communis L.) were placed in the fields to ward off vermin (Bugge 1921; Riste 1916; Soreide 1952; Wille 1786). Thus, systegras and similar names may suggest that folk tradition ascribed Rhodiola rosea abilities to ward off evil. This is confirmed by the widespread belief in the plant's ability to protect against fire (see below). The prefix syster- (and soster-), ie. “sister,” is probably a younger re-interpretation of an old name. b) In most of North Norway, R. rosea is known under names with the pre- fix kalv-or kalve- (“calf”), e.g. kalvegress calf grass”), kalvedans (calf dance”), and others (Alm 1996a; Elvebakk 1979; Heltzen 1834; Hoeg 1974; Lagerberg et al. 1955; Morkved 1996; Oksendal 1977; Qvigstad 1901; Strompdal 1929; NOS). Kalverot (calf root”) is also mentioned from the Bergen area in 1599 (Bring 1758; Holmboe 1953; Lagerberg et al. 1955; Rordam 1873). The name may well derive from North Norway, which was the source of most of the fish exported from the major trade port of Bergen. The origin of the prefix kalv- is unclear. Alm (1996a) suggested that it might be due to the past use of R. rosea as cattle fodder, potentially an important cure for scurvy (see below). An alternative explana- tion is a relation to the old Norse term for the thick muscle on the hind side of the leg (Lagerberg et al. 1955), still known (by a Norse loan-word) as calfin En- glish. If so, kalv- might refer to R. rosea’s thick leaves. At least some of the asso- ciated suffixes support the former interpretation, e.g. kalvegror Ccalf growth”) and perhaps the widespread kalvedans (‘calf dance”), an unlikely acitivity for starving cattle. Reichborn-Kjennerud (1922) suggested that kalvedans could refer to the plant’s dancing and nodding behaviour in wind, but this leaves the — 324 BRIT.ORG/SIDA 21(1) Taste 1. Norwegian and Sami vernacular names for Rhodiola rosea in Norway.Names are given with modern Norwegian and North Sami spellings; deviant spellings used in the original sources are indicated. "Interview" refer to ethnobotanical records made by the author. As far as possible, geo- graphic origin is indicated, using present-day administrative units (counties and municipalities). English translations are literal, but note that some may be folk re-interpretations of older names (see text). A few terms cannot be translated, e.g. the suffix tort, otherwise a frequent vernacular name for Cicerbita alpina (L.) Wallr., and mo/mosott, which are widespread Norwegian terms for a somewhat diffuse disease. Vernacular name English translation Area and source Norwegian Bald aldans Bal dance North Norway: Nordland: Leirfjord Jenssen 1982:43) Bergakonge Rock king Western pees Hordaland: Hardanger, interior area (H@eg 1974:597) Bergbukk Rock buck Southern eo elses: Bygland; Central Norway: Nord-Trendelag: Verdal (Haeg 1974:596) Bergebruse Rock buzz Southern Norway: Aust-Agder: Bykle; Bygland; Valle (H@eq 1974:596 Bergebukk Rock buck Southern Norway: Aust-Agder: Valle (Haeq 1974:596) Berggull Rock gold Northern Norway: Nordland: Beiarn (Vreim 1943:50, footnote, as bergull) Bergkrans Rock wreath Central Norway: S@r-Trandelag: Oppdal (Donali 1988:587, as beerjkrans; Hoeg 1974:597) Berjebruse Rock buzz Western Norway: Mere og Romsdal: Surnadal (H@eg 1974:596-597, as beerjebruse) Bukkablom Buck flower Western Norway: Rogaland: Forsand (H@eg 1974:597) Bukkebruse Buck buzz Nordland: Vesterdlen; Troms: Senja (Ross 1895:66, 71) Feitbokk Fat buck Central Norway: Nord-Trandelag: Lierne (S@rli); Namdalseid (Haeg 1974:596) Fjellbruse Mountain buzz Mare og Romsdal: Nordmare (Ross 1895:66) Fjellbu Mountain plant North Norway: Nordland: Sorfold (Engan 2002:56 Fjyellkaur Mountain curl Norway: Nordland (H@eg 1974:597; Nordhagen 1934:1 24); Saltdal (Nordhagen 1934:1 24); Skjerstad (Ha@eg 1974:597; NOS): Beiarn (Vegusdal 1979:159, as fjellikaur;Vreim 1943-50; NOS Fjellknesk(e) Mountain squeak North Norway: Nordland: Bode (H@eg 1974:597) Fjellknirke Mountain squeak North Norway: Finnmark: Nordkapp (H@eg 1974:597) Fjellkrans Mountain wreath Central Norway: Ser-Trandelag: Midtre Gauldal (Haukdal 1961:141-142); Soknedal (Haeg 1974:597 ALM, ETHNOBOTANY OF RHODIOLA ROSEA Tas_e 1. continued Vernacular name English translation Area and source Norwegian Gnagblomst Harblomster Harkjeks Harvekster Harvokst Harvokster Hedlekaure Heilkaur Heilkaure Itch (squeak?) flower Hair flowers Hair “biscuit” (plant) Hair growth Hair growth Hair growth Heal-curl ? Heal-cur| Heal-curl North Norway: Nordland: Veeray (Haeg 1974:597 Central Norway: Ser-Trendelag: Bjugn: Stjarna (Haeg 1974:596) Western Norway: Hordaland: Fusa (H@eg 1974:596) Western Norway: Sogn og Fjordane: Nordfjord (Krogh 1813:266, 282, as Haarweexter); Mare og Romsdal area (Gunnerus 1766:49, as Haarveexter); Central Norway: Sor-Trandelag: Bjugn; Stjarna (H@eg 1974:596) Eastern Norway: Oppland: Dovre; Lesja; Lom (Haeg 1974:596), Nord-Fron: Sikkilsdalen (Nordhagen 1934:124, footnote); Gudbrandsdalen area (Nordhagen 1934:123); Western Norway: More og Romsdal: Romsdalen area (Nordhagen 1934:123); Central Norway: Se@r-Trendelag: Oppdal (Donali 1988:587; Rise 1947:56) Norway, unspecified (SchUbeler 1888:268, as Haarvokster); Eastern ae ee al area ee 934: bes Oppland: L (Hoeg | 596);Western Norwa ee O Romsdal Gunmen 1766: Hs da 1952:37), Romsdalen area (Nordhage 1934:123); Sunnme@re area (Strom ae 19, as Haar-Voxter); Nesset: Eresfjorden (Haeg 1974:596); Volda (H@eg 1974:596); Fraeena (Haeg 1974:596): Halsa (H@eg 1974:596); Surnadal (Haeg 1974:596; NOS, as harvakkster); Sunndal: Alvundeid (Haeg 1974:596); Central Norway: Graten—Vedey (Dahl 1896:70, 1899:8, 58, as {aar-Voxter); Sor-Trandelag: Oppdal (Donali 1988:587; Rise 1947:50, 56); Roan (Haeg 1974:596): Nord-Trondelag: Meraker (H@eg 1974:596); Stjordal/Meraker Pe 1968:510) Western Norway: Hordaland: Hardanger area (Reichborn-Kjennerud 1922:57): Kvam (NFS Gade-Gron 149) North Norway: Nordland: Rana (unpublished note by Axel Blytt, 1870 Norway, unspecified (Nordhagen 1947:39); North Norway: Nordland: Helgeland? (Reichborn-Kjennerud 192 — TA BLE 1. continued BRIT.ORG/SIDA 21(1) Vernacular name English translation Area and source Norwegian Helkaure Huskall Kalvdans Kalveblomst Kalveblomster Kalvedans Kalvedaude Kalvegras/kalvgras Kalvegress Kalvegror Kalvekal Kalveknark Kalverompe Heal-cur| House man Calf dance Calf flower Calf flowers Calf dance Calf death Calf grass Calf grass Calf growth Calf cabbage Calf squeak ( Calf tail ’) North Norway: Nordland: ia as IPUBIBNE diary by Axel Blytt, 1870-1 he Central Norway: Nord ae Lierne: Nordli (Haeg 1974:595, 597) ees Norway: Nord- Winer ale aed 974:596-597); North Norway: N nea (NFS O.A. H@eg 572); ears (NFS O.A.H@eg 425); Bronney (Strompdal 1929:85, NFS O.A. Hoeg 415; NOS); Vefsn (Oksendal 1977:99; NFS O.A. Haeg 88, 445, 719); Rana (NOS, note by Hallfrid Christiansen); Leirfjord Uenssen 1982:43): Rodey (NFS O.A. H@eg 36); Flakstad (Morkved 1996:18); Troms: Harstad; Ibestad; Troms@: Hille BY (| loeg 1974:597). Finnmark: naa (Haeg 1974: 7 North Norway: Troms: Harstad; rvoy (Heeg 1974: oo Kvaenangen ee ae 2004 North Norway (Elvebakk 1979-47) Central Norway: Nord-Trandelag: Vikna (Hoeg 1974:597); North Norway (Elvebakk 1979:47, as kalvedainjs); Nordland: Vefsn (NFS O.A.H@eg 145); Donna (NFS O.A H@eg 145); am Wage! 1974: 597): Troms: Harstad (Haec ); Ibestad (Haeg 1974:597); ae oes 1974:597- NOS; inter- view 2001); Kvaenangen (interview 1988, 2003); Finnmark: Hammerfest (Ovigstad 1901:311;interview 1998): Kvalsun (interview 2003), Lebesby (Hoeg 1974:597), Garnvik (interview 1998); Berlevag (interview 2003); Batsfjord (interview 1988) North Norway: Nordland: Bindal (Hae 1974:597) North Norway: Nordland: Luray; Rodey; Ofoten area; Troms: Karlsay (Haeg 1974:597) North Norway: Nordland: Rana (Heltzen 1834:8, as Kalvegrees) North Norway: Nordland: Vagan (Merkved 1996:18); Langeya area (Markved 1996:18); Narvik (NOS, note by Hallfrid Christiansen Troms: Troms@ (NOS) North Norway: Nordland: Rana (H@eg 1974:597) North Norway: Troms: Bjarkay (Hoeg 1974:597) North Norway: Troms: Salangen; Tran@y (Hoeg 1974:597 ALM, ETHNOBOTANY OF RHODIOLA ROSEA TABLE 1. continued Vernacular name English translation Area and source Norwegian Kalverot Kalvespreng Kalvestolpe Kalvgras Kalvgress Kalvgror Kalvkaur Kalvkal Kalvlyng Kalvtort Knerke Langrot Longrot Lungerot Morot Mosottrot Oksfot Oksstut Calf root Calf burst Calf post Calf grass Calf grass Calf growth Calf curl Calf cabbage Calf heather Calf tort “Squeek" Long (?) root Long (?) root Lung root Mo root Mosott root Ox foot Ox bull Western Norway: Hordaland: cae! (Bring 1758:64: Holmboe 1953:9-10; Rar 1873:405, diary note of Sivert pains July 5, 1599, as Kalfweroed) North Norway: Nordland: Branney (H@eg 1974:597) North Norway: Nordland: Heray (H@eg 597) North HAG roan oo (NFS O.A. Hoeg 159); Rady (NFS O.A.H North Norway: cee cane (unpublished note by he eels Vestvagoy (Merkved 1 18) North a ie (Morkved 1996:18); Bo (NFS O.A. Haeg 496); Sortland (NFS O.A.Ho@eg 715); Langaya (Merkved 1996:18) Southeast Nopvay te Hole Sandefjord (NFS O.A.Ho 03); North Norway: Nordland: Rana (Hoeg 1 974:597) North Norway: Nordland: Sortland (NFS O.A. H@eg 346) North ee Nordland: Hadsel (Hoeg 1974:597) oe Norway: Nordland: Hadsel (Haeg 1974:597) North Norway: Finnmark: Hammerfest (Haeg 1974:597; interview 2002) Central Norway: Sor-Trandelag: Melhus: Holonda; Midtre Gauldal: Singsas; Holtalen: Alen (Haeg 1974:597) Central Norway: Ser-Trandelag: Melhus: Halonda; irae is Singsas; Holtalen: Alen (Haeg 19 Eastern Nor way:! 1 -Alvdal; Tynset 1974:596-597) Central Norway: Nord-Trondelag: Snasa (H@eg 1974:597) Central Norway: Nord-Trondelag: Snasa (H@eg 1974:597) — Haeg Central Norway: Nord-Trendelag: Nery: Foldereid; Grong: Harran; Haylandet; Snasa (H@eg 1974:597) Central Norway: Nord-Trandelag: Neroy: Foldereid (Hoeg 1974:597) Taste 1. continued BRIT.ORG/SIDA 21(1) Vernacular name English translation Area and source Norwegian Smeerbukk Smerbukk Smorbukk/smorbokk Smerstakk Sestelakjel Sestelokla Sesteloklar Sastergras Sostergress Sesterlok S@stregras Stubberot Systegras Systelykjel Butter buck Butter buck Butter buck Butter skirt Purifying key Purifying keys Purifying keys Sister grass Sister grass Sister onion Sisters grass Stump root Purifying grass Purifying key Eastern Norway: Telemark: Vinje (Halvorsen 1988:197) Norway, unspecified ahaa: eee 08) Eastern Norway: Oppland: La Vestre Slidre (NOS, ease notes by G. Kirkevoll and A. @degaard); Valdres a (Kirkevoll 1940:173); Western ce Hordaland: Fusa; Grat | ', Vaksdal: Vos | several other municipalities (NOS Nereneieree notes by T. Hannaas and N. Lid); Mare og Romsdal (NOS); Romsdal area (Gunnerus 1766:49, as Sme@rbuk, cf. Nordhagen 1934:1 24) Wetera) Norv lay More og Romsdal: Freena; R| AA es ft {ve Pal hae | ieee een Norway: S@r-Trandelag: Roan; Nord-Trandelag: Meraker (Hoeg 1974:596) Norway, unspecified (Schtibeler 1888:268, as Smerstak; cf. Nordhagen 1934:124) Western Norway: Sogn og Fjordane: Jalster (Hoeg 1974:597) Western ane Sogn og Fjordane: Jalster (Haeg 1974:597) Western cone Sogn og Fjordane: Jalster (H@eg 1974:597) Mare og Romsdal oe 952:37); Halsa (NFS Maurit Fugelsoy IIl:7) Western Norway: Sogn og Fjordane: Nordfjord (Krogh 1813:266, as Sastergraes); M Romsdal: Sunnmare area (Stram 1762, as Saster-Grees; Gunnerus 1766:49, as Soster-grees) Western Norway: Sogn og Fjordane: Jalster (Haeg 1974:597) Western Norway: More og Romsdal: Sunnmere area (Lid 1941:75, annotation on a oe in lvar Aasen’s herbarium from 1837 — 39) Norway (Bring 1758: UN by Sivert Grubbe (as Stubberod) in 1 Western Norway: Sogn og Bremanger: Davik (Haeg om aan Eid (Hoeg 1974:595, 597; a0.) ie 29): ae Stryn: Innvik (H@eg 1974 Romsdal: Rauma oreide 19599 Q) Western Norway: Sogn og Fjordane: Naustdal (S@reide 1952:29) ALM, ETHNOBOTANY OF RHODIOLA ROSEA TaBle 1. continued Vernacular name English translation Area and source Norwegian Systelykla Systergras Systerlykkjel’e Systerlykla Systerlyklar Systerose Systerater Systerrot Systregras Takbruse Takdupp Takgull Takkrans Taklauk Tjukke-Nils Trappakall Trappkall Trappros Galbberahta Purifying keys Sister grass Sister keys Sister keys Sister keys Purifying rose Puryifying roots Sister root Sisters grass Roof buzz Roof-nod Roof gold Roof wreath Roof onion Fat Nils Staircase man Staircase man Staircase rose Calf sprout Western poe Sogn og Fjordane: Naustdal (Hoeg 1974:597) Norway (Schubeler 1888:268; Reichborn- Kjennerud 1922:57); Sogn og Fjordane: Leerdal: Borgund; Gloppen; Jalster (Haeg 1974:579); Monae! VINCI, 953:49); More og Romsdal: Volda; @rsta (Haeg 1974:597) Western Norway: Sogn og Fjordane: Farde (Haeg 1974:597) Western Norway: Sogn og Fjordane: Gaular (Ho@eg 1974:597) Western Norway: Sogn og Fjordane: Ferde; Gaular; Jalster (H@eg 1974:597) n Nor lare og Romsdal: Vanylven: Syvde (H@eg 1974: 597) Western aoe Sogn og Fjordane: Selje (S@reide 1952:29) Western ere Hordaland: Lindas: Alversund (H@eg 1974:596-597); Sogn og Fjordane: Fjaler (NOS, unpublished note by H. Tveit); More og Romsdal: Vanylven: Syvde (Haeg | 974:597) Borgund; Mare og Romsdal: Stranda: Sunnylven (H@eg 1974:597) Western Norway: Meng 09) Romseal BInGS (Hoaeg 1974:597;Mo al (H@eg 1974:597) 8 en eee net (Hoeg 1974:597) ata Norway: Nord-Trondelag: Neeray (Haeg 1974:597) Central Norway: S@r-Trendelag: Meldal (Haeg 1974:597) North Norway: Nordland: Beiarn (Vequsdal 1979:159) Western Norway: Rogaland: Kvitsay (Haeg 1974:597) Central Norway: Sor-Trondelag: Afjord: Stokksund (Haeg 1974: 597 Central Norway: S@r-Trandelag: Roan (H@eg 1974:597) Central Norway: Sor-Trondelag: Roan (H@eg 1974:597) North Norway: Finnmark: Alta (interview 1998) 330 BRIT.ORG/SIDA 21(1) Tape 1. continued Vernacular name English translation Area and source North Sami Galbberassi Calf grass North Norway: Troms: Lavangen (Qvigstad 1901:311, as galbe-rasse); Finnmark Deatnu/Tana (interview 1996 Galbberassit Calf grasses North Norway: Troms: Lyngen (Qvigstad 1932:78, as galberaset) prefix unexplained. The few Sami names recorded so far, galbberdhta Ccalf sprout”) and galbberdssi “calf grass”), belong to the same name complex. c) Vernacular names with the prefix hdr- Chair”) occur in parts of south- ern Norway (Dahl 1898; Donali 1988:587; Hoeg 1974; Lagerberg et al. 1955; Nordhagen 1934; Reichborn-Kjennerud 1941; Rise 1947; Schtibeler 1888; NOS). They obviously reflect the use of a decoction of R. rosea as a hair wash. Ver- nacular names incorporating the term kaur or kaure, meaning something curly may also refer to such use (Nordhagen 1934, 1947). d) R. rosea has sometimes appropriated vernacular names more frequently used for other species, e.g. smorbukk “butter buck”), more frequently a name for Hylotelephium telephium (L.) Ohba (syn. Sedum telephium L.),e.g. in inland areas adjacent to the latter’s m ainly coastal distribution (Halvorsen 1988; Kirkevoll 1940), and taklauk (roof onion”), normally Sempervivum tectorum L.,at a few stations (Vegusdal 1979). e) A deviant name, knerke, was recorded from Hammerfest, Finnmark by Hoeg (1974); two informants both explained it as an onematopoeticon: “it squeaks when the flowers are touched”, “if the leaves are touched, one hears a squeaking sound” (translated from Norwegian). This seems to be a well-known tradition in parts of Finnmark; one of my own informants knew the name kalvedans from the same area, but suggested knerke as an alternative name: “That sound when you touch them, it says[knerke]... | think it was a sound-like name we had.” (interviews 1998 & 2002). A few other, sound-based vernacular names are known from North Norway (Table 1). Rosenrot (rose root”), the “official” Norwegian name of R. rosea, has been introduced through floras. lt was first used in the Danish herbal of Paulli 1648), probably based on the German Rosenwurz, and has no root in Norwegian folk tradition (Hoeg 1974; Nordhagen 1934). Only two Sami names for R. rosea have been recorded in Norway: gal bberassi both in Troms and Finnmark, and gdlbberdhta in the latter county. Gdlbberdssi is obviously a Norwegian loan-word, identical to “kalvegras.” Qvigstad (1901, 1932) noted it from Lavangen and Lyngen, Troms. It is still used, and was re- fo. —_— ALM, ETHNOBOTANY OF RHODIOLA ROSEA Goes oo an Oo oA ts pot MPa " rad Stee + O @) t 4 ot | (@] i van) . ff=ut \ : t “ft \ ‘ ! Lay | QO O~ \ 2 ait * ? { 0. : ary ‘ é a ) Oo Re Ke ce ; o PO "m oN ALS ’ 9 mee Ne Hy Oo feels *. * H a a) \ 1 bored i \ i 2 ai 1°) ' 7 ; . r ; . Caw », s, a 7 ay 1 H % , t ' ' 7 c , ’ e oe / \ ~ ? / \ ' A o Ly 4 - } . ¢ p . \ i , \ o~ ! \ \ 4 3 ‘ mu t ) on : i ‘ ‘ ' my 65 Fe uJ 1 ? z oand D | om + ‘ ri ™“ | . - ? H | x . i “. ' ‘ ; : | : ar ? “ ¢ ’ < ‘ A i i ., . Uy ~~ i: » 1’ 5 ' : hy Al i ea H ' ’ | . ' Y é ‘ ‘ ’ / i ’ H Fa | pha r; | 1 7 a , ‘ ? / N. , é 7 - ‘, ’ ‘ vl ? / if ont 7 Fi a ‘ ¢ ‘ ‘ ‘ ~ ra J: | oe Cratul meaning “calf”: N NUT wegian A Norwegian J AaIV /k names with orefix of the eased type, based o on ine Help ste or on SS (dots). B. Names with a prefix uar rec) UdIC5) 332 BRIT.ORG/SIDA 21(1) corded in the Deatnu/Tana area of Finnmark in the 1990s (Alm & Iversen, un- published data). Rhodiola rosea as a Cure for Scur Nordal (1939) studied the contents of vitamine C in various plants species tra- ditionally used to treat scurvy in Norway. Scurvy-grass Cochlearia officinalis L.and cloudberries Rubus chamaemorus L. may have been the most important of these, at least for human use (Alm 1995, 1996b: Eckblad 1989). Rhodiola rosea contains less vitamine C; 12 mg/g (rhizome) and 33 mg/g (fresh leaves) accord- ing to Nordal (1939, 1946). Still, it may have been an important source, in par- ticular for livestock (Alm 1996a). Insufficient feeding of cattle during the win- ter was an established tradition in Norway, and if spring was late, livestock could be heavily affected by scurvy. Cloudberries (and, perhaps less so, scurvy-grass) were reserved for human consumption, but Rhodiola rosea was not. Secondly, due to its preference for cliff habitats, the rhizomes could be collected even in late winter. It is also one of the first plants to sprout in spring, and both rhi- zomes and fresh shoots may thus have served as an important remedy for scurvy in livestock. Records in Elvebakk (1979) and Hoeg (1974) confirm that R. rosea was usually collected during the period of fodder shortage in late spring. As noted above, such use is a likely explanation for the wide-spread vernacular names with kalv- as a prefix, which may thus reflect the species’ importance for making cattle (and calves) survive (Alm 1996a; Heeg 1974). A letter to Olaf I. Ronning, then curator at Troms@ museum, was sent asa response to his popular account (Ronning 1959) of R. rosea and its uses. The comment, based on tradition at the west coast of Senja island, Troms county, confirms the use as cattle fodder: “In my childhood, about the turn of the century [1900], we las] children were frequently told to collect kalvegror, ie., rosenrot [Rhodiola rosea]. It was used as fodder for calves, which had been] born during the winter, in spring. As it was almost always a hortage of fodder [at this time], the kalvegror wasa welcome food an the calves (and a vitamine source). The kalvegror sprouted earlier than other plants (in our area of southern Berg, and the islets) and was easy to get hold of” (letter from Paul Hay, Grylletjord, dated January 20, 1960). Some early data on R. rosea ethnobotany, including its use as an antiscorbutic, were recorded during the Danish-Norwegian king Christian IV's naval expedi- tion to the northern outposts of his kingdom in 1599 (Alm 1996a: Nielsen 1873). At least two of the Danes participating (Sivert Grubbe and Jonas Charisius) wrote diaries. A comparison of the two leaves no doubt that Grubbe was the better botanist; his diary (Bring 1758; Rordam 1873) contains scattered notes on plants seen during the voyage. Despite this, Charisius (1773-76) is more fre- quently cited, often from an 18th century transcript by Hans Paus (extracts e.g, in Hansen & Schmidt 1985). Although the most interesting comment on R. rosea was made during a visit to the NW Kola peninsula, now on Russian territory (but also claimed as ALM, ETHNOBOTANY OF RHODIOLA ROSEA 333 belonging to the king’s territory), it is worth recording here: “On this island Kildin there is found a kind of herb at the shore, which the Sami and Russians call orpin, and [which] is very remarkable to use for scurvy, the root smells of rose, and tastes well in beer” (Charisius 1773-76:74; Hansen & Schmidt 1985:191, translated from Danish). Qvigstad (1901), in his survey of Sami plant names, accepted orpin as an East SAmi term. It is not, however, included in a dictionary of East Sami as used in the Kola Peninsula (Genetz 1891). As noted by Alm (1996a), orpinis the French name of R. rosea, and it is highly unlikely that a similar name should occur in both East Samiand Russian. The suggestion that it was termed thus by the Sami is obviously wrong: educated Russians could perhaps have known this name. Qvigstad (1901) couples orpin with the Norwegian term Stubberod, seemingly as a translation. The latter name is mentioned in Grubbe’s Latin diary, in the entry for May 18, 1599: “Collegimus in isto monte herbas, quas angli vocant orpin, Norvegi Stubberod, praesentissimum re- medium contra scorbutum, in illis locis valde familiare. Capi sesti loco acetarii.” (Bring 1758:40—41)—‘In this mountain we collected some hed watch une eu call orpin, the Norwegians stubberod, an excellent remedy for scurvy, and very freq | Our captain [king Christian IV] used the leaves of this plant as a salad.” Thus, Grubbe certainly did not record orpin as a local name (though it was hardly an English term either, as he believed). Stubberod (stubberot in present- day Norwegian) is clearly identified as a Norwegian name, otherwise unknown, but easily comprehensible; its meaning (stump root”) is a descriptive term for the rhizomes in their early spring state, ie. just at the time they were presum- ably gathered asa cure for scurvy. Pontoppidan (1752) also noted that R. rosea had “en herlig Kraft mod Skiorbug”—“a splendid force against scurvy.” Gunnerus (1766) mentioned that “Radix scorbuticis salutaris’—“the root heals scurvy”; this seems to be the last suggestion of human use as an antiscorbutic in Norway. Rhodiola rosea as Food Both the rhizomes and green parts of Rhodiola rosea are edible. Human con- sumption is well known from other areas, e.g. Siberia, but seems to have been rare in Norway—at least according to our present knowledge. Ruge (1762), vicar in Valdres, advocated plantations of R. rosea as food in times of need, but it is unlikely that this idea was based on local tradition. On the contrary, he referred to Egede (1741), who had noted that the rhizomes were eaten in Greenland. Ruge added that “I have eaten it myself, both fried and roasted as well as cooked, and neither in taste nor effect have I found it unpleasant.” (Ruge 1762:286). Heoeg (1974) recorded a single, modern instance of R. rosea consumption in western Norway; an informant from Uvdal (Buskerud, SE Norway) claimed that the leaves had been ground and mixed in dough. In addition, children could eat the fresh leaves. 334 BRIT.ORG/SIDA 21(1) According to Engan (2002), people at Sorfold in North Norway used ground bark of birch (Betula pubescens Ehrh.) asa flour substitute during times of need. The resulting flour was coarse and hardly suited for dough. In order to improve it,an unidentified alpine herb called fjellbu “mountain plant”) was added. The brief description included fits Rhodiola rosea, and hardly anything else, sug- gesting that it may have had some tradition as an emergency food. No informa- tion is given on the plant part used, only that it was boiled before being added to the flour, Rhodiola rosea as Hair Wash A decoction of R. rosea has been widely used as a hair-wash in Norway, e.g. along the western coast northwards at least to Nordland (Donali 1988:587; Gunnerus 1766, Hoeg 1974; Hukkelberg 1952; Lagerberg et al. 1955; Reichborn-Kjennerud 1922, 1941; Rise 1947; Strem 1762; NOS). This is frequently indicated by its ver- nacular names (see above), as noted e.g. by the latter author: “In the fjords, it is called Soster-Grees lie. sister grass], but otherwise everywhere lat Sunnmare, western Norway] Haar-Voxter [hair growth], because one boils it in water and washes the hair with it, in the belief, that it will grow well thereafter” (Strom 1762:119). Two hundred years later, folk tradition in More og Romsdal remained unchanged: “Sastergras [sister gras] is also called hdrvokster [hair growth] be- cause if one boiled it in water and washed the hair with it, the hair would grow much better afterwards” (Hukkelberg 1952:37). The motivation for using R. rosea as a hair-wash, according to folk tradi- tion, varies slightly. Some claimed that it prevented hair loss (Donali 1988; Rise 1947), and ensured a long, fine hair, others that it stimulated hair growth or healed various complaints affecting the hair, e.g. dandruff. An undated archi- val note, again from More og Romsdal, adds a piece of folk etymology: “Women who were about to loose their hair made a kind of hair oil from rosenrot which made the hair grow well—for this reason [it was] called sastergras [sister grassl.” (NFS Maurits Fugelsoy IIL7). At Narvik in Nordland, North Norway, folk tradition claimed that plants for use as hair wash should be collected in spring or early summer: — “If you manage to collect kalvegror before the cuckoo [Cuculus canorus] cries, and then boils a decoc- tion from it, itis good for washing the hair.” (NOS, unpublished 1940s note by Hallfrid Christiansen). Other Medical Uses Apart from its use a an antiscorbutic and as a hair-wash, Rhodiola rosea has found little use in Norwegian folk medicine. Hoeg (1974) noted that it had been used as a remedy for lung diseases at Volda, W Norway, but also that this might be inspired by its local vernacular name, lungerot (‘lung root”). From the Nordfjord area of W Norway, Krogh (1813) merely noted that the rhizome was sometimes used as a medicine, and considered an adstringent. According to Kirkevoll (940), R. rosea was “much used as an ointment for wounds” in the ALM, ETHNOBOTANY OF RHODIOLA ROSEA 335 Valdres area of interior SE Norway. In Sunnfjord, W Norway, a poultice of R. rosea and ground oats Avena sativa L. was used to treat wrenched or swollen limbs (Reichborn-Kjennerud 1922; NFS Gade-Gron 149). The vernacular name mosottrot, used in Snasa, Nord-Trondelag (Hoeg 1974), also implies a medical use. In folk medicine, mosott was a frequently diagnosed, if rather ill-defined disease, usually treated with magical means, in particular “measuring” the patient’s body with a woolen tread. In Lyngen (Troms, North Norway), Sami folk medicine used a decoction to treat urinary disorder (Qvigstad 1932; Steen 1961):“A decoction of Rhodiola rosea (..) is drunk, and the softened roots [rhizomes] are rubbed at both sides of the joint. When the urine starts to drip, another mouthful is consumed.” (Qvigstad 1932:78; translated from German). In his late 17th century topographical description of Finnmark, district govenor Hans H. Lilienskiold mentioned that the Sami of Finnmark used rose water to treat eye diseases (Qvigstad 1932). No further details are given. Rosa majalis J. Herrmann is the only species of the genus occurring in Finnmark, but it is very rare, and it it thus possible that the cure was a decoction of Rhodiola rosea, with its characteristic, rose-like odour. Folk Veterinary Medicine Apart from its use to heal (or prevent) scurvy, there are few records of Rhodiola rosea being used in folk veterinary medicine in Norway. Heeg (1974) noted that people at Holonda (Ser-Trondelag, Central Norway) had used it as a “horse medi- cine, long ago”—which does not exclude the possibility that even this was as a cure for scurvy. In Hardanger, W Norway, a decoction was given to cattle to treat many kinds of disease, in particular intestinal parasites (Reichborn-kjennerud 1922). The original record, by Lars T. Steine, dated 1920, is found in the NFS archives: “Hedlekaure is the name of a plant that grows preferably in the shade (...) It was used for all kinds of animals; it was boiled and given in drink; it was good for many kinds of livestock diseases in particu- lar intestinal parasites” (NFS Gade-Gron 148). Rhodiola on House Roofs Rhodiola rosea was formerly a frequent sight on turf roofs in Norway. Accord- ing toa widespread tradition, its presence there should protect against fire, e.g. caused by lightning (Fegri 1944; Haukdal 1961; Hoeg 1974; Lagerberg et al. 1955, Nordhagen 1934; Vegusdal 1979). A similar tradition related to Sempervivum tectorum L. is widespread in Europe. S. tectorum is not an indigenous species in Norway, but has a long history of cultivation, and is naturalized at scattered stations in southernmost Norway (Elven 1994; Nordhagen 1941). Both living plants and the associated traditions are likely to have been imported from Cen- tral Europe during the middle ages. Further north in Norway, in areas where S. tectorum does not thrive, Rhodiola rosea has acquired a similar reputation of 336 BRIT.ORG/SIDA 21(1) protecting roofs from fire. According to folk tradition in Gauldal, Central Nor- way, its purpose was to avert the anger of Tor, the Norse god who controlled thunder and lightning: “It was the god Tor who was to be appeased by planting fjellkransen on the roof. When he saw the yellow flower on the roof, he passed by without striking it with fire.” (Haukdal 1971:14D. The occurrence of an unidentified herb “on everyone’s house roof” in Bergen in the early 13th century is mentioned in the saga of the Norwegian king Hakon Hakonssen. Nordhagen (1934) argued convincingly that the unnamed plant had to be R. rosea. Its use on house roofs is also mentioned in a 16th century description of Bergen (Nordhagen 1934, 1941), and remained well-known as long as turf roofs were common, ie. well into the early 20th century. This practice is documented from much of southern Norway, especially along the west coast, northwards to the Salten area of Nordland, North Norway (Donali 1988; Grue 1943; Haukdal 1961; Heeg 1974; Kirkevoll 1940: Leirfall 1968; Nordhagen 1934; Sereide 1952; Strompdal 1929; Vegusdal 1979; Vreim 1943). In Beiarn, Nordland, people believed that the custom had been introduced from Bergen (Vegusdal 1979). Melheim (1953) studied the flora of turf roofs in Hornindal, W Norway in the early 1930s. Rhodiola rosea was still frequent, and always planted if found on house roofs, but people could not any longer give any traditional reason for this practice: “Some said it was because the plant was so beautiful, others be- cause great-grandpa had wished it, others because their neighbours had it like that.” (Melheim 1953:49). Norman (1894), in his flora of Norway north of the Polar Circle, recorded numerous stations of R. rosed on house roofs, noting for one of these that it was “som sedvanlig plantet’—“as usual planted.” Nordhagen (1934) saw several roofs with planted R. roseain Nordland in the 1930s, and recorded vestiges of an oral tradition related to such use. Vreim (1943:50) noted that he had seen “whole roofs of” R. rosea in Beiarn, Nordland. Vegusdal (1979:142) recorded a fading tra- dition in the same area: “Old superstition said that fjelljkauren[R. rosea] on the roof had power to protect from fire. Lately, some have retained the custom to keep up the old tradition at the farm, or because they found it decorative. In the past, there were few turf roofs, at least at @ynes, on which fjelljkaur had not been planted, and they grew well.” This tradition may have been known further north as well, as indicated by the following note from Alta, Finnmark: “In Sami, we call it gal bberahttd. And it used to grow on old turf roofs.” (interview 1998). Sempervivum tectorum may form dense mats of leaf rosettes, which may to some extent protect turf and straw roofs from fire (Fegri 1944). Rhodiola does not, and there is not much reason to believe that the plants as such had any real protective value. However, the belief may have some ecological justifi- cation. In Central Europe, Sempervivum tectorum thrives on old turf roofs, since os = ALM, ETHNOBOTANY OF RHODIOLA ROSEA 337 these provide more humid conditions—and would thus less easily burn—than young or new roofs, which are much drier, just as R. rosea in Norway is often found on old turf roofs. Still, the Norwegian tradition of planting Rhodiola on turf roofs is likely to have been inspired by similar use of Sempervivumtectorum in areas further south, with Rhodiola rosea serving mostly as an apotropaic. Old photographs (e.g. in Nordhagen 1934 and Fegri 1944) frequently show only a single or a few plants placed along the ridge of the roof. On the other hand, Vreim (1943), in a booklet on timber houses and turf roofs, considered this as a useful practice, offering protection of the weakest part of the turf cover. In some cases, e.g. at Meraker (Nord-Trondelag, Central Norway), R. rosea was placed over the door (Haeg 1974), reflecting a widespread European tradition of plants used as apotropaics to guard the house entrance. The use of R. rosea on house roofs is probably rare by now, although many turf roofs still exist. A late 2002 search of the digital photo database at Troms@ museum yielded 908 photographs of turf roofs all over Norway, but nota single one with visible R. rosea stands. People’s belief in R. rosea as an apotropaic, protecting houses from fire, may be the reason for its celebrated mention in the saga of king Hakon Hakonsson. In 1218, his mother, Inga of Varteig, was challenged to prove the royal parentage of her son by an ordeal of carrying hot iron. Prior to the task, Sigarr of Brabant, an employe of Earl Skule, suggested to one of her friends that she could protect herself from burns by salving her hands with the juice of a plant. When ques- tioned where to find this wondrous herb, Sigarr answered “bat vex 4 binum husum ok hver manns hér i Bjorgyn”—“it grows on everyone’s house roof here in Bergen.” As noted by Nordhagen (1934), R. rosea is the only likely candidate. Nordhagen, however, fails to note that the offer was refused; Earl Skule was one of Inga’s enemies, and she may have had reason to consider the suggestion as an attempt at foul play (Hertzberg 1912)—perhaps as a means of persuading her that she had nothing to fear from the ordeal, and thus agree to it. If so, the plan failed. The ordeal was successfully completed, as far as the historical sources can tell without resorting to the protective powers of R. rosea, and Inga’s son Hakon Hakonssen went on to become one of the most illustrious Norwegian kings. DISCUSSION The uses recorded for Rhodiola rosea in Norway are closely reflected in other areas settled by people of Norse origin. Debes (1673) noted that a decoction was used as a kind of “rose water” in the Faroes, presumably as a hair wash; an oint- ment was used on wounds (Nordhagen 1934). An unpublished Faroese dictio- nary from about 1670 lists R. rosea as hjalpirot “help root”), suggesting some kind of medicinal use, e.g. to improve hair growth (Lange 1960). In Iceland, a decoction of the fresh rhizome was used to wash the head as a cure for head- 338 BRIT.ORG/SIDA 21(1) ache (Nordhagen 1934); such use was a part of older school medicine. Rhodiola rosea was also dried, mixed with butter, and used as an ointment on old wounds (Hallgrimson 1964; Mohr 1786). In Greenland, the rhizomes of R. rosea were eaten by the Eskimos or Inuits (Birket-Smith 1928; Egede 1741; Hoygaard 1941; Hughes 1960; Schtibeler 1888); leaves and shoots were also consumed. It was also eaten by the Eskimos of North America. In Alaska, both the rhizomes, stems and leaves were eaten, partly fresh (Porsild 1953), fermented or frozen (Moerman 1998). The Eskimos of Nunivak Island, Alaska, prepared a tea from the flowers (Griffin 2001). The former use of R. rosea to treat (or avoid) scurvy is in accordance with its rather high contents of vitamine C. Hoygaard (1941, table D) found it to be the richest terrestrial plant source of vitamine C available to the Eskimos in his study area at East Greenland. Rhodiola rosea is also found in the mountains of Central Europe, and had acquired some reputation as a medicinal plant in this area as well. It is included in several early herbals, e.g. Fuchs (1543) and Bock (1551), and the rhizomes (Ra- dix rosea) were sold in pharmacies. The comments in the major Danish herbal of Paulli (1648) seems to be based mainly on Central European tradition. Rhodiola rosea does not occur in Denmark, and its use here was based on culti- vated plants—Paulli noted that it was found in some “noble gardens”—possibly supplemented by plant material imported from Norway. Both in Denmark and in Central Europe, R. rosea was used to cure head-ache (Paulli 1648, Lippert 1995). The German-speaking farmers of South Tyrol (Italy) believed that a decoction in milk promoted pregnancy (Marzell 1979). In terms of medical properties, the peoples of eastern Asia have found a much wider use of Rhodiola species than that of R. rosea in Norway and other parts of Europe. In Russian Siberia, R. rosea was considered an aphrodisiac, and supposed to ensure a long life. In Mongolia, it was used to treat cancer and tu- berculosis. In China, both R. rosea (imported) and other Rhodiola species were prized for their medical qualities (Germano & Ramazanov 1999). Asa cure for tuberculosis, R. rosea was also used by the western Eskimos of North America: the disease was treated with raw flowers (Lantis 1959). The Eskimos of Nelson Island, Alaska, used R. rosed to treat sores (Ager & Ager 1980). Rhodiola species, e.g. R. quadrifida (Pall.) Fisch. & Mey,, R. rosea,and R. sacra (Prain ex Hamet) S.H. Fu, are currently the focus of substantial interest in terms of medical and pharmacological properties. Studies of Asiatic species have re- vealed a vast array of chemical compounds (see Brown et al. 2002, Kurkin & Zapesochnaya 1986, Yoshikawa et al. 1995, 1996, 1997), including some with promising antiallergenic effects. The only biochemical study of Norwegian R. rosea so far, by Rohloff (2002), identified various terpenes and volatiles, but failed to identify some constituents (rosiridin, rosiridol) reported from Russian material of R. rosea (Kurkin et al. 1995), and found only neglible amounts of ALM, ETHNOBOTANY OF RHODIOLA ROSEA 339 octadecadienoic acid, heptanol derivates and hexadecanoic acid, claimed by Belov et al. (1994) to be some of the main constituents. Thus, the biochemical characteristics of R. rosea may vary according to geographical origin, and Nor- wegian plants could differ from those found in Asia in terms of pharmacologi- cal properties—which would be in accordance with folk tradition in the two areas. Alternatively, its potential qualities were undetected in Norwegian tra- dition, or other plants were considered more effective. Norwegian R. rosed is rich in the phenylpropanoid rosavin (S. Dragland, pers. comm.), specific to R. rosea and now considered to be one the most important constituents in terms of medi- calactivity (Brown etal. 2002), which might support the latter hypothesis. The sparse or absent use of R. rosea as food in Norway may also have a biochemical explanation; according to Hoygaard (1941), Norwegian R. rosea has a much sharper taste than the plants found in Greenland. At present, the old traditions related to R. rosea in Norway are probably fading away, e.g. its former use as an apotropaic on house roofs. The vernacular names are more likely to survive, at least as long as R. rosea remainsa well known and popular ornamental, especially in rock gardens. ACKNOWLEDGMENTS The referees Joachim Thiede and U. Eggli), Steinar Dragland, and Sida editor Barney Lipscomb provided useful comments. Ernst Hogtun prepared the map. ARCHIVAL SOURCES NFS (Norsk folkeminnesamling/Norwegian folklore collection): (a) Gade-Gron, questionaires on Norwegian folk medicine, distributed 1911; (b) original mate- rial of O.A. Hoeg; (c) undated notes by Maurits Fugelsoy; NOS (Norsk ordbok, seddelarkivet/Norwegian dictionary, card archive). REFERENCES Aasen, |. 1860. Norske Plantenavne. Budstikken 2:9-37. Acer, |.A. and L.P. Acer. 1980. Ethnobot of the Eskimos of Nelson Island, Alaska. Arctic Anthropol. 17:27-48. Aum, 1.1995. Multer i folketradisjonen—navn og bruk. Ottar 206:7-12. Aum, T. 1996a. Bruk av rosenrot (Rhodiola rosea) mot skjorbuk. Polarflokken 20:29-32. Aum, T. 1996b. Bruk av skj@rbuksurt (Cochlearia officinalis) i Nord-Norge. Blyttia 54: 185-192. Aum, T. 2000. Dactylorhiza maculata (L.) Sod—Norwegian and Sami ethnobotany of a magical orchid. J. Eur. Orch. 32:189-221. Betov, V.N., TV. Lavrova, N.G. VAsHKevicH, and ALY. MikHaitov. 1994. Extraction of essential oils from plant raw material by steam distillation. Russ. J. Appl. Chem. 67:154-156. BirkeT-Smith, H. 1928. The Greenlanders of the present day.In:M.Vahl,G.C. Amdrup, L. Bobe, and A.S. Jensen, eds. Greenland. Vol. 2. The past and present population of Greenland. 340 BRIT.ORG/SIDA 21(1) CA. Reitzel, Copenhagen—Humphrey Milford/Oxford University Press, London. Pp. 1-208. Bock, H. 1551.New Kreuterbuch. Strassbourg. Boon-NieRMeuer, E.K., A. VAN DER Bera, G. Wikman, and F.A.C. Wiecanr. 2000. Phyto-adaptogens protect against environmental stress-induced death of embryos from the freshwater snail Lymnaea stagnalis. Phytomedicine 7:1-11. Barina, S. 1758.Samling Af atskilliga Handlingar och Paminnelser, Som Férmodligen kunna gifwa Lius i Swanska Historien. Ill. Lund. Brown, R.P., P.L.GerbarG, and Z.Ramazanov 2002: Rhodiola rosea: A phytomedicinal overview. HerbalGram 56:40—52. Bucce, K. 1921. Eneren Uuniperus communis) i norsk folketro og skik. Tidssk. Hist. Bot. 1: 141-170. Cuarisius, J. 1773-76.Kong Christian den Fierdes Reise til Norge og Vardahus | Findmarken. In: Samlung zur Danischen Geschichte, Munzkenntniss, Okonomie und Sprache durch Johann Heinrich Schlegel. Kopenhagen. Pp. 55-90. DAHL, O. 1896. Biskop Gunnerus' virksomhed fornemmelig som botaniker tilligemed en oversigt over botanikens tilstand i Danmark og Norge indtil hans ded. Ill. Johan Ernst Gunnerus. Tillaeg Il: Uddrag af Gunnerus’ breweksling, seerlig til belysning af hans videnskabelige sysler. Hefte 1. Kongel. Norske Vidensk. Selsk. Skrifter 1895 (3):11-224. DAHL, O. 1898. Biskop Gunnerus’ virksomhed fornemmelig som botaniker tilligemed en oversigt over botanikens tilstand i Danmark og Norge indtil hans dad. Ill. Johan Ernst Gunnerus. Tilleeg Il: Uddrag af Gunnerus’ brevweksling, szrlig til belysning af hans videnskabelige sysler. Hefte 4. F. Breve til Gunnerus. Kongel. Norske Vidensk. Selsk. Skr 1898 (4):1-83 Datt, O. 1899. Biskop Gunnerus’ virksomhed fornemmelig som botaniker tilligemed en oversigt over botanikens tilstand i Danmark og Norge indtil hans dad. Ill. Johan Ernst Gunnerus. Tillaeg Il: Uddrag af Gunnerus’ breweksling, serlig til belysning af hans videnskabelige sysler. Hefte 5. Kongel. Norske Vidensk. Selsk. Skr. 1899 (4):1-164. Deses, L. 1673. Faeroae et Faeroa reserata. Kiabenhavn Donau, |. (ed.) 1988. Oppdaling. Ord og uttrykk. (Oppdalsboka, vol. 3). Strindheim trykkeris forlag, Trondheim. DRraGLanb, S. 2001. Rosenrot, botanikk, i toff, dyrking og bruk. En litteraturoversikt. Planteforsk Gronn forskning 2001 (9):1-28 + 14 pp. Ecketan, F.-E. 1989. Molter som skj@rbuksmiddel i skriftlige kilder. Blyttia 46:177-181. Ecebe, H. 1741. Det gamle Gronlands nye Perlustration. Kiobenhavn Etvesakk, A. 1979. Smakebitar fra nord-norsk folkebotanikk med glimt fra Trandelag og Nordmare. Polarflokken 3:42-60. Etven, R. 1994. Johannes Lid & Dagny Tande Lid: Norsk flora. Ed. 6. Det norske samlaget, Oslo. ENcan, H. 2002. Fra barkebradstider. Arbok for Sorfold 2002:56. Facri, K. 1944, Planteverdenen i folketroen.In:S. Solheim, K. Faegri, P Hovda, and S. Johnsen, eds. Folketru og folkeliv. J.W. Cappelens forlag, Oslo. Pp. 41-55. ALM, ETHNOBOTANY OF RHODIOLA ROSEA 341 Fucus, L. 1543.New Kreuterbuch. Genetz, A. 1891.Kuollan lapin murteiden sanakirja ynna kielennaytteita./ Worterbuch der Kola-lappischen Dialekte nebst Sprachproben. Bidrag till kannedom af Finlands natur och folk 50:I-XLVI, 1-291. Germano, C. and Z. Ramazanoy. 1999. Arctic root (Rhodiola rosea). The powerful new gin- seng alternative Kensington Publishing, New York. GrirFiN, D. 2001. Contributions to the ethnobotany of the Cup’it Eskimo, Nunivak Island, Alaska. J. Ethnobiol. 21:91-127. Grue, T. 1943. Os i Osterdalene. Ostgards trykkeri, Roros. Gunnerus, J. E. 1766. Flora norvegica. Pars prior. Nidrosiae (Trondheim). HAticrimson, H. 1964. De rhodiolae. Flora. Timarit um islenzka grasafraedi 2:77-82. Hatvorsen, R. 1988. En del av kulturarven. Plantenavn i Vinje-dialekten. Blyttia 46:195-198. Hansen, L.|. and T. ScHmior. 1985. Major Peter Schnitlers grenseeksaminasjonsprotokoller 1742-1745.Vol. 3. Norsk historisk kjeldeskrift-institutt, Oslo. HAuKDAL, J. 1961.1 skreddartimen. Folkeminne fra Gauldalsbygdene. Norsk folkemi innelags Skr.87:1-169. HeizMann, W. 1993. alee der ee eee im Altwestnordischen. (Reallexikon der germanischen Altert Aalie gsbande, Band 7).Walter de Gruyter, Berlin - New York. Herzen, A. 1834.Ranens beskrivelse. Manuscript, printed as: Ranens Beskrivelse 1834.Rana museums- og historielag/Lofotboka. HertzeerG, E. 1912. Inga af Varteig og hendes eettlegg. Hist. Tidsskr. 5. Rk. 1:1-28. Haec, O.A. 1974. Planter og tradisjon. Floraen i levende tale og tradisjon i Norge 1925- 1973. Universitetsforlaget, Oslo—Bergen—Troms@. Hoimesog, J. 1953. Tidligere undersokelser over Bergenstraktens plantevekst. Utgitt ved av Knut Feegri. Bergens Hist. For. Skr.59:1- Hayaaard, A. 1941.Studies on the nutrition and physio-pathology of Eskimos, undertaken at Angmagssalik, East-Greenland 1936-1937. Norske Vidensk.-Akad. Oslo, Skr. Mat.- Naturv. KI. 1940(9):1-176. HucHtes, C.C. 1960. An Eskimo village in the modern world. Cornell University Press, Ithaca, New York. Huxkevserc, M. 1952. Helsestell og sjukdomar. Arsskr. Romsdal sogelag 1952:33-38. Hutten, E. 1958.The amphi-atlantic plants and their phytogeographical connections.Kung| Sv. Vetensk.-Akad. Handl.4. Series 7(1):1-340 Hutten, E. and M. Fries. 1986. Atlas of North ucanee vascular plants north of the Tropic of Cancer. Koeltz Scientific Books, KOnigstein. Jenssen, @. 1982. Leirfjord bygdebok. Szerbind |. Helgelandsmalet i Leirfjord. Sprak og sprakkultur der Vefsn og Rana moter yan. Leirfjord bygdeboknemnd. Kirkevolt, G. 1940. Plantor og plantenamn fra Valdres, serleg fra Vestre Slidre. Tidsskr. Valdres historielag 4(3):165-176. Krocu, J.A. 1813. Efterretninger om Provstiet Nordfjord i Bergens Stift i Norge. Topogr.- Stat. Saml. Kongel. Selsk. Norges Vel 2(1):1-291. 342 BRIT.ORG/SIDA 21(1) Kurkin, V.A. and G.G. ZapesocHnaya. 1986. The chemical composition and pharmacological properties of Rhodiola plants. Khim.-farm. Zhurn. 20:1231-1244, KurKIN, V.A., G.G. ZAPESOCHNAYA, aNd A.N. SHCHAVLINSKI. 1985. Terpenoids of Rhodiola rosea rhi- zomes. Khim. Prir. Soedin. 5:632-636. LAGERBERG, T., J. HoumBoe, and R. Norbxacen. 1955.Vare ville planter. Vol. 3. Johan Grundt Tanum, Oslo. Lane, J. 1960. Ordbog over Danmarks plantenavne. Vol. I]. Munksgaard, Kabenhavn Lantis, M. 1959. Folk medicine and hygiene. Anthropol. Papers Univ. Alaska 8:1—75. LeirFatt, J. 1968. Liv og lagnad i Stjordalsbygdene II. Stjardalsboka, vol.|, part Ill. Stjardal og Meraker kommuner. Lio, J. 1941. lvar Aasens herbarium. Nytt Mag. Naturvidensk. 81:57-80. Lippert, W. 1995. Familie Crassulaceae Dickblattgewachse. In: H.E. Weber, ed. Gustav Hedi. llustrierte flora von Mitteleuropa. Band IV. Teil 2A. Blackwell Wissenschafts-Verlag, Ber- lin. Pp. 69-129 Marzett, H. 1979. Worterbuch der deutschen Pflanzennamen. Vierter Band. Sabadilla— Zygophyllum. Anhang:Abies—Zygocactus. Schrifttum, Berichtigungen. S. Hirzel Verlag, Stuttgart—Franz Steiner Verlag, Wiesbaden. MetHeim, A. 1953.Om floraen pa hustak i Hornindal. Blyttia 11:33-61. Mo, E. 1925. Rindalske namn pa dyr og vokstrar. Aarskr. Nordmor historielag 1925:75-86. Moerman, D.E. 1998. Native American Stoner Timber press, Portland, Oregon. Mone, N. 1786. Forsa@g til en Islandsk N ie. Kj@benhavn Merkvep, B. 1996."Varknipa” i nordnorsk tradisjon, med hovedvekt pa forholdene i Troms og Lofoten/Vesteralen. Polarflokken 20:9-18. Nievsen, Y. 1873. Kristian IV's Reise til Finmarken 1599. Kristiania. Norpat, A. 1939.Uber einige norwegische volksmedizinische Skorbut-Pflanzen, und ihren Vitamin-C-Gehalt. Nytt Mag. Naturvidensk. 79:193-231, 81:117-142. Noroat, A. 1946. A pharmacognostical study of Sedum acre L. including a comparison with some other Crassulaceae. Johan Grundt Tanum, Oslo. Norbduacen, R. 1934. Taklauk og fjellkaur. Maal og minne 1934:101-128. NorbHacen, R. 1941. De gamle klosterhager. Foreningen til norske fortidsminnesmerkers bevaring. Arsberetn. 1939:55-94. Norouacen, R. 1947. Kappilaup, kobbeloppe og kappegullkare: norske dialektnavn pa lovetann. Mal og minne 1947:27-69. Norman, JM. 1894. Norges arktiske flora. |. Speciel plantetopografi. Part 1. Kristiania. @«senvat, K. 1977. Plantenamn fra Vefsn. Arbok for Helgeland 1977:94-102. Pautu, S. 1648. Flora Danica. Det er: Dansk urtebog. Pontoppipan, E. 1752. Det forste Fors@g paa Norges Naturlige Historie. Farste Deel. Kia@benhavn. Porsivo, A.E. 1953. Edible plants of the Arctic. Arctic 6:15-34. Qviastap, J. 1901. Lappiske Plantenavne. Nyt Mag. Naturvidensk. 39:303-326. Quiastan, J. 1932. Lappische Heilkunde. Inst. Sammenl. Kulturforskn., Serie B, Skr. 20:1—270. REICHBORN-KJENNERUD, |. 1922.Vare folkemedisinske laegeurter. Kristiania. ALM, ETHNOBOTANY OF RHODIOLA ROSEA 343 ReICHBORN-KUENNERUD, |. 1941.Var gamle trolldomsmedisin. lll. Vidensk.-Akad. Oslo. II. Hist-fil. KI. Skr. 1940 (1):1-221. Rise, O.J. 1947, Oppdalsboka. Historie og folkeminne. |. Johan Grundt Tanum, Oslo. Riste, O. 1916. Primstaven. Nyaars- og tids-rekning, primstav- og “ring’stav. Syn og segn 22:13-22. RoutorF, J. 2002. Volatiles from rhizomes of Rhodiola rosea L. Phytochemistry 59:655—-661. RONNING, O.|. 1959. Rosenrot. Ottar 21:22-24. Rarpam, H.F. 1873. Sivert Grubbes dagbog. Danske Mag. 4 Rk. 2:361-406. Ross, H. 1895. Norsk ordbog. Tillaeg til “Norsk Ordbog" af Ivar Aasen. Christiania. Ruse, H. 1 762.Forslag Om Plandterit gel til at lJaette Hur gers Ned Maanedlige Afhandlinger til en og anden Forbedring i Huusholdningen 1762:259-288. ScHUBELER, F.C. 1888. Viridarium norvegicum. Norges Vaextrige. Vol. 2. Universitetsprogram, Christiania. Soreibe, L. 1952. Systegras—Syftesoke. Folkeminne ved Samuel Roti. Arbok for Nordfjord —31. Spasov, A.A., G.K.Wikman, V.B. MANnorikoy, LA. Mironova and V.V. Neumoin. 2000. A double-blind, placebo-controlled pilot study of the stimulating and adaptogenic effect of Rhodiola rosea SHR-5 extract on the fatigue of students caused by stress during examination period with a repeated low-dose regimen. Phytomedicine 7:85-89. Sprincate, L. 1995. Rhodiola. In: J. Cullen, J.C.M. Alexander, A. Brady, C.D. Brickell, PS. Green, V.H. Heywood, P-M. Jorgensen, S.L. Jury, S.G.Knees, A.C. Leslie, V.A. Matthews, N.K.B.Robson, S.M. Walters, D.O. Wijnands, and PF. Yeo, eds. The European garden flora. Vol. 4. Cam- bridge University Press. Pp. 200—203. STEEN, A. 1961. Samenes folkemedisin. Samiske Saml. 5(2):1—-62. Strom, H. 1762. Physisk og Oeconomisk Beskrivelse over Fogderiet Sondmer, beliggende i Bergens Stift i Norge. Farste Part. Sorge. StROMPDAL, K. 1929. Gamalt fra Helgeland. Norsk Folkeminnelags Skr.19:1-171. VeGuspAL, E. 1979. Gamalt or Beiara. Folkeminne fra Beiarn Ill. Norsk Folkeminnelags Skr. 119:1-179. Vreim, H. 1943. Laftehus. Tomring og torvtekking. Ed. 3. Norske arkitekters landsforening, Oslo. Witte, H.J. 1786. Beskrivelse over Sillejords Praestegield i @vre-Tellemarken i Norge tilligemed et geographisk Chart over samme. Gyldendal, Kabenhavn YOSHIKAWA, M., H. SHIMADA, H. SHiMoDA, H. Matsupa, J. YAMAHARA, and N. Murakami 1995. Rhodiocyanoside-A and rhodiocyanoside B, new antiallergic cyanoglycosides from Chinese natural medicine si-li-hong-jing-tian, the underground part of Rhodiola quadrifida (Pall.) Fisch. et Mey. Chem. Pharm. Bull. 43(7):1245-1247. YosHikawa, M., H. SHiMADA, H. SHimopa, N. Murakami, J. YAMAHARA, and H.Matsupa 1996. Bioactive constituents of Chinese natural medicines. Il. Rhodiolae Radix (1):Chemical structures and antiallergic activity of rhodiocyanosides A and B from the underground part of Rhodiola quadrifida (Pall.) Fisch. et Mey. (Crassulaceae). Chem. Pharm. Bull. 44(11): 2086-2091. WS BRIT.ORG/SIDA 21(1) YosHIKAWwA, M., H. SHIMADA, S. Horikawa, T. Murakami, H. SHimopa, J. YAMAHARA,and H. Matsupa 1997. Bioactive constituents of Chinese natural medicines. lV. Rhodiolae Radix (2): On the histamine release inhibitors from the underground part of Rhodiola sacra (Prain ex Hamet) S.H. Fu (Crassulaceae). Chemical structures of rhodiocyanoside D and sacranosides A and B.Chem. Pharm. Bull. 45(9):1498-1503. NOTES ON “COFFER” FROM THE KENTUCKY COFFEETREE (GYMNOCLADUS DIOICUS, FABACEAE) John P. Spaeth and John W.Thieret Department of Biological Sciences orthern Nenwehy es versi ny Highland Heights, 9,U.S.A ABSTRACT The ceeeara ae seca of the Rennaehsy cotlecties (Gymnocladus dioicus, Fabaceae) were spar- ingly used in pionee ffee. Preparation of the beverage, opinions on its smell and taste, oun of the plant, and history and present-day status of this use of the seeds are discussed. RESUMEN Las semillas del café de Kentucky (G) ladus dioicus, Fabaceae) se usaron en los tiempos de los . Y a peace’ ] “2 « en | ] Pp prep ebida, of y sabor, toxicidad de la planta, historia y es | del uso de las semillas. INTRODUCTION The assertion that the roasted and ground seeds of the Kentucky coffeetree (Gymnocladus dioicus (L.) K. Koch; Fabaceae) (hereafter KCT) can be brewed into a coffee-like beverage and that the Kentucky and Tennessee pioneers made such a beverage from them is repeated frequently in American and even Euro- pean literature on New World edible wild plants (e.g., Allen & Allen 1981; Bean 1973; Browne 1857; Fernald & Kinsey 1958; Loudon 1838; Loz 1890; Nicholson 1883; Peterson 1978; Rogers 1905; Sargent 1889; Saunders 1920; Vannorsdall 1958; Wampler 2000). Most of the reports are quite definite concerning the pioneers: yes, the seeds were indeed so used by them. Some, though, are a bit uncertain, e.g., “Pioneers may have made a coffee substitute from [the] seeds” (Stacy 1992). Braun (1961) noted that the making of “coffee” from the seeds in Kentucky was in the Bluegrass region, where the tree is as common as or even more common than in most other places. The Kentucky coffeetree ranges from New York to South Dakota, south to Virginia, and Oklahoma (Little 1977 [range mapl; Spaeth n.d_). Its seeds (Fig. 1) are borne in large, woody, brown pods (legumes) (4.3)12-16.5(25?) cm long, 4-5 cm wide, and 1-2 cm thick. The pods remain on the tree throughout much of the winter; at a distance the tree appears to have a flock of birds among its leaf- less branches. Each pod containsa fleshy pulp in which are embedded 1-9 dark olive brown, hard seeds 1.1-2 cm long and wide and 0.6-1.3 cm thick. The seeds are rock hard—“adamantine,” as Rogers (1905), with but slight hyperbole, well SIDA 21(1): 345-356. 2004 346 BRIT.ORG/SIDA 21(1) Fic. 1.K ky coff (G ladus dioicus). Left, pods. Lower right, seeds (a U.S. penny included for size compari- son). Upper right, ground seeds, the “coffee.” described them. Later, Rogers (1917) wrote: “How the pioneers ever crushed [the seeds] is a puzzle to all who have tried to break one with a nutcracker.” Living in the Bluegrass region and having access to an ample supply of the seeds, we decided to experiment by preparing a potable drink from them. Next we investigated the matter of toxicity of KCT and the history of beverage use of the seeds. Results of our trials and studies are reported here. THE “COFFEE” We found only four recipes for roasting the seeds. One internet site suggested roasting a cup of seeds, one layer thick, in an oven for 30 minutes (Rolen 2001). No temperature was specified, and 30 minutes did not seem to us long enough for any toxins possibly present in the seeds to dissipate (see “Toxicity” below). Another source, which we followed, gave more details: Brill and Dean (1994) suggested a “safe” way to make KCT coffee: bake the seeds in a “covered roasting pan” at 300° F for 3 hours (similar instructions are in Brill [2002] and Phillips [1998], although Phillips suggested 350° F). On 5 Aug 2003, we put 30 seeds into each of four Pyrex jars labeled 1, 2, 3, and 4, respectively. One layer of seeds covered the bottom of each jar. The jars , NOTES ON “COFFEE COFFECIREE 347 were roasted at 150° C (=300° F). During the roasting, the room had at first an aroma of peanut butter. Jar | was removed after 2.0 hours, jar 2 alter 2) Hours, and jar 3 after 3.0 hours by which time four of the seeds had burst (“popped”). By the time jar 4 was removed (3.5 hours), the room had an aroma of something burning; 11 seeds had popped. The next step was to establish a method of breaking the seeds, which were still rock hard. Without a corn grinder available (and fear that if we did have one, the seeds would break the grinder), we used pliers and a mortar and pestle. The seeds were first cracked open one-by-one with the pliers over the mortar. As they were cracked, most shattered or exploded so the cracking was done under a cloth that covered the mortar. They were then coarse-ground with the pestle, pulverized (Fig. 1) in a coffee grinder, and placed back into their respec- paises wien were covered in plastic wrap to retain freshness. There appeared tobenor p between the ease of cracking the seeds and the roasting times. To make the ‘ coffee,” one heaping teaspoonful of ground seeds was placed in 2/4 of acup (<175 ml) of boiling water. (This probably would have been the method used by pioneers.) The grounds were stirred and allowed to settle to the bottom. Twenty people were granted the opportunity to smell and taste this historical drink; their responses are listed in Table 1. No one claimed to enjoy the taste or the experience. All agreed that an overpowering and persistent af- tertaste remained. When the “coffee” was sweetened with sugar, the brew became “more pal- atable.” The four participants who tasted it rated it as “acceptable,” “like sweet- ened tea,” “still rather unpleasant,” and “could get used to it.” The general con- sensus was that KCT “coffee” is not destined for supermarket shelves. A lot of work, with little pay-off, goes into roasting and grinding the seeds; this may be why some literature rates the brew as a poor substitute for coffee. The second author of the present paper drank about 1/3 cup of sweetened KCT “coffee” ev- ery day for 2 weeks, hoping that, with familiarity, the flavor might improve for him; it did not. t,in contrast, some individuals rate the beverage more highly. Brill (2002) called it “the world’s best caffeine-free coffee substitute”; his wife, too, likes the drink (Brill, pers. comm., Oct 2003). Phillips (1998) wrote that the beans make “a decent cup of caffeine-free coffee.” The brew has been described also as “pal- atable and wholesome” (Thwaites 1905). TOXICITY Anyone wishing to prepare “coffee” from KCT seeds will probably have some interest in reports of the plant's toxicity to various organisms. KCT has long Taste L. Opinions of the taste and smell of “coffee” brewed from ground, roasted seeds of Kentucky coffeetree in a 20-person taste/smell test, October/Novem- ber 2003, Northern Kentucky University. Identical opinions were expressed by 348 BRIT.ORG/SIDA 21(1) several participants. Taste Smell Akin to mud Earthy Horrible Chocolaty Fruity Stale coffee A little bit like coffee Hint of mocha Why bother? Chocolate coffee Like coffee with a touch of chocolate Low-quality coffee Like espre Like bakers chocolate Bitter Like a Like bargain coffee Three-day-old brewed coffee Not worth the effort Hint of sour Really bad Bad after taste Pungent Not at all like coffee been known to possess poisonous properties, both for livestock and humans, though it is by no means a toxic plant of major significance (Kingsbury 1964). The toxin appears to be water soluble; intoxication of livestock has been re- ported after the animals drank water into which KCT pods had fallen. An ex- tract from the leaves is said to poison flies (Bronaugh 1991; Chesnut 1898; Millspaugh 1887), a report that should be investigated. Toxins are found in all parts of the plant, especially fresh growth, and can cause “stomach and intesti- nal disorders with diarrhea, vomiting, irregular pulse, and coma” (Hardin & Arena 1974). For over a century, this poisoning has been attributed to the alka- loid cytisine (e.g, Chesnut 1898), but the presence of this compound has never been confirmed in KCT. Unusual amino acids and an alkaloid are reported from the seeds (Southon 1994), but these compounds are not likely to be of toxico- logic significance (Burrows & Tyrl 2001). One's desire to try making “coffee” from the seeds would probably be tem- pered by statements such as the following: “Only a few cases of human poison- ing have been reported from eating the seeds or using them to make a coffee substitute” (Stephens 1980). “... chewing one or two [seeds] would not be ex- pected to produce toxic effects” (Lampe & McCann 1985). “Eating the seeds” or “chewing one or two” would, of course, be most unwise because they are so hard; chewing them would be like chewing small rocks. oe though, that there is probably little to worry about with the “cof- “The toxin is heat labile” and “[the seeds] are not toxic when parched” (Bur- = »NUTES UN CUFFEE FrEctnee 349 rows & Tyrl 2001). Nevertheless we do not recommend the “coffee” until more is known about the poisonous compounds in the seeds. HISTORY Pioneer times The earliest appearance of the name Kentucky litalics ours] coffeetree—as op- posed to simply coffeetree—we have noted is in an April 1785 entry in one of the diaries of George Washington: “Planted ... eight nuts from a tree called the Kentucke [sic] coffee tree . ...” Jackson & Twohig 1978). According to Hill and Fountain (2003), the Kentucky coffeetree was promoted by early land developers who wanted to get settlers out to ae ae et (which included Kentucky at that time). Coffee, a popular beverage from coastal ports. Land developers atverticed BERENS ce asa place w here a tree grew with beans that could be roasted and | d tk bl the be verage WaS no substi tute for coffee, and the early settlers quickly dropped it as soon as the real thing became available We were unable to verify this account in spite of extensive searching of litera- ture and internet. The earliest link we have found between KCT seeds and the “coffee” is in John Filson’s The discovery, settlement and present state of Kentucke [sic] (Filson 1784): “The coffee-tree greatly resembles the black oak, grows large, and also bears a pod, in which is enclosed good coffee.” This statement, however, does not inspire confidence that the author actually partook of the “good coffee” “en- closed” in the pods or that he knew of the seeds being the basis for the drink. In Pioneer life in Kentucky 1785-1800 (Horine 1948) a letter is quoted from Kentucky pioneer Daniel Drake to his adopted daughter that is definite on the use of the seeds for a beverage: “| We gathered] those wild fruits which were so precious to us in the absence of the cultivated. Some of them were for immedi- ate use, or little thought of except by the children; others had a more perma- nent value, and were stored for winter . ... Among the latter, grapes, nuts, crab apples, and occasionally the hard seeds of the coffee tree, .. of which, by way of change from Bohea tea[Camellia sinensis), we made a substitute for coffee.” This report, even in the absence of any other, would seem to verify the pre-1800 use of a beverage made from the seeds of KCT. Bakeless (1965) wrote about the complaints of aman who, during the 1780's, was a guest at one of the “dreadful” hovels providing food, drink, and bed to Kentucky travellers: Nothing to eat but bear meat and ‘corn-meal dodgers.’ And nothing to drink but whisky ... and coffee ‘com as of an ar oes that grew some eight hundred or one thousand miles north of where coffee tree [Cof r did grow. This beverage was brewed from the pod [sic] of the Kentucky ‘cof- tee tree.” The nee fluid was described by a charitable geographer of the period as ‘not unlike coffee” Another traveler, still more charitable, described it as ‘a pod in which is good coffee-seed. Michaux (1817; French version 1812) wrote that “the name of coffee tree was 350 BRIT.ORG/SIDA 21(1) given to this vegetable by the early emigrants to Kentucky and Tennessee, who hoped to find in its seeds a substitute for coffee: but the small number of per- sons who made the experiment abandoned it, as soon as it became easy to ob- tain from the sea ports the Coffee of the West Indies” (see also Keeler 1900). This, too, has the ring of a reliable report. The extent of use of the seeds for a beverage recorded by Michaux differs much from that given by others. Saunders (1920) wrote that “a century agosuch use of them was quite prevalent in what was then the western wilderness, and travelers’ diaries of the time make frequent mention of the practice.” Medsger (1972) wrote that “the pioneers apparently made much use of the seeds of this tree, which were roasted and ground, then used as a substitute for coffee.” Ac- cording to Clute (1943), KCT “coffee” was “often” resorted to when the supply of arabica coffee was exhausted. We have made a wide search through the lit- erature—literally many hundreds of sources—and have found nothing that cor- roborates these statements. Michaux, we believe, was right in his assessment of the frequency of use. Itis perhaps indicative of the infrequency of use of KCT “coffee” that Porcher (1863), in his Civil War era Resources of the southern fields and forests, did not mention G. dioicus even though he listed various coffee substitutes. The tree’s range does extend into the far northern part of the area covered in the book (Little 1977; Spaeth n.d). Perhaps, too, arabica coffee, in spite of war-induced scarcity, was sufficiently available so that substitutes were generally not im- portant; Dick (1974), in his book on the “southern frontier,” mentioned that “b 1840 black coffee was coming into use as a beverage.” Loudon (1838) noted that the use of KCT seeds for “coffee” had long since been discontinued. Wilson (1905) noted that the “coffee” use had “long since ceased.” A 1902 paper on KCT did not mention the “coffee” even though it considered “economic uses” (Anonymous 1902). Gibson (1913), describing the “coffee” as black and bitter and noting that “a little of it would go a long way with a modern coffee drinker,” wrote that “when the Kentuckians were able to procure coffee they let the wild substitute alone.” Kentucky and Tennessee were not the only places where this beverage was tried. Thomas Nuttall (1821) apparently had drunk the “coffee” in early 1819 he wrote in the record of his journey into the Arkansas Territory,“Among the trees, we still continue to observe the coffee-bean (Gymnocladus canadensis), now loaded with legumes, the seeds of which, when parched ... produce a substi- tute for coffee greatly inferior to the Cichorium [chicory]” At that writing, he was near the confluence of the Ohio and Mississippi rivers. And in southwest- ern lowa during the winter of 1819-1820, KCT “coffee” was drunk by members of S.H. Long’s expedition to the Rocky Mountains (Thwaites 1905). Seeds of KCT were not the only coffee substitute used in early Kentucky. SPAETH , NOTES ON “COFFEE OFFEETREE 351 Another report on the pioneers (Goode 1989), from an area of the state in which KCT is lacking or exceedingly rare (Little 1977), stated that ground parched corn sweetened with honey made a “reasonably tasty substitute for coffee.” Other substitutes included parched “tubers,” rye, wheat, acorns, beans, chestnuts, chicory, chinquapins, cotton, grapes, peanuts, English peas, persimmons, okra, sorghum, sugar cane, “Irish” and sweet potatoes, and dandelion—and even, in desperation, wood shavings (Anonymous n.d.; Howard 1975; March 2000; Mitch- ell 1991). Writing about pioneer Kentucky, Cotterill (1917) mentioned that “Tea and coffee were reserved for the sick and were considered as a mark of effeminacy if taken by people in good health.” Was this, in addition to the scarcity of arabica coffee in those early times, maybe one reason to seek a coffee substitute? One might wonder where the pioneers got the idea of preparing a beverage from these seeds, which seem to us to be a most unlikely source. The seeds cer- tainly do not look like the “beans” of arabica coffee, and the task of preparing them for brewing is daunting. The pioneers might have learned the process from Native Americans; Smith (1928) reported that the Meskwaki Indians cooked the seeds and then ground them and boiled them to make “coffee.” These Indi- ans, however, were in Wisconsin, not Kentucky. But perhaps some Kentucky tribes knew of the use and shared the information with the pioneers. Indians of the Missouri River region (Gilmore 1991; Moerman 1998) knew of KCT but ap- parently did not make a beverage from the seeds although they roasted and ate the seeds and prepared a “food” from the pounded seeds. According to Phillips (1998), “the roasted beans are a reasonably good nut, especially when salted or soaked in salted water and re-baked.” We infer that the roasting time, though not stated, is about !/2 to | hour. It is also possible that some pioneers familiar with arabica coffee but suf- fering coffee deprivation simply tried to make a beverage from KCT seeds, pro- ducing a brew that they liked and about which they spread the word. KCT “coffee” during the Civil War During the early days of November 2003 we were told by a Civil War history buff (who wishes to remain anonymous) that he had heard from a Civil War re-enactor that, during that war, quite limited use was made of KCT seeds as “coffee.” The taste of the “coffee” was reportedly so bad that it made hardtack— those plain flour-and-water biscuits that were often moldy or infested with maggots and weevils (Billings 1887)—seem good by comparison. In her Civil War plants and herbs, Mitchell (1996) noted that the seeds are the basis for a “coffee,” a report apparently based on Anonymous (1995). But she gave no indi- cation that they were actually used for that purpose during the war although, of course, they could well have been. Arabica coffee was apparently a favorite non-alcoholic beverage of soldiers (Show delicious the aroma of it, and how 352 BRIT.ORG/SIDA 21(1) readily each man disposed of a quart” [McCarthy 1882]). Although arabica cof- fee was available at least some of the time (Locke 1870), its supply in the south was sometimes uncertain because of the vicissitudes of war on land and the Lincoln-ordered blockade of southern ports from Virginia to Texas. The George Rogers Clark connection In going through hundreds of KCT websites we came upon one from Storm Lake, lowa,a city that has a “tree museum” in which various “historic” trees are planted (Anonymous 2003b), among them a “George Rogers Clark Kentucky Coffee tree.” The text concerning the tree is as follows: “During the Civil War, George Rogers Clark attempted to develop a coffee bean tree to replace regular coffee which was unavailable during that time. However, since the coffee was not pleasant to the taste, the project failed.” Is this statement credible? Certainly not in the timing of Clark’s posthumous interest; he died in 1818. One wonders, too, how one could “develop” a tree during the short time span of the Civil War. But we decided to investigate further. We directed a query to Storm Lake, hoping to learn the source of their KCT. We were given the name of an individual in Louisville; later checking showed that he and another person were involved in a heritage tree program and that one of them is deceased. Two letters to the first man have gone unanswered; we suspect that he, too, has died. We learned that Clark did send KCT seeds to Thomas Jefferson at Monti- cello. Indeed, in the Kentucky legislation (Anonymous 2003a) that established G. dioicus as the state tree of Kentucky (S.B. 150, approved 8 Mar 1976; later re- scinded), one of the “whereas’s” is this: “WHEREAS, in 1783 General George Rogers Clark did send seeds of the Kentucky coffeetree to Thomas Jefferson at Monticello and which seeds produces [sic] trees still gracing the grounds of Monticello.” In a letter dated 4 Dec 1783 Jefferson wrote to Clark: “We received here about a week ago your obliging letter of Oct. 12. 1783, with the shells and seeds for which we return you many thanks” Jefferson 2003). One can but as- sume that among the included seeds were those of KCT. (Asan aside, this letter is the one in which Jefferson asked Clark if he would like to lead [an explora- tion party] into the country from the Mississippi to California.) We found a brief quote said to be from Clark’s October 1783 letter to Jeffer- son: speaking of KCT, Clark wrote “It makes beautiful shade and we think it will flourish with you” (Anonymous 2003a). Did Clark also mention in the let- ter the “coffee” use of the seeds of the tree? Inquiries to the Jefferson collection at the Princeton University Library (Linda Monaco, pers. comm., 16 Oct 2003) and to the Monticello Library (Bryan Craig, pers. comm., 16 Oct 2003) elicited the response that the letter is apparently lost. Finally, we were referred to a person said to be knowledgeable about GRC’s , NOTES ON “COFFEE 353 correspondence: Julia Parke, former director at Locust Grove, Clark’s Louisville home. In an October 2003 telephone conversation with her, the second author was told that she had known of the 12 Oct 1783 letter and that yes, GRC did mention in the letter not only the horticultural possibilities of the tree but also the use of Kentucky coffeetree seeds for “coffee.” Thus the Storm Lake report of Clark’s interest in KCT had been somewhat exonerated. CURRENT USE OF KCT “COFFEE” In anarticle on various wild-growing substitutes for arabica coffee, Serlin (1977) wrote, “All last winter, along the Ramble in New York City’s Central Park, I’ve seen people pinching coffee costs by scuffing through fallen leaves for precious pods under a stand of Kentucky Coffee Trees.” In October 2003, wondering about the frequency of such foraging by New Yorkers, we telephoned Steven Brill, well- known author and educator on the subject of wild edible plants. He told us that, in his years of visiting Central Park, the only foragers he had seen were those accompanying him on his field trips. According to some of these foragers, the seeds apparently can be used asa chocolaty seasoning in cakes and cookies (Brill & Dean 1994). One seed added to carob-flavored ice cream can make the product “taste more like chocolate than ever” (Brill 2002). We checked with several more people knowledgeable about the subject of edible wild plants. None knew of anyone who had prepared a drink from KCT seeds. Other than the reports of Brill (2002), Brill @ Dean (1994), Phillips (1998), and Serlin (1977) and the recent brewing of “coffee” from KCT seeds here at Northern Kentucky University, we have located no evidence of significant ex- tant use of the beverage. Even if current edible-plant books mention the use, we suggest that KCT “coffee” is mostly a curiosity from a time now past, being in- dulged in only by an occasional one of those individuals who eat their way through the landscape (often with tasty rewards). It poses no serious threat to Maxwell House or Starbucks. CONCLUSION Limited use of KCT seeds asa coffee substitute was made in pioneer Kentucky and elsewhere. It would appear that the many post-pioneer reports of a KCT- seed-based beverage derive ultimately froma few early accounts; Drake's, Long's, Michaux’s, Nuttall’s, and possibly Clark’s. The Kentucky coffeetree is of essen- tially no present-day concern as a beverage plant. ACKNOWLEDGMENTS We thank the following for help: David M. Brandenburg, The Dawes Arbore- tum, Newark, OH; Steve “Wildman” Brill, Mamaronech, NY; Kenneth C. Carstens, 354 BRIT.ORG/SIDA 21(1) Murray State University, Murray, KY; Bryan Craig, Library, Monticello, Charlottesville, VA; James Duvall, Boone County (Kentucky) Public Library: Jennifer Gregory, Archives, Steely Library, Northern Kentucky University, High- land Heights, KY; Miriam Steinitz Kannan, Biological Sciences, Northern Ken- tucky University, Highland Heights, KY, The Lloyd Library, Cincinnati, OH; Patricia B. Mitchell, Chatham, VA; Linda Monaco, The Papers of Thomas Jeffer- son, Princeton University Library, Princeton, NJ; Julia Parke, Louisville, KY: Maggie Whitson, Northern Kentucky University, Highland Heights, KY, Den- nis Worthen, Lloyd Library, Cincinnati, OH; and the 20 brave individuals who took part in our taste/smell tests. REFERENCES Atven, O.N. and E.K. ALLEN. 1981. The Leguminosae. A source book of characteristics, uses, and nodulation. University of Wisconsin Press, Madison. Anonymous. n.d. The CWi Civil War cookbook. Http://www.civilwarinteractive.com/ cookbooknonalcoholic.htm. Accessed 10 Sep 2003. Anonymous. 1902. Coffeetree (Gymnocladus dioicus).U.S.D.A. Forest Serv. Circ. 91. Anonymous. 1995. Coffee substitutes. Herb Quart. 65 (spring):10. ANONYMOUS. 2003a.5.B. 150.Kentucky’s state tree. http://www.kdla.net/statelib/KYtree.htm. Accessed 10 Sep 2003. Anonymous. 2003b. Storm Lake [lowa] Tree Museum. http://www.stormlake.org/city/pages/ treemuseum.htm. Accessed 15 Sep 2003. Baxe.ess, J. 1965. Daniel Boone. Stackpole Company, Harrisburg, PA. Bean, WJ. 1973. Trees and shrubs hardy in the British Isles. Volume 2:D-M. John Murray, London. Bituincs, J.D. 1887.Hardtack and coffee or the unwritten story of army life.George M.Smith & Company, Boston Braun, E.L. 1961.Woody plants of Ohio. Ohio State University Press, Columbus. Bri, S.and E. Dean. 1994. Identifying and harvesting edible and medicinal plants in wild (and not so wild) places. Hearst Books, New York. Britt, S. 2002. The wild vegetarian cookbook. Harvard Common Press, Boston. BronaucH, W. 1991. The biggest coffeetree. Amer. Forests (May/June):35. Browne, D.J.1857.The trees of America; native and foreign. Harper and Brothers, New York. Buurrows, G.E. and RJ. Tyr. 2001. Toxic plants of North America. lowa State Univ. Press, Ames. CHesNuT, V.K. 1898. Principal poisonous plants of the United States.U.S.D.A. Div. Bot. Bull. 20 Ciute, W.N. 1943. Coffee from Kentucky. Amer. Bot. 49:52-53. CoTTerILt, R.S. 1917. History of Kentucky. Johnson and Hardin, Cincinnati. Dick, E. 1974. The Dixie frontier: a social history of the southern frontier from the first transmontane beginnings to the Civil War. Octagon Press, New York. FeRNALO, M.L.and A.C. Kinsey. 1958. Edible wild plants of eastern North America. Revised by R.C. Rollins. Harper and row, New York. »NUTES ON COFFEE Fiison, J. 1784. The discovery, settlement and present state of Kentucke. Printed by James Adams, Wilmington, DE Gisson, H. 1913. American forest trees. Hardwood Record, Chicago. Gimore, M.R. 1991. Uses of plants by the Indians of the Missouri River region. Univ. Ne- braska Press, Lincoln. [Work first published in 19191] Goooe, C.E. 1989. Southern Kentuckians. Historical sketches of Barren and surrounding counties in Kentucky. Published by the author, Glasgow, KY. Haroin, J.W. and J.M. Arena. 1974. Human poisoning from native and cultivated plants. Duke Univ. Press, Durham, NC. Hiv, D.B. and W.M. Fountain. 2003. Kentucky coffeetree: the state tree of Kentucky. http:// www.ca.uky.edu/agc/pubs/for/for8/for8.htm. Accessed 1 Dec 2003 Howard, M. 1975. Recollections of a Maryland confederate soldier and staff officer under Johnston, Jackson and Lee. Morningside Bookshop, Dayton, OH. [Work first published in 1914] Horine, E.F. 1948. Pioneer life in Kentucky 1785-1800. Henry Schuman, New York. Jackson, D. and D. Twouic. 1978. The diaries of George Washington. Volume 4. http:// memory.loc.gov/cgi-bin/query/r?ammem/mgw:@field[DOCID+@lit[wd0414]]. Ac- cessed 1 Dec 2003. Jerrerson, T. 2003. The letters of Thomas silt The Avalon Project at Yale Law School. http:// le.edu/lawweb/avalon/jefflett/let21.htm. Accessed 20 Sep 2003. Keever, HL. 1 900. Our native trees. Charles ceibners Sons, New York. Kinaspury, JM. 1964. Poisonous plants of the United States and Canada. Prentice-Hall, Englewood Cliffs, NJ. Lampe, K.F. and M.A. McCann. 1985. AMA handbook of poisonous and injurious plants. American Medical Association, Chicago. Lirtte, E.L., Jr. 1977. Atlas of United States trees. U.S.D.A. Misc. Publ. 1342. Locke, EW. 1870.Three years in camp and hospital. Geo. D. Russell and Company, Boston Loupon, J.C. 1838. Arboretum et fruticetum britannicum.Volume 2. Printed for the author, London. Loz, MJ. 1890. Le gymnoclade du Canada. Rev. Sci. Nat. Appl. 37:487-488. Marcu, W.C. 2000. Food and rations in the Civil War. http://users.aol.com/cinticwrt/ food.html. Accessed 10 Sep 2003. McCartuy, C. 1882. Detailed minutiae of soldier life in the army of northern Virginia 1861- 1865. Carlton McCarthy and Company, Richmond, VA. Meoscer, O.P. 1972. Edible wild plants. Collier Books, a Division of Macmillan Publishing Company, New York. Michaux, F.A. 1812. Histoire des arbres forestiers de l'Amérique septentrionale. Tome II. L. Haussmann, Paris. Michaux, FA. 1817. The North American sylva, or a description of the forest trees of the United States, Canada and Nova Scotia. Thomas Dobson-Solomon Conrad, Philadelphia. Mittspaucn, C.F. 1887. American medicinal plants. Boericke and Tafel, New York. 356 BRIT.ORG/SIDA 21(1) MitcHett, PB.1991.Desert Storm recalls Civil War food woes. Http://www.foodhistory.com/ foodnotes/leftovers/dstorm.htm. Accessed 10 Sep 2003. MitcHett, P.B. 1996. Civil War plants and herbs. PB. Mitchell, Chatham, VA. Moerman, D.E. 1998. Native American ethnobotany. Timber Press, Portland, OR. NicHoLson, G. 1883. The Kentucky coffee tree. Garden 24:29-30, Nuttatt, T. 1821.A journal of travels into the Arkansa [sic] Territory during the year 1819. T.M. Palmer, Philadelphia. Peterson, L. 1978.A field guide to edible wild plants. Houghton Mifflin Company, Boston Prius, J. 1998.Wild edibles of Missouri. Missouri Department of Conservation, Lefferson City]. PorcHer, F.P. 1863. Resources of the southern fields and forests. West and Johnston, Rich- mond, VA Rocers, J.E. 1905. The tree book. Doubleday, Page & Company, New York. Rocers, J.E.1917. Trees worth knowing. Doubleday, Page and Company, Garden City, NY. Roten, M. 2001. Kentucky Coffeetree Gymnocladus dioicus www.schools.1th5.k12.il.us/ bths-e/2001/kentuckycoffee.html. Accessed 8 Aug 2003. SARGENT, C.S. 1889.The Kentucky coffee-tree. Garden & Forest 11:75-76. Saunoers, C.F. 1920. Useful wild plants of the United States and Canada. Robert M.McBride and Company, New York. Serin, K. 1977. My old Kentucky home-made coffee tree. Flower and Garden 21(10):14- 15,42, 47. Smith, H.H. 1928. Ethnobotany of the Meskwaki Indians. Bull. Milwaukee Public Mus.4:175- 326, plates 37-46. SouTHON, |W. 1994. Phytochemical dictionary of the Leguminosae. Volume 1. Plants and their constituents. Chapman and Hall, London. SpaetH, J.n.d. Gymnocladus. Flora of North America. |n preparation. Stacy, H.P.1992. Kentucky Coffeetree. Page 493 in J.E. Kleber (editor in chief). The Kentucky encyclopedia. Univ. Press of Kentucky, Lexington STEPHENS, H.A. 1980. Poisonous plants of the central United States. Univ. Press of Kansas, Lawrence. THwates, R.G. (ed.). 1905. Account of an expedition from Pittsburgh to the Rocky Moun- tains performed in the years 1819, 1820 ... under the command of Maj. S.H. Long. 4 vols. Arthur H. Clark Company, Cleveland. (1:265.] VANNORSDALL, H.H. 1958. Trees of Ohio. Privately published, Wilmington, OH. WaMpLer, F. 2000. Trees of Indiana. Indiana University Press, Bloomington. Witson, E.H. 1905. Gymnocladus canadensis—the American coffee-tree. Fl. & Sylva 3: 313-315. REFLECTIONS ON THE TAXONOMY AND DISTRIBUTION OF MEDICINAL FLOWERS OF PAKISTAN Mohammad Athar'! M. Akmal Siddiqi! California Department of Food and Agriculture Clinical Research Center 2014 Capitol Avenue Marshfield Clinic Research Foundation Sacramento, CA 95814, U.S.A. 1000 North Oak Avenue atariq@cdfa.ca.gov Marshfield, WI 54449, U.S.A. siddiqi. muhammad@merf.mfldclin.edu ABSTRACT A literature search was conducted to determine the medicinal flowering plants found in Pakistan 1] ] enc, : \; : : vue paren ee df diseases. TI | we period of these plants are described to facilitate their collection at blooming. There are 95 spe- s belonging to 85 genera and 43 plant families. Most of the species belong | to Fabaceae and vaceae (7 species), Lé ae (6 species) peace (17 and 13 species respectively), followed by o and Boraginaceae (4 species). Apocynaceae and Rosaceae contained 3 species each. It is important for the sustainability of Pakistani medicinal plant industry to consolidate the plant-resource data base and hae ore the national and international markets for its medicinal flora. It is imperative that ti oe a be se pai and that priority species of the medicinal plants be salvaged. Pakistan also | research efforts into cultivation, production and conserva- tion of ume pene Institutional pas are urgently needed between all the Pakistani re- earch p g g with medicinal and aromatic plants. RESUMEN S lizé haustiva busqueda bibli Afi de | i licinales de Paki utilizadas como ee contra euleamedstes de diversa nacuraleza Se ae una lista de las ave stribucion y periodo } renciaasu posicion taxonomica. Se in j fnaléeics para facilitar su re 5n en el to optimo de la floracion El listado comprende FET alee ol arer 5 : : oO ~ r L a las familias Fabaceae y Asteraceae (17 y 13 especies respectivamente) Seda de Melvaceas 7 especies), Lamiaceae (5 especies) y Boraginaceae (4 especies); MRecyuaceas ) y Rosaceae presentan 3 especies cada una. ae ss para la industria sostenible pakistani de plantas medicinales cada ee isponibles y enploret los meICados Baton e internacionales que Sims | a ee ] y preserval L la prioridad de las plantas medicinales. Pakistan aedeaie también extender los esfuerzos de inv estigaci6n « a m0 piobiemes sobre en, aaa ’ conservacion de plantas medicinales. \] y un sistema de transferencia institucional [ lai 10 y l i i de desat rollo que intervienen en los procesos industriales de plantas medicals y aromaticas en Pakistan. 'Disclaimer: The views expressed in this article are those of authors and do not necessarily represent those of their respective departments. The publication of this article does not constitute a recommendation or en- dorsement of the use of these flowers as herbal medicine. The authors or their departments make no warranty, expressed or implied, and assume no legal liability for the use of these flowers for medical purposes. SIDA 21(1): 357 — 368. 2004 358 BRIT.ORG/SIDA 21(1) INTRODUCTION The flora of Pakistan is very rich due to the diverse climatic and soil conditions in its different ecological regions. The country has around 6000 species of wild plants; about 400-600 are considered to be of medicinal importance. Pakistan has considerable significance from the point of view of medicinal plants (Nasir et al. 1972) and has been correctly called as the floral emporium of medicinal plants (Rizvi 1998). An estimated 80% of the rural population in Pakistan de- pends on traditional medicines for their primary healthcare needs, the major- ity of that are plants or their active principles (Shinwari &@ Khan 1998). The sustainable harvesting of medicinal plants has great economic poten- tial. Although some medicinal plants, such as Carthamus tinctorius, Crocus sativus and Passiflora edulis, are locally grown but require knowledge about the proper collection and preservation of these important medicinal plants (Rizvi 1998; William @ Ahmad 1999). However, medicinal plants are used both in the indigenous medicinal system and by pharmaceutical industries (Shaheen et al. 2003). Many different flowering species, such as Artemisia absinthium, (Gul-e- afsantee), Matricaria chamomilla (Gul-e-babuna), Viola odorata (Gul-e- banafsha), Rosa damascena (Gul-e-surkh), and Crocus sativus (Gul-e-zafran), are known as potential sources for curing various liver diseases (Bach 1994; Bisset 1994; Duke 1986, Khan et al. 1996). Safflower (Carthamus tinctorius) has been used for centuries in Europe and Asia as a laxative and diuretic (Keville 1995). Passiont lower (Passiflora edulis) is reputed to have anti-spasmodic and seda- tive properties and used widely as an ingredient in herbal remedies (Reynold et al. 1994). The aqueous extract prepared from the calyx of Hibiscus sabdariffa is reported to possess cathartic activity (Haruna 1997). Classification of medicinal plants is organized in different ways depend- ing on the criteria used. In general, medicinal plants are arranged according to their active principles in their storage organs of plants, particularly roots, leaves, flowers, seeds and other parts of plant (Shaheen et al. 2003). These principles are valuable to mankind in the treatment of diseases (Shaheen et al. 2003; Shinwari & Khan 1998). According to the literature there are many Pakistani medicinal plants (Bach 1994; Nasir @ Ali 1972; Rizvi 1998; Shinwari &@ Khan 1998) that could be used for remedies, medicinal preparations and future utili- zation (Bach 1994; Bugti 1998; Mahmood et al. 1996; Nasir & Rafiq 1995; Shinwari & Khan 1998; Zaman & Khan 1970). This paper presents the taxonomy of Paki- stani medicinal flowering plants, their regional distribution, and flowering periods. ~— — MATERIALS AND METHODS A literature search was conducted to find out which Pakistani plants are used as remedies for various disorders or diseases (Bach 1994; Bugti 1998; Duke 1986; ATHAR AND SIDDIQI, MEDICINAL PLANTS OF PAKISTAN 359 Mahmood et al. 1996; Malik & Farooq 1984; Nasir & Ali 1972; Nasir & Rafiq 1995; Rizvi 1998: Shaheen et al. 2003; Shinwari & Khan 1998: Zaman & Khan 1970). The taxonomic position of species with medicinal properties was deter- mined. Their distribution and flowering period was also noted to facilitate their collection at blooming. The plants included herbs, shrubs, vines and trees. The genera were arranged alphabetically within each family. The nomenclature and classification followed Nasir and Ali (1972), and author citations followed Brummitt and Powell (1992). RESULTS AND DISCUSSION Ninety-five Pakistani species distributed among 85 genera and 43 plant families were found to have medicinal properties (Table 1). The largest numbers of species were found in Fabaceae and Asteraceae (17 and 13 species respectively), followed by Malvaceae (7 species), Lamiaceae (5 species) and Boraginaceae (4 species). Apocynaceae and Rosaceae each contained 3 species of medicinal plants. Other families contained only one or two species of medicinal plants (Table 1). Their flowering period and distribution were also worked out and presented in Table 1. The medicinal plants that are commercially exploited in large quantities occur mainly in four ecological regions of Pakistan: alpine and high altitude; temperate mountain forests; sub-tropical foothill forests; and arid and semi-arid scrubs (Ali & Qaiser 1986; Shaheen et al. 2003; William & Ahmad 1999). A number of medicinally important Pakistani plants are found in the moist alpine and high altitude areas, especially in the northwestern valleys (Nasir & Ali 1972; Shaheen et al. 2003; William & Ahmad 1999). Most plants of these areas are Slow growing perennial species, which require several years of vegeta- tive growth before reproduction by seed (Shaheen et al. 2003). Some of the spe- cies are classified as threatened or vulnerable if the current rate of collection continues. Endangered plant species of this area includes Podophyllum hexandrum, Saussaurea costus, Picrorrhiza kurrooa, Aconitum heterphyllum, and Corydalis sp. (Shaheen et al. 2003). Care should be taken in the conserva- tion of plants with medicinal flowers so that their existence is not threatened. The origin of Unani (Greek) medicines goes back to the material medica of numerous early civilizations. Unani medicine traces its origin to Greek medi- cine, which was adopted by the Arabs and thereafter spread to both Europe and Indo-Pak subcontinent (William @ Ahmad 1999). There are about 27 large, Pakistani herbal-manufacturing companies, which produce Unani medicines on a commercial scale (Shaheen et al. 2003). The number of herbal-medicine manufacturers in the non-organized sector runs into the hundreds. The annual gross income of some large herbal manufacturers is comparable to that of mul- tinational companies in Pakistan. About 50,000 traditional healers, including homeopaths, are serving about 60% of the population, aaa those living in rural areas (Shaheen et al. 2003; William @ Ahmad 19 360 Taste 1. The taxonomy, distribution, and flowering period of some BRIT.ORG/SIDA 21(1) of the medicinal flowers of Pakistan. Species Distribution Medicinal Remedies MONOCOTYLEDONS Poaceae OQ mbopogon ivarancusa Found in Karachi, Detoxifier, astringent and (Jones) Schult. Zea mays L. DICOTYLEDONS Acanthaceae J ‘ NI AVGHIOUCUGU VOOICU INCU S — hot dal Amaranthaceae Achyranthes aspera L. = osia cristata L.= Celosia argentea L. var. cristata (L.) Kuntze Ce Anacardiaceae Mangifera indica L. Apocynaceae Catharanthus roseus (L.) G.Don. P g | 4 +t Jacq. F f] . D, f rgularia daemia (Forssk.) Chiov. var. daemia Vinca major L. Asclepiadaceae Calotropis procera (Aiton) W.T. Aiton Asteraceae Achillea millefolium L. Artemisia absinthium L. Multan, Chitral, NWFP Quetta, Gilgit Cultivated in Sindh, Punjab, NWFP Planted in Karachi, Sindh Found in Gilgit, Karachi, Punjab, Baluchistan Grown in gardens Grown in Punjab and Sindh Cultivated and naturalized in the tropics Found in Karachi, Sindh, Lasbella, Peshawar, Rawalpindi Found in Parochial, Abbottabad, More hills Widely distributed in eserts throughout Pakistan Occurs in Gilgit, Swat, Murree, Poanch, Baluchistan, Chagia Grows in Thandiani tonic Astringent, chloretic diuretic and remedy for urinary infection Asthma, bronchitis, gonorrhea, highgrade fever and conjunctivitis Anti-hemorrhoidal Astringent, anti-diarrheal and for irregularity o menstrual cycle Astringent, urinary infection, catarrh, anti-diarrheal, anti- dysentery, veneral diseases Asthma, anti-leukemia, eyes salve and flatulence Anthelmintic,emetic expectorant Fresh flowers are purgative Asthma, Catarrh, cold cough, cholera, and for dyspepsia Hypotensive, haemostatic to arrest bleeding Anthelmintic, anti-scorpion enom and anti-snake venom ATHAR AND SIDDIQI, MEDICINAL PLANTS OF PAKISTAN Taste 1.continued Species Flowering Distribution Medicinal Remedies eriod Artemisia maritima L. Aug-Sep Found in Astor, Dyspepsia, tonic and anti- Baluchistan, Chitral, Swat — helminthic Calendula off lis L. Dec-Apr Cultivated in many parks © Duodenal-gastric ulcers, and gardens of Pakistan ypotensive, emmenag- ogue and cures skin diseases Carthamus tinctorius L. Apr-Jul Distributed NWFP, Emmenagoqgue, laxative, ae histan, (Harnai), sedative, stimulant jab Chrysanthemum Mar-—Jul Shee in Peshawar, ae oo cinerariifolium (Trevir.) Abbottabad and dys Vis.=Tanacetum cinerariifolium (Trevir.) Sch. Bip. Helianthus annuus L. Jul-Sep Widely cultivated in Anti-diarrheal, anti- akistan inflammatory carminative, diuretic Matricaria ct Mal= Jul-Jan Found in plains of Punjab, Analgesic, antiseptic Matricaria recutita L. Pishin carminative, anti- convulsant, diuretic, liver diseases, dyspepsia Silybum marianum (L.) Mar-Apr Found in Lahore,Peshawar, Flower heads are consumed Gaertn. Saidu Sharif,Abbottabad, — for diabetes control Mirpur, Rawalpindi Tagetes erecta L. Jun-Nov Grown in gardens of Anti-dote against wasp Pakistan stings, cure for eczema, juretic Tanacetum gracile Hook.f. Jun-Aug Found in Hunza, Anti-helminthic & Thomson Baluchistan araxacum officina Feb—Apr Widely distributed Aperient, diuretic, stimulant F.H.Wigg. Group ae Baluchistan — stomachic, tonic, detoxicant Xanthium strumarium Ll. — Jul-Aug Found in Karachi, lowers useful for tooth- Gilgit, Bs Baluchistan, ache Swat, NWFP, Hazara Balsaminaceae Impatiens balsamina L. Jul-Oct Cultivated in Karachi, Antibiotic activity, inter- Chitral, Murree costal neuralgia and useful in lumbago Bignoniaceae Millingtonia hortensis L.f.. =Nov—-Mar Cultivated in Sindh,Punjab Cures a Stereospermum suaveolens May-Jun D tereospermum colais (Buch.-Ham. ex Dillwyn) Mabb. Occurs in Rawalpindi District sthm Aphrodisiac, ibecual 362 Taste 1. continued BRIT.ORG/SIDA 21(1) Distribution Medicinal Remedies Species Flowering Bombacaceae Bombax ceiba L. Dec—Mar Boraginaceae Arnebia benthamii (Wall. = Oct-Nov ex G, Don) I.M.Johnst Borago officinalis L Jan-Feb Onosma hispidum Wall. Mar—Jul & G.Don Trichodesma indicum (L.) = Aug-Oct Sm. Brassic ae need ae irik, Mar—May Erysi imum cheiri (L.) aaa BR permur acerifolium Dec-Jul (L.) Willd annabace Humutlus lupulus L. Jul-Aug Caprifoliaceae Sambucus nigra L. Mar-Apr ucurbita ee a ica Roxb. Jun—Oct Fabaceae (Caesalpinioideae) S Bauhinia purpurea L. ep-Nov Bauhinia variegata L Feb Apr Caesalpinia pulcherrima Apr-Sep (L.) Sw. Oct-Dec Cassia alata L.= Senna alata (L.) Roxb. Cultivated as roadside and garden plant in Pakistan Found in Makran, Kaghan, Poonch Reproduced from seeds at Karachi Found in Pishin, Common in Landikotal, Swat, Chitral, Kaghan Occurs in Mangopir Cultivated in gardens Cultivated in Islamabad, Peshawar as an introduced tree Found in Pangi on the upper Chenab Occurs in Parachinar, Nathiagali, Hazara Found in Punjab, Ravi, tlie Doab, Rawalpindi Distr Cultivated in Punjab, WFP. Rawalpind Cultivated in Pakistan Cultivated in gardens of Pakistan Zz Sometimes cultivated in Pakista Diuretic and laxative Angina, fever, pharyngitis Anti-cancer agent (breast or face), corns, sclerosis and tumors Cardiac tonic, stimulant Flowers used as emullient and diuretic Cardiac disorders, emme- nagogue remedy for impotence and paralysis Dehydration, otalgia haematuria, massage Anti-septic, female inflo- scence used as diuretic, emmenagogue, dyspepsia Laxative, anti-pruritic and stimulant of blood circulation Lowers total cholestrol and blood sugar Flowers are used as purgative Flowers are aperient Asthma, bronchitis anti-pyretic, expectorant, anti-malarial salve Useful in skin exture ATHAR AND SIDDIQI, MEDICINAL PLANTS OF PAKISTAN Taste 1. continued 363 Species Flowering Distribution Medicinal Remedies Cassia fistula L. Apr-May Naturalized throughout = Cough, diphtheria, laxative Pakistan. Cultivated in Karachi, Cassia siamea Lamk. = Oct-Dec Cultivated in Karachi, Anthelmintic, anti-hyper- Senna siamea Sindh tensive, asthma, dandruff (Lam.) H.S. Irwin & insomnia, laxative, tranquil- Barneby izer, sedative Delonix regia (Bojer ex May-Jun Planted in Karachi, Anthelmintic Hook.) Raf. Hyderabad, Lahore Tamarindus indica L. Feb—Apr Grown in Sindh, Punjab, — Anti-viral against New Jehlum, Karachi Castle disease virus, astringent and sedative Fabaceae (Mimosoideae) Acacia nilotica (L.) Delile = May—Jun Found cultivated or Useful in jaundice and wild in Sindh, Punjab, palpitations Baluchistan, NWFP Albizia lebbeck (L.) Benth = Jul-Oct ws in Sialkot to Aperient, boils, carbuncle, ae seat Malakand nae erial Apr-Jul Found in : Ae against Prosopis cineraria (L.) Druce Fabaceae (Papilionoideae) Butea frondosa Roxb. = Mar-Apr ed Ne a (Lam Butea monosperma (Lam.) Mar-Apr Taub. Pongamia pinnata L. = Apr-May Millettia pi die (L) Panigra Sesbania Sibel P Aug—Mar Pers Kesbanie sesban (L.) Merr. = Apr-Nov Trifolium pratense L. Feb—Apr lridaceae Crocus sativus L. Oct cee Punjab (in Thal and Cholistan deserts) Cultivated in Punjab, NWFP Cultivated in Punjab, NWFP Cultivated in Sindh, Punjab Alias in Karachi, Kutch, Sindh, Punjab Found ulna and wild in Sindh, Pu Occurs in Sete Swat, Hazara Propagated by bulb in Baluchistan oe Anti-pyretic, appetizer, aphrodisiac, blood purifier, diuretic, tonic, viral hepatitis Astringent, aphrodisiac boil depurative, diuretic, gout, anti-leprosy agent Flowers are used in diabetes Flower juice improves vision (as eye drops) Anti-fertility activity reported Anti-asthmatic, aoe bronchi and expector Beneficial for liver, brain, heart, regulates the menstrual function 364 BRIT.ORG/SIDA 21(1) Table 1.continued Species Flowering Distribution Medicinal Remedies erio Lamiaceae Hyssopus officinalis L. Jun-Sep Cultivated in Kashmir, Used for chest congestion, angi, Upper Chenab flower tea is expectorant Leucas aspera (Willd.) Link Aug—Feb Found in Jammu, Ghat, Anti-tussive, decongestant Ravi, Chenab, Doab for children Mentha longifolia (L.) Huds. Feb-May Common in home gardens Carminati t imum basilicum | Nov—Apr Cultivated in Karachi, Decongestant Baluchistan, Punjab Pervoskia abrotanoides Kar. Sep-Oct Occurs in Baluchistan, Anti-pyretic hitral, Gilgit, Hunza, Quetta, Ziarat Linaceae Li itatissimum L. Feb-Apr Cultivated in Karachi Cardiac and nerve tonic Lawsonia inermis L. Jun Found in Sindh, Magnoliaceae AAi-hoely, -} Malvaceae Abutilon indicum (L.) Sweet Alth a tae lis L Gossyvnitum herb 1C 7s id uM L. Hibiscus annabinus L Hibiscus rosa-sinensis L. ~ Hibiscus sabdariffa L. Urena lobata L. Meliaceae Azadirachta indica A. Juss Melia azedarach L. Throughout the year Feb-—Mar Jul-Oct May-Jul Autumn Winter Apr-Sep Aug-Sep Sep—Dec Mar-Apr Mar—May Baluchistan, Punjab Cultivated in Punjab, FP Widely distributed in Karachi, Sindh, lower hills of Punjab Grows in Azad Kashmir, Peshawar Rawalpindi Cultivated as a crop in Punjab and Sindh Cultivated in Sindh, Karachi, Swat, Punjab, Chitral Grown as ornamental plant in Punjab, Sindh Cultivated in Karachi Occurs in Lahore, Jehlum, Changa Manga Found in Sindh, southern Punjab, lower Baluchis fv Found in Sindh and Punjab Anti-pyretic, sedative, soporific Used in dyspepsia, anti- pyretic, anti-emetic Anti-diarrheal, demulcent anti-hemoptysis, sedative and decongestant Emollient, demulcent, di- uretic, bronchial catarrh and rheumatism Extracted flowers used as ee and fo inducing menstrual flow Gastritis and popular laxative Cardiac tonic, expectorant, anti-pyretic, anti-tussive, decongestant Cathartic activity Aphtho pectorant decongestant sis,e Adrenalgic stimulant, dys- pepsia, also used in skin diseases Poultice to relieve head- ache, nervousness ATHAR AND SIDDIQI, MEDICINAL PLANTS OF PAKISTAN 365 Taste 1.continued Species Flowering Distribution Medicinal Remedies period Moringaceae Moringa oleifera Lam. Feb-Apr Cultivated in Rawalpindi, © Cholagogue, diuretic tonic planted in Sin Musaceae Musa sapientum L. = Feb-Sep Cultivated in Sindh, Anti-hypoglycemic Musa xparadisiaca L. Punjab, Myrtaceae Myrt mmuni Apr-Jun Wild in Baluchistan, As anti-septic, disinfectant NWFP Nyctaginaceae Mirabilis jalapa L. Nov—Jan Found in Karachi, NWFP, Anti-hemorrhoidal Hunza, Gilgit Nymphaeaceae Nelumbo nucifera Gaertn. © May-—Jul Found in Charsada, Cardiac tonic, diuretic, Multan, Shahdara anti-pyretic Oleaceae Jasminum grandiflorum L. Warm season Occurs in Peshawar, Aphrodisiac, astringent Karachi carminative, dysenter hepatitis, suppress excess lactation las! bac (L.) Jul-Oct Occurs in Karachi, Lahore, Anti-pyretic, cardiac tonic, Aiton islamabac lactifuge Paeoniaceae Paeonia emodi Wall. May-Jun Common in moist Anti-diarrheal ex Royle ground. Kaghan, Thandiani, Chitral, Bahrin, Poonch Papaveraceae Papaver rhoeads L. Jun-Sep Cultivated in gardens Bronchitis, hoarseness sedative, sudorific Passifloraceae Passiflora incarnata L. Jul-Sep Cultivated in Karachi Asthma, dysentery, insom- nia, whooping cough Pontederiaceae Eichhorina crassip Apr-Jul Occasionally found Arthritis and gout (Mart.) Solms filling ponds in plains Punicaceae Punica granatum L. May-Jun Commonly grown in Anti-diarrheal, dysentery, Quetta, Sibbi, Karachi, bronchitis Punjab, NWFP 366 Table 1. continued BRIT.ORG/SIDA 21(1) Species Flowering Distribution Medicinal Remedies erio Rosaceae Eriobotrya japonica Jul-Aug ns a in sub- Flowers are expectorant (Thunb.) Lindl. malayan zone Rosa xdamascena Mill. Jan—Jul ee in gardens Anti-HIV, aperient, cardio- of Pakistan active, liver protector Rosa foetida Herrm. Jan—Jul Found in Baluchistan, Anti-diarrheal Kurrum, Quetta, Ziarat eae Bassia sc oxb. Jul-Aug Cultivated in Sindh, Regarded as bronchitis, M cee ifolia cooling, cold, anti-tussive, (L.) Macbr demulcent and tonic Scruphulariaceae Verbascum thapsus L Jun-Aug Common in Chitral, Coughs, diarrhea, febrifuge Mansehra stimulant, pharyngitis Solanaceae Datura metel L. May-Jun Weedy places, Karachi Smoke as anti-asthma Solanum surattense Jun-Nov Throughout Pakistan Paresthesia, carminative Burm olanut virginianum L. Rubiaceae Ixora coccinea L. Jul-Jan Cultivated in Karachi Cure sores, relieve blood, ulcers Tropaeolaceae Tropaeolum magu Dec-—Feb Cultivated in Karachi Natural anti-biotic Verbenaceae Nyctanth bor-tristis Aug-Oct Naturalized in Punjab, Anti-pyretic, faintness, Rawalpindi, NWFP Mardan_ anti-vertiginous Vitex negundo L. Mar-Jun Cultivated in Thai, Swat, Cardio tonic, cholera, Mirpur diarrhea, useful for liver disorders Violaceae Viola odorata L. Mar—May Naturalized in ie gali, Hazara, Kaghan, Swa Liver protector, decongestant Pakistan has the potential for sustainable utilization of its medicinal flora. Basic infrastructure also exists to carry out research and development activi- ties in this field. However, more attention needs to be paid to the systematic propagation, collection and conservation of medicinal plants. It is important that the country consolidates its plant-resource database, explores the interna- ATHAR AND SIDDIQI, MEDICINAL PLANTS OF PAKISTAN 367 tional and national markets for its medicinal flora, and implements large scale cultivation projects and salvage projects for its more important medicinal-plant species. Pakistan needs an expansion of research efforts into cultivation, pro- duction and conservation of medicinal plants. There is also a dire need to de- velop a system of institutional linkages among all the research and development organizations dealing with the Pakistani medicinal and aromatic plant industry. ACKNOWLEDGMENTS Special gratitude is expressed to Joseph H. Kirkbride, USDA, Agricultural Re- search Service, Beltville, MD, for his help in checking the nomenclature and taxonomy of the plants and Mohammad Nawaz, California Department of Health Services, Sacramento, CA, for valuable suggestions and helpful criticism on the manuscript. The authors are also grateful to Zahoor Ahmad, Pakistan Agricultural Research Council, Islamabad, and Prof. M. Iqbal Choudhary, H.EJ. Research Institute of Chemistry, University of Karachi, Karachi, Pakistan for providing many original reprints used in the preparation of this paper. REFERENCES Au, S.1. and M. Qaiser. 1986. Phytogeopraphic analysis of phanerogams of Pakistan and Kashmir. Proc. Roy. Soc. Edinburgh. B89:89-101. Bacu, E. 1994. Flower remedies. Alternat. Med. 232-243. Bisset, N.G. 1994. Herbal drugs and phytopharmaceuticals. CRC Press, Boca Raton, Florida, Brummit, K.R.and C.E. Powett (eds.). 1992. Authors of plant names. Royal Botanic Gardens, Kew, U.K Bucti, R.A. 1998. Prospectus and potentials of saffron (Crocus sativus Linn.) cultivation in Baluchistan. In:N. Haq, K. Anthony, M. Sarwar and Z. Ahmad, eds. Proceeding of Sympo- sium on under Utilized Crops of Pakistan. Plant Genetic Resources Instititute, National Agricultural Research Center, and ICUC, Agha Jee Printers, Islamabad, Pakistan. Pp. 101-103. Duke, J.A. 1986.CRC Handbook of medicinal herbs. CRC Press, Boca Raton, Florida, U.S.A. Haruna, A.K. 1997. Cathartic activity of soborodo; the aqueous extract of calyx of Hibiscus sabdariffa L. Phytother. Res. 9:307-308 Kevicte, K. 1995. Herbs an illustrated encyclopedia. Michael Friedman Publishing Group, U.K. KHAN, H.A., T.O. Sippiau!, and J. Axmap. 1996. Medicinal plants and herbs as potential source of drugs for liver diseases: ethnic and traditional approaches. In: Hakim M. Said, M. Attaur Rahman, M. Abdulla, $.8. Vohra, and M. Athar, eds. Elements and Liver, Proceedings of International Symposium on Trace Elements and Liver Diseases, Hamdard Foundation Pakistan, Karachi, Pakistan. Pp.128-141. MaxHmooo, N., S. PIAcENTE, C. Pizza, A. Broke, A.|. KHAN, and AJ. Hay. 1996. The Anti-HIV activity and mechanism of action of pure compounds isolated from Rosa damascena.Biochem. Biophys. Res. Commun. 229:73-79. = 368 BRIT.ORG/SIDA 21(1) Matik, S.and S. Farooo. 1984. Cultivated trees, shrub and climbers of gardens of Pakistan. Taxonomic studies. PCSIR, Peshawar Laboratory, Peshawar, Pakistan. Nasir, E.and S.1.Au. 1972.An annotated flora of West Pakistan. Stewart Herbarium Gordon College, Rawalpindi, and Department of Botany, University of Karachi, Pakistan. Nasir, YJ. and R.A. Raria. 1995. Wild flowers of Pakistan. Elite Publishers Ltd., Islamabad, Pakistan. REYNOLD, J.E.F., K. Parritt, A.V. Parsons, and S.C. SweeTMAN. 1994. Martidale: the extra pharmaco- poeia. Info Access and Distribution, Singapore. Rizvi, M.A. 1998. Medicinal flowers of Pakistan, Part-lll. Horticultural Society of Pakistan, Karachi, Pakistan. Pp.48-52. SHAHEEN, F., M. Attaur RaHman, K.VASISHT, and M.lasat CHOUDHARY. 2003. The status of medicinal and aromatic plants in Pakistan, In: Medicinal plants and their utilization, UNIDO. Pp. 77-87. SHINWARI,M.1. and M.A. KHAN. 1998. Ethonobotany of Margalla Hills National Park of Islamabad. Department of Biological Sciences, Quaid-i-Azam University, Islamabad, and PASTIC Press, Pakistan. Wiiuiam, J.T.and Z. Admab. 1999. Priorities for medicinal plants research and development in Pakistan. Published by MAPPA, India, and IDRC, Canada. ZAMAN, M.B. and M.S. KHAN. 1970. Hundred drug plants of West Pakistan. Pakistan Forest Institute, Peshawar, and Ferozsons Ltd., Lahore, Pakistan. CUSCUTA (CONVOLVULACEAE)—THE STRENGTH OF WEAKNESS: A HISTORY OF ITS NAME, USES AND PARASITISM CONCEPT DURING ANCIENT AND MEDIEVAL TIMES Mihai Costea'! Francois J. Tardif oe of Plant Agriculture ee oneian Agriculture rsity of Guelph University of Guelph Guelph at NIG 2W1, CANADA Guelph ee N1 1 Wit CANADD coste_amihai@hotmail.com ftardif@uoguelph.ca ABSTRACT We examine the etymology and the first names of Cuscuta. as well as some unexplored aspects in the early history of parasitism concept. The name “Orobanche” was used for the first time by Theophrastus possibly for Cuscuta, however for sure not for the today’s broomrapes. The etymology of the generic name is neither Arab nor Greek, as commonly thought, but Aramaic and/or Hebrew. The ideas of parasitism ae medieval ee are merely a regression of the Greek ancient concept. Whereas the ancient Greeks view the parasites as originating orn their own seeds curing the medieval times le 1] } 1 } andas late as the 17th century, C t literally b ir hosts. Several lesser known uses of these plants dui aS ancient — ae al times are explored in the Old as well as in the New World. RESUMEN S ind la etimologia de | “uscuta, asi como, aspectos no explorados en la historia del soncept ae parasitism : Bomite: ee fue utilizado por la primera vez por f Teotrasto, [ los actuales “jopos de .” La etimologia del nombre genérico no es ni ace ni i priega, como se creia comunmente, aunque puede ser arameo y/o a breo. El concepto con respecto a las plantas parasitas desde la edad media es solamente una regresion al anti go. Mientras que, el punto de vista de los antiguos griegos era que las ee paras asitas se te de sus propias ie en la Edad Me- dia, a finales del siglo oh las Pe antas Cuse ula se consideraron _ literalmente surgian de sus hospedadores. | guos y en la Edad Media son practicamente desconocidos, por lo que éstos se cesta exclomsauo enel Viejo y el Nuevo Mundo. INTRODUCTION Cuscuta is a worldwide distributed genus, that comprises about 150 species of parasitic plants (dodders) (Yuncker 1932). The etymology of the generic name is not clear since it has commonly been said to be either Greek (e.g. Dawson et al. 1994) or Arabic (Austin 1980). The history of these intriguing plants, and of parasitic plants in general, has been traced back to antiquity (Mirande 1900; Kuijt 1969). Such plants must have drawn the attention of both early scholars \Corre pondina author. b . SIDA 21(1): 369 — 378. 2004 370 BRIT.ORG/SIDA 21(1) and farmers. As Theophrastus wrote: “the oddity is that a plant grows exclu- sively on another plant, and not on the ground...” (De causis plantarum 2:175, 1976). Since then, the concept of parasitism has evolved slowly. The early his- tory of parasitism in plants is particularly poorly known. We examine the first names of Cuscuta, some unexplored aspects in the history of parasitism con- cept, and several less known uses of these plants. Was Orobanche one of the first names of Cuscuta? Theophrastus referred to a plant called ‘dpoBayyn” Cvetch strangler” from “orobos” - vetch - Vicia ervilia (L.) Willd.?, and “anchone’ - to strangle) because “it overspreads the whole plant (of vetch) and holds it fast as if it were in coils, for itis thus that (vetch-strangler) strangles the plant” (Inquiry into plants 8.8.4, 1916). It is certain that, when using this name, Theophrastus did not have in mind the broomrapes (today’s Orobanche), because he apparently spoke about these in the next paragraph, under a different name. Thus, he mentioned oaptodapov, which is said to be like another unnamed plant that “springs straight from the roots of cummin” (Cuminum cyminum L.), and both these plants are “somewhat more peculiar in their habits.” This oiuod@pov attaches itself to fenugreek” (Trigonella foenum-graecum L.) and its “root is more or less round” (Theophrastus, Inquiry into plants 8.8.5, 1916). Both these latter plants are prob- ably the actual Orobanche, which cannot be the 6poBayyn, the “vetch-strangler.” It would be tempting to assume that ‘dpofayy’ (the vetch-strangler) is Cuscuta. This would mean that one of the first names of Cuscuta was Orobanche. In sup- port of this hypothesis are the habit of the plant and the chapter about weeds in which Theophrastus mentioned it (Inquiry into plants 8.8.3-5, 1916). Unfor- tunately, the “vetch-strangler” was not further described or mentioned elsewhere by Theophrastus, and Pliny the Elder did not carry on this name. The Labyrinth of names and languages Theophrastus also spoke about the “small Syrian weed kasytas (kaoutag) that grows on trees, thorn-bushes and certain other plants” (De causis plantarum 2:17.3, 1976). Mirande (1900) argued that “kasytas” is probably not Cuscuta. He suggested that since Pliny’s subsequent reference to this plant (“cadytas”) alleg- edly alludes to some aromatic properties, this plant may be Cassytha spp., a coastal plant from Lauraceae resembling Cuscuta in the parasitic habit. It must be noted that Pliny in his description (Natural History 13:129, 1951) amalgam- ated “kasytas” with other, unnamed plant from the account of Theophrastus About this unnamed plant, Theophrastus said that is “sown in the dog days on the thorn-bushes in Babylonia that sprouts the same day and then speedily en- velopes the bush” (De causis plantarum 2:17.3, 1976). Pliny referred to these two plants as if they were a single one, and added that this plant “is used in making spiced wine and it is cultivated for this purpose” (Natural History 13:129, 1951). An even earlier indication to the preparation of a drink can be found in the COSTEA AND TARDIF, CUSCUTA, A HISTORY OF ITS NAME, USES AND PARASITISM 371 Talmud: “Why are not sufferers from ra’athan in Babylon? Because they ... drink beer containing Cuscuta (growing on) the hizme shrub” (Kethuboth 77b; see Epstein 1961). Epstein (1961) indicated that the “hizme shrub” is “Spira (Spina?) Regia.” We could not determine what this spiny shrub was: Tragacantha (As- tragalus tragacantha L.), Rhamnus spp. Paliurus SpP., Berberis SPP, Zizyphus? Therefore, Theophrastus’ “kasytas” may be either Cuscuta or Cassytha, and the etymology of the two names is probably the same. Even in the 16th century, Fuchs (1542) used the name “Cassutha” for a dodder (Fig. 1) that, judging by its mentioned host, was apparently Cuscuta epilinum Weihe. Several authors mentioned that the name has Greek origin (e.g. Dawson et al. 1994). But “kasytas” does not have any meaning in Greek and it would be more logical to assume that the name was introduced. Although Austin (1979) initially suggested that that the name comes from Aramaic, he mentioned later that the name has an Arabic origin (Austin 1980). However, apparently “kushkuut,” “kashuut” or “koshotit” as Arabic names of Cuscuta are early Ara- maic neologisms (Fraenkel 1886; Nakhle Al-Yassu’i 19860). Many ancient Ara- bic words, especially related to agriculture were introduced from Aramaic (Nakhle Al-Yassu’i 1986). Based on the new root “k-sh-w-th,” the word “kushsha” meaning in Arabic “a lock of hair” was probably derived (David Mehall, pers. comm). Since Theophrastus mentioned Babylonia (Assyria) as a place of origin for “kasytas,” this was probably from the geographical source of the name as well. Indeed, the etymology can be retraced in Syriac Aramaic and Hebrew (ksutha and ksuth, respectively), two closely related Semitic languages. The triradical root of the verb K-S-Y (Kaph, Shin, Yodh) means “to cover.” Based on this root, a verbal noun that signifies “cover,” “clothing” or “garment” is con- structed in both languages: K-S-W-T (in Hebrew) and K-S-W-T-A (Kaph, Shin, Waw, Tav, Aleph) in Aramaic (Yona Sabar, pers. comm.; Paul Flesher, pers. comm; Sokoloff 1990). “Ksutha” is often mentioned in the Talmud (see Epstein 1961) and other Jewish-Aramaic texts suchas the Targums VJastrow 1950; Bowker 1969; Sokoloff 1990). The word was introduced into Arabic, Persian and Greek, creat- ing a bridge between these fundamentally different languages and cultures. Before Pliny, Dioscorides said that “Epithumon.... is ye flower of ye harder Thyme, & like to Satureja” (4:179, see Gunther 1959). Pliny mentioned two types of Epithymum: “one which is like the flowers of the thyme and satureia” “qui est flos e thymo satureiae simili”) and another one “that grows without a root, has slender a head like a long mantle, is red in color and is dried in the shade” (“sine radice nasci, capite tenui similitudine pallioli, ruben, siccari in umbra”) (26:55-56). The recognition of these two “types” persisted until medieval age, when Tragus (1552) and Parkinson (1640) considered that they belong to “one kind growing on different herbs.” Between the 16th and 17th centuries, dodders were called differently depending on the author. For example, Casper Bauhin (1623) used both “Epithymum” and Cuscuta. A common practice during that 372 BRIT.ORG/SIDA 21(1) eee eee Fi Ce 5 aa Ng) ta) if on ot AP “ . . aN i SRS HglPOO “pt ‘ae “as if . We a>, S, ¥ x va Ny ena nti: y Le PAS » TAC kts a “Ry en ee i AS Pe . z oer) 4 pk > h \ # 4 OF nag ean ee Re, > eigen ef Pree emt Ss ~ eo; eee gS PPR PSR IN ” oe! és —-, 5 ee Fr ia Sate; YaPhee ix pat ay. Saale -CASSVTHA Hladh$ feader. Fic. 1. Cuscuta spp. (probably C. epilinum). Fuchs, 1542. COSTEA AND TARDIF, CUSCUTA, A HISTORY OF ITS NAME, USES AND PARASITISM 373 time was to create a Latin name based on the hosts the dodders are growing upon, e.g. “Epithymum” (on Thymus spp.), “Epilinum” (on Linum spp.), “Epimajorana” (on Majoran), “Epistoebe” (on Stoebe - Vicia spp.), “Epiurtica” (on Urtica spp.) (e.g. Gerard 1633; Parkinson 1640). Apparently, Ray (1682) and Tournefort (1694-1695) were the first to retain only Cuscuta, which was later officially endorsed by Linnaeus (Sp. Pl. 124. 1753). The parasite, “a spontaneous act of nature” Kuijit 1969) suggested that the earliest reference to parasitic plants belongs to Theophrastus. However, an even earlier record can be found in Aristotle: “for even among land-plants there are some that are independent of the soil, and that spring up and grow, either parasitically upon other plants, or even entirely free (De partibus animalium 4:681b, 2001). Ancient Greeks were well-aware of the parasitic phenomenon. Even rooted, green lianas, like the ivy (Hedera helix L.) were regarded as “taking the food that belongs to the tree” (Theophrastus, De causis plantarum 2:17.4, 1976). And approximately three centuries latter, Pliny wrote: “it is a well-known fact that trees are killed by ivy” Chedera necari ar- bores certum est”) (Natural History 16:243, 1951). The animal-oriented biological treaties of Aristotle have influenced our understanding of plant parasitism for almost 2000 years. Theophrastus was Aristotle’s student and he developed many of the ideas of his teacher. “It may perhaps be that like animals, plants are fond of one another and live together” (Theophrastus, De causis plantarum 2:17.5, 1976). Or, that “some (plants) come from seed as through by a spontaneous act of nature. The latter come from rot- ting earth or from rotting parts of plants: for some are not constituted sepa- rately by themselves but are produced on trees, for example the mistletoe” (Ar- istotle, De generatione animalium 1:715b. 25-30, 1965). Following this idea, Theophrastus observed that maybe some plants “came from a corruption of something in the host” similarly to “some animals that can arise from other animals” (De causis plantarum 2:17.5, 1976). For example, like “those insects (such as lice and bed bugs) that ... live on the flavors of living flesh ... (and are pro- duced) from the moisture from animals” (Aristotle, De partibus animalium 31:556b. 21-28, 2001; also De generatione animalium 1:715b.5, 1965). But it must be emphasized that the Greek spontaneous concept does not preclude the ori- gin of the parasite from seeds: “birds eat its fruit and let their droppings fall on the host tree.” (Theophrastus, De causis plantarum 2:17.6, 1976). The wonder of ancient Greeks came from their belief that since plants “absorb the food already prepared from earth with their roots” they don’t need a stomach, and they don’t produce excrement (Aristotle, De partibus animalium 2.10, 2001). This is why plants “not growing on the ground atall is astonishing” (Theophrastus, De causis plantarum 2:17.5, 1976). In the Babylonian Talmud, the parasite was explicitly depicted as an integral part of its host: “if one plucks Cuscuta from shrubs and 374 BRIT.ORG/SIDA 21(1) thorns ... one is culpable as if it inserts his hand in an animal’s bowels and de- taches an embryo” (Shabbath 107b; see Epstein 1961). Later, Arabians were inspired by Greek science and introduced it into Eu- rope. The ancient meaning of the “spontaneous act” degraded and the parasite was literally viewed as arising from its host. For instance, Francis Bacon, in the England of the early 17th century, believed that mistletoes were an exudation (sap) “that the tree doth excern and cannot assimilate,” and that the idea of birds spreading its seeds is merely a “fable” or a “tale” (Sylua Syluarum 556-557, 1626). This misconception was reflected in the “Uva Barbata” or “Le Raisin Barbu,” a Cuscuta spp. parasitizing Vitis vinifera L.,and considered together with its host either as an entirely different species of grape wine (Tabernaemontanus 1588- 1591), or a “monstrosity,” a metamorphosis of the normal species (Jean Bauhin 1591). As late as 1831 such a blemish was connected to the appearance of the comet in the previous year (Kuijt 1969). In conclusion, the “spontaneous” con- cept regarding the origin of parasitic plants from medieval ages was merely a regression of the ancient Greek concept. Rufinus, a forgotten botanist of the 13th century, was probably the first to observe that “Cuscuta ...is born in meadows and it ties to other plants, which it kills” CCuscute ... nascitur in pratis et ligat alis herbas et necat eas”) (Rufinus approx. 1300 A.D.; see Thorndike 1946). His short note passed unobserved by all botanists and herbalists, and about three centuries later, Parkinson (1640) rediscovered that dodders “spring from their owne seede.” He emphasized the discovery several times to “let all others understand”... it may appeare plainely toany that neither Tyme, Savory, or any other herbe, doe naturally of their owne seede bring these strings or laeces but that they spring from their owne seede, either shed or scattered of themselves upon ground, or coming among the seedes of other hearbs that are sowne” (Parkinson 1640). Cuscuta, the strength of weakness Theophrastus compared parasitic plants with grafted shoots and buds that “get ood that is more readily available and that has been prepared and practically concocted” (De causis plantarum 2:17.6, 1976). A good understanding of the para- sitic phenomenon can be also encountered in the Babylonian Talmud, at ap- proximately 100 BC. Although apparently it takes its food “from air ... Cuscuta derives its nourishment from soil (through its host); for we may observe that when the shrub (on which Cuscuta grows asa parasite) is cut off, Cuscuta dies” (Erubin 28b; see Epstein 1961). But “a plant seeking such food would be weak in its nature” (Theophrastus, De causis plantarum 2:17.6, 1976). Furthermore, such plants must be weaklings since they don’t sprout and grow by themselves (Theophrastus, De causis plantarum 2:17.6, 1976). The same idea of helpless- ness, exemplified with “cadytas” and “viscum,” can be found at Pliny: “because they have no abode of their own and consequently live in that of others” COSTEA AND TARDIF, CUSCUTA, A HISTORY OF ITS NAME, USES AND PARASITISM 375 (“namque cum suam sedem habeant, in aliena vivunt,” Natural History 16:39. 243-244, 1951). Ina sense this is true; today we know that during the parasitiz- ing stage, Cuscuta depends entirely on the nutrients provided by its host (Dawson et al. 1994). Nonetheless, gradually the harmful effect of dodders on their hosts outweighed other considerations. Fuchs (1542) noted that Cuscuta epilinum “grows... over plants, quite often entangling them so thickly with its web that shuts off the sun like a tent.... It forces out growth from the host plant, and strangles it with its filamentous stems, or, by continuous sucking of the mother's (host’s) sap, reduces it to exhaustion” (Historia Stirpium:347). Even today, it is intuitively seductive and deeply rooted in our unconscious fears to imagine plant parasitism in terms of animal parasitology. The hot or cold character of dodders and several old uses Quoting Phoenicians and Arabs, Fuchs and other medieval herbalists (Ruellius 1529; Gerald 1633; Parkinson 1640; Culpeper 1652) believed that the curative powers of dodders depend on the “character of the parent (host): if it invades a warm plant, it strengthens its heating nature, and if it clings to a cold one, it will acquire the cold strength” (Fuchs, Historia Stirpium:349, 1542). This is why during medieval times, dodders were named according to the host they are grow- ing upon (see above) (e.g. Gerard 1633; Parkinson 1640). Among these, “Epithymum’” was said to have the best therapeutic properties because Thy- mus, its host, is dry and hot in the third degree, as indicated by Galen (Tragus 1552; Gerard 1633; Parkinson 1640; Culpeper 1652). “Physitians crying up Epithymum, (most of which comes from Hymettus in Greece, or Hybla in Sicilia, becaus those Mountains abound with Time) he is a Physitian indeed that hath wit enough to chuse his Dodder according to Nature of the Diseas and Humor peccant” (Culpeper 1652). “That dodder growing upon Tares (Vicia spp.), being the most frequent in London, and wherewith our markets are onely in a manner furnished and Apothaecaries shoppes stored from thence ...can have no effectual quality com- parable to Epithymum for ... Tares are hard of digestion and binde the bellye and the nourishement of them engendereth thicke blood apt to turn into melancholie” (Parkinson 1640). Today we know that, indeed, the parasite forms a biochemical continuum with its host, and that its chemical makeup may de- pend on the latter. Primary metabolic compounds, minerals, xenobiotics, sec- ondary products (e.g. alkaloids and cardenolides), herbicides, viruses, and my- coplasmas are translocated from the phloem of the host via the haustoria to the parasite (Dawson et al. 1994). “You are what you eat” couldn’t be more true for these plants. Dioscorides, and later Pliny, recommended “Epythimum” asa purgative, as well as “for melancholicall, & ye puffed up with wind, ye quantity of an ac- etabulum to ye quantity of 4 dragms with honey & salt, and a little Acetum 376 BRIT.ORG/SIDA 21(1) (4:179, see Gunther 1959; also Pliny the Elder, Natural History 26:55). In an Ara- bic text dated before 1000 AD it is written: “If you put an euquia of blood (of the black dog) with a mithqal? of grounded Cuscuta; who will take this will never be reached by the spirits which causes enchantment. The meat (of black dog) in food with salt, is good a antidote against infantile epilepsy” (Alfonso-Goldfarb 1999). Other Arabic uses of Cuscuta have been reviewed by Guigues (1909), Hamarneh (1973) and Levey (1966). These were the main uses of dodders, which were later retained by all medieval herbalists. For example, Gilbertus Anglicus, around 1250 endorsed “Epithymum” as a remedy in a mixture of plants that “purge the head of evil humors” (Getz 1991). Culpeper (1652) noted that it is also good “to purge black or burnt Choller, which is the cause of many Diseases of the Head and Brains, as also for the trembling of the Heart, faintings, and swounings.” Since dodders are under the sign of Saturn, “this helps by Sympa- thy, & strengthensal the parts of the Body he rules: Melancholy, Addust Choller, Trembling, fainting, swooning, Spleen, Hypochondria, Obstructions, Gall, Jaun- dice, Liver, Disury” (Culpeper 1652). ACKNOWLEDGMENTS Many thanks are due to Guy Nesom, Ovidiu Constantinescu and Daniel Austin who reviewed the manuscript and made interesting observations. Numerous linguists kindly helped in our search for meanings and possible etymologies of Cuscuta: Paul Flesher, Yona Sabar, and Michal Sokoloff with the Hebrew and/ or Aramaic, and Safa Jubran, David Mehall, Hanita Brand, Irfan Shadid and Binyamin Abrahamov with the Arabic. Safa Jubran sent us the book she trans- lated with Ana Maria Alfonso-Goldfarb. Sawsan Oran and Mohammed Sawaie gave us names of Cuscuta from Northern Africa. Rabbi Nissan oriented our search toward the Talmud. Yolanda Camacho translated the Spanish abstract. REFERENCES ALFONSO-GoLDFARB, A.M. 1999. Livro do tesouro de Alexandre. Translated by S. Jubran and A.M. Alfonso-Goldfarb. Editora Vozes, Petrdpolis, Brazil Aristorte (384-322BC). 1965.De generatione animalium.Recognovit Brevique Adnotatione Critica Instruxit HJ. Drossaart Lulofs. Oxonii, E Tlyoographeo Clarendoniano. Aristorte (384-322 BC). 2001. De partibus animalium. Clarendon Press, Oxford University Press, Oxford, NY. Austin, D.F. 1979. Plants without beginning or end. Fairchild Trop. Gard. Bull. 34:17-19. Austin, D.F. 1980. Studies of the Florida Convolvulaceae—lll. Cuscuta. Florida Sci. 43: 294-302. Bacon, F. 1626. Sylua syluarum: or A naturall historie: In ten centuries. Written by the Right ) *euquia = 32.76g; 1 mithgal = 3.9g (Safa Jubran, pers.comm. COSTEA AND TARDIF, CUSCUTA, A HISTORY OF ITS NAME, USES AND PARASITISM 377 Honourable Francis Lo. Verulam Viscount St. Alban. Published after the authors’ death, by William Rawley Doctor of Diuinitie, late his Lordships chaplaine. London. BAUHIN, C. 1623. Pinax theatri botanici, sive index in Theophrasti Dioscoridis, Plinii et Botanicorum quia saeculo scripserunt opera. Basileae Helvet. Subtibus & Typis Ludovic Regis. BAUHIN, J. 1591. De plantis a Divis Sanctis. Additae sunt Conradi Gesneri Medici Clariss. Epistolae a Casparo Bauhino. Basileae Apud Conrad. Waldrich. Bowker, J.W. 1969. The targums and rabbinic literature: an introduction to Jewish interpre- tations of scripture. Cambridge University Press, Cambridge, UK. Cucperer, N. 1652. The English physitian:or an astrologo-physical discourse of the vulgar herbs of this nation, Peter Cole, London. Dawson, J.H., L.J. MusseLman, P. Wotswinkel, and I. Dorr 1994. Biology and control of Cuscuta. Rev. Weed Sci. 6:265-317. Epstein, |. (Editor and translator). 1961. The Babylonian talmud. Soncino Press, London. FRAENKEL, S. 1886. Die Aramaeischen Fremdwoerter im Arabischen, Leiden. Fucus, L.(1542).1999.The great herbal of Leonhart Fuchs. De historia stirpium commentarii insignes. G. Meyer, E.E. Trueblood, and J.L. Heller, eds. Vol. 2. Stanford University Press, Stanford, CA. GerARD, J. (1542-1612). 1633. (republished in 1975). The herbal or general history of plants. Dover Publications, New York. Getz, F.M.1991.Healing and society in medieval England: A middle English translation of the pharmaceutical writings of Gilbertus Anglicus. University of Wisconsin Press, Madison. Guicues, P. 1909. Les noms arabes dans Serapion, “Liber simplici medicina.” Journale Asiatique 10:473-546. GunTHeR, R.T. 1959. The Greek herbal of Dioscorides. Hafner Publishing Co. New York. HaAMARNEH, S.K.1973.Al-Birundi’s book on pharmacy and materia medica. Karachi, Hambard National Foundation, Pakistan. Jastrow, M.1950.A dictionary of the Targumim, the Talmud Babli and Yerushalmi, and the Midrashic literature, Pardes Publishing house Inc., New York. KuuT, J. 1969. The biology of parasitic flowering plants. University of California Press, Berkeley, CA. Levey, M.1966.The medical formulary of Aqrabadhin of Al-Kindi. The University of Wisconsin Press, Madison. Miranbe, M. 1900. Recherches physiologiques et anatomiques sur les Cuscutacées. Thesis, fac. Sci. Paris. Naktle Al-Yassu’i, R. 1986. Ghraai’b al-lughah al-arabiyya (Strange things in the Arabic lan- guage). Dara El-Mashreq, Lebanon. ParKINSON, J.(1567—1650). 1640. Theatrum botanicum or an herbal of a large extent. Printed by T.Cotes, London. PLINY THE Evper. (ca. 23-79 A.D.). 1951. Natural History, Vol. IV (Libri I-XVI) and Vol. VII (Libri 378 BRIT.ORG/SIDA 21(1) XXIV and XXVII) translated by W.H.S. Jones and D.Litt. The Loeb Classical Library, Cam- bridge, Massachusetts and Harvard University Press, London. Ray, J. 1682. (reprinted 1962). Methodus plantarum. Wheldon & Wesley, Ltd. and Hafner Publishing Co., New York. Ruetuus, J (1529). 1537. De natura stirpium libri tres. Froben, Hieronymus & Nic, Bazel. Soxotorr, M. 1990. A dictionary of Jewish Palestinian Aramaic of the Byzantine period. Bar Ilan University Press, Ramat-Gan, Israel. TABERNAEMONTANUS, J.T. 1588-1591. Neuw Kreuterbuch. Mit schonen kinstlichen und leblichen Figuren unnd Conterfeyten. Neuw und volkommentlich Kreuterbuch. Das Ander Theyl. Das dritte Theil des Kreuterbuchs. Nikolaus Basse, Frankfurt. THEOPHRASTUS (Ca. 371—-Ca. 287 B.C.). 1916. Inquiry into plants and minor works on odors and weather signs. Vol. |, Il. English translation by A. Hort. The Loeb Classical Library, Heinemann, London THEOPHRASTUS (Ca. 37 1—-ca. 287 B.C.). 1976. De causis plantarum. Vol. I-lll. English translation by B. Einarson and G.K.K. Link. The Loeb Classical Library, Heinemann, Harvard Univer- sity Press, London. THornpike, L. 1946. The herbal of Rufinus. University of Chicago Press, Chicago. Tournerort, J.P. 1694-1695. Elemens de botanique ou methode pour connoitre les plantes. L'Imprimerie Royale, Paris, Tracus (Bock Hieronymus). 1552. De stirpium, maxime earum quae in Germania nostra nascuntur, usitatis, nomenclaturis, propriisque diferentiis neque non temperaturis ac facultatibus Commentariorum libri tres ... interprete Davide Kybero ... His accesserunt praefationes duae :altera Conradi Gesneri ... adjectus est Benedicti Textoris Segusiani de Stirpium differentiis, ex Dioscoride secundum locos communes. Argentinae, Wendel Rihel. YuNCKER, 1.G. 1932. The genus Cuscuta. Mem. Torrey Bot. Club. 18:113-331. HISTORY AND EPONYMY OF THE GENUS NAME AMSONIA (APOCYNACEAE) James S. Pringle D | Se co mg U CUA HCC VDIUIUET IS PO. Box 399 Hamilton, Ontario, Canada L8N 3H8 jpringle@rbg.ca ABSTRACT The genus Amsonia was named for Dr. John Amson, a physician in Williamsburg, Virginia, in the id ] t th LLbLita nr Iry. RESUMEN El genero Amsonia fue nombrado por Dr. John Amson, un médico de Williamsburg, Virginia, en la mitad del siglo diez y nueve. Amsonia Walter is a genus of about twenty species native to North America and eastern Asia, of which the best known is A. tabernaemontana Walter, the eastern bluestar. The name Amsonia was first applied to that species by John Clayton of Gloucester, in colonial Virginia, in the 1750s, and ever since that time uncertainty has persisted and statements have differed as to its derivation or eponymy. Clayton did not include the name Amsonia in the manuscript that he sent to the Dutch botanist Johan Frederik Gronovius, much of which Gronovius (1739) incorporated into his Flora Virginica. At that time Clayton thought that the bluestar might be considered a species of Nerium (oleander), and designated it “Anonymus Suffrutex folits Salicis alternis...Nerii species.” He gave it the name Amsonia in the later manuscript (not extant) on the plants of Virginia that he sent to Peter Collinson in England in 1757, but that work was never published. He also included the name Amsonia with specimens and seeds that he sent to British and European botanists. Still later Clayton proposed a different name for the eastern bluestar, based on the form of the seeds, but that name likewise remained unpublished. By then, the name Amsonia had become inseparably associated with this species, which had quickly become popular in British hor- ticulture following its introduction as Amsonia by Philip Miller in 1759 (Smith 1819; Berkeley & Berkeley 1963, 1982). Linnaeus received a description of the eastern bluestar from his former stu- dent Daniel C. Solander in 1761. Solander, who was living in London at the time, had based this description on plants he had seen in the gardens of Peter ‘Contribution No. 112 from the Royal Botanical Gardens, Hamilton, Ontario, Canade SIDA 21(1): 379 — 387. 2004 380 BRIT.ORG/SIDA 21(1) Collinson and James Gordon, supplemented with information from Clayton’s manuscript. He informed Linnaeus that Clayton had called the species Amsonia, but added that “whence the name was taken | know not,” nor did any of the botanists in London and vicinity of whom he had inquired; they believed “that Clayton named it for someone in North America” (letter from Solander to Linnaeus 16 November 1761, excerpted by Berkeley & Berkeley 1963). In 1762 Linnaeus received a sketch and information on the eastern bluestar from the botanical artist Georg Dionysius Ehret, with a promise of specimens to be sent via Solander (Calmann 1977). Ehret had been approached with regard to illus- trating the work that Clayton had sent to Collinson, and had studied specimens collected by Clayton. Later that year Linnaeus (1762) named the species Tabernaemontana Amsonia. The “Anonymus Suffrutex” was designated Clayton 306 by Gronovius, but no such specimen is in the Clayton herbarium now at BM. The Linnaean her- barium at the Natural History Museum in Stockholm (S-LINN) contains a speci- men of the eastern bluestar from the herbarium of Clas Alstrémer, who had acquired it from Andreas Dahl. It is labeled “Tabernaemontana Amsonia” and “Dahl a Linné P” in Dahl’s handwriting, indicating that Dahl had received it from Linnaeus himself (Lindman 1908). It was perhaps part of a collection made by Clayton and given to Linnaeus by Ehret or Gronovius, or it may have been obtained by Solander from a cultivated plant. Amsonia was published as a genus name by Thomas Walter in 1788. The name of the type species, Amsonia Iabernaemontana Walter, was based on Taber- naemontana Amsonid L. Walter gave no derivation or eponymy for the name. Many recent references state that Amsonia was named for “Charles Amson.” The earliest association of the forename Charles with Amsonia that | have found was by Loudon (1830), according to whom the genus was named for “Charles Amason [sic], an American traveller.” This eponymy for Amsonia was soon adopted in other British and Continental European references. The surname was altered to “Amson” by Paxton (1840), who identified Charles Amson as “a traveller in America.” Nicholson (1884), whose wording has been retained in the successive editions of the Royal Horticultural Society Dictionary of Garde n- ing, identified Charles Amson asa “scientific traveller in America.” It may be pertinent to the credibility of thiseponymy that Loudon almost certainly would have seen Smith’s (1819) account of the Neotropical genus Amasonid Lf. Verbenaceae). As noted by Smith, Linnaeus filius had stated that Amasonia was named “in memory of Amason, a traveller in America” (transla- tion), whom, according to Smith, “M. [Alexandre] De Thies [had] baptized Tho- mas.” Smith was skeptical, because he had “never been able to learn any tidings of sucha person.” Linnaeus lilius provided no forename or further information, and may merely have assumed that South America’s largest river, the name of which is spelled “Amason” in Swedish and some other European languages, was vy PRINGLE, HISTORY AND EPONYMY OF THE GENUS AMSONIA 381 named for an explorer. Loudon, however, having seen statements that Amasonia was named for Thomas Amason, may have followed De Thies’s alleged prece- dent a step further and arbitrarily coined the name Charles Amason, and des- ignated that imaginary person, like Thomas Amason, “an American traveller.” Of Amsonia, Smith (1819) said that “no positive account of the meaning or origin” of the name could be given, noting only that it had originally been be- stowed by Clayton. He speculated that the spelling of the names Amasonia and Amsonia might represent an error, and that both genera might have been named for the British Admiral George Anson, Baron Anson of Soberton. Rafinesque (1838, 1840) accepted this eponymy with none of Smith’s uncertainty and “cor- rected” the spelling to “Ansonia,” but no others have done so. The eponymy given by Loudon and Paxton was probably the basis for Gray’s (1856) statement, in the second edition of his Manual, that Amsonia was “said to be named for a Mr. Charles Amson.” That uncertain wording was retained through the sixth edition of Gray’s Manual. Wood (1861) expanded it to “Dedi- cated to Charles Amson, of S. Carolina?” The speculative addition was not ex- plained, but may have been made because the genus name was published by Walter, whose botanical activity had been confined to South Carolina. Britton and Brown (1896) stated with no such reservation that the genus was named “for Charles Amson of South Carolina.” That wording was soon adopted in other American publications. Woodson (1928) was skeptical about references to “Charles Amson.” He consulted encyclopedic and historical references and made inquiries of histori- cal societies in Virginia and the Carolinas, and found no evidence that any Charles Amson had lived or traveled in Virginia or the Carolinas during the colonial period or contributed to the knowledge of their natural history. Like- wise, using more recent biographical references and publications from that his- tory-conscious region, as well as Internet search engines, I found no record of any Charles Amson in those colonies during the 18th century (except as the supposed eponym of Amsonia). Nor have I found any mention of either of Loudon’s “travellers” in any other context. Consequently, | share Smith’s, Gray’s, and Woodson’s skepticism about their having existed. Had they lived and ex- plored the wilds of the Americas, it seems unlikely that they could so greatly have impressed Clayton and Linnaeus filius, and at least have been known to Loudon, yet so completely have escaped the attention of historians and biogra- phers. A thorough search would be complicated by the apparently indepen- dent origins of the surname Amason in England and Scandinavia. Even if these Amasons existed, however, there is ample evidence, presented below, that the apocynaceous genus Amsonia was not named for either of them. The earliest extant indication that Amsonia was named for a physician is in a letter written 30 December 1759 by Ehret (Berkeley & Berkeley 1963), who had probably seen words to that effect in Clayton’s manuscript or correspon- 382 BRIT.ORG/SIDA 21(1) dence. Apparently not distinguishing between Carolina and Virginia as Clayton's place of residence, he wrote that Amsonia had been named “perhaps for a doctor in Carolina.” An indication of the eponymy of Amsonia by Clayton himself appearsina letter from him to the Philadelphia botanist John Bartram dated | September 1700, in which he said that he had previously that the plant be named Amsonia “for a doctor, here” (William and Mary College Quart. Hist. Mag., ser. 3, 6: 319. 1926; Woodson 1928). Woodson inferred from Clayton’s use of “here” that Amsonia had been named for “a physician of Gloucester County, Virginia.” He added that “regarding his complete name, or the positive spelling of his family name, doubt still remains.” The same letter was probably the basis for the state- ments by Robinson and Fernald (1908) and Fernald (1950), in the seventh and eighth editions of Gray’s Manual, that Amsonia was “Named for Dr. Amson, physician of Gloucester, Virginia, in 1760, and friend of John Clayton.” Combi- nations of “Charles Amson,” from Loudon, Paxton, Gray, and Britton and Brown, and “physician” and “Virginia,” from Robinson, Fernald, and Woodson, have come to prevail in recent publications. Berkeley and Berkeley (1963, 1982) found from a source on Virginia medi- cal history (Blanton 1931) that Dr. John Amson had been practicing in Williamsburg, Virginia, in 1751. They 1982) concluded that “Clayton had named the new genus Amsonia in honor of his friend Dr John Amson of Williamsburg.” I have not found this information repeated by any more recent botanical authors. erkeleys found one other published reference to this Dr. Amson, which noted only that he frequently used the expression “neither here nor there Present-day search engines, although not providing all standard biographical data, do lead to further information about the Dr. Amson who was Clayton’s contemporary in colonial Virginia. Records in the Archives of Colonial Williamsburg, compiled and annotated by Mary A. Stephenson (1961), confirm the Berkeleys finding that John Amson, a “Doctor of Physic,” lived and pone uy Williamsburg in the mid-18th cen- tury. Clayton would have considered Williamsburg, ca. 22 km away across the York River in James City and York counties, near enough to Gloucester to jus- tify the use of “here” in writing to Bartram in Philadelphia, especially since he had spent part of his youth in Williamsburg and members of his family lived there. I have found no definite information on this Dr. Amson’s g ce or his life prior to his practicing medicine in Williamsburg. Records compiled by the Church of Jesus Christ of Latter-Day Saints provide limited data on a few indi- viduals named John Amson who lived in England and Scotland during the first 40 years of the 18th century, but no data on anyone by that name in colonial Virginia. The record perhaps most likely to be relevant is that of John Amson, son of James and Margaret Amson, who was baptized 24 July 1699, at the age of one year, in St. Sepulchre Church, London (where Captain John Smith of early — a) — PRINGLE, HISTORY AND EPONYMY OF THE GENUS AMSONIA 383 Virginia history had been interred). The time and (in view of data presented below) the country fit, but nevertheless it is by no means certain that this was the same John Amson. Although an English origin is not unlikely, the name Amson does not ap- pear in Munk’s Roll of the members of the Royal College of Physicians. How- ever, since the surname is not common, it does seem probable that Dr. John Amson of colonial Virginia was the John Amson, M.D., who received that de- gree from the university at Rheims (now Reims), France, 2 September 1722. Such credentials would have been impressive in colonial America, consistent with the apparent reputation (below) of Dr. Amson of Williamsburg as one of the “best Physicians” in Virginia. This record is from a list of early students from the British Isles at medical schools in continental Europe, compiled by Dr. Harold T. Swan (1996). This compilation was based on manuscript notes that had been in the possession of the medical historian Dr. Robert W. Innes Smith at the time of his death in 1933 (originals at the University of Sheffield). Aside from the data noted above, it states only that this John Amson was English, as distin- guished from Scottish or Irish. According to Dr. Swan (pers. comm. 1] April 2003), Innes Smith’s notes contain nothing further on Amson. He considers it unlikely that Innes Smith found additional information at Reims, because many of the university's records from the relevant time had been destroyed, presumably during the French Revolution. I have not found when Amson arrived in America, if in fact he was of En- glish birth. The earliest record of his presence in Virginia that | have encoun- tered is from 1738, when the estate of the late Dr Charles Brown of Williamsburg retrieved a book that Amson had borrowed from Brown’s library James 1895). A physician in Williamsburg in 1738 could hardly have avoided the contro- versy associated with John Tennent, a “practitioner of medicine” in that city (Blan- ton 1931: Jellison 1963). Tennent’s domestic medical manual, Every Man His Own Doctor, was highly popular, but some physicians questioned his competence and objected to his pursuit of financial recognition from public funds. In particular, his advocacy of the use of Polygala senega L. became obsessive, and his reactions toward those who took issue with him became intemperate. He felt that physi- cians who held medical degrees from European universities were unjustifiably disdainful of those who had obtained their qualifications through apprentice- ship and independent study. | have found no record of Amson’s involvement in this controversy. In one of his statements in his own defense, however, Tennent (1738) implied that medical degrees were awarded too freely at Rheims, alleging that one had inadvertently been bestowed upon a horse. Rheims may by chance have provided an anecdote that Tennent considered suitable for his purposes, but one wonders if someone with a degree from Rheims was among those whom Tennent perceived as a hostile medical establishment. In 1746, Amson bought the property identified as Lots 212-217, Block 36, 384 BRIT.ORG/SIDA 21(1) City of Willamsburg, County of York, from Thomas Jones, nephew-in-law of the naturalist Mark Catesby. These lots at the intersection of Boundary and Scot- land streets had been so numbered by the Trustees for the Land when the city was laid out. Much later they included the site of “Wheatlands,” the home of the artist Georgia O’Keefe during her late teens. The property included at least one house plus outbuildings when it was acquired by Amson. He owned and resided on this property until his death. About 1751 he purchased from Henry Tyler 180 acres of land just outside the Williamsburg city limits (Stephenson 1961). As well as being a physician Amson appears to have been a planter, or gentleman farmer, as men primarily associated with other enterprises or pro- fessions often were in colonial times. At least as early as 1746 he was an alder- man (Stephenson 1954), and in 1750-1751 he was mayor of Williamsburg (Bodie et al. ca. 1999). In 1758 then-Colonel George Washington had been ill for some time, prob- ably with dysentery, although he was concerned that he might have consump- tion. He traveled from Mount Vernon to Williamsburg “to receive the Advice of the best Physicians,” and there consulted Dr. Amson on 15 March (Fitzpatrick 1931-1944, including a letter from Washington to Colonel John Stanwyx, 5 March 1758). Fitzpatrick, in editing Washington’s papers, commented that Washington “seems to have received the proper treatment, for he was able to be back to his command at Fort Loudon April 5.” Amson may also have treated Daniel Parke Custis, whose widow Washington subsequently married; he was listed among the creditors of Custis’s estate, although Dr. James Carter was re- corded as having been the attending physician during Custis’s last illness (Stephenson 1959; Abbot 1988). A few other references to Dr. John Amson of Williamsburg appear in his- torical sources. He was among the “officials of the government of Virginia” who signed an oath of allegiance to King George II (Virginia...1738-1752), probably having done so upon becoming alderman or mayor. York County records indi- cate that as of 21 September 1747 he was paying taxes on five “tithables,” ie., indentured slaves (Stephenson 1961). His name appears in the account book of the Yorktown merchant William Lightfoot (Lightfoot 1740-1764), and he was one of those who proved the will of Philip Lightfoot in 1748 (Tyler 1894). The jurist John Blair recorded having visited Amson’s gardens 24 March 1751 (Tyler 1899), but wrote nothing about the gardens. Neither Ms. Stephenson nor I have been able to determine the date of Amson’s death, although the well-indexed Virginia Gazette usually published obituaries of prominent Williamsburg residents. His will was proved in the General Court of Virginia, of which the records from the relevant period are not extant (Stephenson 1961). His death evidently occurred after November 1761, when he was listed asa creditor of the estate of John Parke Custis, son of Daniel (Stephenson 1959; Abbot 1988), and before July 1765. Records of land owner- PRINGLE, HISTORY AND EPONYMY OF THE GENUS AMSONIA 385 ship include that of a transaction in July 1765 in which Anne Anderson is said to have inherited the six lots and the buildings thereon in Williamsburg plus the 180 acres outside the city from “the late Dr Amson.” Mrs. Anderson, who was at that time the wife of Robert Anderson of Williamsburg, is thought likely to have been Amson’s daughter, or possibly his widow if she had soon remar- ried, but the relationship has not been determined (Stephenson 1961). I have found no mention of any other individuals with the surname Amson in eigh- teenth-century Williamsburg and vicinity. What may be Amson’s only extant writing therefore appears to have been published posthumously, either from a handwritten heirloom or as a reprint from an earlier publication. This (Amson 1776) is a prescription for the treat- ment of whooping cough, which included several botanical ingredients, in quantities deemed appropriate for a boy of ten. The treatment began with an emetic containing ipecachuana and oxymel of squills (Urginea maritima (L.) Baker, a mild source of cardiac glycosides). This was followed first by chamo- mile tea, then by elixir paregoricum (an opiate) in pennyroyal tea and mint water. The next day the treatment called for gum ammoniac (resin from Dorema ammoniacum D. Don) in pennyroyal tea, mixed with the juice pressed from two hundred woodlice in white wine, with saffron and honey. REFERENCES Assor, WW. (ed.). 1988. The papers of George Washington: Colonial Series. Charlottesville: University Press of Virginia. (Amson mentioned passim in Vols.5 and 6.) Amson, J.1776.Doctor Amson’s prescription for the mee ove eid The palo ina Ga- ette, September 27, 1776, p.6. Facsimile on | Berkevey, E.,and D.S. Berketey. 1963. John Clayton: Pioneer as Ammeriesn Beran Chapel Hill: The University of North Carolina Press. (Material on Amson and Amsonia indexed as being on p. 224 is actually on p. 214.) Berkevey, E.,and D.S. Berkeley. 1982. The life and travels of John Bartram: From Lake Ontario to the River St. John. Tallahassee: University Presses of Florida. Bianton, W.B. 1931. Medicine in Virginia in the Eighteenth Century. Richmond: Garrett & Massie. Boole, L.S., eT AL. (compilers). Ca. 1999. Mayors of Will g.In City of Williamsburg web site: http://www.ci.williamsburg.va.us/history/govh7.htm Britton, N.L., and A. Brown. 1896. An illustrated flora of the northern United States, Canada and the British Possessions ... ,ed. 1. New York: Charles Scribner's Sons. CaLMANN, G. 1977. Ehret: flower painter extraordinary. Oxford: Phaidon Press. FerNato, M.L. 1950. Gray's manual of botany, ed.8. New York: American Book Company. Fitzpatrick, J.C., ed. 1931-1944. The writings of George Washington 1745-1799 from the original manuscript sources. Washington: U.S. Government Printing Office. Adapted for Internet: eon Text Center, University of Virginia Library: http:// etext lib.virginia.edu/washingon/fitzpatrick/ 386 BRIT.ORG/SIDA 21(1) Gray, A. 1856. Manual of the botany of the northern United States, ed. 2. New York: Ivison & Phinney. GRONOVIUS, J.F. i: 739-1743. Flora Virginica exhibens plantas quas V.C. Johannes Clayton in Virginia Observavit atque Collegit. Leiden: Cornelius Haak. Reprinted 1946. Jamaica Plain: Arnold Arboretum. (“Anonymus Suffrutex” 1739.) James, E.W.1895. Libraries of Colonial Virginia (continued).William and Mary College Quart. Hist. Mag., ser. 1, 3:246-253. Reprinted 1966. Same information, from York County Wills and Inventories 18, 1732-1740, pp. 476-477, on Internet: http://pastportal.org/ cwdl_new/archive/probates/html/pi0315.htm Jevuson, R.M. 1963.Dr.John Tennent and the universal specific. Bull. Hist. Med. 37: 336-346. LicHTFoot, W. 1740-1764. Account book. In: William Lightfoot Papers, Manuscript MS 52.4, John D. Rockefeller, Jr, Library, Colonial Williamsburg Foundation, Williams! seen in the present study, at http://ead.lib.virginia.edu/vivaead/published/cw/ vivadoc.pl?file=viwc00065.xml LinbMAN, C.A.M. 1908, 1910.A Linnaean herbarium in the Natural History Museum in Stock- holm. Ark. Bot. 7(3): 1-57; 9(6): 1-50. LINNAEUS, C. 1762-1763. Species plantarum ... , ed. 2. Stock (labernaemontana Amsoania 1762.) Loupon, J.C.1830.Loudon’s hortus britannicus, ed. 1.London:Longman, Orme, Brown, Green, and Longman’s. NicHotson, G. 1884-1887. The illustrated dictionary of gardening ... New York: J. Arnot Penman; London: L. Upcott Gill. (Amsonia 1884.) Paxton, J. 1840.A pocket botanical dictionary, comprising the names, history, and culture of all plants known in Britain..., ed. 1. London: J. Andrews and WS. Orr and Co. Rarinesoue, C.S. 1836-1838. New flora of North America. Philadelphia: Published by the author. Reprinted 1946. Jamaica Plain: Arnold Arboretum. (Ansonia [sic] 1838). Rarinesque, C.S, 1840. Autikon botanikon. Philadelphia:The author.Reprinted 1942 Jamaica Plain: Arnold Arboretum. Rosinson, B.L.,and M.L. Fernatp. 1908. Gray's new manual of botany, ed. 7. New York: Ameri- can Book Company. Smith, J.E. 1819. Tabernaemontana. In: Rees, A.,ed. The cyclopaedia ... 35(I). London: Long- man, Hurst, Rees, Orme, & Brown. STEPHENSON, M.A. 1954, (The) Red Lion historical report, Block 18-1, Building 23A Lot 44. Colonial Williamsburg Foundation Library Research Report Series 1423. Adapted for Internet: http://www.pastportal.com/Archive/research%20Reports/html/rr1423.htm STEPHENSON, M.A. 1959. Custis Square historical report, Block 4, Lot 1-8.Colonial Williamsburg Foundation Library Research Report Series 1070. Adapted for Internet: http:// www.pastportal.com/Archive/Research%20Reports/Html/RR1070.htm STEPHENSON, M.A. 1961. Block 36, Lot 212 & 217 historical report.Colonial Williamsburg Foun- dation Library Research Report Series 1620. Adapted for Internet: http:// www.pastportal.com/Archive/Research%20Reports/Html/RR1620.htm g. (Index nolm: Lars Salvius. PRINGLE, HISTORY AND EPONYMY OF THE GENUS AMSONIA 387 Swan, H.T. (ed.). 1996. English-speaking medical students at continental universities [in the 18th century]: The work archive of R.W.Innes Smith. Typescript in the library of the Royal College of Physicians of Edinburgh. Adapted for Internet: http://www.rcpe.ac.uk/ library/English_Students/Rheims/Rheims_A.html| TENNENT, J. 1738. A memorial, humbly dedicated to the learned, impartial, and judicious world. The Virginia Gazette, Number 112, September 22,1738, pp. 1-2,and Number 113, September 29, 1738, pp. 1-2. Facsimile on Internet: http://pastportal.org/browse Tver, L.G, (ed.). 1894. Lightfoot family (concluded). William and Mary College Quart. Hist. Mag,.,ser.1,3:104-111.Reprinted 1966. Adapted for Internet: http://ftp.rootsweb.com/ pub USYEIIVy b Va sch ols/ HTIETTaly quarterly 03n2/pg104-1 11 TERT: Tyer, L.G. (ed.). 1899. Diary of John Blair [part 1]. William and piney ae pat Hist. maSs ser.1,7: hoa 153.Reprinted 1966. maaeee for | tt ts om/ pub/usgenweb/va/schools/wmmary/blair.txt VIRGINIA. SECRETARY OF STATE. 1738-1752. Oath of allegiance to King George Il of Great Britain, 1738-1752.Last page extant in Library of Congress; photocopy in Manuscripts Collec- tions, Virginia Historical Society. Adapted for Internet: http://vhs3.vahistorical.org/cgi- bin/starfinder/16400/x.marc.txt Watter, T. 1788. Flora Caroliniana ... London: J. Fraser. Reprinted 1946. Jamaica Plain: Ar- nold Arboretum. Weesite. Accessed 22 Oct 2003. http://www.familysearch.org/Eng/Search/ frameset_search.asp (Latter-Day Saints genealogical records). Wepsite. Accessed 22 Oct 2003. http://linnaeus.nrm.se/botany/fbo/t/taber/ tabeams.html.en (specimen of Tabernaemontana amsonia at S-LINN). Weesite. Accessed 22 Oct 2003. http://linnaeus.nrm.se/botany/fbo/hand/welcome.html.en (examples of handwriting at S-LINN). Woop, A. 1861.A class-book of botany ... ,ed.s.n. New York:A.S. Barnes & Co. Woopson,R.E., Jr. 1928. Studies in the Apocynaceae.Ill.A monograph of the genus Amsonia. Annals Missouri Bot. Gard. 15: 379-434, plates 50-53. 388 BRIT.ORG/SIDA 21(1) BOOK NOTICES Timber Press Guy STERNBERG and JIM WILSON. 2004. Native Trees for North American Landscapes. (ISBN 0-88192-607-8, hbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $59.95, 552 pp., 513 color photos, 96 line drawings and | map, 8!/2" x 11". This is a book of detailed information on ae ee an native trees iO landscapes and gardens. The authors say they exclusively fo ation of native trees. Most of — the book is devoted to what the aoe call the “Menu of Native Trees. Basically that is the ninety- six trees which are a feat es Specie Each iene oe Ww oe a silhouette of the tree and a leaf, desc ription lea Jest seaso culture proble ems, simi- fo} lar and cee species, and comments. Oh, ie not forget the very nice color photographs that com- pliment each featured species.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 761202-4060, U.S.A. THomas Hopss. 2004. The Jewel Box Garden. (ISBN 0-88192-66-9, hbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, |- 800-327-5680, 503-227-3070 fax). $34.95, 176 pp., 160 color photos, 10" x 10". ‘he Jewel Box Garden is full of exciting plants, gardens, and garden designs in beautiful color. The scans is by David McDonald are a sight for sore eyes. What else can one say? This is one pretty book that any gardener would love to have. Just one look at this gem and you will be hard pressed not to want to use some of the ideas in the book.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 761202-4060, U.S.A. WILLIAM T. STEARN. 2004. Botanical Latin. (ISBN 0-88192-627-2, pbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327- 5680, 503-227-3070 fax). $29.95, 560 pp., 42 line drawings, 6 3/4" x 9 3/4". Botanical Latin was first published in 1966 and this is the first paperback edition. Joser J. Hapa and James W. Wappick (Botanical Illustrations by Jarmila Haldova). 2004. The Genus Paeonia. (ISBN 0-88192-612-4, hbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, |-800-327- 5680, 503-227-3070 fax). $34.95, 228 pp., 36 watercolors and 43 line draw- ings, 81/2" x 11". Review forthcoming in volume 21, no. 2. SIDA 21(1): 388. 2004 CARBON USE BY THE ENDANGERED TEXAS WILD RICE (ZIZANIA TEXANA, POACEAE) Paula Power Robert D. Doyle U.S. Fish and Wildlife Service Baylor University National Fish Hatchery and Technology Center Department of Biology 500 E. McCarty Lane PO. Box 97388 San Marcos, Texas 78666, U.S.A. Waco, lexas 76798, U.S.A. ABSTRACT A pH drift experiment was conducted to dete rmine a HCO, uptake potential of Texas wild rice (Zizania texana Hitchc.). In 15 separate a tora inorganic carbon toalkalinity ratio varied between 0.93-0.99 and acl a v Aiea between : ae 9.21, strongly suggesting Texas wild ric is an obligate CO, plant unable to utilize HCO,. The estimated CO, compensation point for a es wild rice is 2 um ol a from this study provide pl logical evid for th iti lationshi rent ie and biomass pied ity y observ ed in Texas wild ri rice Ca 1996a, 1996b, 2000; Poole & Bowles 1999; Saunders et al. 2001) and provides water resource managers with information they need to manage and protect the San Marcos River ecosystem. RESUMEN Un experimento de la variacion del pH fue utilizado para determinar el potencial de absorcion de HCO, del arroz salvaje de Tejas (Zizania texana GhiCne ). En 15 experimentos separados, la relacion entre “i carbono inorganico y la alcalinidad ntre 0,93—0,99 y el pH final vario entre 8,54—9,21, sugiriendo que el arroz salvaje de Tejas es una een Co, ce incapaz de utilizar el HCO,. El punto estimado de compensacion del CO, en el arroz salve raje de Tejas es 2 umol |. s datos de este estudio proporcionan la evidencia fisioldgica para la relaci6n positiva entre la velocidad actual y la produccion de la biomasa observadas en el arroz salvaje de Tejas (Power 1996a, 1996b, 2002; Poole zi) Pawiee 1999, Saunders et al. 2001) y ofrecen a los encargados de los recursos hidricos la inform nmanejar y proteger el ecosistema del rio San Marcos. INTRODUCTION The San Marcos River arises from springs within the City of San Marcos, Hays County, Texas and flows 8.1 river km ina southeasterly direction before it joins the Blanco River. The source aquifer for the San Marcos River is the Edwards Aquifer, the sole source of municipal, agricultural, and industrial water for 1.5 million people in central Texas. Water is pumped continuously from the Ed- wards Aquifer for human use, but is recharged only during rain events. As wa- ter is pumped during drought periods when recharge is minimal, the level of the aquifer drops, threatening spring flow to the San Marcos River and other local spring systems. The spring-fed San Marcos River provides habitat for five federally listed species: fountain darter (Etheostoma fonticola), San Marcos gambusia (Gambusia georgei), San Marcos salamander (Eurycea nana), Texas SIDA 21(1): 389 — 398. 2004 390 BRIT.ORG/SIDA 21(1 — — slind salamander (Typhlomolge rath buni),and Texas wild rice (Zizania texana). The primary threat to each species is reduced spring flow from the source aqui- fer (U.S. Fish and Wildlife Service 1996). The threat became critical during a drought event in 1990 and all five species were the focus of an endangered spe- cies lawsuit initiated in 199] (Sierra Club v Lujan, No. MO-91-CA-69). One out- come of the lawsuit was creation of a water authority to manage the aquifer for human use and to protect the San Marcos River ecosystem for endangered spe- cies by maintaining adequate spring flow. Defining “adequate” spring flow fora species is a monumental challenge met by research systematically addressing ecological and physiological aspects of the species. The target species for this study was Texas wild rice. This species is en- demic to the San Marcos River, TX and was listed as endangered in 1978 (U.S. Fish and Wildlife Service 1985). Texas wild rice is a perennial, submersed mac- rophyte now commonly found in swiftly flowing water. Under these condi- tions, it produces long, ribbon-like, submersed leaves and reproductive culms. Culms can become emergent and produce wind pollinated, terminal inflores- cences. Nodes along each culm give rise to roots and asexual clones, called tillers. A thorough description of the species and its habitat is provided by Terrell et al. (1978), US. Fish and Wildlife Service (1995), Poole and Bowles (1999), and Saun- ders et al. (2001). Texas wild rice has two distinct phenotypes under wild and cultured con- ditions. Wild plants found in flowing water (>0.46 ms!) in the San Marcos River are perennial and primarily submersed, although reproductive culms are present. Under cultivated conditions in slow moving water (<0.05ms"}), Texas wild rice has low vegetative productivity compared with study plants grown in faster flowing water ranging from 0.29-0.49 ms! (Power 1996a, 2002). Texas wild rice in slow moving water ranging from 0.001-0.12 ms7 is primarily emer- gent, sexually reproductive, and annual to short-lived as is common for other North American Zizania species (Terrell et al. 1978; Power 1996a, 2002). Prima- rily emergent plants with low biomass productivity are especially vulnerable to herbivory because of the potential for loss of reproductive culms prior to seed set and the ultimate loss of the plant because of a shortened life span. Carbon utilization by macrophytes has been shown to influence photo- synthesis in experimental studies (Smith & Walker 1980) and may be an im- portant factor influencing phenotypic variation in Texas wild rice. The rela- tive concentrations of carbon species dissolved in water are influenced by pH. At lower pH propor tionally more CO? is available; as pH rises, equilibrium shifts to HCO3, then CO3~. Macrophytes utilize inorganic carbon in the form of CO2 for photosynthesis and all submersed species are able to extract this form of dissolved inorganic carbon from the water when it is available. In addition, some submersed species show the ability to extract efficiently HCO3 from the water while others exhibit little or no ability to do so (Alen & Spence 1981; Maberly POWER AND DOYLE, CARBON USE BY THE ENDANGERED TEXAS WILD RICE 391 & Spence 1983; Sand-Jensen 1983; Spence & Maberly 1985). In addition to the absolute concentration of dissolved inorganic carbon, the flow environment of the submersed leaves also influences their ability to extract carbon from the water. Flowing water reduces the thickness of the boundary layer surrounding leaves through which gases move predominantly by diffusion. Slower diffusion of CO2 across the boundary mee can limit photosynthesis in still or very slow moving water (Smith &@ Walker | Differential carbon ie combined with carbon uptake potential may contribute to phenotypic variation in Texas wild rice and provide a physiologi- cal explanation for its preference for a flowing water habitat. The purpose of this study was to determine the bicarbonate uptake potential of Texas wild rice. METHODS AND MATERIALS Carbon use by Texas wild rice was examined using plants taken from culture after seedlings had been growing about three months in an outdoor cement raceway. The raceway was supplied with water pumped from the Edwards Aqui- fer and with chemistry similar to that of the San Marcos River. The plants had 4-6 leaves, each of which was 20-30 cm in length. The plants were maintained in aerated river water (4.5 meq! alkalinity) under artificial illumination (250 umol ms! PAR; 14:10 light/dark photoperiod) at laboratory temperature (22° C) until utilized. All experiments were conducted within four days of arrival of plant material at the lab. For comparison data are also presented for Hydrilla verticillata, a sub- mersed aquatic species known to be well adapted for extracting bicarbonate from water due to its unique, C-4 like photosynthetic pathway (Holaday & Bowes 1980). Hydrilla verticillata apical stem fragments were collected from culture ponds at the Lewisville Aquatic Ecosystem Research Facility, Lewisville, ne The pH drift technique described by Allen and Spence (1981) was used to determine the HCO3 uptake potential of Texas wild rice. This method involves continuous measurement of pH ina solution bathing actively photosynthesiz- ing plants in a closed system at constant alkalinity (ALK). The method relies on shifts in abundance of the three species of dissolved inorganic carbon (CO3, HCO3, CO3*) with pH. In a closed system, plants will continue to photosyn- thesize until inorganic carbon can no longer be extracted from the bathing so- lution. When photosynthesis ceases, the final pH of the solution is dependent on the concentration of total dissolved inorganic carbon (Cy) in solution which depends upon the carbon-extracting potential of the plant. The pH endpoint of these experiments varies with a plant’s ability to uti- lize HCO3 and on the alkalinity of the solution. Carbon dioxide obligate plants (those which cannot utilize HCO3 ) cannot shift the pH of the bathing solution past the point where free CO? disappears from solution (pH ca. 9.2 depending 392 BRIT.ORG/SIDA 21(1) on alkalinity). The CO? content of the solution at the pH where photosynthesis ceases is considered the CO2 compensation point for that species. Removal of CO? from the water results in increased pH, but has little impact on Cy, and does not impact ALK. Other aquatic plants are also able to utilize HCO3 asa source of inorganic carbon. Ina closed system, these plants will shift the pH well above the thresh- old where CO) disappears. In this case, the final pH of the solution depends on the total Cr pool available as well as the affinity of the plant for HCO3. The HCO3 content of the solution at the pH where photosynthesis ceases is con- sidered the HCO3 compensation point for that species. The removal of HCO3" from solution has a stronger impact on Cy but still does not impact ALk, since the buffering capacity lost by the consumed HCO; is balanced by the produc- tion of OH”. The Cr:ack ratio at the end of a pH drift experiment is considered a relative measure of a plant's ability to utilize HCO3 (Maberly and Spence 1983). The Cr:aAk ratio of CO2-obligate plants is very close to 1.0, while the ratio of HCO3 users is well below 1.0. Oxygen and pH were continuously measured in a re-circulating closed system. The experimental apparatus consisted of a glass incubation chamber, an acrylic probe block, and a submersible centrifugal pump, all connected by gas tight tubes of butyl rubber (total volume 400 ml) and submerged in a tem- perature-controlled water bath (20° C). The incubation chamber was a glass cylinder 5 cm in diameter x 30 cm long. A glass tube connector was fused to one end, while a ground-glass fitting with another glass tube connector was fused to the opposite end to allow access to the cylinder for plant tissue inser- tion and removal. Water was re-circulated at an approximate velocity of 2 cm s! through the incubation chamber and was sufficient to remove diffusive limi- tations of photosynthesis. The acrylic sensor mounting block allowed both pH and O2 sensors to be screwed in so that the electrode tips were continually bathed by the re-circulating solution. The pH electrode (Ross combination elec- trode) had a sensitivity of 0.01 pH units and was connected toa pH transmitter (VJenco 629). The fast-response, low-consumption Clark-type polarographic oxy- gen sensor (YSI 5331) was connected to an oxygen meter (Cameron Instrument Corp). The oxygen electrodes were calibrated by submersing the probe in No- sparged water and fully aerated water at incubation temperature. pH sensors were calibrated prior to use utilizing pH 7.0 and 10.0 buffers. Outputs from the pH transmitter and the oxygen meter were continuously monitored by a com- puter equipped with data acquisition software (Strawberry Tree Workbench PC). The sensors were monitored continuously and data recorded every 3 min- utes. Calibrations did not drift over the time-scale of the experiments. Alkalin- ity of the bathing solutions were determined by titration with dilute hydro- chloric acid (APHA 1992). POWER AND DOYLE, CARBON USE BY THE ENDANGERED TEXAS WILD RICE 393 Entire Texas wild rice rosettes consisting of 3-6 leaves (0.2-0.3 gdw) were used in the experiments. Light was provided by specialized fluorescent aquaria lights (6700 K) providing saturating light intensities (500 pmol m~? s! PAR). The bathing solutions used for the experiments were either San Marcos River water (Alk = 4.5-5.0 meq!!),a general purpose culture solution (Alk = 0.82 meq ||. Smart and Barko 1985), or intermediate mixtures of the two. Prior to the start of the experiment, the incubation chambers were flushed with a bathing solu- tion which had been bubbled with a gas mixture composed of 350 ppm CO? (atmospheric concentration) and 6% O2 (approximately 30% atmospheric con- centration) in nitrogen. This reduced the O2 concentration to about 75 umol |! while maintaining normal dissolved inorganic carbon and pH levels of the bath- ing solutions. Lowering the initial O2 concentration allows photosynthesis to proceed for extended periods within the closed system without potential com- plications due to O2 supersaturation which increases photorespiration. Photosynthesis was measured as the rate of O2 evolution in the closed sys- tem through time. Corrections were made for total volume of the incubation apparatus, and the results normalized to plant dry weight. Experiments were allowed to proceed until photosynthesis stopped and the pH stabilized. This usually consisted of 2 to 3 hour runs, although longer runs up to 12 hours were made to verify that steady state conditions were achieved in the shorter runs. Concentrations of different carbon species as a function of pH and tem- perature were estimated from equilibrium constants and formulae in Stumm and Morgan (1981). RESULTS Table | shows the results of 15 independent pH drift experi s conducted on Texas wild rice and three experiments conducted on H. verticellata. Alkalinity was measured prior to the experiment for each bathing solution as was the fi- nal pH at the apparent photosynthetic compensation point. Total carbon and CO? were estimated based on these data. The Cr:atk ratio for all experiments was very close to 1.0 and varied between 0.93-0.99, strongly suggesting that this species is a CO? obligate plant which is unable to utilize HCO3 at a rate sufficient to keep up with photosynthetic needs (see Maberly and Spence 1983). For CO2-obligates, the CO2 concentration at the end of the pH drift experiment represents a conservative estimate of the CO compensation point for the spe- cies. In this series of experiments, the values varied from 1-13 umol I! Rates of photosynthesis as a function of pH for two Cr concentrations are shown in Figure 1A. As expected, the rates of photosynthesis at a given pH are higher for the higher Cy solution. Since pH controls the proportion of Cy present as COp, the level of CO2 present in the two solutions was actually very different. However, both data sets show a linear and strong reduction in photosynthesis as the pH of the solution approaches 9.0. In Fig. 1B these data are re-plotted to 394 BRIT.ORG/SIDA 21(1) Taste 1.Summary of pH drift experiments for Texas wild rice and Hydrilla verticillata conducted in closed systems including alkalinity (Alk) of bathing solution, final pH, final concentration of total inorganic carbon (C,),CO, remaining at the end of the experiment and the ratio of C_/Alk. Run ID Alkalinity Final pH G (0, G/Alk (meq!') (mmol I") (umol I") Texas wild rice (San Marcos River) | 8.54 1.225 8 0.99 2 1.24 8.66 1.216 6 0.98 3 1.24 8.76 1.207 5 0.97 4 2.00 8.54 1.978 13 0.99 5 2.00 8.72 1.956 8 0.98 6 2.00 9.14 1.876 3 0.94 7 2.16 9.21 2.006 3 0.93 8 2.04 9.16 1.908 3 0.94 9 2.04 9.14 1.914 3 0.94 10 0.82 8.8] 0.796 3 0.97 11 0.82 8.72 0.801 4 0.98 12 0.83 9.14 0.778 | 0.93 13 2.45 8.80 2.391 9 0.98 14 2.34 8.63 2.308 13 0.99 15 2.40 9.18 2.254 3 0.94 Hydrilla verticillata (pond culture) ] 10.43 0.377 0.01 0.44 2 0.85 10.36 0.414 0.02 0.49 3 0.85 10.23 0.477 0.04 0.56 show the relationship of photosynthesis to actual CO levels. The linear nature of this curve at low CO> is characteristic of CO2-obligate plants and the ex- trapolated X-axis intercept of 2 moll! providesa reliable estimate of the CO2- compensation point of photosynthesis for this species (Sand-Jensen 1983). DISCUSSION The experimental method used in this study identifies the upper pH limit for positive net photosynthesis. Net photosynthesis by Texas wild rice approached zero as pH approached 8.7 when CO) was 0.5% of total dissolved inorganic car- bon. Bicarbonate in the bathing solution at pH 8.7 was 97.5% of total dissolved inorganic carbon. At this pH there was abundant HCO3° available for plant uptake. If Texas wild rice had the ability to remove HCO3 rapidly, positive net photosynthesis would continue alter CO? was exhausted, driving pH above 9.2. During this study, no photosynthesis occurred above 8.7 suggesting submersed leaves of Texas wild rice are unable to take up HCO3 . Carbon dioxide availability to submersed aquatic plants is a function of the interplay between water current velocity, leaf boundary layer thickness and POWER AND DOYLE, CARBON USE BY THE ENDANGERED TEXAS WILD RICE 395 0.30 ———______ ee A e@ C.=4.4mM | eal . A C,=0.8 mM | | 6202 e “=~ 015-4 = e6 e ‘m 0.10 4 @e O ° % (52 a & S rs A OB ps Bg 2 A ds & 0.00 | | Ba = 8.0 8.2 8.4 8.6 8.8 9.0 L pH © ~ 0.30 ” ® 0.25- B * — e @20 = a ee 2 o15 e . ated = CO, compensation i? Oo 0.10 - o° point=2uM 4 fan 0.05 4 me 0.00 - a , , | | | 0.00 0.01 0.02 003 004 005 0.06 £0.07 CO, (mM) Fic. 1. Apparent net piatgs then rate yy Texas it rice ina donee water- -flow ea a ale ae up CO, for photosynthesis driving gupp I pproximately 8.7. B) Estimated CO, compensa tion point for Texas wild rice is 2 umol 1", concentration gradient between surrounding water and leaf surface; as flow increases, boundary layer thickness decreases and CO? availability increases. Using a closed water-flow system Madsen and Sondergaard (1983) found ap- parent photosynthesis was stimulated by increasing velocities and a maximum rate was reached at 0.008-0.0012 ms! In contrast, Madsen et al. (1993) found 396 BRIT.ORG/SIDA 21(1) photosynthetic rates decreased with increasing current velocity and suggested that physical stress imposed on plants by agitation or stretching in flowing water was a key factor in the observed response. Ribbon-like submersed leaves of Texas wild rice are adapted for flowing water and can reduce carbon limita- tion by exploiting the flowing water habitat where boundary layer surround- ing leaves and diffusion distances for CO2 are reduced and leaves are continu- ally bathed with carbon-rich water. In contrast, in slower moving water, photosynthesis by submersed leaves of Texas wild rice are probably carbon lim- ited and few submersed leaves are produced, a greater proportion of biomass is allocated to reproductive parts, and plants are less productive overall (Power 2002). Emergent reproductive culms most likely are not carbon limited because culms obtain CO? from the atmosphere where CO2 is more readily available owing to the higher diffusion rate and current velocity in air relative to water (Madsen & Sand-Jensen 1991; Denny 1993). Texas wild rice is one of five endangered or threatened species which oc- cur in the San Marcos River. As the result of a lawsuit involving the listed spe- cies, a water authority was created by the Texas Legislature to manage the source aquifer for the San Marcos River (Texas Legislature 1993). Flow requirements lor Texas wild rice are important because of the legal requirement to manage the source aquifer for human needs while simultaneously protecting aquatic and wildlife habitat and protecting listed species. The potentially conflicting management goals of providing aquifer water for human use while maintain- ing adequate spring flow for endangered species cannot be attained without information concerning the habitat requirements for the listed species. Numer- ous surveys have identified flowing water habitat as a requirement for Texas wild rice and experimental studies have consistently shown a positive relation- ship between current velocity and productivity as well as biomass allocation tosubmersed plant parts (Power 1996a, 1996b, 2002; Poole & Bowles 1999: Saun- ders et al 2001). Data from this study provide a physiological explanation for these observations and furthers our understanding of habitat requirements for Texas wild rice. ACKNOWLEDGMENTS This research was supported by a Section 6 grant from US. Fish and Wildlife Service and Texas Parks and Wildlife. We would like to thank Kathryn Kennedy, Center for Plant Conservation; Dr. Michael Smart, Lewisville Aquatic Ecosys- tem Research Facility; and Stephen McClintik for technical support. REFERENCES Aten, E.D. and D.H.N. Spence. 1981. The differential ability of aquatic plants to utilize the inorganic carbon supply in fresh waters. New Phytol. 87:269-283. POWER AND DOYLE, CARBON USE BY THE ENDANGERED TEXAS WILD RICE 397 APHA. 1992. Standard methods for examination of water and wastewater. 18" edition. A.E. Greensger, L.S. Clesceri and A.D. Eaton, eds. Amer.Water Works Assoc. and Wat. Pol- lution Control Federation, Washington, D.C. BLANCH S.J.,G.G. Gane, and K.F. Wa ker. 1999. Growth and resource allocation in response to flooding in the emergent sedge Bolboschoenus medianus. Aquatic Bot.63:145-160. Denny, M.W. 1993. Air and water: The biology and physics of life’s media. Princeton Univer- sity Press. Ibestam-Atmouist, J.and L. Kautsky. 1995. Plastic responses in morphology of Potamogeton pectinatus L.to sediment and above-sediment conditions at two sites in the northern Baltic proper. Aquatic Bot. 52:205-216. Hotapay, A.S.and G. Bowes. 1980.C, acid metabolism and dark CO; fixation in a submersed aquatic macrophyte (Hydrilla verticellata). Plant Physiol.65:331-335. Maberly, $.C.and D.H.N. Spence. 1983. Photosynthetic inorganic carbon use by freshwa- ter plants. J. Ecology 71:705-724. Maosen, T.V., H.O. ENevotosen, and T.B. Jarcensen. 1993. Effects of water velocity on photosyn- thesis and dark respiration in submerged stream macrophytes. PI., Cell and Environm. 16:317-322. Maosen, T.V. and K. SAND-JENSEN. 1991. Photosynthetic carbon assimilation in aquatic mac- rophytes. Aquatic Bot. 41:5—40. Mapsen, T.V.and M. Sonpercaard. 1983.The effects of current velocity on the photosynthe- sis of Callitriche stagnalis Scop. Aquatic Bot. 15:187-193. Poote, J.and D.E. Bowtes. 1999. Habitat characterization of Texas wild-rice (Texas wild rice Hitchcock), an endangered aquatic macrophyte from the San Marcos River, TX, USA. Aquatic Conservation: Mar. and Freshwater Ecosystems 9:291-302. Power, P. 1996a. Effects of current velocity and substrate composition on growth of Texas wildrice (Zizania texana). Aquatic Bot. 55:199-204. Power, P. 1996b. Reintroduction of Texas wildrice (Zizania texana) in Spring Lake: Some important environmental and biotic considerations. In: Proceedings from Second Southwestern Rare and Endangered Plant Conference: Rocky Mountain Forest & Range Experimental Station. Power, P. 2002. Resource allocation patterns and p rent velocity in the endangered Texas wildrice (Zizania texana Hitchc.). Sida. 20:571- cisee Ressoorr, A. 1972. The carbon dioxide system of freshwater. A set of tables for easy com- putation of the carbon dioxide system. Printed booklet. Freshwater Biological Labora- yenotypic plasticity in relation to cur- tory, Hillerod, Denmark SAND-JENSEN, K. 1983. Photosynthetic carbon sources of stream macrophytes. J. Exp. Bot. 34:198-210. SAUNDERS, K.S., K.B. Mayes, T.A. JURGENSEN, J.F. TRUNGALE, L.J. KLEINSASSER, K. Azziz, J.R. Fietos, and R.E. Moss.2001.An evaluation of spring flows to support the upper San Marcos River spring ecosystem, Hays County, Texas. River Studies Report No. 16. 398 BRIT.ORG/SIDA 21(1) Smart, R.M.and J.W.Barko. 1985.Laboratory culture of submersed freshwater macrophytes on natural sediments. Aquatic Bot. 21:21-263. Smith, F.A and N.A.Watker 1980. Photosynthesis by aquatic plants: effects of unstirred lay- ers in relation to assimilation of CO2 and HCO3 and to carbon isotopic discrimination. New Phytol. 86:245-259. Stumm, W. and J.J. Morcan. 1981. Aquatic chemistry: an introduction emphasizing chemi- cal equilibia in natural waters. Wiley, New York. Terre, E.E,W.P.H. Emery,and H.E. Beaty. 1978.Observations on Zizania texana (Texas wildrice) an endangered species. Bull. Torrey Bot. Club 105:50-57. Texas Lecistature. 1993. Chapter 626, S.B. No. 1477. 73" Legislature-Regular Session. THomas, A.G. and J.M. Stewart. 1969. The effect of different water depths on the growth of wild rice. Canad. J. Bot.47:1525-1531. U.S. FisH AND Wicovire Service. 1985. Endangered and threatened wildlife and plant taxa for listing as endangered or threatened species. Fed. Reg. 5C(188):39526-39584, U.S. FisH AND WILDLIFE Service. 1996. San Marcos/Comal (Revised) recovery plan. Albuquer- que, New Mexico. Y] THE POST OAK SAVANNA ECOREGION: A FLORISTIC ASSESSMENT OF ITS UNIQUENESS Michael H.MacRoberts and Barbara R. MacRoberts Bog Research 740 Columbia Shreveport, Louisiana 71104, U.S.A. and Herbarium um of Life Sciences Louisiana State University-Shreveport Shreveport, Louisiana 71115, U.S.A. ABSTRACT Q itati hod he f] { the Post Oak Savanna and Oak-Pine-Hickory ecoregions of the West Gulf Coastal Plain, The analyses show that the two areas are nearly identical floristically. Key Worps: Post Oak Savanna ecoregion, Oak-Pine-Hickory ecoregion, biogeography, West Gulf Coastal Plain RESUMEN Se usaron métodos cuantitativos para comparar la flora de las ecoregiones Post Oak Savanna y Oak- Pine-Hickory de la llanura costera del West Gulf. Los analisis muestran que las dos areas son casi idénticas floristicamente. INTRODUCTION A half century ago, Webb (1950) examined the methodology of biogeographers who worked in Oklahoma and Texas and found that not only they did not agree among themselves on the location and description of biogeographic regions, but that regional delineation was subjective. He suggested a remedy: “Quanti- tative methods must be developed before general agreement on the extent of the major [ecoregions] can be reached. As long as personal opinion and indi- vidual judgment are the only bases for judging the extent of the [ecoregionsl, fundamental and insoluble differences of opinion will continue” (Webb 1950:246). This situation remains essentially unchanged today (MacRoberts & MacRoberts 2003a). The purpose of this paper is to describe the botanical simi- larities and differences between the Post Oak Savanna (POS) region or ecoregion and the adjacent Oak-Pine-Hickory (OPH) region or ecoregion (herein region, ecoregion, and vegetation area are used interchangeably). STUDY AREAS The Post Oak Savanna region (POS) of Texas has been considered a distinct veg- SIDA 21(1): 399 — 407. 2004 400 BRIT.ORG/SIDA 21(1) etation area (ecoregion) since Harshberger (1911) mapped it, but it was Shantz and Zon (1924) and later Gould (1962) who gave it its modern shape (MacRoberts & MacRoberts 2003a) (Fig. LD. The POS region has been described repeatedly. It isa gently rolling or hilly region comprising 30,000 to +0,000 sq. km with elevations from 90 to 250 m above sea level. Annual rainfall is 75 to 115 cm with highest rainfall in May and June. The Carrizo Sands extend along the length of the region. Soils on the up- lands are acid sandy loams or sands. Bottomland soils are generally acid sandy loams and clays. Prairies are scattered throughout, notably in the south. The area contains a diversity of plant communities, from hillside pitcher plant bogs, peat bogs, and upland marshes to open xeric sandylands, oak-hickory forests/ woodlands, prairies, and bottomland floodplains. Overstory trees in the uplands are primarily Quercus stellata Wang,, Q. margaretta (Ashe) Ashe ex Small, 9. marilandica Muenchh., and Carya spp. The virtual absence of pine is empha- sized. Understory vegetation includes common grasses Schizachyrium scoparium (Michx.) Nash , Sorgastrum nutans (L.) Nash, Panicum virgatum L., Tridens flavus (L.) Hitche., Bothriochloa saccharoides (Sw.) Rydb., Nassella leucotricha Trin.& Rupr.) Pohl, Chasmanthium sessilifolium (Poir.) Yates. Fur- ther description of the POS region can be found in many sources (e.g., Correll & Johnston 1970; LBJ School of Public Affairs!978; Hatch et al. 1990: Telfair 1999). McBryde (1933) conducted the first major floristic study of the POS region, emphasizing the Carrizo Sands. Recent studies have been on grasslands (Smeins & Diamond 1983), bogs and marshes (MacRoberts & MacRoberts 1998, 2001), creek systems (Telfair 1988), and xeric sandylands (MacRoberts et al. 2002a). What has not been studied is the floristic similarities and differences between the POS region and its adjacent “ecoregions” or “vegetation areas” (MacRoberts & MacRoberts 2003a). The OPH region, to which we mainly compare the POS region, consists of about 175,000 sq. km in east Texas, west Louisiana, south Arkansas, and south- east Oklahoma. It is characterized by the presence of pine and by the frequent co-dominance of pine, oak, and hickory in the forest or woodland canopy. Pre- cipitation is about 100 cm per year. The terrain varies from nearly level to gen- tly undulating. Upland soils are generally acidic sandy loams and sands over sandy loam and clay. Bottomlands are acidic sands and clays and loams. The dominant vegetation is a mixed pine-hardwood forest on the uplands and a mixed hardwood forest on the bottomlands. Pinus taeda L. and P. echinata Mill. occur throughout. Pinus palustris Mill. is confined to the southern part of the OPH region. Principle hardwoods are those that occur over most of the south- east coastal plain: Liquidambar styraciflua L., Quercus spp., Nyssa sylvatica Marsh., Magnolia spp., Ulmus spp., Populus deltoides Bart. ex Marsh., Carya spp., Acer spp., Fagus grandifolia Ehrh., Fraxinus spp., Taxodium distichum (L.) L. Rich. Further description of the OPH region can be found in many sources (e.g., — MACROBERTS AND MACROBERTS, THE POST OAK SAVANNA ECOREGION 401 a naa ee Ke cin ae Fie at 2 ee AL Pas rg ei i. \_% : ae ae Sah 2 tate E > rane Gp tiene aoe 7 i x r ~ a om ie ra we AY Ce \] tea, : m = Z 7 : wt we (he | HH Pp aN f+ eft ten see Correll & Johnston 1970; Hatch et al. 1990; Telfair 1999 and references con- tained therein). The longleaf pine portion of the OPH region (often considered to be a separate ecoregion itself, is not so considered here) in southeast Texas and south-central Louisiana has been by far the best studied portion of the OPH region (see Harcombe et al. 1993; MacRoberts & MacRoberts 2003a for literature). 402 BRIT.ORG/SIDA 21(1) METHODS 1. In order to determine the uniqueness of the POS region, we examined the distribution of the taxa on the Singhurst et al. (2003) list of the flora of the 4,466 ha Gus Engeling Wildlife Management Area (GEWMA) in Anderson County, in the middle of the POS region. This sample contains 830 native taxa from a wide variety of communities and habitats including bogs, marshes, xe- ric sandylands, stream bottoms, and oak-hickory woodlands. Using all avail- able sources (e.g., Hatch et al. 1990; Thomas & Allen 1993-1998; Turner et al. 2003; Flora of North America; Evans n.d.), we determined how many of the taxa on the Singhurst et al (2003) list occur in the OPH region and/or in the coastal region of southeast Texas and Louisiana and how many of them are either con- fined to ‘the POS region or to that region and areas westward. ft idereda key indicator of ecoregional unique- ness (Tahktajan 1986), we studied the distribution of endemic West Gulf Coastal Plain taxa to see how many are unique to the POS region (see MacRoberts et al. 2002b for West Gulf Coastal Plain endemics and their community affiliations), 3. In order to discover the North American affinities of the POS region, us- ing Kartesz and Meacham (1999) we plotted the North American distribution by state or regional area of the 830 native taxa on the Singhurst et al. (2003) GEWMA list. We converted the results into percentage of taxa that occur in each state or regional area (see MacRoberts & MacRoberts 2003b for details of this method). RESULTS 1. Ninety-eight percent of the native taxa found at GEWMaA also occur in the OPH and/or coastal plain regions of Texas and Louisiana. Only two percent are either endemic to the POS region or reach their eastern limit in the POS region. 2. Of the approximately 100 endemic taxa of the West Gulf Coastal Plain, only nine are endemic or near endemic to the POS region. These are Abronia macrocarpa L. Galloway, Brazoria truncata (Benth.) Engelm. & Gray, Hymenopappus carrizoanus B.L. Turner, Lactuca hirsuta Muhl. ex Nutt. var. albiflora (Torr. & Gray) Shinners, Monarda viridissima Correll, Paronychia setacea Torr. & Gray, Polygonella parksii Cory, Rhododon ciliatus (Benth.) Eping, and Valerianella florifera Shinners. All of these taxa are associated witha single plant community: xeric sandylands (MacRoberts et al. 2002b). This commu- nity accounts for fifty percent of the endemics in the West Gulf Coastal Plain and is associated with the Carrizo Sands and other upland sandy areas and ter- races that run the length of the POS region and spread out across the OPH re- gion (McBryde 1933; MacRoberts & MacRoberts 1994, 1995, 1996; MacRoberts et al. 2002a). The remaining West Gulf Coastal Plain endemics occur in either both the POS region and the OPH region or in the OPH region alone. MACROBERTS AND MACROBERTS, THE POST OAK SAVANNA ECOREGION 403 3. The GEWMA flora is overwhelmingly eastern and notably southeast- ern (Fig. 2). Ninety-five percent of the taxa occurring at GEWMA occur in Loui- siana, 92 percent in Arkansas, 89 percent in Oklahoma, 85 percent in Missis- sippi, 81 percent in the Carolinas, and 53 percent in New York and the New England states, but only 28 percent in New Mexico. This is consistent with our previous finding that eastern flora characterizes east Texas to about 95 degrees W. longitude, at which point east begins to give way to west over the next 300 km (MacRoberts & MacRoberts 2003b). GEWMaA is located at about 96 degrees W. longitude on the eastern edge of this broad transition zone, and is thus es- sentially eastern, as the above figures indicate. DISCUSSION Unfortunately, for our floristic analysis there is no operational definition of ecoregion or any of the other “chorionomic” categories (e.g., region, province, district, subdistrict) that biogeographers use (Takhtajan 1986). The best defini- tion we have found for ecoregion is that it is a distinct assemblage of natural communities and species involving a fairly large geographical region (Ricketts & Dinerstein 1999; The Nature Conservancy 2003). Takhtajan (1986) also states that endemism is fairly high, notably at the species and subspecies level. But these statements are of little help since they do not designate whether 25% or 50% difference in species composition constitutes a distinct assemblage or just what percent of the total should be endemic. Nonetheless, our main finding is that the POS region differs only insignifi- cantly from the OPH region. Far fewer than one percent of its taxa are endemic to it and only about two percent of its taxa do not also occur in the adjacent region to its east. As part of our floristic study of the POS region, we examined and com- pared all community classifications for the POS and OPH region to see how many communities described for the POS region were unique to it (Marks & Harcombe 1981; Diamond et al. 1987; Harcombe et al. 1993; Texas Natural Heri- tage Program 1995; Nesom et al. 1997; Turner et al. 1999; Van Kley 1999a, 1999b; Bezanson 2000; Fleming et al. 2002; Louisiana Natural Heritage 2003; Singhurst et al. 2003). While this was not a quantitative analysis, we found the commu- nity structure of the POS region to be virtually identical with the OPH region. No community appeared to be unique to it. Those communities that typify it upland post oak hickory forest/woodland, xeric sandylands, prairies, glades, baygalls, and so forth also occur in the OPH region (e.g., Marietta & Nixon 1983, 1984: Ward & Nixon 1993; Nesom et al. 1997; MacRoberts et al. 2002a). If there is anything distinctive about the POS region, it is that some of these communi- ties (e.g., xeric sandylands, prairies) are more common in that region than they are farther east. Ironically, while the absence of pine is the feature most often noted for the POS region, pine does occur in the POS region, most dramatically 404 BRIT.ORG/SIDA 21(1) Fic. 2. Association of Post Oak $ fl d t f flora at Gus Engeling Wildlife Management A 7 ' h ctat n Les bs L ° £[f C ih VARIA LA AA A I province, or region g J (see text for further explanation). at its western edge in Bastrop County, where Pinus taeda is often a dominant or co-dominant canopy species (Wilson 1990). Neither a botanist nora plant ecologist familiar with the southeastern flora would find much new in the POS. The main difference would be the absence of pines in some areas and the different frequency of communities and thus taxa in others. Even the few endemics to the region are tied to one community type —xeric sandylands—which occurs as far east as southwest Arkansas, central Louisiana, and southeast Texas (MacRoberts & MacRoberts 1994, 1995, 1996; MacRoberts et al. 2002a). The difference between the POS and the OPH regions is one of degree, not of kind. This finding agrees with a number of previous workers who have indicated the lack of a sharp boundary between the OPH and POS and who have questioned the floral uniqueness of the POS altogether (e.g, Tharp 1926; McCarley 1959; Diggs et al. 1999; MacRoberts & MacRoberts 2003a). Basically, the POS region is part of the 300 km ecotone between east and west, just as is the north central Texas prairie and cross timbers region (Diggs et al. 1999; MacRoberts & MacRoberts 2003b). ACKNOWLEDGMENTS Ray C.Telfair and an anonymous reviewer made many useful comments on the paper. MACROBERTS AND MACROBERTS, THE POST OAK SAVANNA ECOREGION 405 REFERENCES Bezanson, D. 2000. Natural vegetation types of Texas and representation in conservation areas. M.A. Thesis, University of Texas, Austin. Corrett, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner Diamonp, D.D., D.H. Riskino, and S.L. Orzeit. 1987. A framework for plant community classi- fication and conservation in Texas. Texas J. Sci. 39:203-221. Diccs, G.M., B.L. Liescome, and RJ. O’Kennon. 1999. Illustrated flora of North Central Texas. Sida Bot. Misc. 16:1-1626. Evans, R.E.n.d. List of the taxa occurring in Region 1 of Texas. Unpublished manuscript. Fiemine, K.M.,J.R. SincHurst, and W.C. Houmes. 2002. Vascular flora of Big Lake Bottom Wildlife Management Area, Anderson County, Texas. Sida 20:355-371. Gouto, F.W. 1962. Texas plants. A checklist and ecological summary. Tex. Agric. Exp. Sta. Misc. Publ. 585:1-112. Harcomee, PA., J.S. GLITZENSTEIN, R.G. KNox, S.L. Orzett, and E.L. Brioces. 1993. Vegetation of the longleaf pine region of the West Gulf Coastal Plain. Proc. Tall Timbers Fire Ecology Con- ference 18:83-104. HarsHBeRGER, J.W. 1911. Phytogeographic survey of North America. (2nd Ed.) Hafner, New York. Hatcn, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Misc. Publ. 1655:1-158. Kartesz, J.T.and C.A. Meacham. 1999. Synthesis of North American flora. Version 1.0. North Carolina Botanical Garden. Chapel Hill. LBJ ScHoot of Pustic Arrairs. 1978. The natural regions of Texas. In: Preserving Texas's Natu- ral Heritage. L.B.J. School of Public Affairs Policy Res. Proj. Rept. 31:17-24. Louisiana Natura Heritace. 2003. Natural communities of Louisiana. Unpublished report, Louisiana Dept. Fish and Wildlife, Baton Rouge. MacRoserts, M.H. and B.R. MacRoserts. 1994. Floristics of a xeric sandyland in western Loui- siana. Phytologia 77:414-424. MacRoseats, B.R. and M.H. MacRoserts. 1995. Floristics of xeric sandhills in northwestern Louisiana. Phytologia 79:1 23-131. MacRoseats, B.R. and M.H. MacRoserts. 1996. Floristics of xeric sandhills in east Texas. Phytologia 80:1—7. MacRoserts, B.R. and M.H. MacRoserts. 1998. Floristics of muck bogs in east central Texas. Phytologia 85:61-73. MacRoserts, M.H. and B.R. MacRoserts. 2001. Bog communities of the West Gulf Coastal Plain: a profile. Bog Research Papers in Botany & Ecology 1:1-151. MacRoseats, B.R., M.H.MacRoserts, and J.C. Carney. 2002a.Floristics of xeric sandylands in the post oak savanna region of east Texas. Sida 20:373-386. MacRoseats, M.H., B.R. MacRoserts, B.A. Sorrie, and R.E. Evans. 2002b. Endemism in the West Gulf Coastal Plain:importance of xeric habitats. Sida 20:767-780. 406 BRIT.ORG/SIDA 21(1) MacRoserts, M.H. and B.R. MacRoserts. 2003a. West Gulf Coastal Plain ecoregions. Sida 20:1247-1276. MacRoserts, M.H. and B.R. MacRoserts. 2003b. The east-west transition of flora in Texas: a biogeographical analysis. Sida 20:1693-1 700. Marietta, K.L. and E.S. Nixon. 1983. Vegetational analysis of a post oak-black hickory com- munity in eastern Texas. Texas J. Sci. 35:196-203 Marietta, K.L.and E.S. Nixon. 1984.Vegetation of an open, prairie-like community in eastern Texas. Texas J. Sci. 36:25-32. Marks, P.L. and P.A. Harcomee. 1981. Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monogr. 51:287-305. McBryoe, J.B. 1933. The vegetation and habitat factors of the Carrizo Sands. Ecol. Monogr 3:247-297. McCarey, H. 1959. The mammals of eastern Texas. Texas J. Sci. 11:385-426, Nesom, G.L., B.R. MacRoserts, and M.H. MacRoserts. 1997. A new plant community type in southeast Texas related to baygalls. Phytologia 83:371-383. Ricketts, T.H. and E. Dinestein. 1999. Terrestrial ecoregions of North America. Island Press, Washington, D.C Shantz, H.L. and R. Zon. 1924. Natural vegetation. Atlas of American Agriculture. Pt. 1.U.S. Dept. Agriculture, Washington, D.C SinGHuRsT, J.R., J. CatHey, D. ProcHaska, H. Haucke, G.C. Kron, and W.C. Hotmes. 2003. Vascular flora of Gus Engeling Wildlife Management Area, Anderson County, Texas. Southeastern Naturalist 2:347-368. Smeins, F.E.and D.D. Diamonp, 1983. Remnant grasslands of the Fayette prairie, Texas. Amer. Midl. Naturalist 110:1-13. TAKHTAJAN, A. 1986. Floristic regions of the world. University of California Press, Berkeley. Tetrair, R.C. II. 1988. Conservation of the Catfish Creek ecosystem: a national natural land- mark in eastern Texas. Texas J. Sci. 40:11-23. Tevrair, R.C. Il. 1999. Introduction: ecological regions of Texas: description, land use, and wildlife. In: R.C. Telfair Il. ed., Texas wildlife resources and land use. Univ. Texas Press, Austin. Pp. 1-39. Texas Natural Heriace Procram. 1995, Plant communities of Texas. Unpublished report. Texas Parks and Wildlife Department, Austin. Tuarp, B.C. 1926. Structure of Texas vegetation east of the 98th meridian. Univ. Texas Bull. 2606:1-99. THomas, R.D.and C.M. ALLen.1993-1998. Atlas of the vascular flora of Louisiana. The Nature Conservancy, Baton Rouge. THe Nature Conservancy. 2003.What is an ecoregion? www.tnc.org. (14. Jan. 2004). Turner, B.L., H. Nicots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida, Bot. Misc. 24:1-888. Turner, R.L., J.E. VAN Key, L.S. Swit, and R.E. Evans. 1999. Ecological classification system for the national forests and adjacent areas of the West Gulf Coastal Plain. The Nature Con- servancy, Nacogdoches. MACROBERTS AND MACROBERTS, THE POST OAK SAVANNA ECOREGION 407 VAN Kuey, J.E. 1999a. The vegetation of the Kisatchie sandstone hills, Louisiana. Castanea 64:64-80. VAN Ktey, J.E. 1999b. The vegetation of the high terrace rolling uplands, Louisiana. Casta- nea 64:318-336. Warp, J.R.and E.S. Nixon. 1992.Woody vegetation of the dry, sandy uplands of east Texas. Texas J. Sci.44:283-294. Weer, W.L. 1950. Biogeographic regions of Texas and Oklahoma. Ecology 31:426-433. Witson,R.E. 1990.The eastward recession of the piney woods of northeastern Texas, 1815- 1989. Texas J. Sci.42:179-189. BRIT.ORG/SIDA 21(1) BOOK NOTICE Timber Press DENISE WILES ADAMS. 2004. Restoring American Gardens: An Encyclopedia of Heirloom Ornamental Plants, 1640-1940. ISBN 0-88192-619-1, hbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Or- ders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800- 327-5680, 503-227-3070 fax). $39.95, 420 pp., 155 color photos, 37 b/w photos, 154 line drawings, 1 table, 81/2" x 11". — The meat of this great book is the “Encyclopedia of Heirloom Ornamental Plants.” The heirlooms are alphabetical under broad categories like Heirloom Trees, Heirloom Ornamental Shrubs, Heirloom Bulbs and Tuberous Plants, Heirloom Roses, ete. Each heirloom plant is listed with most of the fol- lowing descriptive information: L) Scientific name, 2) Common names. 3) Synonym, 4) Family, 5) .6) Date of introduction into North America, 7) Earliest American citation, Nativ ity Jnat ive or exotic] 8) Hardiness zone, 9) Description, 10) Design notes, 11) Remarks, 12) Heirloom varieties, 13) Related species, and 14) Historical commercial sources. Lreally like this book a lot! It is filled with hard to find information, great images of plants from os old catalogs, as well as beautiful colored plates of the past three centuries, and modern color photo- », Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas graphs.—Barney Lipscom 761202-4060, U.S.A. SIDA 21(1): 408. 2004 AMQOUANTITATIVE STUDY OF THE VEGETATION SUR= ROUNDING YELLOW LADY-SLIPPER ORCHID (CYPRIPE- DIUM KENTUCKIENSE, ORCHIDACEAE) POPULATIONS AT FORT POLK IN WEST CENTRAL LOUISIANA Charles M. Allen Sara Thames Colorado State Univ., Fort Polk Oak Ridge Institute for Science and Education 1 23rd Street 1645 23rd Street Fort Polk, Louisiana 71459, U.S.A. Fort Polk, Louisiana 71459, U.S.A. Spencer Trichell and Jeremy White Dept. of Biology University of Louisiana at Monroe onroe, Louisiana 71209, U.S.A. ABSTRACT The vegetation around five populations of yellow lady-sli rchid (Cypripedium kentuckiense Reed) ee at Fort Polk, Louisiana were sampled using circular nese ae Mea een cluded frequency mean density, mean cover percent, mean dbh. i importance value, and diversity (richness). The yellow lady-slippers at Fort Polk were associated with forests composed mainly of American beech (Fagus grandifolia), eastern hophornbeam (Ostrya virginiana), white oak (Quercus alba), horsesugar (Symplocos tinctoria), and witch hazel (Hamamelis virginiana) in the overstory and an abundance of poison ivy (Rhus radicans) and broad beechtern (Thelypteris hexagonoptera) in the understory. RESUMEN Se muestre6 la 10 ] led | 1 L blaci | -ypripe edium kentuckiens¢ Reeden Fort Polk, Louisiana mediante | parcelas anidadas circulares. Las medidas incluyen frecuencia, densidad } a media lioala alr del pee ho, valor de importancia, iY Huversiciad (riqueza). Cypripedium kentuckiense en Fort Poll | | | por haya americana (Fagus grandifolia), Ostrya virginiana, roble t (Quercus all pS aplaces tinctoria, y Hamamelis virginiana en el dosel arboreo y joene ate zumaque sha radicans) y g q — Thelypteris hexagonoptera en el sotobosque. INTRODUCTION Yellow lady-slipper (Cypripedium kentuckiense Reed) is an herbaceous peren- nial in the Orchidaceae family. This species is reported from the southeastern US.in AL; AR; KY; LA; MS; OK; TN; TX; and VA (USDA, NRCS 2002). Globally this species is ranked G3 which is defined as either rare and local throughout its range or found locally (even abundantly at some of its locations) in a re- stricted range (e.g., a single physiographic region) or because of other factors making it vulnerable to extinction throughout its range, or with 21- 100 known extant populations or 3,000 to 10,000 individuals (NatureServe 2003). It is ranked Sl in AL, GA, LA, OK, TX, and VA. An SI species is defined as being SIDA 21(1): 409 — 417. 2004 410 BRIT.ORG/SIDA 21(1) critically imperiled in a state because of its extreme rarity (5 or fewer known extant populations) or because of some factor(s) making it especially vulner- able to extirpation. It is listed as SI/S2 in TN, S2/S3 in KY, $3 in AR, and SU in MS. The S2 ranking is defined as imperiled in the state because of rarity (6 to 20 known extant populations) or because of some factor(s) making it very vul- nerable to extirpation while the S3 ranking is defined as rare and local through- out the state or found locally (even abundant at some of its locations) in a re- stricted region of the state, or because of other factors making it vulnerable to extirpation (21 to 100 known extant populations). The SU ranking is for those species that are possibly in peril in the state but with an uncertain status and more information is needed. Thirteen populations of this orchid are known from east Texas (Liggio & Liggio 1999). In Louisiana, it is reported from 11 parishes with four from unverified reports (Louisiana Department of Wildlife and Fish- eries; Natural Heritage Program 2003). The habitat for this species is described as mature floodplain forests and the slopes of mesic (relatively dry) ravines (Oklahoma Natural Heritage Inven- tory 2001) and rich, mesic to dry, deciduous forests on well-drained alluvium and bases of slopes, or mucky seeps; mostly 0-400m (Flora of North America Editorial Committee 2002). Orzell (1990) describes the habitat for this species as terraces and lower moist slopes in small stream valleys of pine-hardwood forests. This author lists the common trees of the canopy as American beech (Fagus grandifolia), white ash oe aan” black gum (Nyssa sylvatica),and southern magnolia (A liagrandiflora). The understory spe- cies reported by Orzell 1990) include flowering dogwood (Cornus florida), red maple (Acer rubrum), chalk maple (Acer lecoderme), bigleaf snowbell (Styrax grandifolia),and American hornbeam (Carpinus caroliniana). Orzell (1990) also reports that two shrubs species, maple-leaf viburnum (Viburnum acerifolium) and arrow wood (Viburnum dentatum) are frequently associated with this or- chid. Herbaceous plants often found associated with yellow lady-slippers in- clude cranefly orchid (Tipularia discolor), slender wood sedge (Carex digitalis), partridge berry (Mitchella repens), Christmas fern (Polystichum acrostichoides), Walter’s violet (Viola walteri), eared goldenrod (Solidago articulata), broad beech fern (Thelypteris hexagonoptera), and slender wake robin (Trillium grac- ile) (Orzell 1990). The habitat in Louisiana is reported to be mesophytic woods, calcareous forests, and hardwood slope forests (Louisiana Department of Wild- life and Fisheries, Natural Heritage Program, 2003). A search of the literature yielded no quantitative data on the habitat for yellow lady-slipper throughout its range. The objective of this study was to quantitatively describe the habitat for yellow lady-slipper in west central Louisiana. Five distinct groups of yellow lady-slipper are known from the Fort Polk Military Installation in Vernon Parish, Louisiana. Three of the five groups are located at one site while two are located at a different site. The two sites are ALLEN ET AL., POPULATIONS OF , LOUISIANA 411 located about six kilometers apart and each is a well drained ravine in a mesic hardwood forest. Each ravine has a small, intermittent stream at the bottom. The soils at both sites are Eastwood silt loam (Soil Survey Division, Natural Resources Conservation Service, United States Department of Agriculture 2003). These five groups have been observed for the past ten years with four clumps having flowered and one clump not flowering during this period. METHODS Three nested circular plots were sampled with the center of each of the five yellow lady- slipper groups Scrane as the center point of the plot. All trees and shrubs (woody non-vine species taller than 6 ft = 1.83m) were sampled withina 15m radius circle; shrubs and saplings (woody non-vine species shorter than L83m = 6 ft) and woody vines were sampled within a 5m radius circle; and al herbaceous species were sampled within a 1m radius circle. During the sam- pling period, the number of stems was counted and recorded for all species in the samples. For the trees and shrubs within the 15m radius samples, the dbh was measured at the standard 1.37m (4.5 ft) height using a diameter tape and recorded to the nearest 0.1 cm. For herbaceous plants, woody vines, and shrubs and saplings, cover was determined by measuring the area occupied by the in- dividuals of the species in the sample. For the few species with only one or two individuals per sample, the actual area was used. For those species with more than two individuals per sample, the mean area was calculated by summing the area occupied by three or more randomly selected individuals of the spe- cies and dividing by the number of individuals selected. The cover was then calculated by multiplying the actual area or mean area times the number of individuals and then dividing by the area of the plot, 78.035m? for the 5m ra- dius plots and 3.14m? for the lm radius plots. The cover was converted to a per- cent by multiplying by LOO. All data were entered into a Microsoft Excel spread sheet for storage and calculation of variables. The mean diversity (richness) and mean density were calculated for each of the five plots for all plant groups while the mean dbh was calculated for trees and shrubs and mean cover percent was calculated for her- baceous plants, woody vines, and shrubs and saplings. The frequency and mean density were calculated for each species. The mean dbh was calculated for tree and shrub species and the mean cover percent for all other species. The relative values for each of these variables (frequency, mean density, mean dbh, and mean cover percent) were calculated by dividing the value for a species by the total for all species. Each value was converted to a percent by multiplying by 100 and the sum of these relative values was used to calculate the importance value. RESULTS The number of species totaled seventy-eight (78) with nine (9) woody vines, 412 BRIT.ORG/SIDA 21(1) twenty-one (21) herbaceous plants, and forty-five (45) tree, shrub or saplings (Table 1). The mean number of species per sample ranged from 9.00 for woody vines to 23.80 for shrubs and saplings. The mean number of stems per sample (density) averaged 651.40 per sample for all plants and ranged from 77.00 for trees and shrubs to 216.80 for shrubs and saplings. The mean cover percent for all plants was 240.74 percent and ranged from 8.68 percent for woody vines to 159.86 percent for herbaceous plants. The mean dbh per sample was 877.98 cm. The frequency, mean density, mean dbh, and importance value for the tree and shrub species are in Table 2, while the frequency, mean density, mean cover percent, and importance value for the appropriate plant group are in Tables 3, 4,or5. The species are arranged in descending importance value in all four tables. The three most important tree and shrub species are American beech (28.12), white oak (Quercus alba) (27.73), and horsesugar (Symplocos tinctoria) (25.94) (Table 2). Eastern hophornbeam (Ostrya virginiana) was the dominant shrub and sapling species with an importance value of 55.01 followed by witch hazel (Hamamelis virginiana) (31.63) and horsesugar (23.64) (Table 3). The most im- portant woody vine species was poison ivy (Rhus radicans) (128.28) and ac- counted for almost one-half of the total importance value of 300 while crossvine (Bignonia capreolata) (33.42) and common greenbrier (Smilax rotundifolia) (33.34) ranked second and third respectively (Table 4). The dominant herba- ceous species was broad beechfern (Thelypteris hexagonoptera) with an impor- tance value of 125.18 followed distantly by partridge berry (37.19) and longleaf woodoats (Chasmanthium sessiliflorum (20.51) (Table 5). DISCUSSION Allen et al (1994) reports physiognomy data for 4m radius samples in a beech forest area in the Louisiana Arboretum in Evangeline Parish near a yellow lady- slipper population. The diversity (richness) for this beech forest includes a mean of 8.20 species for the shrubs and saplings and 5.10 species of trees and shrubs compared to 23.80 species for shrubs and saplings and 19.00 for trees and shrubs in this study. The density and cover percent in the beech forest for the shrubs and saplings were 76.10 stems and 42.88 percent, respectively, which compared to 216.80 stems and 72.20 percent in this study. The mean trees and shrub den- sity in this study was 77.00 compared to 10.30 in the Arboretum beech forest. The mean dbh in this study was 877.98 cm compared to 50.68 cm at the Arbo- retum. Newman (1995) reports physiognomy data from 4m radius samples ina hardwood slope forest in Caldwell Parish, Louisiana. This site is a short distance south of a yellow lady-slipper population in extreme southern Ouachita Par- ish. The shrub and sapling data from the Caldwell site included a mean diver- sity of 15.82 species compared to 23.80 species in this study; mean density of 67.65 stems compared to 216.80 stems; and mean cover percent of 39.56 percent compared to 72.20 percent. The trees and shrubs from the Caldwell Parish study — ALLEN ET AL., POP VLATIVIND UE POLK, LOUISIANA 413 Taste 1. Community physiognomy variables (diversity, density, cover percent, and dbh) in samples around yellow lady-slioper populations at Fort Polk, La. Diversity (Richness) All Trees & Shrubs & Woody Herbaceous Plants Shrubs Saplings Vines Plants Mean 48.00 19.00 23.80 9.00 10.20 Std Dev. 2.83 2.55 3.77 1.00 2.49 Range 45-5] 16-22 18-27- 8-10 8-14 Total Number 78 33 4l 12 21 Density All Trees & Shrubs & Woody Herbaceous Plants Shrubs Saplings Vines Plants Mean 651.40 77.00 216.80 205.20 152.40 Std Dev. 83.46 20.41 55.95 77 86 88.49 Range 644-706 55-108 175-310 106-317 71-286 Cover Percent DBH (cm) All Shrubs & Woody Herbaceous Trees & Plants Saplings Vines Plants Shrubs Mean 240.74 72.20 8.68 159.86 877.98 Std Dev. 141.53 25.44 2.81 136.79 124.00 Range 113.63- 39.67- 4.65- 34.55- 741.10- 449.00 109.82 11.80 365.49 1011.20 had a mean diversity of 4.20 species compared to 19.00 in the present study; mean density of 8.28 stems compared to 77.00; and mean dbh of 68.03 cm com- pared to 877.98 cm. Some of the larger numbers in this study in comparison to both of these reports can be attributed to the difference in sample size, but the Fort Polk yellow lady-slipper area seems to be more diverse, with a higher veg- etation density, and a higher basal area. The five yellow lady-slipper populations at Fort Polk were associated with a forest composed of American beech, eastern hophornbeam, horsesugar, white oak, and witch hazel in the overstory and an abundance of poison ivy and broad beechfern in the understory. This description is very similar to the qualitative report of associated species by Orzell (1990) except that American hornbeam (Carpinus caroliniana) was not found in the sampled area at Fort Polk. The three most important shrubs and saplings reported from Caldwell Parish by New- man (1995) include American beech, red maple (Acer rubrum), and eastern hophornbeam and the top three reported from the Louisiana Arboretum by Allen et al (1994) include native bamboo (Arundinaria a paw paw (Asimina triloba), and pignut hickory (Carya glabra). All of these species were found in the Fort Polk study except for pawpaw and pignut hickory. The three most important species of trees and shrubs in Caldwell Parish included flower- ing dogwood (Cornus florida), sweet gum (Liquidambar styraciflua), and east- 414 BRIT.ORG/SIDA 21(1) Taste 2, Frequency, Mean Density, Mean dbh, and Importance Value for Trees and Shrubs in 15m radius samples around Yellow Lady-slipper Populations at Fort Polk, La\<4> Mean Mean Importance Species Frequency Density dbh Value Fagus grandifolia 100.00 8.20 107.14 28.12 Quercus alba 100.00 4.80 142.56 27.73 Symplocos tinctoria 60,00 14.00 40.42 25,94 Nyssa sylvatica 100.00 3.60 92.90 20.52 Ostrya virginic 100.00 6.20 46.96 18.66 Liquidambar ee 100.00 3.40 69.76 17.62 Cornus florida 100.00 4.80 38.72 15.9] llex opaca 100.00 3.60 20.38 12.26 Pinus taeda 60.00 1.20 65.60 12.19 Hamamelis virginiana 100.00 3.80 9.80 11.31 sisi alba 60.00 1.80 51.04 11.31 Pricana 100.00 1.60 30.32 10.79 ie jpn 80.00 2.00 31.88 10.44 Acer leucoderme 60.00 2.80 2148 9.24 Carya ovat 60.00 1.20 27.18 7.8] Quercus falcate 60.00 0.60 2844 7.18 Styrax grandifolia 60.00 2.00 7.80 6.64 Arundinaria gigantea 40.00 2.60 1.92 5.70 Rhododendron car 60.00 1.20 1.62 4.90 Vaccinium elliottii 60.00 1.20 1.58 4.90 Vaccinium arboreu 40.00 1.60 6.10 4.88 Aralia spinosa 40.00 1.60 4.28 4.67 Chionanthus virginicus 60.00 0.60 1.84 4.15 Ulmus alata 20.00 0.60 10.04 2.98 Pinus echinata 20.00 0.40 10.24 2/4 Carya spp. 20.00 0.20 3.04 1.66 Crataegus spathulata 20.00 0.20 1.36 1.4/ Crataegus marshalli 20.00 0.20 Ted 1.45 Viburnum rufidulum 20.00 0.20 0.76 1.40 Magnolia grandiflora 20.00 0.20 0.62 1.38 llex decidua 20.00 0.20 0.46 1.36 Morus rubra 20.00 0.20 0.30 1.35 Viburnum dentatur 20.00 0.20 0.22 1.34 Total 1900.00 7700 877.98 300.00 ern hophornbeam, and in the Louisiana Arboretum the three most important species were American hornbeam, flowering dogwood, and American beech. All of these species were in the Fort Polk site except for the notable absence of American hornbeam. The absence of American hornbeam could be explained by the Fort Polk site being slightly drier than the other sites as American hornbeam’s wetland ranking is FAC and eastern hophornbeam’s ranking is FACU-(USDA-NRGS 2002), ALLEN ET AL., PYUPULATIVING UT POLK, LOUISIANA Taste 3. Frequency, Mean Density, Mean Cover Percent, and Importance Value for Shrubs and Sap- lings in 5m radius samples around yellow lady-slipper Populations at Fort Polk, Mean Mean Importance Species Frequency Density Cover Value Ostrya virginiana 80.00 39.00 24.30 55.01 Hamamelis virginiana 100.00 33.60 8.61 31.63 Symplocos tinctoria 80.00 10,20 11.24 23.64 | inium elliottii 80.00 7.00 5.74 14.54 Styrax grandifolia 100.00 12.00 1.85 12.29 ne epee 80.00 2.40 Bl? 11.63 ia gigantea 20.00 12.20 3.22 10.93 Ae leudoderme 60.00 15.40 0.79 10.72 Callicarpa americana 60.00 3.60 3.29 8.74 Viburnum dentatum 100.00 7.00 0.54 8.18 Acer rubrum 100.00 7.00 0.31 7.87 Cornus florida 100.00 6.80 0.32 778 Pinus taeda 60.0 9.40 0.01 6.87 Chionanthus virginicus 100.00 3.40 0.65 6.68 Quercus alba 80.00 5.40 0.14 6.04 Vaccinium arboretum 60.00 2.20 1.49 5.60 Carya ovata 40.00 6.80 0.48 5.48 Prunus serotina 80.00 4.20 0.05 5.37 agus grandifolia 80.00 3.80 0.05 5.18 Carya spp. 60.00 440 0.31 4.98 Aralia spinosa 40.00 2.00 1.28 4.37 pean canescens 60.00 2.80 0.31 4.25 americana 80.00 1.00 0.25 4.17 C egus marshallii 40.00 3.40 0.51 3.96 llex decidua 60.00 1.80 0.37 3.87 Nyssa sylvatica 60.00 2.00 0.06 3:52 Viburnum rufidulum 60.00 1.60 0.05 3.32 Sassafras albidum 60.00 0.60 0.00 2.80 Vaccinium virgatu 40.00 0.80 0.18 2.30 Hypericum hypericoides 40.00 0.60 0.10 2.09 Crataegus spathulata 40.00 0.40 0.14 2.06 Hypericum frondosum 40.00 0.40 Oel2. 2.03 Care canadensis 40.00 0.40 0.05 1.94 ! alata 40.00 0.40 0.01 1.88 Rubus argutus 40.00 0.40 0.01 1.88 Morus rubra 20.00 0.80 0.07 1331 Liquidambar styraciflua 20.00 0.60 0.03 1.16 Quercus velutina 20.00 0.40 0.03 1.07 Ilex eae 20.00 0.20 0.03 0.97 Persea palus 20.00 0.20 0.02 0.96 Quercus cae 20.00 0.20 0.01 0.95 Total 2380.00 216.80 72.20 300.00 416 BRIT.ORG/SIDA 21(1) Taste 4. Frequency, Mean Density, Mean Cover Percent, and Importance Value for Woody Vines in 5m radius samples around yellow lady-slipper Populations at Fort Polk, La Mean M Importance Species Frequency Density Cover Value Ave radicans 100.00 128.20 4.75 128.28 lato 100.00 29.00 0.71 33.42 vile euandiiola 100.00 8.40 1.57 33.34 a rotundifolia 100.00 8.60 0.22 17.84 Smilax pumila 40.00 11.80 0.60 17.16 Parthenocissus quinquefolia 80.00 6.40 0.14 13.66 Smilax tamnoides 80.00 4.40 0.21 13.47 Smilax smallii 100.00 3.40 0.05 13.37 Vitis vulpina 60.00 1.60 0.29 10.75 Smilax glauca 80.00 1.40 0.02 9.78 Lonicera sempervirens 40.00 1.60 04 5.64 Berchemia scandens 20,00 0.40 0.07 3.28 Total 900.00 205.20 8.68 300.00 TABLE 5. Frequency, Mean Density, Mean Cover Percent, and Importance Value for Herbaceous Plants Im radius samples around yellow lady- slipper poriuiens at Fort Polk, La. Mean Mean Importance Species Frequency Density Cover Value Thelypteris ee 40.00 82.00 107.82 125.18 Mitchella repens 100.00 37.60 4.34 37.19 eon jum sessiliflorum 100.00 5.80 11.04 20.51 Dicanthelium boscii 100.00 6.20 4.29 16.56 Carex spp. 40.00 2.40 14.69 14.69 Cypripedium i, 100.00 2.40 4.42 14.14 Solidago ¢ 60.00 3.00 1.43 8.74 pigelia n oe dica 60.00 2.60 1.51 8.53 sceda oligantha 60.00 1.80 1.79 8.18 ee en 40.00 2.00 2.99 7.10 / 0.00 1.20 0.54 701 te pe 40.00 1.20 1.69 5.76 Desmodium spp. 40.00 0.80 0.62 4.83 Elephantopus spp 40.00 0.40 0.14 4.27 Seite tri pe um 20.00 0.80 0.93 3.07 Polystichum acr id 20.00 0.60 0.74 2.82 psa eae 20.00 0.60 0.19 24/7 nutatum 20.00 0.20 0.57 2.45 Ruellia eee 20.00 0.40 0.10 2.28 Viola walteri 20.00 0.20 0.01 2.10 Viola spp. 20.00 0.20 0.01 2.10 Total 1020.00 152.40 159.86 300.00 ALLEN ET AL., POPULATIONS Ur LOUISIANA 417 This study is apparently the first report of quantitative data on the vegeta- tion associated with yellow lady-slippers. The vegetation around other yellow lady-slipper populations throughout its range should be sampled and compared and contrasted to our results to provide data for ecologically sound management decisions to preserve and expand the populations of this threatened species. ACKNOWLEDGMENTS We are indebted to Fort Polk, the University of Louisiana at Monroe, and the two anonymous reviewers. REFERENCES ALten, C.M., MLE. Viorine, and B. Borsari. 1994. Analysis of the woody vegetation of a beech forest area in the Louisiana Arboretum. Louisiana Environm.Professional 10-11(1): 17--26. Flora or NortH America Epitoriat Committee. 2002. Flora of North America: Volume 26, Magnoliophyta: Liliidae: Liliales and Orchidales. Oxford Univ. Press, New York. Liccio, J.and A.O. Liccio. 1999. Wild orchids of Texas. Univ. of Texas Press, Austin. LouIsIANA DEPARTMENT OF WILDLIFE AND FisHeries; NaTurAL HeriTAGe Procram. 2003. Rare Plant fact sheet for Cypripedium kentuckiense C.F. Reed. http: ifstate.la.us/apps/netgear/ clientFiles/lawlf/files/1049978676.pdf NatureServe. 2003, NatureServe Explorer: An online encyclopedia of life [web application]. 2002. Version 1.6 . Arlington, Virginia, USA: NatureServe. Available: http:// www.natureserve.org/explorer. (Accessed: April 28, 2003). Newman, S.D. A. 1995. Ecological analysis of the vegetation of a hardwood slope forest in northern Caldwell Parish, Louisiana. M.S. Thesis, Northeast Louisiana University, Monroe. Oxtanoma Naturat Heritace. 2001. Cypripedium kentuckiense southern lady’s slipper: sta- tus, description, life history, habitat, distribution, field characters, cause of decline, and recovery needs. http://www.biosurvey.ou.edu/ Orzett, S. L. 1990. Inventory of National Forests and National Grasslands in Texas. Texas Natural Heritage Program. Austin. Soi Survey Division, NATURAL RESOURCES CONSERVATION Service, UniTeD STATES DEPARTMENT OF AGRICUL- ture. 2003. Official soil series descriptions, Eastwood series. (http:// ortho.ftw.nrcs.usda.gov/osd/)USDA, NRCS. 2002. The PLANTS database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. 418 BRIT.ORG/SIDA 21(1) BOOK NOTICES Sinauer Associates RICHARD B, PRIMACK. 2004. A Primer of Conservation Biology. Third Edition. (ISBN 0-87893-728-5, pbk.). Sinauer Associates, Inc. Publishers, 23 Plumtree Road, Sunderland, MA 01375, U.S.A. Orders: orders@sinauercom, wwwsinauercom, $44.95, 320 pp., illustrated, 7" x 9" Contents: 1) Conservation and Biological Dive rsity. 2) Threats of Biological Diversity. 3) Conserva- tion at the copHiaiion and Species Levels. 4) Conserving Biological e communities, 3, and we Conserva- tion and Sustainable Development. Appendix: Selected E Information, Glossary, Bibliography, and Index. Barry G. Hatt. 2004. Phylogenetic Trees Made Easy: A How-to Manual. Second Edition. (ISBN 0-87893-312-3, pbk.). Sinauer Associates, Inc. Publishers, 23 Plumtree Road, Sunderland, MA 01375, U.S.A. Orders: orders@sinauercom, wwwssinauer.com, $31.95, 221 pp,, illustrated, 7" x 91/4". According to the author, “This isa ‘cook book’ eas as a tool to aid begit in creati eee netic trees from protein data. There are four eae FOV ide astep step sequence lor building ae netic trees: 1) Tutor mi Create a Tree! 2) Basic Elements in Creat- ing and Presenting Trees. 3) Advanced Elements in Constructing Trees. 4) U sing Alternative Soft- ware to Construct and Present Trees. Two appendices are included: I: File Formats and Their Interconversion. I. Printing Alignments. SIDA 21(1): 418. 2004 CHANGES IN BAYGALL VEGETATION FROM 1986 TO 2001 AT FORT POLK IN WEST CENTRAL LOUISIANA Charles M. Allen and John Pate Sara Thames olorado State Univ., Fort Polk Oak Ridge Insti for Sci and Education 645 23rd Street 1645 23rd Street Fort Polk, Louisiana 71459, U.S.A. Fort Polk, Louisiana 71459, U.S.A. Spencer Trichell and Lacy Ezell Department of Biology Universi aC of Louisiana at Monroe Monroe, Louisiana 71209, U.S.A. ABSTRACT Baygalls area plant community associated with small streams in west central Louisiana. Plots from a 1986 sud) on the effects of sedimentation on bay pall “eae ation were ices ty in 2001. Nested ] 4 Ble min ] ubsample Cs CC ALC CLIC SATIIC alCas Import ATICE Values were la, using frequency, individual density, stem deni: and dbh. In 1986, ae oe nee and control baygalls were significantly different in allvareble sexcept lor woody 5 and woody stem density. In 2001, dbh was the only variable not Sioniticandly different between the control and sedimented ae In the sedimented area between 1986 and 2001, the woody plant variables increased while herbaceous plant variables decreased. os species, Liquidambar styraciflua and Alnus serr ulata showed a large decrease, however, Toxicodendron radicans, Smilax rotundifolia, and Ligust 5 se increased. In the control baygalls, four fern species increased in importance value. RESUMEN Los “Baygalls” idad a! rn ] oeste de Luisiana Las parcelas de un estudio en 1986 sobre Os efectos de la sedimentacion en la vepetaciOn bay gall se volvierona muestrear en 2001.5 lad mismas areas. Los valores de je eee se calcularon aes frecuencia, densidad errand densidad de tallos, y diametro a la altura del pecho. En 1986, los baygalls sedimentados y de control fueron sie unc atvamientee diferentes en todas las variables excepto para la densidad individual de le lenosas y llos lenosos. En 2001, el diametro a la iris del pecho fue la unica variable no significativamente diferente entre el control y los baygalls sedimentados. En el area sedimentada entre EO Y oe las variables de Beales lehosas aumentaron mientras que las variables de plantas I Liquidambar styraciflua y Alnus serrulata mostraron una gran disminucion, sin eabace Toxicodendron clea Smilax rotundifolia, y Ligustrum sinense aumentaron. En los baygalls de control, cuat p helechos incrementaron su valor de importancia. INTRODUCTION Bay gall is a colloquial term that refers to the small drainage systems and to the vegetation developed along the streams (Allen et al. 1990). In the center of most baygalls is a small sandy or gravelly-bottomed stream that is intermittent in SIDA 21(1): 419 - 427. 2004 420 BRIT.ORG/SIDA 21(1 the upper drainage and usually continuous in the lower portions of the drain- age. The smaller streams drain into slightly larger streams and ultimately into a larger watercourse. Baygalls are surrounded by pine forests and are narrow in the upper portion gradually widening downstream to a maximum width of 20-50 meters. Drainage from the surrounding upland pine forest percolates downward until it reaches an impervious layer of clay or rock typically form- ing seepage areas throughout the baygall. Pitcher plant bogs are found in open areas in and along the edge of many baygalls (Allen et al. 1987). The woody vegetation in the baygalls is predominately broad-leaved (dicotyledonous) de- ciduous or evergreen trees or shrubs contrasting with the very common ever- green (gymnospermous) needle-leaved trees of the upland pine forests. The veg- etation developed along the floor of the baygall is sparse and contains a number of ferns, bryophytes, especially peat moss (Sphagnum spp.), and a few shade- tolerant flowering plants. METHODS In the summer of 2001, six baygalls used in a 1986 study (Allen et al. 1990) were examined for resampling. The study area in one of the sedimented baygalls was completely destroyed and thus was excluded from resampling. One of the two transects in two different control baygalls could not be relocated. Two transects were resampled in the one sedimented baygall with a total of 37 subsamples. Four transects in four different control baygalls were resampled with 94 subsamples. The 1986 data from these 37 and 94 relocated samples only were used for comparisons with the 2001] data. In 1986, the starting point for each transect was randomly located in the center of a baygall using random numbers and pacing. The transect extended perpendicular from the baygall stream upslope to the end of the baygall veg- etation or sedimented area. Subsamples were established using metal poles at one meter intervals along the transect. Most of the original metal poles were still in place along the transects. In both sampling periods, nested subsamples were taken along the transects at each of the one meter markers. A 0.09 meter (0.3 meter x 0.3 meter) quadrat was used to sample the herbaceous plants. The sample was taken on the upstream side of each point. The shrubs (woody plants with a dbh (diameter at breast high at approximately 4.5 feet) of less than 4 inches and/or shorter than 25 feet) were sampled using a 1 meter square quad- rat with 1/2 meter on each side of the transect center line. The trees (woody plants 4 inches dbh or larger and/or 25 feet or taller) were sampled using 10 meter x 1 meter quadrat with 5 meters on each side of the transect center line. All species and total number of stems were recorded in each quadrat at each subsample. The dbh was recorded to the nearest 0.1 cm for woody taxa of suffi- cient height Some of the herbaceous taxa could only be identified to genus, family Pn ALLEN ET Al _ LOUISIANA 421 (Poaceae or Cyperaceae), or class. Data were > entered and analyzed USInE Mi- crosoft Excel software. The variables by summing the value for each mee and dividing by the number of subsamples, 37 for sedimented baygalls and 94 for control baygalls. Species rich- ness was calculated using all plant taxa. Individual density (dumber of indi- viduals per subsample) and stem density (number of stems per subsample) were calculated for herbaceous taxa and also for woody taxa. The dbh was also cal- culated for woody taxa. The population variables were calculated for all taxa by summing the value for each subsample and dividing by the number of subsamples, 37 for sedimented baygalls and 94 for control baygalls. The frequency (percentage of subsamples of occurrence) and mean value for individual density and stem density was calculated for all taxa. The mean value for dbh was calculated for the woody taxa of sufficient size. Relative frequency, relative individual den- sity, and relative stem density were calculated for all herbaceous taxa from the sedimented baygalls by summing the values for all herbaceous taxa in the sedimented area and dividing the value for each taxon by the total. The relative values for frequency, individual density, and stem density were calculated iden- tically for the herbaceous taxa from the control baygalls and for woody taxa from both control baygalls and sedimented baygalls. All relative values were then converted toa percentage. The importance value for each herbaceous taxon was calculated by summing the values for relative frequency, relative individual density, and relative stem density. The importance value for each woody taxon was calculated by summing the values for relative frequency, relative dbh, rela- tive individual density, and relative stem density. The total importance value for all herbaceous taxa is 300 and for the woody taxa is 400. Most plants were identified in the field by the senior author, and voucher herbarium specimens for most taxa were collected, mounted, and filed at Ft. Polk. Duplicates of many of the taxa were deposited in the Herbarium of the University of Louisiana at Monroe (NLU). Some taxa were identified in the labo- ratory using Allen, (1980 or 1992); Correll and Correll (1972); Correll and Johnston (1970); Diggs et al. (1999); Godfrey and Wooten (1979, 1981); or Rad- ford et al. (1968). The scientific names are from USDA, NRCS (2002). RESULTS The mean physiognomy community variables (species richness, dbh, woody individuals, woody stems, herbaceous individuals, and herbaceous stems) per subsample for the sedimented and control baygalls for 1986 plus 2001 are pre- sented in Table 1. The standard deviation is also given. The student’s t-test was used to compare variables between control 1986 and control 2001, sedimented 1986 and sedimented 2001, control 1986 and sedimented 1986, and control 2001 and sedimented 2001. Allen et al. (1990) reported that t-test comparisons showed 422 BRIT.ORG/SIDA 21(1) Taste 1. Community Physiognomy data (Species Richness, DBH, and Density) in control (94 Su Poalnipie>) ane sedimented baygalls (37 subsamples) for 1986 and 2001 at Fort Polk, La. All vari ables are significantly different at the 0.05 level between: control 1986 and control 2001;sedimented 1986 a sed ene 2001; control 1986 and sedimented 1986; and control 2001 and sedimented 2001, except where noted, Standard deviation is included in parentheses. — Control Baygalls Sedimented Baygalls 86 2001 1986 2 Species Richness 4.79 (2.36) 6.28(2.41) 7 .32(2.67) 5.00(2.12) No. Woody species 34 42 25 27 No. Herbaceous Species : 23 34 3 DBH (cm) 16° (23.54) 17.71°°(21.94) — 6.19(21.64) 14.19° (15.30) Woody Individual Density 7.13'(4.40) 12.80(5.94) 6.73° (4.34) 10.41(5.59) Woody Stem nsity 8.10°(5.05) 15.87(7.42) 7.35°(5.22) 11.32(6.14) Herbaceous Individual Density 1.94°(3.05) .38°(1.64) 15.05(8.35) 0.16(0.55) Herbaceous Stem Density 4.219 (7.82) 424° (6.10) 22.30(19.41) 0.19(0.62) * Not significantly different between 1986 and 2001 control baygalls ° Not significantly different between 2001 control and 2001 sedimented baygalls “Not significantly different between 1986 and 2001 sedimented baygalls all variables between the control 1986 and sedimented 1986 data to be signifi- cantly different. However, when the data for the relocated samples only are used, the woody individual density and woody stem density were not significantly different between the control and sedimented baygalls in 1986. When the con- trol 1986 data are compared to the control 2001 data, the species richness, woody individual density, and woody stem density were the variables that were sig- nificantly different. A comparison of the 1986 sedimented data to the 2001 sedimented data reveals that all six variables are significantly different. In the 2001 control and sedimented data, dbh was not significantly different while the other five variables were all significantly different. In Table 1, the variables that are not significantly different are indicated by: if between 1986 and 2001 control baygalls; > if between 2001 control baygal is and 2001 sedimented baygalls, and © if between 1986 control baygalls and 1986 sedimented baygalls. The most dramatic changes from 1986 to 2001 occurred in the sedimented baygalls with great increases in dbh (6.19 cm to 14.19 cm), woody plant indi- viduals (6.73 to 10.41), and woody plant stems (7.35 to 11.32). Large decreases occurred in herbaceous individuals (15.05 to 0.16) and herbaceous stems (22.30 to 0.19). The importance values for each woody taxon in the control and sedimented baygalls for 1986 and 2001 are in Table 2. In 1986, the woody taxa with the highest ——" ALLEN ET AL., CHANGES IN BAYGALL VEGETATION AT FORT POLK, LOUISIANA 423 Tase 2. List of woody taxa and importance value from control baygalls and sedimented baygalls for 1986 and 2001. 1986 2001 Taxon control sedimented control sedimented Acer rubrum 17.87 22.27 17.06 7.54 Alnus serrulata 0.00 18.32 4.27 0.00 Aronia arbutifoli 5.59 1.74 8.52 0.00 Selene seanae 7.13 7.74 6.10 5.51 p 1.12 0.00 8.22 15.03 Ca illicarpa americana 2.50 1.74 483 2.11 Cephal ats occidentalis 0.00 26.42 0.00 0.00 Chionanthus VIFgINiCUs 0.56 0.00 1.19 0.00 Cornus florida 0.56 0.00 0.35 0.00 Cyrilla racemiflora 0.00 0.00 0.70 0.00 Gelsemium sempervirens 757. 4.26 5.88 3.90 Hamamelis virginiana 0.00 0.00 0.70 0.00 Hypericum spp. 0.56 1.74 0.35 0.00 [lex coriacea 22.45 0.00 35.00 3.08 llex opaca 7.9) 1.74 5.62 0.00 [tea virginica 1.68 26.94 4.94 0.00 Ligustrum sinense 0.00 774 0.00 4747 Liquidambar styraciflua 5.95 35.10 1.69 7.58 Lonicera japonica 0.00 0.00 0.00 6.5] Lyonia luci 23.00 0.00 3372 0.00 Magiiole weit 52.14 6.78 37.04 20.79 M 19.6] 1.74 13.62 1.05 Morella cerifera 4,76 16.27 0.65 22.91 Nyssa sylvatica 78.30 148.97 51.69 69.28 Parthenocissus quinquefolia 0.00 0.00 0.35 0.00 Persea palustri 23.37 1.74 24.50 3.16 Pinus palustris 0.00 0.00 2.76 0.00 Pinus taeda 22.15 2.52 22.04 49.00 Quercus alba 0.00 0.00 1.34 0.00 Quercus falcata 0.56 0.00 0.00 0.00 Quercus nigra/laurifolia 4.73 22.5] 6.95 9.32 ae seedling 0.00 0.00 0.35 0.00 hododendron spp. 7.12 0.00 4.85 0.00 Hi copallina 0.00 1.74 0.35 0.00 Rubus spp 8.13 6.96 18.60 2.11 Salix nigra 0.00 9.85 0.00 0.00 dum 1.12 0.00 0.00 0.00 Smilax glau 5.88 0.00 8.92 1.05 Smilax Sons 9.46 0.00 8.32 6.27 Smilax rotundifolia 28.43 4.26 5.52 23.70 Smilax smallii 0.00 0.00 1.46 0.00 Sion tamnoides 0.00 0.00 0.00 1.05 n radicans 9.24 16.64 21.31 60.40 Toxicodendron vernix 1.68 0.00 0.85 1.05 424 BRIT.ORG/SIDA 21(1) Tasb_e 2. continued 2001 Taxon control sedimented control sedimented Vaccinium arboreun 0.00 0.00 0.00 1.55 Vaccinium arkansanum 1.12 0.00 0.91 0.00 Vaccinium elliottii 1,96 0.00 2.30 2 Vib lentatu 4.33 4.26 4.23 18.32 Viburnum nudum 11.45 0.00 20.69 8.17 Vitis rotundifolia 0.00 0.00 0.70 0.00 importance value (>20.00%) in the control baygalls were Nyssa sylvatica, Mag- nolia virginiana, Smilax rotundifolia, Persea palustris, Lyonia lucida, Ilex coriacea, and Pinus taeda and in 2001, the top taxa were Nyssa sylvatica, Mag- nolia virginiana, llex coriacea, Lyonia lucida, Persea palustris, Pinus taeda, Toxi- codendron radicans, and Viburnum nudum. In the sedimented baygalls in 1986, the most important taxon was Nyssa sylvatica with an importance value of 148.97%, which was almost five times as much as the second most important taxon. In 2001, Nyssa sylvatica was still the most important taxon, but its im- portance value was only 69.28%. In the sedimented baygalls in 1986, other taxa with importance value greater than 20.00% were Liquidambar styraciflua, Itea virginica, Cephalanthus occidentalis, Quercus nigra/laurifolia, and Acer rubrum. In the same sedimented baygalls in 2001, the taxa with more than 20.00% im portance value were Ioxicodendron radicans, Pinus taeda, Ligustrum sinense, Smilax rotundifolia, Morella cerifera,and Magnolia virginiana. The importance value for each herbaceous taxon in the control baygalls lor 1986 and 2001 are in Table 3. In 1986, the taxon with the highest importance value was Poaceae followed by Rudbechia scabrifolia, Dichanthelium tenue, Viola primulifolia, and herbaceous dicotyledons. In 2001, the five taxa with the highest importance value (in decreasing order) were Dichanthelium tenue, Wood wardia aerolata, Rudbeckia scabrifolia, Carex leptalea,and Chasmanthium laxum. In 2001, there were only three herbaceous taxa observed in the sedimented baygalls (Poaceae 137.91%, Chasmanthium laxum 108.06%, and Mitchella repens 54.03%): this contrasted tremendously with the 43 taxa re- corded in 1986 in all subsamples (Allen et al. 1990) and the 34 taxa in the relo- cated subsamples. The taxa with the highest importance value in 1986 were Poaceae, Lycopus spp., Herbaceous dicotyledons, Juncus diffusissimus, and Juncus coridceus. CONCLUSIONS In 1986, the data indicated that sedimentation had a significant effect on the baygall vegetation community. The student's t test showed four variables to be ALLEN ET AL., , LOUISIANA 425 Taae 3.List of herbaceous taxa and importance value from control baygalls for 1986 and 2001. Taxon 1986 2001 Apteria aphylla 1.89 0.00 Arisaema triphyllum 3.78 8.46 Arnog| ta 0.00 3.47 Athyrium felix-femina 4.01 9.62 Carex folliculata 0.00 15.72 Carex leptalea 0.00 24.50 Chasmanthium laxum 497 17.74 Coreopsis gladiata 10.19 0.00 peraceae 2.70 6.42 Dichanthelium acuminatum 0.00 2.95 Dichanthelium commutatum 1.89 0.00 Dichanthelium dichotomum 0.00 4.22 Dichanthelium tenue 43.36 61.67 Erochtitec hieracitoliq 1.89 0.00 Fupatorium leucolepis 0.00 2.20 iy 9) atoritum rotundifolium 1.89 0.00 Herbaceous dicot 27.29 0.00 Lachnocaulon anceps 3.50 0.00 Lobelia reverchonii 1.89 2.20 Lycopus spp. 0.00 2.20 Melanthium virginicum 2.65 0.00 Mitchella repens 14.29 16.32 Osmunda ci q 5.80 10.32 Poaceae 4772 2.20 Rhynchospora rariflora 4.72 0.00 Rudbeckia scabrifolia 43.98 29.89 Scleria oligantha 0.00 2. Solidago patula 1.89 9,39 Symphyotrichum spp. 0.00 4.23 Viola primulifolia 42.30 9.07 Woodwardia areolata 16.42 52.29 Woodwardia virginica 1.89 2.20 Xyris Spp. 9.08 0.00 highly significantly different between the sedimented and control or non- sedimented baygalls. Species richness and herbaceous individuals and stems had increased in the sedimented baygalls while the dbh had decreased. Sedi- mentation had killed many of the larger woody plants which explained the decrease in the dbh. The increase in species richness and herbaceous individu- als and stems probably occurred because the addition of sedimentation cre- ated a more mesic habitat. When the two areas (control and sedimented) were compared again in 2001, the dbh had increased in the sedimented baygalls and was no longer different 426 BRIT.ORG/SIDA 21(1) (Table 1). A comparison of the sedimented area 1986 data to the 2001 data re- vealed that the species richness had decreased from 7.32 to 5.00 species per sample. The number of woody individuals and stems had almost doubled and the dbh had more than doubled in the sedimented baygalls. The biggest changes inthe sedimented baygalls occurred with the herbaceous plants where the mean number of individuals decreased from 15.05 to 0.16 and the mean number of stems decreased from 22.30 to 0.19 stems. These changes in the sedimented baygall indicate that this area is changing from an open area toa forested area. The increase in trees and shrubs has created more shade and caused a large decrease in the number of herbaceous plants. The number of herbaceous spe- cies decreased from 34 to 3. In the control baygalls between 1986 and 2001, species richness, woody individuals, and woody stems had all increased significantly and the dbh, her- baceous individuals, and herbaceous stems had remained fairly constant. These changes and lack of change could be attributed to normal succession in the baygalls. Nyssa sylvatica is atree that seems to be little affected by sedimentation, as its importance value is very high in the sedimented baygalls in 1986 (Table 2). The tree (Liquidambar styraciflua), shrubs (Alnus serrulataand Cephalanthus occidentalis), and vine (Toxicodendron radicans) seem to grow better in the sedimented areas as indicated by the increase in their values. Tree species (Mag- nolia virginiana and Pinus taeda), shrub species (Persea palustris and Ilex coridcea), and the vine (Smilax rotundifolia) seem to have been affected by sedi- mentation as their values decreased in the sedimented baygall. In 2001 in the control baygalls, the shrubs Ulex coriacea and Lyonia lucida) had increased in importance value and is probably linked to natural succession in the baygalls. The sedimented baygalls had undergone more dramatic changes between 1986 and 2001 where early successional species (Liquidambar styracifluaand Alnus serrulata) showed a tremendous decrease in im portance value; Alnusserrulata was completely absent in 2001 and Liquidambar styraciflua decreased from an importance value of 35.10% to 7.58%. The shade tolerant vines (Toxicoden- dron radicans and Smilax rotundifolia) increased greatly in importance val- ues. The introduced species (Ligustrum sinense) had a large increase in impor- tance value from 7.74% in 1986 to 47.47% in 2001. Most of these changes seemed to be linked to succession in a sedimented area except for the introduced spe- cies filling in the niches normally occupied by native shrubs. The notable change in the herbaceous taxa in 1986 was the increase in the number of weedy taxa in the sedimented baygalls that were not present in the control areas or were present in small numbers (Allen et al. 1990). Some of the weedy taxa include several species of Juncus and Solidago, Diodia teres, Ambro- sid artemisiifolia, and Bidens aristosa. The creation of a new habitat in the sedimented baygalls is the reason for the invasion by the weedy taxa. All of the — ALLEN ET AL , LOUISIANA 427 ferns (Athyrium felix-femina, Osmunda cinnamomea, Wood wardia aerolata,and Woodwardia virginica) were not found in any of the sedimented samples but were present in the control samples. All four showed an increase in importance value between 1986 and 2001 in the control baygalls. The fern taxa in the sedimented baygalls in 1986 apparently decreased because of increased sun exposure caused by the death of trees in the baygall. These fern species prob- ably increased in value in the control baygalls in between 1986 and 2001 as a result of increased shade. In 2001, the sedimented baygalls changed dramati- cally with virtually no herbaceous plants persisting. The dense shade produced by the trees and shrubs greatly decreased the number of herbaceous plants, especially weedy species. ACKNOWLEDGMENTS We are indebted to Fort Polk, the University of Louisiana at Monroe, and the anonymous reviewer. REFERENCES Atven, C.M. 1980. Grasses of Louisiana. Univ. Southwestern Louisiana Press, Lafayette. Aten, C.M. 1992. Grasses of Louisiana 2"? ed. Cajun Prairie Habitat Preservation Society, Eunice, LA. Aten, C.M.,C.H.Staac and S.D. Parris. 1987. Analysis of the vegetation in pitcher plant bogs in two baygalls at Ft. Polk in west central Louisiana. Proc. Louisiana Acad. Sci.50:1-6. Auten, C.M., H.D. Guittory, C.H. Stace, and S.D. Paris. 1990. The effects of sedimentation on the vegetation and flora of baygalls in west central Louisiana. Proc. Louisiana Acad. Sci. 53:33-41. Corrett, D.S. and M.C. JoHNnston. 1970. Manual of the vascular plants of Texas. Texas Re- search Found., Renner. Corrett, D.S.and H.B.Corrett. 1972. Aquatic and wetland plants of the southwestern United States. Environmental Protection Agency, Washington. Dies, G.M., B.L. Lirscome, and R.J. O’Kennon. 1999. Shinners and Mahler's illustrated flora of north central Texas. BRIT, Fort Worth. Goorrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of Southeastern United States: M otyledons. Univ. Georgia Press, Athens, Georgia. Goorrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of Southeastern United States: Dicotyledons. Univ. Georgia Press, Athens, Georgia. Ravroro, A.E., H.E. Adtes, and C.R. Beit. 1968. Manual of the vascular flora of the Carolina. Univ. North Carolina Press, Chapel Hill. USDA, NRCS. 2002. The PLANTS database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. 428 BRIT.ORG/SIDA 21(1 BOOK NOTICES Blackwell Publishing Robert L. ZIMDAHL. 2004. Weed-Crop Competition, A Review. Second Edition. (1S 0-8138-0279-2, hbk). Blackwell Publishing Professional, 2121 State an Ames, IA 50014, U.S.A. (Orders: 1-800-862-6657, 1-515-292-3348 fax, c www.blackwellprofessional.com). $54.99, 224 pp., graphs, 7" x 10" Publisher Comments: “For the past 20 years, the first edition of this text has been widely cited as an authoritative academic reference. The latest edition continues the tradition set by the original book and covers weed science resez — aed) aas been published since 1980. T ‘his. book aims to reduce on instance of resear n—saving scientists and supporting ins s time and money.” “Not aie does the second edition of Weed-Crop Competition review, summarize, and combine current research, it critiques the research as well. This text has the potential to accelerate advance- 1 remains an important factor dss: crop ee . Weed clentists, Crop scientists, plant ecologists, sustainable agric ulturists, a g g will ments in weed- “Crop p competition, whic be well-pleased with this long overdue and much needed new edition.” Contents: 1) Introduction: An historical Perspective. 2) Definition of Plant Competition. 3) Com- petition in the Community. 4) Influence of Competition on the Plant. 5) The Effect of Weed Density. 6) The Effect of Competition Duration. 7) The Elements of Competition. 8) Weed Management Us- ing the Principles of Competition. 9) Methods Used to Study Weed-Crop Competition. 10) Models and Modeling. 11) Conclusion: The Complexity of Competition. Nets R. LERSTEN. 2004. Flowering plant Embryology. (ISBN 0-8138-2747-7, hbk.). Blackwell Publishing Professional, 2121 State Avenue, Ames, IA 50014, US.A. (Orders: 1-800-862-6657, 1-515-292-3348 fax, wwwblackwellprofessional.com). $79.99, 224 pp., b/w photos, line drawings, 7" x 10" Publisher Comments: “This richly illustrated reference text, with more than 350f ‘igures, presents eral angiosperm embryology using examples from economically important plants. The unique oe cus of this book on economically significant plants makes Flowering Plant Embryologya “must-have” for all a int science researchers and students.” Drawing from his career of teaching botany, and his original embryological research, Lersten emphasizes the plant species that affect human liveli no inclu ling weeds, ornamentals, and other cultivated plants that are used for commercial products. Topics are treated witha uniformity of style jes that boosts comprehension, and technical terms are well-defined. The book presents research re sults, hypotheses, and speculations about why things are as they are, and also offers supporting facts and — ific examples. This book ‘Prov idesalirm foundation for thorough ancien oe embryo- logical diversity plants and ane in the library of every plant scientis ents: D caoem tion. 2) Stamen and Androecium. 3) Pollen I Development: Theme and Varia: a tions. 4) Pollen Development: Details of Stages. 5) Carpel and Gynoecium. 6) Ovule and Embryo Sac. 7) Polli 1 Pollen: Stigma Interaction. 8) Pollen Germination, Pollen Tube Growth, and Double Fertilization. 9) Endosperm. 10) The Embryo. SIDA 21(1): 428. 2004 VASCULAR FLORA OF HACKBERRY FLAT, FREDERICK LAKE, AND SUTTLE CREEK, TILLMAN COUNTY, OKLAHOMA Bruce W. Hoagland! and Priscilla H.C. Crawford Phillio 1. Crawford Oklahoma Biological Survey and Oklahoma Biological Survey and Department of Geography Department of Botany and Mi aie University of Oklahoma University of Oklah 111 East Chesapeake St. 111 East Luceahe ae : Norman, Oklahoma 73019, U.S.A. Norman, Oklahoma 73019, U.S.A. Forrest Johnson rene Ok lahor na Biol urv psd of Ok ee 1 East Chesapeake St. a Oklahoma 73019, U.S.A. ABSTRACT The objective of this study is to fill a floristic data gap for southwest Oklahoma. Based upon a review of the Atlas of the Flora of Oklahoma database, it was noted that Tillman County was floristically under-documented. This paper reports the results of an inventory of the vascular plants in Tillman County. A total of 371 taxa of vascular plants in 253 genera and 74 families were collected. The most species were collected from the families Asteraceae (65), Poaceae (67), and Fabaceae (25). There were 123 annual and 248 perennial species. Forty-one species of woody plants were present. Forty-two exotic species were collected representing 11% of the flora. A total of 266 previously unreported spe- cies were documented. Five species tracked by the Oklahoma Natural Heritage Inventory were lo- cated, none of which were Federally listed as threatened or endangered. RESUMEN El objeti ste estudio es completar los datos floristicos del suroeste de Oklahoma. Basados en una revision nde la base ee dates del Atlas se the Flora of Oklahoma, se ae el Tillman County publican los resultados de un inventario de las plantas vasculares en Tillman County. Se colectaron un total 371 taxa de ple scul Ulares Ge 253 géneros y 74 familias. La tae oria de las especies se colectaron de las oak Meee (65), Poaceae (67), y Fabaceae (25). Habia 123 especies anuales y 248 perennes. Estaban eaiaed cuarenta y una especies de plantas pee Se colectaron cuarenta y dos es hi uae el 11% de la flora. Se documentan un total de 266 especies no citadas pre | 7iamente Se especies con seguimiento por el Oklahoma Natural Heritage Inventory, ninguna d listada federalmente como amenazada 0 en peligro e las cuales estaba INTRODUCTION North American botany has had a long tradition of floristic exploration and inventory (Ertter 2000a). Nevertheless, floristic inventories are of continued learracnondinga thor: (40N5)3225-1985 e-mail: bhoagaland@oi J Vv 1700, SIDA 21(1): 429 — 445, 2004 430 BRIT.ORG/SIDA 21(1) value for research, conservation, and management purposes (Palmer et al. 1995). For example, it has been recently documented that new taxa are discovered and described at a rate of 60 per year (Ertter 2000a). Inventories are also crucial to biogeographic research by filling gaps in the geographic distribution of taxa at all levels. Floristic inventories play a role in plant species conservation, both in locating populations of rare and/or undescribed species and bringing their presence to the attention of conservation organizations (Radford et al. 1980; Stuessy & Sohmer 1996). The lack of accurate florisitc data can jeopardize the long term persistence of sensitive species (Ertter 2000a). Finally, floristic in- ventories aid resource managers in locating populations of sensitive species and documenting the arrival of exotic and nuisance species (Barkley 2000). Igno- rance of the presence of exotic species can be detrimental to sensitive species and/or exert adverse economic impacts (Ertter 2000b). The objective of this study was to fill a gap in floristic data for southwest Oklahoma. Based on the Atlas of the Flora of Oklahoma database (AFO:; Hoagland 2003), Tillman County is a floristically under-documented county. Prior to 1996, the year collecting began for this study, only 175 species were re- ported from Tillman County (Hoagland 2003). The first collection gathered in Tillman County was a specimen of Eryngium diffusum by G.W. Stevens on 17 October 1924. Peak years for plant collecting in Tillman County prior to this study were 1936 (30 specimens) and 1940 (45 specimens). Study Area Tillman County (Fig. l) occupies 237,503 hectares and is located within the Subtropical Humid (Cf) climate zone (Trewartha 1968). Summers are warm (mean July temperature = 28.9° C) and humid, and winters are relatively short and mild (mean January temperature = 3.5° C). Mean annual precipitation is 78.7 cm., with periodic severe droughts (Oklahoma Climatological Survey 2003). Physiographically, the study area is located in the Osage Plains section of the Central Lowlands province (Hunt 1974) and within the Central Redbed Plains province of Oklahoma (Curtis @ Ham 1979). The surface geology of Tillman County is predominately red sandstone and shale formed from shallow-ma- rine and alluvial deposits of Permian age (Branson & Johnson 1979). There are eight soil associations in Tillman County (Lamar & Rhodes 1974). The two predominant soil associations are the Tipton-Hardeman-Grandfield, which occurs on the floodplains of the Red and North Fork of the Red River and is nearly level, loamy, sandy soil with loamy subsoil, and the Ford-Tillman, which is a nearly level to gently sloping upland, loamy soil with loamy and clay subsoils. The Clairemont-Asa-Miller Association occupies bottomlands and is deep, nearly level soil which is loamy, clayey throughout (Lamar & Rhodes 1974). The predominant potential natural vegetation in Tillman County (Duck S Fletcher 1943) includes the Mixedgrass Eroded Plains, which would occupy HOAGLAND ET Al Buses MAA es Frederick Lake Frederic Hackberry Flat Suttle Creek Grandfield Rive, pling in Tillman County, Oklahoma. Fic. 1. Location of tt 138,565 hectares (58%) on loamy soil and shallow rocky soil, and 68,635 hect- ares (29%) of tallgrass prairie (Duck & Fletcher 1943). Sandsage grassland oc- cupies 10,360 hectares (4%) and occurs on deep sand deposits along the North Fork of the Red River, Red River, and Otter Creek. Mesquite grassland occupies 4.403 hectares (2%). Bottomland forest occupies 15,540 hectares (6%). METHODS Collections were made at three locations: Hackberry Flat Wildlife Management Area (HF), and Lake Frederick (LF),a municipal reservoir, and Suttle Creek (SC) a Bureau of Land Management (BLM) holding along the Red River (Table LD. At each of these locations, collection sites were established for intensive floristic sampling. Sites were selected following a review of U.S. Geological Survey 1:24,000 topographic maps and field reconnaissance. The predominant vegeta- tion association at these sites was classified according to Hoagland (2000). Col- lections were also made randomly throughout each location and the county. Collections at HF were made from April to October, 1996, and at LF and SC from April through October 2000, from March through October 2001. Vouchers for exotic species (defined as those species not native to North America) were made 432 BRIT.ORG/SIDA 21(1) Taste 1: Location and el ation data fo rsample sites. oordinates represent the north and southern most era and the east an e west most longitude of each study site. Coordinates are presented in decimal deqrees. Site North South East West Max. Elev. Min. Elev. Frederick Lake 34.54°N 34.51°N 98.87°W 98.90°W 381m 360m Hackberry Flat 34.30°N 34.25°N 98.89°W 98.98°W 366m 349m Suttle Creek 34.23°N 34.22°N 98.96°W 98.97°W 358m 332m from naturalized populations only, thus excluding cultivated and ornamental plants. Specimens were processed at the Robert Bebb Herbarium of the Univer- sity of Oklahoma (OKL) following standard procedures. Manuals used for speci- men identification included Waterfall (1973), Great Plains Flora Association (1986), and Diggs et al (1999). Origin, either native or introduced, was deter- mined using Taylor & Taylor (1991) and USDA-NRCS (2003). Nomenclature fol- lows the United States Department of Agriculture-Natural Resources Conser- vation Service (USDA-NRCS 2003). A voucher set was deposited at OKL Sorensens’ Index of Similarity (Pielou 1984) was calculated for pairs of sites in order to determine similarity of floras. mth RESULTS AND DISCUSSION A total of 371 taxa of vascular plants in 74 families and 253 genera were col- lected in Tillman County. Among the angiosperms, 97 were monocots and 270 were dicots. In addition, one fern ally and one gymnosperm was recorded. The Asteraceae (65), Poaceae (67), and Fabaceae (25) had the greatest number of spe- cies. Genera with the greatest number of species were Dalea (6), Cyperus (5), Panicum (5) and Bromus (4). One hundred and ae ea species were an- nualsand 248 were perennials. Forty-one species of woody plants were collected: 20 trees, 13 shrubs, and 8 woody vines. Forty-two introduced species (11% of the flora) were collected in Tillman County. This iscomparable to the number of exotics collected at the Chickasaw National Recreation, located in south-central Oklahoma, where 12% of the flora was composed of exotic species (Hoagland & Johnson 2001). The families with the greatest number of introduced species were Poaceae (17), Brassicaceae (5), and Asteraceae (4). Genera with the most exotic species were Bromus (4) and Echinochloa (3). Species tracked by the Oklahoma Natural Heritage Inventory (ONHD were Abronia fragrans (G5, $253), Argythamnia humilis (G5,S2S3), Cenchrus echinatus (G5, SL), Escobaria vivipara (G5,52S3),and Malvella leprosa (G5, S1S2). Species are ranked according to level of imperilment at the state (S) and global (G) levels on a scale of 1-5; 1 representing a species that is imperiled and 5 one that it is secure (Groves et al. 1995). No Federally listed threatened or endangered species were encountered. — HOAGLAND ET AL, VASCULAR FLORA OF TILLMAN COUNTY, OKLAHOMA 433 Species richness was highest at LF (Table 2) and lowest at HF, which was by far the largest site (Table 3). The low species richness may be due to the fact that 97% of the land cover was classified as disturbed. Although LF was smaller in area than HE which had the lowest species richness, there were more habitat types present. SC, which had the second lowest species richness, was 65% dis- turbed (Table 3). LF was only 10% disturbed. However, the number of annual species, which often indicates the degree of disturbance, was highest at LF not HE Although all three sites were in close geographic proximity, Sorensen Indices were below 0.5 (Table 4). The highest similarity was scored for the comparison of HF and LE which shared 53 species (Table 5). The high number of arena- ceous species at SC may account for the low similarity index values between that site and the other two. Interestingly, the highest similarity was between the largest sites. The AFO database (Hoagland 2003) lists 175 species for Tillman County that were collected prior to 1996. Seventy species in the Atlas database were not collected in this study. There were eight families in the AFO database that were not collected in this study; Acanthaceae (Dyschoriste linearis and Justicia americana), Apocynaceae (Apocynum cannabinum), Cuscutaceae (Cuscuta cuspidata), Dryopteridaceae (Woodsia obtusa), Fumariaceae (Corydalis aurea), Polemoniaceae (Ipomopsis longiflora), and Pteridaceae (Pellaea atropurpurea). Of the remaining species, 107 were reported both in AFO database and in this study. When that number is subtracted from the total of species in the check- list, this study contributed 266 species previously unreported from Tillman County. When the species unique to the AFO and this study are summed, along with the number of shared species, this gives a total of 441 species in Tillman County, a 40% increase in our previous knowledge. Seven habitat types were found at the three primary collecting sites (Table 2). The actual number of habitat types at each site ranged from 3 to 6. A brief description of each habitat type follows. Sandbars and dunes (SB) Sandbars and dunes occurred only at the SC site. Vegetation on sandbars, which were in the Red River channel, was sparse, however Cyperus esculentus, Heliotropium curassavicum, and Tamarix chinensis were present. Sandunes, which occurred along the floodplain and terraces of the Red River, were veg- etated by the Artemisia filifolia/Sporobolus cryptandrus-Schizachyrium scoparium shrubland association (Hoagland 2000). Associated species included Dalea villosa, Prunus angustifolia, Calylophus serrulatus, Eriogonum anuum, Sideroxylon lanuginosa, Rhus aromatica, Vitis acerifolia, and Zanthoxylum hirsutum. Tracked species found in this habitat type were Abronia fragrans and Cenchrus echinatus. 434 BRIT.ORG/SIDA 21(1) Taste 2: Summary of floristic collections at three sites in Tillman County, Oklahoma. Format follows Palmer et al. (1 Taxonomic group Species Native spp. Introduced spp. Hackberry Flat C niferophyta 0 0 0 Magnoliophya Magnoliopsida 86 75 11 a sida 35 25 10 tal 121 100 21 ae ee Coniferophyta 1 | 0 Magnoliophya Magnoliopsida 144 130 14 Liliopsida Al 35 5 Total 185 166 19 Suttle Creek Coniferophyta 1 1 0 Magnoliophya Magnoliopsida 127 119 8 Liliopsida 54 45 9 I 182 165 17 Taste 3: Habitat types mapped at the three primary collection sites in Tillman County, Oklahoma. Area = total area of the site, HT = number of habitat era at the site, SB = sandbars and dunes, a = aquatic, DA = Old fields and disturbed areas, PS = pasture, MG = mixedgrass prairie, MQ = quite shrubland, and BLF = bottomland forest. (HF = cerns oo Management ee ie = Lake Frederick, SC = Suttle Creek). All values are reported in hectare Site Area HT SB AQ DA PS MG MO BLF HF 2,770 0 0 2,690 67 14 0 0 LF 911 6 0 341 95 ) 0 44] 26 SC 16] 5 17 9.6 105 @ 0.1 0 28 Sorensens' Index of Similarity values for three collection sites, Tillman County, Oklahoma (HF = = Boel beies Flat Wildlife Management Area, LF = Lake Frederick, SC = Suttle Creek). Suttle Creek Lake Frederick Hackberry Flat Suttle Creek 1 0.347 0.275 Hackberry Flat 0.352 1 HOAGLAND ET AL OF TILLMAN COUNTY, OKLAHOMA 435 Taste 5: An inter-site comparison of shared species in Tillman County, Oklahoma. Unique refers to species found only at the site listed. W/ = species shared with two sites. Site Total Unique w/LF w/HF Hackberry Flat 12] 45 Lake Frederick 185 87 53 Suttle Creek 182 8&8 63 4] Aquatic and wetland habitats (AQ) Aquatic habitats were found at LF and SC. Aquatic environments at LF were composed of the 341 hectare Lake Frederick and its shoreline. Vegetated areas were on the upper reaches of the lake, where the Polygonum pensylvanicum-P. lapathifolium herbaceous association (Hoagland 2000) was common. Associ- ated species included Eleocharis palustris, Juncus torreyi, Potamogeton pectinatus, Schoenoplectus pungens, and Xanthium strumarium. Wetlands at SC were formed by seeps and springs that emerged at the base of sand dunes and flowed into Suttle Creek. The vegetation in the seeps and springs was best characterized as the Rorippa nasturtium-aquaticum herba- ceous association (Hoagland 2000). Associated species included Eclipta prostrata, Hydrocotyle verticillata, Lobelia cardinalis, Ludwigia palustris, Myosurus minimus, and Ranunculus scleratus. Along broader stretches of Suttle Creek, the Schoenoplectus americanus - Eleocharis spp. herbaceous association (Hoagland 2000) was predominant. Associated species included Amorpha fruticosa, Cephalanthus occidentalis Distichlis spicata, E. palustris J. torreyi, Polypogon monspeliensis, Symphyotrichum subulatum and Typha domingensis. Disturbed areas and old fields (DA) Disturbed areas occurred at all three sites and was the predominant cover type at HF and SC. Disturbed area designations included mowed lawns, roadsides, and other sites exhibiting signs of physical disruption. Common plants in dis- turbed areas included Bothriochloa ischaemum, Cynodon dactylon, Daucus pusillus, Melilotus officinalis, and Mollugo verticillata. Old-fields were charac- terized by Ambrosia trifida, Amaranthus rudis, Cnidoscolus texanus, Conyza canadensis, and Sorghum halepense. Malvella leprosa was the only species tracked by ONHI found in this habitat type. Pasture (PS) This habitat type occurred only at HE where B. ischaemum had been planted on 67 ha in the northwest corner of the site. The pasture was essentially a mo- noculture of B. ischaemum witha few widely scattered individuals of Prosopis glandulosa. oe 436 BRIT.ORG/SIDA 21(1) Mixedgrass prairie (MG) Mixedgrass prairie occurred at all three sample sites, but was most extensive at LE Bouteloua hirsuta - Bouteloua curtipendula herbaceous association was com- mon on coarse, shallow soils. Associated species included Aristida purpurascens, Bouteloua rigidiseta, Crotonopsis elliptica, Leucelene asteroides, Lithospermum tenellum, Opuntia phaeacantha, Schizachyrium scoparium, and Thelesperma filifolia (Hoagland 2000, Crawford 2002). On loamy soils, the Schizachyrium scoparium - Sorghastrum nutans herbaceous association pre- dlominates. Common associates include Andropogon gerardii, A. purpurascens, Bouteloua curtipendula, Panicum virgatum, Sporobolus cryptandrus, and Symphyotrichum ericoides. Argythamnia humilis and Escobaria vivipara were found in this habitat type and also in mesquite shrubland. Mesquite shrubland (MQ) This habitat type, representing the Prosopis glandulosa/Bouteloua sp. shrubland association (Hoagland 2000), was found only at LE Unlike P. glandulosa shrublands throughout Tillman County, the herbaceous vegetation at LF was predominantly native species (Crawford 2002). Dominant grasses included Bouteloud curtipendula and Schizachyrium scoparium. Associated species in- cluded Aristida purpurascens, Bouteloua rigidiseta, Echinacea angustifolia, Erioneuron pilosum, Eryngium leavenworthii, Opuntia phaeacantha, Sorghastrum nutans, Sporobolus cryptandrus, Symphyotrichum ericoides, and Thelesperma filifolia. peed: ~— Bottomland forest (BLF) Bottomland forest occurred at LF and SC. Two vegetation types occurred in this category: Fraxinus pennsylvanica - Ulmus americana forest association and the Populus deltoides/Salix (exigua, nigra) forest association (Hoagland 2000). Associated species included Ampelopsis cordata, Celtis laevigata, Teucrium canadense, and Toxicodend ron radicans. ANNOTATED CHECKLIST Annotated species list for Tillman County, Oklahoma. The first entry is the col- lection number (the prefix BLM = Suttle Creek, a Bureau of Land Management holding on the Red River, HF = Hackberry Flat Wildlife Management Area, and PC = Lake Frederick). Specimens with the prefix M9 or the suffixes BWH or 98 represent specimens collected outside the three areas inventoried), followed by origin (N = native, I = introduced), life history (A = annual, Bi = biennial, P = perennial), and habitat (SB = sandbars and dunes, AQ = aquatic, DA = Old fields and disturbed areas, PS = pasture, MG = mixedgrass prairie, MQ = mesquite shrubland, and BLF = bottomland forest). Voucher specimens were deposited at the Robert Bebb Herbarium at the University of Oklahoma (OKL). HOAGLAND ET AL EQUISETOPHYTA EQUISETACEAE Equisetum laevigatum A. Braun; M9.134;N; P; AQ CONIFEROPHYTA CUPRESSACEAE Juniperus virginiana L;BLMO179,PC-138;N;P;DA, MG MAGNOLIOPHYTA-LILIOPSIDA AGAVACEAE Yucca glauca Nutt.; BLM064; N; P; MG ALISMATACEAE Sagittaria brevirostra Mackenzie & Bush; BLM0348; N; P; AQ CYPERACEAE Carex tetrastachya Scheele; BLM033, HF032;N;P; AQ Carex perdentata S.D. Jones; BLM045; N; P; AQ Cyperus acuminatus Torr.& Hook.ex. Torr; HFO1 58; de Cyperus squarrosus L.;0119-90; N; A; AQ Cyperus croceus Vahl; PC-168; N; A; DA Cyperus esculentus L.; HFO151;1;P; DA Cyperus odoratus L.; BLM0427;N;P; DA Eleocharis montevidensis Kunth.;BLM031;N;P;AQ Eleocharis palustris (L.) Roem. & Schult.; HFO019, Fuirena simplex Vahl; BLM0357;N; P; AO Schoenoplectus americanus (Pers.) Volk ex Schinz &R are oe P: AQ Schoenoplec pungens (Vahl) P ave var. longispicatus eo S.G: Simi; PG-1335NeP: AQ Schoenoplectus maritimus (L.) Lye; BLM0451; N; IRIDACEAE Nemastylis gemin eres NDE 2;N;P; DA, MG Hie HAGeNtep: Cp OOO ING Fe ‘DA, MG Sisyrinchium chilense Hook.; PC-16; N; P; DA JUNG ASE Is marginatus Rostk.; M9.132;N:P; AO Juncus torreyi Cov.; BLM0355, PC-132;N;P; AQ LILIACEAE Allium Canes ee BLM0076; N; P; DA, MG Ce f ; |b ! ler ber ts Erythronium al m Nutt.; PC-232; N; P; BLF Rothe ctorlune bivalve (L.) Britton; HFO166, PC-68; N; P: DA, MG, MQ NAJADACEAE Najas guadalupensis (Spreng.) Magnus; 2085- BWH;N; A; A BORCENE. indrica Host; HFO51;1; A; AQ inno gerardii Vitman; BLM0435, PC-200; Ancropogn glomeratus (Walt.) B.S.P.; BLM0424; fern, Aristida purpurascens Poir.; BLM0369;N;P: DA, MG se purpurea Nutt; BLM0161,HF0104,PC-27; A, eis oa ischaemum (L.) Keng var. songarica (Rupr.ex Fisch.& C.A oo Harlan; C-22;1;P; DA, P Bothriochloa ees ie Rydb.; HF047,PC- 60; N; P; DA, MG, MQ Bouteloua curtipendula (Michx.) Torr; BLM0363, HFO105, PC-172;N; P; MG, M hirsuta Lag.; PC-173; N;P; MG, MOQ pouteieua rigidiseta (Steud.) A. S. Hitchc.; BLMO180, ie PC-21;N;P; MG, MQ Br ath Vahl; BLM046, HFO36, PC-63; |; P: BLF, DA, MQ Bromus commutatus Schrad.; PC-46;1; A; DA Bromus japonicus Thunb. Ex Murr; BLM0509, O1:1:A; Bromus secalinus L.; HFO9; 1; A; DA Buc ee dactyloides (Nutt.) Engelm.; BLM051, 2,PC A Q B itelc Wd Ul Ud Pea tee echinatus L.; BLM032 2;N;A;SB Chloris cucullata Bisch.; BLM0O183; N; P; SB Chloris verticillata Nutt.; HFO103;N;P; DA, MG, MO Cynodon dactylon (L.) Pers; BLM0432, HFQ100, PC-128;|;P: DA, PS Digitaria sanguinalis (L.) Scop.;BLM0450;N;A;DA Distichlis spicata (L.) Greene; BLM031 2; N; P; AO, SB Echinochloa a (L.) Link; BLM0434: 1; A; AQ L.) Beauv,; BLM0431;1;A l é Echinochloa crus-galli ( Echinochloa crus-pavonis (H.B.K.) Schult. var. macera (Wieg.) Gould; PC-194; 1; A; AQ Echinochloa muricata (Beauv.) Fern.; HF0101; N; Elymus canadensis L., PC-114;N; P; BLF, MG, MQ Elymus virginicus L.;BLMO147,HFO110;N;P;MG,MQ Eragrostis cilianensis (All.) Vign. ex Janchen; M0328, HFO168;1; A; DA Eragrostis hirsuta (Michx.) Nees; BLM0449; N; P; Eragrostis hyena ides (Lam.) B.S.P; hie N;A;AQ Eragrostis sessilispica Buckl.; PC-76;N;P; DA Eriochloa contracta Hitchc.; HFO1 See ere Eriochloa sericea (Scheele) Munro ex. Vasey; PC- pe Erioneuron pilosum (Buckley) Nash; BLM028, PC- 9;N:P; MG Hordeum pusillum Nutt.; BLM044, HFO42, PC-29: N; A; DA Leersia oryzoides (L.) Sw.; BLM0430; N; P; AQ Leptochloa fusca (L.) Kunth.; BLMO511, HF0135 ;DA Leptochloa panicea (Retz.) Ohwi ssp. brachiata (Steudl.) N. Snow; HFO152, PC-192;N;P; DA Muhlenbergia asperifolia plea & Meyen ex Trin. Parodi; BLM0437; AQ Nassella leucotri ain ce Pohl;BLM0195, P MG te Neeragrostis ean (Michx.); PC-193;N; A; AQ Pani api L., HFO124, PC-160;N; A; DA Panicum sal fi HiVasey var. filipes (Scribn.) F.R. Waller; PC-169; N; P: MG Pani cael H.B.K.; HFO77, PC-156;N;P:BLF, Panicum rigi ae Bosc ex Nees; HFO122;N;P; MG P CGTHCUTTT virgatum i BLM0488, PC-1 99- N; P: BLF, MG, MQ rae Gy smithii (Rydb.) A.LOove; BLM0486, PC- N; P; AQ, BLF alum distichum L.; HFO150; N; P: DA a. setaceum Michx.; BLM0487 Pennisetum glaucum (L.) R.Br; BLM0429, HFO1 30; |. A; DA, Ph 7/ 1 ~BLMO161;1;A; AO Phalaris ues Walt.; N; A; AQ, MG Poa annua L.; BLM047;1: A; ae Poa arachnifera Torr.; BLM041; Polypogon monspeliensis (L ee ge: I;A; AQ /SB Schedonnardus paniculatus (Nutt.) Trel.; HFOSO; P:DA, MG Setaria parviflora (Poir.) Kerguelén; BLM0359: N; P:DA Setaria viridis (L.) Beauv.; BLMO500;1; A; DA Sorghastrum nutans (L.) Nash; BLM0433;N;P: MG — BRIT.ORG/SIDA 21(1) Sorghum halepense (L.) Pers.; BLM0484, HF041, PC-124;1;P: DA Sporobolus airoides ( Torr.) Torr; HFO79, PC-182:N; L.; HFO86, PC-18;1;P; DA CORNACEAE Cornus drummondii Mey.; 0132-98; N; P; DA, MG CUCURBITACEAE C -ucurb ita foetidissima Kunth.; HFO7O, PC- 131;N P;DA, MG, MOQ 80;1;A;DA ELATINACEAE Bergia texana (Hook.) Walp.; PC-137;N;P; MG cei neauainsibe tryifolia Riddell; PC-188;N;A;DA Arey eae bus (Engelm & A. Gray) Muell,; Arg var. leiosperma Waterfall; PC-78; N; P; MG HOAGLAND ET AL Chamaesyce albomarginata (Torr. & A. Gray) Small; HFO96; N; P; DA Chamaesyce serpens (Kunth) Small; HFO89; N; A; DA Chamaesyce missurica (Raf.) Shinners; BLM0377, PC A; Chamaesyce nutans (Lag.) Small;PC-174;N;A;DA Chamaesyce prostrata (Aiton) Small; PC-108; N; Small: Cnidoscolus texanus (Muell.-Arg.) BLMO1 3B Croton capitatus Michx.; PC-152;N; A; DA, MG Croton monanthogynus Michx.; PC-74; N; A; DA Croton texensis (Klotzsch) Muell.-Arg.; BLM0428; A, Euphorbia dentata Michx.;PC-73;N; A; DA ues a ioe: ioe Pursh; BLM0347, HFO143, PC-3 B Euphorbi Ele ae Lam.; BLMO50, PC-26;N;A; DA Phyl re ae lygonoides Nutt.ex Spreng; PC-98; MG ie fal oe Garden ex.L.; BLMO203;N;P;MG FABACEAE Acacia angustissima (Mill.) Kuntze. var. hirta (Nutt.) B.L. Rob.; HF093, ie 141;N;P; MG, MQ morpha hincosals BLMO1 -P-AQ ee, us lindheimeri ac ex A. Gray; PC-32; G, MQ Astagal us plattensis Nutt.; PC-28; N; P; MG, MO Chamaecrista fasciculata (Michx.) Greene; PC 204; N; A; MG, MQ Dalea aurea Nutt. ex Pursh; PC-116; N; P; MG Dalea candida Willd.; BLM0323, PC-122;N:P:MG, MQ Dalea enneandra Nutt.; BLM0321, PC-115; N; P: MG Dalea lanata Spreng.; BLM0306; N; P; SB alea purpurea Vent.; PC-91; N; P; MG, MQ Dalea villosa (Nutt.) Spreng.; BLM0305; N; P; SB ed illinoensis (Michx.) MacM.;BLM0344, HFO111;N;P;DA iw) Gleditsia triacanthos es BLMO73;N; P; BLF pid Pursh; BLM0310; N; P; SB omar ee (Ort.) Eifert; HFO39, PC- ndgteo miniata Ort. var. leptosepala; BLM0159, 206; N; a Medicago minima (L.) L; PC-31;1; A; DA TILLMAN COUNTY, OKLAHOMA 441 Medicago orbicularis (L.) Bartal.; HFO45;1; A; DA Melilotus officinalis (L.) Lam.;BLM0485, HFO27, PC- 95;1;A;D Mimosa nuttallii (DC.) B.L. Turner; BLMO189, ,PC-90; N:P; MG, MQ Neptunia lutea Benth; PC-100; N; P; MG, MQ Pediomelum cuspidatum (Pursh) Rydb.; PC-83;N; P; MG, MQ Pediomelum linearifolium (Torr. & A. Gray) J Grimes ;PC-95;N; P; MG,M Prosopis ee Torr; BLMO1 76, HFO68, PC- 64;N;P;DA,MG, MQ Vicia sativa L.; BLM053;1;A; DA GENTIANACEAE Eustoma Pees it BLM0341,PC-202;N GERANIACEAE Erodium cicutarium (L.) UHér. ex Aiton; BLM063, PC- A y Salisb. ex G. Don; AQ altar Geranium carolinianum L.; 2087-BWH, HF049; N; A; DA HYDROPHYLLACEAE Nal | am BLM0O153;N; A: MG MHOPTOUTEE Ulay, KRAMERIACEAE Krameria lanceolata Torr.; BLM0O167, HFO57, PC- LAMIACEAE Hedeorma reverchonii A. Gray; PC-126; N; P; MG, Lamium amplexicaule L.;BLMO74, PC-239;1; A;BLF, DA Monarda clinopodioides A.Gray; BLMO197,PC-69; Cry itellariaq driymr ndii oe N; A; MG 1; PC-5; N:P; MG Teucrium canadense L. seen 1,PC-129;N;P: BLF Teucrium laciniatum Torr; BLM0O149, HFO55, PC- 30;N; P: MG, MO LINACEAE Linum perenne L.; BLM069; 1; P; MG, MQ Linum pratense (J.B.S. Norton) Small; PC-103; N; Scutellari iresinosa lor A; MG LOASACEAE Mentzelia nuda (Pursh) Torr. & A. Gray var. stricta Seeiouy ean BLM0325;N; P; SB | t.ex Sims. PC-121;N; lig re PMG LYTHRACEAE Lythrum alatum Pursh; HF04; N; P; AQ Lythrum californicum Torr.& A. Gray; PC-130; N:P; AQ MALVACEAE Malvella leprosa (Ortega) Krapov.- HF03;N;P;DA Sphaeralcea coccinea (Nutt.) Rydb.;PC-7;N;P:MG, MQ MENISPERMACEAE Cocculus carolinus (L.) DC.; BLM0309; N; P; DA MOLLUGINACEAE Mollugo verticillata L.; HFO75;N;A;DA MORACEAE Maclura ene (Raf.) Schneid., BLM0318, HFO69; N; P; BLF, DA Morus alba L: BLMO5SQ7, HFO117;1-P; DA NYCTAGINACEAE Abronia fragrans Nutt.ex Hook.; BLM057;N; P; SB ed lisa he (Walt.) Heimerl;BLMO172,PC-162; Mab apa HFO153;1;P; DA { Purs sh) Heimer; H 072;N;P;MG, MO OLEACEAE Fraxinus pennsylvanica Marsh; PC-150; N; P; BLF ONAGRACEAE Calylophus hartwegii (Benth.) Raven subsp. pubescens (A. Gray) Towner & Raven; PC-33; N:P; MG Calylophus serrulatus (Nutt.) Raven; BLM0202;N; B Gaura coccinea Nutt. ex Pursh; BLM066, HF044- »MG, SB Gaura parviflora Dougl.ex Lehm.;PC-191, HF025; N; A; D Gaura sinuata Nutt. ex. Ser; PC- 158; N; P: MG, MQ se ae ides (Kunth) Raven; 2089-BWH;N; ince repens J.R. Forst.; BLMO198; N; P; AQ Oenothera grandis Britton; BLMO80, 062-97, PC- 1;N;A;MG, SB nothera rhombipetala Nutt. ex Torr. & A. Gray; BLM0378; N; Bi DA, SB Oenothera speciosa Nutt.; HFO37, PC-34; N:P: DA, MG Oenothera triloba Nutt.; M9.026; N; Bi; MG, MOQ BRIT.ORG/SIDA 21(1) Stenosi ae poe (Nutt. ex James) Heynh.; BLM C-89; N; P: DA, MG, MQ OXALIDACEAE Oxalis corniculata L.; PC-246;N; P; DA PAPAVERACEAE Argemone polyanthemos (Fedde) G.B. Ownbey; BLMO185;N; A; SB PEDALIACEAE Proboscidea louisianica (P. Mill) Thell.; HFO88, PC- 211;N;A;DA PLANTAGINACEAE Plantago rhodosperma Decne.;BLM079, PC-25;N; POLYGALACEAE Polygala alba Nutt. BLM0148, PC-48; N; P; MG POLYGONACEAE gonum annuum Nutt.; BLM0354; N; A; SB Fringannitm lonaifoliiim Nutt; PC-] 7S: N: p. MG ee hydropiperoides Michx.; BLM0422:N; oe Polygonum lapathifolium L.; HFO148; N; A; AQ a penaivani icum L.; HFO24; N; A; AQ Polygonum ramosissimum Michx.; HFO160; N; A; DA Rumex altissimus Wood; HF021;N;P: AQ, DA Rumex crispus L.; HFOS5; |; P; AQ, DA, BLF PORTULA ACACEAE Portulaca oleracea L.; HFO137;N;A;DA PRIMULACEAE Androsace occidentalis Pursh; PC-248; N; A; DA Samolus ebracteatus Kunth; BLMO184- N; P: AQ Samolus valerandi L.; BLM0503; N; P; AQ RANUNCULACEAE Anemone berlandieri Pritz; PC-23; N; P; MG Ale one caroliniana Walt.; PC-227; N:P; DA, MG avis carolinianum Walt. ssp. virescens (Nutt.) Brooks; PC-20; N; P; MG, MQ Myosurus minumus L.; PC-234; N; A; AQ Ranunculus sceleratus L.; BLM034; N; A; AO RHAMNACEAE Ziziphus obtusifolia (Hook.) A. Gray; BLMO158; N; ROSACEAE Crataegus viridis L.; M9.129; N; P; BLF HOAGLAND ET AL, VASCULAR FLORA OF TILLMAN COUNTY, OKLAHOMA Prunus angustifolia Marshall; HFO52,PC-139;N;P; DA,MG Rubus trivialis Michx.; BLM0O37; N; P; BLF RUBIACEAE Cephalanthus occidentalis L.; PC-119;N;P; AQ Galium aparine L.; BLM039; N; A; BLF Hedyotis nigricans (Lam.) Fosberg; BLM0191, PC- 82;N;P; MG Houstonia pusilla Schoepf; PC-226; N; A; DA RUTACEAE Zanthoxylum hirsutum Buckl.; BLMO160; N; P; SB ALICACEAE Populus deltoides Marsh. ssp. monilifera (Aiton) w.; PC-19; N; P; BLF Populus de ides Bartr. ex Marsh; HF0128,PC-251; N;P; Salix Nutt.; BLM0308, N; P; SB Salix nigra Marsh; BLM0314, HFO29, PC-120; N;P; AQ, BLF, SB SAPINDACEAE Cardiospermum halicacabum L.; BLM0339; N; A; FDA Sapindus saponaria L. var. drummondii (Hook. & rn) L.D. Benson; HF026, PC-149;N;P;BLF, SB SAPOTACEAE Sideroxylon lanuginosum Michx.; BLMO173, PC 207;N;P; SB SCROPHULARIACEAE Lindernia ae (L.) Pennell; M9.136;N; A; AO P a Nutt; PC-51;N;P; MG \ ica anagallis-aquatica L.; BLMO52; N; P; AQ Veronica peregrina L.; BLM081;N; A; DA SIMAROUBACEAE Ailanthus altissima (P. Mill.) Swingle; 0489-98; |; P; SOLANACEAE Chamaesaracha coniodes Moric.ex Dunal;PC-58; N; P; Physalis angulata L.; HFO123;N;A;DA Physalis longifolia Nutt.; BLMO351, PC-97;N;P:DA Physalis mollis Nutt. var. mollis, HFO133, PC-167, N;P;DA Quincula lobata (Torr.) Raf; HFO13,PC-52;N;P:DA; MQ Solanum dimidiatum Raf.; BLMO153, PC-87; N; P; DA Solanum elaeagnifolium Cav.;BLM0169, PC-50;N; Solanum rostratum Dunal;BLM0493, HFO92;N;A; TAMARICACEAE Tamarix chinensis Lour.; HFO20, PC-198; |; P; AQ, ULMACEAE Celtis laevigata Willd. var. texana (Scheele) Sang.; HFO80, poe 110;N;P;B Ulmus ame aL. BLMO337, PC-253; N; P; BLF URTICACEAE Parietaria pensylvanica Muhl.ex Willd.; BLMO1 70; pact VALERIANACEAE Valerianella radiata (L.) Dufr.; BLM083; N; A; DA VERBENACEAE Phyla lanceolata (Michx.) Greene; BLM0199;N; P; AQ Phyla nodiflora (L.) Greene; BLM0330, PC-127;N; P: AQ Glandularia bipinnatifida (Nutt.) Nutt; PC-12; N; A; DA Verbena plicata Greene; BLM060; N; P; MG Verbena scabra Vahl: BLM0423; N;P; MG VIOLACEAE Viola bicolor Pursh; PC-225;N; A; DA ISCACEAE Phoradendron tomentosum (DC.) Engelm. ex A. Gray; BLM0164, PC-250; N; P; BLF VITACEAE Ampelopsis cordata Michx.; BLM0362; N; P; BLF Cissus Hifol iata (L.) L.;BLM072;N;P; MG Parthenocissus quinquefolia (L.) BLM0350; N; P; BLF Planch.; Vitis acerifolia Raf., 0134-98; N; P; SB Vitis riparia Michx.; 093-98; N; P; SB Vitis vulpina L.; PC-208; N; P; MG, MQ ACKNOWLEDGMENTS This project was funded by a grants from the Oklahoma Department of Wild- life Conservation, and the Bureau of Land Management. We thank Debbie 444 BRIT.ORG/SIDA 21(1) Benesh, Amy Buthod, and Newell McCarty for assistance in the field. Todd Fagin produced the study area map. REFERENCES BarxLey, T.M. 2000. Floristic studies in contemporary botany. Madrono 47:253-258. Branson, C.C., and K.S. JoHNson. 1979. Generalized geologic map of Oklahoma. |n:K.S. et al, eds. Geology and earth resources of Oklahoma. Oklahoma Geological Survey, Norman. Curtis, N.M.and W.E. Ham. 1979.Geomorphic provinces of Oklahoma. |n:K.S. Johnson et al., eds. Geology and earth resources of Oklahoma. Oklahoma Geological Survey, Norman. Diccs,G.M.,B.L. Liescome, and R.J.O’Kennon.1999. Illustrated flora of north central Texas. Sida, Bot. Misc. 16:1-1626. Duck, L.G., and J.B. FuetcHer. 1943.A game type map of Oklahoma. A Survey of the Game and Furbearing Animals of Oklahoma. Oklahoma Department of Wildlife Conserva- tion, Oklahoma City. Ertrer, B. 2000a. Our undiscovered heritage: past and future prospects for species-level botanical inventory. Madrono 47:237-252 Ertrer, B. 2000b. Floristic surprises in North America north of Mexico. Ann. Missouri Bot. Gard. 87:81-109. GREAT PLAINS Flora Associarion. 1986. Flora of the Great Plains. University Press of Kansas, Lawrence. Groves, C.R., M.L. Kuein, and T.F. Breven. 1995. Natural heritage programs: public-private partnerships for biodiversity conservation, Wildl. Soc. Bull. 23:784-790. Hoactano, B.W. 2000. The vegetation of Oklahoma:a classification of landscape mapping and conservation planning. Southw. Nat. 45:385-420 HoacLanb, B.W. 2003. Atlas of the flora of Oklahoma. (www.biosurvey.ou.edu) Oklahoma Biological Survey, University of Oklahoma, Norman. Hoactanp, B.W. and F.L. JoHnson. 2001. Vascular flora of the Chickasaw National Recreation Area, Murray County, Oklahoma. Castanea 66:383-400, Hunt, C.B. 1974. Natural regions of the United States and Canada. WH. Freeman, San Francisco. Lamar, O.W. and C.E. RHoves. 1974. Soil survey of Tillman County, Oklahoma. United States Department of Agriculture. OKLAHOMA CLIMATOLOGICAL Survey, 2003. Oklahoma Climatological Data. (http:// www.ocs.ou.edu/). University of Oklahoma, Norman. Pictou, E.C. 1984. Interpretation of ecological data. Wiley Interscience, New York. Patmer, M.W., G.L. Wave, and P Neat. 1995. Standards for the writing of floras. BioScience 45:339-345. Raprorb, A.E., D.K. Orte, J.R. Massey, and PD. Wuitson. 1980. Natural heritage: classification, inventory, and information. University of North Carolina, Chapel Hill. Stuessy, T.F.and S.H. SoHmer. 1996. Sampling the green world. Columbia University Press, New York. HOAGLAND ET AL, RA Taytor, RJ.and C.S. Taytor. 1991. An annotated list of the ferns, fern allies, gymnosperms and flowering plants of Oklahoma. Biology Dept. Herbarium, Southeastern Oklahoma State University, Durant. TrewarrTHa, G.T. 1968. An introduction to climate. McGraw-Hill, New York. USDA-NRCGS. 2003. The PLANTS database. (http://p! usda.gov/plants). National Plant Data Center, Baton Rouge, LA. WaterFAaLt, U.T. 1973. Keys to the flora of Oklahoma. Published by the author, Stillwater, OK. BRIT.ORG/SIDA 21(1) BOOK NOTICES Blackwell Publishing JON Etrrince. 2004. Research Methodology in Applied Economics. Second Edi- tion. (ISBN 0-8138-2994-1, hbk.). Blackwell Publishing Professional, 2121 State Avenue, Ames, IA 50014, U.S.A. (Orders: 1-800-862-6657, 1-515-292-3348 lax, www.black wellprofessional.com). $74.99, 224 pp,, illustrated, 6" x 9" = Publisher Comments: “Emphasizing research methodology as it applies to economics, Ethridge pro- jor a dural guidelines on designing, coordinating, and conducting research projects. This textbook inte- grates philosophies, concepts, and procedures in research methodology, adding practical tips such es an overview of the conceptual and philosophical basis of research methodology and proce- as how to write a research proposal, how to apply for funding, and how to write re ports that effec- tively present research. This edition updates and increases the use of relevant exam ples for today’s students, faculty, and researchers.” Contents: Part I: 1) Introduction, 2) Research and Methodology, 3) Methodological Concepts and Perspectives, 4) Philosophical Foundations. Part I: 3) Planning the Research, 6) The Research Problem and Objectives, 7) The Literature Review, 8) The Conceptual Framework, 9) Methods and Procedures. Part HI: 10) Reporting the Research. Part IV: Appendices: A) Example of a Research Pro- posal for a SRL Meiers B) Example of a Master's Thesis Proposal: C) Example of a Ph.D f Dissertation Pro] id lor Critiquing Papers; E) Seeking Research Funding. Reference Index BirnGItTTA MALMForS, PHIL GARNSWorTHy, and MICHAEL GROSSMAN, 2004. Writing and Presenting Scientific Papers. (ISBN 0-897676-12-3, pbk.). Nottingham University Press, Manor Farm, Main Street, Thrumpton Nottingham NGI OAX, UK. (Orders: Blackwell Publishing Professional, 2121 State Avenue, Ames, IA 50014, US.A., 1-800-862-6657, 1-515-292-3348 fax, wwwblackwell professional.com). $74.99, 224 pp. graphics, figures, tables, 6 3/4" x 91/2" Contents: “Preface. 1) Communicating Science. 2) Sections of a Scientific Paper. 3) Tables and Figures. 4) Other Types of Scientific Writing. 5) Getting Started in Writing, 6) Improving your Writing. 7) Writing Statistics. 8) Literature Searching and Referencing. 9) Getting a Paper into Print. 10) Oral Presentation and Visual Displays. 11) Poster Presentation. 12) Training Students in Writing and Pre- sentation. 13) Reviewing Papers and Presentations. — SIDA 21(1): 446. 2004 SAW TOOTH OAK (QUERCUS ACUTISSIMA, FAGACEAE) IN NORTH AMERICA Alan T.Whittemore US National Arboretum | New York Ave N Washington, DC (District of Columbia) 20002-1958, U.S.A. ABSTRACT Sawtooth oak (Quercus acutissima Carruth.), native to eastern Asia, is vaely planted in the eastern United States as a source of food for wildlife cee ae and asa landscape tree in developed t fe Itivation has now been conlirmed for areas. Spontaneous reproduction o Alabama, Louisiana, Maryland, Missouri, Mississippi, North Carolina, Pennsylvania, and the District of Columbia. A complete necee eae ss an illustration are provided. Feral a sawtooth ] vl ] | oak are mostly ¢ onfined to o opel rbed areas, and it oe Very SLOV apparently due to limited dispersal of the acorns. a unting ol large ste — of Quet icutissimad in natural areas or revegetating areas for wildlife food is likely to result in the establishment of this exotic species and nabitats, but the use of sawtoot iG oak as a landscape tree in developed areas — its spread into adjacent usually poses much less danger of escape RESUMEN Quercus acutissima CAEFURs nativo del este de Asia, esta eae acu cultivado en el este aa - Esti idos Uni lat PpavOs )) como al lo an paisaje en areas desarrolladas. Su repr pdlere cion espontanea is a ae cultivo se ha coaleiae en Alabama, Louisiana, Maryland, Missouri, Mississippi, North Carolina, Pennsylvania, y el Distrito de Columbia. Se aporta una descripcion completa y una ilustracion. Las poblaciones silvestres de Quercus acutissima estan mayormente conlinadas a areas abiertas alteradas, y se expande muy bellotas. El establecimiento de lenieunniente debido aparentemente a la spol as inoantecls de : su | ES yeeEaCion aalimento grandes paecione - aus CUS de | lel establ | uexpansion en los habitats ea eeonie. pero el uso de Quercus dcutissima como aie de pi aisaje en areas desarrolladas tiene mucho menos peligro de escape. Sawtooth oak, Quercus acutissima Carruth., isa deciduous tree, native to open woodlands in eastern Asia, from northeastern India east to northern Vietnam and north to Japan and Korea (Huang et al. 1999). It was first introduced to the United States in 1862 (Rehder 1940), but it has only become common in culti- vation in the past 50 years. Sawtooth oak has been widely planted asa source of food for wildlife (especially turkeys), because of its fast growth and early, heavy fruiting (Sullivan & Young 1961; Mercer 1969; Hopkins & Huntley 1979; Goelz & Carlson 1997; Stribling 1994). It has also has gained favor in recent years asa landscape tree because of its attractive form, rapid growth, and tolerance of dif- ficult conditions (Spicer 1971; Francis & Johnson 1985; Gilbert &@ Henry 1988; Hensley et al. 1991; Tuttle 1995; Dirr 1998). SIDA 21(1): 447 — 454. 2004 448 BRIT.ORG/SIDA 21(1) Some concern has been expressed about the use of Q. acutissima as wildlife food in the United States. Mercer (1969) said, “The sawtooth oak has aroused some apprehension... Foresters wonder if it might become a “weed” tree. So far none of the plantings studied has increased in number of trees.” Coblentz (1981) suggested that its high germination rate and high resistance to insect damage might indicate that Q. acutissima has the potential to spread widely in the south- ern United States. He noted that the potential for hybridization between sawtooth oak and native North American oaks had not been investigated. He also cited papers indicating that the acorns of sawtooth oak are less nutritious than acorns of native species, and less utilized by North American wildlife, and concluded that long-term management goals will be best met by improved management of diverse native hardwood communities rather than by planting sawtooth oak. Perhaps because of these concerns, many wildlife programs have returned to planting native oaks In recent years, Quercus acutissima has been reported to reproduce outside of cultivation in six states: Alabama (Younghance and Freeman 1996), Louisi- ana (Thomas and Allen 1998), Maryland (Terrell et al. 2000), Missouri (Yatskievych and Summers 1993), Mississippi (Kartesz 1999), and Pennsylvania (Rhoads and Klein 1993). The species is not mentioned in the treatment of Quercus L. for Flora of North America (Nixon et al. 1997). There are almost no published data on the ecology of sawtooth oak in North America, and the very brief description in Rhoads and Klein (1993) is the only description of the spe- cies ina North American identification manual. Field and herbarium work in the eastern United States, and inquiry among active fieldworkers in the area, indicates bo Quercus dcutissima is escaping at sites across the eastern United States. S tion of sawtooth oak outside of cultivation has now been confirmed for se seven states and the District of Columbia. Since the species is becoming widespread and is being collected more frequently, it seems desirable to supply a full description of Q. acutissima, and asummary of its current range and habitat preferences in North America. KEY TO SEPARATE QUERCUS ACUTISSIMA FROM NATIVE OAK GROUPS OF THE EASTERN UNITED STATES 1. Acorns maturing in the first fall after flowering (so all acorns in summer are + the same size, and immature rns are not present on the tree in winter). Tips of veins at leaf margin never projecting as bristles. Axils of major veins on leaf underside without conspicuous tufts of hairs. Bark light to medium gray, splitting into loose or more or less persistent ridges, plates, blocks or strips White oaks (Quercus section Quercus) . Acorns maturing in the second fall after flowering (thus with large and small acorn on single n summer, and immature acorns present in winter). Tips en at leaf margin sleet always projecting as bristles 0.5—-7 mm long. Axils of major veins on leaf underside usually with tufts of stalked 4—-15-rayed hairs 0.3-0.5 mm high. Bark medium to dark gray, splitting into persistent ridges or blocks. <3 WHITTEMORE, QUERCUS ACUTISSIMA IN NORTH AMERICA 449 2. Scales of acorn cup lanceolate or strap-shaped, strongly recurved, 8-10 mm lon Leaf unlobed, its margin with 10-23 bristles on each side ouercus acutissima (Quercus section Cerris Loudon 2. Scales of acorn cup triangular, appressed, 1.5-5 mm long. Leaf not as above either deeply lobed or with 0-3 bristles on each side Black oaks (Quercus sect. Lobatae Loudon, sometimes called Quercus subgenus Erythrobalanus (Spach) Oerst.) =< Quercus acutissima Carruth., J. Linn. Soc., Bot. 6:33. 1862. (Fig. 1). SAWTOOTH OAK. Trees to 30 m tall. Bark medium to dark gray, divided into narrow persistent ridges. Twigs dark brown, puberulent with 1-5-rayed appressed (occasionally spreading) hairs, or glabrescent, 2-3 mm thick. Buds brown, 5-8 mm long, pu- bescent (at least the upper half), scales long-ciliate. Petioles 10-39 mm long. Leaf blade lance-oblong to lanceolate or oblanceolate, 11-21 cm long, 3-6 cm wide, base rounded or truncate; secondary veins each (except the basalmost) reaching the margin at the tip of a tooth and ending ina bristle, teeth LO-23 on each side of the blade, well-developed teeth obtuse to acuminate, each tooth ending ina single bristle 2-5 mm long. Upper surface of blade shiny, with scat- tered inconspicuous simple (rarely 2-4-rayed) hairs; lower surface green, the blade with inconspicuous unbranched appressed hairs, the veins with spread- ing simple hairs, vein axils with small tufts of ca 4-rayed stalked fasciculate hairs. Calyx of female flower fused to the ovary. Anthers retuse. Styles linear, their tips not broadened. Nuts ripe the second autumn after flowering. Peduncle 0-2 mm long. Acorn cup hemispherical, 14-15 mm long, 18-25 mm wide, cov- ering 0.3-0.5 of the nut, its inner surface smooth, hairy. Cup scales narrowly lanceolate or strap-shaped froma short triangular base, 8-10 mm long, weakly costate, free from cup for their whole length and strongly recurved, the scales at the margin of the cup longer but otherwise not differentiated. Nut ovoid to ovoid-cylindrical, 15-20 mm long, 13-17 mm wide. Inner surface of the shell densely pubescent, abortive ovules near the base, seed coat adhering to the fruit wa Flowering in April. Native to Asia, from Korea and Japan south to Vietnam and west to northeastern India. D ined: ALABAMA. Bibb Co.: 8 miS of Centerville along levee dirt road, off of county ‘oad 219, 32° 52' 30" N, 87° 00' 00" W, elev. 465 ft, S.T. Smith s.n., 10 Jul 1999 (AVA). DISTRICT OF COLUMBIA: Spontaneous small tree 5 m tall, trunk 6.5 cm thick, open grassy (partly cleared) margin of Quercus spp. woodland, north side of Beechwood Road near its intersection with Ellipse Road, U.S. National Arboretum. A.T. Whittemore 00-014, 29 Aug 2000 (CAS, NA, MOR, MU, JNC, US). LOUISIANA. Winn Parish: along paved road leading to Blewer’s Pond at Bienville Parish Line, north of LA 126 and NE of Roadheimer, sect. 6, T13N R5W, K.H. Kessler 1864, 20 Sep 1981 (AUA) MARYLAND. Prince George’s Co.: scattered adult trees with frequent saplings, open second-growth Quercus woodland, edge of Beltsville Agricultural Research Center along 1-495 near the Cherry Hill Rd. overpass, north side of College Park. A.T. Whittemore 01-058, 28 Nov 2001. (NA). MISSISSIPPI. Lafayette Co.: open area along Forest Service road, Tallahatchie Experimental Forest rd. 3, N of County Road 244, Holly Springs National Forest, UTM: 2 76 238E, 38 21 127N (GPS); L.M. McCook & M. Hodson 2180, 26 Jun 2000 (MISS, NA). MISSOURI. Franklin Co.: Quercus-Carya woodland below BRIT.ORG/SIDA 21(1) 450 Fig. 1. Quercus acutissima foliage and acorn. Scale bar = 3 cm. WHITTEMORE, QUERCUS ACUTISSIMA IN NORTH AMERICA 451 the nursery, next to Old Gray’s Summit Road, Shaw Nature Reserve, S of Gray’s Summit, AT. Whittemore 01-029, 15 Oct 2001 (NA). NORTH CAROLINA. Orange Co.: disturbed area: along drive to Craige Dormitory, UNC Campus, Chapel Hill, Emily W. oa 863, 2 Apr 1974 (MO). PENNSYLVA- NIA. Lehigh Co.: escape in fallow field 2 mi NW of Newside, R.L. SchaefferJr. 49540, 26 Jul 1955 (PH). These specimens represent areas where the species is definitely reproducing outside of cultivation. Specimens of planted trees have been seen from most parts of the eastern United States. Quercus acutissima is easily distinguished from all native North Ameri- can oaks. No native North American oak has unlobed leaves with numerous marginal bristles, and no native North American oak has the scales of the acorn cup long and reflexed. Sterile specimens of Castanea spp. are sometimes con- fused with Q. acutissima. Species of Castanea almost always have the leaves and buds in two regular ranks and not crowded at the stem apex. The sole excep- tion to this is Castanea dentata (Marshall) Borkhausen, in which the leaves and buds of lateral stems are two-ranked and not crowded apically, as in other Cas- tanea spp., while those of the leading stems are arranged in several irregular ranks and + crowded at the stem apex, similar to stems of Quercus. Fertile ma- terial of Castanea is easily distinguished by characters of the inflorescence (erect and rigid in Castanea, pendent and lax in Quercus) and fruit (the nut in Casta- nea is completely enclosed in a valvate husk that is covered with long spines, while in Quercus the nut is in an unlobed scaly cup). The only exotic oak that resembles Q. acutissima is Q. variabilis Blume, another Asian species that is rarely cultivated in North America. Quercus variabilis is very similar to Q. acutissima, differing only in having glabrous twigs, dense stellate pubescence on the underside of the leaf blade, and bark that is usually somewhat corky. A third species sometimes recognized from Asia, Q. chenii Nakai, is probably a synonym of Q. dcutissimad. Quercus acutissima reproduces spontaneously in grassland, open margins of deciduous woodlands, and other open disturbed areas. Sawtooth oak is espe- cially prolific in mowed meadows. Mowing keeps the seedlings small, but does not seem to harm them otherwise. Seedlings are found only in close proximity to adult trees. Careful searches at several sites in Missouri, Maryland and the District of Columbia showed that almost all seedlings and saplings grow within 20 m of an adult tree, and none was found more than 100 m from an adult, suggesting that spread of the tree is severely limited by short seed dispersal. As with other introduced species (i.e. Lonicera maackii (Rupr) Maxim., Luken & Thieret 1995), sawtooth oak was slow to appear in the North Ameri- can floristic literature. The first literature report of sawtooth oak as an escape in North America (Rhoads & Klein 1993) came 38 years after the first herbarium collection documenting it (Schaeffer 49540, 26 Jul 1955; see specimens exam- ined above), and it is still not treated in many recent floras. This may have de- 452 BRIT.ORG/SIDA 21(1) layed recognition of the plant in some areas, since collectors who encounter it may not be able to key it out or find descriptions of it. Quercus dcutissima isa member of Quercus sect. Cerris Loudon, a group of about forty species native to Eurasia and North Africa. In the past, this group has sometimes been included in sect. Quercus (the white oaks; Nixon 1993), but it is now clear that the white oaks and section Cerris are not closely related (Manoset al. 1999, 2001). Hybridization between Q. acutissima and native oaks, cited as a potential area of concern by Coblentz (1981), is not likely to be a prob- lem. Cottam et al. (1982) attempted numerous crosses between species of sect. Cerrisand various native North American oak species, with little success. They found that it is very difficult to obtain hybrids between species from different sections of the genus, even when all competing pollen is strictly excluded by bagging and emasculation of the bagged branches. Quercus acutissima was not one of the species they used, but they made extensive use of Q. variabilis, which is a very close relative based on numerous morphological (Huang et al. 1999) and molecular (Manos et al. 2001) characters. Cottam et al. (1982) attempted pollinations between Q. variabilis and sixteen species of white and black oaks, and they were unable to obtain a single hybrid from any of these pollinations. It is difficult to predict the long-term performance of sawtooth oak in the vegetation of eastern North America, since the decades that have passed since large-scale planting of the species in North America began are less than a full generation for the species. Even so, because sawtooth oak seedlings are able to establish themselves, mature, and set seed in reasonably natural habitats, Quercus acutissima should be considered naturalized, in the sense of Nesom (2000). Concerns about planting large stands of Quercus acutissima in natural areas seem to be well founded. In such sites, sawtooth oak can be expected to reproduce and spread slowly into adjacent open fields and woodland margins. The use of sawtooth oak as a landscape tree in developed areas usually poses much less danger of escape, primarily because of the short seed dispersal dis- tances of the species. However, trees planted close to disturbed grassland and open woodland may be expected to invade these sites. Furthermore, the heavy acorn drop in autumn and the frequent seedlings in garden beds are undesir- able characteristics in a landscape tree and cultivars with lower seed set would certainly be desirable. ACKNOWLEDGMENTS I would like to thank David E. Boufford, Steve Clemants, Allison Cusick, Lau- rence Dorr, Craig C. Freeman, David Giannasi, Curtis J. Hansen, Robert R. Haynes, William Hess, Marion Lobstein, James Macklin, Bill McClain, Lucile McCook, Cynthia Morton, Guy Nesom, Ann F Rhoads, John Schwegman, Ted Scott, Guy Sternberg, Jil Swearingen, John Thieret, John Townsend, James C. WHITTEMORE, QUERCUS ACUTISSIMA IN NORTH AMERICA 453 Trager, Lowell E. Urbatsch, Michael A. Vincent, Alan Weakley, David Whetstone, Ronald Wieland, and George Yatskievych for providing information, herbarium citations, help in the field, and other kinds of assistance, Andrea Ottesen for drawing the figure, and the curators of the herbaria cited for providing loans or facilities for work at their institutions. REFERENCES Costentz, B.E. 1981. Possible dangers of introducing sawtooth oak. Wildl. Soc. Bull. 9: 136-138. Cottam, W.P., J.M. Tucker, and F.S. Santamour, Jr. 1982. Oak so ail at the University of Utah. State Arboretum of Utah Publ. No. 1. pp.i-xiv Dirr, M.A. 1998. Manual of woody landscape plants.ed 5 Stipes Publishing, Champaign, IL. Francis, J.K. and R.L. JoHNson. 1985. Direct-seeded sawtooth oaks (Quercus acutissima Carruth.) show rapid growth on diverse sites. Tree Pl. Notes—U. S. Forest Service 36:3-5. Giceert, C.F.and D.S. Henry. 1988. Quercus acutissima'Gobbler: Amer.Nurseryman 168(3):122. Goetz, J.C.G.and D.W.Car.son. 1997.Growth and seed production of sawtooth oak (Quercus acutissima) 22 years after direct seeding. Research note SO-386. USDA-Forest Service Southern Research Station, Stonesville MS. Pp. 1-7. Henstey, D.L., S.C. Wiest, C.E. Lone, J.C. Pair and F.D. Gigsons. 1991. Evaluation of ten landscape trees for the midwest. J. Environ. Hort. 9:149-155, Hopkins, C.R. and J.C. Hunttey. 1979. Establishment of sawtooth oak as a mast source for wildlife. Wildl. Soc. Bull. 7:253-258. Huan, C., Y. ZHANG and B. BaRTHOLOMEW. 1999, Fagaceae. In: Z.-Y.Wu and PH. Raven, eds. Flora of China, vol. 4. Science Press, Beijing. Pp. 314-400. Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vas- cular flora of the United States, Canada, and Greenland. First Edition. In: Kartesz, J.T.,and C.A.Meacham. Synthesis of the North American flora. CD-ROM, Version 1.0.North Caro- lina Botanical Garden, Chapel Hill, NC. Luxen, J.O. and J.W. THieret. 1995. Amur honeysuckle (Lonicera maackii; Caprifoliaceae): its ascent, decline, and fall. Sida 16:479-503. Manos, PS., J.J. Doyte and K.C. Nixon. 1999. Phylogeny, biogeography, and processes of dif- ferentiation in Quercus subgenus Quercus (Fagaceae). Molec. Phylogenetics and Evol. 12:333-349 Manos, PS., Z.-K. ZHou and C.H. Cannon. 2001. Systematics of Fagaceae: phylogenetic tests of reproductive trait evolution. International journal of plant sciences. 162:1361-1379. Mercer, J., Jr. 1969. Sawtooth oak holds promise as wildlife plant for Southeast. Soil Con- servation 34:178. Nesom, G.L. 2000. Which non-native plants are included in floristic accounts? Sida 19: 189-193. Nixon, K. 1993. Infrageneric classification of Quercus (Fagaceae) and typification of sec- tional names. Ann. Sci. For. 50 (suppl. 1):25s—34s. 454 BRIT.ORG/SIDA 21(1) Nixon, K.C., RJ. Jensen, P'S. MANos and C.H. Mutter. 1997. Quercus. In: Flora of North America Editorial Committee (eds.), Flora of North America Vol. 3. Oxford University Press, New York. Pp. 445-506 Renper, A. 1940. Manual of cultivated trees and shrubs hardy in North America, exclusive of the subtropical and warmer temperate regions. Ed. 2. New York. RHoaps, A.F.and W.M. Kiein Jk. 1993. The vascular flora of P ylvania: Annotated checklist and atlas. Amer. Philos. Soc., Philadelphia. Spicer, G. 1971. Trees tough enough for the city. Amer. Nurseryman 133(10):7-8. STRIBLING, H.L. 1994, Planting sawtooth oak for wildlife. Circular ANR-851. Alabama Coop- erative Extension Service, Auburn. Pp. 1-2. SuLuvaN, E.G.and W.C. Younc. 1961.An exotic oak, Quercus acutissima, for wildlife food plant- ing. In: E. Conner Jr, ed. Proceedings of the fifteenth annual conference of the South- east Association of Game and Fish Commissions. Southeast Association of Game and Fish Commissions, Nashville, TN. Pp. 136-141. Turtte,G.A.1995. Tree of the month: Quercus acutissima, sawtooth oak. Arbor Age 15(1):28. YatskievycH, G. and B. Summers. 1993. Studies in the flora of Missouri, lV. Missouriensis 14: 2/-42 YOUNGHANCE, S.L. and J.D. Freeman. 1996, Annotated checklist of trees and shrubs of Ala- bama. Sida 17:367-384. SPOROBOLUS COAHUILENSIS (POACEAE): A NEW RECORD FOR THE U.S.A FROM TRANS-PECOS, TEXAS B.L. Turner Plant Resources Center University of Texas Austin, Texas 78 713 USA, Recent collections by the present author in southern Brewster County has re- vealed the presence of at least two new, presumably natural, populations of the poorly known S. coahuilensis Valdes, heretofore known only by a few collec- tions from gypseous soils of south-central Coahuila, Mexico (Fig. 1). Study of the grass collection at SRSC revealed three additional collections of the taxon from Brewster County made by yet others, these all identified as S. pulvinatus Swallen, and mapped as such by Turner et al. (2003). The Reeders of ARIZ sug- gested that an additional collection from along the Rio Grande in Hudspeth County might also belong to the taxon, which proved to be so. All of these col- lections follow: XAS: Brewster Co.: West end of Maverick Mt. near Study Butte, 8 Nov 1991, Powell 5805 (collected by Michael Clark); ca. 3 mi SW of Hen Egg Mt., 2.8 mi W of Hwy. 118 along unpaved South County Road, clayey soils immediately along roadside (ca 29° 27, 103° 33), 31 Oct 2003, B.L. Turner 23-325 (SRSC, TEX) ; 3.4 mi W of Hwy. 118 along South County Road, gypseous silty clays, 26 Dec 2003, [ rner 23-335 (SRSC, TEX); Study Butte, 30 Oct 1906, Warnock 23434 (SRSC); N of Study Butte, 25 t 1968, Warnock 23395 ? (SRSC). Hudspeth Co.: Quitman Mts., S tip of mountains at Indian Hot a Ss ‘seep area along road at base of hill near Rio Grande,” 3400 ft, 12 Oct 1980, Worthington 6804 (UTEP). In the treatment of the Grasses of the Southwestern United States by Gould (1988), the above collections will key to Sporobolus patens Swallen. Gould noted that the latter is“known only from the type collection made at Wilcox, Cochise County, Arizona (Silveus 3504, September 26, 1938).” He also noted that S. pat- ens might prove to be but a variant of S. pulvinatus. Sporobolus coahuilensis differs from both in having more numerous smaller florets on longer, more divaricate, pedicels, as nicely illustrated by Valdes (1978) in his original descrip- tion of the taxon. In the treatment of the grasses of the Trans-Pecos and adjacent areas by Powell (1994), S. coahuilensis will key to S. pulvinatus, the only annual species said to occur in the area concerned. Indeed, as already noted, Powell referred several of the above cited specimens to S. pulvinatus, the latter readily distin- guished from S. coahuilensis by its less open inflorescence and appressed flo- rets on shorter pedicles. In the Trans-Pecos, typical elements of S. pulvinatus SIDA 21(1): 455 — 457. 2004 456 BRIT.ORG/SIDA 21(1) : - C ae Ra, ; Sy a! ak SPOROBOLUS SS cad. coahuilensis Rea ae Fic. 1. Distribution of Sporobolus coahuilensis. are known only from El Paso County, these having been reported by Correll and Johnston (1970) as S. patens, but subsequently referred to S. pulvinatus by Johnston (1990). Turner et al. (2003) mapped the latter as also occurring in southern Brewster County; such plants are accepted herein as S. coahuilensis. In the account of Sporobolus for the Flora of North America (Peterson et al. 2003), S. coahuilensis will key to S. pyramidatus (Lam.) Hitche. Unfortunately, the names S. patens and S. pulvinatus are not accounted for in their treatment; presumably, they regard the two names as synonymous with their concept of S. pyramidatus. Regardless, Peterson (pers. comm...) recognizes S. coahuilensis as distinct, as do most recent workers interested in Mexican grasses (e.g., Ortiz, by annotation, TEX). In the area of Hen Egg Mt. where I first collected S. coadhuilensis the popu- lation concerned was composed of 40 or more very uniform, late-flowering individuals growing with or near the much more abundant earlier flowering S. pyramidatus. By no stretch of my imagination might I have included these un- der the fabric of S. pyramidatus. Subsequent collections of S. coadhuilensis from this same area (on 26 Dec 2003) showed that the latter occurred among or with numerous specimens of S. pyramidatus, there being no discernable intermedi- TURNER, : THE U.S.A. 457 ates between the taxa. During this second visit, approximately nine specimens of S. coahuilensis were detected growing with S. pyramidatus along a transect of some 40 yards along the edge of a recently plowed roadside. ACKNOWLEDGMENTS lam grateful to Charlotte Reeder for calling to my attention that the taxon con- cerned might be S. coahuilensis, and to her husband John, for yet other helpful comments. Thanks also to Richard Worthington at UTEP for the loan of S. coahuilensis from Hudspeth Co., the specimen itself composed of ca. five very uniform individuals mounted upon a single sheet. REFERENCES Correll, D.S.and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Texas. Gouto, FW. 1988. Grasses of the southwestern United States (Fifth edition). Univ. of Arizona Press, Tucson Jounston, M.C. 1990. A list up-dating the manual of vascular plants of Texas, Published by the author, Austin, Texas. Peterson, P.M.,S.L.HatcH, and A.S. Weaktey. 2003. Sporobolus, in Fl.N. Amer. 25:115-139. Powell, A.M. 1994. Grasses of the Trans-Pecos and adjacent areas. Univ.of Texas Press, Austin. Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003 Atlas of the plants of Texas 2:859. Vatoes, J. 1978. Sporobolus coahuilensis (Gramineae), a new species from Coahuila, Mexico Phytologia 41:81-84. BRIT.ORG/SIDA 21(1) BOOK NOTICE DALE LINDGREN and ELLEN WiLbr. 2003. Growing Penstemons: Species, Cultivars and Hybrids. (SBN 0-7414-1529-1, pbk). Infinity Publishing.com, 519 West Lancaster Avenue, Haverford, PA 19041-1413, U.S.A. (Orders: 877-BU Y BOOK, 1-610-520-2500, 1-610-519-0261 fax, www.buybookssontheweb.com, Info@buybooksontheweb.com). $17.95, 149 pp., illustrated, 8 1/4" « 10 3/4". Growing Penstemons begins with a history of Penstemons and the American Penstemon Society. The book is more than an alphabetical listing of species anc nee with descriptions. Also included for Penstemons, methods of propa- are chapters on Penstemon hybrids, selecting, | gation, and creating your own hybrids. Fight appe adic: prov ide information from Penstemons for beginners to hunting for Penstemons in the wild.—Barney Lipscomb, Botanical Research Institute of lexads, 509 Pecan Street, Fort Worth, Texas 761202-4060, U.S.A DoucLas Crasr. 2004. Both: A Portrait in Two Parts. ISBN 0-375-422066-8, pbk.). Pantheon Books, 1745 Broadway, New York, NY LOO19, U.S.A. (Orde: Ss www.pantheonbooks.com, www.randomhouse.com). $24.00, 320 pp., illus- trated, 53/4" x 7 1/2" Publisher Comments: “Both is the enchanting account of a remarkable filty-year relationship: Dwight Ripley, the child heir toan American railroad fortune, and Rupert Barneby, the product of a wealthy, baronial English upbringing, shared an obsession with botany from the moment they met at an ex- clusive boys’ boarding school in England. Together they embarked ona lifelong pursuit of rare plants, first in Europe and then in the United States, where they migrated in the late 1930s. Every spring they explored the American Southwest ina sputtering pees AON ering new pperies sae culiivas: } ing the spoils at their renowned home gardens. Barneby | he became a world authority on legumes. But the two men had other interests as well: fen were intimates in the expatriate circles that included W. H. Auden and Peggy Guggenheim, and early col- lectors of painters such as Jackson Pollock and Joan Miro. Ripley, a prescient artist himself, whose ost in a trunk for several decades before a rediscovered, startling work in colored pencil was | used his fortune to bankroll much of the avant-garde art scene of the early 1950 Sida Review coming in volume 21, no. 2 SIDA 21(1): 458. 2004 NEW REPORTS OF EURYBIA AND ASTERSSTR. (ASTERACEAE: ASTEREAE ) FROM CALIFORNIA, IDAHO, AND WYOMING Luc Brouillet Herbier Marie-Victorin Institut recherche en biologie végétale, Université de Montréal, 4101 Sherbrooke St. £, Montreal, Quebec, CANADA, H1X 2B2 luc.brouillet®@umontreal.ca Ab Ss, RAG | am reporting two new records of Eurybia merita (A. Nelson) GL .Nesom for California (Siskyou County), and new records of Aster alpinus L. subsp. vierhapperi Onno for Idaho (Lost River Range) and Wyoming (Beartooth Pass). RESUMEN Se hacen dos nuevas citas de Eurybia merita (A. Nelson) G.L .Nesom de California (Sisk you County), y dos nuevas citas de Aster alpinus L. subsp. vierhapperi Onno de Idaho (Lost River Range) y Wyo- ming (Beartooth Pass). INTRODUCTION While reviewing herbarium material in preparation of the treatments of Aster L.s.str.and Eurybia (Cassini) S.F Gray for the Flora of North America project, | came across specimens that represent new records for California, Idaho, and Wyoming. Eurybia merita (A. Nelson) G.L. Nesom CALIFORNIA. Siskyou Co.: Metcalf’s ranch, northeast base of Mt. Eddy, in moist soil in the forest, 3900 ft, 30 Jul 1936, Heller 12206 (MO); Mountains. 6000 ft, 5 Aug 5 1882, Pringle 14590 (MO). The two specimens I was able to examine are both typical E. merita, with purple- margined phyllaries and subserrate leaves, among other features. The Heller specimen was originally determined as Aster sp. and later annotated to Aster radulinus A. Gray by AG. Jones. The Pringle specimen was identified as Aster sibiricus L. var, (=A. montanus Rich.,a form approaching A. radulinus A. Gray). Identification of both specimensas E. radulina (A. Gray) G.L. Nesom is not un- expected given that E. merita had not been reported for California (e.g., Allen 1993) and that taxonomic confusion exists between the two species. Yet, exami- nation of a wide range of specimens in the course of preparation of the Eurybia treatment for FNA reveals that the two taxa are distinct (I have yet to find hy- brid or intermediate material) and clearly identifiable using the following com- bination of features: SIDA 21(1): 459 — 461. 2004 460 BRIT.ORG/SIDA 21(1) Stems ascending to erect, + densely villous distally; leaf margins coarsely serrate (teeth mucronate), cauline often clasping; phyllaries without purple margins; rays white (sometimes purplish) Eurybia radulina or = serrate, cauline subauriculate or slightly clasping; phyllaries graduated, purple on margins; rays purple (sometimes pale) Eurybia merita In the flora of the Pacific Northwest, Cronquist (1955) stated that smaller forms of E. radulina (as Aster) with purple rays and anthocyanic involucres were dif- ficult to distinguish from E. merita (as A. sibiricus var. meritus). He then men- tioned ecological and geographical separation to advocate segregation of the two taxa and also observed that larger forms of the latter are very distinct from E. radulina. Part of the problem may lie (I have been unable to verify this) with the fact that some small individuals of E. merita may have been misidentified as E. radulina, causing confusion, and from the fact that the ranges of the two species (as exemplified by the records cited above) may not be as disjunct as initially envisioned by Cronquist and others. Though | did not find anthocyanic specimens of E. radulina during my study (admittedly a small sample of all material potentially available), the distribution of purplish coloration on the phyllaries of this species (if they ever are purplish) may differ from the typi- cally purplish phyllary margins of E. merita. Cronquist (loc. cit.) did not ad- dress this issue. Further complicating the problem is the confusion between E. merita and E. sibirica, a species that barely reaches the conterminous United States. The definition of the range of characters of E. merita due to its inclusion within E. sibirica may have rendered its delimitation more difficult and thus less efficient. The recognition that E. merita is a species distinct from E. sibirica is crucial to our understanding of the former, Given our current knowledge of the distribution of Eurybia merita in Cali- fornia, it appears to be of conservation concern in the state. Further study of E. radulina specimens in California herbaria may yield further localities for E. merita and provide more precise data as to its habitat and distribution there. Aster alpinus L. subsp. vierhapperi Onno IDAHO. Custer Co.: Lost River Range, Challis National Forest, ridge between two forks of upper Grouse Creek, ca. 1.5 mi W of Grouse Creek Mt.,ca 19 mi N of Dickey; T12N R21E $23 NE1/4, 9900 ft, 14 Aug 1984, Moseley 533 (RM). WYOMING, Park Co.: Beartooth Pass, US-212, 10940 ft, 5 Sep 1979, Semple & Brouillet 4432 (MT, WAT). Aster alpinus subsp. vierhapperi has not been reported from the floras of Idaho (Davis 1952) and Wyoming (Dorn 1977: Nelson & Hartman 1994; R.L. Hartman, pers. comm. 2003) (see also, e.g,USDA-NRCS 2002, NatureServe 2003). The Moseley specimen was identified as Aster sibiricus var. meritus, while the Semple and Brouillet specimen was correctly identified but went unreported. The species is easily recognized by its rosette leaves, single heads with subequal, foliaceous phyllaries, and obconic, flattened, 2-nerved, puberulent, BROUILLET, ANT ASTER 461 apically glandular cypselae. Yet misidentifications abound with other species of asters, as well as, mostly, with species of Erigeron. The Idaho and Wyoming populations fill the gap between the southern Canadian populations of this species in Alberta and the Colorado locations (e.g, Hartman & Nelson 2001). All southern populations are at high elevations in the mountains, which may explain the rarity of records. Given that the Wyoming population is near the border with Montana and given the elevation of mountain ranges there, it is expected that collections of this taxon have been or will be made in this state; it should actively be sought there. Examination of Erigeron and other “aster” material from Idaho, Wyoming, Montana, and Colorado may yield further lo- cations. Given the current state of our knowledge, Aster alpinus should be con- sidered of conservation concern in Idaho and Wyoming. CONCLUSION The discovery of these new state records in well known states such as Califor- nia, Idaho, and Wyoming from examination of herbarium specimens under- scores the importance of herbaria and the value of revisionary and floristic work such as the Flora of North America project to our understanding of the continent's biodiversity. REFERENCES Aten, G.A. 1993. Aster. In: J.C. Hickman, ed. The Jepson manual: higher plants in California. University of California Press, Berkeley. Pp. 205-209. Cronauist, A. 1955. Part 5: Compositae. In: C.L. Hitchcock, A. Cronquist, M. Ownbey & J.W. Thompson, Vascular plants of the Pacific Northwest. University of Washington Press, Seattle. Davis, RJ. 1952. Flora of Idaho.Wm.C. Brown Co., Dubuque, lowa. Dorn, R.D. 1977. Manual of the vascular plants of Wyoming. Garland Publ. Inc., New York. 2 vols. Hartman, R.L.,and B.E.Netson, 2001.A checklist of the plants the vascular plants of Colorado. Rocky Mountain Herbarium, Laramie, Wyo. (obtained from http://www.rmh.uwyo.edu/ colorado/index.html, September 2003). NatureServe, 2003. NatureServe version 1.8 (1 July 2003) (http://www.natureserve.org/). Arlington, Va. (consulted September 2003). Newson, B.E. and R.L. Hartman. 1994. Checklist of the vascular plants of ey ae Mountain Herbarium, Laramie, Wyo. (obtained from http://wwyv cies/index.htm, September 2003). USDA-NRCS. 2002. The PLANTS database, Version 3.5 (http://plants.usda.gov) .National Plant Data Center, Baton Rouge, LA 70874-4490 USA (consulted September 2003 ait 462 BRIT.ORG/SIDA 21(1) THE NEW YORK BOTANICAL GARDEN Institute of Systematic Botany 200th Street and Southern Blvd. Bronx, New York 10458-5126 THE RUPERT BARNEBY AWARD The New York Botanical Garden is pleased to announce that Karen Redden, currently a graduate student in the Department of Biological Sciences, George Washington University, Washington, D.C., is the recipients of the Rupert Barneby Award for the year 2004. Ms. Redden will be studying the systematics of a diverse group of legumes centered around Dicymbe, paloue, paloveopsis, Heterostemon, and Elizabetha that are concentrated in the Guiana Shield area. The New York Botanical Garden now invites applications for the Rupert Barneby Award for the year 2005. The award of US $1,000 is to assist research- ers to visit The New York Botanical Garden to study the rich collection of Leguminosae. Anyone interested in applying for the award should submit their curriculum vitae, a detailed letter describing the project for which the award is sought, and the names of 2-3 referees. Travel to the NYBG should be planned for sometime in the year 2005. The application should be addressed to Dr. James L. Luteyn, Institute of Systematic Botany, The New York Botanical Garden, 200th Street and Kazimiroff Blvd., Bronx, NY 10458-5126, U.S.A. and received no later than December 1, 2004. Announcement of the recipient will be made by December 15. Anyone interested in making a contribution to THE RUPERT BARNEBY FUND IN LEGUME SYSTEMATICS, which supports this award, may send their check, payable to The New York Botanical Garden, to Dr. Luteyn. SIDA 21(1): 462. 2004 NOTES ON THE DISTRIBUTION OF PSEUDOGNAPHALIUM LUTEOALBUM (ASTERACEAE: GNAPHALIEAE) Guy L.Nesom Botanical Research Institute of Texas 509 Pe eet Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT The historical presence of Ps lium luteoalbum in Colorado and apenas is id mented by collections from the 19%) century, and itis first reported from Louisiana by a tion. The species is common in the southern half of California and locally common in ae n ‘loti but uncommon or rare in other states. Historical collections from ballast also record its presence in New York, Oregon, and Washington, but it has not persisted in any of those states. RESUMEN La pases msn? ee Feuaoes oo luteoalbum en ecleieee BE gee laren esta XI staes] } una coleccion reciente. La especie es frecuente en la mitad sur de Calif ornia y localmente en el sur de Florida pero poco frecuente 0 rara en otros estados. Las colecciones historicas procedentes de lastre también senalan su presencia en New York, Oregon, y Washington, pero no ha persistido en ninguno de estos estados. Pseudognaphalium luteoalbum (L.) Hilliard & Burtt has been known in the United States from California, Oregon, Washington, Nevada, Arizona, Utah, Florida, and New York (as sumrnarized by Kartesz 1999), and recent reports have expanded the range to include New Mexico, Texas,and Arkansas (Nesom 2001, 2002). A recent collection from Louisiana, as well as historical collections from Colorado and Pennsylvania, are reported here. LOUISIANA. Winn Parish: roadside along Hwy 84, 0.3 mi W of Cooley Methodist Church, ca. 13.5 mi WSW of Winnfield, area of Kisatchie National Forest, sandy soil in area of pine woods, ca. 100 plants scattered ae 50 feet of roadside, 10 Apr 2004, Nesom G2004-8 (BRIT, LSU, LSUS, MO, NY, NCU, UC, US). COLORADO. [Fr nt Co}: Canon City, along ditches, 22 Jul 1872, Redfield 188.5 (NY). PENN- SYLVANIA. aes hia a Philadelphia, Girard Point, in ballast, Aug 1877, Parker s.n. (NY). NEW YORK. N.Y, ballast filling, at northern terminus of 8th Ave., Jul 1879, Brown s.n. (NY). Mitchell and Tucker (1997) listed Pseudognaphalium luteoalbum for New York as a non-native, non-persistent species; the 1879 collection from an early bal- last site cited above perhaps is the basis for that citation. An early record of a ballast waif in Philadelphia, Pennsylvania, is added here. Historical collections from Oregon (Portland in 1902) and Washington (Bingen) also record intro- ductions through ballast, and the species apparently has not persisted in those 5 places (Cronquist 1955; Chambers & Sundberg 1998). SIDA 21(1): 463 — 464. 2004 464 BRIT.ORG/SIDA 21(1) The cited collection from Colorado indicates that the species was present in that state more than a hundred years ago; it seems remarkable that it has not been documented there since then (e.g, Weber & Wittman 1994) but it perhaps is an indication, as probably true also for Oregon, Washington, New York, and Pennsylvania, that winter cold limits the distribution. Pseudognaphalium luteoalbum iscommon in the southern half of Califor- nia and locally common in southern Florida but uncommon or rare in other states. Relatively scattered and recent collections, however, indicate that it is naturalized and actively expanding its range into inland regions, and it prob- ably should be expected to occur abundantly across the Gulf Coast region, once established. In the U.S.A, plants flower April through October and have been collected from roadsides, fields, gardens, ditches, riparian areas, and other dis- turbed sites. The species apparently is native to Eurasia and has been reported to occur in Africa, Australia,and New Zealand. It has been collected from scat- tered localities in Mexico. ACKNOWLEDGMENTS lam grateful to the staff at NY for their help during a recent visit, and | enjoyed “my mother on the April 10th afternoon of cudweed observa- =, the company o tions in Louisiana. REFERENCES Campers, K.L.and S. Sunpserc. 1998, Oregon vascular plant checklist: Asteraceae. Oregon Flora Project. Oregon State Univ., Corvallis. Cronouist, A. 1955. Vascular plants of the Pacific Northwest. Part 5: Compositae. Univ. of Washington Press, Seattle. Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. In: Kartesz, J.T, and CA. Meacham. 1999. Synthesis of the North American flora, Version 1.0. North Carolina Bo- tanical Garden, Chapel Hill, North Carolina. Mitchel, R.S. and G.C. Tucker. 1997. Revised checklist of New York state plants. New York State Bull. 490. New York State Museum, Albany. Nesom, G.L. 2001. New records in Pseudognaphalium (Gnaphalieae: Asteraceae) for the United States. Sida 19:1185-1190. Nesom, G.L. 2002. New records in Asteraceae from A 403-404. Weeer, W.A. and R.C. Wittman. 1994, Catalog of the Colorado flora: A biodiversity baseline. Univ. Press of Colorado, Niwot, Colorado. abama and Arkansas. Sida 20: DICHANTHELIUM SCOPARIUM AND MUHLENBERGIA GLABRIFLORIS: NEW TO THE FLORA OF OHIO Richard L. Gardner James S.McCormac The Nature Conservancy, Ohio Chapter Ohio Department of Natural Resources 6375 Riverside Drive, Suite 50 Division of Natural Areas and Preserves Dublin, Ohio 43017, U.S.A. 1889 Fountain Square, F-1 Email:rgardner@tnc.org Columbus, Ohio 43224, U.S.A. David Minney The Nature Conservancy, Ohio Chapter 6375 Riverside Drive, Suite 50 Dublin, Ohio 43017, U.S.A. ABSTRACT Two native grasses are reported new to Ohio's flora, Dichanthelium scoparium (Lam.) Gould and Muhlenbergia glabrifloris Scribn. Both species share a similar coastal plain and Mississippi gla § : embayment distribution pattern. They were found at localities less than 4.5 km apart and in an area . National Plant Data Center; ,Baton Rouge, LA 70874-4490 USA. PHALARIS ARUNDINACEA (POACEAE: AVENEAE) ASP BCIES NEW TOURE CAS AND A KEY TO PHALARIS IN TEXAS Stephan L. Hatch Dale A. Kruse S.M. Tracy Herbarium (TAES) S.M. Tracy Herbarium (TAES) Department of Rangeland Ecology Department of Rangeland Ecology and Management and Management Texas A&M University, 2126 TAMU Texas A & M University, 7126 TAMU College Station, Texas 77843-2126, U.S.A. College Station, Texas 77843-2126, U.S.A. s-hatch@tamu.edu dakruse@taexgw.tamu.edu Jennifer Pluhar Box 931 Canyon, Texas 79015, U.S.A. jjpluhar@arn.net ABSTRACT Th u f Phalaris arundinacea L., Reed canarygrass, in Texas is reported and a key to the genus Phalaris in Texas is provided. RESUMEN Se reporta la presencia de Phalaris arundinacea L., en Texas y se provee una clave para el genero Phalaris en Texas. During the week of May 20-23, 2002, while compiling data from specimens at the herbarium of West Texas A & M University (WTS), a misidentified speci- men of Phalaris arundinacea _. was found in the WTS collection. The sheet, originally identified as Phalaris caroliniana Walt., was collected by Larry C. Higgins in Hemphill County, Texas on June 29, 1978. Visits were made by the authors to the original collection site to determine whether the species still oc- curred in the state. In July 2002, the species could not be found, however on July 23, 2003, the species was located and persisting in the vicinity of Lake Kiowa on the Shannon Ranch in Hemphill County, Texas. At that time collec- tions were made, and later accessioned at the S. M. Tracy Herbarium (TAES) of Texas A & M University in College Station, Texas. All specimens were identi- fied using Chase (1951) and McGregor et al. (1986), and subsequently verified using TAES specimens. The North American distribution of P. arundinacea is documented in nu- merous sources. Chase (1951) indicated a range from Canada extending south to New Mexico, Oklahoma, Missouri, Kentucky, and North Carolina. Allred (1993) cited occurrences in San Juan, Rio Arriba, Mora and Lincoln counties, SIDA 21(1): 487 - 491. 2004 488 BRIT.ORG/SIDA 21(1) New Mexico. McGregor et al. (1986), reported the species as common in the northern Great Plains but rare in Oklahoma. Phalaris arundinacea, was not listed in Louisiana by Allen (1992) nor in Texas by Correll and Johnston (1979), Gould (1975) and Hatch et al. (1990). Jones et al. (1997) listed P arundinacea L. var. picta L. as being in the state, however this is a cultivated variety. No records for the species, in Texas, were found in collections at the University of Texas (TEX/LL) or at The Botanical Research Institute of Texas (BRIT). Inquiries were sent to curators of herbaria in Oklahoma (OKL, OKLA, WOH) and New Mexico (NMCR, SNM, UNM). None reported having collections of P arundinacea from Texas. Phalaris arundinacea was collected from the SE end of Lake Kiowa, NE of Lake Marvin in east-central Hemphill County, Texas. Lake Kiowa is a small to moderate size lake, ca. 7 hectares, which is fed by Boggy Creek and the Dry Fork of Boggy Creek, just N of the Canadian River. At the site, P arundinacea is lo- cally abundant as an understory species along the SE margin of the lake in as- sociation with Spartina pectinata Link, Juncus, Carex, Salix, Baccharis, Cephalanthus occidentalis L., Scirpus pallida (Britt.) Fern., [ypha, and other veg- etation associated with mesic to wetland habitats. Although the water level had dropped by the time of collection, there was evidence to indicate the site was inundated earlier in the season. The typical habitat for this species is meadows, stream banks, ditch banks, lake margins, or floating in water, Correll and Correll (1972), Arnow (1987) and Yatskievych (1999). The Hemphill County material appears to be at the southern limits of its distribution where it grows under the canopy of shrubs and trees. This makes finding the species more difficult com- pared to the populations in its northern distributions. This species may have been introduced to Hemphill County when the lakes, where it is found, were created. This species is distinguished from the other Phalaris species in Texas, by the presence of an obvious rhizome, being perennial, and having two reduced florets that are equal in length. Phalaris species in Texas are described in Gould (1975), with the exception of P arundinacea. Phalaris arundinacea L., Sp. PI. 55. 1753. (Fig. 1). Strongly rhizomatous perennials. Culms 50-160 cm tall, erect. Leaves basal and cauline. Sheath margins open. Ligules 5-9 mm long, membranous, obtuse (some- times lacerate), decurrent. Blades to 35 cm long, to 16 mm wide, flat. Panicles 6- 16 cm long, contracted, cylindrical (compact and sometimes lobed basally). Spikelets +-6.5 mm long. Glumes 4-0.5 mm long, subequal, laterally compressed Cxeeled), mid-vein wingless, 3-veined, apiculate. Sterile floret lemmas |-2.5 mm long, subulate, appressed pubescence, brownish. Fertile floret lemmas 3-4+ mm long, ovate, glabrous to pubescent. Fertile floret paleas 2.7-3.0 mm long, ap- pressed pubescence. Chromosome numbers 2n = 14, 28, 42, 56, and aneuploid June. ull, counts. Flowering period May- HATCH ET AL., PHALARIS ARUNDINACEA NEW FOR TEXAS AA h Ufo ] 4; Fal Fic. 1. Ph creeping rhizome; f, upper part a cults pec a fertile lemma, palea, and anthers; c, panicle, interrupted Belge e leaf sheath ligule; blade, and node; e, habit, showing Herbert L. Mason. University of C BRIT.ORG/SIDA 21(1) Specimens examined. TEXAS. Hemphill Co. the old Boy Scout Camp 12 mi E of Canadian, 29 Jun 1978, Higgins 12099 (WTU),; Lake Kiowa, 12 mi E of Canadian, 23 Jul 2003, Hatch and Hatch 8390 (TAES). A KEY TO THE SPECIES OF PHALARIS IN TEXAS 1. Plants perennial; rhizomes present. 2. Lower florets 2, about equal in length, 1-2.5 mm long P. arundinacea 2. Lower florets 1 or 2, unequal in length, one 0.5 mm long, longer one 1-2 mm long 1. Plants annual; rhizomes absent. 3. Reduced florets 1, scale-like 3. Reduced florets 2, scale-like. 4. Reduced florets broad, more than one-half the length of the perfect floret P. aquatica P. minor . canariensis 4. Reduced florets subulate, less than one-half the length of the perfect floret. 5. Reduced florets 0.4-0.8 mm long 5. Reduced florets 1-5 mm long. 6. Panicles 2-7 cm long; culms to 70 cm tall 6. Panicles 6-15 cm long; culms 60-150 cm tall P. brachystachya P. caroliniana P angust ACKNOWLEDGMENTS The authors thank the following: Angela Huff, Texas Agricultural Extension Service, Hemphill County, for establishing contact with the owner of the Shan- non Ranch. Lonnie and Sue Cox, managers of the Shannon Ranch, for provid- ing access to the property and transportation while on-site. David Sissom and Richard Kazmaier, the former and current curators of WTS for providing ac- cess to the collection, and the loan of specimens to TAES. The curators of BRIT and TEX/LL for access to their collections. Curators of the following herbaria for their assistance in searching their collections for species records in the re- gion; NMCR, OKL, OKLA, SNM, UNM, and WOH. REFERENCES ALLEN, C.M. 1975. Grasses of Louisiana. University of Southwestern Louisiana. Lafayette. ALLRED, KW. 1993. A field guide to the grasses of New Mexico. Agric. Exp. Sta. Las Cruces. Arnow, L.A. 1987. Gramineae. In: S.L. Welsh, N.D. Atwood, L.C. Higgins, and S. Goodrich. A Utah flora. Great Basin Naturalist Mem. Brigham Young University, Provo, UT. Pp. 684-788. Cuase, A. 1951. Hitchcock's manual of the grasses of the United States, 2" ed. USDA. Misc. Publ. No. 200. U.S.Government Printing Office, Washington, D.C CorreLt, D.S. and H.B. Corrett. 1975. Aquatic and wetland plants of southwestern United States. Environmental Protection Agency, U.S.Government Printing Office, Washing- tOnZOX, Corrett, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner HATCH ET AL., PHALARIS ARUNDINACEA NEW FOR TEXAS 491 Gould, FW. 1975.The grasses of Texas. Texas A & M University Press, College Station. Hatch, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Bull. MP-1655, College Station. Jones, 5.D.,J.K. Wiper, and PM. Montcomery. 1997.Vascular plants of Texas. University of Texas Press, Austin. McGrecor, R.L. and T.M. Barkiey (eds.). 1986. Flora of the Great Plains. University of Kansas Press, Lawrence. YATSKIEVYCH, G. 1999. Steyermark’s flora of Missouri, Volume I. The Missouri Department of Conservation, Jefferson City. SILENE CONOIDEA (CARYOPHYLLACEAE) NEW TO TEXAS Monique Dubrule Reed Biology Department Herbarium TAMU 3258 Texas A&M University College Station, Texas 77845-3258, U.S.A. Silene conoidea L., Conoid Catchfly or Cone Catchfly, is an annual or biennial native to Europe. It can be weedy in temperate regions and has been recorded in the US. from California, Colorado, Delaware, Idaho, Maryland, Missouri, Mon- tana, Nevada, Oregon, and Washington (USDA, NRCS 2004). The species was not listed for Texas by Correll and Johnston (1970) or by Jones et al. (1997).This is the first report of its occurrence in the state. In March of 2003, two plants were found growing in a vegetable garden at Peaceable Kingdom Farm near Washington, in Washington County. Vaccaria hispanica (P. Mill.) Rauschert, uncommon in the area, was collected nearby. The garden owners suspect the seeds of both may have been introduced ina load of turkey manure. Voucher specimen: U.S.A. TEXAS. Washington Co.: Peaceable Kingdom Farm, Washington, Mt. Falls School Road, vegetable garden, 27 Mar 2003, Loring, Winston-Mize, and Mize 03-62 (TAMU). Images of the Washington County plants may be seen at http://www.csdl.tamu.edu/FLORA/cgi/ gallery_query?q-=silene+conoidea. Silene conoidea somewhat resembles S. antirrhina L. (native to North America) and S. gallica L. (introduced), but it is larger in all respects. The leaves are lan- ceolate to oblanceolate, to 1 cm or more wide. The flowers are pink to purplish or white, with the petal blades 8-12 mm long and entire to toothed or notched. The most distinctive feature is the synsepalous calyx, which is 18-30 mm long and strongly inflated-conic at maturity, with about 20—30 strong ribs (Gleason & Cronquist 1991; Hickman 1993). Silene conica L., another European species, is very similar but has petal blades only 3-6 mm long and a mature calyx only 12-17 mm long (Gleason & Cronquist 1991). It has been collected in the U.S. from Delaware, Maryland, Mas- sachusetts, Michigan, New Jersey, New York, Ohio, Oregon, and Washington (USDA, NRCS 2004). It may eventually be found in Texas. Though the owners of Peaceable Kingdom do not plan to let S. conoidea recur on their property, this is a species known to be weedy and it should be watched for. SIDA 21(1): 493 — 494. 2004 494 BRIT.ORG/SIDA 21(1) ACKNOWLEDGMENTS Thanks go to J.K. Morton (WAT) for his confirmation of the identification and to Amanda Neill (BRIT) for reviewing the text. REFERENCES Corrett, D.S. and M.C. JoHnston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner. Gieason, H.A.and A. Cronauist. 1991.Manual of the vascular plants of northeastern United States and adjacent Canada, 2nd ed. New York Bot. Garden, Bronx. Hickman, J.C. (ed.) 1993. The Jepson manual: higher plants of California. University of Cali- fornia Press, Berkeley. Jones, S.D., J.K. Wiprr, and PM. Montcomery. 1997. Vascular plants of Texas:a comprehensive checklist including synonymy, bibliography, and index. University of Texas Press, Aus- tin. USDA, NRCS. 2004. The PLANTS database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA. NOTEWORTHY COLLECTIONS OF CYPERUS DRUMMONDII (CYPERACEAE) FROM TEXAS David J.Rosen Tracy Herbarium Department ofRange land Ecology and Management A&M University College eee ion nid 77843-2126, U.S.A. ABSTRACT Recent collections of Cy ul lii Torr. & Hook. from Texas coastal prairies known as “Nash Prairie” and “Bailey's Prairie” are repo — ae with ecological notes and a synopsis of its Texas distribution RESUMEN Se citan colectas recientes de Cyperus drt idii Torr. & Hook. de las prad teras de Texas onocidas como “Nash Prairie” y “B Bailey s Prairie”. Se anaden notas ecolégicas notes y una sinopsis de su distribuci6én en Texas. Carter et al. (1999) recognized Cyperus drummondii Torr. & Hook. as distinct from C. virens Michx. and provided maps of the distribution of both taxa in the southeastern United States. Cyperus drummondii has been scarcely collected in Texas. Beside the type collection by Thomas Drummond, Denton (1978) re- ported only two other collections from Texas, Cory 50890 (US) and Hall s.n. (F). In addition to these, Carter et al. (1999) reported Jones 719 (US). In Texas, this taxon appears to be rare and restricted to the upper portion of the Gulf Prairies and Marshes as defined by Gould (1975). Recently, while conducting surveys of botanical resources on private property in Brazoria County, collections of Cyperus drummondii were made from populations encountered on large tracts of moderately disturbed to relatively undisturbed coastal prairie. These collec- tions fill gaps in the distribution of this poorly understood taxon. Since perti- nent works (Torrey 1836; Kikenthal 1935-1936; Denton 1978; Carter et al. 1999: Tucker et al. 2003) include scant information on the distribution and ecology of C.drummondii in Texas, the following notes and synopsis of its distribution are provided. Distinguishing Characteristics Specimens were easily distinguished from C. virens by key characters and dia- grams provided by Carter et al. (1999). Upon critical examination, the ratio of achene length to scale length is sufficient to separate the taxa. Cyperus drum- mondii is also taller, has fewer and shorter primary inflorescence bracts, and SIDA 21(1): 495 — 497. 2004 496 BRIT.ORG/SIDA 21(1) fewer sessile to sub-sessile primary peduncles. As Carter et al. (1999) suggested these characters are best observed in the field. Habitat Cyperus drummondii was rare to locally common in poorly drained, fine sandy loam and clayey soils of prairie depressions mapped as Fdna fine sandy loam (Crenwelge et al. 1981). These large tracts were topographically intact as was evident by numerous pimple mounds. A composite list of closely associated species for all sites includes: Axonopus fissifolius (Raddi) Kuhlmann, Boltonia diffusa ELL, Croton capitatus Michx. var. lindheimeri (Engelm. & Gray) Mull, Cyperus entrerianus Boeck., C. haspan L., Diodia virginiana L., Eryngium yuccifolium Michx., Helianthus angustifolius L., Hydrolea ovata Nutt.ex Choisy, Juncus brachycarpus Engelm., Leersia hexandra Sw, Ludwigia linearis Walt., Panicum hemitomon Schult. P. hians El. P. virgatum L., Paspalum floridanum Michx. var. floridanum, P. plicatulum Michx., Polygonum hydropiperoides Michx., Rhynchospora caduca EIL, R. corniculata (Lam.) Gray, R. indianolensis Small, Rudbeckia nitida Nutt. var. texana Perdue, Sesbania drummondii (Rydb.) Cory, Solidago tortifolia ELL, Tridens strictus Nutt.) Nash, Tripsacum dactyloides (L.) L.,and Vernonia missurica Raf. Specimens examined. TEXAS. Brazoria Co.: Nash Ranch: hay meadow W of CR 25, about 8.7 mi N of its intersection with Hwy. 35 in West Columbia, 25 Aug 2003, Rosen & Carr 2590 (SBSC, TAES), and 19 Sep 2003, Rosen 2631 (TEX, VSC); Nash Ranch; Head of the Creek Pasture W of Hwy. 35, about 2.7 mi Sof its iota with FM 1462 in Damon, 25 Aug 2003, Rosen & Carr 2605, (BRIT, VSC): Bailey's Prairie; W of FM 521, approximately 4 mi SW of its intersection with Hwy. 35, W of Angleton, 17 Dec 2003, Rosen & Lange 2684 (MICH, VSC). Goliad Co.: 8 mi S of Coleto Creek, S of Victoria on Hwy. 77, hydric roadside ditch, frequent, 06 Jan 1988 Jones 719 (TAES). Harris Co.: Houston, 1872, Hall sn. (F mixed with Cyperus virens). Orange Co.: Growing in shallow water, 6.5 mi W of Orange, 16 Nov 1945 Cory 50890 (LL). ACKNOWLEDGMENTS Fieldwork was facilitated by the Nature Conservancy of Texas. Thanks to Di- ane Schenke and Bill Carr for offering the opportunity for access to the Nash Ranch, and to the curatorial staff at BRIT, F TAES, and TEX/LL for their assis- tance. Thanks to Richard Carter, Steve Hatch and an anonymous reviewer for their helpful comments. REFERENCES CARTER, R., D.K. ALEXANDER, C.T. Bryson, and A. Lazari. 1999.The taxonomy of Cyperus virens and Cyperus drummondii in the Southeastern United States. Sida 18:1049-1063. Crenwelce, G.W.,J.D.Crout, E.L.Grirrin, M.L.GoLbeNn, and J.K. Baker. 1981.Soil Survey of Brazoria County, Texas. United States Dept. of Agriculture, Soil Conservation Service. Denton, M.F. 1978. A taxonomic treatment of the Luzulae group of Cyperus . Contr. Univ. Michigan Herb. 11:197-271. ROSEN, LL 497 Gouto, FW.1975.Texas plants—A checklist and ecological Summary. MP-585 Revised, Texas Agricultural Experiment Station, College Station. KUKENTHAL, G. 1935-1936. Cyperus. In: A. Engler and L. Diels, eds. Das Pflanzenreich IV. 20 (Heft 101):1-671. Torrey, G.C.1836. Monograph of North American Cyperaceae. Ann. Lyceum Nat. Hist. New York 3:1-448. Tucker, G.C., B.G. Marcks, and J.R. Carter. 2003. Cyperus. In: Flora of North America Editorial Committee, eds. Flora of North America North of Mexico, Vol. 23. Oxford Univ. Press, New York. Pp. 141-191. RUPPIA CIRRHOSA (RUPPIACEAE) IN NORTH CENTRAL TEXAS Tiffany L. Morgan Walter C. Holmes Brazos River Authority Baylor University Waco, Texas 76714-7555, U.S.A. Waco, Texas 76798-7388, U.S.A. tiffanym@brazos.org walter_holmes@baylor.edu Ruppia cirrhosa (Pentagna) Grande has been recently collected from Lake Gran- bury in Hood County and Possum Kingdom Lake in Palo Pinto County, both in North Central Texas. The species was found at multiple locations in Lake Gran- bury in association with Stuckenia pectinata (L.) Boerner. In Possum Kingdom Lake the plant was collected in only one location, a shallow area called the Pea- nut Patch. In both reservoirs, the species was submersed, had long stems rooted in the sediment in water less than two meters deep, and was abundant in the locations where it was found. Voucher specimens: TEXAS. Hood Co.: Lake Granbury, approximately 0.3 mi ENE of Mallard Pointe, 6 Aug 2003, Morgan 599 (BAYLU), Lake Granbury, approximately 0.05 mi SW of intersection of Port Ridglea Drive and East Port Ridglea Court, 6 Aug 2003, Morgan 562 (BAYLU), Lake Granbury, ap- proximately 0.09 mi SW of terminus of East Port Ridglea Court, Morgan 564 (BAYLU); Palo Pinto Co.: Possum Kingdom Lake at the Peanut Patch $ of Caudill Mountain, N 32.89253, W 98.5067], 27 Aug 2003, Morgan 601 (BAYLU). Lake Granbury has a normal water surface elevation of 693 feet above mean sea level. At normal water surface elevation, Possum Kingdom Lake is 1,000 feet above mean sea level. Lake Granbury lies on the dividing line of the Black- land Prairie Ecoregion and the Oak Woods and Prairie Ecoregion, while Pos- sum Kingdom Lake lies in the Rolling Plains Ecoregion. The terrain around both reservoirs is varied, with rolling topography near the headwaters, and becomes increasingly rugged towards the dam. Lake Granbury is slightly saline with an annual average chloride concentration of 642 mg/L, an annual average sulfate concentration of 228 mg/L, and an annual average total dissolved solids con- centration of 1269 mg/L. Possum Kingdom Lake is also saline with an annual average chloride concentration of 995 mg/L, an annual average sulfate concen- tration of 364 mg/L, and an annual average total dissolved solids concentra- tion of 1,801 mg/L. Ruppia is known to frequent water with high calcium and sulfur concentrations (Haynes 2000). In North America, Ruppia has been treated as part of the Najadaceae (Steyermark 1963), the Potamogetonaceae (Thorne 1992), the Ruppiaceae (Correll and Johnston 1970, Haynes 2000), the Zannichelliaceae (Small 1933), and the Zosteraceae (Fernald 1950). We are following Haynes (2000), the most recent account of the North American species available. SIDA 21(1): 499 — 500. 2004 500 BRIT.ORG/SIDA 21(1) The genus consists of about ten species of nearly worldwide distribution, with two species, Ruppia cirrhosa and R. maritima L., recognized in North America (Haynes 2000). Both are known to occur in Texas (Turner et al. 2003). Ruppia maritima is limited to the Gulf Coast region of the state with almost all records being from coastal counties, while R. cirrhosa is known from nine coun- ties that are widely scattered in the Panhandle, South Texas Plains. along the Pecos River, and the northeast (Van Zandt Co.). The two species may be distin- guished by peduncle characteristics (Haynes 2000). The peduncle length in R. cirrhosa is greater than 30 mm and the peduncle has five or more coils, while in R. maritima the peduncle is 2-25 mm long and has less than five coils. Haynes (2000) also mentions that R. maritima is primarily of coastal occurrence and R. cirrhosa is mostly of inland occurrence. This is the first report of the genus and species for the area treated in Diggs et al. (1999). ACKNOWLEDGMENTS We thank Doug Amburn, Randy Johnston, Kyle Lewis, Craig Goen, and Mike Cox for their assistance in searching for the specimens on Lake Granbury and Kay Barnes for her helpful reviews. REFERENCES Corrett, D.S. and M.C. Jonnston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner Diccs, G.M., Jr., B.L. Liescome, and R.J. O’KeNNon. 1999. Shinners and Mahler's illustrated flora of north central Texas. Sida, Bot. Misc. 16. Botanical Research Institute of Texas, Fort Worth. Fernato, M.L. 1950. Gray’s manual of botany 8" ed. American Book Co., New York Haynes, R.R. 2000. Ruppiaceae. In: Flora of North America editorial committee, Flora of North America. vol. 22. Oxford University Press, New York. SMALL, J.K.1933.Manual of the southeastern flora. University of North Carolina Press, Chapel Hill. STEVERMARK, J.A. 1963. Flora of Missouri. lowa State University Press, Ames. THorne, R.F. 1992. Classification and geography of the flowering plants. Bot.Rev. (Lancaster) 58:225-348. Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. vol. 2. Sida, Bot. Misc. 24. Botanical Research Institute of Texas, Fort Worth. OSMORHIZA BIPATRIATA (APIACEAE) IN TEXAS: TAXONOMIC STATUS AND CONSERVATION CONSIDERATIONS Jun Wen Porter P. Lowry II Department of Botany Missouri Botanical Garden The -, ed of Natural History -O. Box 299 Lake Shore Dr. St. Louis, Missouri 63166-0299, U.S.A. Chicago, Ai inois 60605-2496, U.S.A and and Département de ene et Evolution oe of stra ic & Evolutionary Muséum National d'Histoire Naturelle m, Instit te of Botan ny Case pe 39, 57 rue Cuvier “Chi inese Academ\ ces 16 rue Buffon Beijing 100093, CHINA; ee am org 75231 Paris, FRANCE; pete.h bot.org James C. Zech Department of Bio en 5ul Ross State Unive Alpine, Texas 79832, U.S.A. ee ross.edu ABSTRACT The paper evaluates thet taxonomic an i conservation status of Osmor ued bipatr iata as ee & R.H. Shan. T] sahighly restricted distribution on n Mt. Biveunite of the Trans- Pecos region in Texas, as ill as in aaa Nuevo Leon, Mexico. Its habitat on Mt. Livermore is highly restricted within the mesic Pinus eae community, where it is associated with several other taxa that are also rare in Texas, uch as Polemonium pauciflorum S. Watson subsp. hinckleyi Genel) Wherry and Aquilegia iuekane A. Gray. The population of O. bipatriata on Mt. Livermore is small (< 20 individuals), although apparently healthy, with a full range of age structure. Application of the IUCN Red List criteria shows that O. bipatriata is an endangered species [B2ab(iv)], We recommend careful moni- toring of O. bipatriata and its associated habitat, and restricted access to its only known locality within the United States. RESUMEN En este articulo se evaltia ENS estatus taxonomico y ade conservacion de Osmorniza bipatriata Gene ] stance & R.H. Shan. | £ independiente. Tiene una di SbuciGe rouy res aaeail Mt. Livermore de la region Trans- — en TERS asi como en eyCodluatla y NUK c eon, MeEIeo ou Babitay en el Mt. PUVSHeks esta my taxa fie también son rarosen Texas, tales como Palemnouit ium pauciflorum S. Watson oo hinc ite yi (StandL) Wherry y Aquilegia longissima A. Gray. La poblacion de O. bipatriata en el Mt. Livermore es | pequenia ( Sint vidios) ludabl una estructura de edad con el rango completo. La aplicacion de lose criterios IUCN para la Lista Roja muestran que O. bipatriata es una especie amenazada [B2ab(iv)]|. Recomendamos un control cuidadoso de O. bipatriata y su habitat asociado, y acceso restringido a su nica localidad conocida en los Estados Unidos. SIDA 21(1): 501 — 506. 2004 502 BRIT.ORG/SIDA 21(1) INTRODUCTION Osmorhiza Raf. (Apiaceae subfam. Apioideae) is a small genus of approximately 11 species, including nine that occur in North America (two of which are also disjunct in South America) and one each in Asia and the central Andes. Mem- bers of the genus are characterized by their deep brown to black, linear to ob- long, fusiform to clavate fruits that are slightly compressed laterally and have a caudate basal appendage and filiform ribs, which in most species bear numer- ous retrorse bristles (Lowry & Jones 1984). Several recent molecular phyloge- netic analyses have shown that Osmorhiza is monophyletic and that it is most closely related to the Old World genera Myrrhis Mill. and Geocaryum Coss. (Downie et al. 2000; Wen et al. 2002: Yoo et al. 2002). These studies have also indicated that all the New World members of Osmorhiza evolved from a com- mon ancestor that arrived from Asia, where the genus appears to have origi- nated (Wen et al. 2002). Within the New World clade, a group of largely west- ern North American species forms a well supported subclade that probably diversified relatively rapidly. Two of these species, O. berteroi DC. and O. depauperata Phil. have subsequently dispersed into the Great Lakes area, east- ern North America, and southern South America, and another, O. mexicana Griseb,, has an interrupted range from northern Mexico to northern Argentina, while the remaining species occupy relatively narrow geographic ranges in the western United States (Lowry & Jones 1984; Wen et al. 2002). Specimens attributed to one of these species, Osmorhiza bipatriata Con- stance & R.H. Shan, were first collected on September 28, 1935, by LC. Hinckley on Mt. Livermore in the Davis Mountains of the Trans-Pecos region in Texas (Hinckley 408, SRSC). Constance & Shan (1948) described this sweet cicely (herein referred to as the bipatriate sweet cicely) from the Davis Mountains and northern Mexico as a species new to science based on a later collection by Hinckley (26 Jul 1937, L.C. Hinckley s.n; holotype at NY and isotypes at ARIZ and GH). Constance & Shan chose the epithet to reflect the presence of O. bipatriata in two countries as well as the two nationalities of its authors. Lowry S Jones (1984) reported several morphological intermediates between O. bipatriata and O. mexicana collected from sites in northern Mexico, including on Cerro Potosi in Nuevo Len, where the two taxa occur sympatrically. These intermediates prompted them to reduce O. bipatriata to a subspecies of O. mexicana. Recently, Turner et al. (2003) recognized the taxon asa variety of O. mexicana because he chose, somewhat arbitrarily, not to use the rank of sub- species in his work (B.L. Turner, pers. comm.). — TAXONOMIC STATUS Phylogenetic analyses conducted in the last several years using sequence data from the nuclear ribosomal ITS regions and the chloroplast ndhF gene and trnL- WEN ET AL., OSMORHIZA BIPATRIATA IN TEXAS 503 F regions (Downie et al. 2000; Wen et al. 2002; Yoo et al. 2002) suggest that Osmorhiza mexicana and O. bipatriata do not form a monophyletic group and indicate that they show considerable divergence in their sequence profiles of these markers (e.g., 1.415% divergence between the ITS sequences). In the ITS phylogeny, the positions of the two taxa are unresolved, although both are closely related to other western North American members of Osmorhiza (Downie et al. 2000; Wen et al. 2002). The chloroplast data set suggests that O. bipatriata is most closely allied to O. depauperata (Yoo et al. 2002). The avail- able data thus do not support the inclusion of O. bipatriata within O. mexicana at an infraspecific level. The phylogenetic data suggest instead that it should be treated asa distinct species, as initially proposed by Constance & Shan (1948), using either the phylogenetic (Nixon &@ Wheeler 1990) or the lineage (de Queiroz 2000) species concept. While the presence of morphological intermediates in northern Mexico would appear to be incompatible with the recognition of O. bipatriata using the classical morphological species concept, as supported by Cronquist (1978), the phylogenetic data suggest that these intermediates may represent interspecific hybrids or perhaps may have resulted form introgres- sion between O. bipatriata and O. mexicana. Detailed analyses, however, will be required to assess the origin of these intermediates. Osmorhiza bipatriata differs from its congeners by its small (9-11 mm long) and glabrous fruits, which are linear-fusiform and taper into a short beak at the apex, and by its very short styles [0.5-0.75 mm long (Lowry & Jones 1984)]. Based primarily on fruit morphology, Constance & Shan (1948) placed O. bipatriata in subgen. Glycosma, but commented that it possessed characters intermediate with members of the typical subgenus, in particular with regard to its involucel, styles and stylopodium, which are similar to those of O. mexicana, which belongs to sect. Mexicanae of subgen. Osmorhiza (see Con- stance & Shan 1948; Lowry & Jones 1984). Our molecular phylogenetic studies (Wen etal. 2002; Yoo et al. 2002) show that O. bipatriata possesses a large num- ber of unique molecular nucleotide substitutions, suggesting its antiquity. Osmorhiza bipatriata is thus perhaps best described as a relict species, which has survived in mesic habitats in the northeastern Chihuahuan Desert, which have undergone a gradual drying since the late Tertiary (Graham 1993). Anillustration of Osmorhiza bipatriata is found in Lundell & collaborators (1961: plate 42). Lowry & Jones (1984) provided comprehensive information on its morphology and distribution and cited specimens from both Texas and Mexico deposited in various herbaria. Below are some additional specimens from Texas that we have examined, but which were not seen by Lowry & Jones (1984). — a U.S.A. Texas. Jeff Davis Co.: Madera Car yon, Mt. Livermore, 28 Sep 1935, in fr, Hinckley 408 (SRSC); in wet soil of spring in Madera Canyon, Mt. Livermore, 23 Jul 1945, in young fr, Hinckley 3489 (SRSC); at the upper spring of Madera Canyon, NW slope of Mt. Livermore, 7300 ft, 12 Sep 1947, in fr, Warnock AIA CD Ny & Hinckley 4147 (SRSC); rare at upper spring in Madera Canyon on Mt. Livermore, Davis Mountains 504 BRIT.ORG/SIDA 21(1) igneous soil, 7500 ft, 11 Sep 1947, in fr, Warnock & Hinckley 7479 (SRSC); infrequent in igneous soil on upper Madera Canyon of Mt. Livermore, 7500 ft, 23 Aug 1968, Warnock 23015 (SRSC); upper Mad- era Canyon, Tobe SDInE, NW es Mt. Livermore, aust above al anes pOLeSneS wall, in deep mesic can- yon under Pi Juni associated with Rham- nus betulifolia, ome poicarpes ereophilus Aguile egia longisima Polemonium pauciflorum subsp. hinckleyi, Salv arizonica, Ptelea trifoliata, Galium sp., and Thalictrumsp.,? N30°38. 364! W104°10.746), 7242 [t., 3 Aug 2003, in fl & fr, Wen et al. 7265 (F). = CONSERVATION CONSIDERATIONS Osmorhiza bipatriata has a highly restricted distribution, with only a few lo- calities recorded on Mt. Livermore, Texas, and in Coahuila and Nuevo Leon, Mexico. Until recently, all of the collections from Texas were made by L.C. Hinckley and Barton Warnock (perhaps from the same locality), the most re- cent of which dates from 1968 (Warnock 23015, SRSC). During a recent study trip, we were able to relocate O. bipatriata on the northwestern slope of Mt. Livermore. A single, small population was found on 23 August 2003, compris- ing about 15 individuals, including 5-6 young seedlings. It occupied a small microhabitat with moist, rich soil in a deep mesic canyon under Pinus strobiformis-Quercus gambelii-Juniperus deppeana forest that differs consid- erably from most of the surrounding vegetation, which is primarily dominated by ponderosa pines and various oak species. The plants of O. bipatriata on Mt. Livermore were highly heterogeneous phenologically, with some individuals observed in flower, some in young fruit, and others in mature fruit, in addition to a few that were just emerging from the ground after a recent rain shower, which suggests that O. bipatriata is particularly sensitive to water availability. Suitable habitat for Osmorhiza bipatriata is apparently rare in the Davis Mountains. Although the population we studied appears to be healthy, as indi- cated by the presence of a full age structure, from young seedlings to mature adult plants, it is small (less than 20 individuals) and only persists ina restricted microhabitat. Despite three days searching the mountain for comparable habi- tat, we were unable to locate any other populations. We thus recommend care- ful monitoring of the only known population in the United States, which may require controlled access to ensure its continued survival. Osmorhiza bipatriata also occurs sympatrically with two other rare taxa in the Davis Mountains: Po- lemonium pauciflorum S. Watson subsp. hinckleyi (Standl.) Wherry; and Aq- uilegia longissima A. Gray. We also observed two additional rare species (Quercus depressipes Trel. and Aralia bicrenata Wooton & Standl.) at a higher elevation on the same slope of Mt. Livermore, the summit of the Davis Mountains. Osmorhiza bipatriata is not currently listed in the Texas Threatened and Endangered Species list (Texas Parks & Wildlife Department 2003) nor in the Threatened and Endangered Species System [TESS (U.S. Fish and Wildlife Service, 2003, http://endangered.fws.gov)], Application of the IUCN Red List criteria (IUCN, 2001; see also http://www.iucn.org/themes/ssc/redlists/ WEN ET AL., OSMORHIZA BIPATRIATA IN TEXAS 505 RLcats 2001booklet.html) shows, however, that O. bipatriata is an endangered species [B2ab(iv)] because it has an area of occupancy estimated to be less than 500 km? and because it is both known from fewer than five localities (cf. Lowry & Jones 1984: fig. 17) and appears to have a declining number of subpopula- tions, as indicated by our observations on Mt. Livermore. Most species of Osmorhiza have a relatively wide distribution (cf. Lowry & Jones 1984). Along with O. glabrata Phil, a central Andean endemic, and O. brachypoda Torr, found only in California and Arizona, O. bipatriata has one of the most restricted ranges in the genus. In the case of O. bipatriata, this may at least in part reflect the fact that its fruits lack the caudate appendages and retrorse bristles characteristic of most members of the genus, which are pre- sumed to play a role in dispersal (Constance & Shan 1948; Lowry & Jones 1984), as suggested for the widespread O. berteroi, which Cruden (1966) concluded was bird dispersed. In conclusion, Osmorhiza bipatriata is a distinct species with a long evo- lutionary history. It appears to be among the rarest plant species in the United States, with a single known population of less than 20 individuals occurring in a rare habitat in the Davis Mountains of west Texas. Careful studies need to be conducted to investigate its reproductive biology, seed germination and dis- persal biology, ecological preferences, and conservation genetics to develop a suitable management plan for this relict species. Comparative analyses of its congeners with both widespread and restricted distributions should also pro- vide insights into the biology of O. bipatriata that may be useful for its conser- vation. Furthermore, it isnecessary to compare the Texas population with those in Coahuila and Nuevo Leon, Mexico in order to verify that they are indeed con- specific, as suggested by their morphology. The evolutionary consequence or impact of the possible hybridization and introgression between Osmorhiza bipatriata and O. mexicana also needs to be assessed. ACKNOWLEDGMENTS We thank the Nature Conservancy of West Texas for granting access to the Davis Mountains Preserve, authorizing our field study on Mt. Livermore in August, 2003, and providing lodging during the field work; we are grateful to John Karges for helpful advice, Patricia Manning for field assistance, Mike Powell for her- barium assistance, permission to examine the specimens at SRSC, and valuable comments on the manuscript, B.L. Turner for discussion and his open- mindedness of accepting the species status of Osmorhiza bipatriata, and George Schatz for assistance with the IUCN Red List analysis. Support for the study was provided by the Field Museum of Natural History, the Chinese Academy of Science (a grant of Excellent Overseas Chinese), and the National Science Foundation (DEB 0108536 to JW). 506 BRIT.ORG/SIDA 21(1) REFERENCES ConsTANce, L. and R.H. SHan. 1948. The genus Osmorhiza (Umbelliferae), a study in geo- graphic affinities. Univ. Calif, Publ. Bot. 23:111-156 Cronauist, A. 1978. Once again, what is a species? In: L.V.Knutson, ed. Biosystematics in agriculture. Allenheld Osmun, Montclair. Pp. 3-20. Cruden, R.W. 1966. Birds as agents of long-distance dispersal for disjunct plant groups of the temperate western Hemisphere. Evolution 20:51 7-532. De Queiroz, K.2000.The general lineage concept of species and the defining properties of the species categories. In: R.A. Wilson, ed. Species: New interdisciplinary essays. MIT Press, Cambridge, Massachusetts. Pp. 49-89. Downie, S.R.,D.S.Katz-Downie, and K. Spauik.2000.A phylogeny of Apiaceae tribe Scandiceae: evidence from nuclear ribosomal DNA internal transcribed spacer sequences. Amer. J. Bot. 87:76-95, GRAHAM, A. 1993. History of the vegetation: Cretaceous (Maastrichtian)-Tertiary. In: Flora of North America Editorial Committee, ed. Flora of North America north of Mexico. Ox- ford Univ. Press, New York. 1:57-70. IUCN. 2001.IUCN Red list categories and criteria: Version 3.1.IUCN Species Survival Com- mission, Gland, Switzerland and Cambridge, UK. Lowey Il, PP.and A.G. Jones. 1984. Systematics of Osmorhiza Raf. (Apiaceae: Apioideae). Ann. Missouri Bot. Gard. 71:1128-1171. LUNDELL, C.L.and Cottasorarors. 1961.Flora of Texas, vol. 3. Texas Research Foundation, Renner. Nixon, K.C.and Q. D. WeeLer. 1990.An amplification of the phylogenetic species concept. Cladistics 6:211-223. TEXAS Parks AND Witotire DerartMeNT. 2003. Texas threatened and endangered species. Dis- tributed by Texas Parks and Wildlife Department, Austin. Turner, B.L.,H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Vol. 1. Sida, Bot. Misc. 24. Botanical Research Institute of Texas, Fort Worth. U.S. FisH AND WitoLire Service. 2003. Endangered and threatened plants. http:// endangered fws.gov. Wen, J., P.P. Lowry Il., J. Watck, and K.-O. Yoo. 2002. Phylogenetic and biogeographic diversi- fications of Osmorhiza (Apiaceae). Ann. Missouri Bot. Gard. 89:414-428. Yoo, K.-O., PP. Lowry Il, and J. Wen. 2002. Discordance of chloroplast and nuclear ribosomal DNA data in Osmorhiza (Apiaceae). Amer. J. Bot. 89:966-971. FIRST RECORD OF CLITORIA MARIANA (LEGUMINOSAE) IN WESTERN TEXAS Brian P. Oswald! Mohammad Bataineh Arthur Temple College of Forestry Arthur Temple College of Forestry Stephen F Austin State University tephen F. Austin State University Nacogdoches, TX 7596S U.S.A. Nacogdoches, TX 75965 U.S.A. boswald@sfasu.edu mb_96064001@yahoo.com Amanda Rountree Arthur Temple College of Forestry Stephen F Austin State University Nacogdoches, TX 75965 U.S.A. rountree_amanda@hotmail.com The genus Clitoria includes 35 species that are mostly common in warmer parts of the world. Common names of pigeon-wings or butterfly pea are used for the entire genus (Correll & Johnston 1996). Clitoria mariana L. isa perennial herb found throughout the United States, including eastern Texas. This specific spe- cies is reported as frequent in eastern and central Texas and infrequent in south- ern Arizona from the Chiricahua to the Baboquivari Mountains, thus creating a wide gap in the distributional range of the species (Correll & Johnston 1996; Turner et al. 2003). In Arizona it is found under junipers and live-oaks in rich soils (Kearney & Peebles 1964). Confirmed identification of this species was made during a recent study ol Pinus ponderosa L. and adjacent communities within the Davis Mountains of west Texas. The species was found at the bottom of a slope at approximately 1900 m elevation where the soils were the richest in the general area under ponderosa pine, with associated species including Mexi- can Pinyon (P. cembroides Zucc.), Alligator juniper Juniperus deppeana Steud.), and Texas Madrone (Arbutus xalapensis Buckl.). This is the westernmost find- ing of Clitoria mariana in Texas. The isolated finding reported here supports similar conditions described for southern Arizona. Clitoria mariana may be found in other scattered locations across the Madrean Province. Voucher Specimen: U.S.A. TEXAS. Jeff Davis Co.: TNC Davis Mountains Preserve, Madera Canyon, W 104° 7' 30" N 30° 39 30", elevation 1900 m, ponderosa pine community, growing in sandy loam soil, 14 Aug 2003, Mohammad Bataineh and Amanda Rountree 1(SRSC). ACKNOWLEDGMENTS We gratefully thank Billie L. Turner and A. Michael Powell for their identifica- tion of the specimen and for their extensive assistance in plant identification in ‘Corresponding Author SIDA 21(1): 507 - 508. 2004 508 BRIT.ORG/SIDA 21(1) the field. Our special thanks are extended to the Sul Ross State University her- barium (SRSC) for specimen storage and we also thank Chris Davis and Allan Pringle for their fieldwork. REFERENCES Correll, D.S.and M.C. JOHNSTON. 1996, Manual of the vascular plants of Texas. The University of Texas at Dallas. Kearney, T.H.and R.H. Peesies. 1964. Arizona flora. University of California Press, Berkeley. TURNER, B.L., H. NicHoLs, G.C. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Volume 1, Dicots. Sida, Bot. Misc. 24. Botanical Research Institute of Texas, Fort Worth. BOOK REVIEW JOHN Bork. 2001. Natural comy ds i therapy: f ing agents from plants & other natural sources. (0-9648280-1-4, pbk.) Oregon Medical Press, LLC. 325 10th Avenue North, Princeton, Minnesota 55371, US.A. (Orders: 800-610-0768; wwwompress.can). 00, 521 pp., b/w charts, figures, graphs, and tables, 13 appendices, 8 !/2" x 1x<¢ Ht Seldom does one encounter a seminal work wherein the author tackles the enormously comple immune system, the intricacies of the se ent ge nen bass process iaNeNved in meen com- bined with a most articul of botanical and other sources, which possess eee anti- ae ee ook is divided into three parts: or I: Cancer at the Cellular Level, is comprised of Chap- ters 2 through 6, and is preceded in Chapter | which presents background information related to Parts I and II, notably, the seven strategies for cancer inhibition; Part ll: Cancer at the Level of the Organism, and Part II: Clinical Considerations he topics included in Part I consist of the following: Mutations, Gene Expression, and Prolif- eration (Chp. 2); Results of Therapy at the Cellular Level (Chp. 3); Growth Factors and Signal Trans- duction (Chp. 4); Transcription Factors and Redox Signaling (Chp. 5); and Cell-to-Cell Communica- ) tion (Chp. 6 Several interesting and salient points are stressed. The central driving force behind the trans- formation of a normal cell into a cancer cell is mutation; especially through the expression of oncogenes and decreased expression of tumor suppressor genes Both nor a ane cancer cells po erate in response to the same signals, but in cancer the At the cellular level, successful cancer therapies (1) can cause cancer meals to assume more ae morphology and function, (2) prevention of cancer cells to enter the cell cycle, (3) induction of cell death through apoptosis, and (4) induction of cell death through necrosis. The first three of these can be accomplished by a variety of natural compounds: To avoid apoptosis and promote prolifera- ealt tion, cancer cells override the contro nor mally g | cells. Often cancer cells produce their ow th fact lucti and protein moieties. Of the three major transcription factors, cancer weet rely on abnormally low P53 ee activity and abnormally high NF-Kappa Band AP-1 activity. Intercellular communication and cell- matrix interaction are vital processes linking a cell to its environment. Cancer cells frequently ex- hibit a heen ant forms of communication. iets E-cadherin expression and increasing gap junc- eatment tion communication can provide useful strategies i Part II consists of Chapters 7 through 12. Topics covered include: overview of Angiogenesis (Chp. 7), Natural Inhibitors of Angiogenesis (Chp. 8), Invasion (Chp. 9, Metastasis (Chp. 10), The Immune System (Chp. 11), and Natural Compounds that Affect the Immune System (Chp. 12). A number of valuable observations were noted. While angiogenesis is a normal part of wound healing, it is the major mainstay supporting tumor growth. The difference between wound healing and tumor growth is th4a fact that angiogenesis in wound ces is finite and self-limiting but completely unchecked in tumors. A large number of compounds are capable of inhibiting angiogenesis, especially antioxi- dants, anticopper complexes, and vitamins A and saan including en have shown real promise. Cancet cells peace three i atrix com- id I matrix C San geae and variant Sha ponents surface proteins that alee migration. Metastasis isa ie step process, name ntravasation, (2) migration through circulatory channels, (3) arrest at a new location, (4) ex- oye and (5) cell proliferation and angiogenesis. it is highly likely that natural compounds (1):cell det << SIDA 21(1): 509. 2004 510 BRIT.ORG/SIDA 21(1) could play a significant role in slowing down or abrogating metastasis. The response of the immune system to cancer involves two major processes: (1) innate immune response segs nein of the tu- mor antigens, and (2) adaptive immune responses. Ideal results have been achieved when an immunostimulant in combination with materials that PEcren immune system evasion. Beledse of cytokines (IL-2 and the interferons) plus the addition o est potential for therapy together with inhibitors of PGE 2 or other specific commuinosumpreceine compounds. art HI consists of Chapters 13, through 23. Topless covered include: Back- ground for Part III (Chp. 13), Trace Metals (Chp. 14), Vitamin C and A chp. 15), So ere hp. 16), oS moniaceae asi como Ps familias. Se discut lit cias. Un punto déb ébi idelacl adistica molecular es el intento de clasificar es upos usando ae iaoe aS ae DNA. Se pueden obtener mejores resultados Hay una necesidad creciente de nuevas revisiones taxonomicas en las Polemoniaceae y otras familias de igual comportamiento. INTRODUCTION In the nineteenth century, Gilia was treated asa catchall genus for the temperate herbaceous Polemoniaceae that did not fit into the well-defined genera Polem- SIDA 21(2): 531-546. 2004 532 BRIT.ORG/SIDA 21(2) onium, Phlox,and Collomia. Gilia in that era was consequently very | and was subdivided into numerous sections (Bentham & Hooker 1873- 1876; Gray 1886; Peter 1897). In the early twentieth century as the plants became better known morpho- logically and in the field, it became apparent that many of these sections were only remotely related to ther. Milliken (1904) started the process of reclassi- fication by segregating Navarretia and Linanthus from Gilia. Her Gilia remained heterogeneous with six subgenera. Subsequent authors continued the process of pruning by spinning off Eriastrum, Leptodactylon, Allophyllum, Ipomopsis, etc. as separate genera, while retaining a polymorphous core genus Gilia. Grant’s (1959) treatment recognized a core genus Gilia composed of five interrelated sections (sects. Giliastrum, Giliandra, Gilia, Arachnion, Salt ugilia) Section Giliastrum has been subdivided subsequently into two or three smaller sections (Grant 1999) (Table 1). In 1959 | thought that the disparate elements had all been removed from Gilia, and Alva Day thought so too, but this was not the case. In later studies using pollen-morphological characters, Day (1993a, b) found a small group of species in Gilia sect. Saltugilia (the G. leptalea group) that did not belong in Gilia. It was not clear in 1993 where these species did belong. Day placed them in a section Kelloggia of Gilia for holding purposes, and later she and I trans- ferred them to Allophyllum (Grant & Day 1999). All these groupings and regroupings were made by taxonomists working within the conceptual framework of traditional or evolutionary taxonomy, us- ing numerous phenetic characters, and changing the system gradually and pro- gressively. In the year 2000, Porter and Johnson published a radically different classification of the Gilia complex and of the family as a whole. Their system was arrived at by the approach of molecular cladistics; they used DNA sequence variation in selected organellar genes as evidence, and interpreted this evidence according to cladistic concepts. In the Porter and Johnson (2000) system, the genus Gilia of Grant (1959, 1999) is broken up into seven genera, as shown in Table |. With regard to one species group, the former Gilid leptalea group, both parties agree that it should come out of Gilia, but do not agree on where to put it (Table 1). The new mono- typic genus Lath rocasis was set up for Gilia tenerrima (Porter & Johnson 2000); [did not at first accept Lathrocasis but do so now in this paper (Table 1). The genus Gilia of Porter and Johnson (2000) consists of Gilia sects. Giliaand Arachnion plus part of Gilia sect. Saltugilia. And their genus Gilia is separated at the tribal level from the other sections of Gilia (from Gilia subg. Greeneophila). How do we explain the large differences between the two contemporane- ous and up-to-date classifications of Gilia s. |? Porter and Johnson (Johnson et al. 1996, Porter 1998; Porter & Johnson 2000) claim that Gilia s. lis polyphyl- etic; their subdivided system is intended to correct the situation. I have argued aaa GRANT, :GILIA AND LATHROCASIS 533 Taste 1. Comparison of two current classifications of Gilia and certain gilioid taxa. Grant system (1999, 2001, this paper) Porter and Johnson system (2000) Tribe in PJ system Gilia subgen. Gilia Sect. Gilia Genus Gilia Gilieae Sect. Arachnion Genus Gilia Gilieae Sect. Saltugilia Genus Saltugilia Gilieae Gilia subgen. Greeneophila Sect. Giliastrum Genera Giliastrum, Dayia, Bryantiella Loeselieae Sect. Giliandra Aliciella Loeselieae Sect. Gilmania Aliciella Loeselieae Sect. Campanulastrum Linanthus campanulatus group Phlocideae Lathrocasis (formerly in Gilia) Lathrocasis Gilieae Allophyllum leptaleum group Navarretia leptalea group Gilieae (formerly in Gilia) (Formerly in Gilia) elsewhere (Grant 2001, 2003a, b) that the claim of polyphyly is not supported by the evidence, except in the Gilia leptalea group. The polyphyly issue will be discussed again later in this paper. The difference between the alternative treatments of the Gilia complex can be adequately explained as a result of using the very different approaches of taxonomy and cladistics. The differences in working concepts and methods can be summarized briefly here (see Grant 2003a for review). First, the system- atic units of taxonomy are similarity groups or taxa, those of cladistics are in- ferred phyletic lineages or clades. Second, taxonomy uses a traditional defini- tion of monophyly: any group descended from a close common ancestor; whereas cladistics defines monophyly as a group consisting of all the descen- dants of the common ancestor. A given group can be monophyletic by the taxo- nomic definition but non-monophyletic by the cladistic definition. Third, taxonomy employs any and all characters that are useful in distin- guishing taxa. Phenetic cladistics sets some restrictions on the characters used. Molecular cladistics uses one or a few preselected DNA segments; the data are valuable but the database is very narrow. Fourth, taxonomy and molecular cla- distics sample different parts of the overall genomes. The phenetic characters used in taxonomy are expressions of the chromosomal genome. The organellar DNA used in molecular cladistics is cytoplasmic in origin in the case of chloro- plast and mitochondrial genes and is encoded ina special kind of chromosome site in the case or ribosomes. Finally, there is always a subjective element when a cladogram is transformed into a system of taxa. Isa given clade going to be treated as a genus or a section? These factors inevitably bring about some differences between taxonomic and molecular cladistic systems of the same plant group. They account for the differences in the tribal classification of the Polemoniaceae of Porter and 534 BRIT.ORG/SIDA 21(2) Johnson (2000) and Grant (2003b). And they will explain most of the differ- ences in the treatment of the Gilia complex. This brings us to the next question. In cases of incongruence between taxo- nomic and molecular cladistic systems of classification, which system comes closest to the goal of expressing natural or phylogenetic relationships? The short answer is: sometimes one system or approach, sometimes the other. | have found the following modes in comparisons of taxonomic with mo- lecular cladistic systems in the Polemoniaceae and other plant groups. (1) Each system is acceptable by its own standards. (2) A cladistic author applies the cladistic definition of monophyly to a taxonomic system which is natural by the taxonomic definition of monophyly, and the cladistic author then falsely accuses the taxonomic treatment of being non-monophyletic. (3) The molecu- lar cladograms reveal a relationship which taxonomists had not noticed and which leads toa desirable revision in the taxonomy. (4) The DNA evidence is in conflict witha pattern of variation in reliable phenetic characters. This is likely to occur when the DNA evidence is derived from cytoplasmic organelles, espe- cially chloroplasts, which are semi-independent of the chromosomal genome which determines most taxonomic characters. A chloroplast DNA cladogram may give a very distorted picture of the organismic relationships ina plant group. Many plant groups have been studied with respect to both DNA markers and morphological or other phenetic characters. Sometimes the two kinds of characters are in agreement, sometimes they are not, and the latter situation is common. Rieseberg et al. (1996) list 34 seed plant genera in which discordance is found between DNA markers and phenetic characters. The type of DNA that is most common in the list of unreliable markers is chloroplast DNA. Unreli- able chloroplast markers are found for example in: Helianthus (Rieseberg 1991: Rieseberg et al. 1991), Quercus (Whittemore & Schaal 1991), Eucalyptus (McKin- non et al. 1999), and Gossypium (Cronn et al. 2002). In Helianthus (Rieseberg 1991; Rieseberg et al. 1991), Quercus (Whittemore S& Schaal 1991), Gossypium (Cronn et al. 2002), and Phlox (Ferguson & Jansen 2002) it is possible to compare the reliability of chloroplast DNA with that of ribosomal DNA. In each case ribosomal DNA is more concordant with phe- netic character variation than chloroplast DNA is. The various sources of incongruence between taxonomic and molecular cladistic systems, listed above, all occur in the Gilia complex and other Polemoniaceae. Examples will be given in this paper Old taxonomic treatments are currently being revised by cladistic, mainly molecular cladistic, methods, but cladistic systems are not always right. There isa continuing need for up-to-date taxonomic treatments. Such treatments pro- vide a choice for those who use classifications. With this goal in mind, I have recently revised and updated the tribal classification of the Polemoniaceae (Grant 2003b), and am doing the same here for the genus Gilia. GRANT, :GILIA AND LATHROCASIS 535 MATERIALS AND METHODS [assembled as many characters as I could that distinguish major subgroups in Gilia s. |. These are mostly gross morphological features but also micromor- phological or biochemical. Among the microscopic characters that are useful in Gilia is corolla vena- tion. The lower part of the corolla in Gilia has five sets of veins, one set for each corolla lobe, and each set has three parallel veins. The veins of a set branch above or distally. In some sections of the genus, the veins remain separate distally, while inother sections they anastomose distally (Day & Moran 1986; Day, pers.comm.). The flavonoids in the genus fall into three groups, designated as types A, B, and C, and these types vary among the sections (Smith et al. 1977). Type A fla- vonoids are kaempferal, quercitin, and myricetin; type B is 6-methoxyflavonal; and type C is C-glycosylflavone (Smith et al. 1977). A number of studies of DNA sequence variation have been made in the Polemoniaceae (listed in Porter & Johnson 2000; and Grant 2003b. For the pur- pose of this study I used mainly the papers of Johnson et al. (1996), Porter (1997, 1998), and Johnson and Weese (2000) which have the best coverage of the Gilia complex. Johnson et al. present cladograms of cpDNA matk, Porter of rDNA ITS, and Johnson and Weese of rDNA ITS, cpDNA trnL, and matK. Recently Johnson et al. (2004) have published a survey of the fine struc- ture of the seed coat in Gilia and related genera (see their SEM photographs). The descriptions in the formal classification consist mainly of diagnostic characters. These serve to show the evidence supporting the classification. Good complete descriptions are given by Porter and Johnson (2000) ANALYTICAL KEY TO THE MAIN GROUPS OF GILIA AND LATHROCASIS 1. Glandular hairs with tiny black tips; pollen white; seeds one per locule in capsul enus Lathrocasis KI ies es where present with amber or col inal gland I| Fi cream; seeds generally several or many per locule in capsule Genus Gilia 2. Pollen blue; stamens inserted in sinuses of lene fell annuals colle ia alas 3. Pubescence of long fine intertwined whit y hairs; stipitat lar hairs often present also Gi fas sect. Arachnion 3, Cobwebby pubescence not present, pubescence consisting of multicellular trichomes and stipitate glandular hairs. 4. Plants scapose Gilia sect. Saltugilia 4. Plants not scapose, cauline leaves ranging from large on lower stems to mall on upper stems Gilia sect. Gilia 2. Pollen generally yellow or cream, but blue in one species; stamens inserted in corolla tube, throat, or sinuses; perennials and annuals Gilia subgen. Greeneophila 5. Plants usually eraicnlig from ERS with elm spreading, but sometimes single-st g p rotate; ) ilaginous when w 6. Small canes with wiry stems and small flowers Gilia sect. Campanulastrum 6, Perennials and some annuals; flowers showy or small Gilia sect.Giliastrum 536 BRIT.ORG/SIDA 21(2) 5. Plants scapose with a basal rosette and a central leader stem; corolla funnel- ple or sometimes trumpet-shaped; seeds not mucilaginous when wet or only slightly so, ; oe leaves pinnate with a broad strap-shaped rachis and short lobes Cilia Sect. Giliarndra 7, Lower leaves with a broad blade and lobed margin, the lobes sharp- tippec ____ Gilia sect. Gilmania 7 CLASSIFICATION Genus 1. Gilia Ruiz & Pavon, Prod. Fl. Peruv. 25, t. 4.1794. Type: Gilia laciniata Ruiz & Pavon, Herbaceous perennials, biennials, or annuals, sometimes with a woody or soft woody base. Basal leaves well developed and upper cauline leaves much reduced in size, or moderately reduced in sect. Gilia. Generally spring blooming. Flow- ers usually in loose or glomerate cymes, or sometimes solitary or in ea heads. Calyx lobes equal in length. Corolla frequently moderate-sized and showy, or small in many species. Pollen pores zonocolporate. Seeds small, an- gular, and sandy-colored, usually numerous in the capsule. Ancestral basic chro- mosome number x= 9 present in all sections; polyploidy common. Other char- acters vary between the subgenera and sections. Distribution and taxa.—Widespread in western United States and Canada and northern Mexico; also in tem perate South America. In arid or semiarid habi- tats, frequent in deserts. About 78 species WN eu Subgenus 1. Gilia Leaves pinnately dissected, often bipinnate or tripinnate, but once pinnate or lin- ear in reduced forms. Pubescence varies among the sections. Stipitate glandular hairs often present; they are medium-sized with a large terminal g gland that is yellow or amber. Corolla usually funnelform, sometimes long-tubed and sub- salverform. Corolla veins anastomosing in distal part of corolla (see Materials and Methods for explanation of this character.) Stamens inserted in corolla lobe sinuses Pollen blue. Seeds generally numerous in capsules, mucilaginous when wet. Fla- vonoids of type A and/or C present, but not type B. (See Materials and Methods lor explanation of these types.) Basic chromosome number x = 9 throughout. jee) Section 1. Gilia Plants with leafy stems, the leaves being the largest on lower stems and smaller but well developed on upper stems. Pubescence of multicellular trichomes and medium-sized stipitate glandular hairs. Inflorescence an open cyme, ora capi- tate head in some species. Corolla concolored or bi- or tricolored with purple spots on the throat and yellow tube. Corolla veins anastomosing. Flavonoids of type A found (see Materials and Methods for explanation). Distribution and taxa—Cismontane California to British Columbia and Baja California, and in Peru and Chile. Ten species: G. achilleaefolia, G. angelensis, G. capitata, G. clivorum,G. laciniata(S. Amer), G. lomensis(S. Amer),G. mil lefoliata G. nevinii, G. tricolor, G. valdiviensis(S. Amer). GRANT, :GILIA AND LATHROCASIS 537 Section 2. Arachnion A.D. & VE. Grant, Aliso 3:214, 1956. Type: Gilia latiflora A. Gray. Plants scapose with a basal leaf rosette and central leader stem. Pubescence of long fine intertwined white cobwebby hairs; medium-sized stipitate glandular hairs of ten present also. Inflorescence an open cyme. Corolla often bi- or tricol- ored with contrastingly colored lobes, throat, and tube. Corolla veins anasto- mosing. Flavonoids of type C present. Distribution and taxa.—Mountains and deserts of western North America, especially numerous in the Mojave desert; also in temperate South America. Twenty-five species: G. aliquanta, G. austrooccidentalis, G. brecciarum, G. cana, G. clokeyi, G. crassifolia (S. Amer.), G. diegensis, G. flavocincta, G. inconspicua, G. interior, G.jacens,G. latiflora, G. leptantha, G. malior, G. mexicana, G. minor, G. modocensis, G. ochroleuca, G. ophthalmoides, G. salticola, G. sinuata, G. tenuiflora, G. tetrabreccia, G. transmontana, G. tweedyi. The basic taxonomic treatment is that of Grant and Grant (1956). Section 3. Saltugilia V.E.@ A.D. Grant, Aliso 3:84, 1954. Type: Gilia splendens Douglas ex HL. Mason & A.D. Grant, Madrono 9:212. 1948. Genus Saltugilia L.A. Johnson in Porter & Johnson, Aliso 19:69. 2000. Type designated by Johnson: Saltugilia grinnellii (Brand) L.A. Johnson. See Grant & Wendt (2003) for discussion of type. Plants scapose with a basal leaf rosette and central leader stem. Pubescence of straight multicellular trichomes and stipitate glandular hairs, or with genicu- late multicellular trichomes in G. stellata. Inflorescence an open cyme. Corolla usually concolored. Corolla veins anastomosing. Flavonoids of types A and C. Distribution and taxa.—Central cismontane California to northern Baja California, and to western parts of desert. Woodland and desert habitats. Seven species: G. australis, G. caruifolia, G. latimerii, G. scopulorum, G. splendens, G. stellata, G. yorkii. Gilia latimerii (T.L. Weese & L.A. Johnson) VE. Grant, comb. nov. Saltugilia latimeri TL. Weese & L.A. Johnson, Madrono 48:198. 2001. Related to G. australis. Comment.—The species in sect. Saltugilia fall into two groups. The G. splendens group (G. splendens, G. caruifolia, G. australis, G. latimerii) is a natural interre- lated group of woodland and desert species with mostly large flowers. The sec- ond group (G. scopulorum, G. stellata, G. yorkii) consists of small-flowered, mostly desert species which are similar to the G. splendens group in gross mor- phological characters. This was the basis for grouping them together in the same section in earlier studies (Grant & Grant 1954; Grant 1999). However, the molecular evidence throws some doubt on this assumption. The G. splendens group and the second group form separate clades in the cla- dograms for chloroplast genes matK and trnL and ribosomal ITS Johnson et al. 1996; Johnson & Weese 2000). A new character, sculpturing of the seed coat, also differs between the two groups Johnson et al. 2004). Porter and Johnson (2000) treat the G. splendens group as a segregate ge- nus, Saltugilia, and leave the second group (G. scopulorum etc.) in their genus 538 BRIT.ORG/SIDA 21 = 2) Gilia. I of course believe that these groups should be treated at the rank of sec- tion rather than genus. Otherwise I agree that there is a problem concerning the closeness of the relationships between the two groups and within the sec- ond group itself. More morphological, breeding, and molecular studies are de- sirable to clarify the relationships. In the meantime we have the practical prob- lem of making a place for the second group in the classification system. In the present system, the species of the second group are retained in the sect. Saltugilia until we know better what to do with them. a Subgenus 2. Greeneophila Brand, Pflanzenreich 4(250):144. 1907. Type: Gilia rigidu Benth. Leaves with broad blades, or pinnately divided, or linear in reduced forms. Small or tiny glandular hairs, short-stalked, the glands translucent and colorless, or herbage sometimes glaucous in sect. Ca lastrum. Corolla varying in form: campanulate, rotate, funnelform, or trumpet- shaped. Corolla veins anastomos- ing or non-anastomosing (see section descriptions). Stamens often inserted in corolla tube or throat, but sometimes in sinuses (see section descriptions). Pol- len yellow or cream, but blue in one species (in sect. Giliastru m). Seeds gener- ally numerous in capsules; mucilaginous or non-mucilag (see section de- scriptions). Flavonoids of type B (6-methoxyflavonols) present in sects. Giliastrum and Gilmania; no data for the other sections. Basic number x = 9 present in all sections; x = 9 and 8 in sect. Giliandra. Section 4. Giliastrum Brand, Pllanzenreich 4(250):147. 1907. Type: Gilia rigidu Benth. Giliastrum Rydb., FL Rocky Mts., ed. 2, 699, 1066. 1922. Bryantiella J.M. Porter, Aliso 19:70. 2000. TyPE: Gilia palmeri S. Wats., Proc. Amer. Acad. Arts 24:61. 1889. Dayia J.M. Porter, Aliso 19:71. 2000. Typr: Gilia scabra TS. Brandegee, Zoe 5:166, 1903. a. a Q Perennial herbs with a soft woody base and some annuals, stems branching from base. Leaf consisting of a broad blade with serrate margin, or blade cleft and with lobes, or reduced toa narrow linear rachis with narrow lobes. Corolla generally campanulate or rotate, rarely funnelform (in G. scabra); large or small. Corolla violet, blue, pink, or white, sometimes with a yellow tube. Corolla veins separate and non-anastomosing, except in G. rigidula where they do anastomose (see Materials and Methods). Stamens inserted in corolla base or throat. Pollen usually yellow, sometimes white, blue in one species (G. scabra). Seed coat mu- cilaginous when wet. Basic number x = 9; n = 6 and 12 occur in G. insigne. Distribution and taxa—Colorado and Kansas to Texas and northern Mexico and Baja California, also in temperate South America. Often in semiarid or arid plains and deserts. Twelve species: G. castellanosii (S. Amer.), G. foetida (S. Amer), G. glutinosa (S. Amer.),G. incisa, G. insigne, G. gypsophylla, G. ludens, G. palmeri, G. purpusti, G. rigidula, G. scabra (includes Dayia grantii J.M. Porter pending further study), G. stewartii. See Turner (1994) for a treatment of the Texas and Mexican species. Porter and Johnson (2000) treat Giliastrum as a genus. GRANT, :GILIA AND LATHROCASIS 539 Gilia castellanosii (J.M. Porter) V.E. Grant, comb. nov. Giliastrum castellanosii J.M. Por- ter, Aliso 19:75. 2000. Comment.—Gilia scabra of Baja California was poorly understood for many years. Brandegee (1903), who described it, stated that it was related to Gilia flori- bunda in section Siphonella. These are old names for Linanthus nuttallii. When compiling a list of species names in the 1950s, I followed Brandegee and listed Gilia scabra asa synonym of Linanthus nuttallii (Grant 1959, p. 140). 1 did not see herbarium material until much later. Recently Porter has studied this spe- cies in the field and laboratory, and gives a full description of its morphology (Porter and Johnson 2000). He also presents molecular evidence as noted be- low. Porter proposes a new genus, Dayia, for D. scabra and the closely related D. grantii.G. (or D.) scabra seems to fit into sect. Giliastrum, though it does differ from other species of Giliastrum in having funnelform corollas and blue pol- len. Alva Day also views G. scabra asa member of sect. Giliastrum (pers. comm.). The molecular evidence consists of sequence variation for the chloroplast gene matK and ribosomal ITS Johnson et al. 1996, Porter 1997; Prather et al. 2000). Molecular evidence could help greatly to clarify the relationships of Gilia scabra, but in fact only raises more questions. In the DNA cladograms, Gilia scabra forms a clade consisting of itself and Loeselia glandulosa. This result is puzzling. Gilia sect. Giliastrum is only distantly related to Loeselia (Grant 2003b). Gilia scabra does not have the phenetic characters of Loeselia. In the cladograms Gilia scabra is adjacent to a Giliastrum clade, but Loeselia glandulosa seems very much out of place. The possibility of mislabelling plant material suggests itself. The assays of Gilia scabra and Loeselia glandulosa should be repeated. For the present it seems best to treat G. scabra as a member of sect. Giliastrum. In the future, with more study, it might be assigned toa new section, Dayia, related to sect. Giliastrum. Gilia palmeri of Baja California and G. glutinosa of Peru and Chile have been treated as a related amphitropical species pair in sect. Giliastrum (Grant 1959). Porter proposes to treat them as a new bitypic genus, Bryantiella (Porter & Johnson (2000). The phenetic characters to support this change are not im- pressive. Porter has some molecular evidence from cpDNA and rDNA to sup- port this proposal but this is unpublished (Porter & Johnson 2000, p. 71). think these two species belong in sect. Giliastrum, and Alva Day (pers. comm.) is of the same opinion. Section 5. Giliandra A. Gray, Proc. Amer. Acad. 8:276. 1870. Type: Gilia stenothyrsa A. Gray. Aliciella sect. Giliandra J.M. Porter, Aliso 17:27. 1998. Aliciella Brand, Pflanzenreich 4(250):150. 1907. TYPE: Gilia triod E Woody-based perennials, short-lived perennials, biennials, and annuals. Plants scapose with a basal leaf rosette, central leader stem, and cymose inflorescence. Lower leaves leathery, pinnate, witha strap-shaped rachis and short lobes. Flow- 540 BRIT.ORG/SIDA 21(2) ers showy in the perennial and biennial species, mostly small and inconspicu- ous in the annual species. Corolla in the large-flowered species funnelform or sometimes trumpet-shaped; blue, red, or pink. Corolla veins anastomosing (see Materials and Methods). Stamens inserted in corolla tube or sinuses. Pollen mostly yellow or cream-colored, rarely blue. Seeds not mucilaginous or only slightly so when wet. Two basic numbers, x = 9 and 8; n = 8 is common in the perennial and biennial species; polyploids are common in the annual species. Distribution and taxa.—Colorado Plateau, Rocky Mountains, and adjacent plains for the perennial and biennial species; Mojave desert and neighboring deserts for the annual species. Nineteen species. G. caespitosa, G. formosa, G. haydenii, G. heterostyla,G. humillima,G. hutchinsifolia, G. leptomeria, G. lottiae, G. mcvickerae, G. micromeria, G. nyensis, G. pentstemonoides, G. pinnatifida, G. sedifolia, G. stenothyrsa, G. subacaulis, G. subnuda, G. tenuis, G. triodon. Gilia humillima (Brand) A.G. Day ex V.E. Grant, comb. nov. Aliciella triodon var. humillima Brand, Pflanzenreich 4(250):150. 1907. Aliciella humillima J.M. Porter, Aliso 17:41.1998 Comment.—Porter (1998) has recently revised sect. Giliandra, and treats it as a genus, Aliciella. He includes the Gilia latifolia group in Aliciella, whereas | assign it toa neighboring section, Gilmania. Porter's (1998) treatment contains much information about the geographical distribution, habitats, chromosome numbers, and other features of the species. Section 6. Gilmania (H.L. Mason & A.D. Grant) VE. Grant & A.D. Grant, Aliso 3:299. 1956. Type: Gilia latifolia S. Wats. Gilia subgen. Gilmania H.L. Mi oes A.D.Grant, Madrono 9:205. 1948. Aliciella subgen. Gilmania J.M. Porter, Aliso 17:43. Woody-based perennials and annual herbs. Plants scapose with a basal rosette, central leader, and cymose inflorescence. Lower leaves with a broad blade, lobed margin, and sharp-tipped lobes. Corolla funnelform, pink. Corolla venation not recorded. Stamens inserted in corolla tube. Pollen yellow. Seeds not mucilagi- nous or only slightly so when wet. Basic number x = 9. Distribution and taxa.—Deserts from southeastern California to Utah. Two species: G. latifolia (annual) and G. ripleyi (perennial). Section 7. Campanulastrum Brand, Pflanzenreich 4(250):144. 1907. Typr: Gilia He aes A. Gray. Gilia subgen. Campanulastrum H.L. Mason & A.D. Grant, ee 9.1948. Tintinabulum Rydb.,, Fl. Rocky Mts., ed. 2, 698, 1065. 1922. Typr: Gilia filiformis in A. Gray. Gilia subgen. Tintinabulum H.L. Mason & A.D. Grant, Madrono 9:220. 1948. Small annuals. Stems very slender and wiry, branching from base and spread- ing. Pubescence glandular-puberulent, or commonly glabrous in G. filiformis. Leaves small and linear. Flowers solitary. Corolla campanulate, small, yellow or cream. Veins non-anastomosing (A. Day, pers. comm.). Stamens inserted in co- rolla throat or tube. Pollen yellow. Seeds mucilaginous when wet. Basic number x = 9, diploids. GRANT, :GILIA AND LATHROCASIS 541 Distribution and taxa.—Desert mountains, California to Utah and Arizona. Three species: G. campanulata, G. filiformis, G. inyoensis. Comment.—Some phenetic characters of sect. Campanulastrum relate it to sect. Giliastrum, other characters relate it to sects. Giliandra or Gilmania. The molecular cladograms for ribosomal ITS and chloroplast genes trnL and matk show a Campanulastrum clade adjacent to a Giliandra clade Johnson and Weese 2000). These authors list the species under generic names, Linanthus and Aliciella. The question is how to express the relationships in the taxonomic system. Sect. Campanulastrum does not fit neatly into any one of the other sections in subgen. Gilia. Including the Gilia campanulata group in sect. Giliastrum as in Grant (1959) is not the answer. Segregating it asa genus Tintinabulum (Rydberg 1922; Grant 1999) obscures the relationship. Treating this group as a section in subgen. Greeneophilia seems to be the best solution. Genus 2. Lathrocasis L.A. Johnson, Aliso 19:67. 2000. Tyre: Gilia tenerrima A. Gray. Small annuals with small flowers. Stems branching from base and ascending, Leaves linear, with one or two lateral lobes, or simple. Pubescence of tiny stipitate glandular hairs with a black dot-like head. Corolla broad-throated funnelform, white or bluish with yellow spots in throat. Corolla veins branching but not anas- tomosing Johnson & Weese 2000; Day, unpubl.). Stamens inserted in corolla throat. Pollen white. Pollen exhibiting an unusual zonocol ndition with the pores in a broad equatorial band (Grant & Day 1999). Seeds rounded, | per locule, mucilaginous when wet. Flavonoids not reported. 21n = 36, x = 9. See Johnson and Weese (2000) for a more detailed morphological description. Distribution and taxa.—One species, L. tenerrima. Western mountains from Sierra Nevada, California, to Montana, Wyoming, and Utah. Comment.—Gilia tenerrima possesses a unique combination of characters making it difficult to place in the system. On the basis of some characters, Day and I formerly placed this species in or near the Gilia campanulata group (Day 1993a: Grant & Day 1999). However, the more recent molecular evidence does not support this assignment Johnson & Weese 2000). A cladogram for rDNA ITS shows G. tenerrima as a clade adjacent to the sections Saltugilia, Arachnion, and Gilia. Cladograms for chloroplast genes trnL and matK agree with the ribosome cladogram Johnson and Weese 2000). Gilia tenerrima is widely separated from G. campanulata in all three cladograms. It also differs from the G. campanulata group in seed coat sculpturing Johnson et al. 2004). In fact, G. tenerrima falls outside the range of variation of the genus Gilia as described in this paper. It differs from Gilia as described here in the type of glandular pubescence, seed shape and number, pollen color, and distribution of pores on the pollen grains. 542 BRIT.ORG/SIDA 21(2) Gilia tenerrima resembles Allophyllum in a number of phenetic characters (Grant 1999; Grant & Day 1999). However, this indication of relationships is not supported by the molecular evidence. Gilia tenerrima and Allophyllum fall in separate major clades in the molecular cladog of Johnson and Weese (2000). Johnson and coworkers set up a new monotypic genus, Lathrocasis, for G. tenerrima (Porter & Johnson 2000; Johnson & Weese 2000). This is a good so- lution for the taxonomic problem and is followed here. Future study of L. tenerrima should include cytotaxonomic work. The few populations that have been chromosome-counted are tetraploid. Diploids could well turn up with further exploration and they might shed some light on the ancestry of the known tetraploid form. ~— DISCUSSION Phylogenetic Relationships in the Gilia Tribe The tribe Gilieae as defined by Grant (2001, 2003b, this paper) contains the temperate herbaceous members of the family with zonocolporate pollen, as contrasted with other temperate herbaceous groups which have pantoporate pollen. The tribe consisted of the genera Gilia, Ipomopsis, Eriastrum, Langloisia, and Tintinabulum in the recent treatments (Grant 2001, 2003b). In the present treatment, Tintinabulum is reduced to a section of Gilia (sect. Campanulastrum), and Lathrocasis is taken up, resulting in a tribe composed of Gilia, Lathrocasis, Ipomopsis, Eriastrum, and Langloisia. The genera fall into two grades with respect to the basic chromosome num- ber, which is x = 9 in Gilia and Lathrocasis, and x =7 in Ipomopsis, Eriastrum, and Langloisia. X = 9 is the ancestral condition in the Polemoniaceae and x = 7 is derived (Grant 1959). Gilia is regarded as basal in the tribe and the seven-paired genera as ad- vanced. The latter do exhibit some advanced phenetic characters, such as brac- teate flowering heads in Eriastrum, bilateral corollas in Langloisia, and hum- mingbird and hawkmoth flowers in Ipomopsis. The summer-blooming habit of Eriastrum and lowland species of Ipomopsis may be an advanced trait. The woody-based perennials in Gilia sect. Giliastrum appear to be basal within Gilia. The other sections represent branches in a series of radiations. The California-centered annual gilias (sects. Gilia, Arachnion, Saltugilia) are one such major branch. Section Giliandra with x =9 and also the reduced num- ber x = 8 is another. The seven-paired genera Eriastrum and Ipomopsis can be viewed as off- shoots of one or two sections of perennial gilias in subgen. Greeneophila. The small desert genus Langloisia seems to be an offshoot of Eriastrum. Lathrocasis (x = 9) appears to be related to the California-centered annual gilias. The molecular cladistic approach of Porter and Johnson (2000) and Johnson GRANT, :GILIA AND LATHROCASIS 543 et al. (2004) leads to a very different classification, as mentioned in the intro- duction (see also Grant ae First, om s. |.is broken up into numerous smaller genera (Table 1). Second, t gregate g igned to three diff tribes (Table 1). Third, these tribes contain mixtures of genera with different ancestral roots as indicated by phenetic characters. For example, Porter and Johnson (2000) group the equivalent of Grant's Gilia subgen. Gilia together with Allophyllum and Collomia in their tribe Gilieae (Table 1). This isa non-monophyletic group- ing according to strong phenetic evidence (Grant 1998, 2001, 2003b). The tribe Loeselieae of Porter and Johnson (2000) is also non-monophyletic, containing amixture of Loeselia and most of Gilia subgen. Greeneophila (Table 1). Loeselia and Gilia have different roots and are assigned to different subfamilies in the taxonomic system (Grant 2003b). Gilia Is Not Polyphyletic Johnson et al (1996) state that Gilias. lis polyphyletic, repeating the statement several times for emphasis. The same conclusion is stated in other molecular systematic papers (Porter 1998; Porter & Johnson 2000; Weese & Johnson 2001). The authors do not present an explicit verbal justification for their claim. How- ever, it is clear from the context that the basis for their conclusion is a broad incongruence between the existing taxonomic classification of Gilia s. |. and their molecular cladograms. The comprehensive family-wide cladograms of Johnson et al. (1996) were the forerunner of a reclassification of the family as a whole including the Gilia complex. The molecular evidence consisted of the sequence variation in the chloroplast gene matK. Johnson et al. (1996) made the assumption that the cla- dograms for matK provide a reliable guide for the phylogeny of the species in the family. The clades were assigned informal taxonomic names. Porter and Johnson (2000) set out to construct a phylogenetic classifica- tion system based on molecular evidence; and in practice they took up the cpDNA matK clades of Johnson et al. (1996) and transformed these into formal taxonomic groups with taxonomic names. The primary and secondary molecu- lar clades became subfamilies and tribes respectively. Third-order clades be- came genera or small sets of genera. Evidence from studies of other DNA re- gions played a supporting role: rDNA ITS (Porter 1997, 1998) and rDNA ITS and cpDNA trnL Johnson & Weese 2000). Full descriptions of phenetic characters were attached to the taxonomic groups of Porter and Johnson (2000), but it is difficult to see what effect, if any, the phenetic characters had on the circumscription of the groups. In cases of conflict between molecular and phenetic evidence, the molecular evidence rou- tinely prevails. A comparison of the Porter and Johnson (2000) system with other systems 544 BRIT.ORG/SIDA 21(2) before 2000 is thus a comparison of a cladistic system based primarily on DNA data with a taxonomic classification based primarily on phenetic characters. In the molecular cladograms of Johnson et al. (1996), clades containing the subgroups of Gilias. | are scattered in different positions on the graph, and other genera such as Eriastrum and Ipomopsis lie between them. This topology indi- cates non-monophyly in cladistics, which uses the cladistic definition of mono- phyly; and it is probably the basis for Johnson et al’s (1996) conclusion that Gilia is “extremely” polyphyletic. However, the topology is quite consistent with the concept of monophyly used by taxonomic students of Gilia. This isan example of a “false accusation” of polyphyly, as mentioned in the introduction. The pattern of the molecular cladograms is consistent with the phyloge- netic hypothesis that Gilias. lisa basal multisection genus in the tribe Gilieae. The sections differ in molecular as well as phenetic characters. Some sections of Gilia have given rise to derived genera such as Eriastrum and Ipomopsis, and these lie between sections of Gilia in the cladograms. The same pattern is seen in molecular cladograms of other plant groups and is often misinterpreted as evidence for polyphyly (see Grant 2003a). Actually, the molecular evidence is in reasonably good agreement with the taxonomic classifications of Gilias. 1. The big incongruence is between the taxo- nomic classification and the molecular-based system. This suggests that the incongruence, or much of it, has developed in the process of converting mo- lecular clades into taxa. — The Genus Concept in Gilia The goal in both evolutionary taxonomy and molecular cladistics is to circum- scribe genera so that they are natural or monophyletic. Beyond this basic goal, it is possible, in either school, to adjust the boundaries in various ways ranging from lumping to splitting. The two schools also apply different criteria for cir- cumscription: significant phenetic differences between genera in evolutionary taxonomy, distinctive molecular clades in molecular cladistics. The results are seen in current treatments of Gilia where one school’s sections are another school’s genera. Alva Day and Land other earlier evolutionary taxonomists such as Herbert Mason (Mason & Grant 1948) have favored a broad multisection genus Gilia because the broad circumscription expresses the interrelationships of the sub- branches. Splitting the sections off as a series of segregate genera (Aliciella, Giliastrum, etc.) obscures their interrelationships. Having one generic name (Gilia) fora related set of sections, rather than a different generic name for each subdivision, reinforces the sense of interrelationship, and in addition reduces the memory burden, and facilitates preliminary identification in the field or herbarium. Gilids. l.is more difficult to define diagnostically than its constituent sec- GRANT, > GILIA AND LATHROCASIS 545 tions, but it can be defined. Gilia as treated here is a genus of temperate herba- ceous Polemoniaceae that has zonocolporate pollen, a basic chromosome num- ber of x = 9,a spring-blooming habit, and generally angular seeds. ACKNOWLEDGMENTS AlvaG. Day furnished valuable information and opinions. Tom Wendt was help- ful in the U.T. Herbarium. Rancho Santa Ana Botanic Garden and Field Museum made loans of herbarium specimens and J.M. Porter donated some specimens. Karen A. Grant read the manuscript. Their help is gratefully acknowledged. REFERENCES BentHAM, G.and J.D. Hooker. 1873-1876. Genera plantarum. Vol. 2.Reeve and Co., London. Branpecte, T.W. 1903. Notes and new species of lower California plants. Zoe 5:155—-174. Cronn, R.C.,R.L. SMALL, T. HASeLcorn, and J.F. Wenpet. 2002. Rapid diversification of the cotton genus (Gossypium, Malvaceae) revealed by analysis of sixteen nuclear and chloroplast genes. Amer. J. Bot. 89:707-725. Day, A.G. 1993a. Gilia. In: J.C. Hickman, ed. The Jepson manual: higher plants of California, pp. 828-837. Univ. of California Press, Berkeley, California. Day, A.G.1993b. New taxa and nomenclatural changes in Allophyllum, Gilia,and Navarretia (Polemoniaceae). Novon 3:331-340. Day, A.G.and R. Moran. 1986. Acanthogilia,a new genus of Polemoniaceae from Baja Cali- fornia, Mexico. Proc. Calif. Acad. Sci.44:111-126. Fercuson, C.J. and R.K. Jansen. 2002. A chloroplast DNA phylogeny of eastern Phlox (Polemoniaceae): implications of congruence and incongruence with the ITS phylog- eny. Amer. J. Bot. 89:1324-1335. Grant, A.D. and V. Grant. 1956. Genetic and taxonomic studies in Gilia. 8. The cobwebby gilias. Aliso 3:203-287. Grant, V. 1959. Natural history of the phlox family. Martinus Nijhoff, The Hague. Grant, V. 1998. Primary classification and phylogeny of the Polemoniaceae, with com- ments on molecular cladistics. Amer. J. Bot. 85:741-752. Grant, V. 1999. Classification of the genus Gilia (Polemoniaceae). Phytologia 84:69-86. Grant, V.2001.A guide to understanding recent classifications of the family Polemoniaceae. Lundellia 4:12-24. Grant, V. 2003a. Incongruence between cladistic and taxonomic systems. Amer. J. Bot. 90:1263-1270. Grant, V,2003b. Taxonomy of the Polemoniaceae:the subfamilies and tribes. Sida 20:1371- 1386. Grant, V. and A.G. Day. 1999. Transfer of some species from Gilia to Allophyllum and Tintinabulum, and the effects of the transfer on the generic definition of Gilia (Polemoniaceae). Phytologia 84:368-382. Grant, V.and A.D. Grant. 1954. Genetic and taxonomic studies in Gilia. 7. The woodland gilias. Aliso 3:59-91, 546 BRIT.ORG/SIDA 21(2) Grant, V. and T. Wenor. 2003. Proposal to reject the name Gilia grinnellii (Polemoniaceae). Taxon 52:145-146. Gray, A. 1886. Synoptical flora of North America. 2nd ed.Vol.2.|vison, Blakeman, and Taylor. New York. JOHNSON, L.A., KH. Huish, and J.M. Porter. 2004. Seed surface sculpturing and its systematic significance in Gilia (Polemoniaceae) and segregate genera. Internat. J. Plant Sci. 165:153-172 JOHNSON, L.A, L. ScHuttz, D.E. Soutis, and PS. Soutis. 1996. Monophyly and generic relationship of Polemoniaceae based on matK sequences. Amer. J. Bot. 83:1207-1224 Jounson, L.A. and T.L.Weese. 2000. Geographic distribution, morphological and molecular characterization, and relationships of Lathrocasis tenerrima (Polemoniaceae). Western N.Amer. Naturalist 60:355-373,. Mason, H.L. and A.G. Grant. 1948. Some problems in the genus Gilia. Madrono 9:201-220. McKinnon, G.E., D.A. Steane, B.M. Potts, and R.E. pete 1999, Incongruence between chloroplast and species pl! Amer. J. Bot. 86:1038- 1046, Mivuxen, J. 1904. A review of Californian Polemoniaceae. Univ. Calif. Publ. Bot.2:1-71. Peter, A. 1897. Polemoniaceae. Die nattirlichen pflanzenfamilien 4(3):40-54. Porter, J.M. 1997, Phylogeny of Polemoniaceae based on nuclear ribosomal internal tran- scribed spacer DNA sequences. Aliso 15:57-77. Porter, J.M. 1998. Aliciella, a recircumscribed genus of Polemoniaceae. Aliso 17:23-46. Porter, JM. and L.A. JoHnson. 2000. A phylogenetic classification of Polemoniaceae. Aliso 19:55-91, Prather, L.A., C.J. Fercuson, and R.K. Jansen. 2000. Polemoniaceae phylogeny and classifica- tion; implications of sequence data from the chloroplast gene ndhF. Amer. J. Bot. 87:1 300-1308. RiesegerG, L.H. 1991. Homoploid reticulate evolution in Helianthus: evidence from riboso- mal genes. Amer. J. Bot. 78:1218-1237. RieseBerG, L.H., S.M. BeckSTROM-STERNBERG, A. Liston, and D.M. Arias. 1991. Phylogenetic and sys- tematic inferences from chloroplast DNA and isozyme variation in Helianthus sect. Helianthus (Asteraceae). Syst. Bot. 16:50-76 RieseserG, L.H., J. WHiTton, and C.R. Linoer. 1996. Molecular marker incongruence in plant hybrid zones and phylogenetic trees. Acta Bot. Neer. 45:243-262, SmitH, D.M., C.W. Grennie, J.B. Harsorne, and C.A. Wittiams. 1977. Flavonoid diversification in the Polemoniaceae. Biochem. Syst. Ecol. 5:107-115. Turner, B.L. 1994. Taxonomic overview of Gilia, sect. Giliastrum (Polemoniaceae) in Texas and Mexico. Phytologia 76:52-68. Weese, T.L. and L.A. JoHNson. 2001. Saltugilia latimeri: a new species of Polemoniaceae. Madrono 48:198-204. Wuittemore, A.T. and B.A. ScHAaAt. 1991. Interspecific gene flow in sympatric oaks. Proc. Natl. Acad. Sci. 88:2540-2544, | AA | (KA cr INF eae). = A NEW CITHAREXYLUM (VERBENACEAE) FROM ISLA SOCORRO, REVILLAGIGEDO ARCHIPELAGO, MEXICO José Luis Leon de la Luz Fernando Chiang Centro de Investigaciones Bioldgicas fo to de Botdnica del Noroeste (CIBNOR) Instituto de Biologia, UNAM al 128 Apdo. Postal 70-233 La Paz, Baja California Sur, 23000, MEXICO México D.F.04510, MEXICO jlleon04@cibnor.m ABSTRACT A new species of Verbenaceae, Citharexylum danirae Leon de la Luz et Chiang, is described and illustrated. It is known only from the type collection at the north face of Evermann volcano in the remote Socorro Island, the largest of the Revillagigedo Archipelago, in the tropical Mexican Paci Ocean. Its relationship to C. caudatum L. and C. affine D. Don is discussed. ic RESUMEN Se describe e ilustra una ie de la familia de las Verbenaceas, C ical danirae Leon de la Luz et Chiang. Hasta hoy es ole conocida del ejemplar tipo, procedente de la vertiente norte del volcan Evermann, en la remota Isla Socorro, en el archipiélago de las Islas Coes ubicada en el Pacifico tropical de México. Se discute su relaci6n con C. caudatum L y con C. affine D. Don. Socorro Island is part of the Revillagigedo Archipelago, a group of three islands scattered off the tropical west coast of Mexico (18°45'N, 111°00'W). Socorro is located 400 km south of the southern tip of the Baja California peninsula, and about 580 km west-southwest of Cabo Corrientes, Jalisco, on the nearest west central Mexican mainland coast. The floristic knowledge of the island and the archipelago has grown con- tinuously since the late 19th century as result from sporadic botanical explora- tions, mainly describing new taxa and distributional records. Several floristic checklists also have been published Johnston 1931; Miranda 1960; Levin & Moran 1989). The latter is the most complete, cataloguing the flora of the is- lands, particularly that of Socorro, which is the largest and more diverse in spe- cies (160 taxa of vascular plants), plant communities, and the richest in plant endemism (27 percent). From three botanical explorations to Socorro Island by the first author (from 1988-1990), almost 180 duplicates were sent to Dr. Geoffrey Levin, then botanist at the San Diego Museum of Natural History (SD) and head of the Vas- cular Flora of Socorro Island Project, to contribute to the floristic compilation then in preparation. Of these collections, twelve taxa represent new distribu- tional records (Levin & Moran 1989). The expeditions provided a total of 25 SIDA 21(2): 547-551. 2004 BRIT.ORG/SIDA 21(2) collecting days on Socorro Island that included the first-ever collections on the north side of Evermann Volcano (1150 m). The collections made on February 25th 1990 on the north face of Evermann Volcano revealed several shrubby species, such as Spermacoce nesiotica (B.L. Rob.) G. A. Levin, Chiococca alba(L.) Hitchc., Dodondea viscosa Jacq., Zanthoxy- lum insulare Rose, Lepechinia hastata (A. Gray) Epling subsp. socorrensis Moran, and Rhamnus sharpii M.& L.A. Johnston. These species occur in the scrubland that covers the middle elevations of the island. A terrestrial orchid (cf. Habenaria), collected in its vegetative stage, was never identified and repre- sented the only member of this family with this habit on the island. Another collection (Leén de la Luz 4518) had remained undetermined in the HCIB her- barium until recently. As it is known only from one specimen it is presumed to be rare on the island. It was collected at 805 m, the highest part of the Mixed or Tropical Scrubland plant community, according to Miranda (1960) and Leon de la Luz et al. (1996). The specimen was initially thought to have only inmature flowers, but a recent thorough examination revealed a couple of mature flowers, whose dis- section enabled its placement as Citharexylum. This is only the third genus thus recorded for Verbenaceae on Socorro Island; the other being Verbena and Lan- tana (Levin & Moran 1989). The specimen was compared with descriptions of Citharexylum species in floras and monographs available for Mexico: Veracruz (Nash & Nee 1984), the west-central sector (Rzedowski @ Rzedowski 2002), the Sonoran Desert (Shreve & Wiggins 1964) the Baja California Peninsula (Wig- gins 1980), and the material examined by Moldenke (1958). Although seed and fruiting material was unavailable, it was determined that the specimen repre- sent a new spec cies differing from other taxa in both leaf and flower structure. The new species is herein described. Citharexylum danirae Leon de la Luz & Chiang, sp. nov. (Fig. 1). Type: MEXICO. Isa SOCORRO: Revillagigedo Archipelago: N side slopes of Evermann Volcano, mixed or tropical scrubland, 805 m elevation, 25 Feb 1990, José Luis Leon de la Luz 4518 (HOLOTYPE: HCIB). Frutex ad 3 m altus; folia opposita 7-10 cm longa, 4-6 cm lata, ovata, apice acuta, glabrata; ongls, inflorescentiae racemosae 12-18 cm longae, pedicellis 1-2 mm longis, bracteatis, bracteis 2 2mm foliaceis, oblongo-lanceolatis; corolla albidi, 5 mm longa; calycis tubo ca. + mm sepalis parvis, petalis parvis; stamina robusta ca. 1.5 mm longa, filamenta parva, robusta; ovarium pyriforme, robustum, stylo parvo, ad apicem furcato, lobis 2, stigmatosis, parvis; fructus et semina ignota. Shrub to 3 m high, stems tetragonal, nodes with prominent leaf and bundle scars; leaves opposite, ovate, 4-6 cm wide, acute acuminate at the apex, rounded at the base, the margins entire, the leaf blades subglabrous with sparse simple trichomes (less than 0.5 mm), the upper surface lustrous when young, less so with age, with 3-4(-5) parallel veins per side; petioles 15-25 mm long, canaliculate; racemes terminal, few branched, 12-18 cm in length, 20-35 LEON DE LA LUZ AND CHIANG, A NEW SPECIES OF CITHAREXYLUM FROM MEXICO Fic. 1. Cith J 4 uy Rages act Qthi \n; * £ Flower. c) Flower bud and bract. d) Flowering branch. Taste 1.Morphological comparison between Cithare vingm da BRIT.ORG/SIDA 21(2) irae, C. caudatum, and C. affine. Characters Citharexylum caudatum C. affine C. danirae Habit shrubby to arborescent = shrubby to arborescent — shrubby Leaf size 8-17 cm x 2-6cm 7-10 cm xX 4-6 cm Leaf shape oblong-elliptic ovate, lanceolate-oblong ovate Leaf apex short acuminate to acute acuminate abruptly acuminate Leaf base cuneate rounded rounded Petioles length Inflorescences —2cm racemes axillary and terminal ] racemes terminal 1.5-2.5cm racemes terminal Pedicels Timm 1-2 mm Bracts length up to 1 mm up to 1mm 2-2.55mm Corolla white, 3-4 mm pale blue, 4-6 mm white-greenish,5 mm Corolla lobes 1-2 mm, glabrous 2-4 mm, glabrescent 1 mm, glabrous Pistil slender slender stout Anthers slender slender stout (+ 1.5 mm) Calyx atanthesis — tubular-campanulate, cyathiform, 3 mm tubular, 4mm mm flowered; pedicels usually alternate along the axis, 1-2 mm; bracts oblong-lan- ceolate, up to 2 mm, longer than the pedicels; calyces to 4 mm in length at an- thesis, shallowly 5 toothed, 5 nerved; corollas white greenish, twice as long as the calyx tubes, + 5mm in length, the short lobes 5, entire, not ciliate; stamens 4, anthers stout, 1.5 mm; staminode |; ovary stout, ovoid, stout, styles 2, short; stigmas discoid-capitate; mature fruit and seeds unknown. Etymology.—The new species is named in honor of Miss Danira Leon, the first author’s much-loved daughter. This new species is close to C. caudatum L.,a widespread species in tropi- cal Mesoamerica and northern South America, whose main resemblance is the shiny character of the upper leaf surface due to small shiny scales, although such scales seem to be only present in this new taxon at low density on young leaves. Superticially, the shape of the leaves resembles those of C. affine D. Don, but the flowers differ significantly. A comparison of the new species with the two species mentioned above is presented in Table 1. The type locality has been explored botanically only once. This island sec- tor probably harbors more interesting plants. ACKNOWLEDGMENTS We thank all the CIBNOR colleagues for those memorable expeditions to Socorro Island from 1988 to 1990. Thanks also to Ira Fogel for proofing the text and Os- car Armendariz for the beautiful drawings. Critical revision by Sida, Contribu- tions to Botany editorial staff, particularly Jim Henrickson, enhanced the qual- ity of the manuscript. LEON DE LA LUZ AND CHIANG, A NEW SPECIES OF CITHAREXYLUM FROM MEXICO 551 REFERENCES Jounston, |.M. 1931. The flora of the Revillagigedo Islands. Proc. California Acad. Sci. ser. 4, 20:9-104 LEON DELA Luz,J.L., A. Breceoa, and R. Coria. 1996.Las comunidades vegetales en Isla Socorro, México. Sida 17:215-230. Levin, G.A. and R.V. Moran. 1989. The vascular flora of Isla Socorro, Mexico. San Diego Soc. Nat. Hist. Mem. 16:1-71. Miranpa, F. 1960. La Isla Socorro: Vegetacion. Monogr. Inst. Geofisica, Univer. Nac. Auton. México 2:129-152. Moupenke,H.N. 1958. Materials toward a monograph of the genus Citharexylum.Phytologia 6:242—256(I); 262-320(II); 332-368(III); 383-43 2(IV); 448-505(V). Nasu, D.E.and M. Nee. 1984. Verbenaceae. Fasciculo 41 de Flora de Veracruz. Instituto Na- cional de Investigaciones sobre Recursos Bidticos. Xalapa, Veracruz. Pp. 1-154. SHreve, F.and I.L. Wicains. 1964. Vegetation and flora of the Sonoran Desert, 2 vols. Stanford University Press. Stanford, CA. R7eDowski, J.-and G.C. pe Rzepowski. 2002. Familia Verbenaceae. Fasciculo 100 de la Flora del Bajio y de regiones adyacentes. Instituto de Ecologia, Centro Regional del Bajio. Patzcuaro, Michoacan. Pp. 1-145. Wicains, LL. 1980. Flora of Baja California. Stanford University Press, Stanford, CA. 552 BRIT.ORG/SIDA 21(2) BOOK REVIEW DouG as Crase. 2004. Both: A Portrait in Two Parts. (ISBN 0-375-42266-8, hbk.) Pantheon Books, New York. (Orders: http://www.random house.com/pan- theon/catalog/). $25.00, 303 pp., 6 drawings, 34 photographs, 6" x 8". Imagine two urbane, erudite, amusing (in several languages) characters. Make one an English aristo- crat and the other heir to an American railroad fortune. Give them a group of friends who: are the idd brightest members of the avant-garde on both side of the Atlantic Ocean. A pS parts for such luminaries as Juan Miro and Cary Grant. It sounds like the makings of a piece by Noel Coward or Evelyn Meee Wh } harming biography, Both: A Portrait in Two Parts by Douglas Crase. It is the ory of o intertwined Tes of the eminent botanist, Rupert Charles Barneby, and the aie -artist, Harry cay Dillon Ripley. They met as students at Harrow, drawn together by a mutual love of Latin and a mania for ieee: plants. The relationship persisted through Uni- versity and beyond, resulting in t inheritance of Barneby by his father. Ripley was an orphan. wo men remained never a 48 years. uthor says that Ripley “was the enabling influence of Barneby’s early direction as a bota- The nist th antes his money, his affection, and the magnificent garden at his estate in Sussex.” This place contained three greenhouses, every kind of special habitat, a herbaceous border, and a private her- barium. On the other side, Barneby with his inbred noblesse, was a buffer for the shy Ripley, who suffered from uncontrolled blushing, leading from embarrassment to several visits to the United States, the couple moved to a — in 1939. Botany and cours continued to be at the center of their lives, but Ripley, the artist, also financed the Tibor de Nagy Gallery, a non-commercial enterprise, which became one of the most influential galleries in New York, when after the Second soe Wat New uae was the saat leader in the arts. Ripley and Barneby I | g world of literature, music, and paintin g. All of this is affectionately and gracefully recounted by Mr. Crase, a former MacArthur Fellow. e is not a botanist, but he was a friend of Barneby’s and writes knowingly of his hard work and aunt Although Ripley was regarded as something of a dilettante, he too left an impressive legacy. However, it is the relationship which is subject of this delightful book.—Ruth Ginsburg, Bo- tanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A. SIDA 21(2): 552. 2004 SPOROBOLUS (POACEAE: CHLORIDOIDEAE: CYNODONTEAE: ZOYSIEAE: SPOROBOLINAE) FROM NORTHEASTERN MEXICO Paul M. Peterson Jesus Valdés-Reyna Department of Bota Departamento de Botanica National Museum Patil istory Univerisidad Autonoma Agraria ‘Antonio Narro” mithsonian Institution Buenavista “Saltillo iene DC 20013-7012, U.S.A. Coahuila 25315, MEXICO peterson.paul@nmnh.si.edu jvaldes@narro.uaaan.mx Juan Javier Ortiz-Diaz ta de Botanica Un iersida! de Aut6noma de Yucatan Mérida, Yucatan 97000, MEXICO odiaz@tunku.uady.mx ABSTRACT A taxonomic treatment of Sonus R. Bh for nletEbeastee México (Coahuila, Nuevo Leon, and Tamaulipas), is given.S Sporobolus are recognized in the study area. Sporobolus aimides subsp. regis is endemic to Coatmilacanas atrovirens and S. een are endemic to México. Keys for determining the species, descriptions, distributions ined, illustrations, synonymies, and a brief discussion indicating relationships among all native and adventive species of Sporobolus i in northeastern México are provide RESUMEN Se presenta un estudio taxondmico de Sporobolus E Br. para el noreste de México (Coahuila, Nuevo P Pe! P Leon y Tamaulipas). S ndi y una subespecie de Sporobolus para el area de estudio. S l ides subsp. regis es wedenies para Coahuila y S.atrovirens y S. spiciformis son dene para México. Se incluyen claves para determinar las especies, descripciones, distribuciones | ilustraciones, sinonimias, y una discusion breve indicando l laci las | ivas y adventicias de Sporobolus para el noreste de México. Si Northeastern México (Coahuila, Nuevo Leon, and Tamaulipas), covers an area of 291,955 km2 or 15 % of the total land of the country. This area includes por- tions of two natural regions known as the Chihuahuan and Tamaulipan Deserts. These regions are considered a center of origin and diversification of arid and semi-arid plant species. As part of the current revision of the grass flora of northeastern México, an examination of the taxonomy and distribution of the species of Sporobolus, was begun to aid the agriculture and livestock industries. This study treats 17 species and one variety, for a total of 18 taxa. Sporobolus R. Br. is a worldwide genus of more than 160 species occurring in the tropics, subtropics, and warm temperate regions (Clayton & Renvoize SIDA 21(2): 553-589. 2004 554 BRIT.ORG/SIDA 21(2) 1986; Peterson et al. 1997; Watson & Dallwitz 1992). There are 72 native species of Sporobolus in North, Central, and South America; 27 native in the United States and Canada; and 26 native in México (Espejo-Serna et al. 2000; Peterson et al. 2001, 2003, 2005). The genus is characterized by having single-flowered spikelets, 1-nerved lemmas, fruits with free pericarps or “modified caryopses” as proposed by Brandenburg (2003), and ligules witha line of hairs (Peterson et al. 1995, 1997). Species of Sporobolus generally inhabit dry, saline or alkaline sandy to clay loam soils in prairies, savannahs, and along disturbed roadsides (Peterson et al. 1997) The subtribe Sporobolinae, as currently circumscribed, consists of three New World genera: Calamovilfa (A. Gray) Hack., Spartina Schreb., and Sporobolus (Peterson et al. 2005). In the New World the Sporobolinae share most of the same character trends as for the tribe Zoysieae, i.c., spikelets with a single floret, spiciform inflorescences of numerous deciduous racemelets disposed along a central axis, lemmas usually rounded and rarely with apical awns, and glumes often modified and oddly shaped, but differ by having modified caryposes (pericarps reluctantly free in Spartina), spikelets oriented abaxially along the aixis (lemma is facing the rachis), lemmas that are similar in texture to the glumes, and paleas that are relatively long and about the same length as the lemma (Peterson et al. 2005). Within Sporobolus, Stapf (1898) first divided the genus into two sections: Chaetorhacia Stapf and Eusporobolus Stapf. Pilger (1956) then divided the lat- ter section, which he elevated to Sporobolus subg. Sporobolus (Stapf) Pilg., into six groups based on life form and characteristics of the glumes and panicles. Based on caryopsis morphology, Bor (1960) divided Sporobolus into five rather unnatural groups (Baaijens & Veldkamp 1991). Working on the Malesian spe- cies, Baaijens & Veldkamp (1991) divided Sporobolus subg. Sporobolus into five sections based on overall morphology with special attention given to inflores- cence branching. More recently, Weakley & Peterson (1998) recognized the Sporobolus floridanus complex to include five species in the southeastern United States and based on nuclear ribosomal DNA ITS sequences, Ortiz-Diaz & Culham (2000) presented evidence to support the recognition of at least 10 clades within Sporobolus. Recent major revisions of Sporobolus include Boechat & Wagner (1995) for Brazil, Simon & Jacobs (1999) for Australia, and Peterson et al. (2003) for the United States and Canada. The following taxonomic treatment contains a se for determining the species, descriptions, distribution, specimens examined, illustrations, and syn- onymies for all native and adventive species of Sporobolus in northeastern Mexico. This study is based on the examination of herbarium specimens from ANSM, COCA, MEXU, MO, NMSU, TEX, UAT, and US, including the type speci- mens of most of the species studied. PETERSON ET AL., 555 TAXONOMIC TREATMENT a aan R. Br, Prodr. 169. 1810. Type: Sporobolus indicus (L) R. B ted L.K.G. Pfeiffer, Nom. Bot. 2:1274. 1874, also by Nash, IIL FI. N. US. (ed 2); 1194, 1913) Plants annual or perennial; sometimes rhizomatous, rarely stoloniferous. Flow- ering culms 10-250 cm tall, erect rarely mat-forming, caespitose (often form- ing large clumps), glabrous; leaf sheaths longer or shorter than the internodes usually with smooth margins, occasionally ciliate; ligule ciliate, a line of hairs; blades 3-70 cm long, 1-15 mm wide at base, filiform or linear, flat, involute, or terete, not pungent, cauline without auricles. Inflorescence a panicle 0.5-80 cm ong, 0.3-30 cm wide, exserted or partially included in upper sheath; rachis smooth; primary branches appressed, spreading, divaricate, or reflexed from the main axis, solitary or loosely whorled, sometimes with capillary branches terminating ina spikelet; secondary branches appressed or spreading; pedicels erect, rarely secund, glabrous, scaberulous or scabrous; cleistogamous spikelets occassionally present, in axillary inflorescences. Spikelets 1-4(-7) mm long, solitary, laterally or dorsally compressed, sometimes terete; disarticulation usu- ally above the glumes, commonly above the upper glume, occasionally below with the lemma and palea falling as a unit; glumes shorter or longer than the florets, very unequal, smooth, glabrous; lower glume without midvien or 1- veined; upper glume about the same length as the lemma, usually awnless, 1- veined; florets 1 per spikelet; lemma entire, awnless, glabrous or hairy, pubes- cent to pilose, l-veined, rarely 3-veined, membranous with glabrous veins; palea glabrous, smooth, membranous, margins not enfolding the fruit, 2-veined, of- ten splitting as grain matures; lodicules 2 or sometimes absent, truncate; sta- mens 2 or 3; anthers yellow, reddish-purple, or olivaceous-plumbeous; stigmas 2. Modified caryopsis a follicoid fruit with a free pericarp, commonly swelling and mucilaginous when wet; hilum punctiform; embryo with an epiblast, scutellar tail, and elongated mesocotyl internode (formula P+PF), endosperm hard. Base chromosome number, x = 9, and 10. Named from Greek Sporo, ‘seed’, and bolos, ‘a throw’, referring to the free seeds. Comments.—The following five species included in this study have been placed in four different sections of Sporobolus: S. atrovirens, S. indicus, and S. jacquemontii (sect. Sporobolus), S. virginicus (sect. Virginicae Veldkamp), and S. purpurascens|sect. Triachyrum (Hochst. ex A. Braun) Veldkampl. In addition to this, we recognize three prominent “groups” within the northeastern Mexican species of Sporobolus: 1) Airoidae - culms tall and densely caespitose; leaf blades as viewed in cross section (Annable et al. 1992) with bundle sheath extensions, first order vascular bundles with flattened adaxial ribs, second order vascular bundles with triangular pls and adaxial furrows above the third order vascu- lar bundles; spiklelets d d; modified caryopses plump; includes: S. diroides, S. buckleyi, and S. wrightii; 2) Cryptandrae—Leaf blades as viewed 556 BRIT.ORG/SIDA 21(2) in cross section (Annable et al. 1992) with round adaxial ribs with furrows be- tween each adjacent vascular bundle, distinctive fan-shaped bulliform cells, and lack bundle sheath extensions; spikelets laterally compressed; glumes with scabrous nerves; endosperm translucent yellow or orange; includes S. contrac- tus, S. cryptandrus, S. flexuosus, S. giganteus, and S. nealleyi;, 3) Pyramidatae - Panicle branches whorled or sub-whorled; embryo pandurate witha scutellum edge; includes S. coahuilensis and S. pyramidatus. These three groups parallel those presented by Ortiz-Diaz & Culham (2000) where their group E corre- sponds to our Cryptandrae and their group G, in part, corresponds to our Airoidae. Our Cryptandrae group is identical to the Sporobolus cryptandrus complex presented by Annable et al. (1992), and their Sporobolus airoides group includes S. palmeri Scribn. and S. splendens Swallen that do not occur in the study area. Morphologically, S it d S. spiciformis do not appear to be allied with other northeastern Mexican species of Sporobolus. Sporobolus compositus appears to be allied with S. neglectus Nash and S. vaginiflorus (Torr. ex A. Gray) Alph. Wood from the United States and Canada, whereas S. spiciformis is per- haps related to S. phleoides Hack. from Argentina. Sporobolus compositus, S. neglectus, and S. vaginiflorus all have contracted panicles that are included in the uppermost sheath and an embryo nearly as long as the modified caryopsis. Sporoblous spiciformis and S. phleoides also have contracted panicles with lat- erally compressed spikelets and upper glumes 1/4-1/2 as long as the floret. KEY TO THE SPECIES OF SPOROBOLUS IN NORTHEASTERN MEXICO 1. Plants with rhizomes. 2. Panicles 3-10 cm long, 0.4-1.6 cm wide, contracted, spikelike, dense; culms 10- 65 cm tall 16.S. virginicus 2. Panicles 30-40 cm long, 10-15 cm wide, open, diffuse, subpyramidal; culms 80- tall b 1. Plants without rhizomes. 3. Plants annua 4. Lower panicle nodes with 1-2 branches; spikelets 0.7-1.2 mm long; lemmas ; m lon 2.S.atrovirens 4. Lower panicle nodes with 7-20 branches (whorled); spikelets 1-1.8 mm long; lemmas 1.2-1.7 mm lon 5. Pedicels 0.1-0.5(-1) mm long, appressed 5. Pedicels (2-)3—6(-8) mm long, widely spreading 3. Plants perennial. 6. a apt usually more than 2.6 mm long. wer panicle nodes with 3-5 branches 7. ee panicle nodes with 1 or 2 (—3) branches. @ Panicl wl ad av ill feelee toe 1b.S. airoides subsp. regis 14.S. pyramidatus 4, S.coahuilensis 13. S. purpurascens . conspicuous tuft of hairs at the summit 5.S. compositus 8. Panicles all terminal, elongated; sheaths with a conspicuous tuft of hairs at the summit. PETERSON ET Al 9. Culms 35-100(-120) cm tall, 2-4(-5) mm diameter near the base; mature panicles 0.2-0.8(-1) cm wide; anthers 0.3-0.5 mm long contractus 9. Culms 100-200 cm tall, (3-)4-10 mm diameter near base; mature panicles 1-4 cm wide; anthers 0.6-1 mm long 6. Spikelets 1-2(-2.9) mm lon Lower sheaths Se laterally compressed and keeled 3. S.buckleyi 10. ie sheaths rounded. se Sel a collar with a conspicuous tuft of white hairs. . Panicles contracted, spike-like; branches appress 13. Culms 35-100(-120) cm tall, 2-4 bas 9. S.giganteus sed —5) mm diameter near the 6.5. contractus 13. in. 100-200 cm tall, (3-)4-10 mm diameter near the base 9.S. giganteus 12. Panicles open, branches spreading at least from the middle of the rachis to the apex, the siete sometimes included 14. Culms 10-40 cm tall,0.7-1.2 mm di near base, the hard and knotty; blades a spreading at right angles ____ Ce S.nealleyi 14. Culms 30-100(-120) cm tall, 1-3 mm diameter near base, the base not hard and knotty;blades erect or ascending, not stiff 15. Panicles usually exserted, branches divaricate and flexu- ous, usually tangled between branches or panicles;lower glumes 0.9-1.5 mm long 8.5. flexuosus 15. Panicles usually included at the base, branches appressed or ascending, not markedly flexuous, not tangled; lower glumes 0.6-1.1 mm lon 7. S.cryptandrus . Leaf sheaths oe oad glabrous (sparsely appressed pilose in S. airoides subsp — 16. Pancles ie 3 cm wide, spikelike, the | hes appressed to main axis. 17. Pancles 9-17 cm long, 3-5(-10) mm wide, whitish; glumes unequal, more than two thirds as long as the floret ___ 15. S.spiciformis 17. Panicles 7-60 cm long, 5-12 mm wide, not whitish; glumes about equal, less than two thirds as long as the floret 18. Spikelets 2-2.6(-2.7) mm long; upper glumes usually 1/ 2-2/3 as long as the florets, the apex acute to obtuse, entire 10. S. indicus 18. Mee 1.4-1.8(-2) mm long; upper glumes usually less 1/2 as long as the florets, rarely longer, the apex trun- cate, erose to gemuculate 11.S, jacquemontii 16. Panicles 3-25 cm wide ( s only 0.3 cm wide in immature 5. pyramidatus), open, the branches widely spreading at least from the middle of the rachis to the apex 19. Panicle branches widely divaricate and flexuous, usually tangled within branches or panicles 19. Panicle branches widely Open or erect, Het elise: 20. oan 10s 40 cm tall, deli II;1 8.S. flexuosus if +1 [5h with few cili- ate hairs on the margins and summi it. 21. Panicle branches arranged in whorls at lower nodes; 558 BRIT.ORG/SIDA 21(2) mature panicles pyramidal (immature and extreme forms spikelike); upper glumes 1.2-1.8 mm long 14.8. pyramidatus 21. Panicle branches oy or paired but not in whorls e lower nodes; mature panicles oblong to ovoid; upper glumes 0.4-0.7 mm long 2.S.atrovirens . Culms 30-100(-120) cm tall, robt Ist perennials, often forming large tough leafy tussocks; leaf blades cauline; sheaths usually bearded with long hairs on the margins and summit. 22. Panicles 10-45 cm long, the branches naked near the base; pedicels 0.5-2 mm long, usually erect, spreading 1.S.airoides 22. Panicles 20-60 cm long, the branches densely flow- ered near the base; pedicels 0.2-0.5 mm long, ap- pressed 17.8. wrightii NR [o) la. § iakeare airoides (Torr) Torr. subsp. airoides, Pacific Railr. Rep. Parke, Bot. 7:21 1856. (Fig. 1, A & B). Agrostis airoides Torr, Ann. Lyceum Nat. Hist. New York L151. 1824, non (Poir.) Racpall Vilfa airoides (Torr) Trin. ex Steud., Nomencl. Bot. (ed. 2) 2:766 1841. TYPE: U.S.A. COLORADO: on the branches of the Arkansas, near Rocky Mountains, E. James sn. (HOLOTYPE: NY-327612! IsoTYPE: US-76255 fragm. & photostat ex herb. Torrey!). Sporobolus diffusissimus Buckley, Proc. Acad. Nat. Sci. Philadelphia 14:90. 1862. TYPE: U.S.A. TEXAS: Nestern Texas, 1849, C. Wright 726 (sOTYPE: US-3198038 fragm.!). _ robolus schaffneri Mez, Repert. Spec. Nov. Regni Veg. 17(19-30):295, 1921. Type: MEXICO. SAN UIS POTOSI: Ww Schaffner s.n. GSOTYPE: US-87214 fragm.!) bolus tharpii Hitchc., Proc. Biol. Soc. Wash. 41:161 1928. Type: U.S.A. TEXAS: Padre Island, 4 Sep 1927, B.C. Tharp 4772 (HOLOTYPE: US-1299827!). Densely caespitose perennials. Culms 35-120(-150) cm tall, erect, stout, glabrous below the nodes; base diameter 1-2(-3.5)mm wide, rounded: internodes glabrous. Leaf sheaths 2/3 to about as long as the internodes above, glabrous, shinny, some- times with a few long hairs near the summit, these hairs up to6mm long; ligules 0.1-0.3 mm long; blades (3-)10-45(-60) cm long, (1-)2-5(-6) mm wide, flat to involute, glabrous below and scaberulous to scabrous above; margins mostly smooth to scaberulous. Panicles (LO-)15-45 cm long, 15-25 cm wide, open, dif- fuse, subpyramidal, often included in the uppermost sheath; branches 1.5-13 cm long, ascending to widely spreading 30-90° from culm axis; secondary branches mostly spreading and not floriferous on lower 1/4 to 1/3; pulvini in axils of primary branches glabrous; pedicels 0.5-2 mm long, spreading, glabrous to scabrous. Spikelets 1.3-2.8 mm long, spreading, purplish or greenish:.glumes 0.5-2.4(-2.8) mm long, lanceolate to ovate, membranous, unequal: lower glume 0.5-18 mm long, often appearing without a midvein, the apex acute; upper glume 1.1-2.4(-2.8) mm long, apex acute to obtuse; lemmas |.2-2.5 mm long, ovate, membranous, glabrous, the apex acute; paleas 1.1-2.4 mm long, ovate, membranous, glabrous, the apex acute to obtuse; stamens 3; anthers L.1-1.8 mm — PETERSON ET AL 559 ee an a Pe Tag f iy \ Peet Ni if ‘ La, 1A] i; t z P +/ if, Wy if | eS | Ve ge. Xx { y if airoides. A. Habit. B. Spikelet. S bol iroid bst regis C. Habit. D. Inflorescence Fic. 1 Sp Lat iY rf apical portion. E. Spikelet with stamens. 560 BRIT.ORG/SIDA 21(2) long, yellowish to purplish. Modified caryopses 1-1.4 mm long, ellipsoid, red- dish-brown, striate. 2n = 80, 90, 108, 126. Distribution and habitat.—Sporobolus airoides occurs throughout the arid portions of northwestern North America and in México as far south as Puebla. It is common in dry to sandy gravely flats or slopes usually associated with alkaline soils, occurring with Atriplex canescens (Pursh) Nutt, A. confertifolia (Torr. & Frém.) S. Watson, Larrea tridentata (Sessé & Moc. ex DC.) Coville, Sarcobatus vermiculatus (Hook.) Torr. Distichlis spicata subsp. stricta (Torr.) Thorne, and Ambrosia dumosa (A. Gray) W.W. Payne; 50-2400 m. Flowering June through November. In northeastern México, S.diroides isa halophytic spe- cies forming the alkali zacaton giasiauds where the edaphic effects of a di- verse combination of chloride « d sulfates acc lates to form very saline conditions. Specimens examined. MEXICO. Coahuila: Municipio de Castanos, Paso de San Lazaro, Sierra de la Gavia, 37.6 mi S de Monclova, carretera 57, Peterson et al. 9985 (ANSM, US); Municipio de Cuatrociénegas, Juntoa Nuevo Atalaya, Brigada Ill 4a (COCA); Cuatrociénegas, Brigada III 6 (COCA), km N de la Poza de la Becerra, J.A. Davila s.n. (ANSM); 11 km E de Cuatrociénegas, X. Hernandez 2036 (ANSM), Areas salinas S of Cuatrociénegas, J.S. Marroquin 1351 (ANSM); Cerca de la Poza ‘EL Bonito’, J.S. Marroquin s.n. (MEXU); Dunas yesosas, cerca de la Poza El Bonito, J.S. Marroquin s.n. (ANSM); SE de Cuatrociénegas, Sin collector 6 (COCA), 45 km S of Cuatrociénegas, Peterson et al. 10002 (ANSM, US), Pacers: de la Slenke de San Marcos 24 mi S Cuatrociénegas, Peterson et al. 10008 (ANSM, US); Municif Cepeda, Ejido La Rosa, carretera 40 Saltillo-Torreén, 20 km NE de General Cepeda, S. Vasquez 82 (ANSM); heen de Juarez, Disaat de BIegO 04, ca Martin, P. De la Garza s.n.(ANSM); Municipio de Ocampo, Laguna La Leche le Ocampo rumbo a Sierra Mojada, M.A. Carranza 630 (A NSM Sierra La Enciede: rancho Puerto del Aire, ee pean 780 (A NOM), Sierra del Pino, Ejido Acebuches, Canon La Vaca, M.A. Carranza 967 (ANSM); Ran tal La Rueda, 87 km NE de Ocampo, brecha Ocampo-Boquillas del Car- men, : ee rd s.n. ONaD Sierra El Pino, 9.2 km S of Rancho El Cimarron along the eastern slope, Peterson & Annable 10618 (ANSM, US); 35.5 km NW of Monclova and 11.3 km E of Sacramento on road to Cuatrociénegas, Peterson A al. 8363 (ANSM, US); 4 km S de Laguna del Rey, de la Planta Quimica, Peterson et al. 8371 (ANSM, US); 4 km S of Laguna El Rey Ch emical, Plant, Peterson et al. 8374 (ANSM, US); Laguna La | ee Valdés- Reyna 1330 (ANSM); Rancl tal Santa Teresa de La Rueda, aproximadamente 87 km NE de Ocampo, brecha Ocampo- Boqiilles del Carmen, M. oe s.n.(ANSM); Municipio de Parras, Rancho el Tunal, aproximadamente 25 km ESE de Parras de | a Fuente, A. Rodriguez 1171 (ANSM); Ejido 4 de Marzo, F Roing s.n.(ANSM): 9 km S of Parras on Negras, L. Sta ae 189 (MEXU), seed de Ramos Arizpe, Canada el Diente, Sierra de la oe sine 1300 m., J.A. Villarreal 5183 (ANSM); Municipio de pa ancia 10 km de Sacramento rumbo a Een A. Rodriguez 1229 (ANSM); M ipio de S 53.2 km S of Saltillo on México hwy 54 and 9.6 km E on road to La Ventura, Peterson et al oe (ANSM, sas 47 km S e Saltillo on México hwy 54 to Concepcion del Oro, Peterson et al. 10034 (US); R Los Angeles, 48 km S de Saltillo, carretera 54 Saltillo-Concepci6n ff Oro, Zacatecas, i. Sierra s.n. (ANSM); 2 km N del ejido La Encantada, carretera Saltillo-Concepcion del Oro, Zacatecas, Valdés- Reyna 1511(ANSM); 6 km W de General Cepeda. Carretera a Parrasa orilla del ie Valdés-Reyna 1575 (ANSM); Buenavista, 6 km S de Saltillo por la carretera Saltillo-Zacatecas, Valdés-Reyna 1890 (ANSM); 3 km S de la Ciudad de Saltillo, fraccionamiento Parque de la Canada, Valdés-Reyna 2282 (ANSM); Ejido La Colorada, aproximadamente 15 km S de Saltillo, carretera 54, Saltillo-Concepcion del Oro, Zacatecas, Valdés-Reyna s.n.(ANSM); Buenavista, 7 km S de Saltillo, carretera 54 Saltillo- Q. PETERSON ET Al 561 E ion del Oro, Zacatecas, J.A. Villarreal 1769 (ANSM); km 34 carretera Saltillo-Concepcion del Oro, Tacateeas Entrada al rancho Los Angeles, S. Villarruel sn. (ANSM); Municipio de Torreon, S of Torreon between Jimulco and Juan Eugenio, Peterson & Valdés-Reyna 8478 (US). Nuevo Leon: Municipio de Doctor Arroyo, Ejido Lagunita y Ranchos Nuevos, M.E. Pérez 107 (ANSM); Ejido La Escondida, 5 km N carretera 102 Doctor Arroyo-Galeana, J.A. Villarreal 6518 (ANSM); Municipio de Galeana, along highway 57, about 12 miN of San Roberto, K. Allred 5515.5 (ANSM, NMCR); 5.6 mi E of jtn of hwy 57 on hwy 58 towards Linares, Peterson & Knowles 13296 (US); 13.4 mi E of hwy 57 on hwy 58 at crossing of Rio Potosi, pated : Knowles 13312 (US), Near La Trinidad, (Highway 57), 71 mi S of Saltillo, T. Soderstrom 373 (MEXU, US). Tamaulipas: Municipio de Bustamante, Poblado Bustamante, J.G. Galvdn 30 (COCA), vane de Llera, Ejido Portes Gil J. Barrientos 97 (COCA); Municipio de Tula, pe Francisco Medrano, J. Iribe 126 (COCA); Ejido 5 de Mayo, P. Moya 64 (COCA). KEY TO THE SUBSPECIES OF SPOROBOLUS AIROIDES 1. Sheaths generally glabrous or minutely scabrous on abaxial surface; primary branches of the panicle glabrous, without tufts of hairs in the axils below _____ 1a. S.airoides subsp. airoides . Sheaths pilose i eee surface; primary branches of the panicle with a tuft o hairs in the axils 1b. S. airoides subsp. regis 1b. Sporobolus airoides subsp. regis (IM. Johnst.) Wipff &S.D. oe Sida 16:164. 1994. (Fig. 1, C & D). Sporobolus regis LM. Johnst. J. Arnold Arbor. 24:393. 1943. TYPE: MEXICO. Sern Municipio de Ocampo, salt flat 4 km SE of Laguna del Rey, abundant, 1040 m, 18 Sep 1942, R. Stewart 2653 (HOLOTYPE: GH, ISOTYPE: US-90729 fragm_l). Rhizomes elongate, knotty with internodes 10-12 mm long, 3-4 mm in diam- eter. Culms 80-130 cm tall, erect, densely leafy. Leaf sheaths and collars pilose on abaxial surface with hairs 2-4 mm long, glabrous with age; ligules 0.1-0.3 mm long, fimbriate or densely ciliolate; blades 10-30 cm long, 3-4 mm wide, usually loosely involute. Panicles 30-40 cm long, 10-15 cm wide, open, diffuse, subpyramidal, exserted or partly included below, the branches with a tuft of trichomes in the axils; glumes 1-18 mm long; lemmas 1.9-2.5 mm long; paleas as long as the lemma. Distribution and habitat —Sporoboluls airoides subsp. regis is an endemic species known only from Coahuila, México on saline flats of Laguna del Rey, the type locality, and Salinas del Rey. Sporobolus airoides subsp. regis is distin- guished from S. airoides subsp. airoides by its pubescent sheaths and tuft of trichomes in the axils of the panicle branches. Wipff and Jones (1994) consid- ered these characters not significant to warrant specific rank, however, they felt that they are significant to warrant infraspecific recognition. Specimens examined. MEXICO. Coahuila: Municipio de Ocampo, 12 km S of Salinas del Rey, Henrickson 14152 (TEX). 2. Sporobolus atrovirens (Kunth) Kunth, Revis. Gramin. 1:68. 1829. (Fig. 2, A- C). Vilfa atrovirens Kunth, Nov. Gen. Sp. 1:138. 1816. a atrovirens (Kunth) Roem. & Schult., Syst. Veg. 2:361. 1817. TYPE: MEXICO. DisTRITO FEDERA valle Mexicana prope El Penon del Marques, Humboldt and Bonpland s.n. (HOLOTYPE: P- Sonal ISOTYPE: BM!) Small caespitose perennials, sometimes appearing annual. Culms 7-30 cm tall, 562 BRIT.ORG/SIDA 21(2) se aa Fic. 2. 5; boll i A Habit. B. Ligule.C. Spil | itl lified yopsis. 5 bol huilensis.D. Habit. E Coetl «lee c.tal ond ° t Liqule. F. Inflorescences. G g PETERSON ET AL., 563 erect, with 1-2 nodes above the base; base diameter 0.5-0.8 mm. Leaf sheaths short, ciliolate; collar glabrous or with few hairs; ligules 0.1-0.2 mm long: blades 3-15cm long, 2-4 mm wide, lanceolate, flat, glabrous, borne near base. Panicles 5-20 cm long, 3-10 cm wide, oblong to ovoid, completely exserted; primary branches spreading 70-120° from culm axis, naked below on lower 1/4-1/3, the lowermost solitary or paired at the nodes; pulvini in the axils of the pri- mary branches glabrous, yellowish; pedicels 0.5-L6 mm long. Spikelets 0.7-1.2 mm long, greenish or purplish to plumbeous; glumes without nerves, apex ob- tuse; lower glumes 0.3-0.5 mm long, the apex obtuse or erose; upper glumes 0.4-0.7 mm long, the apex obtuse to acute; lemmas 0.7-1.2 mm long, the apex acute to obtuse; paleas 0.9-1.2 mm long, the apex obtuse; stamens | or 2; anthers 0.4-0.8 mm long, purplish. Modified caryopses 0.5-0.8 mm long, pyriform or quadroid, somewhat laterally flattened, light brownish. Distribution and habitat.—Sporobolus atrovirens is a rare species for north- eastern México usually found occupying xeric habitats in gypsum soils at 1O- 1500 m. It has been reported in México in Aguascalientes, Baja California, Coahuila, Durango, Hidalgo, Guanajuato, Jalisco, México, Oaxaca, Puebla, San Luis Potosi, Tamaulipas, Tlaxcala, Veracruz, Yucatan, and Zacatecas. In the Sierra La Lagunita at 1450 m, S. atrovirens (Peterson et al. 16690) was found growing on slopes with a dominant vegetation of Pinus pseudostrobus LindL., Juniperus flaccida Schltdl. Juglans, Agave, and Muhlenbergia dubia E. Fourn. Other associ- ated native and adventive species from this site include: Chloris, Tragus berteronianus Schult. Urochloa meziana (Hitche.) Morrone & Zuloaga, Paspalum dilatatum Poir, Erioneuron avenaceum (Kunth) Tateoka, Eragrostis intermedia Hitche., Oplismenus hirtellus (L.) P. Beauv, Bromus anomalus Rupr. ex E. Fourn., Schizachyrium, and Panicum bulbosum Kunth. This latter site was perhaps un- usual for this species since it was in a heavily wooded and shaded environment. Specimens examined. MEXICO. Coahuila: oe Saltillo, Los Cerritos, near ae Oct 1912, i) Lyonnet sn. (TEX). Nuevo Leon: Municipio de Monterrey, 5 mi SW of Hidal 15 airline mi NW of Monterrey), B.L. Turner & J. Cr enna 6285(TEX). Sierra 2 Lagunita, 9.5 mi SE of Aramberri on road towards Agua Fria, 24° 03' 37.9" N, 99° 45' 35.8" W, 19 Sep 2002, Peterson et al. 16690 (ANSM, US). Tamaulipas: Municipio de Jaumave, Altas Cumbres km 160 carretera Victoria- Jaumave, 950 m, M. Martinez 727 (ANSM, TEX, UAT); Rio Guayalejo, 2 km E del ejido San Vicente rumbo a Jaumave, LO m, A. Mora 5358 (UAT); Municipio de Tula, 30 km al SW de Tula, cerca del limite de San Luis Potosi and Tamaulipas, Gonzalez Medrano F. 4422 (TEX). aa 3. opsroelus buckleyi Vasey, Bull. Torrey Bot. Club 10:128. 1883. (Fig. 3. A & B). U.S.A. TEXAS: 1883, S.B. Buckley s.n. GSOTYPE: US-556873)). Caespitose perennials. Culms 40-100 cm tall, erect, glabrous, base flattened, internodes glabrous; base diameter 0.7-3 mm wide. Leaf sheaths 4/5 as long as the internodes to longer than the internodes above, glabrous, margins occasion- ally hairy near summit, sometimes with a line of hairs from one margin to the next, the hairs up to L.2 mm long, lower sheaths strongly laterally compressed pa 564 BRIT.ORG/SIDA 21(2) DAN Cathy Fequate. XSF Fic. 3. Sporobolus buckleyi. A. Habit. B. Spikelet with modified caryopsis. Sporobolus spiciformis. €. Habit. D. Ligule. E Spikelet with stamens. PETERSON ET AL., 565 and keeled; ligules 0.2-0.4 mm long; blades 12-35 cm long, 4-12 mm wide, flat, glabrous below and scaberulous above; margins smooth to scaberulous. Panicles 15-50 cm long, 7-22(-30) cm wide, open, diffuse, ovate; primary branches as- cending to widely spreading mostly 2-17 cm long, not floriferous on lower 1/4 -1/2; secondary branches appressed to loosely spreading; pulvini in axils of rimary branches glabrous; pedicels 0.2-1.2 mm long, mostly apressed, scaberulous. Spikelets 1-2 mm long, purplish or brownish; glumes 0.5-1.8 mm long, narrow lanceolate to lanceolate, membranous, unequal, prominently keeled, scaberulous along the distal portion of the keel; lower glumes 0.6-lmm long, the apex acuminate to acute; upper glumes 1.1-1.8 mm long, the apex acute; lemmas 1.2-2 mm long, lanceolate, membranous, glabrous, the apex acute; paleas 1.2-2 mm long, ovate, membranous, often splitting in two between the veins at maturity, glabrous, the apex acute; stamens 3; anthers 0.2-0.4 mm long, purplish. Modified caryopses 0.6-1 mm long, ovoid, slightly flattened, reddish brown. 2n = 40. Distribution and habitat.—In northeastern México, S. buckleyi is a com- mon species of the Tamaulipan desert scrub on loamy soils near margins of woods sometimes in partial sunlight associated with Acacia, Quercus, and Prosopis thickets and thorn scrub; 40-700 m. Flowering April to November. Lane pom MEXICO. Nuevo Leon: Montemorelos, Ojo de Agua. Matorral xerofito. M. A. ti 82300 ), Monterrey. C. G. Pringle 2520 (MEXU); Municipio de Allende, 6.1 km S de Allende on ee 85 -o Montemorees oll higawayc on pt Lice near asociation de abaeeies oan storage), just N of Canoas, Petersor g rral reve S3 Rodrigues 102 (ANSM); Si de Picachos, Rancho El Gall oan Villarreal 8014 (AN uadalupe, Guadalupe, E. Cantt s.n. (ANSM); Municipio de Linares, Rancho El ae ce ee en la carretera Linares-San Roberto, km 12, M. Castillo 92 (COCA), Los Fresnos, J. sie s.n. (ANSM), Municipio - whey Facultad de Agronomia, Universidad Aut6noma de Nuevo n, km 17, M. Castillo 26 (CO! ,A lasalida del eDHEDO de Zacatecas rumbo eer a Fria en las orillas del rio etn pyaa regu 80 (COCA); M S atarina, Canon El care Sierra Meet Ou ental, Dene 20 km 5 de Monterey J. Valdés-Reyna 1969 (AN UAT): M S go, 4km y de Los Cavazos, I. Cabral 128 (ANSM), Rio San Juan, I. se (ANSM): Caen Monterrey A la carretera a Zuazua, PA. Garcia 1864 (COCA). Tamaulipas: Municipio de Abasolo, a La Esperanza.) Iribe 149 (COCA), peeal de Aldama, La Muralla, R. Carranco 127 (COCA), Rancho EI Rosario, M. Cervera 211 (COCA); Ejido El Nacimiento, propieda privada, R. Diaz 319 (UAT), Piedras ee Galvan 235 (COCA); Municipio de Antiguo Morelos, Ejido Las Flores, J. Ramos 12 (COCA), Ejido El Refugio J. Ramos 122 (COCA), Municipio de Casas, Rancho José Roberto, R. Carranco 278 (COCA); Ejido Lazaro Cardenas. A orilla de la via de FEC.C., J. Ramos 207 (COCA), i aniciplo. de Comes Farias, Eudok el Nacimiento, M poe 310 (ANSM); Ejido Sabinas, M. Crespo 422 (ANS J. Iribe 136 (COCA); 15 mi W of Gonzalez pewara Mante, M.C. Johnston 49.29 (MEXU); ce Re Ortiz de Dominguez, P. Moya 36 (COCA); Municipio de Guémez, Rio Corona, J. [ribe 298 (COC i ee de Hidalgo, Ejido Nicolas Bravo, a fate 100 (COCA), Ejido El oe . Iribe 2 jido La Colombina, J. Iribe 262(C ancho La Purisima, J. Iribe 303 (CO ); Municipio : ae io Los Mimbres, 20 mi W fe Rio Purificacion, Peterson & Valdés- me 15939 (US): Municipio de Jiménez, Ejido Sor Juana Inés de la Cruz, R. Carranco 410 (COCA); Ejido Sor Juana Inés de la Cruz, R. Carranco 411 (COCA), 7 mi S of Santander Jiménez, M.C Johnston 4393-A, 4393-B (TEX- 566 BRIT.ORG/SIDA 21(2) LL, MEXU); Municipio de Llera, Rancho el Lloradero, A. Brito, s.n.(UAT); km 152 carretera Llera-Mante, RK. on ranco 272(C Oc A); km 157 carretera Llera-Mante, La Mina, R. Carranco 272a (COCA); Camino al Ejido Lucio Blanco, J.F lribe 285 (COCA); Municipio de Mainero, Ejido Boreal Real, M. Cervera 343 (COCA); Municipio de Padilla, km 65 carretera Victoria-Matamoros, A.. Brito 63 (COCA), Municipio de San Carlos, Ejido Puerto Rico, R. Carranco. 295 (COCA); Cerro del Diente, R. Carranco 47 (COCA): Cerro del Diente, J. A. Franco 92 (COCA); Cerro del Diente, J. Iribe 94 (COCA); Municipio de San Fernando, La Joya, R. Carranco 101 (COCA); Municipio de Soto la Marina, Rancho San Alfonso, ] Barrientos 6(COCA), San José de las Rusias, M. Cervera 84 (COCA). Tramo San José de las Rusias-5 ¢ Mayo, J.G. Galvdn 161 (COCA), Ejido Verde Chico, J. Tribe 337 (COCA); Los Eslabones, J.E. Lopez 44 (COCA); Municipio de Tampico, Laguna del Chairel, M.G. Torres s.n. (COC A): Municipio de Tula, Pu- erto de La Virgen, A. Brito, 50 (COCA); Municipio de Victoria, Lil P , M. Cervera 305 (COCA); Ejido Santa Ana, M. Cisneros 31, 32 (COCA); Municipio de Villagran, L oe M. Cervera 160 (COCA). fay) 4. Stes coahuilensis Valdés-Reyna, Phytologia 41:81. 1978. (Fig. 2, D-G). FE: MEXICO. COAHUILA: 58 air mi SW of Cuatrocienegas, near 26° 17'N, 102° 40' W, 815 m, 15 oon 1976, J. Henrickson & B. Prigge 15363 (HOLOTYPE: LLI lsoTYPES: CSLA, MEXU)). Annuals. Culms 15-60 cm tall, ascending, glabrous. Leaf sheaths shorter than the internodes, glabrous; ligules 0.5-1 mm long, ciliate, the hairs 0.5-1 mm long; blades 4-12 cm long, 1.5-6 mm wide, flat, spreading, evenly distributed, sparsely ciliate-pustulate above. Panicles 6-22 cm long, (-1)5-13 cm wide, open, some- times contracted, branches ending in a spikelet, lowest branches whorled, in verticels of 7-20; pedicels (2)3-6(-8) mm long, widely spreading, capillary. Spikelets 1.1-1.5 mm long; glumes thin, the apex acute; lower glumes ca. 0.5 mm long; upper glumes 1.4-1.5 mm long; lemmas 1.3-1.4 mm long, the apex acute; paleas |-1.3 mm long, hyaline. Modified caryopses 0.6-0.9 mm long, oblong, light brown; embryo 0.2-0.4 mm long. Distribution and habitat.—In México, S. coahuilensis is known only from Central Coahuila, near Las Delicias and Cuatrocienegas. This species has re- cently been found in Texas (per. comm. B.L. Turner). Comments.—Sporobolus codhuilensis appears closely related to S. pyramidatus, a widespread species, and can be separated from the latter spe- cies by having long capillary pedicels and wider panicles. Specimens examined. MEXICO. Coahuila: Municipio de Cuatrociénegas, SE de Cuatrociénegas, Brigada IIL 1 (COCA); cerca 45 km SW of Cuatrociénegas, 760 m., Peterson et al. 10000 (ANSM, US). 5. Sporobolus compositus (Poir.) a var. Compose Cire. Div. Agrostol. US.D.A. 35:6. 1901. (Fig. 4, A-C). A -omposita Poir, ue ea 254. 1810. Vilfa composita (Poir.) P Beauv, Ess. Agrostogr. 16, ‘147, 181. ig. Muhlenbergia composita Trin. ex jn Kunth, Enum. PI. 1:229. 1833, nom. inval. TYPE: U.S.A. “CAROLINA”: L.A.G. Bosc s.n. (HOLOTYPE: not known). Agrostis aspera Michx., Fl. Bor-Amer. 1:52. 1803, nom. illeg. non Weber. Vilfa aspera P. Beauv. Ess. Agrostogr. 16, 147, 181. 1812. Muhlenbergia aspera (P. ee acta Gram. unifl. sesquifl. 193. 1824. sean asper (P. Beauv.) Kunth, Reévis. Gram. 1:68. 182¢ 2 U.S.A. ILLINOIS: A. X sn. (HOLOTYPE: P-MICH; lsoryPe: US-76431 fragm. & see 0 ex a Agrostis nae Torr, Fl.N. Middle United States 1:90. 1823. Vilfa longifolia Torr) Torr, N. Amer. 567 PETERSON ET AL., datus. D. Habit. F. Spikelet with A. Habit. B. Inflorescence. C Spikelet 5} iy Fic. 4. $ modified caryopsis. 568 BRIT.ORG/SIDA 21(2) Gram. 1:4. 1834. Sporobolus longifolius (Torr) Alph. Wood, Class-book 1861. TyPE: U.S.A. PENNSYLVANIA: (HOLOTYPE: not known). Vilfa hookeri Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 6,4C1-2):106. 1840. peas asper var. hookeri (Trin.) Vasey, Descr. Cat. Grass. ee 43, 1885. Type: US.A. TEXAS: T Drummond 306 (isotypes: US-997649 fragm.!, US-997663 y, Proc. Acad. Nat. Sci. Philadelphia 14:95 1862. nom. illeg a non H Hook er “f TYPE USA. TEX we middle Texas, S.B. Buckley s.n. (LECTOTYPE: PH, designated by Hitchcock, Bot. (ed. 1861) 1861:775. Man. Grass. U.S. 958. 1935, but without citing a specific sheet in a specific herbarium: ISOLECTOTYPE: US frag. & photo ex PH)). Sporobolus pilosus Vasey, Bot. Gaz. 16(1):26. 1891 = obol 1 pi asey) Hitchce., Proc. Biol. Soc. Wash. 41161. 1928. Sporobolt “var. pits Vases) Hitchc, N. Amer. Fl. 17(7):488. 1928. TyPE: U.S.A, KANSAS: ee 1890, B.B. Smyth 217 (HOLOTYPE: US-556890!: ISOTYPES: US-746585!, US-75038). Caespitose to solitary-stemmed perennials. Culms (20-)30-130(-150) em tall, glabrous below the nodes, internodes glabrous; base diameter 0.7-5 mm wide. Leaf sheaths 3/4 to 1 1/2 as long as the internodes above; upper (terminal) sheaths 1-6 mm wide, mostly glabrous, summit with a few hairs up to 3 mm long; ligules 0.1-0.5 mm long; blades 5-70 cm long, 1.5-10 mm wide, flat to folded or involute, glabrous below and glabrous to scaberulous above, sometimes pi- lose just above the ligule, rarely pilose throughout; margins mostly glabrous. Panicles 5-30 cm long, 0.4-16(-3) mm wide, terminal and axillary, narrow, sometimes densely spike-like, the base usually included in the uppermost sheath; primary branches 0.4-6 cm long, appressed, usually floriferous to base: secondary branches appressed; pedicels 0.3-3.5 mm long, appressed, glabrous to scaberulous. Spikelets +-6 (-10) mm long, stramenious to purplish-tinged: glumes (1.2-)2-5(-6) mm long, lanceolate, membranous to chartaceous, usu- ally with a greenish midvein that is scrabous towards the apex, subequal; lower glumes (1.2-)2-4 mm long, the apex acute; upper glumes (2-)2.5-5(-6) mm long, the apex acute; lemmas (2.2-)3-6(-10) mm long, lanceolate, membranous to chartaceous, rarely 2- or 3-veined, glabrous to minutely pubescent or long, ovate to lanceolate, membranous to chartaceous, the apex acute to obtuse; stamens 3: anthers 0.2-3.2 mm long, yellow to orangish. Modified caryopses 1-3 mm long, ellipsoid and laterally flattened, often striate, reddish brown. 2n = 54, 88, 108. Distribution and habitat.—A rare species in northeastern México, only known from a single collection initially determined by EW. Gould. Roadsides, railroad right of ways, beaches, cedar glades, pine woods, live oak-pine forests, prairies and many other partially disturbed, semi-open sites; 0-1600 m. Flow- ering August to November. 5 Specimen examined. MEXICO. COAHUILA. Municipio de Muz J. A. Santos s.n.(ANSM). iz, La Boquilla, Rancho La Eneantada, 6. Sporobolus contractus Hitchc., Amer. J. Bot. 2:303. 1915. (Fig. 5, AD). Sporobolus cryptandrus var. strictus Scribn., Bull. Torrey Bot. Club 9:103. 1882. Sporobolus strictus (Scribn.) Merr, Cire. Div. Agrostol. U.S.D.A. 32:6. 1901, non Franchet. TyPr: U.S.A. ARIZONA: Banks of Rillita [Rillito] Brook, near Camp Lowell, 15 Jun 1881, C.G. Pringle sn. GSOTYPE: US-825284!), PETERSON ET AL., imm AZ es 5 LAA rr as ae 7 iS ae —_f > T 4 T 74° 76° 78° soe Fic. 5. Distribution map of Eriocaulon species. PUNEKAR ET AL., NEW SPECIES OF ERIOCAULON FROM INDIA 633 Acaulescent rosulate herbs, to 16.5cm nig: Roots fibrous. Leaves rosulate, 7-11, linear, broad at the base, tapering -acuminate, shorter than sheaths up to L7 cm long and 0.25 cm broad, glabrous. Peduncles 1-5, twisted, glabrous, up to 16.5cm long, 6-ribbed. Sheaths up to 2.7 cm long, glabrous; limb lanceolate, entire, acute. Heads ovoid to quadrangular, 5 mm across, white. Receptacles de- pressed globose, pilose. Involucral bracts spreading, obovate, rounded and notched at apex, straw colored with blackish tinge, papery, glabrous, 1.2 x 1 mm. Floral bracts obl blackish towards apex, hyaline towards base, 1.8 x 1mm. Male flowers: pedicels 0.4 mm long: sepals obovate, connate into a spathe of 1.2mm long and 0.8mm across, 3-lobed, lobes truncate to obtuse and dorsally hoary at apex, hyaline with slight blackish tinge; stipe of corolla 0.5-0.7 mm long; petals 3, unequal, laterals linear, minute, middle oblong to narrowly elliptic, obtuse at apex, | x 0.3 mm, all densely hoary inside, glabrous outside, gland dotted; stamens 6, anther lobes black. Female flowers: pedicels 2-3 mm long; sepals 2, free, oblan- ceolate, conduplicate, keeled, straw colored with blackish tinge, dorsally hoary and obtuse-rounded at apex, 1.5mm long; petals 3, free, hyaline, unequal, later- als smaller, 1 x 0.5 mm, hairy at apex, middle larger, 1.5 x 0.2 mm, ventrally hoary in upper half, dorsally sparsely hairy at apex, all oblanceolate, obtuse at apex, gland dotted, stipitate between sepals and petals; ovary stipitate, ovoid, 0.2 x 0.3mm; style trifid. Seeds ellipsoid, acute, 0.4 x 0.25 mm, yellow; cells of seed coat transversely elongated, aligned in vertical rows, appendages 2-4 from transverse radial walls, setiform, dilated at apex. Eriocaulon konkanense is similar to E. odoratum Dalzell but differs from the latter in having white heads, involucral bracts notched at 2 or 3 places at apex, female sepals 2, obtuse to rounded at apex, seeds yellow, with 2-4 append- ages from transverse walls and none from vertical walls. Distribution.—This species is so far restricted to the coastal lateritic pla- teau near MIDC area, Airport road, Ratnagiri district, Maharashtra (Fig. 5). Habitat and Ecology—Common on coastal lateritic plateau in association with Eriocaulon parviflorum, Eriocaulon richardianum, Exacum pumilum, Fimbristylis sp., Pentanema indicum, Rhamphicarpa longiflora, Utricularia reticulata, etc. Flowering & Fruiting —August-Novem ber. Etymology—The specific epithet of the new species is based on the type locality viz., Konkan Coast of Maharashtra State. eolate, cuneate, acute toacuminate, dorsally hoary and KEY TO NEW SPECIES AND ALLIED SPECIES 1. Sepals of male flowers free. Heads spinulate; involucral bracts usually acuminate; sepals of female flowers longer than floral bracts E. anshiense 2. Heads not spinulate; involucral bracts usually obtuse or subacute; sepals of fe- male flowers as long as floral bracts E. eurypeplon 634 BRIT.ORG/SIDA 21(2) 1. Sepals of male flowers connate into a spathe. 3. Petals of male flowers subequal: anther lobes white E. talbotii 3. Petals of male flowers unequal; anther lobes black. 4. Sepals of female flowers 2 E. konkanense 4. Sepals of female flowers 3. 5. Petals of female fl b |, glandul Is with appendages ___ E.odoratum 5. Petals of female flowers unequal, eglandular; seeds without any appendages E. kanarense ACKNOWLEDGMENTS The authors are grateful to the Director, Botanical Survey of India, Kolkata for providing facilities and to PSN. Rao, Joint Director, Botanical Survey of India, Western Circle, Pune for constant encouragement. One of us (PL) is also thank- ful to the Keeper, Royal Botanic Gardens, Kew for facilities. Our thanks are also due to N.P. Balakrishnan, Ex-Joint Director, Botanical Survey of India, Coimbatore for his valuable suggestions regarding the species. Help rendered by Melanie Thomas (Wilmot-Dear), Royal Botanic Gardens, Kew for Latin di- agnosis and to Lourdes Rico, Royal Botanic Gardens, Kew for the Spanish ab- stract is gratefully acknowledged. We also wish to thank the Karnataka Forest Department for various help rendered by them and also to Mahesh Shindikar, Ecology Laboratory, Dept. of Botany, University of Pune, who prepared the dis- tribution map. REFERENCES Ansari, R. and N.P. BALAKRISHNAN. 1994. The family Eriocaulaceae in India. Bishen Singh Mahendra Pal Singh, Dehra Dun, India. Bote, PV.and M.R. AtmeiDa. 1986 [1987]. Material for the flora of Mahabaleshwar-7. J. Bom- bay Nat. Hist. Soc. 83:570-602. Das Das, S.and N.P. Sinch. 2001. Taxonomic studies on Eriocaulaceae in Karnataka State. J. Econ. Taxon. Bot. 25:449-484, Gaikwao, S.P.and S.R. Yapav. 2001. Eriocaulaceae in Maharashtra In:T. Pullaiah, ed. Biodiversity of India 1:256-341. Gatkwan, S.P.,M.M. Saroesal, and S.R. Yapav.2002.A new species of Eriocaulon L.(Eriocaulaceae) from Maharasntra, India. Rheedea 12:133-136. Gaikwao, S.P, M.M. Sarvesai, U.S. Yapav, and S.R. Yapav. 2003. A new species of Eriocaulon L. (Eriocaulaceae) from Karnataka, India. Rheedea (under publication). KARTHIKEYAN, S., S.K. Jain, M.P. Nayar, and M. Sanuappa. 1989. Florae Indicae Enumeratio: Monocotyledonae. Flora of India Series 4. Botanical Survey of India. KHANNA, KK., V. Mupaat, and A. Kumar. 2000. A new species of Eriocaulon (Eriocaulaceae) from Madhya Pradesh, India. Nord. J. Bot. 20:41 3-414. KosHy, K.C. and P. PusHpancanan. 1993. On the occurrence of Eriocaulon minimum Lam. in India. Rheedea 3:117-119. PUNEKAR ET AL., NEW SPECIES OF ERIOCAULON FROM INDIA 635 Massertey, D.J. 1997. The plant book. ed.2. (Reprinted, 2000). Cambridge University Press. MyrTHona, S., Y. Kumar, and R.R. Rao. 1983 [1984]. Eriocaulon echinulatum Mart. (Eriocaulaceae)—a new record for India. J. Bombay Nat. Hist. Soc. 80:667-668. Praveer, A.K.and C.N. Suni. 2003. Eriocaulon ansarii Pradeep & Sunil (Eriocaulaceae), a new species from India. Candollea 58:321-323. PUNEKAR, S.A., P. LAKSHMINARASIMHAN, and M.K.V. Rao. 2002 [2003]. Eriocaulon balakrishnanii (Eriocaulaceae),a new species from Western Ghats of India. Nord. J. Bot. 22:609-611. PUNEKAR, S.A. and P. LAKSHMINARASIMHAN. 2002. Eriocaulon peninsulare (Eriocaulaceae), a new species from India. Nord. J. Bot. (under publication). Punekar, S.A.,N.V. Macpure, and P. LaksHMINARASIMHAN. 2003, Five new species of Eriocaulon L. (Eriocaulaceae) from Western Ghats of India. Rneedea (In press). Sreedevi, B.and M.S. BiINoJ Kumar. 1999. Eriocaulon melanocephalum Kunth (Eriocaulaceae)— a new record for India. J. Econ. Taxon. Bot. 23:742-743. Yaoay, S.R.,S.P.Gaikwap, and M.M. Saroesal. 1998.A new species of Eriocaulon L.(Eriocaulaceae) from India. Rheedea 8:145-147. BRIT.ORG/SIDA 21(2) BOOK NOTICES Ropert J. BLAck and Epwarpb FE. GILman. 2004. Landscape Plants for the Gulf and South Atlantic Coasts. (ISBN 0-8130-2722-5, pbk.). The University Press of Florida, 15 NW 15" Street, Gainesville, FL 32611-2079, U.S.A. (Orders: l- 800-226-3822, fax: 352-392-7302, www.upf.com). $24.95, 240 pp., 21 b/w drawings, 513 color photos, Ll tables, hardiness zone map, 7" x LO". — Publisher Comments: “Robert J. Black and Edward F Gilman tell gardeners how to conquer these hurdles and establish aesthetic and functional plants that can adapt toa coastal site. Beginning with basic gardening precepts, they present step-by-step information on specific plants, planting, best landscape uses, cultural needs, and plant maintenance. They address watering, fertilization, prun- ing, and other care concerns of salt tolerable plan “Combined with colorful photographs of more than 400 plants, Black and Gilman’s expert advice will be indispensable for all southern coastal homeowners, landscape architects, landscape personnel, retail nursery personnel, extension agents, designers and installers, landscape Maintenance and for inland gardeners who face problems that result from salt water intrusion. Chapter 6 is the Plant Selection Guide listing the plants alphabetical by genus. Plants are listed in four categories: 1) Salt-tolerant trees, 2) Salt-tolerant shrubs, 3) Salt-tolerant vines and ground covers, and 4) Salt-tolerant palms. Each entry includes a color photograph of the plant Curt VAN Duzer. 2004. Floating Islands A Global Bibliography. (ISBN 0-9755424- 0-0, hbk.) Cantor Press, 12117 Winton Way, Los Altos Hills, CA 94024-6431, USA. (Orders: 1-800-247-6553, WWww.cantor press.com). $44.95, 428 PP. yx O" Publisher Comments: “This book is a oe treasury of information about one of nature’s marvels: Sou ish ance ae eo ay contains more than eye books and articles in twenty eferenced, and there are both thematic ae penne aphic igaies All aspects of floating islands are sees oe oe of float- ing islands, the causes of their buoyancy, their role in the ecology of lakes and wetlands, their flora and fauna, their role in the dispersal of plants and animals, and methods for controlling and manag- ing them. Works are also cited on artificial floating islands uoed for agriculture, human habitation, ality; and flox islands in literature, myth, and leg- wildlife habitat, and improvement of water qué end. The book includes the text and an English nee with detailed notes, of G.C. Munz’s rare 1711 thesis on floating islands, Exercitatio academica de insulis natantibus, as well as photographs of several floating islands.” SIDA 21(2): 636. 2004 NOMENCLATURE AND TYPIFICATION IN THE GENUS USNEA (LICHENIZED ASCOMYCETES)— IV. USNEA STUPPEA & USNEA SUBSTERILIS James C. Lendemer Isabelle |. Tavares Bota Ce PORIIret University Herbarium The Academy of Natural Sciences of Philadelphia University of California 1900 Benjamin Franklin Pky. 1001 VL B Philadelphia, Pennsylvania 19103, U.S.A. Berkeley, California 94720-2465, U.S.A. lendemer@acnatsci.org ABSTRACT This, the fourth part of our series discussing typification and nomenclature in the lichenized asco- ] ] ; H C4] hay } (Day \RA ] T aT ] ] o It / sag | om and U. substerilis Motyka. Both names are lectotypified and the differences between the two taxa are discussed. RESUMEN ] ] ee ]; L r 1 i } 415 Tye ; ] on ey ] +p ] Esta, c la tipificacion y | genero de ai i | (Paes } TT] L [ cr Motyka y U. substerilis Motyka. Ambos nombres son lectotipificados y se discuten las diferencias entre los dos taxa. INTRODUCTION Some researchers may consider our approach to the typification of Usnea spe- cies overly cautious and technical; however, we would like to stress that the taxo- nomic study of Usnea has been (and still is) considered extremely difficult and complicated. Thus it would seem logical that the first step towards a clear and thorough revision of the genus would be a careful, precise review of the appli- cation of the published names with regard to their types. Unfortunately, many species of Usnea have not been properly typified, and often recent lectotypifications have not taken into account the fact that Motyka’s monograph includes the lectotypifications of numerous taxa. It should also be remembered that Motyka’s treatment remains the only complete revision of the genus to date. The two species treated here belong toa series of closely related taxa that remains much confused and poorly understood. The first species, U. stuppea (Rasanen) Motyka, has been placed in synonymy with U. substerilis Motyka by Halonen et al. 1998). Here, we reject the recent lectotypification of U. stuppea by Halonen et al. (1998) because of an earlier typification by Motyka (1936). In order to clarify the typification (and taxonomy) of U. stuppea we also exam- ined the type material of U. substerilis. As a result we have concluded that the — SIDA 21(2): 637-642. 2004 638 BRIT.ORG/SIDA 21(2) synonymy of U. stuppea with U. substerilis should be reconsidered. To clarify the application of the name U. substerilis we have also chosen a single thallus as the lectotype from among those lectotypified by Clerc (1987). I. Usnea stuppea (Rasdnen) Motyka When Rasanen (1933) described U. comosa var. stuppea he cited only a single locality with a short diagnosis!. Later, when Motyka (1936) treated the taxon in his monograph he elevated Rasanen’s epithet to the specific rank and lectotypified the name ona specimen in the Rasanen herbarium. Unfortunately, there are three packets in the Rasanen herbarium with the same label data. Two of these were labeled by Rasanen himself (the packet numbered “1” was selected by Halonen et al. (1998) as the lectotype) and one is a specimen of Gyelnik’s Lichenotheca Exsiccati 17. Only one of these packets retains an annotation by Motyka and thus this is the only packet we can conclusively state was reviewed by him. Because we consider Motyka’s (1936) use of the term “type” to be effec- tive lectotypification we consider the packet annotated by Motyka to be the lectotype. Halonen et al. (1998) selected as the lectotype the packet labeled by Rasanen and not annotated by Motyka, apparently because they were not aware of Motyka’s previous lectotypification. The lectotypification of Halonen et al. (1998) thus had no standing since it was predated by Motyka’s lectotypification. The lectotype selected by Motyka consists of two thalli (marked “A” and “C”) mounted on a card with one thallus (marked “B”) annotated by Motyka as a different taxon. The thalli marked “A” and “C” agree both with Rasanen’s scant original description and Motyka’s (1936) later description. The thallus marked “B” contains usnic and norstictic acids (.M. Brodo, annotation) and was given the manuscript name Usnea lapponica var. americana by Motyka. Thus, in or- der to clarify Motyka’s lectotypification and the application of this name we select the thallus marked “C” as the “second-step” lectotype (Greuter et al. 2000, Art. 9.14, Ex.6 Usnea stuppea (Rasdnen) Motyka, Lich. gen. Usnea 1:262. 1936, (Figs. 1-3). Usnea comosa var, stuppea Rasanen, Ann. Missouri Bot. Gard. 20:9. 1933, Type: CANADA. BRITISH COLUMBIA: Hazelton, on Picea abot PP 1931, Kujala s.n. (LECTOTYPE, here designated: H (Rasanen Herbarium; packet marked “2,” thallus on right marked “C”) The type collection of U. stuppea is a mixture of more than one taxon; we have made an effort to examine some of the duplicates distributed in Gyelnik’s Lichenotheca Exsiccati. This examination revealed that some duplicates are a mixture of U. lapponica and U. stuppea while others include other taxa. Like- wise the duplicate card in the Rasanen herbarium (packet marked “1”) that was cited by Halonen et al. (1998) is also a mixture of U. lapponica and U. stuppea. 'Rasdnen (1933: 9)."Thallus erectus aut suberectus, brevior fruticulosus, 5-7 cm. longus, laevigatus vel leviter verrucosus, sorediosus, pallido | i Medulla laxa, stuppea, K- AND U. SUBSTERILIS 639 Fics. 1-3. Usnea stuppea. Fig. 1.1 1, thalli ked A and C=U. stuppea, thall ked B = U. Japponica. Fig. 4 ee Pe | £1 L l = | | het Alat, L 7 nN 41 I . g ft 7 Fr Sea | bk hal +)\Eia 2.0 TAL] hall Webel 1 4 r i La t ? J lacking isidiomorphs. Scale bar = 1 cm. The duplicate of Lichenotheca Exsiccati 17 in Rasanen’s herbarium consists of one large thallus of U. stuppea. Il. Usnea substerilis Motyka When Motyka (1930) first described U. substerilis he did not designate a type specimen. Later, however, he selected an exsiccatum of Arnold Lichenes Exsiccati 1538b in W as the lectotype (Motyka, 1936). While treating some of the species of the U. fragilescens-group, Clerc (1987) also selected part of an exsiccatum of Arnold Lichenes Exsiccati 1538b in W as the lectotype noting that it was a mixture of several taxa. While reviewing the status of U. stuppea we also attempted to confirm the typification (and taxonomy) of U. substerilis. A loan of the type material from W revealed that the packet selected by Clerc bore no annotation by Motyka, a situation similar to that of U. stuppea. This 640 BRIT.ORG/SIDA 21(2) case differs from U. stuppea, however, because no specimen matching the data given for the lectotype by Motyka (1936) with Motyka’s annotation was located in W. Thus, because no specimen annotated by Motyka could be found his lectotypification must be superseded by that of Clerc (1987). In order to avoid any doubt as to the application of the name U. substerilis we have thus chosen to further clarify the lectotypification of Clerc (1987) and select a single thallus from among the thalli selected as the lectotype by Clerc. one oo Motyka, oe ee 56% ioe WwW eases ou 1930. (Figs. 4-6). Typr: rroeden terkoffel prope St. Ulrich, 1889, ae - Lichenes Exsiccati ene ECTOTYPE, sont ae W! (thallus figured herein). Halonen et al. 1998) placed U. stuppeain synonymy with U. substerilis Motyka without discussion; however, we prefer to maintain U. stuppeaasa distinct taxon based ona number of differences in internal and external anatomy. First, in the type of U. substerilis the papillae on the primary branches are raised (tall), rounded at the top, and worn off to some degree closer to the base of the thallus. They are also better defined in shape, and gradually sparser, closer to the tips of the main branches. On the secondary branches the papillae also gradually be- come sparser, larger, and more infrequent towards the tips of the branches. The type of U. stuppea however, possesses papillae that are evenly distributed from the base to the tips of the main branches. The papillae themselves are less raised (shorter), and more blunt. On the secondary branches the papillae are nearly absent except close to the point of attachment to the main branch. The size, shape, and ontogeny of soralia have also been considered valu- able characters in distinguishing species (Herrera-Campos et al. 1998; Halonen et al. 1998; Ohmura 2001) and the soralia of U. stuppea and U. substerilis differ in a number of characters. Those of U. substerilis are raised above the cortex (not excavate) and produce few to many small isidiomorphs. As the isidiomorphs are abraded away with age the soralium becomes excavate and larger in size. This is contrasted with the soralia of U. stuppea which are not distinctly raised above the cortex and do not produce isidiomorphs. Instead, the soralia produce large coarse soredia, and, with age the soralia become larger in size and considerably deeper (more excavate). As discussed by Tavares (1987) and Ohmura (2001) cortical anatomy is also a valuable character for distinguishing Usnea taxa. The cortex of U. substerilis is harder, more rigid, and considerably thicker than that of U. stuppea and hasa dusty gray-brown (subpruinose) appearance in the herbarium. Likewise, the cortex of U. stuppea tends to be softer (occasionally sinking into slight foveae or depressions on the main branches) and lacks the granular appearance of U. substerilis. The branches of U. substerilis are also distinctly shorter than those of U. stuppea. It isimportant to note that both U. stuppea and U. substerilis dif- fer from material currently referred to U. lapponica Vainio (ie. thallus “B” on pL LENDEMER AND AND U. SUBSTERILIS 641 4 Fics. 4-6. Usnea substerilis. Fig. 4. Detail of lectotype y poi substrate. (N I f 1 tall raised ill ) Fig 5 Lectotype thallus marked No. 13 in packet f attachment to the by P. Clerc. Fig. 6. Detail of lectotype thallus: secondary branch and fibrils showing raised soralia with isidiomorphs (indicated by arrows). Scale bar = 1 cm (fig. 4, scale identical for fig. 6), .5 cm (fig. 5). — the lectotype card of U. stuppea) by the presence of a subpruinose (U. substerilis) or subglabrous (U. stuppea) cortex. The cortex of the material here referred to U. lapponica is lighter in color (yellower in the herbarium) than those of the other two taxa and glabrous instead of subpruinose or subglabrous. ACKNOWLEDGMENTS We wish to thank the curators of the following herbaria for loaning material for this study: BP, FH, H, NY, LBL, S, W. We are grateful for the comments of Gerry Moore and an anonymous reviewer. REFERENCES Cierc, P. 1987.Systematics of the Usnea fragilescens aggregate and its distribution in Scan- dinavia. Nordic J. Bot. 7:479-495. Cierc, P.1997.Notes on the genus Usnea Dill.ex Adanson. Lichenologist 29:209-215. Greuter, W,, J. McNeil, E.R. Barrie, H.M. Burbet, V. Demoutin, T.D. Fitigueiras, D.H. NIcoLson, P.C. SILva, 642 BRIT.ORG/SIDA 21(2) J.E. Skoa, P. TREHANE, N.J. TuRLAND, and D.L. Hawksworth (eds.). 2000. International code of botanical nomenclature (St. Louis Code) Adopted by the Sixteenth International Bo- tanical Congress St. Louis, July-August 1999. Koeltz, KOnigstein Hatonen, P,, P. CLerc, T. Gowaro, ILM. Bropo, and K. Wutrr. 1998. Synopsis of the genus Usnea (lichenized Ascomycetes) in British Columbia, Canada. Bryologist 101:36-60. Herrera-Campos, M.A., P.Crerc, and T.H. Nasu Ill. 1998. Pendulous species of Usnea from the temperate forests of Mexico. Bryologist 101:303-329. Moryka, J. 1930. Materialy do flory porostow Slaska (matériaux pour la connaissance des lichens de Silésie). Wydawncitwa Muzeum Sdaskiego w Katowicach 3:1-28. Morya, J. 1936. Lichenum generis Usnea studium monographicum, pars systematica. 1: 1-304 Oxmura, ¥. 2001. Taxonomic study of the genus Usnea (lichenized Ascomycetes) in Japan and Taiwan. J. Hattori Bot. Lab. 90:1—96. RASANEN, V.J.P.B. 1933.Contributions to the lichen flora of North America. Ann. Missouri Bot. Gard. 20:7-21. Tavares, |. 1987. The taxa of Usnea (Lichenes) described by Michaux from eastern North America. Mycotaxon 30:39-68. NOMENCLATURE AND TYPIFICATION IN THE GENUS USNEA (LICHENIZED ASCOMYCETES)— Ill. USNEA ALATA & USNEA SULCATA James C.Lendemer Isabelle |. Tavares Botany Department University Herbarium The ney OR NanG Sciences of Philadelphia University of Californic n Franklin Pky. 1001 VLSB #2465 Philadel, i ia, ae 9103, U.S.A. Berkeley, California 94720-2465, U.S.A. ee iorg ABSTRACT — This paper, the third in a series presenting cases of confusion of nomenclature and typification in the genus Usnea, discusses two names of pendent, apotheciate, angulose Usnea species from South America. The first, Usnea alata Motyka, was not correctly typified by ie ee 937) at ne time of description and is lectotypified here. The second, Usnea suieate Moby posi a lectotypification that is in con ie with the } | ctoty con ere. A lectotype is also selected for the name U.angulata f. er rugined iene Inner Ww which isconsidereda synonym of U. sulcata var. su cata RESUMEN ] J é ie A ] ] Este ¢ articulo > tipificacion en el género Usnea, discute dos sone de especies de Usnea colgantes, con apotecios, angulosas de Sur América. El primero, Usnea alata Motyka, no fue tipificado correctamente por Motyka (1937) en el momento de su descripcion y se lectotipifica a El is Saas ee sulcata Moen, ha sido mal | protolog aplicado debido a una lectotipificacion que esta en lectotipificada aqui. También se selecciona un lectotipo para el nombre U. cnailuid f. Jem ruginead Krempelhuber, que es considerado un sindénimo de U. sulcata var. sulcata. I. Usnea alata Motyka Usnea alata Motyka is one of the pendent, angulose, apotheciate species of Us- nea known to occur in South America. Though Motyka (1937) clearly intended a specimen in the Vainio herbarium (TUR) to serve as the type, no specimen annotated as the type by him has been located there. As noted by Alava (1986) there are in fact two specimens matching the collection data given in the protologue. One of these specimens (TUR-VAINIO #000492) represents a col- lection not distributed in Vainio’s Lichenes Brasiliensis Exsiccati and the other (TUR-VAINIO #00493) is a duplicate of Lichenes Brasiliensis Exsiccati #395. Since Motyka did not indicate if he intended the exsiccati collection to serve as the type there is no way to conclude with certainty which of these two speci- mens Motyka might have regarded as the type. Thus, we have chosen to lectotypify the species using the specimen distributed in Vainio’s exsiccati in- stead of simply assuming its status as the holotype. It should be noted that Her- SIDA 21(2): 643-650. 2004 644 BRIT.ORG/SIDA 21(2) rera-Camposet al. (1998) did not effectively lectotypify U. alata when they stated “TYPE: BRAZIL, Minas Geraés, Chequeira, 1885(TUR holotype)...” because a single specimen was not cited and two specimens are present in Vainio’s herbarium. The specimen here selected as the lectotype agrees well with Motyka’s protologue and all duplicates of this collection reviewed by the first author are conspecilic with the specimen selected as the lectotype. oo Usnea alata Motyka, Lich. gen. Usnea 2(1):395-396. 1937. (Figs. 1-2). Tyer: BRAZIL. MINAS GERAES: Chequeira, in arbore, 1885, Vainio s.n. = Lichenes Brasiliensis Exsiccati #395 (LECTOTYPE, here designated: TUR! (Vainio Herbarium #00493); ISOLECTOTYPE: S! Though U. alata was considered a distinct taxon by Herrera-Campos et al. (1998) it seems likely that some authors would consider it conspecific with U. sulcata Motyka (as redefined here). These two taxa differ in a number of respects, how- ever, including the type of angulation of the branches, structure and shape of the fibrils, size of the apothecia, and overall appearance. Furthermore, much confusion has apparently resulted from the fact that Motyka changed his in- terpretation of U. alata prior to the publication of the monograph and after he annotated most of the specimens cited therein. That Motyka re-interpreted U. alata shortly before its publication is evidenced by the fact that many speci- mens in S that were annotated by Motyka as U. alata or “U. angulata var. alata” were cited by him as paratypes of U. sulcata var. neutra Motyka. Indeed, these specimens are not referable to U. alata in the sense of the type because they possess apothecia that are generally smaller than those of the type, the branches are angulose (having parallel ridges or sharp foveae) instead of alate, the fibrils are long, slender, regular, and abundant, and the chemistry of the type of U. alata apparently differs from that of U. sulcata var. neutra. Incidentally, Motyka (1937) reported the type of U. alata to have a KOH- medulla; however as re- ported by Herrera-Campos et al. (1998) the type actually contains norstictic and connorstictic acids.) It is tempting to consider the possibility that U. alata represents the non-sorediate, fertile counterpart to U. paradoxa Motyka (as de- fined by Lendemer & Tavares 2003). a IL. Usnea sulcata Motyka While the first author was working with U. angulata Acharius, a number of problems in typification and taxonomy were encountered involving taxa re- cently placed in synonymy with U. angulata by other authors (Awasthi 1986; Herrera-Campos et al. 1998; Ohmura 2001). One such synonym is U. sulcata Motyka. Motyka (1937) described U. sulcata without the mention of soralia and with the description of small pruinose apothecia. As Tavares (2002) has noted, this taxon (originally described as apotheciate) was lectotypified with a soraliate specimen that lacked apothecia and the species was then placed in synonymy with U. angulata (Awasthi 1986). Usnea angulata in our opinion isa ALATA AND IIL CHICATA 645 ver Fics. 1-2. Usnea alata Fig 1 Lectotype thallus Fig 2. Detail of | lval I h witl “winged second- L L L | ra el | es 4h hk cl 1 H | Cralok nc 7 J w ? 646 BRIT.ORG/SIDA 21(2) soraliate species from eastern North America and northern Mexico. Apparently these names were placed into synonymy because the lectotype selected by Awasthi (1986) is densely soraliate (not apotheciate) and thus superficially simi- lar to U. angulata. All later workers with the exception of Tavares (2002) have continued to include U. sulcata as a synonym of U. angulata. Motyka’s designation of the type specimen of U. sulcata was simply “Typus in Museo Botan. Univ. Fennicae in Turku.—Locus classicus: Brasilia, Minas Geraés, Sitio, 1885 Vainio.” As noted by Tavares (2002) there are four specimens in TUR-VAIN matching this description, one of which (TUR- VAIN 00450) was selected by Awasthi as the lectotype. One of us JCL) has examined all four of these specimens; three of the four thalli are soraliate and thus not suitable can- didates for lectotypification. A fourth thallus, though not soraliate, is small, poorly developed, and lacking apothecia. This fourth specimen isa possible can- didate for lectotypification (since it does not possess soralia); however, it is too poorly developed to allow positive identification as U. sulcata and does not pos- sess apothecia, a feature Motyka described in the protologue. It would seem, therefore, that none of the specimens in TUR-VAIN matching Motyka’s pub- lished data are ideal for lectotypification. It should be noted that Vainio (1890) stated that all of the specimens of “U. angulata” from Sitio were sterile. Interest- ingly, though the specimens at Turku that were collected at Sitio are not apotheciate, all of the other specimens we have examined that Motyka cited in the protologue are either apotheciate, pycnidial, or lack both apothecia and pycnidia but are not soraliate. Likewise, with the exception of the specimens distributed by Vainio in his Lichenes Brasilienses Exsiccati, all of the exsiccati specimens (that we have examined) cited by Motyka in the protologue also are either apotheciate or pycnidial. There is, however, a specimen in Motyka’s her- barium (now at LBL) labeled as having come from Sitio that is apotheciate and it is this specimen that we select as the lectotype. The comparisons Motyka (1937) made between U. sulcata and other taxa also serve to confirm that the lectotype selected by Awasthi conflicts with the published diagnosis. Motyka contrasted U. sulcata with U. paradoxa Motyka, a soraliate taxon (see Lendemer @ Tavares 2003) and placed in synonymy with U. sulcata a previously described form anda variety of U.angulata(U.angulata forma ferruginea Krphb. and U. angulata var. rubiginosa Hillmann), both of which are based on apotheciate specimens. Unfortunately, the type material of U. angulata var. rubiginosa Hillmann was destroyed during World War II and thus is not available for study. It is retained here as a questionable synonym. Usnea sulcata Motyka var. suleata, Lich. gen. Usnea 2(1):478-480. 1937. (Figs. 5-6). Type: BRAZIL: MINAS GERAES: Sitio, 1885, Vainio s.n. (LECTOTYPE, here designated: LBL! #300.7 (fragment ee here). Us lata Ac] ferruginea Krempelhuber, Flora 61(28):4 37. 1878. TYPE: Lorentz & omnia (LECTOTYPE, here destonated Mh: ISOLECTOTYPE: UC). LENDEMER AND , ALATA AND II CINCATA yi Fics. 3-6. Usnea sulcata var. neutra. Fig. 3. Lectotype thallus. Fig. 4. Detail of | 79.5.1) lh Af lnrtnt toe oT h. Usnea sulcata Fig 6. Lecto- var. sulcata. F g “winged type thallus. Scale bar = 0.5 cm. vt VG} 648 BRIT.ORG/SIDA 21(2) @) = Usnea angulata Acharius var. rubiginosa Hillmann, Repert. Spec. Nov. Regni Veg. 27U16- 25):291. 1930. TYPE: BRAZIL: Sellow s.n. (HOLOTYPE: B, destroyed). The lectotype packet (LBL 300.7.) of U. sulcata contains fragments of several thalli, four of which possess apothecia. The fragment selected here as the lecto- type contains norstictic, caperatic, a nd connorstictic acids by TLCtUR.C. Harris. pers.comm.) and thus is chemically similar to the type of U. alata Motyka. It is important to note that the lectotype selected here differs in appearance from the soraliate specimen previously selected as the lectotype (by Awasthi 1986) and possibly does not represent part of the same gathering. Since we have shown here that U. sulcata Motyka is in fact an apotheciate species from South America the previously accepted synonymy with U. angulata Acharius should be rejected. Usnea angulata f. ferruginead Krempelhuber was described without the designation of a type; thus here we have chosen to select the specimen in Krempelhuber’s herbarium (M) as the lectotype. It should be noted that the red coloration which Krempelhuber’s epi- thet suggests is not actually a pigmentation of the cortex such as that seen in U. michauxii LL. Tavares or U. pensylvanica Motyka but instead is simply a discol- oration of the entire collection. No other collection with similar discoloration has been seen by us. Motyka (1937) also described U. sulcata var. neutra Motyka on the basis of its medulla having a negative KOH reaction. Later, Rizzini (1952) elevated this taxon to specific rank. Subsequent authors have not discussed this taxon; when re-evaluating the status of the other apotheciate angulose taxa in South America it became clear that U. sulcata var. neutra was also in need of revision. The type specimen was indicated by Motyka (1937) to be in his personal herbarium (now at LBL); however no such specimen was located in a loan of specimens from LBL of the apotheciate angulose taxa discussed here. Furthermore, it was also clear that there exists much confusion in the use of this name because prior to publishing the name U. sulcata var. neutra Motyka included specimens later cited as paratypes of U. sulcata var. neutra in his concept of U.alata Motyka. In order to alfix the usage of the name U. sulcata var. neutra to a specimen, we have chosen to lectotypify the name with one of three specimens from S match- ing the data given in Motyka’s protologue. All of these specimens differ chemi- cally from the type U. sulcata var. sulcata (see below). ey Usnea ees Motyka var. neutra Motyka, Lich. gen. Usnea 2(1):480-481. 1937. (Figs. 3-4). Usnea neutra (Motyka) Rizzini, Revista Brasil. Biol, 12(4) page #. 1952. TYPE: BRAZIL. a AS GE RAS: Ciade de Caldas, 1879, Regnell sn. (LECTOTYPE, designated here: S! #1100 fragment ligured here). — Of the eee lent to the first author from S, three specimens matched the data given in Motyka’s protologue for the type of U. sulcata var. neutra Motyka. Only the specimen selected here as the lectotype was actually identified as U. AI ATA AND SUICATA 649 sulcata var. neutra (R. Santesson, undated annotation) and marked “TYPUS” (not in Motyka’s hand). It is important to note, however, that at present we do not consider U. alata and U. sulcata var. neutra to be synonyms because they differ both chemically and morphologically. We also prefer to retain U. sulcata var. neutra as distinct from U. sulcata var. sulcata because the type specimens of the two taxa also differ on chemical and morphological grounds. (The lecto- type of U. sulcata var. neutra contains the stictic acid complex in addition to usnic and norstictic acids whereas the lectotypes (selected here) of U.alataand U. sulcata var. sulcata lack the stictic acid complex.) As noted above, Rizzini (1952) elevated Motyka’s epithet to specific rank; we however choose to follow Motyka’s original placement pending further study. It should be noted that U. sulcata var. neutra Motyka is not synonymous with U. angulata var. neutra Motyka ex Rasdnen, a soraliate taxon. The latter name will be treated in a fu- ture publication (Lendemer in prep.). ACKNOWLEDGMENTS We wish to thank Orvo Vitikainen (H) for providing data associated with the lectotype of U. angulata as well as Paul Silva (UC) for providing nomenclatural advice; also, we thank Soili Stenroos (TUR) for providing access to the Vainio specimens from Sitio, and Jan Bystrek (LBL) for locating the material from Motyka’s herbarium. Also, we wish to thank James Macklin (PH) and Richard Moe (UC) for reviewing drafts of the manuscript, the curators of B, LBL, M,and TUR, for loaning material to the first author, and R.C. Harris for performing TLC on several of the specimens cited here. REFERENCES Avaya, R. 1986.Edvard August Vainio's journey to Brazil in 1885 and his Lichenes Brasilienses exsiccati. Publ. Herbarium, Univ. Turku 1:1-174 Awastui, G. 1986. Lichen genus Usnea in India. J.Hattori Bot. Lab. 61:333-421, Greurter, W.,J. McNeil, ER. Barrie, HM. Burbet, V. DemouLIN, T.D. FiticueiraAs, D.H. NIcoLson, P.C. Sitva, J.E. Skos, P. TREHANE, NJ. TurLaNo, and D.L. Hawkswortn (eds.). 2000. International code of botanical nomenclature (St. Louis Code) Adopted by the Sixteenth International Bo- tanical Congress St.Louis, July-August 1999. Koeltz, KOnigstein Herrera-Campos, M.A., P. Clerc, and T.H. Nasu Ill. 1998. Pendulous species of Usnea from the temperate forests of Mexico. Bryologist 101:303-329. HILLMANN, J. 1930. Einige bemerkungen Uber die roten Usneen. Repert. Spec. Nov. Regni Veg. 27(16-25):287-291. KrempeLHuser, A. 1878. Lichenes, collecti in republica Argentina a Doctoribus Lorentz et Hieronymus, determinati et descripti a Doct. A.de Krempelhuber, Monacoensi. Flora 61(28):432-439. Lenoemer, J.C.in. prep. A revision of the pendent angulose species of the genus Usnea in the Americas. 650 BRIT.ORG/SIDA 21(2) Lenbemer, J.C. and Il. Tavares. 2003. Nomenclature and typification in the genus Usnea (Lichenized Ascomycetes) Il: Usnea paradoxa and Usnea fallax. Proc. Acad. Nat. Sci. Phila- delphia 153:181-185. Motyka, J. 1937. Lichenum generis Usnea studium monographicum. Pars Systematica 2(1):305-560. Oxmura, Y. 2001. Taxonomic study of the genus Usnea (lichenized Ascomycetes) in Japan and Taiwan. J. Hattori Bot. Lab. 90:1-96. Rizzini,C.1.1952. Species Organenses generis lichenum Usneae.(Omnes acidum usnicum praebentes). Revista Brasil. Biol. 12:337-348. Tavares, 1.1. 2002. Notes on lectotypification of Usnea fulvoreagens, U. gigantea, U. sulcata, and U. cavernosa. Constancea 83: (http://ucjeps.berkel lu/constancea/83/tavares usnea.html) Vainio, E.A. 1890. Etude sur la classification naturelle et la morphologie des lichens du Bresil. Acta Soc. Fauna FI. Fenn. 7:1—247. RESURRECTION OF A LITTLE-KNOWIN-SPECIES OF OENOTHERA SECT. OENOTHERA IN NORTHEASTERN MEXICO (TRIBE ONAGREAE: ONAGRACEAE) Warren L.Wagner Department of Botany, MRC-166 P.O. Box 37012 Smithsonian Institution Washington, DC (District of Columbia) 20013-7012, U.S.A. wagnerw@si.edu ABSTRACT Oenothera aa ee sect. Oenot] Nutanti ) was recently placed in synonymy with the 1 and variable O. pubescens Willd. ex — A series of recent col- lections from Coahuila, Rees a Tamaulipas, and Zacatecas, México, show that O. pennellii isa distinctive, subacaulescent, apparently perennial species restricted to northeastern Mexico at rela- tively high elevations. It is largely allopatric from the widespread, caulescent, annual or biennial O. pubescens. Like O. pubescens, O. pennellii is a permanent translocation heterozygote species. Key Worps: Oenothera sect. Oenothera subsect. Nutantigemma, evening-primrose, Mexico, Onagraceae RESUMEN Oenothera penne Munz (perteneciente a Oenothera subsect. Nutantige mma) fue colocada le O. pubescens Willd. ex Spreng variable y de amplia distribucion. Las colecciones procedentes de Coahuila, Nuevo Leon, 1 moans y Zacatecas en México, muestran que O. pennellii es una ee distinguible por su habito sub-acaulescente y que aparentemente esta restringida al noreste de México en elevaciones aaa ane donde es ampliamente alopatrica con O. pubescens, especie caulescente, anual y de amplia dist Oenothera pennellii Munz was described in 1939 and was maintained in Munz’s treatment of all North American Onagraceae (1965). Following detailed stud- ies of subsect. Raimannia, Dietrich and Wagner (1988) placed O. pennellii in synonymy with the widespread and variable O. pubescens Willd. ex Spreng. They considered it a rare (five known collections) but noteworthy variant among several in O. pubescens (Dietrich & Wagner 1988: 77). Subsequent to that study, many new specimens have been collected in high-elevation areas (2,000- 3,600 m) in the northeastern Mexican states of Coahuila, Nuevo Leén, Tamaulipas, and Zacatecas; most of these collections are deposited at TEX/LL. What previously seemed a sporadic variant now appears to represent a distin- guishable species restricted to I n Mexicoat relatively high elevations, and largely allopatric from O. pubescens, which ranges from Arizona and New Mexico to Guatemala, and in South America in the Andes of Colombia, Ecua- SIDA 21(2): 651-655. 2004 652 BRIT.ORG/SIDA 21(2) dor, and Peru. The geographical ranges of the two species overlap only in Sierra 8 E } del Carmen in Coahuila, but the single collection of O. pubescens from there was made several hundred meters lower than the lowest known populations of O. pennellii. Ocnothera pennellii characteristically has sma —_— | flowers (petals 6-8 mm long) and is subacaulescent, with stems 0-2(-4) cm long. In contrast, O. pubescens usually has petals (6-)15-25(-35) mm long and is always caulescent with conspicuous stems up to 100 cm long, even in adverse moisture regimes. Oenothera pubescens never flowers from the basal rosette but has flowers formed in the leaf axils near the tips of the stems, as is typical in the genus. It is also = annual or biennial, whereas O. pennellii appears to be a short-lived perennial from an enlarged taproot. On this basis lam here resurrecting this regional en- demic to species status. Like O. pubescens, O. pennellii is a permanent translo- cation heterozygote (PTH) species. Pollen fertility is about 50% judging from several collections McDonald 2064, Hinton 18858, 20452) examined. Permanent translocation heterozygosity has been very important in the evolution of the genus Ocnothera and several other genera of the Onagraceae. The metacentric chromosomes with pycnotic, condensed proximal regions (Kurabayashi et al. 1962; Cleland 1972; Raven 1979) have been associated with the regular occur- rence of rings of chromosomes, resulting from reciprocal translocations. The wu phenomenon of reciprocal translocations reaches an endpoint of development in the specialized system known as PTH. The best known species possessing this system are the members of Oenothera subsect. Oenothera, in which the structure and mechanisms were worked out (Cleland 1972; Harte 1994; Deitrich et al. 1997). In addition to the translocations, t — ne system requires balanced le- thals, which prevent the formation of the homozygous combinations (most easily observed as ca. 50% infertile pollen), self-pollination, and alternate dis- junction of the chromosomes during meiosis. —_ Oenothera pennellii Munz, Leal |. W. Bot. 2:156, 157. 1939. (Fig. 1). TYPE: MEXICO. NUEVO LEON: Sierra Madre Oriental, Mt. “El Infiernillo,” Pablillo, SE of Galeana, 2,750-2,900 m, 29 Jun 1934, EW. Pennell 17139 (HOLOTYPE: US-01640419 ISOTYPE: PH not seen, POM). = Acaulescent or subacaulescent, short-lived perennial herb from fairly stout tap- p 5 root; stems occasionally present, 1-4 cm long. Rosette and cauline leaves 2-6(- 10) x 0.2-1.5 cm long, oblong-lanceolate, sinuate-pinnatilid to occasionally re- motely serrate on smaller leaves, strigillose and hirtellous, especially on the veins. Flowers axillary, arising among the basal leaves or on the short stems. Floral tube 17-30 mm long, nodding prior to anthesis, reddish purple, sparsely hirtellous, the hairs appressed or spreading, and occasionally also strigillose. Sepals reflexed in pairs at anthesis, 5-6 mm long, oblong-lanceolate, pubescent, free tips ca. 0.2 mm long. Petals 6-8 mm long, about as broad, yellow, changing to reddish orange when wilted, slightly notched apically with short tooth in WAGNER, RESURRECTION OF OENOTHERA PENNELLII 653 it B. Habit (Hinton et a) 20452 & Wendt & Ad, icz 5230), 1 bud (P If 17139, a B. Leaf (Pennell 17139 holotype). C I (Hinton et al 20452) D Flower with part of floral tube, but not ovary (Hinton et al. 18858).E. Capsule (Pennell 17139 holotype) F. Seed (Pennell 17739, holotype). — 654 BRIT.ORG/SIDA 21(2) notch. Staminal filaments 6-7 mm long, subequal; anthers 2-3.2 mm long; pol- len ca. 50% fertile. Style 20-35 mm long; stigma surrounded by the shedding anthers at anthesis, the lobes ca. 1.5 mm long. Capsules 16-28 mm long, cylin- drical, thin-walled, sessile, hirtellous and strigillose, ca. 3mm in diameter. Seeds 1-12 mm long, light brown, globose-obovoid, with median ridge and shallowly regularly pitted surface. Distribution—Oenothera pennellii occurs in open areas in mixed conifer (Pinus, Pseudotsuga, Abies) and Quercus forests and in subalpine Pinus forest, Coahuila, Nuevo Leon, Tamaulipas, and Zacatecas, Mexico, from 2,000 to 3,600 m. Flowering from May through September. Asoo Examined: MEXICO. Coahuila: Madera del ¢ Carmen, upper end of Dos Canyon, at road fork to Campo Uno, 23 Jun 1976, Fryxell 2715 (LL, NY); Sierra Madera del Carmen, at Campo El Tres, an ae logging camp in the high country, pee oe N, 102°36' WI, 7 Aug 1974, Wendt & Adamcewicz 523C (TEX): Ocampo, along old logging road, of mill at Campo 4 adjacent to sawdust pile, [28°59 N, 102°33' W], 28 May 1975, Riskind & Patterson 1826 (LL); Sierra de Parras, Purpus 4624 (UC); Arteaga, Sierra Zapaliname, 27 Jun 1990, Hinton et al. 20452 (TEX); Sierra del Coahuila, 2 Jun 1985 Hinton et al. 18858 (TEX); Sierra del Arteaga, Canon de la Carbonera, Las Vigas, [25°20'N, 100°39' WI, s rreal & Car ranza 3776 (TEX). Nuevo Leon: Picacho de San Onofre, ladera este, 5 Jul 1985, ead 1662 (TEX). Tamaulipas: filo y lado este de Pena Nevada, 5 Jul 1985, McDonald 1630 (TEX); Miquihuana, sien de pina Nevada, ceja y ladera S y SO[23° 35'N, 99° 46' W], 3600 m, 22 Aug 1986, McDonald 2064 (MO, TEX [2]). Zacatecas: Sierra Madre Oriental, Mt. “El Temoroso,” N of Aranzazu, l7 Jul 1934, Pennell 17466 (US). The five collections cited by Dietrich and Wagner (1988) as the low-growing phenotype are a mix of short-stemmed Oenothera pubescens and O. pennellii. Three of the collections represent O. pennellii and are cited above. The collec- tion from Sierra del Carmen, Coahuila (Henrickson 11643[MO)), is O. pubescens but was collected in a heavily grazed area and thus had very short stems. Culti- vated material of this collection at MO has stems up to 30cm long and is atypi- cal of O. pubescens only in having flowers much smaller than usual (petals ca. 6 mm). Another collection (Moore 3157 from Hidalgo, MO) also is O. pubescens and occurs well outside the range of O. pennellii. The substrate for Oenothera pennellii is largely unknown. Wendt & Adamcewicz 523C give the substrate as rhyolite, and Pennell 17466 gives it as gravelly andesite, both volcanics. With the information available it is not pos- sible to ascertain if O. pennelliiis restricted to volcanic substrates; however, Guy Nesom (pers. comm.) indicates that the type locality is gypseous. Since lime- stone is very common throughout this region it should be looked for on this substrate as well. ACKNOWLEDGMENTS Ithank Tom Wendt for calling my attention to the large number of recent col- lections in TEX of Onagraceae, particularly Oenothera, which allowed me to appreciate the distinctions between O. pubescens and O. pennellii. 1 thank Alice WAGNER, RESURRECTION OF OENOTHERA PENNELLII 655 Tangerini for her excellent illustration and Denise Mix for assistance with the collections and comments on the draft manuscript. | also appreciate the reviews by Peter Hoch and Guy Nesom, which improved the clarity and information content of the paper. REFERENCES CLELAND, R.E. 1972. Oenothera cytogenetics and evolution. London: Academic Press. Dietrich, W.and W.L.Waaner. 1988. Systematics of Oenothera section Oenothera subsection Raimannia and subsection Nutantigemma (Onagraceae). Syst. Bot. Monogr. 24:1—91. Dietrich, W., W.L. Wacner, and PH. Raven. 1997. Systematics of Oenothera sect. Oenothera subsect. Oenothera (Onagraceae). Syst. Bot. Monogr. 50:1-234. Haare, C. 1994. Oenothera: Contributions of a plant to biology. Monographs on Theortical and Applied Genetics vol. 20. Berlin: Springer-Verlag. KurasayasHi, M., H. Lewis, and PH. Raven. 1962. A comparative study of mitosis in the Onagraceae. Amer. J. Bot. 49:1003-1026. Munz, PA. 1939. Interesting western plants IV. Leafl.W. Bot. 2:156-158. Munz, PA. 1965. Onagraceae.N. Amer. FI. II. 5:1-278. Raven, PH. 1979. A survey of reproductive biology in Onagraceae. New Zealand J. Bot. 17:575-593. 656 BRIT.ORG/SIDA 21(2) BOOK NOTICES Books from the Hunt Institute for Botanical Documentation GAVIN D.R. Bripson, JAMES J. WHITE, and LUGENE B. BRUNO. 2003. American Botani- cal Prints of Two Centuries. (ISBN 0-913196-75-4, pbk.) Hunt Institute of Botanical Documentation, 5th Floor, Hunt Library, Carnegie Mellon Uni- versity, Pittsburgh, PA 15213, U.S.A. (Orders: 412-268-2434; 412-268-5677 fax; email: huntinst@andrewcmucedu; huntbot.andrew.cmuedu). $25.00, 239 pp., 114 figs (106 color). Pictorial stiff paper cover, 8" x 10 1/2" Two centuries of botanical prints is a wide range to cover in so short a book, but this task is m andere quite wellin American Botanical Prints of Two Centuries. Starting with the 19th century, the authors take us through a period of utilitarian prints designed for the sole purpose of botanical research. We see the evolution of the botanical print as one of necessity to one of ar co technology improved, artists were free to elaborate and put their own personal touch on a piece. ce with the 20th century, we begin to see the effects of light and color. We see more styl- ized works, things we would want to hang on the wall. = American Botanical Prints of Two Centuries is laid out very well. There is no table of contents, but there is a very well-organized catalogue of prints including biographical information on the art- ists on pages 150-19 It is impossible to merely glance at the prints displayed in this book. Every picture is a work of art, even the utilitarian apes Wo knew a Lousiet oF vo could be so enjoyable and beautiful to look at!—Abra Al Texas, 509 Pecan Street, Fort Worth, TX 76102- 4060, U.S.A GaviIN D.R. BRIDSON (Compiler), SCARLETT T. TOWNSEND, ELIZABETH A. POLEN, and ELIZABETH R. Smitu (eds.). 2004. BPH-2. Periodicals with Botanical Content: Constituting a Second Edition of Botanico-Periodicum Huntianum (Vol- ume 1 A—-M, Volume 2 N-Z). (ISBN 0-913196-78-9, pbk.) Hunt Institute of Botanical Documentation, 5th Floor, Hunt Library, Carnegie Mellon Uni- versity, Pittsburgh, PA 15213, U.S.A. (Orders: 412-268-2434. 412-268-5677 fax; email: huntinst@andrew.cmu.edu; huntbot.andrewcmuedu). $130.00 (2 vol. set), 1470 pp., 81/2" x 11". Congratulations to the team at the Hunt Institute of Botanical Documentation for a second edition ot Botanico-Periodicum-Huntianum or BPH-2. From an editor who has already used the second edi- tion, here is a big thank you for an incredible job. The first edition of BPH was published in 1968 and recorded periodicals published down to the end of 1967. In 1991, B-P-H/S (Botanico-Periodicum- ea iy ah me sda was pub sere With B-P-H/S, the iecues 2 titles jumped to over NJ +] ingthe total to over 33 ,000, about 21,000 additional titles since 2 1967." ‘BPH-2isa ee but not wholly comprehensive listing by title sal penoulesis from ee to 200 that mere ain (or, in some period of their histor lud the Zee and bibliogra- phy.” Good job thou good and faithful eee Burney Lipscomb, aie Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A pale SIDA 21(2): 656. 2004 TAXONOMY OF HYMENOXYS SUBGENUS MACDOUGALIA (ASTERACEAE: HELENIEAE: TETRANEURINAE) Mark W. Bierner School of Biological Sciences and Plant Resources Center The University of Texas Bierner@mail.utexas. a ABSTRACT A taxonomic treatment is presented for Hymenoxys subg. Macdougalia, which includes a single spe- cies, Hymenoxys bigelovii. The treatment includes a discussion of the original circumscription of Hymenoxys bigelovii (as Actinella bigelovii), the later description of the genus Macdougalia to ac- commodate this taxon, and the eventual placement, based on chemical, cytological, and morpho- logical evidence, of Macdougalia within Hymenoxys as a monotypic subgenus. The treatment also includes synonymies of Hymenoxys subg. Macdougalia and Hymenoxys bigelovii, lectotypilication of Actinella bigelovii, and a description and range map for Hymenoxys bigelovii. RESUMEN Se presenta un tratamiento taxondmico de Enenee): une Macgousai que eye) una roe especie, Hvmenorve bi gelovii Fl H. bigelovii (como Actinella bigelovii), la ie del género Macdougalia Oe are este taxon, y la colocacién eventual, basada en pruebas quimicas, citologicas, y morfoldgicas, de Macdougalia en Hymenoxys como un subgénero ae El tratamiento incluye también sinonimias de Hymenoxys subg. Macdougalia y Hymenoxys bigelovii, lectotipificacion de Actinella bigelovii, y una descripcion y mapa de distribucion de Hymenoxys bigelovii Hymenoxys Cass. subg. Macdougalia (A. Heller) Bierner comprises only one species, Hymenoxys bigelovii (A. Gray) K.L. Parker. This taxon was originally described as Actinella bigelovii A. Gray, Actinella Pers. being the generic name commonly used at that time (e.g., Torrey & Gray 1842) for taxa now placed in Tetraneuris Greene and Hymenoxys. When describing Actinella bigelovii, Gray (1853) made no comments that would indicate any hesitation on his part as to its placement in Actinella. Later (1883), when Gray positioned Actinella bigelovii in Actinella section Hymenoxys, he commented, “Connects [section] Hymenoxys with [section] Euactinella, and with section Dugaldea [sic] of Helenium.” This statement indi- cates to me that Gray may not have been completely comfortable with his place- ment of this taxon. His section Hymenoxys equates today with Hymenoxys subg Hymenoxys, Hymenoxys subg. Phileozera (Buckley) Cockerell (in part), and Hymenoxys subg. Picradenia (Hook.) Cockerell (Bierner 200D, his section Euactinella equates today with Tetraneuris, Hymenoxys subg. Phileozera (in SIDA 21(2): 657-663. 2004 658 BRIT.ORG/SIDA 21(2) part), and Hymenoxys subg. Rydbergia (Greene) Bierner (Bierner 2001; Bierner & Turner 2003), and his section Dugaldea [sic] of Helenium equates today with Hymenoxys subg. Dugaldia (Cass.) Bierner (Bierner 2001). Heller (1898) seems to have been even less sure about its placement, as in- dicated by his comment, “In habit it is more like the genus Tetraneuris, but has a different involucre, and while its involucre is somewhat similar to that of the genus Picradenia |= Hymenoxys subg. Picradenial, there is a wide difference in habit.” His solution was to describe the genus Macdougalia to accommodate this one taxon, a circumsct iption followed by Cockerell (1904), Rydberg (1915), and Robinson (1981). Other workers, such as Turner and Powell (1977) and Karis and Ryding (1994), felt there was no clear basis for maintaining Macdougaliaas a separate genus and submerged it in Hymenoxys. Spring et al. 1994) began their study of chemical components of glandu- lar trichomes in Hymenoxys and related genera by recognizing Macdougalia as a genus separate from Hymenoxys. By the end of the study they were of the opinion, based on sesquiterpene lactone and monoterpene glycoside chemis- try, that Macdougalia should be incorporated into a broader concept of Hymenoxys. Likewise, Bierner and Jansen (1998), who began their study of DNA restriction site variation in Hymenoxysand related genera recognizing Macdougalia as a distinct genus, concluded that it is in fact congeneric with Hymenoxys. The relationship of Macdougalia to Hymenoxys is further supported by similarities of flavonoid chemistry (e.g., Wagner et al. 1972) and chromosome number, 2n = 30 being the chromosome number of H. bigelovii (Speese & Baldwin 1952: Strother 1966; Bierner unpublished and Parker & McClintock unpublished— see representative specimens) and the predominant number among the diverse taxa ol Hymenoxys(e.g Speese & Baldwin 1952; Beaman & Turner 1962: Strother 1966; Sanderson 1973; Turner et al. 1973). In 2001, Bierner formally recognized Macdougalid as a subgenus of Hymenoxys. While Spring et al. (1994) were confident about the association of Macdou- galia with Hymenoxys, they were less sure about its relationship to other taxa within Hymenoxys. The phenogram prepared from sesquiterpene lactone data placed H. bigelovii (as Macdougalia bigelovii) closest to H. hoopesii of subg. Dugaldia and H. rusbyi of subg. Picradenia. The strict consensus tree prepared by Bierner and Jansen (1998) placed H. bigelovii (as Macdougalia bigelovii) in the clade containing taxa of Hymenoxys subg. Dugaldia, Hymenoxys subg. Picradenia,and Hymenoxys subg. Plummera, but no clear association with any species in particular was apparent. Morphology also supports the placement of Macdougalia in Hymenoxys. The stems, peduncles, receptacles, ray florets, and disc florets of Hymenoxys bigelovii are very similar to those of the Hymenoxys species in general. Con- versely, a substantial number of morphological differences support the recog- nition of Macdougalia as a subgenus. BIERNER, TAXONOMY OF HYMENOXYS SUBGENUS MACDOUGALIA 659 Hymenoxys bigelovii usually has all simple leaves that are eglandular or sparsely glandular. Among the other taxa of Hymenoxys, only H. hoopesii has all simple leaves, and only H. texana has leaves that are weakly to moderately glandular (all of the other taxa have distinctly glandular leaves). As inessentially all of the perennial taxa of Hymenoxys (and Tetraneuris as well), the basal leaf bases of H. bigelovii are persistent and tend to thicken the caudices distally as the plants age. The basal leaves of H. bigelovii, however, decay down to the veins so that the tops of the caudices usually appear to be encased in a stringy cocoon. I have observed this decay to the veins occasion- ally in other taxa of Hymenoxys, but it is unusual and never creates the appear- ance of a stringy cocoon. The phyllaries of Hymenoxys bigelovii are in two unequal series, as they are in most of the taxa of Hymenoxys (those of subg. Dugaldia and subg. Ryd bergia are in two or three subequal series). The outer phyllaries of H. bigelovii are basally connate only slightly to 1/5 their lengths; the outer phyllaries of the other Hymenoxys taxa with two unequal series (except for H. texana) are basally connate 1/4 to 2/3 their lengths. The inner phyllaries of H. bigelovii are narrowly lanceolate to narrowly obovate, have aristate apices, and very dis- tinctly exceed the outer in length; inner phyllaries of the other Hymenoxys taxa with two unequal series are usually obovate, have acuminate to usually mucr- onate apices, and surpass the outer in length only slightly or not at all. The relationship of Hymenoxys bigelovii to other taxa of Hymenoxys re- mains unclear Its relatively large involucres (13-20 mm high by 23-32 mm wide) might suggest a connection to H. hoopesii, H. brandegeei, or H. grandiflora, but the phyllaries are very different. Perhaps a clue rests with some unusual popu- lations of H. richardsonii var. richardsonii from Fremont County, Wyoming (e.g, Fisser 661 and 699 [RMI, and Dorn 3516 [NY, RMI). The phyllaries look so much like those of H. bigelovii that | was convinced when I first saw the specimens that these plants represented an undescribed species belonging to subgenus Macdougalia, even though the plants had divided leaves (blades are only rarely divided into three segments in H. bigelovii) and were well separated geographi- cally from H. bigelovii. When I was able to see them in the field, however, it was obvious that they are indeed plants of H. richardsonii var. richardsonii, but with narrower, longer, aristate inner phyllaries. Yet, this illustrated to me that it is not a long morphologic leap from the involucres of H. bigelovii to those of some other Hymenoxys species. TAXONOMY Hymenoxys subg. Macdougalia (A. Heller) Bierner, Lundellia 4:39. 2001. Macdougalia A. Heller, Bull. Torrey Bot. Club 25:629. 1898. TYPE SPECIES: Actinella bigelovii A. Gray, Pl. Wright. 2:96. 1853. (= Hymenoxys bigelovii) Hymenoxys bigelovii (A. Gray) K.L. Parker, Madrono 10:159. 1950. Basiony™: Actinella 660 BRIT.ORG/SIDA 21(2 bigelovii A. Gray, Pl. Wright. 2:96. 1853. Type: U.S.A. NEW MEXICO. Grant or Sierra Co.: “Cop- per r Mines - on anes mountains’ ‘Mlpetoly pe label), 17 cay 1852, Bigelow s.n. (LECTOTYPE [per pre- ISOLECTOTYPES: NY!, US-27506!, US-275071, US-27508!). The ae notation in the pelea “On mountains near the copper mines, and near the Mimbres, New Mexico; April, June,” indicates that Gray was looking at more than one Bigelow collection when he wrote the description. Furthermore, the type sheet at GH contains three Bigelow specimens, one with the notation “Copper Mines” (far right), an- other with the notation “Near the Mimbres June 1852” (center), and another with the nota- tion “Copper Mines - on the mountains, April 17.1852.” (far left). The specimen to the far left was already annotated as the lectotype when | borrowed it (no name or date on the label), and | likewise annotated it as the lectotype. I did so because there is no doubt at all that Gray was using this specimen; it is the only plant on the sheet that has cauline leaves that are divided into three segments, and the original con includes, “..foliis angustissime linearibus rigidis integerrimis paucisve caulinis trilobis..” In addition, collection informa- tion beneath the specimen includes, “Actinella Rigcloviin n. sp. (Pl. Wr).” The oe bears no collection number; specimens at NY and US that appear to be part of the type collection bear the number 637. = Actinea Ai las Bigelowii] (A. Gray) Kuntze, Rev. Gen. PI. 1:303. 1891. = Macdougalia bigelovii (A. Gray) A. Heller, Bull. Torrey Bot. Club 25:629. 1898. = Actinea bigelovii (A. Gray) A. Nelson, Univ. — Publ. Sci., Bot. 1:139. 1926, nom. superflu. Ac ined ee ee A. Nelson, Univ. Wyoming Publ. Sci., Bot. 1:140. 1926. TYPE: U.S.A. ie no Co. “Rocky hillsides, among Yellow Pine Flagstaff Ariz.” (holotype label), 2 Jun 1922 Hanson 42 (HOLT YPE: RM-LO07 33! PROBABLE ISOTYPE: MO-895987)). Polycarpic perennials. Caudices sparingly branched, thickened distally, usu- ally encased ina stringy cocoon-like covering formed by the veins of the decay- ing leaf bases. Aerial stems 1-5, erect, usually unbranched distally, green throughout to purple-red-tinted distally to purple-red-tinted throughout, 20- 70cm, sparsely to densely pubescent, often tomentose proximally, eglandular or sparsely dotted with sessile glands. Leaves basal and cauline, alternate, lin- ear to linear-lanceolate to linear-oblanceolate, simple and entire or blades rarely divided into three segments, glabrous or sparsely to densely pubescent, eglandular or sparsely dotted with impressed glands; basal leaf bases expanded, clasping, persistent, sparsely to densely long-villous-woolly. Heads 1-5 per plant, usually borne singly but sometimes in paniculiform arrays. Peduncles (1.5-)6- 20(-29) cm, expanded apically, moderately to densely pubescent, densely to- mentose distally beneath the involucres, sparsely to moderately dotted with sessile glands. Involucres hemispheric to broadly campanulate, 13-20 x 23-32 mm. Phyllaries in 2 series; outer phyllaries 13-19, basally connate only slightly tol/Stheir lengths, green throughout or yellow to yellow-green proximally and green distally, often purple-red tinted on the margins, lanceolate to narrowly lanceolate or obovate to narrowly obovate, 7-11 x 1.3-2.8 mm, weakly to mod- erately keeled, apices acute to acuminate, abaxial faces sparsely to densely pu- bescent, sparsely to moderately dotted with sessile and impressed glands, adaxial faces glabrous or sparsely pubescent, eglandular or sparsely dotted with sessile glands; inner phyllaries 13-18, free, bodies yellow to yellow-green and scale-like, usually green distally, often purple-red tinted at the apices, narrowly BIERNER, TAXONOMY OF HYMENOXYS SUBGENUS MACDOUGALIA 661 Fic. 1. Known distribution of Hymenoxys bigelovii. lanceolate to narrowly obovate, 8.5-12.6 x (..1-)1.5-2.2 mm, distinctly surpass- ing the outer, not keeled or weakly to moderately keeled, apices aristate, abaxial faces glabrous or sparsely pubescent, eglandular, adaxial faces glabrous, eglandular. Ray florets 13-15, pistillate, fertile; corollas yellow, extending well beyond the phyllaries, 13-26 x 5.4-9.5 mm, lobes 3, abaxial faces glabrous or sparsely pubescent, eglandular, adaxial faces glabrous, eglandular. Disc florets 100-250+, bisexual, fertile; corollas yellow, cylindric to cylindric campanulate, 5.7-7.4 x 0.7-0.9 mm, lobes 5, glabrous or sparsely pubescent, eglandular. Re- ceptacles hemispheric to globoid to ovoid, paleae none. Cypselae narrowly obpyramidal, 4.2-4.7 x 0.9-1.2 mm, densely pubescent with straight, forked, antrorse hairs, eglandular; pappi 9-11(-15), obovate- to oblanceolate-aristate, 4.7- 7.3 x 0.7-1.3 mm. Chromosome number, 2n = Flowering and Distribution.—Flowering May to June. Roadsides, edges of juniper-pine and pine forests, 1375-2470 m. Central to eastern Arizona and western New Mexico (Fig. 1). Representative specimens examined. UNITED STATES. ARIZONA. Apache Co.: 7 mi N of hwy 264 on rd to Sawmill, 24 Jun 1965, Strother 402 (TEX). Coconino Co.: Hwy 89, ca 13 mi N of IH 40 Gct in Flagstaff) at turnoff to Sunset Crater National Monument, 20 May 1989, Bierner 89-25 (ARIZ, TEX); 7 mi NE of Strawberry, 29 May 1966, Lehto 6311 (ASU); 26 mi E of Strawberry, 2n = 30, 16 Jun 1947, Parker & McClintock 6851 (ARIZ); ca 10 mi S of Flagstaff on hwy 79, 14 Jun 1965, Strother 339 (TEX). Gila Co.: eee Pass, Mazatzal Mts, 15 May 1935, Collom 299 (ARIZ); 20 mi SW of Young, Sierra May 1947, Parker 531 (ARIZ); 5 mi N of Young, Pleasant Valley, 14 Jun 1947, Parker & aa k er tle TEX). Greenlee Co.: 20 mi N of Clifton, 7 Jun 1935, Maguire et al. 11865 (ARIZ). 662 BRIT.ORG/SIDA 21(2) Navajo Co.: Hwy 60, 12.8 mi SW of hwy 260 (jct in Show Low), 2n = 1511, 19 May 1988, Bierner 88-54 (ARIZ, TEX); Lakeside, White Mts, 9 Jun 1928, Harrison 5467 (ARIZ, LL). Pima Co.: 3 mi up Mt. Lem- mon Rd past entrance sign to Coronado Forest, 31 May 1967, Mears 1663b(TEX). NEW MEXICO. Catron Co.: just W of Luna, 23 Jun 19605, Strother 394 (TEX). Cibola Co.: Near Bluewater Canyon Dam, 12 mi S of Bluewater, 20 May 1936, Parker & Parker 3481(ARIZ); Grant Co.: near Santa Rita Copper Mines (Santa Rita del Cobre), 22 mi FE of Silver City, 18 May 1936, Parker & Parker 3452 (ARIZ). Note.—More than 70 specimens were examined for this treatment. Those listed above were chosen as representative of the geographic distribution and mor- phologic variation of Hymenoxys bigelovii. ACKNOWLEDGMENTS lam grateful to the following institutions for loans of specimens: ARIZ, ASU, LL, NY, RM, TEX. Lalso thank John Strother for his help with the description of Hymenoxys bigelovii and José Panero for his help with preparation of the Spanish abstract. Guy L. Nesom and A. Michael Powell provided helpful review comments. REFERENCES Beaman, J.H. and B.L. Turner. 1962. Chromosome numbers in Mexican and Guatemalan Compositae. Rhodora 64:271-276. Bierner, M.W. 2001. Taxonomy of Hymenoxys subgenus Picradenia and a conspectus of the subgenera of Hymenoxys (Asteraceae: Helenieae: Tetraneurinae). Lundellia 4:37-63. Bierner, M.W. and R.K. Jansen. 1998. Systematic implications of DNA restriction site variation in Hymenoxys and Tetraneuris (Asteraceae, Helenieae, Gaillardii Lundellia 1:17-26. Bierner, M.W. and B.L. Turner. 2003. Taxonomy of Tetraneuris (Asteraceae: Helenieae: Tetraneurinae). Lundellia 6:44—96. Cockerelt, TDA. 1904. The North American species of Hymenoxys. Bull. Torrey Bot. Club 31:461-509. Gray, A. 1853. Plantae Wrightianae Texano-Neo-Mexicanae: an account of a collection of plants made by Charles Wright, A.M., in an expedition from Texas to New Mexico, in the summer and autumn of 1849, with critical notices and characters of other new and interesting plants from adjacent regions, &c. 2:96. Smithsonian Institution, Washington. Gray, A.1883.1.Contributions to North American botany. |.Characters of new Compositae, with revisions of certain genera, and critical notes. Proc. Amer. Acad. Arts 18:1—73. Hever, A.A. 1898. New and interesting plants from western North America.—IV. Bull. Torrey Bot. Club 25:626-629. Karis, PO. and O. Ryoinc. 1994. Tribe Helenieae. In: K. Bremer, ed. Asteraceae: cladistics & Classification. Timber Press, Portland, Oregon. Pp. 521-558. Rosinson, H. 1981.A revision of the tribal and subtribal limits of the Heliantheae (Asteraceae). Smithsonian Contr. Bot. 51:1-102. Ryosere, PA. 1915.Macdougalia. North American Flora 34:109-110. The New York Botani- cal Garden, New York. BIERNER, TAXONOMY OF HYMENOXYS SUBGENUS MACDOUGALIA 663 SANDERSON, S.C. 1973. In: IOPB chromosome number reports XL. Taxon 22:285-291. Speese, B.M. and J.T. Batowin. 1952. Chromosomes of Hymenoxys. Amer. J. Bot. 39:685-688. SPRING, O., B. ZITTERELL-Haip, M.W. Bierner, and T.J.Masry. 1994. Chemistry of glandular trichomes in Hymenoxys and related genera. Biochem. Syst. Ecol. 22:171-195. STROTHER, J.L. 1966. Chromosome numbers in Hymenoxys (Compositae). Southw. Naturalist 11:223-227. Torrey, J.and A. Gray. 1842. A flora of North America: containing abridged descriptions of all the known indigenous and naturalized plants growing north of Mexico; arranged according to the natural system. 2(2):381-383.Wiley & Putnam, New York. Turner, B.L. and A.M. Powett. 1977. Helenieae-systematic review. In: V.H. Heywood, J.B. Harborne, and B.L.Turner, eds. The biology and chemistry of the Compositae. Academic Press, London. Pp.699-737. Turner, B.L., A.M. Powett,and TJ.Wartson. 1973.Chromosome bers in Mexican Asteraceae. Amer. J. Bot.60:592-596. Waener, H., M.A. lvencar, L. HorHammer, and W. Herz. 1972. Flavonol-3-glycosides in eight Hymenoxys species. Phytochemistry 11:3087-3088. 664 BRIT.ORG/SIDA 21(2) BOOK NOTICE Davip J. WisHart. 2004. Encyclopedia of the Great Plains. (ISBN 0-8032-4787-7, k.). The University of Nebraska Press, PO. Box 84555, Lincoln, NE 68501- 4555, U.S.A. (Orders: 402-472-3584, Fax: 402-472-6214, www.unp.unl.edu) $75.00, 919 pp., b/w photos, 8 1/2" x 11" Publisher Comments: “The Great Plains is a vast expanse of grasslands eras from the Rocky eco to the sansa River and from the Rio Grande to the coniferous forests of Canada—an area hundred miles from north to south and more than five ae miles from east to west. The Ge Plains region includes all or parts of Texas, New Mexico, Oklahoma, Kansas, Colorado, Nebraska, Wyoming, South Dakota, North Dakota, Montana, Alberta, Saskatchewan, and Manitoba. The region, once labeled “the Great American De is now more a ten called the “heart- and,” or, sometimes, “the breadbasket of the world.” Its i lands, sparse population, enveloping horizons, and dominating sky convey a sense of expansiveness, even empti- ness or loneliness, a reaction to too much space and one’s own ee laguna in it.” “With 1,316 entries contributed by more than one t {breaking ref- erence work nae ae is ees an eacene ss the Great Plains- fren its henicramsentel climate to its images ter, its folklore, and its politics. Thoroughly illus- trated, annota sted index ne hi kabl lium of information and oe will prove the definitive and indispensable resource on the au Plains for many years to ome.” What an impressive book this is. The 27 chapters range from African penne to Water with each chapter being introduced oO a major essay or synthesis of the topic. I explored the index to see what was listed under “Cotton.” There is a nice summary of Cotton oe mhirsutum)andan impressive b/w photo of a ea Fete in West Texas. | looked up to see what information was avail- able on the flora and there is a relatively short entry, yet a nice summary, provided under the sae ter, Physical Environment. “More than 2,900 sp f vascular cael from some 730 genera in 159 families grow in the Great Plains.” The actor of this entry cite e other than the Flora of the Great Plains 1986). at ipscomb, Botanical Research ane y Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A = SIDA 21(2): 664. 2004 ERIGERON PEREGRINUS AND ERIGERON GLACIALIS (ASTERACEAE: ASTEREAE) Guy L.Nesom Botanical Research Institute of Texas 09 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. gnesom@brit.org ABSTRACT a bs Be: 1 s . ] _ Fig Erigeronglacialis (Nutt.) A. Nels. (=E E. oe sli ex ue Greene. It has most recently been treated as a perenne subsp. F. glacialisand E. ; d but the two species are mostly distinct even wh incl ntact in Washin son nl British Columbia; intermediates are , uncommon or absent as E. gl lacialis at the Ve 1 edge of its sass is conneuqus or Cs inter- mittently sympatric with E. Alaska Cronq.) Nesom, comb. nov, is proposed in order to complete the al eiaes tron If Shanna recogni- tion of two subspecies within E. peregrinus is to be followed, the name E. pereg callianthemus oe is incorrect, the precedent for a name at that aie set by E. a a. angustif folius 19 RESUMEN a a acialis(Nutt. ) A. Nels. (= E t} p diferent de E. peregrinus (Bank RinSy Cee: Ha sido tratada recient t E. peregrinus subsp. illianthemus. Se encuentran i FE glacialis y E. peregrinus ee las dos especies son muy Mises incluso cuando estan en cua en Washington y en Columbia Britanica; los lacialisen el limite Oeste de su area vive contiguo o quizas L intermitentemente simpatrico con E. pee hacia el HONS pasta ee = propoue Erigeron glacialis var. hirsutus (Cronq.) Nesom, comb. nov, p Sise sigue el reconocimiento de Cronquist de dos subespecies en E. peregrinus, el nombre E. peregrinus op callianthemus (1943) es incorrecto, el precedente de un nombre en ese rango esta establecido por E. salsuginosus subsp. angustifolius 1906). A taxon previously known as Erigeron salsuginosus (Richards. ex R. Br.) A. Gray was united by Cronquist (1943) at subspecific rank with E. peregrinus (Banks ex Pursh) Greene. Cronquist observed that Erigeron (Aster) glacialis (Nutt.) A. Nels. is the oldest name at specific rank to replace the misapplied E. salsuginosus but that the type of E.callianthemus Greene is more representative of the taxon, thus he used the latter for the name at subspecific rank (see nomenclatural summary below). He later (1947) recognized several varieties within each subspecies. Erigeron peregrinus subsp. callianthemus is widely distributed in the mountains of the western U.S.A. and southwestern Canada, while E. peregrinus subsp. peregrinus occurs in coastal and near coastal habitats from southern Alaska to the northwestern conterminous U.S.A. Their ranges are largely dis- SIDA 21(2): 665-672. 2004 666 BRIT.ORG/SIDA 21(2) tinct (Fig. 1) but the occurrence of intermediates in Washington and British Columbia suggested to Cronquist (1947) that the two taxa were best treated within a single species, a taxonomic arrangement that has since been followed in accounts of North American Erigeron. Douglas et al. 1998, p. 252) also noted that the subspecies “intergrade” where ranges overlap. My observation, in con- trast, is that intermediates do occur but that even where they overlap or are in close contact in Washington and British Columbia, populations of ‘good’ subsp. peregrinus (without signs of intermediacy) and ‘good’ subsp. callianthemus are more common. The distribution of subsp. peregrinus runs from the Kamchatka area of the Russian Far East (Commander Islands: Botschantzev 1959; Czerepanov 1995) across the Aleutian Islands, southern-central Alaska, southwestern Yukon, and southward along the Alaskan-British Columbian archipelago into western Washington and northwest Oregon (Fig. l; more detailed maps in Hultén 1950, 1968; Cody 2000). From the Yukon region, it trails southward in a relatively thin line along the Pacific; as noted by Douglas et al. (1998), it is “common in and west of the Coast-Cascade Mountains, rare east of the Coast-Cascade Moun- tains.” Calder and Taylor (1968, p. 533) observed that in the Queen Charlotte Islands “a few collections lof subsp. peregrinus sensu stricto] show a tendency towards ssp. callianthemus as the involucral bracts are not conspicuously vil- lous and tend to be slightly glandular.” Subspecies callianthemus is widely distributed and relatively abundant in montane regions throughout the western U.S.A. as well as in southwestern Alberta and British Columbia. At the western edge of its range from British Columbia northward into Alaska, it is contiguous and perhaps intermittently sympatric with subsp. peregrinus. Subsp. callianthemus is absent from the Queen Charlotte Islands (Calder & Taylor 1968) and absent or rare in other in- sular portions of the Alaskan-British Columbian archipelago. It is rare in south- western Yukon and southeastern Alaska, apparently growing within the range of subsp. peregrinus. Scoggan (1979) recorded subsp. peregrinus for Alberta, but this probably is based on plants such as Breitung 16978 (BRIT) from Waterton Lakes National Park, which have aie stipitate-glandular and also sparsely villosulous on the proximal half, “determined by A. Cronquist” as E. peregrinus subsp. callianthemus var. scaposus. Typical subsp.callianthemus is the common form in the same region. In the U.S.A, subsp. peregrinus occurs in Whatcom, Skagit, Snohomish, King, and Pacific counties, Washington, and Clatsop Co., Oregon. Erigeron peregrinus var. thompsonii occurs in Grays Harbor Co., Washington. Typical subsp. callianthemus is known from Whatcom, Snohomish, and King coun- ties, and it also occurs in Clallam and Jefferson counties of the Olympic penin- sula, Washington, immediately north of Grays Harbor Co. In Oregon, it occurs in Tillamook Co., immediately south of Clatsop Co. a NESOM, ERIGERON PEREGRINUS AND ERIGERON GLACIALIS Oo Co A A ‘ ® ar gy ‘ a Ya Zia A Erigeron peregrinus © Erigeron glacialis 7 J F Jj = Bg Pe Pee ed errr glacialis g 668 BRIT.ORG/SIDA 21(2 In the view here, subsp. peregrinus and subsp. callianthemus are reason- ably treated as separate species, FE. peregrinus and E. glacialis, apparently occa- sionally hybridizing but not intergrading in a sense that would imply the ex- istence of a zone of intermediacy reflecting continuous gene exchange. The two species can be identified by the following morphological contrasts. Phyllaries eglandular or sparsely sessile-glandular at the apices, rarely sparsely dular over the surface, sparsely to moderately villous-hirsute on surfaces, margin Eri usta cllate rigeron peregrinus Phy y and evenly stipitate-glandular, without other hairs or rarely sparsely willow: on rene and margins of outer phyllaries Erigeron glacialis Cronquist (1955) noted other distinctions of E. peregrinus (vs. E. glacialis): leaves often toothed (vs. usually entire), often soft-pubescent (vs. usually glabrous), peduncular hairs rather loose (vs. close), but these are less diagnostic. Choice of specific over infraspecific rank in this case emphasizes three factors: (1) Reproductive isolation, although incomplete, apparently exists between E. glacialis and E. peregrinus, as noted above. (2) The distinction between E. glacialis and E. peregrinus is analogous to that between other closely related species of Erigeron, where relatively small but consistent and conspicuous differences in vestiture are significant (e.g. E. compactus-E. consimilis, E. ursinus-E. gracilis, E. caespitosus-E. abajoensis, E. tracyi(E. colomexicanus)-E. flagellaris, E.engelmannii-E. pumilus, and E. flettii- E.algidus-E. simplex/grandiflorus). (3) Erigeron glacialis may be as closely related to E. howellii (A. Gray) A. Gray as to E. peregrinus. These three species form a morphological unit and ap- parently are more closely related among themselves than to any other species. Erigeron howellii, which is essentially endemic to an area along the Columbia River in Oregon and Washin but otherwise glabrous stems (vs. stems strigillose, more densely so distally), more consistently spatulate basal and lower cauline leaves (vs. leaves linear- oblanceolate to broadly lanceolate or spatulate), consistently white rays (vs. rays blue to rose-purple or pink, less commonly white to pale blue), and habi- tats mostly at lower elevation. Erigeron howellii and E.glacialisare nearly iden- tical in involucral vestiture, and a reasonable hypothesis is that the narrow en- demic and E. glacialis are sister species. This is not intended as a “cladistic argument” for recognition of E. glacialis at specific rank, and it is clear that E. glacialis is more strongly differerentiated from E. howellii than from E. peregrinus, but it adds a line of evidence for consideration in this admittedly subjective decision concerning choice of rank. gton, differs from E. glacialis in its distally strigillose Erigeron peregrinus (Banks ex Pursh) Greene, Pittonia 3:166. 1897. Aster peregrinus Banks ex Pursh, Fl. Amer. Septent. 2:556. 1814. Type: U.S.A. ALASKA. Unalaschka, D. Nelson s.n. (HOLOTYPE: BM). NESOM, ERIGERON PEREGRINUS AND ERIGERON GLACIALIS 669 Phyllaries moderately to densely hirsute to villous-hirsute on the surfaces, margins ciliate; ray corollas purplish to pink or white; upland habitats Erigeron peregrinus var. peregrinus Phyllaries very sparsely villous-hirsute to glabrous on the surfaces, margins ciliate; ray corollas white; sphagnum bogs Erigeron peregrinus var. thompsonii Erigeron peregrinus (Banks ex Pursh) Greene var. peregrinus Erigeron unalaschkensis Less., Linnaea 6:122. 1831. Erigeron peregrinus (Banks ex Pursh) Greene var. dawsonii Greene, Pittonia 3:166. 1897. Erigeron peregrinus var. dawsonii was described from the Queen Charlotte Is- lands, where var. peregrinus is abundant. Calder and Taylor (1968) did not rec- ognize var. dawsonii, noting that populational variants include plants of both varieties and numerous intermediates. Cronquist (1955, p. 188) noted that “A phase of ssp. peregrinus, resembling var. peregrinus but perhaps properly to be segregated, occurs on Saddle Mt. in Clatsop Co., Oreg.” Chambers (pers. comm.) observes that these plants “com- bine the genetic traits of glacialis and peregrinus,” with villous hairs most abun- dant on the outer phyllaries, dense glandular indument on the inner phyllar- ies. They are perhaps “best interpreted as derived from a history of gene exchange between E. peregrinus and E. glacialis, along with segregation and recombination of the genes affecting the principal morphological differences in pubescence. ‘Good’ peregrinus has not yet been found in this part of the state, but ‘good’ glacialis is present nearby.” Erigeron peregrinus (Banks ex Pursh) Greene var. thompsonii (Blake ex J.W. son) Cronquist, Brittonia 6:144. 1947. Erigeron thompsonii Blake ex J.W. Thompson, Rhodora 34:238. 1932. TYPE: U.S.A. WASHINGTON. GRAYS HARBOR CO. open bog Pie Quinault, 10 Jul 1931, J.W. Thompson 7336 (HOLOTYPE: US! ISOTYPES: GHI, K, MOI, aC From details and wording of the protologue and description, it seems clear that Blake wrote both; he was not credited by Thompson, however, other than being cited as sole author of the name and the authorhip must be attributed to Thompson as “ex” rather than “in.” Cronquist (1947, p. 148) observed that var. thompsonii “is in a sense intermedi- ate between E. glacialis and E. peregrinus]and intergrades both ways.” The taxo- nomic status and evolutionary relationships of this taxon, which is endemic to a small area of the Olympic peninsula in western Washington, need to be in- vestigated in more detail. Erigeron glacialis (Nutt.) A. Nels., Bot. Gaz. 37:270. 1904. Aster glacialis Nutt., Trans. Amer. Philos. Soc. n. ser. 7:291. 1840. ear salsuginosus (Richards. ex R. Br.) A. Gray var. glacialis (Nutt.) A. Gray, Synopt. Fl. N r., pt. 2:209. 1884. TyPE: U.S.A. Nuttall’s protologue ObsEeSe at the aout of A. sacialiow was “with the preceding” species, Aster andinus Nutt. [ re 1 ind.) Nesom], which was noted to have been collected “on ‘he highest summits vor the Rocky Mountains, near the line of perpetual snow, in 42°. About ten thousand feet above the level of the sea. Near summit of Thornberg’s Ridge, where we made an ineffectual attempt to cross the Northern Andes, in August, still deeply buried in snow.” Gray saw the specimen—“Nutt!” as indicated in Torrey and Gray (1841, p.155) and later 670 BRIT.ORG/SIDA 21(2) noted (1884, p. 209) “first coll. by Nuttall in Wyoming.” According to Graustein (1967), it is likely that Nuttall’s group was on or near Hyndman Peak (Blaine Co, Idaho), at the place they called Thornberg’s Ridge or Thornberg’s Pass on 12-13 Aug 1834, this confirmed by the expedition narrative reproduced in McKelvey (1955). (HOLOTYPE: PH; HOLOTYPE fragment: CAS). Stems densely strigillose with loosely appressed, slightly crinkled hairs, most densely strigillose close beneath the heads; leaves glabrous to short- villous on both surfaces on glacialis var. glacialis Stems hirsute to hirsute-villous; leaves hirsute to hirsute- villous on tothe urfaces rigeron alacialls var. hirsutus Erigeron glacialis (Nutt.) A. Nels. var. glacialis Erigeroncallianthemus Greene, Leafl. Bot. Observ. Crit. 2:197. 1912. Erigeron peregrinus (Banks ex Pursh) Greene aes callianthemus (Greene) Cronq., Rhodora 45:264. 1943. Erigeron ursh) Greene var. cucallianthe misenc: Brittonia 6:145, 1947, Eri- geron peregrinus (Banks ex Pursh) Greene var. callianthemus (Greene) Crong,, Univ. Wash. Publ. Biol. 17(5):188. 1955 (in clave). Aster sdlsuginosus Ric jards. ex R. Br. var. angustifolius A. Gray, Bot. Calif. 1:325. 1876. Erigeron salsuginosus (Richards. ex R. Br.) A. Gray var. angustifolius (A. Gray) A. Gray, Proc. Amer. Aca Arts 16:93. 1880. Erigeron angustifolius (A. Gray) I Rydb, Bull. Torrey Bot. Club 24:295. 1897, Erigeron salsuginosus (Richards. ex R. Br.) A. Gray leg Ek ae Gray) Piper, Contr. x Pur US. Natl. Herb. 11:565. 1906. Erigeron peregrinus rsh) Greene var. angustifolius (A, és Crongq,., Brittonia 6:147. 1947. suginosus Richards. ex R. Br. var. scaposus Torr. & A. Gray, Fl. N. Amer, 2:503. 1841. Eri- geron peregrinus (Banks ex Pursh) Greene var. scaposus (Torr. & A. Gray) Cronq., Brittonia 6: 146.1947. E eas var. scaposus (Torr. & A. Gray) Breitung, Canad. Field- Naturalist 71:69. 1 Aster sa As earlier observed (Nesom 1992, p. 190), within Erigeron peregrinus subsp. callianthemus, var. scaposus and var. angustifolius can be recognized apart from var. callianthemus “only as arbitrarily distinguished and intergrading popula- tions.” Douglas et al. 1998) also noted that the varieties “often grow together and show a continuous variation.” Cronquist (1947, p. 148) regarded var. scaposus as a “reduced alpine phase” with the dwarfing “probably genetically controlled,” but he observed that it “intergrades profusely with var. eucallianthemus ... and both are often present in the same collection.” Var. angustifolius, also, was seen by Cronquist to be Scene inter grading with other expressions of the species. li lat infraspecific rank within E. peregrinus, the taxon E. peregrinus subsp. callianthemus (sensu Cronquist) includes all varietal taxa, but if the widespread entity identified by Cronquist as E. peregrinus var. callianthemus is interpreted to include either var. scaposus or var. angustifolius or both, it should be recognized that both latter names at va- rietal rank have precedence over var. callianthemus (var. scaposus the oldest). If any of these varieties are to be recognized within E. glacialis, the type of E. glacialis represents the alpine form treated as var. scaposus. In the taxonomic alternative proposed here, all three of these taxa are included within Erigeron glacialis var. glacialis. If Erigeronglaci NESOM, ERIGERON PEREGRINUS AND ERIGERON GLACIALIS 671 Further complicating the nomenclature is the observation that the earliest name at subspecific rank in this whole complex is Erigeron salsuginosus subsp. angustifolius (A. Gray) Piper, from 1915, rendering Cronquist’s combination in 1943 based on E. callianth incorrect (superfluous, because it included the type of “subsp. angustifolius”). Thus, if one desires to follow Cronquist’s concept of recognizing two subspecies within E. peregrinus, the one he treated as “subsp. callianthemus” requires a new combination, based on Gray’s original Aster salsuginosus var. angustifolius. Erigeron glacialis (Nutt.) A. Nels. var. hirsutus (Crong.) Nesom, comb. nov. Eri- geron peregrinus (Banks ex Pursh) Greene var. hirsutus Cronq,, Brittonia 6:147. 1947. TYPE: U.S.A. CALIFORNIA. [TUOLUMNE Co]: Yosemite National Park, vicinity of Lake Tenaya, 8300 ft, Jun 1902, H.M. Hall and E.C. Babcock 3506B (HOLOTYPE: UC). Variety hirsutus is restricted to the seven southernmost counties in the range of the species in California (Fresno, Inyo, Madera, Mariposa, Mono, Tulare, and Tuolumne cos.) and in Mineral Co., Nevada. I also have seen plants of typical E. glacialis and intergrades toward var. hirsutus from Mono, Tulare, Fresno, and Inyo cos., but they apparently are less common than those identified as typical var. hirsutus. ACKNOWLEDGMENTS I thank Bob Kiger for pointing out that Erigeron peregrinus var. callianthus is not the correct name when it includes var. scaposus and var. angustifolius (as | have used it in the past). His comments precipitated the presentation of this taxonomic alternative, which I have anticipated for more than a decade. Ken Chambers further pointed out that the name E. peregrinus subsp. callianthemus is incorrect, as it was preceded at that rank by “subsp. angustifolius.” Ron Hart- man gave advice on geography of the E. glacialis type collection, Kanchi Gan- dhi reviewed aspects of the nomenclature, James Macklin provided informa- tion on the type collection at PH, and Ken Chambers reviewed the whole manuscript and arranged a loan of pertinent specimens from OSU. Review com- ments by David Murray also were very helpful. REFERENCES BotscHaNntzey, V.P. 1959. Erigeron. Flora U.R.S.S. 25:191—288. Translated from Russian, 1999, Smithsonian Institution Libraries, Washington, D.C. Catoer, J.A.and R.L.Tayior. 1968. Flora of the Queen Charlotte Islands. Part 1:Systematics of the vascular plants. Canada Dept. of Agriculture, Monograph No. 4, Part 1. Ottawa, Canada. Copy, WJ. 2000. Flora of the Yukon Territory (ed. 2). NRC.CNRC, NRC Research Press, Ottawa. Cronauis, A. 1943. The identity of Aster salsuginosus. Rhodora 45:262-264. Cronauist, A. 1947.A revision of the North American species of Erigeron, north of Mexico. Brittonia 6: 121-302. 672 BRIT.ORG/SIDA 21(2) Cronauist, A. 1955. Vascular plants of the Pacific Northwest, Part 5: Compositae. Univ. of Washington Press, Seattle. CzerePANOV, S.K.1995.Vascular plants of Russia and adjacent states (the former USSR).Cam- bridge Univ. Press, Cambridge, England Douatas, G.W., G.B. Stratey, D. Meipincer, and J. Posar. 1998. Illustrated flora of British Colum- bia. Vol. 1, Gymnosperms and Dicotyledons (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks, Ministry of Forests. Victoria, British Columbia. GrausTEeiN, JE. 1967. Thomas Nuttall, naturalist: Explorations in America 1808-1841. Har- vard Univ. Press, Cambridge, Mass Hutten, E. 1950. Flora of Alaska and Yukon. Vol. X. C.W.K. Gleerup, Lund. Hutten, E. 1968. Flora of Alaska and neighboring territories. Stanford Univ. Press, Stanford, California. McKetvey, S.D. 1955. Botanical exploration of the trans-Mississippi West, 1790- nold Arboretum of Harvard University, Jamaica Plain, Mass. Nesom, G.L. 1992. Taxonomic notes on Frigeron (Asteraceae: Astereae) of California, Ne- vada, and Arizona. Phytologia 73:186—202. Scocaan, H.J. 1979. The flora of Canada. Part 4-Dicotyledoneae (Loasaceae to Compositae). Natl. Mus. Nat. Sci. Publ. Bot. No. 7(4). — 850. Ar- A NEW SPECIES OF ERIGERON (ASTERACEAE: ASTEREAE) FROM NORTHWESTERN CALIFORNIA Guy L.Nesom Thomas W. Nelson Botanical Research Institute of Texas 44 Fairway Drive 509 Pecan Street Eureka, are 95503-6413, U.S.A. Fort Worth, Texas 76102-4060, U.S.A. gnesom@brit.org ABSTRACT Erigeron maniopotamicus G.L. Nesom & T.W Nelson, sp. nov, a member of the E. eatonii alliance, ap- Ee is endemic to E50) -central Hum boldt com and ey geheeue Trinity County, Cali- ie robustior | FP I E. eatoniior E. lassenianus RESUMEN Erigeron maniopotamicus G_L. ee & os Nea ape nov, un puembre ae) grupo e eatonii, ] aparentemente es endémico del condado de Trinity, California. a simpatrico con E. robustior pero quizas esta mas cercanamente relacionado con EF. eatonii o E. lassenianus. In preparation of a taxonomic treatment of Erigeron by Nesom for Flora of North America, three collections (HSC) from Humboldt Co., California, appeared di- vergent from known taxa of the genus. During a study of the Frigeron eatonii A. Gray alliance, Strother (1987, by annotation) identified these collections as “FE. decumbens var. robustior vel aff.,” but he did not comment on them in a re- lated publication (Strother & Ferlatte 1988). Nesom (2004) noted that they might represent an undescribed entity. One of the three original HSC collec- tions was made by a coauthor of the current report (Nelson), and in order to investigate their identity, he subsequently made two additional collections of the same entity from the same area. These plants occur within the geographic range of E. robustior (Crong.) Nesom but are morphologically distinct. They are out of the known range of any other species of the E. eatonii alliance and are here formally described as a distinct species. Erigeron maniopotamicus G.L. Nesom & T.W. Nelson, sp. nov. (Fig. 1). Tyre: US.A. ALIFORNIA. Humboldt Co.: Board Camp Mt,, off Forest Service Road 49N38, 1 mi from jet with FS Road 1, T4N, R4E, Sec. 28, UTM 450779 E 438981, small, dry, rocky, open meadow, 4860 ft, — 2004, T.W. Nelson 9253 and S. Carothers (HOLOTYPE: HSC; ISOTYPEs: BRIT, NY, OSC, U Differta E. robustiore 1j lavil eet | Tete jori} foliis latioril Peer hl 7 : } minoribus, et phyllariis Aliptic -oblanceolatis vel oblongi a E. eatonii var. plantagineo caudicibus plerumque non ramosis, foliis cami non redactis, et SIDA 21(2): 673-678. 2004 674 BRIT.ORG/SIDA 21(2) y Avni Resonrs Inottute wy PLANTS OF THE NORTH COAST RANGES OF CALIFORNIA Erigeron maripslaenius Nesam & Uelssn Hu t County. T4N,R4E, Sec. Ur N “505979 E 438981. Elev. iséo'. “ Board Camp Mountain Road off Forest PAS Sa RR ee ae i Service Road 49N38 1 mile from junction wach Fo mane Service Road 1, ry, rocky, bare nous Alesem L Melsor * i sada rViae WW 1 polar v Evigeven Ut VAN EEE altess 18 June 2004 Thomas W. Nelson & Sydney Carothers ISOTYPE OF: Fic. 1. Habit of Erigeron maniopotamicus (isotype, BRIT). phyllariis elliptici-oblanceolatis vel oblongi-oblanceolatis abrupte acuminatis. Plants perennial, taprooted (roots [3-]5-8 mm thick); caudex usually simple, rarely with branches to 20 cm long. Stems (including peduncles) 2-6(-8), 10- 22(-27) cm long, basally ascending to decumbent-ascending, sparsely to densely strigose, eglandular, stems and leaves usually basally purplish. Leaves basal and cauline, basal persistent into anthesis, oblanceolate to spatulate-oblanceolate, NESOM AND NELSON, A NEW SPECIES OF ERIGERON FROM CALIFORNIA 675 3-10 cm long, (3-)5-12(-14) mm wide, strongly to weakly 3-nerved, margins entire, cauline gradually reduced distally or not, usually continuing relatively unreduced to near heads, elliptic-oblanceolate to elliptic-lanceolate, sometimes narrowly lanceolate or oblanceolate, hirsute-pilose to weakly pilose on both surfaces, eglandular. Heads l(-4) on peduncles 0.5-1(-5) cm long, held well be- yond leaves at peak anthesis, from branches near midstem or slightly more dis- tal; involucres (5-)6-7 mm high, 9-12(-14) mm wide (pressed), phyllaries in 2- 3(-4) equal to subequal series, elliptic-oblanceolate to oblong-oblanceolate, abruptly acuminate, usually each with an orange midnerve, with narrow scari- ous margins, hirsute-strigose to hirsute-pilose, densely villous at base, eglandular to sparsely glandular. Ray florets (16-)21-33, corollas 10-12 mm long, 1.5-2.5 mm wide, laminae white to pinkish or purplish, not coiling (or weakly so) or reflexing. Dise florets: corollas 3.2-3.8 mm long, throat not indurate or inflated. Cypselae 2-2.5 mm long, 2-nerved, sparsely strigose; pappus of 16-20 bristles 2.5-3 mm long, with a few outer setae ca. 0.1 mm lon Etymology, habitat, and phenology.—Known only from east-central Humboldt Co. and adjacent Trinity Co., California. The Mad River (whence the epithet) more or less dissects the geographic range of the species. Though the river’s name may have been meant to convey the madness (as “angriness”) of the torrents and rapids, the epithet here implies that the river is deranged (mad as “crazy”). There apparently is no psychological assessment of the subject, but we like the audile flow of the syllables. Populations of Erigeron maniopotamicus occur at elevations of 1350-1500 meters on a tan-colored, rocky, non-serpen- tine soil sharply distinct from typical regional soils surrounding the sites. The sites are relatively dry and might be described as “barrens” because few other plant species occur there, and they often are bordered by somewhat stunted woods of mixed conifer, mostly Douglas fir. Serpentine outcrops are scattered in the area, especially on ridges, but apparently do not influence the ‘erigeron barrens.’ Flowering June through August. Conservation significance.—Because of the limited distribution and few known collections of this species, a more detailed assessment of its distribu- tion and ee would be valuable. Additional collecti xamined: CALIFORNIA. Humboldt Co.: near Mad River Buttes, Pilot Creek Quad, 40° i 15", 123° 44' 41", meadow, 4739 ft, 28 Jun 1980, Baker 2479 (HSC), Baker 2499 (HSC); Jack Rabbit Valley, along Swayback Ridge 4-wheel drive road 1 mi S of jct with Forest Service road, Blocksburg Quad, TIN, R5E, NE 1/4 of NW 1/4, Sec. 26, meadow on metasediments, 4900 ft, 10 Jun 1980, Nelson and Nelson 5395 (HSC); along Swayback Ridge Road, which branches off Forest Service Road 1NO8 ca. 1/4 mi past gate, TIN, R5E, Sec. 23, dry open hillside, 4450 ft, 18 Jun 2004, Nelson 9249 and Carothers (BRIT, HSC, NY, OSC, RM, UC, US); Twin Lakes vicinity, Snow Camp and Twin Lakes trip, with Mr Murphy, frequent locally in = ane open places, 26 Jun 1951, Tracy 19581 (UC 223009, photocopy). Trinity Co.: Kitten-Pum [Kettenpom] Valley, 26 Jun 1893, Blaninenpten (UC 87679, photocopy). [Trinity Co. or Humboldt Co.|: Mad River, 1 Jul 1893, Blankinship s.n. (UC 87680, photo- copy). 676 BRIT.ORG/SIDA 21(2) Frigeron maniopotamicus occurs within the range of E. robustior, which is known from Humboldt, Trinity, and Mendocino cos. and is recorded as grow- ing in sagebrush-scrub, glades and meadows, and lower montane coniferous forest, sometimes in seeps and sometimes over serpentine, at elevations of (200- )700-1500 meters. In the range of E. maniopotamicus, E. robustior is not com- mon—it grows in loamy soil in openings and along edges of moist fir-oak woods and has not been observed in close proximity to E. maniopotamicus. No inter- mediates between E. maniopotamicus and E. robustior or any other species have been encountered in the present study. Populations of the new species are rela- tively large (over 200 plants) and are relatively uniform in morphology. The description of Erigeron decumbens subsp. robustior Cronq. by Cronquist (1947) (see Nesom 2004 for its treatment at specific rank) probably included measurements of E. maniopotamicus, judging from the low ranges of involucral size; one of the specimens that he cited as subsp. robustior, Blankinship s.n.-UC 87679, is identified here as E. maniopotamicus. Our com- parisons are primarily drawn from study of collections at NY and HSC. In his review of the present manuscript, John Strother also reviewed collections at UC and sent photocopies of two that are referable to the new species. Erigeron maniopotamicus and E. robustior are contrasted morphologically in the follow- ing couplet. a. Tpeer Reeve ICE, (3- ea 8mm es ale 10- 22( ck a basal leaves blanceolate to spatulate \reduced to near heads; involucres (5-)6-7 mm high, 9-12 mm wide; phyllaries lit nese to oblong-oblanceolate, abruptly acuminate Erigeron maniopotamicus a. ea ae val 2-3 orn ele ele (15- ie a cm OM, basal leaves lin- ced OF NONe Neal heads; ae 6- 85 mm high, (12-)14-20 mm wide; ohillatee narrowly oblanceolate to lanceolate, acute-acuminate rigeron robustior The Erigeron eatonii alliance (sensu Strother & Ferlatte 1988) is recognized by the following set of features: taprooted; caudex branches generally slender, plants not caespitose; stems erect or basally ascending to decumbent, some- times purplish at the base, proximal internodes not elongate; leaves basal and cauline, linear to oblanceolate, (1-)3-nerved; heads commonly more than one. Distinctions among the taxa often are subtle but discretely defined geographic ranges give confidence that the morphological differences reflect evolutionary differentiation. Erigeron eatonii comprises a group of contiguous varieties sometimes intergrading at points of contact (see Strother and Ferlatte 1988 for maps). Eri- geron eatonii var. villosus (Cronq.) Crong. and E. eatonii var. lavandulus Strother & Ferlatte are exceptions: var. villosus occurs north of all other varieties except var. lavandulus, which is sympatric with var. villosus, and one or both of these probably is justifiably treated at specific rank. Erigeron canaani Welsh occurs NESOM AND NELSON, A NEW SPECIES OF ERIGERON FROM CALIFORNIA 677 at the southwestern extreme of the range of E. eatonii var. eatonii and may be better treated at varietal rank within E. eatonii. Among other species of the al- liance, E. jonesii Cronq. and E. lassenianus Greene also occupy essentially allo- patric ranges; they are discontinous in morphology from contiguous taxa. The ranges of E. robustior and E. decumbens are relatively isolated on the western margin the alliance. There are no sympatric taxa of the Erigeron eatonii alliance that appear to have a sister relationship, and the sympatry of E. maniopotamicus and E. robustior suggests that their relationship also is more distant than “sister.” The closest relationship of the new species may be closer to E. eatonii itself, perhaps with E. eatonii var. plantagineus (Greene) Cronq., which is the segment of the species geographically closest to E. maniopotamicus. The closest approach of var. plantagineus to E. maniopotamicus is in the northeast corner of Sisk you Co. and Shasta Co. (Strother & Ferlatte 1988). The two taxa are similar in sizes of involucres, florets, and cypselae but differ conspicuously in habit, particularly in features of caudex and size and distribution of cauline leaves. Erigeron maniopotamicus might be treated at varietal rank within E. eatonii, but the nature of its relationship there would be ambiguous, and it is morphologically and geographically disjunct from var. plantagineus. The leafy stems of E. maniopotamicus are more like those of E. lassenianus (which approaches the range of E. maniopotamicus in northeastern Trinity Co.and Tehama Co.), and it is possible that E. maniopotamicus has genetic elements from that species and from E. eatonii. In any case, the choice here of taxonomic rank for E. maniopotamicus is admittedly somewhat arbitrary. Morphological contrasts between E. maniopotamicus and E. eatonii var. plantagineus are provided in the following couplet. a. Caudex branches common! ally slend line | lly strongly reduced in size from the basal, absent or reduced near heads; phyllaries narrowly lanceolate to narrowly oblanceolate, apically acute; pappus bristles 6-12__ Erigeron eatonii var plantagineus | J L |: | = | J J a Caudex us ally unbranc unreaucea to near rede phyllaries elliptic-oblanceolate to oblong: oblanceolate, apically abruptly acuminate; pappus bristles 16-20 Erigeron maniopotamicus To place Erigeron maniopotamicus in a broader perspective, the following key distinguishes all of the California taxa of the E. eatonii alliance. > Involucres 6-10.5 mm high, (12-)14-23 mm wide; phyllaries eglandular. 2. Involucres 6-8.5 mm high, (12-)14-20 mm wide; disc corollas 3.5-4.5 mm long; cypselae (1.8-)2-3.2 mm long Erigeren robustior 2. Involucres 7-10.5 mm high, (14-)17-23 mm wide; disc ae 6.8mm long ypselae 4-4.5 mm long eatonii var. nevadincols — . Involucres 4.2-5.6(-7) mm high, 6-12(-14) mm wide; ere genie eglandular. 3. Phyllaries densely minutely glandular Erigeron lassenianus 678 BRIT.ORG/SIDA 21(2) 3. Fea eaaeee eglandular, auline leaves continuing relatively unreduced to near heads; phyllaries el- ie ee to oblong-oblanceolate, apically abruptly acuminate Erigeron maniopotamicus 4. Cauline leaves absent or bracteate near heads; phyllaries narrowly lanceolate to specail oblanceolate, apically acute. pena Sieg uBHaNY absent or relatively short and thickened; involu- cres 8-12(-16) mm wide; disc corollas 3.5—5 mm;cypsela gas 3.5mm long Erigeron eatonii var. sonnei 5. Caudex branches commonly present, usually slender; involucres (9-)11- 12(-14) mm wide; disc corollas 3-4 mm long; cypselae 1.8-2.3 mm long Erigeron eatonii var. plantagineus ACKNOWLEDGMENTS This study originated from observations made possible by a loan to Nesom of the Erigeron collection at HSC. John Strother and Ken Chambers made con- structive comments that much improved the manuscript. Strother also searched UC-JEPS collections for possible additional material of E. maniopotamicus and sent photocopies of the Blankinship and Tracy collections cited here. Tiana Franklin and Amanda Neill (BRIT) provided the digital image. REFERENCES Nesom, G.L. 1992. Erigeron and Trimorpha (Asteraceae: Astereae) in Nevada. Phytologia 73:203-219. Nesom, G.L. 2004. Taxonomic reevaluations in North American Erigeron (Asteraceae: Astereae). Sida 21:19-40 STROTHER, J.L. and W.J. Fervarte. 1988.Review of Erigeron eatonii and allied taxa (Com positae: Astereae). Madrono 35:77-91. A NEW SPECIES OF SCUTELLARIA (LAMIACEAE) FROM GUERRERO, MEXICO Billie L. Turner James L. Reveal Plant Resources Center University of Maryland The University of Texas College Park, eet elite iia e SA. Austin, Texas, 78712, U.S.A. The New York Botanical Bronx, New York 10458. 5126, USA ABSTRACT A new species, Scutellaria petersonaie B.L. Turner & J.L. Reveal is described from the state of Guer- rero, Mexico. It is closely related to S. hintoniana of the section Crassipedes, but amply distinct. RESUMEN Se describe una nueva se ea poulelland pelesonne . L. porns & J.L. Reveal del estado i Guer- rero, México. Esta muy S.} Crassipedes, pero es diferente. Scutellaria petersoniae B.L. Turner & J.L. Reveal, sp. nov. (Fig. 1). Type: MEXICO. GUERRERO: Sierra Madre del Sur, along the Milpillas-Atoyac road via Puerto del Gallo, ca. 58 mi SW of Mexico Hwy 95, ca. 20.5 mi SW of Carrazal del Bravo and 1.8 mi NE of Yerba Santa in a mixed deciduous forest 17 Oct 1975, ].L. Reveal, K.M. Peterson, R.M. Harley,& CR. Broome 4282 (HOLOTYPE: TEX; ISOTYPES: to be distributed). Gimilic S 1] 1 : EN ee | } oe } } a F aT } 7: et laminis in petiolum gradatim Aeecessentibiis Ge laminarum abrupte petiolacum): Perennial herbs to 50 cm high, arising from fusiform tuberous roots. Primary stems much-branched from the base, moderately appressed-pubescent with upswept small hairs. Leaves opposite throughout, gradually reduced upwards, those at mid-stem mostly 3.0-4.5 mm long; petioles 0.5-1.2 cm long; blades ovate, undulate, somewhat dentate to nearly entire, gradually tapering upon the peti- oles, the upper surfaces moderately short-pubescent to glabrate, the lower sur- faces, venose, glandular-punctate, pubescent along the major veins. Flowers 2 at each of the uppermost several nodes. Pedicels 4-5 mm long. Calyx 5-6 mm long, 3-5 mm wide, pubescent like the stems. Corollas red, 3.0-3.5cm long, gla- brous within at the very base for ca. 3 mm, pubescent thereafter with down- swept hairs for ca.6 mm; upper lip 0.8-1.0 cm long; lower lip 0.4-0.8 cm long. Upper stamens exserted from the tube for 8-10 mm; filaments attached ca. 4 mm below the corolla’s orifice; anthers pale lavender, ca. 0.8 mm long. Mature nutlets not examined. Scutellaria petersoniae is obviously very closely related to S. hintoniana Epling (not to be confused with S. hintonianum Henrickson), differing mainly in vestiture and leaf shape. Scutellaria hintoniana is known only from the state SIDA 21(2): 679-681, 2004 BRIT.ORG/SIDA 21(2) UNIVERSITY OF Ex HERB, HORT. ROT, REG. KEW senteMlae in pedertonineg : | DA. Temes + TReveal " | is if — | P Ae Ll aff ZL 2 AE hI : SGA os JO 1 ' e % f P 4 ‘ Fic. 1, Scutellaria petersoniae, holotype. TURNER AND REVEAL, A NEW SPECIES OF SCUTELLARIA FROM MEXICO 681 — of Mexico in oak woodlands, while S. petersoniae is seemingly confined to east- ern Guerrero, an area well known for its amalgamation of unusual species. Etymology.—Scutellaria petersoniae commemorates Kathleen M. Peterson, gifted teacher and skilled botanist (p. 1239 of the current issue, Reveal 2004). With the present description, Epling’s previously monotypic section Crassipedes now contains two taxa, both confined to the Pacific slopes of west- ern Mexico. These two species are distinguished within the genus by their hab- its (rhizomatous herbs) and elongate, pubescent, red corollas. The following couplet should help distinguish the two taxa: Stems moderately to densely pilose with spreading hairs; blades abruptly petiolate S.hintoniana Stems moderately pubescent with short upswept appressed hairs; blades ee upon the petioles S. petersoniae ACKNOWLEDGMENTS Thanks to Tom Wendt for scanning the holotype, and to Gayle Turner for the Latin diagnosis. Guy Nesom and Richard Olmstead are thanked for their help- ful reviews. REFERENCE Eptina, C. 1942. The American species of Scutellaria. Univ. Calif. Publ. Bot. 20:1-141. Reveal, J.L. 2004. Kathleen M. Peterson, 1948-2003. Sida 21:1239-1243. 682 BRIT.ORG/SIDA 21(2) BOOK NOTICE Lavishly Illustrated Book on Tropical & Subtropical Trees MarGARET BARWICK. 2004. Tropical & Subtropical Trees an Encyclopedia. (0-88192- 661-2, hbk.). Timber Press Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $69.95, 484 pp., 2,305 illustrations, 1,981 in color, hardiness zone map, 8 1/4" x 11 5/8". Publisher Comments: “Margaret Barwick’s pine ean researched book represents the first comprehen- sive compendium of botanical and practical it this prev tes panies topic. The culmination of more than forty- ee years as a hands on ae and | | gner, the work reflects her passion for the subject and her appreciation of what fellow gardeners, both is and pro- fessional, a“ to aad about trees ieeneetOn for ‘tropical and subtropical climates.” “Li din detail with ere on the size, shape of the tree, its cultivation Hecifiremiente ornamental value, and rate of growth. The book is spe with the finer ei ne dows fruit, tha ge, bark, texture, and fragrance.” With text that is bot} to-read, the book is lavishly illustrated with more oe 2500 en many taken in the wild.” Each ene is represented by a type species, usually chosen as the most typical and well known of the genus. The type species is illustrated by photographs of the overall form, the flowers, fruit, bark and, i possible, the natural habitat. An incredible amount of concise information is summa- rized in sidebars for each t Z| roughout the book one will find fascinating facts about the plants (from trees that can kill t to love, sex, and beauty ).—Barney Lipscomb, Botanical Research Insti- tute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 21(2): 682. 2004 RECOGNITION OF PHRAGMITES AUSTRALIS SUBSP. AMERICANUS (POACEAE: ARUNDINOIDEAE) IN NORTH AMERICA: EVIDENCE FROM MORPHOLOGICAL AND GENETIC ANALYSES Kristin Saltonstall Paul M. Peterson and Robert J.Soreng Horn poigunaporataly Department of Bota ity aryland Center NaN Heald re istory C eau eataeaEl Science © Smithsonian Institution PO Box 775 Washington, District of Columbia (DC) 20013-7012, U.S.A. Cambridge, Maryland 21613, U.S.A. peterson.paul@nmnh.si.edu kristin.saltonstall@aya.yale.edu soreng.rob@nmnh.si.edu ABSTRACT f pI a +t IW ey ed . rary tal] PM Dor kr © is described. The ne new cabspecies: can be ee from the introduced and Gulf Coast North panei can lineages of P australis by having caducous leaf sheaths, ligules 1.0-L7 mm long, lower glumes .0-6.5 mm long, upper glumes 5.5-11.0 mm long, jn 8. ve Pe 5mm Mone: ane Py Possessing chlo- roplast DNA haplotypes of A-H,S, Z, AA. Then PCA analysis and when bivariate plots of the morphological eatin: are compared. Additional work is needed to morphologically distinguish the introduced from Gulf Coast lineages. i berlandieri is lectotypified. A key and distribution to the ae lineages are include L RESUMEN Se describe una nueva ue nativa ea er subsp. americanus Saltonstall, PM. Peterson & Soreng. I la Gulf Coast de Norte América de P.australis por aa vainas dics nee ae 1.0-L.7 mm, a inferiores de 3.0-6.5 mm, glumas superiores de 5.5-11.0 mm long, ie os 8.0-13. oe mm, ¥ por tener aia del DNA pase A-H, 5S, Z, AA. La nueva sub da en un a So L L 1 Pe | © 7 CATA As Cc / iL 6 : 16, - CAlAa: 1 : eas 1 | ul If Coast. Se lectotipifica Ph vague: berlandieri. Se ineiaven una clave y ape de distribucion de los tres linajes. Phragmites Adans. is a cosmopolitan genus found throughout the world and is currently placed in the tribe Arundineae with Arundo L.,, Hakonechloa Makino ex Honda, and Molinia Schrank, the latter three genera all introduced in North America (Soreng et al. 2004; Zuloaga et al. 2003). Phragmites is an erect peren- nial grass, 2-5 m tall, that can form dense stands. A number of species, subspe- cies, and varieties have historically been described in the genus Phragmitesand today four species are recognized: P. australis (Cav) Trin. ex Steud, P. karka (Retz.) Trin. ex Steud., P mauritianus Kunth, and P japonicus Steud. All temperate sub- species and varieties are now included under the designation P. australis (Clay- ton 1968). Using five specimens collected in Texas and Mexico, Fournier (1877) SIDA 21(2): 683-692. 2004 684 BRIT.ORG/SIDA 21(2) distinguished a North American Phragmites (P. berlandieri E. Fourn.) from that found elsewhere in the world. Based on measurements of glumes from 28 Euro- pean specimens and many North American specimens, Fernald (1932) sup- ported this distinction of a North American variety, P communis var. berlandieri (E. Fourn.) Fernald. Recent genetic studies indicate that three genetic lineages of Phragmites are found in North America (Saltonstall 2002, 2003a,b). A lineage endemic to North America is found across much of Canada and in the United States, from New England and the Mid-Atlantic states across to the Pacific coast and into the southwest (Fig. la). Regional structuring can be found within this native lineage, with east coast, midwestern, and western populations showing differ- ent chloroplast DNA haplotypes (A-H, S, Z, AA, Saltonstall 2003a). Another lineage is found in the southern United States from Florida across to the Gulf of California, and this lineage is also found in Central America and in Asia (Fig. Ib). It is characterized by chloroplast haplotype | (hereafter referred to as the Gulf Coast lineage). A third lineage, chloroplast haplotype M, is EurAsian in origin and was likely introduced to North America since European coloniza- tion. It is found across the continent, both in areas where Phragmites was his- torically present and also in places (such as the southeastern US) where Phragmites is not native to the flora (Fig. lc; Saltonstall 2002). Today, this intro- duced lineage is the most common type of Phragmites in North America and can be found in a variety of habitats including both brackish and freshwater marshes, inland fens, along the banks of rivers and lakes, and along roadsides. With the recognition that both native and introduced populations of Phragmites may be present, many state and regional management authorities are now revising their Phragmites management strategies with a goal of pre- serving native populations while controlling introduced ones. This has also encouraged Natural Heritage programs to consider listing native Phragmites as a rare or threatened plant ina number of states. However, the appropriate level of taxonomic classification of the different lineages has not yet been clarified. A number of qualitative characters have been suggested for distinguishing the native and introduced lineages including culm color, culm texture, and ad- herence of leaf sheaths to culms (Blossey 2002). While these characters appear to be correlated with ecological characteristics, they are problematic in that they are subject to observer judgment and may require observation at different times of the year. Although genetic testing can provide definitive information as to the lineage of a population, quantitative measurements of morphological features may provide a means of confirming origin in conjunction with quali- tative cl teristics. Robichaud and Catling (2003) performed such an analy- sis using Phragmites specimens collected in southern Ontario and found sig- nificant differences in the length of the lower glume between the native and }@n) SALTONSTALL ET AL., RECOGNITION OF 685 Sources Fic. 1. Distributi fa) Nati b)Gulf Coast, 1c) f Ph ling et al.2004, COMREED 1999, Saltonstall 2002. hee 4 introduced population lineages. However, that study was limited by its geo- graphic scope. This study quantifies differences in size seen in the ligules, lower and upper glumes, and lemmas of native, introduced, and Gulf Coast popula- tions of Phragmites from North America. We formally recognize the native lin- eage that occurs in the USA and Canada as P. australis subsp. americanus Saltonstall, RM. Peterson & Soreng. ‘Although not documented across the Gulf Coast except for in the Mississippi river delta (Saltonstall 2002), introduced Phragmites may already have invaded these regions and certainly has the potential to spread into them. The distribution of introduced Phragmites is not known south of the U.S. border and thus is not included in this figure. 686 BRIT.ORG/SIDA 21(2) METHODS The genetic lineage of specimens was determined prior to taking morphologi- cal measurements. DNA extractions were done using a CTAB extraction proto- col (Doyle & Dickson 1987). Lineages were identified either by sequencing two noncoding chloroplast gene regions, trnT(UGU)-trnL(UAA)S and rbcL-psal, to determine the chloroplast DNA haplotype (Saltonstall 2002) or using an RFLP diagnostic assay on the abovementioned chloroplast regions that distin- guishes the three North American Phragmites lineages (Saltonstall 2003c). Ligules were measured using a Nikon PZ500 dissecting microscope fitted with a 0.1-10 mm micrometer. All samples were obtained from live populations in 1999-2003 throughout the range of Phragmites in North America. Several leaf blades per sample were initially observed to see if there was variation in the ligule lengths. Since no within-plant or within-population variation was detected only one leaf blade was examined for the majority of specimens. Ligule length was calculated by measuring the ligule (both the membrane and its hairy margin) at the center of the leaf blade to the nearest 0.05 mm. Measurements of lower and upper glumes and lemmas were taken from a single inflorescence per clone. Ten glumes and ten lemmas were measured from each specimen. Samples were obtained from live populations during 1999-2003 or from herbarium specimens and cover the geographic range of all Phragmites lineages in North America. Measurements were made to the nearest 0.5 mm using a ruler. From each specimen, ten spikelets showing visible rachilla hairs were selected from the middle part of the inflorescence. Upper and lower glume and lemma lengths were measured from the articulated base to the tip. A com- plete data set of the morphological characters used in this study is available from KS upon request. Data were analyzed using the PROC MIXED procedure in SAS 8.2. Tukey’s comparisons were used to distinguish significant differences between popula- tion types. Since the majority of samples measured for ligule length were dif- ferent from those measured for glume and lemma lengths, the data were ran- domized and treated as groups representing variation within each of the three genetic lineages. Bivariate comparisons were plotted to illustrate these differ- ences between lineages. A Principal Components Analysis (PCA) was performed using PC-ORD (Version 4, McCune & Mefford 1999) using a correlation matrix of standardized data for the variables. RESULTS AND DISCUSSION The morphological characters measured in this study clearly distinguish na- tive from introduced and Gulf Coast Phragmites lineages. This mirrors the dis- tinctiveness seen at the genetic level between the lineages, where all native North American haplotypes shared five unique mutations not seen in any other haplo- types (Saltonstall 2002). Native specimens have longer ligules, glumes, and lem- SALTONSTALL ET AL., ACWUNTEIVIN VE 687 Taste 1.Mean values and their significance level for ligule, glume and lemma lengths by Phragmites lineages: Native (N), Introduced (I), and Gulf Coast (GC). Structure Population Type Sample size Mean + SE (mm) Significant Difference (p<0.01) Ligule Native 28 1.26 + 0.04 GC Introduced 20 0.69 + 0.03 N Gulf Coast 14 0.57 + 0.04 N Lower glume Native 28 46+0.1 ,GC Introduced 17 34+0.1 N, GC Gulf Coast 15 3.9+0.1 N,| Upper glume Native 28 7340.2 LGC Introduced 17 5.8 + 0.2 N Gulf Coast 15 6.3+0.1 N Lemma Native 28 11.1 40.2 GC Non-Native ie 9.2+0.2 N Gulf Coast 15 10.1 +0.2 N mas than both introduced and Gulf Coast specimens (Table 1; Ligule—F259=120.21, Lower glume—F257=37.59; Upper glume—F257=21.01; Llemma—F? 58=17.07, p<0.0001 for all comparisons). Of the four characters measured, the ligule is the most definitive in separating the native from the other two lineages (Fig. 2a-c). The length of the lower glume is also a good way of distinguishing native from introduced specimens, although some overlap is seen (Fig. 2d). The Gulf Coast lineage, although significantly different from others at sev- eral measurements, is intermediate between the other two types when com- paring these four characters (Table 1, Fig. 2a-d). Thus at this time, it remains difficult to distinguish morphologically and it appears premature to conclude that this lineage is a different species Jones et al. 1997). It appears more similar to introduced than native Phragmites for all morphological measurements, in addition to being genetically more closely related to the introduced haplotype M than the native haplotypes (Saltonstall 2002, 2003a). Additional characters that distinguish this lineage morphologically have yet to be identified. Although not verified quantitatively, the leaf internode distance of the Gulf Coast plants appears to be shorter than both the introduced and native lineages (Saltonstall pers. obs.). The syntype of P. berlandieri VL. Berlandier 1446, US-82049 ex P) was included in our morphological survey and falls within the Gulf Coast lin- eage. Fernald (1932) did not indicate if he used one of the syntypes designated by Fournier (1877) in his study. Clearly Fernald was referring to the native lin- eage in his study since the lower glumes range from 4-6 mm long and the upper glumes range from 6-8.5 mm long. To avoid confusion in the future, particu- larly if one chooses to use the name P. berlandieri to include the Gulf Coast lineage, we formally lectotypify P berlandieri E. Fourn., Bull. Soc. Bot. France Fig. 2 a) T= a era Oe aE 6 A E A a aka A 7 a, Aa -* ol ls feheg ROA AA D3. U8 o = 2 ° ald 1 (9 it ae ere Sven eevee eee 0 0.5 1 1.5 Ligule (mm) c) 14 ._____ 7 12 A = 0 eal . padard E . Hog 822 E = 6 a a4 2 QO + = = 0 0.5 1 1.5 Ligule (mm} Fic. 2. Bi t p f hological data for Native ( £ £ a £ 2 Do hi teh) Qa Qa i) 2 | | = E @o | iS | = | oD .— | feb) | Q | Qa | > | 2 ), Introduced ( Lower glume (mm) W) and Gulf Coast (C) Phragmites individuals. 889 (2)L@ WdIS/D¥O'LINa SALTONSTALL ET AL., RECOGNITION OF 689 Taste 2. Eigenvector loadings for the principal components (PC). Relative eigen values, percent of variance, and cumulative percent of variance are also listed. PC1 PC2 PC3 PC4 Ligul -0.4030 0.8859 0.2293 0.0143 Lower glume -0.5285 0.0584 0.7301 -0.4292 Upper glume -0.5489 0.2219 -0.0574 0.8038 L -0.5069 0.4032 0.6411 -0.4116 Eigenvalue 2.996 0.636 0.267 0.101 % of variance 74,904 15.905 6.676 2.515 Cumulative % of variance 74.904 90.809 97,485 100.000 24-178. 1877. Tyee: U.S.A. Texas: Laredo, 1828, J.L. Berlandier 1446 (LectoryeeE: P; ISOLECTOTYPE, the large specimen on the sheet that includes a culm with a com- plete inflorescence: US-82049 ex W!). The PCA confirmed and enhanced the above mentioned results further. The first two PC’s accounted for 90.8% of the total variation in the data (Table 2, Fig. 3). The first axis alone accounts for 74.9% of the total variance and has negative loadings for the majority of Native specimens and positive ones for introduced and Gulf Coast specimens. Thus more negative values along PCI indicate larger morphological structures (Fig. 2), as seen in the native specimens. Analysis of nuclear microsatellite DNA indicates that there is little evi- dence for hybridization between the native and introduced lineages since alle- les considered diagnostic for each of the two lineage types were rarely found in the alternative lineage. Further, this nuclear DNA dataset strongly supports the genetic differentiation seen in the chloroplast DNA between the native and in- troduced lineages (Saltonstall 2003b). Although Phragmites has been said to be self-incompatible (Gustaffson & Simak 1963), little is known about the mat- ing system of this genus and it is not known if hybrids between population types can occur. The morphological data clearly support separation of the na- tive lineage from the introduced/Gulf Coast lineages. We have demonstrated that the native lineage has morphological features (longer ligules, glumes, and lemmas) and unique genetic mutations that differentiate it from the introduced/ Gulf Coast lineages. We chose to recognize the native lineage formally as Pharagmites australis subsp. americanus. The following key using morphologi- cal and genotype features is given to separate these three lineages. KEY TO THE LINEAGES OF PHRAGMITES AUSTRALIS IN NORTH AMERICA 1. Ligules 1.0-1.7 mm long; lower glumes 3.0-6.5 mm long; upper glumes 5.5-1 1.0 mm long; lernmas 8.0-13.5 mm long; leaf sheaths caducous with age; culms ex- posed in the winter, smooth and shiny; rarely occurs in a monoculture; chloroplast DNA haplotypes A-H, S, Z, AA (see Saltonstall 2002, 2003a)_ P. australis subsp. americanus (Native lineage) BRIT.ORG/SIDA 21(2) 2 | 1 1 A a A Ho Ag mS. = A A a 6 o' Te) AA A a A K A on 0 8 O : mo A A A O i. Ay A Ne A as A -2 T T T T aa aa era | -6 -4 -2 0 2 4 Axis | lysis of morphological data for Phrag tralis: Native (_.), Introduced (MM) and Gulf Coast (C) individuals. 1. Ligules 0.4-0.9 mm long; lower glumes 2.5-5.0 mm long; upper glumes 4.5-7.5 mm long; lemmas 7.5-12.0 mm long; leaf sheaths not poet with age, culms not exposed in the winter, smooth and shiny or ei andn n occurs as a monoculture; chloroplast DNA haplotypes | o 2. Culms smooth and shiny; southern California, pone New Mexico, Texas to Florida, throughout Mexico and Central America; chloroplast DNA haplotype | Paustralis var. berlandieri (E. Fourn.) C.F. Reed oe Coast lineage 2.Culms ridged and not shiny; southern Canada from British Columbia to Que south throughout the Continental United States; chloroplast DNA haplot P australis ipo lineage) Nae) Phragmites australis subsp. americanus Saltonstall, PM. Peterson & Soreng, SP. NOV. TyPe: U.S.A. MONTANA. Fergus Co. near the mouth of Dog Creek, 12 Sep 1883, Frank Lamson Scribner 378 (HOLOTYPE: US-824621)). A Phragmite australi (Cav.) Trin.ex Steud. vagina caduca cum aetate, ligulis 1.0-1.7 mm longis, glumis inferioribus 3.0-6.5 mm longis, glumis superioribus 5.5-11.0 mm longis, lemmatibus 8.0-13.5 mm longis, recedit. SALTONSTALL ET AL., RECOGNITION OF 691 Plants usually do not occur as a monoculture. Culms exposed in the winter, smooth and shiny, sometimes purplish at the nodes and internodes. Leaf sheaths caducous with age; ligules L.0-L7 mm mm long. Spikelet lower glumes 3.0-6.5 mm long, upper glumes 5.5-11.0 mm long; lemmas 8.0-13.5 mm long. Distribution —(Fig. la). This subspecies is known to occur in southwestern Northwest Territories east and south to California, Arizona, New Mexico, and east to northern Texas, Oklahoma, northern Arkansas, West Virginia and North Carolina, and north to Newfoundland and Quebec. Specimens examined (included in the genetic and morphological data sets): CANADA. BRITISH COLUMBIA: Osoyors Lake, J. Grant s.n. (US-2432752). UNITED STATES. COLORADO: La Salle, PA. Rydberg 2511 (US-908102). IOWA. Fayette Co.: B. Fink se? 230468). IDAHO: St. Anthony, E.D. Merrill@ E.N. ie ha ee INDIANA. Fulton Co.: W of Rochester, C.C. Deam 30010 (US- 1062053). KA watomie Co.: J.B.S. Norton 922 ae ae MAINE: Lake Anagunticook, Harford, J.C. sae re (Us $8068 MICHIGAN. Allegan Co.: Kalamazoo River near Douglas, WF Wright 125 (US-430189). MINNESOTA: ke Mellissa, H.L. Boll Seoue 908078). MONTANA: banks of the Missouri River, FL. Scribner a cs 153245). NORTH DAKOTA. Banson Co.: Leeds, J. Lunell s.n. (US-898853); NEBRASKA. Thomas Co.: Sand Hills near Plummer Ford, PA. Rydberg 1631 (US- 207984). NEW JERSEY: New Durham, W.M. Van Sickle s.n. (US- Ane NEW MEXICO: Dremionds Ranch near Roswell, J.D. Tinsley 12 (US- eae NEVADA. Nye Co.: Amargosa Drainage Basin, J.C. antes te (US-2876499). OKLAHOMA: E of Woodward, H.E. eee 1030 (1722877). OREGON: Klam . Klammoth Lake, E.I. pen 813 (US-273602). SOUTH DAKOTA: Canning, D. cis 10 0 908084). UTAH: Rabbit Valley, L.F Ward 534 (US-153247). WASHINGTON. Okanogan s of the Okanogan River, A.D.E. Elmer 519 (US-352294). WYOMING. Fremont Co.: Musk- =a Creek, L.O. Gooding 519 (US- 899997), ACKNOWLEDGMENTS We thank Susan Pennington for help with the literature cited and specimen citations, Dan Nicolson and Alain Touwaide for correcting the Latin diagnosis; and Bernd Blossey, J.K. Wipff and an anonymous reviewer for their comments on the manuscript. This is Contribution 3787 from the University of Maryland Center for Environmental Sciences. REFERENCES Biossey, B. 2002. Phragmites: common reed. Morphological differences between native and introduced genotypes. Accessed June 20, 2004. http://www. invasiveplants.net/ I PS Vee t y I CATLING, P-M., G. Mitrow, L. Black, and S. Caren. 2004. Status of the alien race of common reed Phragmites australis) in the Canadian maritime provinces. Bot. Electronic News 324. Crayton, W.D. 1968. The correct name of the common reed. Taxon 17:168-169. Comreed. 1999. COMREED Database. E. Haber [ed.]. National Botanical Services, Ottawa, ON, Canada. Accessed June 20,2004. 24.43.80.21/nbs/IPCAN/reedmap.htm| Dovte, JJ. and E.E. Dickson. 1987. Preservation of plant samples for DNA restriction endo- nuclease analysis. Taxon 36:715-722. — 692 BRIT.ORG/SIDA 21(2) FerNato, M.L. 1932. Phragmites communis Trin. var. Berlandieri (Fournier), comb.nov. Rhodora 34:211-212. Fournier, M.E. 1877.Sur les Arundinacées du Mexique. Bull. Soc. Bot. France 24:177-182. Gustarsson, A.and M. Simak. 1963. X-ray photography and seed sterility in Phragmites com- munis Trin. Hereditas 49:442—450, Jones, S.D.,J.K. Wiper, and PM. Montcomery. 1997.Vascular plants of Texas. University of Texas Press, Austin. McCune, B.and M.J.Merrorb. 1999. PC-ORD. Multivariate analysis of ecological data.Version 4. MJM Software Design, Gleneden Beach, OR. RosicHaup, L.and PM. Car.ine. 2003. Potential value of the first glume length in differentiating native and alien races of common reed, Phragmites australis. Bot. Electronic News 310. SALTONSTALL, K. 2002. Cryptic invasion by a non-native genotype of the common reed, Phragmites australis, into North America. Proc. Natl. Acad. U.S.A. 99:2445-2449, SALTONSTALL, K.2003a.Genetic variation among North American Populations of Phragmites australis: implications for management. Estuaries 26:444-451, SALTONSTALL, K.2003b. Microsatellite variation within and among North American lineages of Phragmites australis. Mol. Ecol. 12:1689-1702. SALTONSTALL, K. 2003c. A rapid method for identifying the origin of North American Phragmites populations using RFLP analysis. Wetlands 23:1043-1047. SoreNG, R.J., G. Daviose, PM. Peterson, F.O. Zutoaca, E.J. Juoziewicz, T.S. Fitaueiras, and O. Morrone. 2004 (23 March). Suprageneric classification in catalogue of New World grasses (Poaceae). http://mobot.mobot.org/W3T/Search/nwgc.html (originally published in 2000). ZULOAGA, F.O., O. Morrone, G. Dawinse, T.S. Ficcueiras, PM. Peterson, RJ. Sorena, and E.J. Juoziewicz. 2003. Catalogue of New World grasses (Poaceae): Ill. subfamilies Panicoideae, Aristidoideae, Arundinoideae, and Danthonioideae. Contr. U.S. Natl. Herb. 46:1-662. LAS ESPECIES DE NOTOTRICHE (MAIVACEAE) DE ECUADOR Magda Chanco Carmen Ulloa Ulloa Herbario San Marcos, M de Historia Natural Missouri Botanical Garden Universidad Nacional Mayor de San Marcos Apartado 14-0434 Lima, PERU St. Louis, Missouri 63166-0299, U.S.A. mchancoe@unmsm.edu.pe armen.ulloa@mobot.org ABSTRACT Four species of Nototriche are recognized for Ecuador, all are poe N. hartwegii is re-established. A key, descriptions, and illustrations of the species are pro RESUMEN Se reconocen cuatro ee de Nototric ne para el Ecuador, todas endémicas. Se restablece N. de las especies. hartwegii.S £ Nototriche Turcz., es un género netamente andino sudamericano Giese distribuye en el paramo de Ecuador y punas de Pert, Chile, Bolivia y norte de Argentina. Comprende alrededor de 130 especies distribuidas entre los 3500 y 5000 metros de elevacion. La mayor concentracion de especies se encuentra en Pert con aproximadamente 65 especies. La monografia del género de hace mas de un siglo (Hill, 1909) presento 62 especies. Varios trabajos Petes de Krapovickas (1950, 1951, 1953, 1957a, 1957b, 1973), Fryxell (1990) y Chanco (1992) han aumentando el numero de especies conocidas para el género y una nueva monografia actualizada para la serie Flora Neotropica esta en camino (Krapovickas y Chanco, en prep.). En la monografia de la familia para el Ecuador, Fryxell (1992) presento tres especies: N. ecuadoriensis Fryxell, N. jamesonii A.W. Hill y N. phyllanthos (Cav.) A.W. Hill, las mismas que fueron registradas por Jorgensen y Leon-Yanez (1999). Recientes colecciones realizadas en el sur del Ecuador indican la presencia de una cuarta especie, N. hartwegii A.W. Hill. Esta fue considerada por Fryxell (1992) como un sinénimo de N. jamesonii, pero nuestros estudios del material tipo indican que se trata de una entidad distinta, muy diferente del resto de colecciones de N. jamesonii. En este trabajo se restablece la especie N. hartwegti A.W. Hill para la flora de Ecuador, que actual te cuenta con Cuatro especies endémicas, duplicando el numero de endémicas registrado por Valencia et al. (2000). Ademas se proporciona su estado de conservacion segun las categorias de la UICN (2001). Las especies de Notot riche son hierbas de apariencia acaulescente, que forman cojines compactos de hojas densamente arrosetadas en las ramificaciones apicales de los tallos. Los tallos son por lo general subterraneos, profundos, robustos, lenosos y estan recubiertos por los restos de las hojas. Las hojas son SIDA 21(2): 693-703. 2004 694 BRIT.ORG/SIDA 21(2) flabeladas a ovadas, palmati o pinnatipartidas, glabrasad bi de tricomas estrellados; parte de las estipulas y el peciolo estan adheridos y forman una vaina o vagina laminar, membranosa. La porcion libre del peciolo y de las estipulas varia segun las especies. Las flores solitarias con pedicelo muy corto se insertan en el punto o debajo donde se separan los apices libres de las estipulas, carecen de involucelo; el caliz es gamosépalo, partido en 5 dientes; la corola es vistosa, consiste de 5 pétalos obovados; el androceo es monadelfo con una columna estaminal estrecha, con las numerosas anteras que forman una cabezuela en su apice; los estilos son hasta 14, mas largos que la columna estaminal, los estigmas son clavados. Los frut pos, variadamente hirsutos, dehiscentes, compuestos de hasta 14 meticarpos: cada uno con arista apical larga o corta. Todas las especies crecen en las partes mas altas de los Andes en habitats rocosos y arenosos sobre los 3500 m, hasta mas de 5000 m de elevacion. Dos especies, N. pinnata (Cav.) A.W. Hill y N.acaulis (Cav.) Krapov, tipificadas por colecciones realizadas, segun las descripciones, por Née en el volcan Chim- borazo, provienen probablemente de Pert, ya que de otra manera no se conocen en Ecuador. CLAVE PARA DETERMINAR LAS ESPECIES 1. Hojas glabras pto los apices de los I6bulos con uno o més cilios largos, lamina eee con 20 o mas lobulos ligulados; caliz glabro por fuera, menos los es y el nervio medio de los dientes con algunos cilios, viloso en los dientes por p dentio Nototriche e arena aes Fryxell bulada 1.H } | | fi-i treeless LS lo con menos de 20 ldbulos obtusos; caliz densamente estrellado-tomentoso en es superfici 2. Superficie ee de la lamina, estipulas y peciolo glabra o esparcidamente pubescente, vernicosa; superficie superior densamente tomentosa Nototriche phyllanthos (Cav. ) i Hill 2. ei ea e inferior de la lamina, estipulas y pecfolo densamente caer 3: eee oa: vilosa con pelos largos y sedosos; parte libre del Lee de 2~3 mm de largo; ramas aéreas aes formando a dros de hasta 15 cm de largo x 4—5 cm de diametr pear pale AW. Hill 3. Lamina estrellado-velutina con ee cortos; parte libre =e ee de e largo; ramas aéreas hojosas formando rosetas laxas de hasta 5 nae x 5 cm de ancho No Seg canon AW. Hill Nototriche ecuadoriensis Fryxell, Contr. Univ. Michigan Herb. 17:164. 1990. (Figs. 1 TIPO: EC vas eolopaay apo, road San Miguel (Salcedo), Puert oe | m San Miguel (Cerro Verde Filo), bunch grass paramo z coand rocky escarpment (78°25 WO 50) S) 3050- 4050 m, | Oct 1976, Ollgaard & Balslev 9929 (HOLOTIPO: AAUE IsoTiPos: Fl, MO!, NY!). Hierba perenne, acaulescente, arrosetada o formando cojines compactos; eje subterraneo lefioso, ramificado, en cuyos apices de las ramas se agrupan las hojas 695 F Fic. 1A | Bey i ae | Pa eee lik Jnl rete] ae tt 1, ] AyE, N. ecuadoriensis. B y F, N. hartwegii.€ y G, N. jamesonii.D y H, N. phyllanthos. formando rosetas de hasta 4,5 cm de diametro. Peciolo y estipulas forman una vagina de 8-11 mm de largo x 3-4 mm de ancho, glabra. Parte libre de las estipulas, ancha, obtusa, de 5 mm de largo x 3 mm de ancho, las superficies glabras, los apices con algunos cilios. Parte libre del peciolo glabro, de 1-2 mm de largo. Lamina pinnatipartida, de 8-12 mm de largo x 7-10 mm de ancho, los lobulos de la lamina con divisiones de segundo orden forman ldbulos ligulados de apice subagudo, ambas superficies totalmente glabras excepto los apices de los lébulos con 1 6 mas cilios. Caliz de 9-12 mm de largo, los dientes triangu- lares, de 4,5 mm de largo x 3mm de anchoen la base, el exterior glabro 0 cercano a ellocon algunos pelos largos y suaves en el nervio medio y en los bordes de los dientes, el interior glabro menos los dientes con pelos largos dirigidos hacia el Apice. Nectarios internos basales 5, aislados, mas anchos que altos, ca. 1 mm de ancho. Corola glabra, de un violeta palido (purptreo en seco); tubo de la corola de 4 mm de largo x 2 mm de ancho. Pétalos de 10-12 mm de largo x 5,5-6 mm de ancho, con una pequena escotadura apical; cada pétalo se continua sobre el tubo de la corola formando dos pequenas “alas” ca. 2,5 mm de largo. Tubo estaminal glabro, mas corto que los pétalos, de 4-6 mm de largo, las anteras dispuestas en una cabezuela globosa, subsésiles de color violaceo. Estilos 7 apenas sobresalen de las anteras, los estigmas capitados. Mericarpos de 6 mm 696 BRIT.ORG/SIDA 21(2) de largo (incluidas las aristas de 3 mm) x 3mm de ancho, con el dorso y el apice largamente hirsutos. Material adicional estudiado: ECUADOR. Pichincha: Slopes north-west of north peak of saan 4400 m, 00.308 078.00W, PJ. Grubb et al. 578 (K, NY). Napo: Paramo de Papallacta, sector El P 4060 m, 28 Oct 1984, Freire 33 (QCA). Distribucién y conservacion.—Especie endémica de la Cordillera Oriental, en los paramos del volcan Antisana, Papallacta y Cerro Verde, entre 3950 y 4400 m. Esta especie es conocida solo de cuatro poblaciones y se la considera como Vulnerable (VU). Notrotriche ecuadoriensis se distingue del resto de especies ecuatorianas por tener las hojas totalmente glabras excepto algunos cilios en el apice de los lobulos y por la lamina pinnatipartida con lobulos ligulados. Por la presencia de un tubo de la corola es afin a N. jamesonii, se diferencia porque esta ultima tiene la lamina flabeliforme con lobulos obtusos y por otros caracteres indicados en la clave. Nototriche hartwegii A.W. Hill, Trans. Linn. Soc. London 7:221-222, pl. 29, fig. 14, 1909. (Figs. 1B, F y 2). Tipo: ECUADOR: CANAR: “Monte Assuay ad Las Cruces,” 4550 m, sees 918 (HOLOTIPO Ki IsoTIPOS: BM (como foto digital), CGE n.v., G como foto digital, NY!, P Hierba perenne, acaulescente, pulviniforme, lanosa; eje subterraneo lenoso, ramificado, en cuyos apices de las ramas se agrupan numerosas hojas formando masas compactas cilindricas. Estipulas y peciolo forman una vagina de 6-7 mm de largo x 3-4 mm de ancho. Parte libre de las estipulas linear, de apice subagudo, de 5-9 mm de largo x 0,7-1 mm de ancho. Parte libre del peciolo de 2-3 mm de largo. Peciolo, estipulas y vagina densamente cubiertos por pelos estrellados lar- 20 y suaves en ambas superticies, en los margenes con pelos estrellados pedi- Lamina flabelada de base cuneiforme, de 10-15 mm de largo x 9-18 mm de ancho en la parte superior, 3-partida, cada segmento se divide en 3 o mas lobulos, el segmento medio entero o poco dividido, los lobulos oblongos de apice obtuso, la superficie superior densamente pilosa con pelos estrellados lar- gos y suaves (lanosa), la inferior igualmente pilosa pero en menor proporcion. Caliz de 15-20 mm de largo, los dientes de 11-15 mm de largo x 2,5-5 mm de ancho en la base, el exterior todo piloso con pelos similares a la hoja, el interior glabro menos los apices de los dientes. Nectarios internos basales 5, aislados, triangu- lares, de 1,5 mm de alto x 2 mm de ancho. Corola violacea con base blanquecina; tubo de la corola ausente. Pétalos de 20-25 mm de largo x 10-13 mm de ancho, la base con pelos estrellados largos. Tubo estaminal de 12-15 mm de largo, las anteras sésiles reunidas en cabezuela alargada, 2/3 del tubo estaminal, con pelos estrellados largos y caedizos. Estilos y estigmas en numero de 12. Mericarpos 12, uniovulados, de 9 mm de largo (incluidas las aristas de 5 mm) x 3mm de ancho, con pelos estrellados cortos en el dorso y apices con cilios largos. 697 ae 5 J Pv la planta en cojin 698 BRIT.ORG/SIDA 21(2) Material adicional estudiado: ECUADOR: “Quito,” Jameson s.n.(K). Azuay: Parque Nacional Cajas. Road Cuenca-Sayausi-Molleturo, 4110-4350 m, 4 Ene 2000, Jorgensen et al. 2114 (MO), Parque Nacional ile sendero paragiiillas, 21 Nov 2000, 4100-4400 m, Jorgensen et al. 2422 (MO), 13 Ene 2003, 4100- 180 m, Ulloa et al. 1184 (MO, USM). Distribucién y conservacién.—Endémica de los paramos del sur del pais entre 4100 y 4400 m en Canar y Azuay (la coleccion de Jameson tiene una localidad como “Quito” que no necesariamente corresponde a la provincia de Pichincha). Esta especie fue recientemente recolectada luego de 150 anos. Si bien hoy en dia la especie se encuentra protegida dentro del Parque Nacional Cajas, se conoce solo una poblacion y su area de distribucién es muy pequena por lo que se le considera como En Peligro (EN). Nototriche hartwegii se distingue por la lamina flabelada con base cuneiforme y poco lobulada; por las numerosas hojas agrupadas en los apices de las ramas formando cilindros compactos; y por la densidad de pelos largos estrellados (lanosos) que dan a toda la planta una apariencia verde-gris. Nototriche jamesonii A.W. Hill, Trans. Linn. Soc. London 7:228. 1909. (Figs. 1C, G). Tipo: ECUADOR: “Quito,” Jameson 154 (LECTOTIPO: Ki ISOLECTOTIPOS: G n.v. (foto F-neg. 23747 en F MO, NY), GH n.v). Nototriche chimborazoensis acne Candollea 16:82. 1957. Tipo: ECUADOR: Chimborazo, 4700- 5100 m. Rauh & Hirsch E-3 G)). Hierba perenne, a ee eje subterraneo lenoso, ramificado con los extremos apicales cubiertos de hojas dispuestas en rosetas. Peciolo y estipulas forman una vagina herbacea de 6-7 mm de largo x 2,5 mm de ancho. Parte libre de las estipulas linear-oblonga, de 3-5 mm de largo x 0,8-1,2 mm de ancho. Parte libre del peciolo de 6-19 mm de largo x 1,2-1,5 mm de ancho. Peciolo, estipulas y vagina con superficies pilosas (la vagina con la superficie inferior glabra), con pelos estrellados cortos, los margenes con pelos largos ciliados. Lamina flabeliforme, de 7-9 mm de largo x 9-11 mm de ancho, 3-fida, segmento medio mas desarrollado que los laterales, los segmentos divididos en 3 6 mas lobulos oblongo-obtusos; superficie superior de la lamina pilosa velutina, con pelos estrellados cortos, la superficie inferior igualmente pilosa pero menos densa que la superior. Caliz campanulado, de 8-11 mm de largo, dividido hasta cerca de la mitad en 5 dientes de apice obtuso, exteriormente todo piloso con pelos estrellados similares a los de la lamina, el interior glabro menos los apices de los dientes con pelos estrellados mas grandes. Nectarios internos basales 5, mas anchos que altos, cerca de 1 mm de ancho x 0,8 mm de alto. Corola violacea: tubo de la corola de 2,5-4 mm de largo, piloso. Pétalos de 15-18 mm de largo x 7-9 mm de ancho, la base con pelos estrellados. Tubo estaminal de 8-10 mm de largo, piloso con pelos algo rigidos y esparcidos, las anteras subsésiles dispuestas en una cabezuela oblonga. Estilos y estigmas cerca de 10. Mericarpos 10, de 6 mm de largo (incluidas las aristas de 2 mm de largo) x 2,5 mm de ancho, el dorso con pelos estrellados pequenos, parte superior y aristas con pelos estrellados largos y ciliados. Material adicional estudiado: ECUADOR: Sin localidad, “14.000 ft,” Pearce s.n. (K). Bolivar: Volcan Chimborazo, wind-swept gravel paramo, with open cushion plant association, 4200 m, 01.285 078.55W, 04 Ago 1979, ante Nielsen 18745 (AAU, F, MO, NY); Superparamo area W of volcan Chimborazo, ca. 33 km N of Guaranda, 4120 m, 01.305 078.56W, 29 Nov 1989, ee 13397 (NY, QCA), 16 Ene 1985, Luteyn & Cotton 11080 (NY, QCA); Chimborazo, sin col. 4200 m (LIL). Chimborazo: aie Chimborazo prope Totorillas ad nives perpect,” 4600 m, 7 Jul 1876, cee 3948 (K, NY(2)), 4200 m, André 3953 (K); Southern slope of Mount Chimborazo, sandy ground, 4600 m, 18 Jul 1939, Asplund 8379 (F, LIL, NY, P), 4260 m, 23 Jun 1989, Dorr & Valdespino 6437 (NY, QCA); Arenales del Chimborazo, super paramo, 4300 m, 19 Oct 2001, Endara et al. 361 (QCA), 25 Oct 1987, Ramsay & Merrow-Smith 997 (K, QCA), 10 May 1939, Penland & Summers 698 (F), 6 May 1927, Rorud s.n. (F). Pichincha: Cayambe mountain, N - 7 m, 24 Oct 1960, Pennington 22C (kK, NY). Tungurahua El Arenal, On the Via Whymper m from Cruce Arenal, 4150 m, 17 Abr 1983, ae 42143 aa, 03 Sep 1983, ee oe - (AAU). Distribucién y conservacion.—Especie endémica en los paramos del norte y centro del pais entre 4000 y 5100 m. La especie esta bien representada en varios paramos por lo que se la considera de Preocupacion Menor (LC). Nototriche jamesonii se distingue por la disposicion de las hojas en los extremos de las ramas formando rosetas laxas. La porcion libre del peciolo es bastante larga en relacién con las otras especies. La presencia del tubo de la corola es un caracter que comparte con N. ecuadoriensis (ver observaciones de dicha especie). Nototriche phyllanthos (Cav.) A.W. Hill, Bot. Jahrb. Syst. 37:579. 1906. (Figs. 1 D, y 3). Sida phyllanthos Cav, Diss. 5:276-277, tab. 127 fig. 4.1788. Malvastrum phyllanthos (Cav.) A. Gray, U.S. Expl. Exped. Phan. 152.1854. Tipo: “Pérou,” J. de Jussieu s.n. GHOLOTIPO: P-JU No. 12282 icroficha; MA nv, PI Sida saxifraga Bonpl., Pl. Aequinoct. 2:116. 1813. Tipo: ECUADOR: Cerro Antisana, Bonpland 2257 Ore TIPO: P como microficha, foto F-neg 35507; IsOTIPOS: B-W No. 12730 como microfich, P!). Si Pl. Aequinoct. 2:115-117, tab. 116. 1813. Malvastrum EUnet ene) ) A Gray. U.S. Expl. Bepedt Phan.1:152. 1854. Nolcrichem chinchensis (Bonpl.) A.W. Hill, Bot. Jahrb. Syst. 37:579. 1906; A. WHill, Trans. Linn. Soc. London, Bot. 7:231, pl. 29 ao pl. 30 fig. 9. 1909. Tipo: ECUADOR: Pichincha, in monte Ruccu Pichincha, Bonpland 3032 (HOLOTIPO: P; IsoTIPO: B-W No. 12731 como microficha). Hierba perenne, acaulescente, arrosetada o formando densos cojines; eje subterraneo lenoso, ramificado, los pices cubiertos de numerosas hojas verde- incano. Estipulas y peciolo forman una vagina membranosa, de 12-15 mm de largo x 3-4 mm de ancho. Parte libre de las estipulas triangular o lineal- subulada de apice agudo, de 8-10 mm de largo x 2-3 mm de ancho. Parte libre del peciolo de 4-6 mm de largo x 15-2 mm de ancho. Peciolo, estipulas y vagina con pelos estrellados pequenos en la superficie superior, glabrescente en la in- ferior,en los margenes con pelos estrellados mas grandes. Lamina flabeliforme, profundamente 3-partida, de 6-12 mm de largo x 10-18 mm de ancho, segmentos 700 BRIT.ORG/SIDA 21(2) Fic. 3. Nototriche ei original publicado como ida PINION en ais aoa Neeiale v. 2 (Humboldt y Bonpland, 1813, pl. 11 701 3-5-lobulados, los lobulos ovado-oblongos de apice obtuso con margenes frecuentemente enrollados, la superficie superior de la lamina incano- tomentosa, la inferior glabra o subglabra y vernicosa. Caliz campanulado, de 9- 10 mm de largo, los dientes de 5 mm de largo x 3 mm de ancho en la base, el apice subagudo, el exterior todo piloso con pelos similares a los de la lamina excepto los margenes con pelos estrellados mas grandes, el interior glabro excepto los dientes con pelos estrellados similares a los margenes. Nectarios internos basales 5, de 1,2 mm de alto x 1,5 mm de ancho. Corola violacea; tubo de la corola ausente. Pétalos de 16-25 mm de largo x 8-10 mm de ancho, la base con abundantes pelos estrellados largos. Tubo estaminal de 9-10 mm de largo, glabro, las anteras dispuestas en una cabezuela oblonga, 4mm de largo. Estilos y estigmas 7. Mericarpos 7, de 7 mm de largo (incluidas las aristas de 1,5 mm) x 2 mm de ancho, dorso estrellado-ciliado. Material adicional estudiado: ECUADOR: Localidad desconocida, 1943, Paredes s.n. (F). Bolivar: 28- 29 km NE Guaranda, Guaranda-Ambato hwy. Superparamo, 4185 m, 01.36S 079.00W, 25 Jun 1989, Dorr & Valdespino 6482 (AAU, CTES, F K, MO, NY, QCA, QCNE). Chimborazo: Volcan Chimborazo, 4750 m, 07 Feb 1988, Molau & Eriksen 2986 (AAU, QCA); 4400 m, 01.305 078.52 W, 12 Jul 1997, Neill & et al. 10788 (MO, QCNE). ee Paramo Ilinizas, 4 mi W Magdalena, 4600 m, 39S 078.40W, 2 Abr 1991, Bensman 362 (MO); Imbabura: Paramo de Zumbagua, 4000 m, 21 Jun 1986, Urgilés 19 (QCA); Napo: Quito- ie mete Guamani). 4150- Pe oe en ae oes (AAU, FE, MO, NY); Volcan Antisana, 60 km SE Quito, paramo, 4200 m, 00.285 78.05W, 14 Jun 1991, Gentry & Ortiz 74363 (MO), 1] Ene 1979, Holm-Nielsen 20651(AAU, MO), Lago Mauca-Machay, m, 02 Nov 1979, Holm-Nielsen 20721 (AAU), 28 Noviembre 1998, Vargas & Narvdez 3114 (MO, QCNE), 28 November 1998, Neill et al. 11494 (MO, QCNE), 16 Jul 1960, Grubbet al. 510 (NY), 19 Jul 1960, Grubb et al. 571 (NY), 23 Jun 1979, Black 68 (AAU), Santapamba, 4200 m, 17 Sep 1979, te 174 (AAU), ses Chuzalongo Chico, 4400 m, 14 May 1979, Black 5 (AAU), Laguna Micacocha, 01 Jul 1979, Lojtnant & Molau 15388 om oe “In montibus Antisana et Pichincha,” ane 917 (NY); “Sive Andium Quitensium,” May 1859, Jameson s.n (K); “Andes Quitenses,” Couthouys.n.(NY(2)), Jame- son s.n. (NY), Spruce 6542 (Kk); ene Pichincha, 4400 m, 00.10S 078.34W, 13 May 1995, Sklenar & eee 264 (AAU), 31 Ago 1939, Asplund 8601 (NY, P), Loma de las Antenas, 2800 m[?], 00.07S 78.30W, 10 Sep 1995, Clark & Fishman 1456 (QCNE, MO), 25 Jun 1934, Heinrichs 705 (NY), 27 Abr ane Holm oe 550 (F); Volcan Guagua Pichincha, oe 4650 m, 09 Ene 1988, Molau et al. 2387(AAU, MO, QCA ue ee, 1996, Clark 2522 (MO,QCNE), 25 May 1985, Nowak & Marcillo 47 (QCA), Volcan Pichincha, 4500 m, 17 Ago 1923, Hitchcock est (NY, US), 3 Jul 1876, André 3878 (K, NY), 15 Abr 1930, ae 2396 (LIL, P), Mexia 7653 (MO, NY, UC), 15500 ft, Jameson 97 (NY), Jameson s.n. (NY), Jameson s.n. (K), Desconocido 105 (NY), Desconocido 8 (K), Antisana, paramo 4350 m, Il Sep 1986, Ehrenburg 102 (QCA), 16000 ft, 16 Jul 1939, Balls B7287 (F, K); Volcan Cayambe, 4550-4660 m, Ol Mar 1988, Molau & Eriksen 3227 (AAU, QCA), 3 Dic 1993, Freire-Fierro et al. 2587 (AAU, NY, QCA), ene et al. 24902 (AAU, MO), 19 Mar 1995, Clark 470 (QCNE, MO); near border with Cotopaxi, 4200-4400 m, 7 Mar 1972, Harling 11170 (NY); Tungurahua: Paramo, ca. 2 km NW of the mountain Carihuairazo, 3500-4500 m, 01.245 78.47 W, 23 Abr 1995, Clark 716 (QCNE, MO). Distribucion y conservacién.—Especie endémica de los paramos del norte y centro del pais entre 3500 y 4650 m. Esta ampliamente distribuida y por lo tanto se la considera de Preocupacion Menor (LC). 702 BRIT.ORG/SIDA 21(2) Brako y Zarucchi (1993) registraron esta especie para Pert sobre la base del ejemplar tipo y de colecciones del departamento de Huancavelica. Los ejemplares de Rauh P-374(a) y P-450 corresponden a N. ulophylla (A. Gray) A.WHill, una especie muy distinta de N. phyllantos. En cuantoa la colecci6n tipo, apenas tiene lainscripcion que dice “Perou,” pero bien podria corresponder a alguna localidad ecuatoriana ya que en tiempos de la colonia parte de Ecuador estaba bajo la jurisdiccion del Virreinato de Pert. De otra manera no habiendo registros para Pert, pensamos que la especie esta restringida a Ecuador. Fryxell (1992) cita como holotipo de Sida pichinchensis una muestra de Humboldt y Bonpland s.n. sobre la base de una microficha del herbario de Willdenow en Berlin. Sin embargo, el Ing. Antonio Krapovikas (com. pers.) nos indica que él considera el holotipoa la coleccion, por élestudiada, de Bonpland 3032 incorporada en el herbario general de Paris ya que ese material es el que el autor debe haber usado para describir la planta mas no aquel de Berlin. Nototriche phyllanthos es una especie bastante comun, facil de distinguir por el envés de las hojas glabro y con un brillo singular. AGRADECIMIENTOS Agradecemos a los curadores de los herbarios por habernos permitido estudiar o haber enviado en préstamo ejemplares de sus instituciones. C. Aedo (MA), B.Klitgaard (BM), G. Lewis (K), V. Noble (BM), P. Barriga y C. Quintana (QCA), L. Ramella (G), nos ayudaron a obtener fotograf {as de material depositado en esos herbarios y F Keusenkothen escane6 la figura 3. A Paul Fryxell y Antonio Krapo- vickas por la minuciosa revision del manuscrito. M. Chanco agradece al Ing. Antonio Krapovickas del Instituto de Botanica del Nordeste, Corrientes, Argen- tina, por las facilidades y el material botanico de estudio brindados durante la estadia en su Institucion. El trabajo de campo de C. Ulloa en el sur del Ecuador se llevéa cabo con subsidios de la Nacional Geographic Society Jjargensen 6327- 98 y Ulloa 7744-02). REFERENCIAS Braxo, L.y J.L. Zarucchi. 1993. Catalogue of the flowering plants and gymnosperms of Peru. Monogr. Syst. Bot., Missouri Bot. Gard. 45:1-1286. CHanco, M. 1992. Dos nuevas especies de Nototriche (Malvaceae) del Peru. Novon 2: sy bor Hit, A.W. 1909. A revision of the genus Nototriche Turcz. Trans. Linn. Soc. London, Bot. 7:201-266, pl. 27-30. Fryxett, PA. 1990. New species of Malvaceae from South America. Contr. Univ. Michigan Herb. 17:163-171. FRYXeLL, PA. 1992. 118. Malvaceae, In:G. Harling & L. Andersson, eds. Fl. Ecuador 44:1-141. JorGeNsen, PM. y S. Leon-YANez (eds.). 1999. Catalogue of the vascular plants of Ecuador. Monogr. Syst. Bot., Missouri Bot. Gard. 75:1-1181. Krapovickas, A. 1950. Una nueva especie del género Nototriche (Malvaceae). Bol. Soc. Ar- gent. Bot. 3:170-173. Krapovickas, A. 1951. Notas citotaxondmicas en Nototriche (Malvaceae). |. Bol. Soc. Argent. Bot.4:107-116. KRrapovickas, A. 1953. Notas citotaxonomicas sobre Nototriche (Malvaceae). Il. ( Argent. Bot. 5:51-74. Krapovickas, A. 1957a. Tres especies nuevas de Nototriche (Malvaceae) de Peru. Bol. Soc. Argent. Bot. 6:233-238. Krapovickas, A. 1957b. Sobre las especies poligamas de Nototriche (Malvaceae). Lilloa 28: as ). Bol. Soc. Krapovickas, A. 1973. Las especies de Nototriche publicadas por Cavanilles en “Icones et descriptiones plantarum.” Bol. Soc. Argent. Bot. 15:285-286. UICN. 2001. Categorias y criterios de la lista roja de la UICN: Versi6n 3.1. Comisién de Supervivencia de Especies de la UICN. UICN, Gland, Suiza y Cambridge, Reino Unido. VaLencia, R., N. Pitman, S. LEON-YANEZ, y P.M. JarGeNsen (eds.). 2000. Libro rojo de las plantas endémicas del Ecuador 2000. Herbario OCA, Pontificia Universidad Catdlica del Ecua- dor, Quito. 704 BRIT.ORG/SIDA 21(2) BOOK NOTICES Two Books on Agapanthus from Timber Press Win SNOEVER. 2004. Agapanthus: A Revision of the Genus. (0-88192-631-0, hbk.). Timber Press Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, US.A.(Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $39.95, 372 pp., 101 color photos, 9 color illus., 7 tables, 6 drawings, 6" 9". +] is from southern Africa with about LO species, but there are hundreds of culti- vars. the author has Caen acu years of research and scholarship in prepar ing the “definitive” } Agapan The author breaks down his research i ntro- ‘1 duction; 2) Genome Size; 3) Taxonomy and Nomenclature; 4) Classification of the Galare 5) De- scriptions of the G Cult d 6) Cultivation. An appendix on Breeders/Intro- ducers and Their eee is followed by Relerences and an Index = om Nn HANNEKE VAN Dyk. 2004 Agapanthus for Gardeners. (0-88192-656-6, hbk.). Tim- ber Press Inc. 133 S.W Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, l- 800-327-5680, 503-227-3070 fax). $24.95, 96 pp., 147 color photos, | color map, 2 line drawings, 4 color illus. 81/2" x 81/2". Publisher Comments: “Agapanthus for Gardeners is an informative and practical guide to growing this popular ee It includes descriptions of all species and oo nea mane a selection of 80 cultivars that have proven themselves in the garden and in containers; information on choos- ing and buying plants nar te requirements, propagation, and pests and diseases; and a discus- sion of Agapanthus as a cut flower.” “T)] Illustrated with over 75 photographs, this is a useful companion to Wim Snoeijer’s more schol- arly and exhaustive Agapanthus: A Revision of the Genus, also published By Timber Press.” s small colorful book is truly a little jewel on Ene Flower of Love.” It covers everything from lta ee propagating to all tl SIDA 21(2): 704. 2004 ACERCA DELA IDENTIDAD DE BACCHARIS SUBSCULPTA (ASTERACEAE: ASTEREAE) Arturo Granda Paucar Daniel A. Giuliano to. de Biologia-Herbario Dpto. Cientifico de Plantas Vasculares Universidad Nacional Agraria La Molina Museo de La Plata Apdo. 456, Lima, PERU Paseo del Bosque s/n a.granda@lycos.com B1900FWA La Plata, ARGENTINA giuliano@museo.fcnym.unlp.edu.ar ABSTRACT AL ] A -] R 1 chr, a supposedly Peruvian endemic, Ho is considered a Sone of the more en B. ae Sch. Bip. ex Griseb. A lectotype is se- lected for the name of the new synonym. Key worpbs: Bacchari i Ipta, B pulchella, Asteraceae, Peru RESUMEN Después del andlisis de los tipos y protologos, Baccharis subsculpta Hochr. uma especie peruana Spare endémica, es considerada un sinonimo de B. pulchella Sch. Bip. ex Griseb. de mas iciOn. Se eleceiond un lectotipo para el nombre del nuevo sinénimo PALABRAS CLAVE: Bacchari } Ipt r B pulchella Asteraceae, Pert El género Baccharis L. (Asteraceae) es reconocido, entre las Astereae, por su primacia en cuanto al numero de especies que lo integran, todas exclusivamente americanas, como por su complicada taxonomia. Sus aproximadamente 360 especies ocupan un drea bastante extensa que abarca toda Sudamérica hasta alcanzar la zona sur de Estados Unidos de América, pero se extiende ademas por ambas norteamericanas mas alla del paralelo 40 N. Se desarrollan en condiciones ecolégicas muy dispares, habitando desde lugares humedos hasta sitios muy secos y desde el nivel del mar hasta las mayores alturas andinas (Cuatrecasas 1969); empero, su mayor diversidad se encuentra focalizada basicamente en la region neotropical. La delimitacion de sus especies generalmente se muestra confusa y la interpretacion cabal de las mismas ofrece serias dificultades, que radican en el extremo polimorfismo de muchas de ellas (o en una cierta continuidad morfol6gica en otras que forman complejos 0 alianzas pecies), asi como en la gran cantidad de entidades Aysdease que las mas de las veces corre pareja con la superficialidad de las diagnosis e ilustraciones que se han publicadoa su respecto. A todo ello se suma su peculiar diecia, que supone una cierta diferencia en la facies de los individuos pistilados y estaminados de una misma especie— que puede inducir a confusion en el observador poco agudo-—, y la distribucion SIDA 21(2): 705-710. 2004 706 BRIT.ORG/SIDA 21(2) — geogralica, que en algunas de sus especies es muy vasta, poniendo de relieve una gran capacidad adaptativa, mientras que en otras es de extension muy limitada senalando su condicién de endémicas. Por otro lado, sucede también que varias especies son halladas en el campo en poblaciones relativamente grandes, otras en cambio, se manifiestan como individuos escasos; como consecuencia, existen entidades que estan bastante bien colectadas y algunas, por contraste, resultan poco representadas en herbario. Estos factores apenas aludidos han contribuido a que la trayectoria taxonomica experimentada por algunas de las especies de Baccharis resultara erratica pues, a menudo, una entidad tempranamente descripta, con el transcurso del tiempo, sufrio diversas interpretaciones y determinaciones erroneas, esta situaciOn motivo que los autores posteriores que se ocuparon de alguna manera del género, utilizaran y difundieran otros epitetos para designarla. Lo dicho queda en evidencia al apreciarse la abultada sinonimia que reunen ciertas especies, v. gr. Baccharis salicifolia (Ruiz & Pav.) Pers., B. rhexioides Kunth, B. linearis Ruiz & Pav) Pers., ete. Baccharis pulchella Sch. Bip. ex Griseb. es un taxon distribuido desde Pert central y meridional hasta el centro de la Argentina. Desde que fuera original- mente descripta por A. Grisebach en 1879—sobre la base de una coleccion del viajero francés Gilbert Mandon proveniente de Sorata, Bolivia (Fig. 1), esta especie ha recibido la atencion de otros botanicos en distintas ocasiones, quienes le aplicaron cada cual denominaciones diferentes, no teniendo en cuenta la plasticidad morfologica de la especie; ésta presenta marcado polimorfismo en varios caracteres, a saber: pubescencia (existen desde plantas densamente tomentosas hasta casi glabras), ancho de la hoja (varia desde 0,5 cm hasta 3 cm), disposicion de los capitulos (reunidos en cimas corimbiformes difusas—o aun capitulos solitarios—o bien, agrupadas en racimos foliosos bien definidos). En efecto, el estudio de los tipos respectivos no ha dejado duda alguna sobre la identidad de las mismas y, en consecuencia, que se trata de nombres que han originado un listado de sindnimos en décadas recientes (Ariza Espinar 1971, 1973). Precisamente, el asunto que nos ocupa, y que estimamos conveniente dara conocer a continuacion, ilustra un caso adicional de sinonimia en Baccharis pulchella que recientemente hemos advertido. En el intento de determinar ciertos ejemplares peruanos colectados poco tiempo atras por uno de nosotros (A. G.) en el departamento de Lima, nos encontramos frente a la duda de asignarles este nombre~—al que parecian ajustarse bien de acuerdo a literatura moderna (Cabrera 1978; Giuliano 2000) y al estudio de ejemplares de herbario— 0, por otro lado, aplicarles el binomio Baccharis subsculpta Hochr., correspondiente a una rara planta peruana supuestamente endémica de territorio limeno con la que guardaban indiscutibles semejanzas morfologicas segtin comprobamos al observar un fototipo de la misma (Fig. 2), y a mas del detalle muy sugerente de provenir del locus typicus de la especie antedicha, la GRANDA Y GIULIANO, IDENTIDAD DE BACCHARIS SUBSCULPTA 707 NEW HAVEN CAMBRIDGE n 4 Fic. 1. lsotipo de Baccharis pulchella Sch. Bip ex Griseb. (Mandon 185, NY [foto LP]). F F “cuesta de Puruchuco”. Estos antecedentes nos llevaron de inmediato a sospechar de la verdadera identidad de B. subsculpta, de modo que intentamos reunir todos los elementos de juicio a nuestro alcance para aclarar nuestra presuncion de un posible caso de sinonimia; dispusimos de los protologos y fototipos de ambas entidades con el fin de estudiarlos y realizar un analisis comparativo. A juzgar 708 BRIT.ORG/SIDA 21(2) Fic. 2. Lect tip de Baccharis subsculpta Hochr. (Matthews 758, K [foto LP]) por las coincidencias entre ambas diagnosis, como asi por la innegable similitud entre los respectivos ejemplares originales—para B. pulchella especialmente en el caso del isotipo en NY, ya que los isotipos de K y LP presentan hojas mas breves y angostas, demostrando su variabilidad intraespecifica—, resulta indudable que B. pulchella es la misma entidad que ulteriormente Hochreutiner volvio a denominar B. subsculpta. Conviene hacer notar que c j iormente, la existencia de la diecia ha moti vado, dentro de Baccharis, la Pope ce de nuevas especies basadas just istilados. Tal es el caso de B. eee descripta en base a especimenes correspondientes al pie estaminado exclusivamente, mientras que B. pulchella lo fuera a partir de ambos tipos de individuos. Sin embargo, como parte de nuestro analisis, jem plares de diferentes poblaciones de la especie B. pulchella, los cotejamos con a protologo y con fotos e imagenes del material original peruano de B. subsculpta (su presunto sinonimo) y, a nuestro juicio, es imposible hallar la mas minima diferencia de ningun tipo que permita fundamentar separacion alguna. Debe mencionarse que, si bien Hochreutiner incluy6o a su Baccharis subsculpta en la seccion Oblongifoliae DC., este taxon infragenérico era altamente heterogéneo y artificial segan su concepcion original (cfr. Candolle 1836); luego de la redelimitacion que se la misma wae Cuatrecasas (1967), queda claro que la especie no corres} absoluto a dicha seccion. Asimismo, en el protdlogo de B. subsculpta se menciona que la especie es muy cercana a B. sculpta Griseb., y que también guarda semejanzas con B. sphaerocephala Hook. & Arn. y con B.grandicapitulata Hieron, no obstante, esta aparente similitud es solo superficial, ya que B. subsculpta no presenta afinidades con ninguna de esas especies, pertenecientes a distintas secciones del género. Por todo lo expuesto, proponemos a Baccharis subsculpta Hochr. como un nuevo sinonimo de la muy extendida B. pulchella Griseb., lo cual documentamos a COntInNUuUaCION Baccharis care Sch. Bip. ex Griseb., Symb. Fl. Argent. 181.1879. Tipo: BOLIVIA. Dpto. LARECAJA: viciniis Sorata, Nov 1858-Mar 1859, Mandon 185 (HOLOTIPO: GOET, ISOTIPOS: K [foto LPI, LP NY [foto LPI). = Baccharis subsculpta Hochr., Bull. New York Bot. Gard. 6(2):292. 1910. Syn. nov. Tipo: PERU. “Cuesta of Perruchuca[Puruchucol,” Matthews 758 (LEctoTIPO, aqui designado, K, foto LP!); “In Peruvia interiore,” Matthews 564 (sintipo NY, foto LP!). Obs: La seleccion del ejemplar Matthews 758 como lectotipo de la especie se fundamenta nee en que dicho espécimen proviene de una localidad lici itada, ademas del hechod id é j ivo de la mi J ee Ejemplares examinados: ARGENTINA. Proy. Catamarca: Dpto. Belén, Falcone & Castellanos 3528 (LP). Dpto. Tinogasta, Cabrera et al. 24649 (LP). Prov. Cordoba: Dpto. Calamuchita, Hu nziker 7217 (CORD, LP); idem, Roig 1111 (LP). Prov. Jujuy: Dpto. Santa Barbara, Cabrera et al. 24075, 25503 (LP). Proy. La Rioja: Dpto. Vinchina, Biurrun et al. 5123 (IZAC, LP). Dpto. no identificado, Morello 5162 710 BRIT.ORG/SIDA 21(2) (LP). Prov. Mendoza: Dpto. Las Heras, Sleumer 516 (LIL, LP). Dpto. Tunuyan, Ruiz Leal 23050 (LP). Proy. Salta: Dpto. Santa Victoria, Adler 6 (LP). Dpto. La Vina, Burkart 13285 (LP. SD. Prov. San Juan: Dpto. Angaco, Kiesling & Sdenz 4127 (LP, SD. Dpto. Sarmiento, Kiesling & Sdenz 4177 (LP. SD. Prov. Tucuman: as to. ey Venturi 913 (SD. Dpto. Chicligasta, Venturi 4614 (LP). o. Cochabamba: Prov. Quillacollo, Hensen 402 (SI). Dpto. La Paz: Prov. B. Saavedra, Beck 1 10 Xs sie Buchtien s.n. (LP 69942), PE to, Apurimae: Prov. Abancay, Ferre yra 9800 (US, USM). Dpto. Cusco: Prov. Cusco, So- lomon ee USM); Ferreyra 20841 (USM). Prov. Urubamba, Zamalloa 61 (LP). Dpto. Lima: Prov. Chancay, Cerrate 6339 (MO, USM,). Prov. Canta, Granda 1238 (MOL); idem, Granda & Alegria 1293, 1318, 1319, 1984, 2232 (MOL). Prov. Huarochiri, Cerrate & Tovar 1901 (USM AGRADECIMIENTOS Agradecemos a Charlotte Taylor y a Guy Nesom por las valiosas sugerencias y aportes que condujeron a la version definitiva del texto. El primer autor desea expresar su reconocimiento a las autoridades y al personal del Herbario del Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos (USM) por las facilidades prestadas; no olvida, por supuesto, al Bach. José Alegria por la lectura diligente del manuscrito. REFERENCIAS Ariza Espinar, L. 1971. Novedades sobre Baccharis (Compositae) de Argentina. Kurtziana Ariza Espinar, L. 1973. Las especies de Baccharis (Compositae) de Argentina Central. Bol. Acad. Nac. Ci. 50:175-305. Caprera, A. 1978. Baccharis. En: A.L. Cabrera, dir. Fl. prov, Jujuy 13(10):207-247. Coleccidn Cientifica del Instituto Nacional de Tecnologia Agropecuaria, Republica Argentina. CANDOLLE, AP. De. 1836. Baccharis. Prodr. 5:398-429. Paris (Treuttel & Wurtz). Cuatrecasas, J. 1967. Revision de las especies colombianas del género Baccharis. Revista Acad. Colomb. Ci. Exact. 13(49):5-102. Cuatrecasas, J. 1969. Baccharis.En:Prima flora Colombiana 3.Compositae-Astereae.Webbia 24:233-299 GiuLiANo, D.A. 2000. Subtribu Baccharidinae: Baccharis En: A.T. Hunziker, ed. Fl. Fanerog. Ar- gent. 66:6-67, GAMOCHAETA COARCTATA, THE CORRECT NAME FOR GAMOCHAETA SPICATA (ASTERACEAE: GNAPHALIEAE) John F. Pruski Guy L.Nesom Missouri Lae ical Garden Botanical Research Institute of Texas ox 299 09 Pecan Street St. Louis, reas 63166-0299 U.S.A. Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Gamochaeta spicata “(L san en ee sed illegiti later | (Gnaphali } Lam.1788, non P. Miller | stol il | x -haet spicat Cabr,, nom. nov, Cabrera’s “nomen novum,” h lso is illegiti I ; hali oarctatum Willd.,a replacement name e for Crea m spicatn iam is me Einst le gitimate name OE L ee D ant. Details in the a(Willd.) Kerg, the correct name in Gamochaeta for this species. Gnaphalium spicatum P. Miller isa synonym of Pterocaulon virgatum (L.) DC. Gnaphalium spicatum (Forssk.) Vahl is a synonym of Ifloga spicata (Forssk.) Sch. Bip. RESUMEN Gamochaeta spicata “(Lam.) Cabr.” esta basado en un homonimo posterior ilegitimo (Gnaphalium spicatum Lam. 1788, non P. Miller 1768) y debe ponSemiee como pale: naeta ae Cabr., nom. nov. El “nomen novum’ a ces sin snabalge, spicatum - cam., es el primer Porabre peti para la Willd., un nom planta ace L ee Paral a clavifieacion de lat t coezete ta WRG. )Kerg, el es | detall sé le | y citacion de los especimenes. Gnaphalium spic geen LP Miller es un sindnimo de Peroeaulon virgatum (L.) DC. S Gnaphalium spicatum (Forssk.) Vahl es siné le Ifloga spicata (Forssk.) Sch. During work preliminary to treatments for Flora Mesoamericana and Flora of North America, we noted that the species previously identified as Gamochaeta (Gnaphalium) spicata (Lam.) Cabr. (Compositae: Gnaphalieae) should be called Gamochaeta coarctata (Willd.) Kerg. Kerguélen’s combination (1987) was made preliminary to the Flora of France, after Flora Europaea (Holub in Tutin et al. 1976), and it has not been widely used. For example, Gamochaeta spicata was used in Freire (1995), but Gamochaeta coarctata was correctly used subsequently by Freire & Iharlegui (1997) and Cabrera et al. (2000). Typification of Gamochaeta coarctata, however, has been confused, and because the species is a worldwide weed (e.g., Cabrera 1961; Drury 1971; Kerguélen 1987; Nesom 2004a, 2004b), we provide details of typification necessary for convincing use of the name. In essence, the name Gnaphalium spicatum was validly published by three different authors for three different species from three different continents; these species are now referred to three different genera. Gnaphalium spicatum Lam. (1788) is the only one of the three that refers toa Gamochaeta species. It was used as a legitimate name by Cabrera (196D), but it is an illegitimate later hom- SIDA 21(2): 711-714. 2004 712 BRIT.ORG/SIDA 21(2) onym of Gnaphalium spicatum P. Mill. (1768). Vahl’s combination (1790), based on Chrysocoma spicata Forssk., added the third Gnaphalium spicatum. Willdenow (1803) provided a replacement name (Gnaphalium coarctatum) for Lamarck’s (1788) illegitimate Gnaphalium spicatum. Cabrera (1961) resurrected Gamochaeta from synonymy within Gnaphalium, but Kerguélen (1987) was the first to correctly use the first legitimate name for Lamarck’s type in Gamochaeta. The nomenclature and taxonomy of the three diff names Gnaphalium spicatum are as follows: 1. Gnaphalium spicatum P. Miller, Gard. Dict., ed. 8, Gnaphalium no. 24. 1768. SYNTYPE: JAMAICA: Sloane 38 Volume 5: eee digital mage: is is Brose me a aug ca lection from the Chelsea Botanic Garden, England is preserved in th [ os Current name.—Pterocaulon virgatum (L.) DC. Distribution.—southern United States to South America (Cabrera & Ragonese 1978). Miller’s protologue reads “24 Gnaphalium (spicatum) foliis lanceolatis decurrentibus tomentosis, floribus spicatis terminalibus lateralibusque.” Ad- ditionally, Miller 1768) cited “Elichrysum caule alato, floribus apicatis. Sloan. Cat. Jam. 125,” in direct reference to Sloane’s polynomial “Helichrysum caule alato, floribus spicatis” (Sloane, 1696:125) as illustrated in Sloane (1707:tab. 152, fig. 5). Miller stated that the plant “grows naturally in Jamaica, and in other of the hot parts of America” and that it “flowers in July and August, but never perfectly sets seed in England.” Gnaphalium spicatum P. Miller, was not cited in the monograph of Ptero- caulon (Cabrera & Ragonese 1978) but was listed by Jackson (1893) as a syn- onym of Pterocaulon virgatum (L.) DC. Based on our observation of the Sloane syntype (BM) of Gnaphalium spicatum P. Miller, of the cited original illustra- tion (Sloane 1707) of Gnaphalium spicatum P. Miller, and of the lectotype of Gnaphalium virgatum L. (LINN 993.29, IDC microfiche 177. 578.11.5!), we treat Gnaphalium spicatum P. Miller asa synonym of Pterocaulon virgatum (L.) DC. 2. Gnaphalium spicatum (Forssk.) Vahl, Symb. Bot. 1:70. 1790, hom. illegit., non P. Miller 1768. Chrysocoma spicata Forssk., Fl. Aegypt. 73. 1775. Ifloga spicata (Forssk.) Schultz-Bip. in Webb & Berthelot, Hist. Nat. Iles Canaries, vol. 3 Q, sect. 2):310. 1836-1850 1845]. Type: EGYPT: 1761-1763, Forsskal s.n. (HOLOTYPE: C, IDC 2200 27.1.1). Current name.—Ifloga spicata (Forssk.) Schultz-Bip. Distribution.—Canary Islands, northern Africa east to Pakistan (Davis 1975:L00-LOL. This widespread herb less than L5 centimeters tall with densely crowded and spiralling filiform leaves and axillary capitula was described from Egyp- tian material. This species was referred to Gnaphalium (Vahl 1790) and in turn to Ifloga (Schultz-Bipontinus 1844-1850). Davis (1975) and Anderberg (1991) treated this species as Ifloga spicata Forssk.) Schultz-Bip. 713 3. Gnaphalium spicatum Lam., Encycl. Méth. Bot. 2:757. 1788, hom. illegit., non P Miller 1768. Gnaphalium coarctatum Willd., Sp. PI. 3(3):1886. 1803, nom sae L. var. spicatum Klatt, Linnaea 42:140. 1878, nom. et stat. nov. Gamoch fae spicata Cabr.,, Bol. Soc. Argent. 9:380. 1961, nom. illegit. Gamochaeta coarctata (Willd.) Kerg., Lejeunia 120:104. 1987. Type: URUGUAY. Montevideo, Commerson s.n. [HOLOTYPE: P-LAM, IDC micro- iche 6207.325.1.1! possible IsoTYPE: P (photographs F!, GH!, TEX!, all photographs from C.B.G.N. negative 37573) Cabrera (1961:380-381) cited a sheet (“Des environs de Buenos Ayres... Commerson”) seen in P as “type” of Gnaphalium spicatum Lam. Drury (1971) cited the same sheet as lectotype. The locality on the label of this sheet (Buenos Aires, Argen- tina) conflicts with that of Lamarck’s protologue (“Monte-video”), and the lec- totype selection by Drury is rejected here. Although Montevideo, Uruguay, and Buenos Aires, Argentina, are 200 km apart and on different banks of the Rio La Plata, these two sheets are, nevertheless, conceivably of the same gathering. The sheet labeled “Buenos Ayres” is listed here as a possible “isotype” not lectotype. Current name.—Gamochaeta coarctata (Willd.) Kerg. Distribution.—cosmopolitan weed. The names listed in the above citation of Gnaphalium spicatum Lam. are homotypic. Gnaphalium coarctatum Willd. 1803) is treated as having been origi- nally anomem novum for Lamarck’s plant, and it dates from 1803. In providing a replacement name for Lamarck’s Gnaphalium spicatum, Willdenow (1803:1886) cited the locality as “Monte Video” (echoing Lamarck’s citation) for the species. On the same page, Willdenow treated the name Gnaphalium spicatum (Forssk.) Vahl as referring to an Egyptian species, citing Chrysocoma spicata Forssk. asa synonym (see #2, above). In the Gnaphalium treatment, Lamarck (1788) did not specifically refer to the earlier Gnaphalium spicatum P. Miller. Cabrera’s (1961:380) intended transfer of Lamarck’s illegitimate name to Gamochaeta (as “Gamochaeta spicata (Lam.) comb. nov.”) is to be treated as hav- ing been originally a nomem novum. Gamochaeta spicata Cabr., however, is il- legitimate (vidi ICBN Art. 52.1) because the “available” and legitimate Gnaphalium coarctatum should have been adopted. Kerguélen (1987) treated Gnaphalium coarctatum and Gnaphalium spicatum Lam. as homotypic and supplied the correct (in Gamochaeta) combination for this plant. Gamochaeta spicata was treated as a synonym of Gamochaeta americana (P. Miller) Wedd. by Nesom (1990), but G. coarctata commonly differs from G. americana by stems being up to 60 (vs. to 25) cm tall, basal leaves wider (vs. narrower) than 8 mm, involucres 3-4 (vs. 4.5) mm high, and phyllaries blunt and straight (vs. acute, cuspidate, and reflexed), as noted by Drury (1971). ACKNOWLEDGMENTS We appreciate the reviews and comments by John Strother and Kanchi Gandhi. 714 BRIT.ORG/SIDA 21(2) REFERENCES AnberberG, A.A. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Bot. 104:1-195. Casrera, A.L. 1961. Observaciones sobre las Inuleae-Gnaphalineae (Compositae) de América del Sur. Bol. Soc. Argent. Bot. 9:359-386. Casrera, AL. and A.M. Raconese. 1978. Revision des género Pterocaulon (Compositae). Darwiniana 21:185-257. Casrera, A.L., JV. Crisci, G. DetuccHi, S.E. Freire, D.A. GUILIANO, L. IHARLEGUI, L. KATINAS, A.A. SAENZ, G. SaNcHO, and E. Urtusey. 2000. Catalogo ilustrado de las Compuestas (= Asteraceae) de la Provincia de Buenos Aires, Argentina: Sistematica, Ecologia y Usos. Secretaria de Politica Ambiental, Buenos Aires. Davis, PH. (ed.). 1975. Compositae. Flora of Turkey and the East Aegean Islands, vol. 5. Uni- versity Press, Edinburgh Drury,D.G. 1971. The American spicate cudweeds adventive to New Zealand (Gnaphalium section Gamochaeta - Compositae). New Zealand J. Bot.9:157-185. Freire, S.E. 1995. 280. Asteraceae, parte 2. Tribu IV. Inuleae. Flora Fanerogadmica Argentina. Fasciculo 14:1—60. PROFLORA, CONICET, Cordoba. Freire, S.E.and L.IHartecul. 1997. Sinopsis preliminar del género Gamochaeta (Asteraceae, Gnaphalieae). Bol. Soc. Argent. Bot. 33:23-35. Ho.us, J. 1976. Gamochaeta. In: Tutin, T.G., V.H. Heywood, N.A. Burges, D.H. Valentine, S.M. Walters & D.A.Webb, eds. Flora Europaea, vol.4. Cambridge Univ. Press, Cambridge. P. 127. Jackson, B.D. 1893. Index kewensis, vol. 1: A-J. Clarendon Press, London. Kerauelen, M. 1987. Données taxonomiques, nomenclaturales et chorologiques pour une révision de la flore de France. Lejeunia 120:1-264 Lamarck, J.B. 1788. Encyclopédie méthodique: botanique, vol. 2. Panckoucke, Paris. Miter, P. 1768. The gardeners dictionary, ed. 8. Rivington Printers, London. Nesom, G.L. 1990. The taxonomic status of Gamochaeta (Asteraceae: Inuleae) and the spe- cies of the United States. Phytologia 68:186-198. Nesom, G.L. 2004a. New species of Gamochaeta (Asteraceae: Gnaphalieae) from the east- ern United States with comments on similar species. Sida 21:717-741., Nesom, G.L.2004b. New distribution records for Gamochaeta (Asterac phali in the United States. Sida 21:1175-1185. ScHuuTZ-Birontinus, C.H. 1844-1850. Compositae. In: PB. Webb & S. Berthelot. 1836-1850. Histoire naturelle des Iles Canaries, vol. 3 [Botaniquel], part 2, sect. 3. Bureaux, Paris. Pp. 203-473 + figs.82—136B. Stoane, H. 1696. Catalogus plantarum quae in insula Jamaica. D. Brown Printers, London. Stoane, H. 1707. A voyage to the islands Madera, Barbados, Nieves, S. Christophers and Jamaica, with the natural history of the herbs and trees. 2 vols. B.M. Printers, London Steuoet, E.T. 1840. Nomenclator botanicus, ed. 2, vol.|: A-K. Tubingae, Stuttgart. VAHL, M. 1790. Symbolae botanicae, vol. 1. Printed privately, Copenhagen. Wittoenow, C.L. 1803. [Syngenesia, vol. 3, part 3, pages 1477-2409]. Caroli a Linné species plantarum. Nauk, Berlin. A ae OF AGOSERIS APARGIOIDES TERACEAE: LACTUCEAE) Gary |. Baird Department of Biology, Brigham Young University-ldaho, 525 S. Center St. Rexburg, Idaho, 83460-1100, U.S.A, bairdg@byui.edu ABSTRACT A brief review of ; presented. The species is redefined to exclude A. hirsuta, a pr grassland species found lone the Coast Ranges of California. Agoseris dpargioides occurs on coastal dunes along the Pacific coast from central California to Washington. Three varieties are recognized: var. dpargioides, var. east woodiae, and var. maritima (E. Sheld.) G.l. Baird, comb. et stat. nov. RESUMEN le defi excluir A. hirsuta, Se cesome a una breve revision de A . La especie s especie que se encentra en los pe astos a lo nee de las cordilleras costeras ide California. Agose ris oe s aparece en las dunas costeras a lo largo de la costa del Pacifico desde el centro de Califor- nia hasta Washington. Se reconocen tres variedades: var. apargioides, var. east woodide, y var. mar- itima (E. Sheld.) G.l. Baird, comb. et stat. nov In 1816, the Romanzov Expedition (1815-1818) spent the month of October at the Spanish presidio on the north side of what is now San Francisco, California (Chamisso 1836). At this time, L. A. von Chamisso, a member of the expedition, collected the type of Agoseris apargioides (Less.) Greene. This species occurs on coastal dunes along the Pacific coast from central California to Washington. In the protologue, Lessing (1831) noted a peculiar feature of this plant: due to its preference for dune habitats, the stems often are buried by drifting sands, leav- ing a terminal rosette of leaves exposed. These buried stems become pseudo- rhizomatous, a feature unique within the genus In his monograph of Agoseris, Jones (1954) mistakenly interpreted the type description of A. apargioides as belonging to an allied but separate species, A. hirsuta (Hook.) Greene. Agoseris hirsuta is a grassland species found along the Coast Ranges of California. It is strictly acaulescent and only rarely occurs ad- jacent to the Pacific shore (and then not on dunes). Jones (1954) incorrectly treated A. hirsuta as “A. apargioides ssp. apargioides” and A. apargioides as “A. apargioides ssp. maritima.” This last subspecies was based on A. maritima E. Sheld., the type collected at Coos Bay, Oregon. Within ssp. maritima, Jones (1954) recognized two varieties: “var. maritima” and “var. east woodiae,” this last vari- ety based on A. east woodiae Fedde, the type collected at Bodega Bay, California. Jones’ (1954) view of A. apargioides has been the basis for all recent treatments of this species (e.g., Jones 1960; Chambers 1993). In my recent monograph of Agoseris (Baird 1996), it became apparent that Jones (1954) had misinterpreted A. apargioides. The name A. hirsuta is here rees- tablished as the correct name for the inland species. Agoseris apargioides is rede- fined to include just those plants that occur along the Pacific coastal strand. Three variants within A. apargioides are morphologically and geographically distinct. SIDA 21(2): 715-716. 2004 716 BRIT.ORG/SIDA 21(2) Agoseris apargioides (Less.) Greene, Pittonia 2:177. 1891. Troximon apargioides Less. Linnaea 6:501. 1831. Type: U.S.A. CALIFORNIA. San Francisco, Oct 1816, Chamisso s.n. Perennials, + caulescent, buried stems pseudo-rhizomatous. Leaves linear to spathulate, entire to lobate, or filiformly pinnatifid, 3-15 cm x 1-30 mm, glabrous to villous, mostly prostrate to reclining; lobes 3-5(-7) pairs, filiform to spath- ulate. Scapes 7-45 cm, villous to tomentose, glandular or not. Involucres obconic to hemispheric, 1.5-2.5 cm tall; phyllaries in 2-4 series, glabrous to tomentose and + glandular; receptacle naked. Florets 25-200; corollas yellow; tubes 2-5.5 mm; ligules 3-16 x 1-3 mm; anthers 1.5-4.5 mm. Achenes 5-12 mm, beaked; body fusiform to obconic, 3-5 mm; beak 3-8 mm. Pappi of bristles, 4-9 mm. x = 18. KEY TO VARIETIES 1. Corolla ligules 3-6 mm long, involucres non-glandular var. maritima 1. Corolla ligules 8-16 mm long, involucres + glandular. 2. Leaves oblanceolate to spathulate, dentate to lobate;involucres densely villous to tomentose ar. eastwoodiae 2. Leaves linear to oblanceolate, entire to dentate or filiformly pinnatifid, aati cres glabrous to villous var. apargioides Agoseris apargioides var. apargioides. Occurs from San Francisco south to Pt. Sur. Agoseris apargioides var. eastwoodiae (Fedde) Munz, Aliso 4:100. 1958. = Agoseris eastwoodiae Fedde, Bot. Jahresb. 31:808. 1904. Type: U.S.A. CALIFORNIA. Sonoma Co.: Bodega Point, 4 Jul 1900, East wooc = sn Occurs from Pt. Reyes north to about Pt. Arena. Agoseris apargioides var. maritima (E. Sheld.) G.1. Baird, comb. et stat. nov. = Agoseris maritima E. Sheld., Bull. Torrey Bot. Club 30:310. 1903. = Agoseris apargioides subsp. maritima (E. Sheld.) Q. Jones ex Crongq., Vasc. Pls. Pacif. Northw. 5:24. 1955. TyPE: U.S.A. OREGON. Clatsop Co.: Clatsop Beach, 21 Aug 1902, Sheldon 11250. Occurs from Humboldt Bay, California, to Neah Bay, Washington. ACKNOWLEDGMENTS [thank the curators and staff at BM, CAS, DS, GH, JEPS, K, NY, ORE, OSC, POM, UC, US, WILLU, and WTU for their assistance with loan materials. 1 appreciate the prompt and excellent comments of Guy Nesom (BRIT). REFERENCES Bairo, G.I. 1996. The systematics of Agoseris (Asteraceae: Lactuceae). Ph. D. diss., Univ. of Texas, Austin. CHameers, K.L. 1993. Agoseris. In: Hickman, J.C.ed. The Jepson manual: higher plants of Cali- fornia. Berkeley, Los Angeles & London. Pp. 190-193. CHAMIssO, L.A.VON. 1836.A voyage around the world with the Romanzov exploring expedi- tion in the years 1815-1818 in the brig Rurik, Captain Otto von Kotzebue. Translated and edited by H. Krantz. 1986. Honolulu. Jones, Q.1954.Monograph of Agoseris, tribe Cichorieae. Ph. D.diss., Harvard Univ., Cambridge. Jones, Q. 1960. 153. Agoseris Raf. In: Abrams, L. and R.S. Ferris, eds. Illustrated flora of the Pacific States, vol.4: Compositae. Stanford. Pp. 562-567. Lessina, C.F. 1831.De plantis in expeditione Romanzoffiana: observatis disserere pergitur. Synathereae Addendae. Linnaea 6:501. NEW SPECIES OF GAMOCHAETA (ASTERACEAE: GNAPHALIEAE) FROM THE EASTERN UNITED STATES AND COMMENTS ON SIMILAR SPECIES Guy L.Nesom Botanical Research Institute of Texas Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Gamochaeta argyrinea Nesom, sp. nov, is doc ted from 19 states, primarily in the southeastern US. os and fom EEO Bio It isa common and abundant species of ruderal habitats and has u all 1 | Gamochaeta purpurea, which hasa similar but en is most closely similar to G. ustulata, another A. and ] BOE An range. Gamochaeta argyrinea af mmonly identified as G. purpurea but native to the Pacific coast region of the aGAcent Canada. Gaiaarhacs chionesthes Nesom, sp. nov, is described from localities in Atego Louisiana, Mississippi, Alabama, Georgia, Florida, South Carolina, and North Carolina—these plants also have been identified previously primarily as G. purpurea. A key and distribution maps are pro- vided for the six species of Gamochaeta in the U.S.A. with strongly bicolored leaves: G. argyrinea, G. ustulata, G. chionesthes, G. purpurea, G. simplicicaulis, and G. coarctata. The name Gamochaeta americana has been misapplied to G. coarctata, but G. americana sensu stricto has not been docu- mented for the U.S.A. it occurs in the Antilles, Central America, Mexico, and South America and is reported to occur elsewhere as an adventive. In order to further clarify its identity, a technical de- scription and commentary are provided for G. americana. RESUMEN Se coments Camestacts argyrinea Neca! ay nov, de 19 eooes paineipaimente ae aed ee US eualmente dentreo del nonce pro amplio de icannnaetn purpurea, que tiene un rango geogralico similar pero mas amplio. Gamochaeta argyrinea es oe mas eo a G. ustulata, otra especie eae comunmente como G. purpurea pero nativa de la region a del Pacifico de nada adyacente. Gamochaeta chionesthes Nesom, sp. nov, se describe de localidades de eee feces Mississippi, Alabama, Georgia, Florida, Carolina del Sur, y Carolina del Norte— estas panes EIPIEN nae sido eae a G. purpurea. Se olrece una clave y haeta de U.S.A. con hojas bicolores: G. argyrinea, G. ustulata, G. NORESENES i. purpurea G. simplicicaulis, y G. coarctata. . El HOEUOE re neemociaete G.coarctata, pero G para o> A. estaen las Antillas América Central, México a del Sur y se cita de otros lugares d y americana para clarificar su identidad. Studies of North American Gamochaeta Wedd. (Godfrey 1958; Nesom 1990) have identified plants with leaves strongly bicolored (persistently white-pannose abaxially with a closely matted vestiture completely obscuring the epidermis, evidently green to grayish green adaxially) mostly as Gamochaeta purpurea SIDA 21(2): 717-741. 2004 718 BRIT.ORG/SIDA 21(2) (L.) Cabr. Review of North American plants shows that more species are present than recognized in these earlier studies and also that some names have been misapplied. Taxonomic studies of Gamochaeta in New Zealand (Drury 1971; Webb 1988), where all the species are non-native and include a number of those found in the U.S.A., have been more discerning, at least in some respects, than previous studies of North American species. In the present study, two species that have been identified mostly as Gamochaeta purpurea are described as new to science. One is distributed over a large part of the eastern U.S.A. and also is known from Puerto Rico; the other is known from Gulf Coast states of the U.S.A. Neither of new species keys un- ambiguously ina recent overview of Gamochaeta (Freire & [harlegui 1997), and a survey of the genus in South America and Central America indicates that neither of these species now recognized in North America has received a name, although it is possible that one or both is native to South America. It also is possible that both are weeds widely distributed on several continents. Hypoth- eses regarding the nativity of all species of Gamochaeta occurring in the U.S.A. are provided in an accompanying discussion (Nesom 2004). All of the species in North America with strongly bicolored leaves are reviewed here. The distinctiveness of Gamochaeta as a genus was emphasized by Cabrera (1961 and later floristic treatments of South American species, e.g., 1963, 1971, 1974, 1978), Dillon and Sagastegui (1991a, 1991b), Cabrera and Freire (1998), and by other botanists who have treated it (e.g., Nesom 1990; Anderberg 1994; Freire & tharlegui 1997). Gamochaeta is distinguished by its combination of small heads ina spiciform capitulescence, concave post-fruiting receptacles, truncate collecting appendages of the disc floret style branches, small achenes with minute, mucilage-producing papilliform trichomes on the surfaces, and pap- pus bristles basally connate in a smooth ring and released as a single unit. A widespread new species from eastern U.S.A. related to Gamochaeta ustulata Godfrey (1958) noted variation within what he identified as Gnaphalium purpureum and | have observed Gamochaeta (Gnaphalium) purpurea sensu stricto growing in close proximity to a “variant” or intermixed with it in many localities in eastern North America. Intergradation apparently occurs rarely if at all and the two entities can be consistently and accurately distinguished, both in the field and herbarium. In fact, the variant is more similar and prob- ably more closely related to Gamochaeta ustulata (Nutt.) Holub, a species ap- parently native to western North America (see comments and description be- low) than to G. purpurea. The plants of eastern North America are recognized here as a previously undescribed species and documented to occur in 19 states. aaa argyrinea Nesom, sp. nov. (Figs. 1, 2, 3, 4, 6). Type: U.S.A. NorTH CARo- NA, Pender Co: Hwy 421 at junction with Hwy 210, between towns of Currie and — ne roadsides and grassy median strip, in sandy soil; in close association purpurea sensu stricto, Gamochaeta pensylvanica, Gamochaeta antillana, and G -haet NESOM, NEW SPECIES OF GAMOCHAETA 719 PLANTS OF NORTH CAROLINA Pender County. Hwy 421 at junction with Hwy 210. between Currie and Rocky pola Roadsides and grassy bese seaa Ganochaela coarctal Common. Guy Nesont WMGT 16 i ee, 28 April 2001 Fic. 1. Habit of Gamochaeta argyrinea. BRIT.ORG/SIDA 21(2) O Botanical Research institute of Texas Cents oy IFORGIA ACEAE Meriwether Co’ te near jot with Bay 109. aah Post Office on Hwy 74/85, nes soil, Gameckacta cearciata and G, chionesthes present and ees Sey site: (7. poe UNCOMMON, bundant plants with involucres lighter Comi esi ne Snel TYPE COLLECTION: | 14 April 2004 GJ., Nesom GASCOI—14 Fic. 2. Habit of Gamochaeta chionesthes. NESOM, GAMOCHAETA 721 coarctata, all growing along the roadside, 28 Apr 2001, G. Nesom WMGT-14 (HOLOTYPE: BRIT; ISOTYPES: AKU, BM, BRIT, CANB, CANU, F GA, GH, K, LP, MEXU, MO, NCU, NSW, NY, P, RB, S, TEX,UC US USE) Differt a Gamochaeta purpurea radicibus plerumque fibrosis, foliis caulinis oblanceolatis vel oblanceolati- eee trichomatis paginaram tgliantum adaxialium muon DUS ad Baek a apich capitulis 3-3.5 mm altis, phyllariis ee amini g p d api- ces, et secant bisexualibus (3-)4- Plants annual to winter eae densely fibrous-rooted, rarely slender-taprooted. Stems decumbent-ascending from the base, 12-40 cm tall, simple or few- branched, closely white-pannose, the vestiture usually of individually evident trichomes, less commonly nearly cloth-like. Leaves basal and cauline, basal persisting and green through flowering, basal and lower cauline oblanceolate to oblanceolate-oblong or oblanceolate-obovate, 1.5-5(-8) cm long, 5-12(-18) mm wide, gradually reduced in size upward, not clasping or decurrent, bicolored, closely white-pannose abaxially, persistently very sparsely arachnoid adaxially (sometimes necessary to examine at 10x). Capitulescence cylindric in early sea- son, 1.5-5 cm long, 10-12 mm wide (pressed), later producing axillary glomer- ules from lower nodes and elongating, becoming strongly interrupted and up tol8cm long (but still narrowly cylindric). Involucres campanulate, 3-3.5 mm, imbedded at base in cottony tomentum,; phyllaries in 4-6 series, outermost ovate-acute to ovate-lanceolate, tawny-transparent, 1/3-4/5 as long as the in- ner, inner elliptic-oblong to oblong, stereome ca. 2/3 the length, lamina apically truncate-rounded and apiculate, flexing slightly outward at maturity, hyaline- translucent and slightly brownish-tinged, often purplish tinged around the stereome/lamina junction; receptacles shallowly concave. Florets: bisexual 4- 5(-6), all corollas purple- to yellow-brown-tipped; pistillate ina broad zone. Cypselae oblong, 0.5-0.6 mm long, tan, papillate. Flowering Mar-Jun(-Jul, -Oct). Roadsides, fields, lawns, open woods, sand to sandy clay, almost always in open, disturbed areas; ca. 0-250 m. USA: Ala- bama, Arkansas, Delaware, Florida, Georgia, Kansas, Kentucky, Louisiana, Mary- land, Mississippi, Missouri, North Carolina, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, West Virginia; Puerto Rico, Argentina?, New Zealand?, Australia?, Portugal? In the southeastern U.S.A,, Gamochaeta argyrinea commonly grows with G. coarctata and G. chionesthes. The epithet (from Gr. argyreos, silvery) alludes to the silvery appearance of the plants, espe- cially when growing in colonies. R U.S.A. ALABAMA. Marion Co.: Hamilton, vacant lot, sandy loam, 22 — 1966, Shimmer 31207 (SMU). ARKANSAS. Cleburne ee Tumbling Shoals, ae bluffs, 500 ft, 19 May 1951, Demaree 39517 (SMU). DELAWARE. Sussex Co.: 4 mi SE of Laurel, open ground at Moore’s (Trussum) Pond, 19 May 1939, Tatnall 4163 (DOV). FLORIDA. Leon Co.: near Sen fallow field, 31 Mar 1955, Godfrey 53123 (S oe See Clarke Co:: Senda. alone River Noth moist ene 700 ft, 2 May 1947, Cronquist 4369 (SMU). KA Chero me ings, 1345S, R24E, Sec 24, 5 May ce ae ee as KENTUCKY. Whitley Co.: N Teellico 722 BRIT.ORG/SIDA 21(2) meadow, Il Jul 1937, Smith and Hodgdon 3808 (GH). LOUISIANA. Lincoln Parish: Dubach, low ground by hwy, sandy clay, | May 1955, Shinners 19976 (SMU). MARYLAND. Calvert Co.: St. Leonard, District No. 1, 3 Aug 1956, Seymour 16837 (MO). MISSISSIPPI. Rankin Co.: 1 mi E of Rankin-Madison County line, Hwy 43, sandy weedy field, 2 May 1970, Jones 18632 (SMU). MISSOURL. Howell Co.: 3.5 mi N of cee rocky, brushy pastureland, 25 Apr 1992, Summers 4827 (MO). NORTH CAROLINA. David- ca. 12 mi ESE of Lexington on Hwy 64, 0.7 mi E of jet. Hwy 109, loamy an f roadbank below sees road cut, area of white oak- eee! woods, 30 May 2001, Nesom GX1 (BRIT, NCU, MO, US). Wake Co.: E. pas a iew Gardens, fallow field, 20 Apr 1948, Godfrey 48072 a OKLA- HOMA. Delaware Co.: 8.5 mi SE of Jay, open woods hillside, chert rock soil, 22 Apr 1967, sits lens 10543 (SMU). PENNSYLVANIA. Northumberland Co.: Herndon, in dry soil along roadside, 30 Sep 19 Moldenke 4186 (NY). SOUTH CAROLINA. Berkeley Co.: 0.5 mi N of Honey Hill, sandy soil along route 45, 16 Apr 1971, Churchill s.n. (SMU). TENNESSEE. Knox Co.: Knoxville, U.T. campus, lawn, 19 Apr 1968, Morton 2861 (SMU). TEXAS. Brazos Co.: near Peach Creek, open areas in woods, 7 Apr 1974, Fryxell 2367 (SMU). VIRGINIA. Charles City Co.: 7.5 mi W of Rustic, 30 Apr 1970, Ware 2870 (V DB). WEST VIRGINIA. Cabell Co.: base of Robert's Hill, Milton, | May 1938, Wan 699 (MO). PUERTO RICO. Barranquitas region, wet place, 700 m, 14 Nov 1979, Liogier 30027 (NY, UPR); Cuilarte Forest, on slope, 900 m, : Jan 1980, Liogier 30271 (NY, UPR). Ciales: Los Tres Picachos, Rt 149, km 35.5, dirt road through ffee plantation, disturbed wet mountain oe ca. 600-750 m, 14 Mar 1992, Axelrod 4213 (NY, UPRRP). Bante Bo. Anon, Toro Negro Forest Reserve, trail along SE side of Monte Jayuya, wet mountain forest, ca. 1200 m, 24 Apr 2003, Axelrod oo near Cerro de Punta, Jayuya, in thickets, 1200 m, 10 Apr 1982, Liogier 33089 (NY, UPR); rte 143 at Cerro de la Punta, roadsides and thickets, 1000 m, 29 May 1988, Taylor 8116 (UPRRP). ae Barrio Lapa, Las Tetas de Cayey, summit area of E. peak, 820-830 m, low exposed thicket near brink of cliff, 31 Mar 1988, Proctor 44634 (SJ). A population in Davidson Co., N.C. (Nesom GX1, as cited above) is a variant— the plants tend to produce small tubers or cormlike swellings. Plants of other populations in the same area often produce offsets that are nearly rhizome- like (e.g, Davie Co., N.C, Nesom GX2, BRIT). Essential differences that distinguish Gamochaeta argyrinea from G. purpurea are given in the following couplet. a. Basal leaves persistent and green at flowering;cauline leaves oblanceolate to oblan ceolate-oblong or oblanceolate-obovate, trichomes of adaxial leaf surfaces filiform from base to tip; involucres 3-3.5 mm high; inner phyllaries with lamina oblong, apically truncate-rounded and apiculate; bisexual florets 4-5(-6) per head Gamochaeta argyrinea a. Basalleaves lly withered and becoming deciduous at flowering; cauline leaves spatulate, trichomes of adaxial leaf surfaces with basal cells expanded and vitreous; involucres 4-4.5 mm high; inner phyllaries with lamina triangular, apically acute but not apiculate; bisexual florets 3-4 per hea Gamochaeta purpurea —_) Plants common in the Pacific coast region have usually been identified as Gamochaeta purpurea, but they are more similar to G. argyrinea and are iden- tified here as G. ustulata. New Zealand plants identified by Drury (1971) and Webb (1988) as G. purpurea var. ustulata have measurements characteristic of G. ustulata, but plants in Portugal identified as G. ustulata by Afonso (1984) are more likely G. argyrinea. Gamochaeta ustulata (Figs. 3, 4, 7) is recognized by its fibrous rooted habit, large, weakly bicolored leaves persistently arachnoid on NESOM, NEW SPECIES OF GAMOCHAETA 723 3. Invol f I ies (left to right). (Top) G. argyrinea, G. ustulata, G. coarctata, G. americana, (bot- tom) G. purpurea, G. chionesthes, and G. simplicicaulis, tw =) | | Phyll hol FG fi i ter, middle, and inner phyllary for each species. (A) G. argyrinea, se uae (C) G ade. (D) G. americana, (E) G. purpurea, (F) G. chionesthes, and (G) G. simplicicaulis. 724 BRIT.ORG/SIDA 21(2) the adaxial surfaces, and large brownish capitula in a thick, usually continu- ous capitulescence. It differs from G. purpurea in its longer duration, thicker and shorter stems, wider and more compact capitulescence, larger capitula,and prominently brown, blunt-apiculate inner phyllaries with a subterminal keel. Gamochaeta ustulata and G. argyrinea differ primarily by features in the fol- lowing couplet. a Capitulescence 12-18 mm wide (pressed), uninterrrupted or rarely so and only at the base, mostly 1—6(-8) cm long; involucres 4.5—5 mm high; outer phyllaries and often lamina of inner phyllaries dark brown, commonly the whole involucre with a dark brown cast; mid phyllaries with a low, subterminal keel; cypselae 0.7-0.8 mm ong Gamochaeta ustulata a. Capitulescence 10-12 mm wide (pressed), early uninterrupted and 1.5—5 cm long, later producing axillary glomerules from lower nodes and elongating, becoming strongly interrupted and up to 18 cm long; involucres 3—3.5 mm high; phyllaries greenish-silvery to stramineous or golden, sometimes light brown but the whole involucre not with a dark brown cast; phyllaries not keeled; cypselae 0.5-0.6 mm long Gamochaeta argyrinea Gamochaeta ustulata (Nutt.) Holub, Folia Geobot. Phytotax. 11:83. 1976 (non Nesom, Phytologia 68:196. 1990). Gnaphalium purpureum var. ustulatum (Nutt.) Boivin, Naturaliste Canad. 87:34. 1960. Gnaphalium ustulatum Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:404. 1841. LECTOTYPE (Nesom 1990): USA. CALIFORNIA: Near St. Barbara in Upper Cali- fornia, [March or April 1836], T. Nuttall s.n. (BM, GH-photo!, TEX-photol). Gnaphalium pannosum Gandoger, Bull. Soc. Bot. France 65:42. 1918 (non A. Gray 1883; non Schultz- Bip. 1845). SYNTYPEs: USA. WASHINGTON. Klickitat Co. on damp ground, Columbia River, 10 May 1893, WN. Suksdorf 1580 (NY-2 sheets, US!); WASHINGTON. Chehalis Co.: near Montesano, 200ft, 8 Jun 1898, A.A. Heller 3919 (NY-2 sheets!). Images of both the Suksdorf and Heller collections can be seen on NY and US internet sites. Plants annual to biennial or short-lived perennial, fibrous-rooted. Stems erect toascending, commonly decumbent-ascending and rhizome-like, 10-40.cm tall, densely white-pannose. Leaves basal and cauline, basal persistent but often withering by flowering, spatulate to oblanceolate, 2-5 cm long, 6-12(-35) mm wide, apiculate, not clasping or decurrent, continuing upwards little reduced until into the capitulescence, weakly bicolored, adaxial surfaces sparsely to densely arachnoid with persistent trichomes, closely white-pannose abaxially. Capitulescence |-6(-8 or more) cm long, 12-18 mm wide (pressed), uninterrrupted or rarely so and only at the base. Involucres campanulate-ur- ceolate, 4.5-5 mm high, base imbedded in cottony tomentum; outer phyllaries broadly triangular-ovate, half as long as the inner, all with a brown or green- ish-brown cast, inner with lamina dark brown, apically abruptly obtuse and (on the middle phyllaries) with a low, subterminal keel and apiculum, some- times purplish at the stereome-lamina junction; receptacles shallowly concave. Florets: bisexual (3-)4-6, all corollas yellowish- or sometimes purplish-tipped; pistillate numerous in a broad zone. Cypselae oblong, 0.7-0.8 mm long, tan to brownish, papillate. NESOM, NEW SPECIES OF GAMOCHAETA 725 Flowering Apr-Jul(-Oct). Mostly in coastal and near-coastal localities: dunes and other sandy sites, ocean bluffs, less commonly in clay-loam, fields, roadsides and roadcuts, ditches, cliffs, pine woods, chaparral slopes, tidal marsh edges; 0-650(-1050) m. California, Oregon, and Washington; southwestern Brit- ish Columbia. A new species from the Gulf coastal plain Another species with strongly bicolored leaves, known to me from eight states of the U.S.A. Gulf coastal plain, appears to be undescribed. Among species oc- curring in North America (including Mexico), it is similar in general aspect to Gamochaeta argyrinea because of the basally decumbent-ascending stems, white-pannose vestiture, bicolored leaves (gray-green abaxially), the basal in a persistent rosette, and mostly oblanceolate cauline leaves, but conspicuous de- tails of the vestiture, phyllary morphology, and cypselae are different. It keys to the area of G. purpurea and G. americana (P. Miller) Wedd. in Freire y Iharlegui (1997), emphazing the bicolored leaves and acute to acuminate inner phyllar- ies. Concepts of G. americana by Cabrera (1963, 1971, 1974), Cabrera and Freire (1998), and others may represent or include this North American species, judg- ing from illustrations of involucral and phyllary morphology, but G.americana sensu stricto, as interpreted here (see below), does not occur in the U.S.A. It is possible or even likely that the new species is native to South America, as it is known by relatively few and recent collections in the U.S.A. Gamochaeta chionesthes Nesom, sp. nov. (Figs. 2, 3, 4, 8). TYPE: U.S.A. GEORGIA. Meriwether Co.; town of Gay, mowed lawn of U.S. Post Office on Hwy 74/85, near Jct with 109, loamy soil, Gamochaeta coarctata , G. argyrinea, and G. chionesthes present and abundant at this site, 14 Apr 2004, G. Nesom GASCO4-14 (HOLOTYPE: BRIT, ISOTYPES: CAN GH, K, LP LSU, MO, NCU, NY, P. TEX, UNA, US, USF). Tite ] sea) ] 12] a ara | “ eres | eae | VILL textiloideo, foliis basalibus numerosis persistentibus, oe equips oblanceolatis vel oblanceolati- spathula involucris 3-3.5 mm altis, f1 purE d ay phyllariis Snnibis oe aut laminis brunneis, et cy peelis pupureis. Plants annuals to winter annuals, fibrous-rooted. Stems erect to decumbent- ascending from the base, 10-45 cm tall, simple or rarely few-branched, closely white-pannose, the vestiture sheath-like, like a continuous covering by a thin, closely appressed, polished cloth formed of filiform trichomes usually not in- dividually evident. Leaves basal and cauline, basal persisting and green through flowering, basal and lower cauline oblanceolate to oblanceolate-spatulate, 2- 6(-7) cm long, 5-13 mm wide, gradually reduced upward in size to linear bracts into the lower part of the capitulescence, not clasping, strongly bicolored, light green above but persistently lightly arachnoid with extremely closely appressed, nearly microscopic trichomes, closely white-pannose abaxially. Capitulescence cylindric in early season, mostly 3-5(-7) cm long, 10-12 mm wide (pressed), 726 BRIT.ORG/SIDA 21(2) later producing axillary glomerules from lower nodes and elongating, some- times strongly interrupted and up to 20 cm long. Involucres campanulate-cy- lindric, 3.5-4 mm long, base imbedded in cottony tomentum and lightly arach- noid on the lower 1/4-1/2; phyllaries in 4-5 series, all apically acute to acute-acuminate, outermost ovate, |/3 as long as the inner, inner oblong-lan- ceolate, with green stereome ca. 3/5 the length of the phyllary, lamina apically acute, not apiculate, lightly striate, slightly flaring outward at maturity, pur- plish coloration ape or faint and present only at stereome apex and distal margins; llowly concave. Florets: bisexual 2-4, all corollas brown- ish- yellow to ‘purple distally, sometimes purple only on adaxial surface of bi- sexual corolla lobes; pistillate numerous in a broad zone. Cypselae oblong, 0.5- 0.6 mm long, purple, papillate. Flowering (Mar-)Apr-May(-Jun). Disturbed, open sites, especially road- sides, clearings, fields, flood plains, low pastures, lawns and almost any other place that has been mowed, sandy, loamy, and clay soil; 0-200 m; Alabama, Ar- kansas, Florida, Georgia, Louisiana, Mississippi, North Carolina, South Caro- lina. Gamochaeta chionesthes is abundant and common at least in Georgia and Alabama, where it often grows with G. coarctata and G. argyrinea. The epithet (Gr. chioneos, snowwhite, and esthes, clothing) alludes to the bright white, cloth- like covering of the stems and abaxial leaf surfaces. Collections examined. U.S.A. ALABAMA. Baldwin Co.: Gulf Shores State Park, arid inland sand hills with Quercus spp., Pinus glauca, and shrubs, 11 Apr 1966, IItis 25208b (WIS); Spanish Fort, sandy field by Ala. 225, 0.5 mi N of jet US 31, 27 Mar 1971, Kral 41865 (MO). Bibb Co.: Cah: as Riverat Hwy 24, 2 mi SE of Blocton, rocky woods and low, sandy soil of floodplain, 12 May 1977, Sessler 1329 (VDI 3) Chilton Co.: S of Clanton on Interstate Hwy 65, 4.4 mi S of jet with Hwy 31/22, grassy roadsides, 17 May 2001, Nesom AL2K1-10(BRIT). Choctaw Co.: low pasture beside Tombigbee River bridge of Ala Hwy 10, 22 Jun 1966 [past mature fruit], Clark 3084 (NCU); 8.8 mi N of Toxey, longleaf pine hills, 15 Apr 1967, Kral 28371 (VDB). Conecuh Co.: Repton, sandy loam of clearing in pine flatwoods by US 84,5 May 1988, Kral 74710 (V DB). Coosa Co.: roadside on dirt road N of Peckerwood Creek, ca. 1 miS of Talladega Springs, 29 Apr 1967, Clark 11391 (NCU); site of Sears Chapel Methodist Church on Hwy 31, 2.3 mi N of jet Hwy 22 in Rockford, open field, cemetery, and roadside, rocky soil, 17 May 2001, Nesom AL2K1-7 (BRIT, CANB, LSU, MISS, UARK, UNA). Greene Co.: by small ditch in Forkland, 6 May 1967, Naugle G264 (LSUS). Lee Co.: 2-3 mi W of junction Ala Hwys 22 and 280, N of Phoenix City, shortleaf pine-oak, sandy soil, 10 May 1969, Lazor 3161 (NCU). Pike Co.: roadside of Banks Hw 5 mi N of Troy, 24 Apr 1967, Shirah 39 (NCU). Talladega Co.: ca. 3 mi S of Cues Pure (at i of Hwy 280 and 8) on Ala. Hwy 8, area of cutover gues pine-mixed hardwood, gravelly clay soil, very com- mon along roadside, 17 May 2001, Nesom AL2K1-3 (BRIT, MISS, UARK, UNA, US, USF). Winston Co.: downtown Haleyville, abandoned homesite near Central Bank, 7 Jun 1975, Whetstone 4507 (NCU). ARKANSAS. Hot Springs Co.: Malvern PO., Magnet Cove, rocky ridges, 600 ft, 24 Apr 1973, Demaree 66292 (MO). FLORIDA. Baker Co.: 5 mi W of Glen St. Mary, grassy roadside shoulders, 1 May 1959, Godfrey 58540 (V DB). Escambia Co.: N edge of Pensacola, open grassy bank of field, 1 May 1982, Correll 53821 (NY, USF); just N of Pensacola, F side of US 29 just S of its jct with Burgess Road, 20 May 1981, Wilhelm and Ladd 8859 (USF). Leon Co.: Tallahassee, common in vacant lots, 6 Apr 1958, Godfrey 56420 (NY-3 sheets). Washington Co.: dry — US Hwy 90, 3.2 mi W of Chipley, 12 May 1967, Ward 6486 (NCU). GEORGIA. Appling Co.: alon y 121 in town i Surrency, 0.2 mi S of jet Hwy 341/27, front lawn and road border of deserted house, oi Apr 2004, Nesom GASC04—38 (BRIT). Ba- NESOM, NEW SPECIES OF GAMOCHAETA 727 con Co.: along Hwy 32, 4 mi W of jet Hwy 23/4-1 in Alma, mowed roadside and lawn-like area between truck stop store and highway, sandy soil, 15 Apr 2004, Nesom GASC04—34 (BRIT). Bulloch Co.: ca. 15 mi SW of Statesboro along Hwy 321, at jct with Hwy I-16, mowed roadside area, gravelly soil, 15 Apr 2004, Nesom GASCO4—45 (BRIT). Coffee Co.: E side of city of Douglas on Hwy 32, near jet Hwy 221/135, lawn area of business, 15 Apr 2004, Nesom GASC04—32 (BRIT). Cook Co.: weedy area between highway and a railroad at Cecil, 30 Apr 1970, Faircloth 6624 (NCU). Fayette Co.: N side of Fayetteville, along Hwy 314, ca. 2 mi N of jct with Hwy 85, roadside, mowed, beside strip of pine woods, 14 Apr 2004, Nesom GASCO4—4 (BRIT, GA). Fulton Co.: S side of Atlanta area on Hwy 279 (Old National Highway) 2.5 mi S of I-85/285, saa ee Flat Shoals Road, closely mowed lawn area of quick-stop food store, 14 Apr 2004, Nesom GASCO4—2 (BRIT). Grady Co.: upland pine woods along- aa a logging trail on Balfour's Nickelville forest, 6. a mi S of Whigham, 11 Apr 1970, Faircloth 6541 reene Co.: 3 mi SW of Greenboro on Hwy 44, at jet with I-20, lawn area of commercial ae bodeane highway, 16 Apr 2004, a GASC04—57 (BRIT, GA). Lee Co.: 7 miE of Leesburg on wy 32 at jet with Hwy 91 to Albany, W side of junction, large mowed field on N side of road, drier oe roadsides, compacted sand, 15 Apr 2004, Nesom GASC04—26 (BRIT, GA, LP, MO). Montgomery 2.4 mi Eof Ailey on US 80, 18 May 1976, Solomon 5562 (MO). Morgan Co.: S side of Madison, just a Hwy 129/24 at jet with I-20, lawn area of motel, sloping toward southeast, 16 Apr 2004, Nesom GASC04—58 (BRIT). Newton Co.: S side of Covington, at jct of I-20 and Hwy 142, grassy area beside access road from I-20 to Hwy 142, 16 Apr 2004, Nesom GASC04—59 (BRIT). Screven Co.: Georgia welcome station, N side of Hwy 301 ca. 0.2 mi W of South Carolina state line, mowed lawn area, 16 r 2004, Nesom GASC04—48a (BRIT). § rCo.: Sside of Americus, at jct Hwy 280/49 and Hwy 19, mowed lot beside pecan orchard, eee oam, 15 Apr 2004, ae GAS SCO4—22 (BRIT, GAD Tatnall Co.: ca. 7 mi SW of Mendes on Hwy 169, at ject Hwy 121, mowed area with lawn grass, beside store, 15 Apr 2004, Nem GASCO4 41 (BRIT). Turner Co.: F side of Ashburn on Hwy 107, mowed, lawn- ae area between hand 15 Apr 2004, Nesom GASCO4—29 (BRIT, NCU). Warren Co.: ca NW of Norased on Hwy 278, at jet of 1-20, grassy roadside, 16 Apr 2004, Nesom GASCO4— oa LOUISIANA. Lincoln Par.: [Ruston], Illinois Central Railway right of way, clay soil, 3 May 1984, Wise 39 (DOV). Natchitoches Par.: W of Natchitoches near jct of Interstate Hwy 49 and La Hwy 6, hard- packed sandy soil in front of gas station complex, past flower and fruit, 5 Jul 2004, Nesom GA04—63 (BRIT). MISSISSIPPI. Covington Co.: right-of-way, US Hwy 49, 11.8 mi N of Hattiesburg, 8 May 1966, Temple 2746 (NCU). Scott Co.: Raworth Recreation Area, between Forest and Morton, sticky dark clay soil, open areas in loblolly pine forest, | May 1970, Jones 18493 (VDB). NORTH CAROLINA. Bladen Co.: 0.2 mi E of Cape Fear River on NC 41, dry roadside [alongside] flood plain forest, 16 May 1976, Solomon 1895 (MO). Duplin Co.: jct of Hwy 40 and Hwy 117, just N of Magnolia, grassy roadsides, edge of ditch, edge of woods, sandy soil, intermixed with G. purpurea, G. coarctata, an : G. argyrinea, 28 Apr 2001, Nesom WMGT 6 (BRIT, MO, NCU, US). SOUTH CAROLINA. Aiken Co.: S side of New Ellenton, at jet of Hwy 278 and Hwy 19, roadside area, sandy soil, lo Apr 2004, Nesom eae ey) (BRIT, LSU, NCU, USCH). Aiken Co.: area of Beech Island (SE of Augusta) near jet of Hwy 278 and Hwy 125, at Beech Island Avenue, ca. 7 mi SE of N. Augusta, roadside and ditch edges, sandy soil, 16 Apr 2004, Nesom GASC04—54 (BRIT, F, MISS, US, USCH). Allendale Co.: along Hwy 301, 5.2 mi WSW of jet Hwy 125/278 in Allendale, 8 mi ENE of Savannah River and state line, grassy roadside median of 4-lane highway, sandy soil, 16 Apr 2004, Nesom GASCO4—50 (BRIT, LP, USCH). Bamberg Co.: ca. 2 i NE of Ulmer, at jet of Hwy 301 and Hwy 321, broad lawn area of Connelly Motel, 16 Apr 2004, Nesom eee 51(BRIT, K, USCH). Barnwell Co.: SE side of Barnwell on Hwy 64, 1.8 mi ESE of jct Hwy 3 in Barnwell, mowed, sandy field beside lookout tower, 16 Apr 2004, Nesom GASC04—52 (BRIT, SCH) As noted above, Gamochaeta chionesthes is similar to G. argyrinea, but the former is distinct in significant features, including habit and phenology, vestiture, capitulescence and involucral morphology, and cypselar color. The 728 BRIT.ORG/SIDA 21(2) early-season branches of G. chionesthes lie nearly flat, with capitulescences on ascending branch tips. The early branches of G. argyrinea are basally decum- bent-ascending to ascending, but the capitulescences are more quickly borne on stems completely erect or with erect distal portions. The difference in habit is particularly evident in mid-April, because G. chionesthes is several weeks later in phenological development and usually lies nearly flat at the same time that G. argyrinea is producing abundant, erect flowering branches. Because of this, it is easy to distinguish the two species when they grow intermixed or in close proximity, as is often the case. Additionally, the stems and leaves of G. chionesthes are stiff, almost brittle-feeling, compared to the softer ones of G. argyrined. The cauline vestiture Gamochaeta chionesthes is cloth-like, formed of fili- form trichomes usually not individually evident but united in a continuous covering like a thin, closely appressed, polished cloth; the abaxial leaf vestiture sometimes is similar. In the closely pannose cauline vestiture of G. argyrined and G. purpurea, the trichomes usually are individually evident in their longi- tudinal orientation; the cauline vestiture of G. coarctata also usually is cloth- like, similar to that of G. chionesthes. The “tightening” of the individual tri- chomes apparently is accentuated during drying, because this feature is more easily observed on herbarium specimens. The phyllaries of Gamochaeta chionesthes are evenly graduate in length, all apically acute and usually the whole lamina or at least the apex of each is brown, usually giving the whole involucre a distinctly brown hue; purplish coloration is absent or faint and present only at stereome apex and distal mar- gins. The brown hue of the involucres is a good “field” character even without a lens; with a field lens, the difference in phyllary shape between G. chionesthes and G. argyrined is easily evident. And finally, if mature cypselae are present, the purple ones of G. chionesthes are distinct from those of similar U.S.A. spe- cies, which are tan. Gamochaeta chionesthes is contrasted individually with both G. purpurea and G. argyrinea in the following pair of couplets. In the context of all U.S.A. species with strongly bicolored leaves, G. chionesthes also is identified in a key further below. — Gamochaeta chionesthes contrasted with G. purpurea: a. Cauline vestiture a white, closely appressed, cloth- like ¢ covering, trichomes usually not ae evGent basal leaves abundant and | ;cauline leaves oblan eolat surfaces aire not hairy but with ie minutely thin, cl losely a sed, sheath-like or cloth-like covering (some- times incomplete) without individually avieat trichomes; involucres 3-3.5 mm high; florets mostly yellowish at the apices; inner phyllaries with brown-tinted lamina—purplish coloration, when present on phyllaries, aa and pe at the ape and distal margins of the stereome; cypselae purple Sane chionesthes NESOM, GAMOCHAETA 729 a. Cauline vestiture densely but loosely pannose or pannose-tomentose; basal leaves often few or not persistent; cauline leaves distinctly spatulate; adaxial leaf surfaces loose staennoid trichomes with basal cells expanded and vitreous; involuc vu! plish-tipped;i Inner phy WatiIlS with whitish or ee silvery lamina at maturity, often purplish when young; cypselae tan Gamochaeta purpurea Gamochaeta chionesthes contrasted with G. argyrinea: a. Cauline vestiture a white, closely appressed, cloth-like covering, trichomes usually not individually evident; phyllaries evenly graduate in length, all phyllaries apically erect, acute to acute-acuminate, and with brown apex or lamina; bisexual florets 2— 4;cypselae purple Gamochaeta chionesthes . Cauline vestiture closely pannose with trichomes individually evident, minutely fili- form and enaivaliay oriented, the vents Hbcommmenly cloth-like; phyllaries unevenly graduate in length, outer iddle ph te to ovate-lanceolate, apical) cle to a ae inner ohyllaries with lamina oblong with apex slightly spr 1, truncate-r nd apiculate, hyaline-translucent and slight brown; ee ere 5(- sn tan Gamochaeta argyrinea Identity of Gamochaeta purpurea sensu stricto Gamochaeta purpurea (Figs. 3, 4, 5) is distributed widely through the world, but the name has been used in various contexts in the U.S.A, often to identify any plant of Gamochaeta with bicolored leaves. A narrower, more accurate con- cept is documented here. Gamochaeta purpurea (L.) Cabr., Bol. au Argentina Bot. 9:377. 1961. Gnaphalium purpureum L., Sp. PL. 854. 1753, Type: Linnaeus noted “Habitat in Carolina, Virginia, Pensylvania.” A Kalm collection a fiche!) may be part of the type material. Two speci- mens in the Clayton Herbarium (BM) were annotated by James Reveal in 1990 as syntypes: U.S.A. In Virginia, J. Clayton 385 (GH- ae internet images! at ). Accessed March 2 Gnaphalium rosaceum LM. Johnston, Contr. Gray Herb. ser. 2, 68:99. 1923. Gamochaeta rosacea (1.M. : hnst.) Anderb., Opera Bot. 104:157. 1991. Tyre: MEXICO. SAN 878, CC. Parry and E. Palmer 426 (HoLotyre: GH). cna heteroides Klatt, Linnaea 42:137. 1878. Tyr: MEXICO: locality unspecified, Ehrenberg 972 (GH fragment and drawings!). _UIS Potosi: region of San Luis Potosi, Plants winter annual or annual, fibrous-rooted or slender-taprooted. Stems erect to basally ascending-decumbent, 10-40(-50) cm, densely but loosely pannose or pannose-tomentose. Leaves basal and cauline, oblanceolate-spatulate to spatulate, basal and lower cauline 1-6 cm long, 5-14 mm wide, persisting or withering at flowering, similarly shaped leaves continuing into at least the lower part of the capitulescence, sometimes closely sinuate on the margins, usually strongly bicolored, closely white-pannose abaxially, adaxial surfaces loosely and sparsely arachnoid, the trichomes with basal cells expanded and vitreous, often glabrescent but at least the basal cells of each trichome persisting. Capitulescence initially a continuous cylinder 1-4(-5) cm long, (5-)10-15 mm 730 BRIT.ORG/SIDA 21(2) ae eer g c ¥ ai = Bi: a, 8 2 a & 7 ‘a _ 3 ae Ss 4 “ues be. ; ai x SMe cae = a A ee " G Pon, \ ae ee / some NN 3 OB ES, / 3 gy a ae a ——, oe 2 ar x _ 2 ~t as ae | \ 3 > Gamochaeta purpurea Fic. 5. State-level distribution of Gamochaeta purpurea. wide, the arrangement becoming interrupted and elongate, of widely separated, bracteate glomerules, with lower axillary glomerules often on long peduncles. Involucres turbinate-cylindric, 4-4.5 mm long, base imbedded in cottony to- mentum and lightly arachnoid on the lower 1/3-1/2; outer phyllaries ovate- triangular, inner triangular-lanceolate, apically acute, usually with a promi- nently striate texture, lamina purplish when young, becoming whitish or slightly silvery at maturity; receptacles shallowly concave. Florets: bisexual flo- rets 3-4, all corollas usually purplish-tipped; pistillate numerous in a broad zone. Cypselae oblong, 0.6-0.7 mm long, tan, papillate. 2n = 14, 28, but identi- ties of vouchers for these chromosome reports need to verified. Flowering Apr-May(-Jun). Open, usually disturbed habitats, roadsides, fields, woodland clearings and edges, almost always in sand; Canada (Ontario); USA (Arkansas, Arizona, Alabama, Connecticut, Delaware, District of Colum- bia, Florida, Georgia, Illinois, Indiana, lowa, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachussetts, Michigan, Mississippi, Missouri, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Rhode Island, South Caro- lina, Tennessee, Texas, Virginia, West Virginia); Hawaii; Mexico (Nuevo Le6n, San Luis Potosi, Veracruz, Puebla, Michoacan, Chihuahua, Sonora), Central America (Nicaragua), Antilles (Hispaniola); also reported in South America and as adventive in other parts of the world. In the western U.S.A., G. purpurea has — NESOM, NEW SPECIES OF GAMOCHAETA 731 Gamochaeta argyrinea eee | dictrihuti ££ 1 eee : With . kou deni . Fic. 6.C County- I t , the lt -ayil l r ¢ That liad | ). South Carolina: Geaiuti Alabama, Mississippi and Aikaieae Georgia reflects the route of a reaecting il) made in wnat 2004. The only known record for Pennsylvania P been recorded only from Arizona (see comments and documentation in Nesom 2004). Gamochaeta purpurea commonly occurs in disturbed sites, but at least in the eastern and southeastern USA, it seems more often to occur in more nearly natural sites, such as woodland edges and clearings, it is true to sandy soil, and it is my impression that in the last 50 years, G. purpurea probably has become much less common. Gamochaeta purpurea is uncommon compared to G. argyrinea, G. chionesthes, and G. coarctata (personal observation), in the range of the latter three. The species is rare or extirpated in the northeastern US.A. (summary in Kartesz 1999), where the latter three do not occur. Gamochaeta purpurea was excluded from Wisconsin’s flora by Wetter et al. (2001), but Cochrane (pers. comm. 2004) notes that an undated collection from Sheboygan by Charles Goessl (WIS, photocopy-BRIT!) probably should be accepted as a valid record, although it probably was a garden weed or waif; it remains the only collection for the state. The species is described by Voss (1996) as “doubt- less adventive” in the few southern counties of Michigan from which it is known. Gamochaeta purpurea is recognized by its mostly spatulate cauline leaves, BRIT.ORG/SIDA 21(2) 732 \ _-—— a ; en ie Gamochaeta ustulata | ) ir cae nae 1. Fic. 7, County-level distribution of Gamochaeta ustulata. NESOM, NEW SPECIES OF GAMOCHAETA 733 Fic.8. County-| J common than indicated by the map, la 1 * rd } va hi +h Ae] bey H J Als} hi - _ L fl ll 1 it probably extends across South Carolina. The loop shown in Georgia and at eee. L L L £ 1: 1 7%. . | itt ¢¢h | J in every county sampled. involucres 4-4.5 mm high, and inner phyllaries with whitish-hyaline, erect, apically acute lamina. The purple-tipped corollas are visible through the trans- lucent lamina. The base of the trichomes on the adaxial leaf surfaces also is a diagnostic character—the basal cells of each trichome are expanded and glassy (use a lens), compared to comparable trichomes of most other species, which are evenly filiform to the very base. The pronouncedly spatulate leaves are dis- tinctive and with experience, one can usually distinguish this species from oth- ers by leaf shape, but it is admittedly a subtle difference and the leaf dimen- sions of G. purpurea are similar to those in other species. In localities from Maryland northward, plants of G. purpurea are clearly annual, usually produc- ing a small basal rosette and very shallow fibrous roots or a filiform taproot; southward and southwestward from Maryland, the basal rosette tends to be larger and the fibrous roots denser, and plants may be winter annual in this region, or at least the longer growing season is reflected in the plant habit. | have identified Gamochaeta purpurea in various localities in Mexico (state BRIT.ORG/SIDA 21(2) Fic. 9. County-level distribution of Gamoct implicicaul citations above) at elevations of 1300-2800 meters. Phyllaries of these plants characteristically are distinctly brownish on the lamina and apices, but the plants otherwise are similar to those of the U.S.A Gamochaeta coarctata vs. Gamochaeta americana in the USA. Plants of the U.S.A. previously identified as Gamochaeta americana (e.g., Nesom 1990, 1999) are here identified as Gamochaeta coarctata (Willd.) Kerg. (Figs. 3, 4, 10). Godfrey (1958) correctly identified these plants as Gnaphalium spicatum, although the latter name is now treated as a synonym. Gamochaeta coarctata occurs mostly in the southeastern states and also apparently is naturalized in California (see Nesom 2004). — coarctata (Willd.) Kerg., L ejeunia 120:104. 1987. Gnaphalium spicatum Lam., Encycl. Meth. Bot. 2:757. 1788, hom. illegit. (non P Miller 1768). esa acta um “ie nom nov, Sp. PI. 3(3):1886. 1803. Gnaphalium radians Benth. var. spicatum (Lam.) Klatt, nom. illeg., Linnaea 42:140. 1878. Gnaphalium purpureum L. var. spicatum (Lam.) oie in Mart. nom. illeg., Fl. Bras. 6(3):125. 1882. Gamochaeta spicata Cabr, nom. nov. illeg., Bol. Soc. Argent. 9:380. 1961. HOLOTYPE: [URUGUAY]. “In Monte-video,” Commerson s.n.(P-LAM, IDC mi- crofiche 6207.325.L.1)). Cabrera (1961, p. 380-381) cited as “lectotype” of Gnaphalium spicatum Lam. “Des environs de Buenos Ayres,” without date, Commerson s.n. Lamarck’s protolo gue, NESOM, NEW SPECIES OF GAMOCHAETA 735 LAs ete hits tf. } tates Atl + ele aL +t H lind |} | A Fic. 10. County | here, the distrit t £ s ralte, * “snd in 1 90NA)\ } spread. C i however, specified the locality as “Monte-video,” as does the P-LAM sheet cited above as holo- type. See Pruski and Nesom (2004) for discussion of the typification of this species. Plants winter annual or biennial(?) herbs, fibrous-rooted. Stems basally decum- bent-ascending, 15-35(-50) cm tall, usually several from the base, white- pannose, the tomentum usually sheath-like. Leaves basal and cauline, basal in a persistent rosette, spatulate to oblanceolate-obovate, (1.5-)3-8(-12) cm long, 6-15(-22) mm wide, cauline gradually or little reduced in size above the basal, not clasping or decurrent, slightly succulent and often becoming crenulate- margined upon drying, strongly bicolored, the adaxial surfaces green and glabrate tocompletely glabrous, closely white-pannose abaxially. Capitulescence usually a dense and continuous cylinder 2-20 cm, becoming branched and highly interrupted in older plants with elongation of branches at lower nodes. Involucres cylindric-campanulate, 2.5-3 mm high, shiny and completely gla- brous from base to apex; phyllaries: outermost elliptic-obovate to broadly el- liptic ovate with rounded to obtuse apices, often slightly but distinctly pur- plish or rosy, 1/3-1/4 as long as the inner, inner witha distinctly brown-hyaline, apically rounded to blunt, apiculate lamina; receptacles shallowly concave. Flo- rets: bisexual 2-3, all corollas usually purplish-tipped; pistillate numerous ina broad zone. Cypselae oblong, 0.5-0.6 mm long, tan, papillate. 2n = 28. 736 BRIT.ORG/SIDA 21(2 Flowering Apr-Jun. Ditch banks, roadsides, lawns, fields, gardens, sidewalk cracks, shaded spots around buildings; 0-150 m; U.S.A. (Alabama, Arkansas, California, Florida, Georgia, Louisiana, Mississippi, North Carolina, South Caro- lina, Texas, Virginia); Mexico, Antilles lamaica, Puerto Rico), South America, Europe, Japan, Taiwan, Australia, New Zealand. In the southeastern USA, Gamochaeta coarctata commonly grows with G. argyrinea and G. chionesthes. Collections documenting the occurrence of this species in Arkansas, Califor- nia, and Virginia are cited in Nesom (2004). Plants of Gamochaeta coarctata are recognized by their persistent, slightly succulent, strongly bicolored leaves (green and glabrate or usually completely glabrous on the adaxial surfaces), involucres 2.5-3 mm high, completely gla- brous and evidently purplish-tinted, outer phyllaries elliptic-obovate to broadly elliptic ovate with rounded to obtuse apices, and 2-3 bisexual florets per head. It usually can be distinguished at a glance from G. argyrinea and G. chionesthes because of the distinctive vestiture and involucres. The status of Gamochaeta americana Gamochaeta americana (Figs. 3, +) is another species with strongly bicolored leaves. Its occurrence in the U.S.A. has not been documented, which is remark- able in view of its widespread distribution in the Antilles and Mexico. The name has been incorrectly used to identify species of the U.S.A. (Nesom 1990). The type collection of Gamochaeta americana was made from Jamaica (below), and I have studied collections from Jamaica, Puerto Rico, Hispaniola, and Cuba (all at NY) and from South America, Central America, and Mexico (various herbaria). It is reported from New Zealand (Drury 1971; Webb 1988) and Australia (Everett 1990). Gamochaeta americana was designated lectotype of the genus by Cabrera (1961, p. 362). Gamochaeta americana (P. Mill.) Wedd., Chlor. Andina 1:15]. 1856. Gnaphalium americanum P. Mill, Gard. Dict. ed. 8, no. 17.1768. Gnaphalium purpureum L. var. americanum (P. Mill) Klatt, Linnaea 42:140. 1878. LECTOTYPE (Fawcett & Rendle 1936, p. 206): JAMAICA. 1731, Houstoun s.n. (BM, N Y-photo!). Gnaphalium guatemalense Gandoger, Bull. Soc. Bot. France 65:42. 1918. Gamochaeta guatemalensis (Gandoger) Cabr., Bol. Soc. Argent. Bot. 9:371. 1961. Type: GUATEMALA. Alta Verapaz, Tuerckheim (not seen). Gamochaeta irazuensis Nesom, Phytologia 68:199. 1990. Type: COSTA RICA. Volcan Irazt, 10,000- 11,330 ft, | Dec 1937-1 Jan 1938, PH. Allen 702 (HOLOTYPE: FI). Plants annual to short-lived perennial herbs from a slender, short but lignescent taproot, shallow fibrous roots, or a short, fibrous-rooted rhizome or caudex re- gion. Stems usually erect from the base, less commonly decumbent-ascending, 10-40(-65) cm tall, arising singly or less commonly with 2-3 shoots, decum- bent stems often developing adventitious roots, loosely lanate-tomentose (not sheath-like). Leaves basal and cauline, basal obovate-oblanceolate, 3-7(-10) cm long, 4-12-16) mm wide, usually withering or withered by flowering and not NESOM, NEW SPECIES OF GAMOCHAETA 737 persisting in a rosette, lower and midcauline often distinctly subclasping (but not auriculate), often slightly decurrent, cauline oblong to oblong-oblanceolate, sometimes narrowly revolute, adaxial surfaces glabrous to glabrate, sometimes more densely and persistently hairy, abaxial surfaces densely white to gray-white pannose. Capitulescence a continuous spiciform cluster 1-6 cm long, 8-12(-15) mm wide (pressed), usually becoming much longer and strongly interrupted by development of axillary clusters from lower nodes. Involucres campanulate- cylindric, 4-4.5 mm high, shiny and completely glabrous from base to apex; phyllaries in 4-5 series, outermost ovate to lanceolate-ovate with broadly acute to nearly obtuse apices, 1/4-1/2 as long as the inner, inner oblong-lanceolate, with light green stereome 2/3 the length of the phyllary, lamina dark brown- hyaline, apically broadly acute, sometimes nearly rounded, to (less commonly) obtuse apiculate, erect, without purplish coloration or slightly purplish only around the stereome-lamina junction; receptacles shallowly concave. Florets: bisexual ca. 3-6, all corollas yellowish distally; pistillate numerous in a broad zone. Cypselae oblong, 0.5-0.6(-0.7) mm long, tan, papillate. Flowering all year, perhaps most abundantly Jun-Sep. Llanos, openings in pine, pine-oak, and deciduous woods, fallow fields, pastures, roadsides and other disturbed sites, commonly in wet or moist soil; 1250-3200 m. Mexico (Chiapas, Chihuahua, Colima, Dist. Federal, Durango, Guerrero, Hidaldo, Jalisco, México (Edo.), Michoacan, Morelos, Nuevo Leon, Oaxaca, Puebla, San Luis Potosi, Sinaloa, Sonora, Veracruz); Central America (Costa Rica, El Salvador, Guatemala, Hon- duras, Nicaragua, Panama); Antilles (Cuba, Hispaniola, Jamaica, Puerto Rico); South America; Galapagos Islands. Gamochaeta americana is generally recognized by erect stems, strongly bicolored leaves with glabrate to glabrous adaxial surfaces, the cauline oblong to oblong-oblanceolate and relatively narrow, commonly subclasping and slightly decurrent, heads clustered in an interrupted spike, involucres shiny and completely glabrous, usually with a distinctly brownish cast. The lamina of the inner phyllaries characteristically are dark brown and sharply delim- ited in color from the proximal portion of the phyllaries; often the whole in- volucre is infused with brown color. Considerably more variation is expressed within Gamochaeta americana, at least as it is understood here, than in similar species in the U'S.A. In Central America and Mexico, plants of G. americana commonly appear to be short- lived perennial, or at least biennial, because of the development of fibrous roots from a short caudex or rhizome or adventitious roots from the lower portion of decumbent stems. In the Antilles (including the type locality, Jamaica), South America, and scattered through the Mexican and Central American range are plants with shallow fibrous roots or a thin taproot—these plants apparently distinctly annual. Leaves of Gamochaeta americana are characteristically strongly bicolored, 738 BRIT.ORG/SIDA 21(2) with adaxial surfaces glabrous to glabrate, but in western Mexico (mostly Durango, Chihuahua, and Sonora), there is a tendency for the adaxial surfaces to be persistently hairy, sometimes resulting in a weakly bicolored appearance. Similar plants also appear sporadically in other parts of Mexico, and Dillon and Sagastegui (1991) noted that similar variation also occurs in South America. In Cordillera de Talamanca region of Costa Rica, a distinctive race with dense, silvery-white vestiture on abaxial leaf surfaces and with sheath-like cauline vestiture is being described as a new species (Nesom & Pruski in prep.). Identification of Gamochaeta species in the U.S.A. with strongly bicolored leaves The following key distinguishes the six species of Gamochaeta with strongly bicolored leaves currently known to occur in the U.S.A. Gamochaeta americana is included so that the key might be used in regions outside of the U.S.A., but its position in the first couplet directs the key toward contrasts of species known within the U.S.A. The abaxial leaf surfaces of all of these are white-pannose with a closely matted vestiture completely obscuring the epidermis, while the adaxial surfaces are evidently green to grayish green, glabrous to sparsely arach- noid. Five of these species occur primarily in the eastern and southeastern U.S.A. Gamochaeta ustulata occurs mostly in coastal and near-coastal Pacific locali- ties in the western U.S.A. and British Columbia. From observations in herbarium and field, Gamochaeta species in the U.S.A. appear to be remarkably constant in diagnostic morphological features, par- ticularly in the vestiture of stems, leaves, and phyllaries and in the shape, size, and coloration of the phyllaries. The species common grow closely intermixed, but intermediates that might indicate hybridity are not common, or at least they are not often observed. In contrast, the capitulescence may vary in degree of compactness or openness to a greater degree than has been generally recog- nized. In G. purpurea, G. argyrinea, and G. coarctata, for example, the capitulescence begins as an essentially uninterrupted cylinder but may elon- gate greatly and become highly interrupted and branched. Keys that utilize a difference of this sort in capitulescence morphology (e.g., Freire & Tharlegui 1997) probably are recognizing artificially separated taxa. 1. Basal leaves usually withering or withered by flowering and not persisting in a ro- sette, cauline oblong to oblong-oblanceolate, lower and midcauline often distinctly subclasping (but not auriculate), often slightly decurrent Gamochaeta americana 1. Basal leaves usually persisting in a rosette at flowering (except in G. simplicicaulis), cauline spatulate to oblanceolate, oblanceolate-oblong, or oblanceolate-obovate none clasping or decurrent. 2. Basal and lower cauline leaves on elongate internodes, usually withered or ab- sent at flowering, clusters of small leaves usually produced in cauline axils; stems strictly erect, (30-)50-85 cm tall;inner phyllaries apically acute; flowering mostly July-August amochaeta simplicicaulis Basal and lower cauline leaves on relatively congested nodes, usually green and persistent at flowering, clusters of small leaves absent in cauline axils; stems erect N NESOM, NEW SPECIES OF GAMOCHAETA 739 to decumbent-ascending, mostly less than 50 cm tall; inner phyllaries apically acute to obtuse or rounded; flowering mostly April-June 3. Adaxial leaf surfaces usually completely glabrous; involucres 2.5-3 mm high completely glabrous, with evident purplish coloration; outer phyllaries ellip- tic-obovate to broadly elliptic-ovate, apically rounded to obtuse; ite flo- ts 2-3 mochaeta coarctata 3. Adaxial leaf surfaces with persistent vestiture (sometimes necessary to view at 10x);involucres 3-4.5 mm high, base imbedded in cottony tomentum and often lightly arachnoid on the lower 1/5-1/2, with or without purplish col- oration; outer phyllaries ovate, apically acute to acute-acuminate; bisexual flo- rets 2-6 per head. 4. Stems usually apy ing hairy but with a white, closely apt d,cloth- like covering, trichomes usually not individually evident; involace: 3-3.5 mm high; all phyllaries apically acute to acute-acuminate and brownish; bisexual florets 2-4 per head; cypselae purple mochaeta chionesthes 4. Stems usually closely white-pannose with trichomes individually evident minutely filiform and longitudinally gl re 3-4.5 mm high; inner phyllarie apic | ute or oblong an nt-apiculate, at least outer and mid ohyllaries not apically brownish; bisexual florets 3-6 per head; cypselae tan. Cauline leaves most! late, trichomes of adaxial surfaces with basal cells expanded and esis involucres 4-4.5 mm high; inner phyllaries with lamina triangular, apically acute but not apiculate; bisexual florets 3-4 per head; fibrous-rooted or slender-taprooted Gamochaeta purpurea 5. Cauline leaves oblanceolate to oblanceolate-oblong or oblanceolate- obovate, trichomes of adaxial surfaces filiform from base to tip; involu- cres 4.5—5 or 3-3.5 mm high;inner phyllaries with lamina oblong, apically truncate-rounded and apiculate; bisexual florets (3-)4-6 per head; fi brous-rooted, rarely slender-taprooted. 6. Capitulescence 12-18 mm wide (pressed), uninterrrupted or rarely so and only at the base, mostly 1-6(-8) cm long; involucres 4.5-5 mm high; outer phyllaries and often lamina of inner phyllaries dark brown, the whole head eclaaiaelny, with a dark brown or eile brown cast; mid phyllaries with a low subterminal keel; cypsela 0.8 mm long Gamochaeta ustulata Capitulescence 10-12 mm wide (pressed), early uninterrupted and 5-5 cm long, later producing axillary glomerules from lower nodes delongating,| lyi pted and up to 18cm long; involucres 3-3.5 mm high; phyllaries ch Ur to stramineous or golden, sometimes light brown but never dark brown, the who Be nad not with a dark b 0.6 mm lon t; phyllaries not keeled; cypselae 0. ae argyrinea ACKNOWLEDGMENTS Staffs of GH, MO, NCU, NY, and US were helpful during recent study at their institutions, ARIZ, ASU, DOV, HSC, LSUS, MO, TEX, UPRRP, USCH, and USF sent loans of specimens, Nancy Elder (University of Texas Life Science Library) helped with literature, and Tom Zanoni (NY), Emily Wood (GH), and Ted Cochrane (WIS) provided specimen data. Review comments by John Pruski and 740 BRIT.ORG/SIDA 21(2) Susana Freire are appreciated. It also seems appropriate to acknowledge the obsessive compulsion of people and agencies in the southeastern U.S.A. to mow, mow, mow to nearly dirt level, creating a continuous and expansive habitat of fields, lots, lawns, and roadsides that Gamochaeta coarctata, G. argyrinea, and G. chionesthes spread across. These species and other colonizers can only be expected to become more abundant. REFERENCES Aronso, R.M.L. 1984. Contribuigao para o conhecimento do género Gamochaeta Weddel em Portugal continental e insular. Bol. Soc. Brot., Sér. 2, 57:113-127. ANpeRBERG, A.A. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Bot. 104:5-195, Casrera, AL. 1961.Observaciones sobre las Inuleae-Gnaphalinae (Compositae) de América del Sur. Bol. Soc. Argent. Bot. 9:359-386. Casrera, A.L. 1963. Flora de la Provincia de Buenos Aires. Parte VI. Compuestas. Coleccion Cientifica del ILN.T.A. Buenos Aires, Argentina. Caprera, A.L.1971.Flora Patagonica. Parte VIl. Compositae. Collecion Cientifica del ILN.T.A., Buenos Aires, Argentina. Caprera, A.L. 1974. Flora llustrada de Entre Rios (Argentina). Parte VI: Dicotiledoneas Metaclamideas (Gamopétalas) B: Rubiales, Cucurbitales, Campanulales (Incluso Compuestas). Collecion Cientifica del |.N.T.A., Buenos Aires, Argentina. Casrera, A.L. 1978. Flora de la Provincia de Jujuy. Parte X.Compositae. Coleccion Cientifica del |.N.T.A., Tomo XIll, Buenos Aires, Argentina. Caprera, A.L. and S.E. Freire. 1998. Flora del Paraguay—27. Compositae V. Conservatoire et Jardin botaniques, Ville de Genéve. Missouri Botanical Garden. Ditton, M.O. and A. SacAstecui A. 1991a. Flora of Peru. Family Asteraceae: Part V. Fieldiana, Bot. (n. ser.) 26:1—70. Ditton, M.O.and A. Sacdstecui A.1991b. Sinopsis de los generos de Gnaphaliinae (Asteraceae- Inuleae) de Sudamerica. Arnaldoa 1:5-91, Drury, D.G. 1971.The American spicate cudweeds adventive to New Zealand (Gnaphalium sect. Gamochaeta-Compositae). New Zeal. J. Bot.9:157-185. Everett, J. 1990. Gnaphalium. Harden, G.J., ed. Flora of New South Wales, Volume 3. Royal Botanic Garden, Sydney, Australia. Pp. 206-210. Fawcett, W.and A.B. Renbte. 1936. Flora of Jamaica. Vol. VII. Dicotyledons:Families Rubiaceae to Compositae. British Museum (Natural History), London. Freire, S.E.and L. lHartecul. 1997. Sinopsis preliminar del género Gamochaeta (Asteraceae, Gnaphalieae). Bol. Soc. Argent. Bot. 33:23-35 Goorrey, R.K. 1958. A synopsis of Gnaphalium (Compositae) in the southeastern United States. Quart. J. Florida Acad. Sci. 21:177-184. Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. In: Kartesz, J.J. and CA. NESOM, NEW SPECIES OF GAMOCHAETA Meacham. 1999. Synthesis of the North American flora. North Carolina Botanical Gar- den, Chapel Hill, NC. KerGueLen, M. 1987.Données taxonomiques, nomenclaturales et chorologiques pour une révision de la flore de France. Lejeunia 120:1-264. Nesom, G.L. 1990. The taxonomic status of Gamochaeta (Asteraceae: Inuleae) and the spe- cies of the United States. Phytologia 68:186-198. Nesom, G.L. 1999. Gamochaeta simplicicaulis (Asteraceae: Gnaphalieae) in four southeast- ern states and new for North America. Sida 18:1259-1264. Nesom, G.L. 2004. New distribution records for Gamochaeta (Asteraceae: Gnaphalieae) in the United States. Sida 21: 1175-1185. Pruski, J.and G.L. Nesom. 2004. Gamochaeta coarctata, the correct name for Gamochaeta spicata (Compositae: Gnaphalieae). Sida 21:711-715. Voss, E.G. 1996. Michigan flora. Part Ill: Dicots (Pyrolaceae—Compositae). Cranbrook Inst. Science Bull. 161 and Univ. of Michigan Herb. Ann Arbor. Weer, C.J. 1988. Gnaphalium. In: Webb, CJ., W.R. Stokes, and PJ. Garnock-Jones. Flora of New Zealand, Volume IV. Naturalized Pteridophytes, Gymnosperms, Dicotyledons. Botany Division, D.S.I.R., Christchurch, New Zealand. Pp. 234-249. Werter, M.A., 1.S. CocHrane, MLR. Buack, H.H. Iutis, and PE. Berry. 2001. Checklist of the vascular plants of Wisconsin. Dept. of Natural Resources, Tech. Bull. 192. Madison, Wisconsin. — BRIT.ORG/SIDA 21(2) BOOK REVIEW Gardener’s Guide for Hemerocallis JOHN P.Peatand Tep L. Petit. 2004. The Daylily: A Guide for Gardeners. (0-88192- 666-3, hbk.). Timber Press Inc. 133 S.W Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $29.95, 200 pp., 200+ color photographs and illustrations, 7 3/4" x 91/4". Publisher Comments: “Few perennials are as tough and versatile as daylilies (genus Hemerocallis), and even fewer offer daylilies’ enormous range of color, shape, and growing characteristics. The ease of hybridizing daylilies is a major attraction for the enthusiast. Any backyard gardener can hybrid- ize daylilies, but this blessing of easy breeding can also be something of a curse to the newcomer. Tens of thousands of new daylilies are bred each year. How to choose and grow daylilies amidst this profusion? John Peat and Ted Petit have come to the rescue in this authoritative overview of all as- pects of daylily history, cultivation, and breeding. Inspired by R.W. Munson Jr's classic treatment, Hemerocallis, they fully describe the history of the modern daylily. In the heart of the book, they detail the various types of hybrids and provide indispensable advice for growing all of them well. More than 200 beautiful color y graphs and illustrations round out the work.” Hemerocallis is an ornamentally and economically important genus of ca. 15-30 species na- tive to eastern Asia. Growers and hybridizers have developed an amazing number of cultivars over the years. According to the authors there are more than 50,000 icles cays ane enone the ones to include in the aii wasa hae ult task, Actually tl | their that were introduc 1992. So if you're looking for a cultivar older than 1992 youare out of luck. Besides the many colorful pics the new cultivars, there is practical information on landscaping with daylilies, horticultural requirements, and daylily pests and diseases. IF you love daylilies you will enjoy this book.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 21(2): 742. 2004 A NEW SPECIES OF PTILIMNIUM (APIACEAE) FROM THE ATLANTIC COAST Alan S.Weakley Guy L.Nesom University of North Carolina Herbarium (NCU) Botanical Research Institute of Texas (BRIT) North pa oe Botanical Garden 0 e Roe gels ee 3280 Fort Worth, Texas 76102-4060, U.S.A. UPHVET SHY OF INOFUT rolin Chapel Hill, North Colina pe eo U.S.A. ABSTRACT Ptilimnium ahlesii Weakley & Nesom, sp. nov, is described from localities in eight ties of North Carolina, South Carolina, and Georgia in tidal freshwater marshes within 20 fale of the Atlantic coast. The species has been previously identified as P. capillaceum but differs from the latter in its conspicuously large ee and fruits, which at maturity are 2.7-4.2 mm long and 1.8-2.2 mm wide (vs 1.4-2.5 mm long and 0.8-1.2 mm wide in P. capillaceum), in its less-divided leaves, and in its in- florescences with fewer ee lets per umbel and fewer flowers per umbellet. The new species also flowers and fruits earlier than P. copicceum Mey early June, Le May EERE VS. Seance any AU gust, July-September, respectively). Pt brackish marshes, while P capillaceum has a wider geographic range and ecological Blerdnce ae the two species grow in close proximity in their area of sympatry. RESUMEN Se describe Ptilimnium ahlesii Weakley @ N sp. nov, Mies varias ulocaleades en ocho condados de Carolina del norte, Carolina - SoryiGesnanen| d banda de 20 omens en 7 costa atlantica. La Spscle ha sido previamente identificada como P. dae pero mm los Oval 10S yf de longitud y 1.8-2.2 mm de anchura (contra 1.4-2.5 mm de nna y 0.8-1.2 mm de ae enP. capillaceum), por sus hojas menos divididas, y por sus inflorescencias con menos umbélulas por umbels neues flores por umbélula. La nueva Se también florece antes que P. capillaceum junio, finales de mayo-julio, vs. mediados de junio-principios de agosto, julio- sent genie aie eee Ptilimnium siecle es tara y restringida a charcas de agua dulce o ligeramente salobres, mientras que P. eo miee tiene un kanes pore. y tolerancia ecoldgica mas amplia, pero las dos especies crecen muy p n impatria Anexceptionally large-fruited form of Ptilimnium has been observed and col- lected in coastal counties of North Carolina, South Carolina, and Georgia (Figs. 1, 2,and 3). This plant apparently drew the attention of the late Harry E. Ahles, then working at the University of North Carolina Herbarium (NCU), and speci- mens at NCU include annotations in his hand indicating his opinion that they represented an undescribed species to be named “P. macrospermum.” In the Caro- linas “Guide” (Radford et al. 1964), these plants were included in the key and species summaries as “Ptilimnium macrospermum Ahles.” The following year, it was mapped in the Carolinas “Atlas” in Brunswick, New Hanover, and Pender SIDA 21(2): 743-752. 2004 744 BRIT.ORG/SIDA 21(2) counties, North Carolina, and Georgetown and Jasper counties, South Carolina (Radford et al. 1965). Three years later, in the first printing of the Carolinas “Manual,” this entity was placed as a synonym of P. capillaceum (Michx.) Raf. with the notation that the name was “a nomen nudum’ attributed to Ahles, but was confusingly included in the key as “P. macrospermum’ (Bell 1968, in Rad- ford et al.); in later printings it was also removed from the key, presumably con- firming the intent of Bell not to include the taxon. Bell (pers. comm., June 2004) states that he was uncertain of the taxonomic distinctiveness of the taxon, be- cause of the paucity of herbarium material available, and as it had not been validly published he decided not to include it in the “Manual” (see Sorrie 1997 for examples of other unpublished names by Ahles). The name in similar form has resurfaced in a widely used database (Kartesz 1999), as “Ptilimnium macrospermum Kartesz, sp. nov. ined.” Additionally, the taxon has been considered to be of conservation concern by the network of Natural Heritage Programs and has been tracked as a rare species under the unofficial name “Ptilimnium sp. I’ in North Carolina and Georgia (Franklin 2004, Georgia Natural Heritage Program 2004). The potential conservation importance of the putative taxon demands that this taxonomic and nomencla- tural confusion be resolved, and the species has been the subject of field and herbarium study by the authors since 1990. With these additional observations, we conclude that this plant is a distinctive and undescribed species. We are pleased to provide it with a valid name, allowing it to proceed in polite com- pany, rather than under a twice-naked name (“Ptilimnium macrospermum’), a non-name (“Ptilimnium sp. I”), or inappropriately ignored under Ptilimnium capillaceum (Michx.) Raf. The new epithet validated here honors the original insight of Harry E. Ahles regarding this, as well as his general contributions to the understanding of the flora of the Carolinas through his voracious collecting. Ptilimnium ahlesii Weakley & Nesom, sp. nov. (Figs. 1-3). Type: NORTH CAROLINA. Brunswick Co. tidal freshwater marsh of the Brunswick River, just E of the Brunswick River and just N of the US 74-76 causeway, ca. 2 mi W of Wilmington, growing with Typha angustifolia, C hyalinole pis, Pontederia cordata, Pentandra virginica, Sagittaria lancifolia, Rumex verticillatus, and Taxodium distichum, 34 14' 04.7'N, 77° 59°05.7" (NAD 83), Alan S. Weakley 7317 and Richard J. LeBlond, 10 Jun 2004 (HOLOTYPE: NCU; IsoTYPEs: BRIT, CLEMS, GH, MO, NY, TEX, UGA, US, USCH. = ea a a ] li] ee } +1 Ptilimnio capillaceo (Michx.) Raf lateralibus fructus valde evolutis, sed differt foliis divisionibus 2-3 (versus 3) per nodum, umbellulis 5-8(-13) per umbellam, floribus 5-8 per umbellulam, fructibus majoribus (2.7-4.2 mm longis versus 1.4-2.5 mm), geographica ac habitatione multo magis restrictis, et tempore praecoci florendi ac fructificandi. Annual, glabrous, caulescent herbs from a short system of fibrous roots, the stems erect, 4-13 dm tall, unbranched or in vigorous individuals branching above mid-stem. Lower stem leaves (often withered by anthesis) phyllodial (re- WEAKLEY AND NESOM, A NEW SPECIES OF PTILIMNIUM : e e t | i i ; f (@) = ey) =-o =n = 0 = pri ARL jo June Deo ac qeahh = Fic. 1. Hol yt cauline leaves 746 BRIT.ORG/SIDA 21(2) nin any O Pee So a SD 8 Fic? Infl £ Dei), y hlocis /f; eee Yu +h 1 ones +h otdad? 1 Tt andthe CA iets J ! 1£ ra j hall { 1+nD t duced to the hollow rachis and lacking lateral segments, thus closely resem- bling the quill-leaves of P fluviatile, P. nodosum, and P. viviparum) to simply pinnate, the segments (when present) flattened, to 2 mm wide; mid- and up- per-stem leaves pinnately decompound, 3-21 cm long, the segments filiform to distinctly flattened, with 2-3 divisions at primary nodes on the rachis; petioles 3-25mm long, broadly winged, ca 5-ribbed. Inflorescence umbellate, distinctly and irregularly flat-topped (the rays variable in length), once-compound, the umbels terminal and axillary on the upper half of the stem, peduncles 5-14. cm WEAKLEY AND NESOM, A NEW SPECIES OF PTILIMNIUM 747 Hur tcm Fic. 2. Frui t Let £ lly + ) £ Deli, y hl. ii, from an isotype. 748 BRIT.ORG/SIDA 21(2) long; primary rays 5-13, 5-35 mm long, pedicels (2-)4-10, (3-)5-10 mm long; involucre of linear bracts, entire or 3-parted; involucel of linear bracts. Calyx teeth deltoid, acute to subacuminate, persistent; petals white, obovate, 0.6-0.8 mm long; stylopodium conic, the styles mostly slightly shorter than the sty- lopodium, ascending to erect-ascending or widely spreading; anthers light pur- plish. Carpophore bifid at the apex. Fruit elliptic-ovoid, slightly compressed laterally, 2.7-3.5 mm long, L.8-2.2 mm wide, the mericarps loosely connate and usually separated at maturity; dorsal ribs 3, narrow and sharp-edged, the lat- eral ribs similar but with a broad, rounded, corky-thickened extension, the ex- tensions of the adjacent mericarps connate and forming a broad, conspicuous band around the fruit; oil tubes dark rusty-brown. In tidal freshwater marshes: flowering May to early June, fruiting late May to July. Collections examined. UNITED STATES. GEORGIA. Chatham: s Co.: als National Wildlife Ref- uge, along Hwy 17, fresh-water marsh, 14 Jul 1966, B 90 (NCU, distributed as “Ptilimnium lel mum Ahles”). NORTH CAROLINA. Brunswick Co.: Gia Jun 1898, Ashe s.n. (NCU). Co.: | mi N of US 17 on Wrightsboro Road, marsh, 2 Jun 1949, Raf 4353 (NCU); es ae on the Cape Fear River on US 17, 29 Jun 1963, Ahles 58397 (NCU) P r Co.:swamp on Northeast Cape Fear River near Rocky Point—Hampstead Road, | Jun 1950 ae 5142 (NCU). SOUTH CAROLINA. Beaufort Co.: 0.5 mi W of Co. Road 111 on Co. Road 33, cat-tail swamp near Trichinham Plantation, 27 Jun 1956, Bell 3767 (NCU). Berkeley Co.: Cooper River at the mouth of Durham Creek, low salinity, 7 Jun 1990, McAninch 23 (NCU). Colleton Co.: 1.4 mi NE of Ashepoo on US Hwy 17, swamp, 26 Jun 1956, Bell 3703 (NCU). Georgetown Co.: tidal marsh of Baruch Plantation, 20 Jun 1967, pal moses Baruch Plantation, fresh-water marsh affected by tidal action, 17 Jun 1969, Batson s.n. (USCH); Baruch Plantation, slough in freshwater marsh, 27 Jun 1969, Batson s.n. — (USCH); occasion: al i in sunny spots in wet swampy woods, side of South Island Road (S-18) abou mi W of Intracoastal Waterway, 16 May 1990, Nelson 9024 (USCH); Sampit River near US 17A, a mi W of Georgetown, fresh-water marsh, 13 Jun 1957, oa Soe CU). Relationships within the genus Ptilimnium is a relatively small genus centered in the southeastern and south- central United States. The five to seven species are separable into subgroups on the basis of differences in morphology and chromosome number, although ap- parent dysploid differences in chromosome number suggest that the situation may be more complex than the current assessment allows. Plants with leaves reduced to a winged petiole and extended rachis (with lateral segments sup- pressed, the structures sometimes referred to as “phyllodia”) are diploids based on x = 6 (2n = 12; Easterly 1957; Bell & Constance 1960). These plants have been regarded as comprising from one to three species, P nodosum (Rose) Mathias, P viviparum (Rose) Mathias, and P fluviatile (Rose) Mathias) (Kral 1981; Kress et al. 1994; Bartgis 1997); the appropriate taxonomic disposition remains ine and is under additional study. Ptilimnium costatum (EIL) Raf. hasd y petiolate leaves with short, crowded, verticillate segments, long styles, and a tetraploid chromosome complement based on x =8 (2n = 32; Easterly 1957). One population of P.costatum from Illinois has been counted as 2n = 22 (Bell & Constance 1960), this perhaps a tetraploid (x = 6, 2n = 24) having lost one pair of chromosomes. WEAKLEY AND NESOM, A NEW SPECIES OF PTILIMNIUM 749 Ptilimnium ahlesii, P capillaceum, and P.nuttallii are similar in their rela- tively short styles (shorter than or equalling the stylopodia) and primarily dip- loid chromosome number based on x = 7 (2n = 14; Easterly 1957, Bell and Con- stance 1960). Tetraploid plants (2n = 28) of P. capillaceum were reported by Easterly (1957) from central Georgia (Unadilla County). These were found ap- parently intermixed with diploid plants of the same species and with plants of P. nodosum “nearby in the same ditch.” Easterly did not comment on any mor- phological difference between the conspecific diploids and tetraploids. One population of P capillaceum from Florida has been counted as 2n = 16 (Con- stance et al. 1976). It is notable that the basal and lower stem leaves of P. ahlesii, produced in the spring and typically withering prior to flowering and fruiting, have few to no leaflets, and therefore closely resemble the reduced “quill leaves” of the “Harperella” group of Ptilimnium (P. fluviatile, P. nodosum, and P. viviparum). This supports the broader circumscription of Ptilimnium as in- cluding Harperella Rose that has been generally followed in recent decades. Ptilimnium texense Coulter & Rose was originally described as a species (Coulter & Rose 1909) but later (Mathias & Constance 1944-45; Easterly 1957) interpreted to be of hybrid origin and of intermediate morphology between P. capillaceum and P nuttallii. This interpretation has been accepted and repeated by subsequent workers (e.g., Correll & Johnston 1970; Correll & Correll 1972), although Mathias & Constance (1961) placed it in synonymy, without comment, under P. costatum. Study of Ptilimnium collections at LL,TEX shows only a single entity in Texas (interpreted here as P. costatum) other than P.capillaceum and P. nuttallii. In the freshwater tidal marshes west of Wilmington, North Carolina, three species of Ptilimnium grow in close proximity to one another: Ptilimnium ahlesii, P capillaceum, and P. costatum. They are morphologically distinctive, no intermediates have been seen. Observations over the past decade further suggest that the flowering times of the three species are largely or completely non-overlapping, with P ahlesii flowering first (May to early June), followed by P. capillaceum (mid-June to early August), and then P costatum (late July to August). The phenologic separation provides additional evidence corroborat- ing the status of P ahlesii as a distinct taxon and not merely a large-fruited form of P.capillaceum Of its closest relatives, Ptilimnium ahlesii is most similar to P capillaceum, particularly in its divided involucral bracts, minute calyx teeth, dorsal fruit ribs narrower than the intervals, and lateral ribs with expanded, corky mar- gins encircling the fruit like a band (P. costatum also shares the distinctive fruit morphology). The two species are separated by the following contrasts (and see Figs. 1 and 2). 1. Mature fruit elliptic-ovate, 2.7-4.2 mm long, 1.8-2.2 mm wide; umbels flat-topped, irregular, the rays variable in length; umbellets 5—8(-13) per umbel, the rays 5-35 750 BRIT.ORG/SIDA 21(2) Fic. 4 AA fd n 4° “1 ° Lin £€ +h + h, North Carolina, South Carolina, and Georgia) of Ptilimnium ahlesii. mm long; flowers 5-8 per umbellet; petals 0.6-0.8 mm long; leaf segments of mid- stem leaves 15-30(—40), capillary to linear, 0.5-2.0 mm wide; flowering May-early June, fruiting late May-July; tidal freshwater marshes, rare in North Carolina, South Carolina, and Georgia Ptilimnium ahlesii . Mature fruit ovate to orbicular, 1.4-2.5 mm long, 0.8-1.2 mm wide; umbels gener- ally rounded to hemispheric, the rays generally nearly equal in length; umbellets (5-)9-21 per umbel, the rays 5~21 mm long; flowers usually 10 or more per umbellet; petals 0.3-0.6 mm long; leaf segments of mid-stern leaves 50 or more, capillary, usually less than 0.5 mm wide (except in submersed leaves); flowering June-Au- gust, fruiting July-September; ditches, marshes, other wet places;common and wide- spread in eastern North America, ranging from Massachussetts and New York to Missouri and Kansas, south to Florida and Texas Ptilimnium capillaceum Descriptions by various authors (e.g., Fernald 1950; Mathias and Constance 1944- 45, Easterly 1957) of P.capillaceum with fruit longer than 3mm almost certainly included observations of P. ahlesii. Annotations by both Constance and Easterly of NCU collections of P. ahlesii as P capillaceum appear to confirm this. Associ- ated with the smaller number of umbellets and flowers borne by P ahlesii, as noted in the key, plants produce about 45-70 flowers/fruits per inflorescence, compared to those of P capillaceum, which usually produce over 100. Depauper- ate plants of P capillaceum, however, can have reduced inflorescences. WEAKLEY AND NESOM, A NEW SPECIES OF PTILIMNIUM 751 It appears that this species is narrowly distributed ecologically and geo- graphically, to tidal marshes associated with the tidal reaches of rivers in south- eastern North Carolina, South Carolina, and eastern Georgia (Fig. 4). ACKNOWLEDGMENTS We thank the staffs of LL-TEX, NCU, and US for help with herbarium studies at their institutions and staff of USCH for a loan of specimens. Richard LeBlond (North Carolina Natural Heritage Program) assisted with field studies. Ronald L. Hartman and an anonymous reviewer provided helpful suggestions. REFERENCES Bartals, R.L. 1997. The distribution of the endangered plant Ptilimnium nodosum (Rose) Mathias (Apiaceae) in the Potomac River drainage. Castanea 62:55-59. Bet, C.R. 1968. Apiaceae. In: A.E. Radford, H.E. Ahles, and C.R. Bell. 1968. Manual of the vas- cular flora of the Carolinas. Univ. North Carolina Press, Chapel Hill. Pp. 761-789. Bett, C.R. and L. Constance. 1960. Chromosome numbers in Umbelliferae. Il. Amer. J. Bot. 47:24-32. Constance, L.,T.-l. CHUANG, and C.R. Beit. 1976.Chromosome numbers in Umbelliferae.V. Amer. J, Bot. 63:608-625. Corrett, D.S. and M.C. Jounston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner, Texas. Corrett,D.S.and H.B. Corrett. 1972.Aquatic and wetland plants of the southwestern United States. Environmental Protection Agency, Washington, DC. Coutter, JM. and J.N. Rose. 1909. Supplement to the monograph of the North American Umbelliferae. Contr. U.S. Natl. Herb. 12:441-451. Easterly, N.W.1957.A morphological study of Ptilimnium. Brittonia 9:136—-145. FerNato, M.L. 1950. Gray’s manual of botany (ed. 8). American Book Co., New York. Franktin, M.A. (ed.).2004. Natural Heritage Program list of rare plant species of North Caro- lina: 2004. N.C. Natural Heritage Program. Georcia Natural Heritace Procram. 2004. Special concern plant species in Georgia. [http:// georgiawildlife.dnr.state.ga.us/content/specialconcernplants.asp] Kartesz, J.T. 1999.A synonymized checklist and atlas with biological attributes for the vas- cular flora of the United States, Canada, and Greenland. First edition.|n:Kartesz, J.T.,and C.A.Meacham. 1999. Synthesis of the North American flora, Version 1.0.North Carolina Botanical Garden, Chapel Hill, North Carolina. Krat, R.1981.Notes on some quill-leaved umbellifers. Sida 9:1 24-134. Kress, W.J.,G.D. Mappox, and C.S. Roeset. 1994. Genetic variation and protection priorities in Ptilimnium nodosum (Apiaceae), an endangered plant of the eastern United States. Conserv. Biol. 8:271-276. Matuias, M.E. and L. Constance. 1944-45. Umbelliferae. N. Amer. Fl. 28B:43-295, Matuias, M.E. and L. Constance. 1961. Umbelliferae. In: C.L. Lundell, ed. Flora of Texas, Vol. 3. Texas Research Foundation, Renner, Texas. Pp. 263-329. BRIT.ORG/SIDA 21(2) Raproro, A.E., H.E. AHies, and C.R. Bett. 1964. Guide to the vascular flora of the Carolinas. The Book Exchange, Univ. of North Carolina, Chapel Hill. Raproro, A.E.,H.E. Axes, and C.R. Bett. 1965. Atlas of the vascular flora of the Carolinas. North Carolina Agric. Exp. Stat. Tech. Bull. No. 165. Sorrie, B.A. 1997. Notes on Lycopus cokeri (Lamiaceae). Castanea 62:119-126. NEW COMBINATIONS IN NORTH AMERICAN CARYOPHYLLACEAE Ronald L. Hartman Richard K. Rabeler Rocky Mountain Herbarium University of a Herbarium Department of Botany 3600 Varsity D niversity of Wyoming Ann Arbor, Michigan 481 A on U.S.A. Laramie Wyoming 82071-3165, U.S.A. rabeler@umich.edu rhartman@uwyo.edu ABSTRACT The ee new BCom enanOns in the Caryophyllaceae are proposed: Arenaria lanuginosa var. saxosa, Eremogc risiae, Minuartia nuttallii vars. fragilis, gracilis, and gregaria, and Paronychia pete var. minima. Key worps: Caryophyllaceae, Arenaria, Minuartia, Eremogone, Paronychia. RESUMEN Se proponen las siguientes combinaciones nuevas en Caryophyllaceae: Arenaria lanuginosa var. saxosa, Eremogone ferrisiae, Minuartia nuttallii vars. fragilis, gracilis, y gregaria, y Paronychia chartacea var. minima. During the course of our work on the treatments of Arenaria, Minuartia, and Paronychia for Flora of North America (Hartman, Rabeler, and Utech, in prep:; Rabeler, Hartman, and Utech, in prep.; and Hartman, Thieret, and Rabeler, in prep., respectively), we found the following combinations necessary to consis- tently treat infraspecific variation. Arenaria nee (Michx.) Rohrb. var. saxosa (A. Gray) Zarucchi, R.L. Hart- man & Rabeler, comb. et stat. nov. BASIONYM: Arenaria saxosa A. Gray, Smithsonian ae Knowl. 5 (Art. 6):280 [Plantae Wrightianae, pt. 2:18]. 1853, priority of the autonym es- tablished by A. saxosa var. cinerascens B.L. Rob., Proc. Amer. Acad. Arts 29:293. 1894 The varietal combination often used for this plant, Arenaria lanuginosa var. cinerascens (B.L. Rob.) Shinners, is not correct because of Art. 11.6 of the Inter- national Code (Greuter et al. 2000) which states that “an autonym is treated as having priority over the name or names of the same date and rank that estab- lished it.” Minuartia nuttallii (Pax) Briq. var. Hasilts (Maguire & A. i Meus Rabeler Ss rtman, comb. nov. BAsIO subsp rE H. Holmgren, Madrons 8:260. 1946. Arenara nuttallii var. pane (Maguire & or Palmprend CL. Hitchc., Vasc. Pl. Pacific North W. 2:258. 1964. Minuartia nuttallii subsp. fragilis (Maguire & A.H. Holmgren) McNeill, Rhodora 82: 499. 1980. meatal nuttallii (Pax) Briquet var. gracilis (B.L. Rob.) Rabeler & RL. Hart- ,comb. nov. BASIONYM: Arenaria nuttallii Pa Rob., Proc. Amer. Acad SIDA 21(2): 753-754, 2004 754 BRIT.ORG/SIDA 21(2) Arts 29:304. 1894. Arenari tallii subsp. gracilis (B.L. Rob.) Maguire, Madrono 8:261. 1946. Minuartia nuttallii subsp. gracilis (B.L. Rob.) McNeill, Rhodora 82:499. 1980. Minuartia nuttallii (Pax) Briquet var. gregaria (A. Heller) Rabeler & RL. Hart- man, comb. et stat. nov. BASIONYM: Arenaria gregaria A. Heller, Bull. S. Calif. Acad. Sci. 2:67. 1903. Alsinopsis gregaria (A. Heller) A. Heller, Mublenbergia 8:96. 1912. Arenaria nuttallii Pax subsp. gregaria (A. Heller) Maguire, FI. Calif. 492.1914. Arenaria nuttallii var. gregaria (A. Heller) Jepson, Madrono 8:261. 1946. Minuartia nuttallii subsp. gregaria (A. Heller) McNeill, Rhodora 82:499, L980. While the four infraspecific entities traditionally recognized within Arenaria (Minuartia) nuttallii Pax are often distinct, occasional specimens, especially of varieties fragilisand gracilis, do intergrade where their ranges overl ap, espe- cially in Oregon and Nevada. All have been treated as varieties in at least one earlier treatment of this group and we feel that this is the appropriate rank. Paronychia chartacea Fernald var. minima (L.C. Anderson) R.L. Hartman, comb. et Stat. NOV. BASIONYM: Paronychia chartacea Fernald subsp. minima L.C. Anderson, Sida 14:436. L991. Postscript.—After additional study of Eremogone macradenia (S. Watson) Ikonn., we now believe that E. macradenia should be split into two taxa with E. macradenia var. ferrisiae (Abrams) R.L. Hartman & Rabeler being elevated to species rank (Hartman & Rabeler 2004). Eremogone ferrisiae (Abrams) R.L. Hartman & Rabeler, comb. nov. Basionywt Arenaria macradenia S. Watson subsp. ferrisiae Abrams, Ill. Fl. Pacif. States 2:151. 1944. Eremogone mac i nia (S. Watson) Ikonn. var. ferrisiae (Abrams) R.L. Hartman & Rabeler. Sida 21:240. 2 ACKNOWLEDGMENTS We wish toacknowledge James Zarucchi (MO) for discovering that the Arenaria combination was required and to Kanchi Gandhi (GH) for confirming this fact. Curators of F MICH, and RM are thanked for providing specimens of the Minuartia nuttallii complex. REFERENCES Greurer, W., J. MCNEILL, R. Barrie, H.-M. Buroer, V. Demouun, T.S. Fitcuerias, D.H. Nicotson, PC. SiLva, JE. Skoe, P. TREHANe, NJ. Turtanp, and D.L. HawkswortH (eds.). 2000. International code of botanical nomenclature (Saint Louis Code) adopted by the Sixteenth International Botanical Congress, St. Louis, Missouri, July-August 1999, Regnum Veg. 138: xviii + 1-474, Hartman, R.L. and R.K. Rasever. 2004. New combinations in North America Eremogone (Caryophyllaceae). Sida 21:237-241. SOLIDAGO SECT. PTARMICOIDEI, A NEW COMBINATION TO REPLACE A “RANKLESS” NAME USED BY TORREY AND A. GRAY (ASTERACEAE: ASTEREAE) John C.Semple Kanchi N. Gandhi Department of Biology Gray Herbarium University of Waterloo Harvard pees Waterloo, Ontario, CANADA N2L 3G! Cambridge, M f ts 02138-2020, U.S.A. jcsemple@sciborg.uwaterloo.ca gandhi@oeb.harvard.edu ABSTRACT Within the treatment of Solidago, Torrey and Gray (1842), there are no textual indicators of the ranks intended for subdivision of the genus, and all of the Torrey and Gray names for subdivisions of Sol- idago are rankless. The new combination Solidago sect. Ptarmicoidei is made. RESUMEN 1QA9 En el tratamiento de Solidago, Torrey y Gray (1842), no hay indicadores textuales de los rangos que se Has acen para ie subdivision del género, y todos los nombres de Torrey y Gray para las subdivisiones de c rango. Se hace la nueva combinacion Solidago sect. Ptarmicoidei. Discussions concerned with the ranks of names of subdivisions of genera pub- lished by Torrey and Gray (1842) led to the conclusion that by and large such names are rankless unless a rank was explicitly indicated within the text John Strother, John Pruski, John McNeill, pers. comm. with K.N. Gandhi). Torrey and Gray used the symbol § to represent either a subgenus or section. Unless a rank is explicitly indicated, names marked by the symbol § are rankless as are lower level names marked by indicators such as *, **, *** etc. Within the treatment of Solidago (Torrey and Gray 1842, pp. 195-231), there are no textual indicators of the ranks intended, and thus all of the Torrey and Gray infrageneric names in Solidago are rankless. In Gray (1884), sectional and subgeneric ranks are clear. In his preface (vol. 2(1). 1878), Gray used the symbol § and large-type capital letters for names of sections and subgenera. He used nouns for names of subgenera and adjectives for names of sections. Additionally, Gray did mention subsection and further lower divisions (he did not mention series or subseries); he used the symbols * ** etc.and small-type capital letters for names of subsections. By implication, a * ** etc. denoted the rank of subsection because those symbols are placed at the beginning of the diagnostic statement and sectional name immediately fol- lows the statement. One may dispute this; we go by implication. For example, Solidago |rankless] Corymbosae Torr. & A. Gray (in 1842) can be taken as Sol- SIDA 21(2): 755-757. 2004 756 BRIT.ORG/SIDA 21(2) idago subsect. Corymbosae (Torr. & A. Gray) A. Gray (in 1884) because Gray (1884) listed it as “***** .. CORYMBOSAE” [small capitals]. Nesom (1993) summarized the infrageneric taxonomy of Solidago and lectotypified some names. He treated the corymbiform, oligoneurate species in a separate genus Oligoneuron Small. He accepted Torrey and Gray names with ranks generally accepted at that time by other authors following the rationale for doing so presented by Holmgren (1979) and Jones (1980). Nesom did not vali- date Solidago sect. Corymbosae Torr. & A. Gray. One of the criteria for valid pub- lication of a name is that the author (in this case Nesom) must accept and use the name. Nesom cited “Solidago sect. Corymbosae T. & G.” as a synonym of Oligoneuron sect. Ptarmicoidei (House) Nesom; therefore, he did not validate the former name. Semple, Ringius and Zhang (1999) followed Nesom’s infrageneric nomenclature, except that they treated Oligoneuron as Solidago sect. Corymbosae Torr. & A. Gray with synonyms but did not list full biblio- graphic data for each name. Had they done so, they might have inadvertently validated the name at sectional rank. Solidago L. sect. Ptarmicoidei (House) Semple & Gandhi, comb. noy, Based on As- ter sect. Ptarmicoidei House, N.Y. State Mus. ae 7 710. 1924, nom. et stat. nov. based on Unamia Greene (1903). Oligoneuron sect. Pt i CH )G.L. Nesom, Phytologia 75:27, 1993. TyPE: Inula alba Nutt. = Aster ptarmicoides Torr. & A. Gray = Solidago ptarmicoides (Torr. & A. Gray) Boivin. [Solidago asteroides Semple (in Semple, ew oe Ener U. Waterloo Biol. Ser. 39:72. 199 fl l name for t] he much older Aster ptarmicoides Tie & A. Gray is accepted as legitimate (see Voss 1996)| Solidago [rankless] Corymbosae Torr. & A. Gray, Fl. N. Amer. 2(2):208. 1842. Solidago subsect. Corymbosae (Torr. & A. Gray) A. Gray in A. Gray et al, Syn. Fl.N. Amer, 1(2):159. 1884. TYPE: S. corymbosa Elliott. Names of subdivisions of genera are automatically typified by the species name from which the name of the subdivision is derived. For group Corymbosae, Torrey and A. Gray included S$. corymbosa. Nesom (1993:26) erred in designating Solidago rigida L. as the lectotype aa Small, FL SE. U.S. 1188. 1903. Solidago subg. Oligoneuron (Small) House, N.Y. State Bull. No, 254. 693. 1925. LEcToTYPE [Nesom 1993:26}: Oligoneuron rigidum (L.) Small = ie rigida L. Unamia Greene, Leafl. Bot. Observ. 1:6. 1903. Type: Inula alba Nutt. = Unamia ptarmicoides (Torr. et A. Gray) Greene, nom. superfl. for Unamia. alba (Nutt.) Rydb. Oligoneuron ser. Pt crmcoidei House L. Nesom, ae he 1993. Aster sect. Ptarmicoidei ouse, N.Y. State Mus. Bull. 254. 710. 1924. TyPE: In tt. = Aster ptarmicoides Torr. & A. Gray = Ee eeareein ms a & A. Gray) Boiv ae nee album(Nutt.) G.L. Nesom Oligoneuron ser. Xanthactis G.L. Nesom Phytologia 75:28. 1993. Type: Oligoneuron riddellii (rank) Rydb. = Solidago riddellii Frank. gq. O = Other nomenclatural decisions made by Nesom (1993) related to the rankless Torrey and Gray names include: Solidago sect. Solidago Solidago subsect. Maritimae (Torr. & A. Gray) G.L. Nesom, Phytologia 75:12. 1993. SEMPLE AND GANDHI, SOLIDAGO SECT. PTARMICOIDES, A NEW COMBINATION 757 Based on Solidago {rankless] Maritimae Torr. & A. Gray, Fl.N. Amer. 2(2):211 1842. LECTOTYPE [Nesom 1993:12]: Solidago sempervirens L.[Nesom made this subsectional name inadvertently] ec a eae Raf., Fl. Tellur. 2:42. 1836 [1837]. LEcTOTYPE [Nesom 1993:12]|: Solidago Br i ee Uliginosae Mackenzie in Small, Man. SE. Fl. 1345, 1347. 1933. TyPE: Solidago uliginosa Nutt. cited under Solidago uniligulata (DC.) Porte Solidago subsect. Triplinerviae (Torr. & A. Gray) G.L. Nesom, Phytologia 75:8. 1993. Solidago [rankless] Triplinerviae Torr. & A. Gray, Fl. N. Amer. 2(2):222. 1842; A. Gray, Syn. FI. N. Amer. 1(2):155. 1884. Lectotype [Nesom 19938]: Solidago canadensis L. [Nesom made this subsectional name inadvertently] pee subg. Brachyactis Raf., FI. Tellur. 2:42. 1836. TYPE: Solidago juncea Aiton. Nesom (1993: 8) d in designating a LT. Raf. designated S. juncea as the type. Sly suby, Triactis Raf., Fl. Tellur. 2:42. 1836 [1837]. TyPE: Solidago retrorsa Michx. Nesom (1993:11) erred in designating a LT; Raf. designated S. retrorsa as the type. pre: ee Serotinae Rydb., Fl. Rocky Mts. 868. 1917. TYPE: Solidago serotina Aiton (1789), Solidago gigantea Ait. Solidago [rankless] Serotinae Mackenzie in J. K. Small, Man. SE. FI. 1345, 1350. 1933, non Rydb. (1917). TyPE: Solidago serotina Retz, (1781), non ae (1789) =? S. tortifolia Ell. 1824 ACKNOWLEDGMENTS This research was supported by a Natural Sciences and Engineering Council of Canada Discovery Grant to JCS. Guy Nesom and John Strother are thanked for their useful review comments. REFERENCES Gray, A.1878.Synoptical flora of North America. vol. 2, part.1.|vison, Blakeman, Taylor & Co, New York. Gray, A. 1884. Synoptical flora of North America. vol. 1, part. 2. Ivison, Blakeman, Taylor & Co., New York. Hotmoren, N.H. 1979. Subgeneric and sectional names for Intermountain Penstemon (Scrophulariaceae). Brittonia 31:358-364. Jones, A.G. 1980. A classification of the New World species of Aster (Asteraceae). Brittonia 32:230-239. Nesom, G.L. 1993. Taxonomic infrastructure of Solidago and Oligoneuron (Asteraceae: Astereae) and observations on the phylogenetic position. Phytologia 75:1-44. Sempte, J.C.,G.S. Rincius, and J.J. ZHANG. 1999 (October). The goldenrods of Ontario: Solidago L.and Euthamia Nutt. 3rd Ed. Univ.Waterloo Biol. Ser. 36:1—90. Torrey, J.and A.Gray. 1842. Solidago L.Flora of North America. 2(2):195-231.Wiley & Putnam, New York. Voss, E.G.1996. Michigan flora. Part Ill. Dicots (Pyrolaceae-Compositae). Cranbrook Inst. Sci Bull. 61 758 BRIT.ORG/SIDA 21(2) TWO BEAUTIFUL BOOKS FROM THE ANTIQUE COLLECTOR’S CLUB Margaret Mee’s Passion and Painting MARGARET Mer. 2004. Margaret Mee’s Amazon: Diaries of an Artist Explorer. (ISBN 1-85149-454-5, hbk.) Antique Collectors’ Club Ltd., Eastworks, 116 Pleas- ant Street, Suite 6OB, Easthamton, MA 01027, U.S.A. (Orders: 413-529-0861, Fax: 413-529-0862, wwwantiquece.com) $59.50, 319 pp., color figures & plates, O12 Se. orthcoming in volume 21, no. 3. a Review Legendary Plant Hunter George Forrest BRENDA McLean. 2004. George Forrest: Plant Hunter. (1-85149-461-8, hbk) An- tique Collectors’ Club Ltd., Eastworks, 116 Pleasant Sreet, Suite 60B. Easthamton, MA 01027, US.A. (Orders: 413-529-0861, Fax: 4 3-529-0862, www.antiquecc.com). $59.50, 250 pp., 80 color and 80 b/w illustrations, 8 1/4" x] Publisher Comments: “George Forrest was a legendary plant collector in the heyday of the British Empire. His career spanned the decades before and after the First World War. Risking his safety and — health, he discovered hundreds of new species, introduced many plants to our gardens, and became one of the most ling plant collectors in the Sino-Himalaya. As many plants were named after him, he is well known: in gardening circles yet this is the first biography of Forrest, and the first book on him for fift “This book odedi to celebrate the centenary of Forrest's first setting out for the mountains of Yunnan, south-west China, brings Forrest to life, drawing on his own letters and those of his con- poraries. It tells of Forrest’s adventures and his escape from death at the hands of war ring Lamas. tem] It shows the influence of his patrons, describes the excitement of his plant and animal discoveries, and reveals his rivalry with other cay hunters, Reginald Farrer ee Frank Kingdon Ward.” | like to read about explorers, adventurers, and dis | rge Forrest was such a per- son. He was a naturalist. Appendix 9 in the book lists some 175 genera rol plants with a forrestii spe- cies named alter George Forrest. There is a dragonfly (lemnogomphus forrestii) named after George Forrest. Then there are birds named after George Forrest. Not just one but six different genera with species ues as resti. * wait, ag are some mammals that bear the species name forresti, not to mention the 2 from Forrest's mammal collection at the Natural History Museum, London fein 10). George Forrest realized he had the supper t ou mi aie people and I'm r thank you by / sure he thanked them in many different ways, but Forrest did the b naming many of his newly discovered pene a enjo the | listory of Forrest: Plant Hunter~is for you.— YU If Barne eats Bot ailee keceant h Institute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4060, TSA SIDA 21(2): 758. 2004 MISCELLANEOUS NOMENCLATURAL CHANGES IN ASTEREAE (ASTERACEAE) John C. Semple peeae: of Biology University of Waterloo Waterloo, ate , CANADA NAL 3G1 jcsemple@sciborg.uwaterloo.ca ae ABSTRACT ai HOHOWNE new name and combinations are proposed: Heterotheca subaxillaris subsp. latifolia. 1 Multiradiatae, and § concolor var. devestitun RESUMEN Se proponen un nombre nuevo y H I I illaris subsp. latifolia, Sol- I idago subsect. Multiradiatae, y Symphyotrichum concolor var. devestitum. The proposed new name and combinations were determined to be needed dur- ing work to prepare the treatments of Heterotheca (Semple 2005), Solidago (Semple & Cook 2005) and Symphyotrichum (Brouillet et al. 2005) for the Flora North America project. Beton subaxillaris (1am.) Britt. @ Rusby subsp. latifolia (Buckley) Semple, : .et Stat. nov. BASIONY™M: Heterotheca latifolia Buckley, Proc. Acad. Sci. Phila. 13:459 ne Heterotheca subaxillaris (Lam.) Britt. & Rusby var. latifolia (Buckley) Gandhi & Tho- mas, Sida Bot. Misc. 4:110. 1989. Type: TEXAS. Llano Co. Buckley s.n. (HOLOTYPE: PH, not seen). The weedy Heterotheca subaxillaris complex has been treated asa single species without varieties (Nesom 1990) and as four separate species: H.chrysopsides DC,, H. latifolia, H. ps phila Wagenknecht, and H. subaxillaris (Semple 1996 and earlier authors). Harms (1964) presented data supporting Wagenknecht’s (1960) division of the complex into four species and provided a general range map of the four taxa. Nesom (1990) opted to merge all four species into a single undi- vided species because he did not find taxonomically significant differences between the four putative species. Semple (1996) continued to followed Wagenknecht and Harms. However, after considerable study of members of the complex, | have adopted Nesom’s position that only one species should be rec- ognized (Semple 2005). Nonetheless, while much of the supposed differences between regional taxon do not appear to be supported, some phytogeographic patterns in variation occur within the complex and two infraspecific taxa can be recognized based on differences in phyllary traits. Typical Heterotheca subaxillaris is for the most part confined to the outer coastal plain from New York to northern Mexico, where they can occur further inland. These plants SIDA 21(2): 759-765. 2004 760 BRIT.ORG/SIDA 21(2) have a well developed tuft of larger hairs near the phyllary tips (Figs. [A-H) and are treated here as subsp. subaxillaris. This combination of distinctive morphology plus the geographic isolation of much of the range of the morphotype warrant subspecies level recognition following Semple (1974). In contrast, plants of subsp. latifolia have phyllaries without such large hairs con- centrated near the tip (Figs. LI-L). Mid series phyllaries of subsp. subaxillaris on average are slightly wider (0.62 mm wide) compared to the average width of those of subsp. latifolia (0.57 mm wide) with the narrowest occurring in Ari- zona and New Mexico plants (‘psammophila’). However, the ranges in widths overlap so much that the trait cannot be used as a diagnostic difference. Within each subspecies there appear to be patterns to the phyllary varia- tion but these are not sufficiently strong that additional infraspecific taxa can be justifiably recognized at this time. There isa general geographic trend to the variation in tufted hairs in subsp. subaxillaris. Plants with the most hairs in the apical tuft occur from Mexico to Florida (Figs. [A-D), while the number of hairs generally decreases from Florida to New Jersey and New York (Figs. E-H). Plants with a few hairs near the apex occur scattered across the range of the species and may represent introductions of subsp. subaxillaris or the occasional more hairy than normal individual of subsp. latifolia. Plants occurring in an arch from Arizona and adjacent New Mexico through Mexico into trans-Pecos Texas tend to be more ly glandular than plants from elsewhere in the range (Fig. LL) in the United States; these have been treated as H. psammophila Wagnknecht. Plants from northeastern Mexico tend to have more glands and more hairs on the mid series phyllaries and are more likely to be weakly peren- nial; these have been treated as H. chrysopidis. Plants treated previously as H. latifolia tend to have fewer hairs and fewer glands on the phyllaries than other members of subspecies latifolia. These occur across the southern prairies and through the south eastern U.S. on the Piedmont as shown in Harms (1965), al- though the weedy nature of the species appears to be facilitating dispersal into the range of subsp. subaxillaris on occasion. Additional study may find new evidence supporting recognition at varietal or subspecies level for the psammophila and chrysopsides morphs, but my recent efforts have not discov- ered such evidence. Solidago subsect. Multiradiatae Semple, subsect. nov. Tyre: Solidago multiradiata L. Solidago ser. Multiradiatae Juz., Fl. URSS 25:47. 1959. nom. invalid, no Latin diagnosis. Sol- idago subsect. Multiradiatae Juz.) Semple, Sida 20:1605. 2003. non valid name. TyPE: Sol- - 4 ie q; - ¢ Cnlid P| -| : ] a3 4 Pee : t ] : WL, [eo] Solidagini sect. I j Members of subsect. Multiradiatae are distinguished from other species in sect. Solidago by having a somewhat rounded-cor y mbiform apit ulescence. Other North America members of the section have variously paniculiform SEMPLE, NOMENCLATURAL CHANGES IN ASTEREAE 761 Fic. 1. Phyllary tip variation in Heterotheca subaxillaris. A-H. Subsp. subaxillaris. A-B. Mexico. A. Veracruz, Turner 15372 (TEX). B. Tamaulipas, LaSueur 496 (TEX). C-H. United States. C. Texas, Cameron Co., King 2 (MINN). D. Louisiana, Cameron Par., Dutton & Pritchett 2549 (WAT). E. Florida, Orange Co., Semple et al. 2557 (WAT). F. North Carolina, Brunswick Co., res (MINN). G. New Jersey, Gloucester Co., Long 60507 (MINN). H. New York, Queens Co., Semple 2031 (WAT).1- L. Subsp. /atifolia. 1. Mexico, Chihuahua, SLU) eae eS So see J-L. United States. J. Georgia, Elbert Co., Coile 1626 (WAT). K. Oklahoma, Caddo Co. rizona, Maricopa Co., L. 179116 (WAT; U “psammophila”). capitulescences[S. subsect. Humiles (Rydberg) Semple]. Like most other taxa in the genus, the phyllaries have a single vein. In robust plants the capitulescence becomes more paniculiform as branches develop from upper stem leaves. Two other species besides the nomenclatural type are included in the subsection: S. leiocarpa DC. (Fig. 2B) and S. spithamaea M.A. Curtis ex A. Gray (Fig. 2C). These are both narrowly distributed endemics in eastern North America (Semple & Cook 2005). Solidago leiocarpa (synonym: S. cutleri Fern.) includes tetraploids 762 BRIT.ORG/SIDA 21(2) [tiradiata, Yukon, Chmielewski et al ((3824 (WAT) B. Solidago leiocarpa, New Hampshire, Ringius 1584 (WAT). C. Solidago spithamaea, North Carolina, Semple & Suripto 9669 (WAT). Scale bars = 1 cm. Fic. 2 Solidago subsect Multiradiatae, capitulescences A Solid found at disjunct high elevations in eastern New York, the Green Moutains in Vermont, the White Mountains in New Hampshire, and on Mt. Katadhin anda few higher peaks in Maine (Beaudry 1963; Morton 1981; Magee & Ahles 1999). Solidago spithamaea includes hexaploids restricted to the highest granitic out- crops in the mountains of western North Carolina (Cronquist 1980; Semple et al. 1984; Semple & Cook 2004). Solidago multiradiata includes diploids and tetraploids and is widely distributed across northern North America from the Maritimes to Alaska across Canada and down the cordillera at higher eleva- tions in the western United States to California, Arizona, Nevada, and New Mexico (Semple et al. 1999). Other goldenrods that can have a rounded- corymbiform capitulescence occur in Solidago sect. Ptarmicoidei Semple & Gandhi, but these have striate phyllaries ( (Semple & Gandhi 2004). Solidago wrightii A. Gray sometime can also have a rounded somewhat corymbiform capitulesence, but the species has short-petiolate mid stem leaves and often glandular phyllaries (Semple & Cook 2005). Kanchi H. Gandhi (pers. comm.) noted that my combination proposed last year (Semple 2003) was based on an invalid basionym because the protologue lacked a Latin diagnosis. Symphyotrichum concolor (.) Nesom var. devestitum (S.F Blake) Semple, comb. OV. BASIONYM: Aster concolor L. var. devestitus S.F. Blake, Rhodora 32:145. 1930. Type: U.S.A. FLORIDA. Bay Co. Lynn Haven, in dry sandy open soil, 13 Oct 1921, Billington 80 (HOLOTYPE: US-1116195!) SEMPLE, NOMENCLATURAL CHANGES IN ASTEREAE 763 Fic.3.M i I £5 i lor. A Flowering heads. Florida. B—C Variety concolor. B. Mid stem leaf sur- face; Georgia, Semple & Chmielowski 6213 (WAT). C. Phyll Florida, Suwanee Co., Semple, Brouillet & Canne (WAT). D-G. Variety divestitum, Florida; E, Bay Co. | Semple & Godfrey 3118; D, F-G, Santa Rosa Co., Semple, pear Canne 3880 (WAT). D. Mid stem and leaf. E—F. Mid stem leaf surfaces. G. Phyllary. Scale bar = 1 mmin C, E-F and 1 cmin D Symphyotrichum concolor is characterized by its narrow, elongate capitulescence of usually many, violet-rayed heads (Fig. 3A). The var. concolor has more mod- erately to densely silky pubescent leaves and phyllaries (Figs. 3B-C), while var. devestitum has glabrous to very sparsely pubescent stems and glabrate to sparsely pubescent leaves and phyllaries (Figs. 3D-G). Plants similar to the type of var. devestitum occur in the western Florida Panhandle. The involucres of such Florida plants tend to be larger than those of diploid var. concolor Blake 1930). Very sparsely pubescent-leaved individuals also occur in southern Miss., 764 BRIT.ORG/SIDA 21(2) Ala., Ga., and S.C. These may prove to belong in var. devestitum pending more detailed study. Semple (1984) described and illustrated the distribution of dip- loids 2n = 8) and tetraploids (2n = 16) in S. concolor under the name Virgulus concolor, but did not discuss varieties and noted incorrectly that no obvious differences occurred between diploids and tetraploids. In Florida, the range of tetraploids is contained within the range of var. devestitum; tetraploids are cur- rently unknown outside Florida. Cytovouchers of tetraploids in WAT have gla- brous to sparsely pubescent leaves and belong in var. devestitum. The tetrap- loid condition may account for the larger involucres. ACKNOWLEDGMENTS This research was supported by a Natural Sciences and Engineering Council of Canada Discovery Grant to the author. Joanna Stinson assisted in collecting data on phyllary variation in the Heterotheca subaxillaris complex. Guy Nesom is thanked for his helpful review of the manuscript. REFERENCES Beauory, J.R. 1963. Studies on Solidago L. VI. Additional chromosome numbers of taxa of the genus Canad. J. Genet. Cytol. 5:150-174. Biake, S.F. 1930. A glabrous variety of Aster concolor.Rhodora 32:144-145. 1930. Cronauist, A. 1980. Vascular flora of the southeastern United States. |. Asteraceae. Univer- sity of North Carolina Press. Chapel Hill. Harms, V.L. 1965. Biosystematic studies in the Heterotheca subaxillaris complex (Compositae- Astereae). Trans. Kansas Acad. Sci.68:122-124 Morton, J.K.1981.Chromosome numbers in Compositae from Canada and the U.S.A. Bot. J. Linn. Soc. 82:357-368. Nesom, G.L. 1990. Taxonomy of Heterotheca section Heterotheca (Asteraceae: Astereae) in Mexico, with comments on the taxa in the United States. Phytologia 69:282-294. Macee, D.W. and H.E. AxLes. 1999. Flora of the northeast: a manual of the vascular flora of New England and adjacent New York. University of Massachusetts Press. Amherst. Sempte, J.C. 1974. The phytogeography and systematics of Xanthisma texanum DC. (Asteraceae): proper usage of infraspecific categories. Rhodora 76:1-19. St oo mc Cytogeographic studies on North American asters. |. range surveys of V.concolor, V.georgianus, V.grandiflorus, V. novae-angliae, V.oblongifolius, iaieis and V. walter’. Amer.J. Bot. 71:522-531. Semete, J.C. 1996. A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae: Astereae): the prairie and montane goldenasters of North America. Univ.Waterloo Biol. Ser. No. 37:1-164. Sempte, J.C. 2003. New names and combinations in Solidago (Asteraceae: Astereae). Sida 20:1605-1616. Sempe, J.C. 2005. Heterotheca Cass. In Flora North America Editorial Committee, eds. Flora of North America. Vol. 20. Asteraceae. Oxford University Press. SEMPLE, NOMENCLATURAL CHANGES IN ASTEREAE 765 Sempte, J.C.and R.E.Coox.2004. Chromosome number determinations in fam.Compositae, Tribe Astereae. VIl. Some eastern North American and Eurasian taxa.Rhodora 106:xxx— Xxx. IN press. Semete, J.C.and R.E.Cook. 2005. Solidago. pp. xxx-xxx. In Flora North America Editorial Com- mittee, eds. Flora of North America. Vol. 20. Asteraceae. Oxford University Press. Semete, J.C., G.S. Rincius. and JJ. ZHanc. 1999. The goldenrods of Ontario: Solidago L. and Euthamia Nutt. 3rd Ed. Univ. Waterloo Biol. Ser. 36:1-90. Semete, J.C., G.S. Rinaius, C. Leeper, and G. Morton. 1984. Chromosome numbers of golden- rods, Euthamia and Solidago (Compositae: Astereae). Il. Additional counts with com- ments on cytogeography. Brittonia 36:280-292. Addendum Brittonia 37:121-121.1985. WAGENKNECHT, B.L. 1960. Revision of Heterotheca sect. Heterotheca (Compositae). Rhodora 62:61-/76, 97-107. 766 BRIT.ORG/SIDA 21(2) BOOK NOTICES/BOOKS RECEIVED RicaRD SOLE and BRIAN GoopWin. 2000. Signs of Life: How Complexity Pervades Biology. (O-465-01928-5, pbk.). Basic Books, 387 Park Avenue South, New York, NY 10016, U.S.A. (Orders: 212-340-8100, Fax: 212-340-8115, www.basicbooks.com) $17.50, 322 pp., b/w photos, drawings, and diagrams, 6"x 91/4". Publisher Comments: “In every major field of biology, from molecular genetics and neurobiology, Oo}? through animal behavior and ecology, to evolution, extinction, and economics, there are well-known phenomena that today’s standard theories are | | plain: Why don't cells in identical en- vironments with identical one live identical lives? How do suche simple creatures as ants and termites ma such complex behavior as building nests? Why did all the animal kingdom’s basic body ria oo in a single geological era, and no new ones since? Yet, as Ricard Sole and Brian Goodwin show, various tools of complexity theory can offer us new ways to understand these phe- nomena. A tour of biology such as you've never seen be ore, ae ife isaboute: plaining the unex- aa by using new ideas to answer questions y mt help us with.” 2 e authors sae out that the real aim of this book is to ie how the new ideas are exerting their il luence in a variety of biological areas. Signs of Life is divided up into 10 chapters: 1) nonlinearity, Chaos, ae os nee; 2) ak Cc soi ae Disorder; 3) Genetic Networks, Cell Dif- ferentiation, and Development; 4) Physi nthe Edge of Chaos; 5) Brain Dynamics; 6) Ants, Brains, and Chaos; 7) The Baroque of Nature: 8) Li ife on the Edge of Catastrophe; 9) Evolution and Extine- tion: and 10) Fractal Cities and Market Crashes. Biodiversity of Deserts —— SARA OLDFIELD (Photography by Bruce Coleman Collection). 2005. Deserts: The Living Drylands. (ISBN 0-262-15112-X, hbk.). The MIT Press, 5 Cambridge Center, Cambridge, MA 02142-1493, US.A. (Orders: 800-405-1619, mitpress- orders@mit.edu, http://mitpress.mit.edu). $29.95, 160 pp., 180 illustrations, color throughout, 12 1/2" x 91/2". Publisher Comments: “Deserts represent the ultimate ch allenge to life on Earth. Their lack of water | (=) and extreme temperatures make survival difficult for both w ‘ildlife e and people, yet deserts are ric in animal and plant life and culture. Their unique species and ancient civilizations include fragile treasures needing protection in a rapidly changing world. Deserts are among our planet's last great wilderness regions, and they continue to offer scientific puzzles and new discoveries. Deserts: The Living Drylands isa celebration of the world’s least understood ecosystems. In this richly illustrated book —leaturing 180 color photographs—wildlife expert Sara Oldfield leads readers on a journey to some of the most remote places on earth, from the rolling sand dunes of the Arabian Peninsula’s Empty Sea to the ancient rock formations of central Australia. Deserts is both a vivid inventory of spectacular images, facts, and stories and an invaluable source of reference. It brings to life our planet s pee ar a habitats at a time when they are more in need of protection than ever. = a book with incredibly beautiful and often unbelievable photographs Once you open the book you will not want to close it. If you think you've seen the desert, think again. The seven chapters take you through a survey of deserts of the world: Africa, The Middle East, Asia, Australia, The Americas, and ends with the future of drylands.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A SIDA 21(2): 766. 2004 A NEW COMBINATION IN ANTENNARIA (ASTERACEAE: GNAPHALIEAE) FROM NORTH AMERICA Randall James Bayer CSIRO, Pane MigusDy Centre for Plant B Australian Nab onal Herbarium GPO Box Canberra, 2601, hiner randy.bayer@csiro.au ABSTRACT Antennaria pulcherrima subsp. eucosma (Fernald & Wiegand) R,J. Bayer, comb. nov., is based on Antennaria eucosma (Fernald & Wiegand) from North America. The new combination is essential as a result of the expansion of the circumscription of A. pulcherrima to include A. eucosma. RESUMEN Antennaria pulcherrima subsp. eucosma Gone & ain RJ. Bayes come nov., se aes en J j Antennaria eucosma (Fernald & WV resultado de la extension de la Gren cicn de A. pees para incluir A. eucosma. INTRODUCTION In preparation for publication of my treatment of Antennaria Gaertn. for the Flora of North America North of Mexico, | came to re-evaluate Antennaria pulcherrima (Hook.) Greene, an amphimictic species, occurring as diploid Qn = 28) and tetraploid (2n = 56) cytotypes from Colorado to Alaska, east to Ontario, and parts of western Quebec. It appears to be uniformly tetraploid throughout its range in Canada and Alaska (Urbanska 1983), but four diploid populations are known from Montana (one population), Wyoming (one population), and Colorado (two populations) (Bayer & Stebbins 1987). Apparently, A. pulcherrima arose in the U.S. Rockies as a diploid and presumed derived autotetraploid cytotypes spread north into Canada and Alaska post glaciation. It is generally found in moist willow thickets throughout its range (Urbanska 1983; Bayer & Stebbins 1987). Porsild (1943) suggested that A. pulcherrima is restricted to cal- careous soils; this has never been empirically demonstrated (Urbanska 1983). Antennaria eucosma Fernald & Wiegand is a narrow endemic restricted to lime- stone and serpentine barrens on the island of Newfoundland and Anticosti Is- land, Quebec. Like the Canadian populations of Antennaria pulcherrima, A. eucosma is amphimictic and tetraploid (Urbanska 1983; Bayer & Stebbins 1987). Its morphological similarity to A. pulcherrima is obvious and the two are sepa- SIDA 21(2): 767-769. 2004 768 BRIT.ORG/SIDA 21(2) rated primarily by the presence of prominent flags! in A. pulcherrima and their usual absence in A. eucosma. Porsild (1943, 1965), who had quite a narrow species concept in Antennaria, treated A.eucosmd asasynonym of A. pulcherrima. Urbanska (1983), who stud- ied the ecology, cytology, and distribution of the Pulcherrimae group, while rec- ognizing it as a distinct species, stated that, A. eucosma may represent nothing more than a specialized group of populations of A. pulcherrima with a distinct ecology. It has the same chromosome number as boreal populations of A. pulcherrima and its range is parapatric with that of A. pulcherrima. Further- more, she suggested that Antennaria eucosma may have arisen via quantum speciation and genetic drift in geographically marginal populations of A. pulcherrima (Urbanska 1983). Ledyard Stebbins and I recognized A. eucosma as a distinct species (Bayer & Stebbins 1987, 1993), and we remarked on the prob- lem of its circumscription as a distinct species (Bayer & Stebbins 1993). Because Antennarid eucosma is also morphologically very similar to, and not always consistently distinct from, A. pulcherrima, it is best to recognize A. eucosmd as an intraspecific taxon under A. pulcherrima. Stebbins and l applied the same set of taxonomic criteria when evaluating A. luzuloides Torr. & A. Gray subsp. luzuloides and A. luzuloides Torr. & A. Gray subsp. aberrans (EE. Nelson) RJ.Bayer & Stebbins (= A. microcephala A. Gray); the two subspecies differ in their ecology, but are not adequately morphologically differentiated to warrant recognizing them as distinct at species rank (Bayer & Stebbins 1993). A scenario whereby marginal populations of tetraploid A. pulcherrima in At- lantic Canada, which may already have been adapted to calcareous soils, be- came genetically isolated and subsequently morphologically and ecologically differentiated from the central populations of A. pulcherrima, seems apparent. TAXONOMY ee pulcherrima (Hook.) Greene subsp. eucosma (Fernald & oe Bayer, comb. nov. Ant aFernald& Wiegand Rhodora 13:23. 1911. TyPr: CANADA. NEWFOUNDLAND: Flora of Westen Newfoundland, Region of Port au Port Bay, dry limestone barrens, upper slopes and tablelands, altitude 200-300 m, Table Mountain, 16 Aug 1910, MLL. Fernald & K.M. Wiegand 4144 (HOLOTYPE: GH: IsoryPes: CANI, NY) inaria carpatica (Wahlenb.) R.Br. var. humilis Hook., Fl. Bor. Amer. 1:329. 1834. KEY TO THE SUBSPECIES OF ANTENNARIA PULCHERRIMA Distal cauline leaves usually flagged; eh corollas 4.5-6.5 mm; staminate corollas mm; wet habitats in willow thickets at subalpine elevations or subarctic in western North America from eee to Alaska, east to Ontario, parts of weste Juebec ree pulcherrima Flags are flat, linear, scarious appendages of the leaf tips that are similar to the tips of the phyllaries, not to confused with ordinary subulate or blunt leaf ti ps that are ¢ ntially and herbaceous BAYER, A NEW COMBINATION IN ANTENNARIA 769 Distal cauline leaves mostly not flagged (sometimes flagged just proximal to heads); pistillate corollas 3-4.4 mm; staminate corollas 3-4 mm; limestone substrates in willow thickets of western Newfoundland and Anticosti Island, Quebec subsp. eucosma ACKNOWLEDGMENTS I thank curators at CAN, GH, and NY for making specimens available for study. Appreciation is extended to Kirsten Cowley, Edward Cross, Guy Nesom, John Strother,and Matthew Unwin for their critique and helpful suggestions concern- ing the draft manuscript and to J. Tupac Otero for editing the Spanish summary. REFERENCES Bayer, R.J.and G.L. Stessins. 1987.Chromosome numbers, patterns of distribution, and apo- mixis in Antennaria (Asteraceae: Inuleae). Syst. Bot. 12:305-319. Bayer, R.J. and G.L. Stessins. 1993.A synopsis with keys for the genus Antennaria (Asteraceae: Inuleae: Gnaphaliinae) for North America. Canad. J. Bot. 71:1589-1604. Porsitp, A.E. 1943. Materials for a flora of the continental Northwest Territories of Canada. Sargentia 4:1-79. Porsito, A.E. 1965. The genus Antennaria in eastern arctic and subarctic America. Saetryk af Bot. Tidssk. 61:22—-55. Ureanska, K.M. 1983.Antennaria carpatica (Wahlb.) Bl. et Fing.s.l.in North America. |.Chro- mosome numbers, geographical distribution and ecology. Ber. Geobot. Inst. E.7.H., Stiftung Rubel 50:33-66. 770 BRIT.ORG/SIDA 21(2) BOOK NOTICES/BOOKS RECEIVED Annual Review of Phytopathology DesoraH P. DELMER, HANs J. BOHNERT, and SABFEHA MERCHANG (editors). 2003. An- nual Review of Phytopathology: Volume 42, 2004. (ISBN 0-8243-1342-9, hbk; ISSN 0066-4286). Annual Reviews Inc., 4139 El Camino Way, P.O. Box 10139, Palo Alto, CA 94303-0139, U'S.A. (Orders: www.AnnualReviews.org, 800-523-8635, 650-493-4400, 650-424-0910 fax). $173.00 (USA), $178.00 (Intl), 498 pp., 6" x 9", Contents of Volume 54 of Annual Review of Phytopathology: « The Accidental Plant Pathologist * Tobacco Mosaic Virus: A Model System for Plant Biology « Assessment and Management of Soil Microbial Community Structure for Disease Suppression « Analysis of Disease Progress as a Basis for Evaluating Disease Management Practices «Evolution of Plant Parasitism Among Nematodes « Lessons Learned from the Genome Analysis of Ralstonia solanacearum . ela ae ane! Resistance i in Wheat and Barley to Fusarium Head Blight “Cc na of Citrus-Associated Bacteri «Systemic Acquired Resistan «Molecular Aspects of Plant " irus Transmission by Olpidium and Plasmodiophorid Vectors * Microbial Diversity in Soil: Selection Microbial Populations by Plant and Soil Type and Implications for Disease Suppressiveness * Microbial Dynamics and Interactions in the Spermosphere « Biological Control of Chestnut Blight with Hypovirulence: A Critical Ana «Integrated Approaches for Detection of Plant Pz ee ee iaand ae “ol Bacterial Diseases * Nematode Molecular Diagnostics: From Bands to Barcodes Type Ill Secretion System Effector Soe as Double Agents in Bacterial Disease and Plant Defense «Plant Virus Satellite and Defective Interfering RNAS: New Paradigms for a New Century * Chemical Biology of Multi-Host/Pathogen Interactions: Chemical Perception and Metabolic Complementation > «Index and Errata SIDA 21(2):770. 2004 NOMENCLATURE OF THE VIRGINIA-BLUEBELL, MERTENSIA VIRGINICA (BORAGINACEAE)! James S. Pringle Royal Botanical Gardens PO. Box 399 Hamilton, Ontario, CANADA L8N 3H8 jpringle@rbg.ca ABSTRACT Mertensia virginica (L.) Pers. ex Link is the correct name for the Virginia-bluebell. It should not be replaced by M. pulmonarioides Roth. RESUMEN Mertensia virginica (L.) Pers. ex Link es el seciaes correcto para la especie usualmente asi conocida. No debe reemplazarse por M. p Virginia-bluebell or Virginia-cowslip, Mertensia virginica (L.) Pers. ex Link, is native to eastern North America and widely cultivated as an ornamental pe- rennial. It has consistently been called M. virginica in floras covering its natu- ral range, including all editions of Gray’s Manual of Botany (Gray 1848, 1856, 1862, 1863, 1867; Gray et al. 1890; Robinson & Fernald 1908; Fernald 1950) and continuing to the most recent state and provincial floras and checklists. That name was accepted in Williams's (1937) monograph on Mertensia in North America and in Al-Shehbaz’s (1991) monograph on the Boraginaceae of the southeastern United States. It is also the accepted name in horticultural refer- ences including Hortus III (Staff...1976), the most recent edition of The Royal Horticultural Society Dictionary of Gardening (Huxley et al. 1992), The Plant- Book (Mabberly 1997), and The European Garden Flora (Richardson 2000). In recent years, nevertheless, this species has with increasing frequency been called M. pulmonarioides Roth in horticultural literature and in seed and nursery catalogues. All or most post-1990 use of the name M. pulmonarioides is probably derived directly or indirectly from Index Hortensis (Trehane 1989), which is an exception among standard references in that the name M. pulmonarioides is accepted and M. virginica is listed as a synonym. Following the publication of Index Hortensis, the name M. pulmonarioides was accepted in the 1997 edition of The American Horticultural Society A-Z Encyclopedia of Garden Plants (Brickell & Zuk 1997) with M. virginica as a synonym. The name M. pulmonarioides was also accepted in the 1998/99 edition of the RHS Plant Finder (Lord et al. 1998), although in the 2003/04 edition (Lord et al. 2003) M. ~ ‘Contribution No. 118 from the Royal Botanical Gardens, Hamilton, Ontario, Canada SIDA 21(2): 771-775. 2004 772 BRIT.ORG/SIDA 21(2) virginica isthe accepted name and M. pulmonarioides is cross-indexed as a syn- onym. Schmid (2002), in An Encyclopedia of Shade Perennials, commented that with taxonomists having “switched” the long-established name M. virginica to M. pulmonarioides, the latter name would “show up more and more often.” Nomenclatural matters are not discussed in Index Hortensis, and the name M. virginica was used in the literature cited for the species. The name M. pulmonarioides may have been taken from the original component of Index Kewensis Jackson 1894), in which M. pulmonarioides was listed as an accepted name and M. virginica as a synonym. No taxonomic publication that specifi- cally discusses the nomenclature of Virginia-bluebell was cited in the exten- sive bibliography compiled by Al-Shehbaz (1991) or in The Kew Record of Taxo- nomic Literature Relating to Vascular Plants or any of the other bibliographies | consulted. In view of the confusion now associated with the nomenclature of this species, it seems appropriate to present such a study here. Virginia-bluebell was named and described at the rank of species by Linnaeus (1753) in the first edition of Species Plantarum, as Pulmonaria virginica. Two specimens are relevant to the typification of this name. One, des- ignated the type by Williams (1937) and accepted as such by Charles E. Jarvis in the Linnaean Plant Names Typification Project (M. Grant, pers. comm.), is from Linnaeus’s own herbarium (LINN) and was collected in North America by Pehr Kalm. It bears the Sprague Catalogue number 184.5. From a photograph in the library of the Arnold Arboretum, this specimen is unequivocally identi- fiable as Virginia-bluebell. Linnaeus cited an earlier description from Gronovius’s (1739) Flora Virginica, which had been based on a manuscript by John Clayton. The speci- men Clayton 339, now in the herbarium of the Natural History Museum (Lon- don) (BM), is from Gronovius’s herbarium. According to data on the herbarium sheet, it was taken froma plant sent from Virginia to England by Mark Catesby and subsequently cultivated in the garden of Peter Collinson. An image can be seen on the museum's web site, with the designation “Linnaean type status: origi- nal element.” Its identity as Virginia-bluebell is likewise unmistakable. The genus Mertensia was segregated from Pulmonaria in 1797 by A.W. Roth, who gave Virginia-bluebell the new name M. pulmonarioides. Roth did not mention the name P. virginica L.and may not have been aware that he had described the same species. The name M. pulmonarioides should therefore be considered a heterotypic synonym. Pulmonaria virginica L., with the species epithet virginica retained, was transferred to Mertensia in 1829 by J.H.F Link, who cited M. pulmonarioides Roth as a synonym. Link attributed the combi- nation M. virginica to Persoon, probably on the basis of herbarium annotations or correspondence, as there is no record of Persoon’s actually having published the binomial. The authorship of the binomial may therefore be cited as “(L.) Pers. ex Link” or simply as “(L.) Link.” The combination is occasionally attrib- — PRINGLE, NOMENCLATURE OF MERTENSIA VIRGINICA 773 uted to Persoon, following Fernald (1950) or Gleason and Cronquist (1991), who routinely omitted the name of the publishing author in such contexts, but un- der current rules of nomenclature that is incorrect. The binomial M. pulmonarioides Roth was published before M. virginica (L.) Pers. ex Link, but as long as the taxonomic rank is not changed, priority is based on the date of publication of the species epithet rather than on that of the binomial. Adoption of the earliest available species epithet is retroactively required under Articles 11.4 and 52 of the International Code of Botanical No- menclature, 2000 ed. (Greuter et al. 2000). In this case, the earliest available epi- thet for the species described by Linnaeus is virginica, the priority of which dates from 1753, the starting point of botanical nomenclature. During the interval 1797-1829 three homonyms of the genus name Mertensia were published: Mertensia Willd. for a genus of ferns; Mertensia Thunb. ex Roth for a genus of red algae; and Mertensia Kunth for a genus of Ulmaceae. These illegitimate homonyms were used only briefly and by few authors. A search of Broun’s (1938) Index to North American Ferns and the In- ternet resources AlgaeBase and The International Plant Names Index (websites) indicated that the species epithet virginica was not used under any of those homonymic genus names. Nor did the IPNI or any other index disclose any pre- 1829 use of the combination M. virginica for any other species in the boraginaceous genus. In summary.—The traditional use of the name M. virginica is compatible with its typification. The epithet virginica in this context has priority from the starting point of botanical nomenclature, 1753, whereas that of pulmonarioides is from 1797. The binomial M. virginica (L.) Pers. ex Link isnot a homonym of a name previously used for any other species. The familiar name Mertensia virginica (L.) Pers. ex Link is the correct name for the Virginia-bluebell; it should not be replaced by M. pulmonar ioides Roth. ACKNOWLEDGMENTS lam grateful to Mike Grant and W George Schmid for very helpful correspon- dence, and to my colleague Dennis Eveleigh for useful information and valued suggestions. I am also grateful to the staff of the Gray Herbarium/Arnold Ar- boretum Library for access to historic literature and a photograph of the Lin- naean herbarium specimen. Crinan Alexander and Guy Nesom are thanked for reviewing the manuscript. REFERENCES AL-SHEHBAZ, ILA. 1991. The genera of Boraginaceae in the southeastern United States. J. Arnold Arbor. Suppl. Ser. 1:1-169. Brickett, C.and J.D. Zuk (eds.-in-chief). 1997. The American Horticultural Society A-Z ency- clopedia of garden plants. Macmillan Publishing Company, New York. 774 BRIT.ORG/SIDA 21(2) Broun, M. 1938. Index to North American ferns. Published by the compiler, Orleans, MA. FerNao, M.L. 1950. Gray’s manual of botany, ed. 8. American Book Company, New York. Republished 1970.D.Van Nostrand Company,New York.Republished 1987.Dioscorides Press, Portland, OR. Geason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada, ed. 2. The New York Botanical Garden, Bronx. Gray, A. 1848. A manual of the botany of the northern United States, ed. 1. James Munroe & Co,, Boston and Cambridge. Gray, A. 1856. Manual of the botany of the northern United States, ed. 2.lvison & Phinney, New York. Gray, A. 1862. Manual of the botany of the northern United States, ed. 3. vison, Phinney & Co., New York. Gray, A. 1863. Manual of the botany of the northern United States, ed. 4. Ivison, Phinney, Blakeman & Co., New York. Gray, A. 1867. Manual of the botany of the northern United States, ed. 5.lvison, Blakeman Taylor & Co., New York. Gray, A., S. Watson and J.M. Coutter (revisers). 1890 ["1889"]. Manual of the botany of the northern United States, ed. 6. American Book Company, New York Greurer, W., J. McNeict, F.R. Barrie, H.M. Buroet, V. DEMOULIN, T.D. Fiticueiras, D.H. NICoLson, PC. Sitva, J.E. Skoa, P. TReHANE, NJ. Turtano, and D.L. HawkswortH (eds.). 2000. International code of botanical nomenclature (St. Louis Code) Adopted by the Sixteenth International Bo- tanical Congress St. Louis, July-August 1999. Regnum Veg. 138. Koeltz, Konigstein Gronowus, J.F.1739. Flora Virginica exhibens plantas quas V.C. Johannes Clayton in Virginia observavit atque collegit, pars prima. Cornelius Haak, Leiden. Republished 1946. Ar- nold Arboretum, Jamaica Plain, MA. Huxtey, A., M. GrirritHs, and M. Levy (eds.). 1992. The new Royal Horticultural Society dictio- nary of gardening. The Stockton Press, New York; The Macmillan Press, London. Jackson, B.D. 1893-1895. Mertensia. In: Index Kewensis [original 2 vols.].Clarendon Press, Oxford. 2(3):213. 1894. Link, J.H.F. 1829. Handbuch der Erkennung der nutzbarsten und am hdufigsten vorkommenden Gewachse. Erster Theil. Haude und Spenerschen Buchhandlung (S.J. Josephy), Berlin Linnaeus, C. 1753. Species plantarum, ed. 1. Lars Salvius, Stockholm. Republished 1957- 1959. The Ray Society, London. Lorb, T., et al. (eds.). 1998. The RHS plant finder 1998-99. Dorling Kindersley, London Loro, T., et al. (eds.). 2003. Royal Horticultural Society plant finder 2003-2004. Dorling Kindersley, London Maseertey, D.J. 1997. The plant-book, ed. 2. Cambridge University Press, Cambridge. RicHARDSON, J.E. 2000. Mertensia Roth. In: The European garden flora. Cambridge University Press, Cambridge. 6:134-135. Rosinson, B.L. and M.L. Fernaco. 1908. Gray's new manual of botany, ed. 7. American Book Company, New York. PRINGLE, NOMENCLATURE OF MERTENSIA VIRGINICA 775 Roth, A.W. 1797. Catalecta botanica. Fasc. 1.J.G. Muller, Leipzig ScHmio, W.G. 2002. An encyclopedia of shade perennials. Timber Press, Portland, OR. STAFF OF THE LiperTY Hype Baitey Hortorium. 1976. Hortus third. Macmillan Publishing Com- pany, New York. TREHANE, [R.]P. 1989. Index hortensis. Volume 1:Perennials. Quarterjack Publishing, Wimborne, Dorset, U.K. Weesite. Accessed 19 July 2004. www.algaebase.org/ Weesite. Accessed 19 July 2004. www.ipni.org/index.html| Wiuams, L.O. 1937.A monograph of the genus Mertensia in North America. Ann. Missouri Bot. Gard. 24:17-159. 776 BRIT.ORG/SIDA 21(2) BOOK NOTICES/BOOKS RECEIVED Wild Orchids of the Southeastern United States PauL Martin Brown with drawings by Stan Foism. 2004. Wild Orchids of the Southeastern United States, North of Peninsular Florida. (ISBN 0-8130- 2749-7, pbk.) University Press of Florida, 15 Northwest 15th St, Gainesville, FL 32611-2079, U.S.A. (Orders: www.upf.com, 800-226-3822). $27.95, 394 pp., color ae b/w illus. 6" x 9" Orchids of Florida is a Publisher Comments: “In more than 100 years of orchid research in Florida, Wilk the first field g ae to be published for this orchid-rich state. Providing more than 400 color photos s by Stan Folsom, distribution maps, and detailed descriptions of each spe- as well as line draw a ble even the novice to easily identify any of the orchids found growing in cles, this guide Ss the wild. “Illustrated keys are provided to aid in identification. Each of the 117 species and varieties has atull page of text, a line drawing, and distribution om with a facing page of full-color photographs. ) treated. A final chapter enabling the reader Species deemed as naturalized, escaped, or waifs are als¢ to use this book in the nearby southeastern ar Plain states makes it an even more valuable re- source.” From the color photos and line drawings to the simple but general range maps, Paul Martin Brown has given us another good Wild Orchids book of the USA, this time from the southeastern United States.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 21(2): 776. 2004 CUIBF CUDWEED Al SPECIFIC RANICIN PSEUDOGNAPHALIUM (ASTERACEAE: GNAPHALIEAE) Harvey E. Ballard, Jr. Danielle Sky Feller Department of Envi | & Plant Biology Robert K. Godfrey Herbarium Ohio University Biology Unit 1, Room 100 Athens, Ohio 45701-2979 U.S.A. Florida State University Tallahassee, Florida 32306-4370, U.S.A. Guy L.Nesom Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT ae m saxicola is treated at specific rank as Pseudognaphalium saxicola (Fassett) H.F.Ballard Feller, ,comb. nov, It is likely an pecuniee Gam Ne of Pseudognaphalium obtusifolium but is msistently different ina number of ters. No intermediates between P saxicola a any other taxon have been ob Oe SUMEN ~ L Bs 2 ] t t ¥ ] eo) i D 1 L k . b Paty a ea each To i ? I ena] oaiaie & Fe ‘fie comb. nov. derivad lutivo de Pseud hali btusif es bastante diferente en un nimero de caracteres morfoldgicos. t Now se a observado inkemaedies entre P saxicola y cualquier otro taxon. The “Cliff Cudweed,” Gnaphalium saxicola Fassett, is endemic to a relatively small area of Wisconsin. It was initially described asa distinct species but sub- sequently reduced in rank to a variety of Pseudognaphalium (Gnaphalium) obtusifolium (L.) Hilliard & Burtt. Cronquist (1946, p. 12D noted that “At least until a larger series of specimens demonstrates its morphological and genetic discontinuity, it seems better treated as a variety of G. obtusifolium.” We find that treatment at specific rank most accurately reflects the biological and evo- lutionary situation and transfer of G. saxicola to the genus Pseudognaphalium is necessary to reflect newly emerging understanding of natural evolutionary groups in the Gnaphalieae. The new combination will be available for use in the forthcoming “Flora of North America” treatments of Asteraceae. Pseudognaphalium saxicola (Fassett) H. anes & pene comb. nov. Gnaphalium saxicola Fassett, Rhodora 33:75. 1931 } tusifolium var. saxicola (Fassett) Cro Rhodora 48:121. 1946. Pseudognaphalium com m var. saxicola (Fassett) Kartesz, Syn- thesis N. Amer. Fl, Nomencl. Innov: no. 32. 1999. TYPE: U.S.A. WISCONSIN. ADAMS Co.: Coldwater SIDA 21(2): 777-779. 2004 778 BRIT.ORG/SIDA 21(2) Canyon, os of ne Wisconsin River, sandstone ledges, 22 Sep 1929, N.C. Fassett, EM. Uhler, ~TI] and \ HOLOTYPE: WIS ISOTYPE: GH!). Plants annual, Sica Stems erect, 4-15(-30) cm tall, filiform, per- sistently tomentose with a loose, envelope-like, transparent haze of extremely thin hairs, doubling the stem width, eglandular. Leaves cauline, 4-6, elliptic- oblanceolate to oblanceolate, gradually narrowed to the base, not clasping or decurrent, 5-30 mm, largest at midstem, sessile, green on both surfaces, within veiny reticulum evident, thinly arachnoid-tomentose to glabrate, eglandular, Heads 2-4 in a terminal, capitate cluster, commonly immediately subtended by uppermost cauline leaf, sometimes several clusters in a corymbiform array. Involucres turbinate, 4-5 mm; phyllaries narrowly triangular to narrowly ob- long-triangular, acute, in 3(-4) gradate series, whitish to slightly tawny. Pistil- late florets 25-28. Bisexual florets 6-7. Cypselae smooth, without raised ridges or papillae. Flowering Qul-)Aug-Sep. Mostly bare sandstone cliff faces, ledges, and cracks, S- to E-facing but commonly shaded; 200-300 m; Wisc. (Adams, Co- lumbia, Richland, Sauk, and Vernon cos.). It seems likely that Pseudognaphalium saxicola is an evolutionary deriva- tive of P obtusifolium but it consistently diverges from the latter in many re- spects. Full details of complementary studies conducted by Ballard and Kowal (1992) and Feller (2000) will be submitted as a combined manuscript for pub- lication but are summarized here in support of the nomenclatural transfer. Phe- netic and preliminary greenhouse comparisons of P. saxicola with P. obtusifolium, P helleri(Britt.) Anderb.,and P micradenium (Weatherby) Nesom have distinguished P saxicola by the following: annual duration and absence of a basal rosette; shorter stature (commonly only a few centimeters tall in fruit); loose, partially det bby-tomentose pubescence on stems, and com- plete absence of glands or elendulae hairs on stems and leaves; fewer leaf nodes with shorter and broader, thinner and more membranous, essentially glabrous leaves; larger and more open inflorescence; few, small heads with relatively few florets; and more slender, uniformly narrowly acute-tapering, semi-translucent phyllaries. Depauperate individuals of P obtusifolium over its whole geographic range may sometimes be as short as 5-10 cm, approaching the habit of P saxicola: such plants differ from P saxicola, however, in their close and more dense stem vestiture, bicolored and relatively narrow leaves, larger heads with greater num- ber of pistillate florets, and broader phyllaries with rounded apices. No inter- mediate specimens between P. saxicola and any other taxon have been con- firmed in hundreds of collections. Genetic studies of Pseudognaphalium saxicola populations using Inter- Simple Sequence Repeats indicate that the taxon is strongly and perhaps obli- gately apomictic; isolated greenhouse flats set abundant seeds, supporting this hypothesis. BALLARD ET Al COMB. NOV. 779 ACKNOWLEDGMENTS The authors gratefully appreciate the assistance of curators of herbaria at Har- vard University (GH), Philadelphia Academy of Sciences (PH), Smithsonian Institution (US), University of Michigan (MICH), University of Wisconsin-Mil- waukee (MIL) and University of Wisconsin-Madison (WIS) for facilitating stud- ies of herbarium collections of Pseudognaphalium. June Dobberpuhl provided help with development of the project and secured funding from the Wisconsin Bureau of Endangered Resources and the US. Fish & Wildlife Service for it. Staff of Wisconsin State Parks were helpful in gaining access to sandstone outcrops for searches. REFERENCES Battarp, H.E., Jk.and R.R. Kowal. 1992. Status survey and taxonomic study of Gnaphalium saxicola Fassett. Report, Bureau of Endangered Species, Department of Natural Re- sources, Madison, Wisconsin. Cronaquist, A. 1946. Notes on the Compositae of the northeastern United States. Ill. Inuleae and Senecioneae. Rhodora 48:116-125. Fetter, D.M.S. 2000. A study of the distribution, ecology, and genetic diversity of Pseudo- gnaphalium saxicola (Fassett) comb. nov. (Asteraceae), a rare annual plant endemic to Wisconsin. Masters Thesis, University of Wisconsin. 780 BRIT.ORG/SIDA 21(2) BOOK NOTICES/BOOKS RECEIVED Natural Resources Management PE Frouiiott, L.A Bojorquez,and M. HERNANDEZ-NArRVAEZ. 2001. Natural Resources Management Practices: A. Primer. (ISBN 0-8138-0913-4, pbk.) lowa State Press, A Blackwell Publishing Company, PO Box 570, Ames, IA, 30010-0570, U.S.A. (Orders: 800-862-6657; www.iowastatepress.com). $34.99, 256 pp., b/w figures, tables, 5 appendices, 6" x 9". The topic of natural resources practices are of ever- Sead interest to many. Authors, Ft eek ui Bojorquez and Hernandez-Narvaez, provide a wonderfu of many natural resource arenas in their book entitled Natural Resources Management Pr ractices: A Primer. The easy- to- understand text would t be appropl late for underg1 aduate of natural resources management. In addition, it is well suited for anyone with an interest in a practices that help preserve wildlife, outdoor recreation areas and that help protect soil, water, for- ests and other natural products and resources. ls, rangeland, tim- he eX i practices ft Waller, pest control, soil con- > te ber, ee wildlife ‘che ries, outdoor recreation areas, wilderness, fire anc servation, and ne of disturbed lands. There is also an excellent discussion of the decision- making process when considering various eapaeement cae in the integrated natural resources management chapter. Each chapter ends witha review of the main features of each management topic. Throughout the text the reader is encouraged to ae of management in a broad ecosystem- based approach and to mere potential impacts from changes in practices regarding one use on other uses or activities in an area. Basic backgrounds for each management topic areas are intro- duced prior to discussion of management techniques and greatly enhance understanding of the m gas t methodology discussed. The “real world” examples of selected management techniques are incredibly helpful and interesting. Many such real world applications are referenced to journal ae W nf na oe eee i agin ona technic or ne - interest. The py f management meth- ods. heres isa brief introduction to experi tal design ane statistical analysis of plot studies per- formed to measure potential impacts of management practices. These plot studies allow researchers to better predict the sons a} am: nese ment mele od by testing that method on amall als ar- eas. Compute that may arise from application of a Management technique in ina virtual way. Geogr aphic | Inform ma- tion Systems (GIS) are discussed as a possible tool in evaluating and guiding management methods. The authors used English units in a number of examples. English units may increase the un- derstanding of examples for many readers, as they are familiar with feet and inches, but metric units srovided (Systeme Internationale) are considered more appropriate in academia The authors have = values in an appendix to allow conversion to metric values if Natural Resources Management Practices: A. Primer eee a very informative introduction to many facets of natural resources management. The text also provides a well-rounded overview to related topics of data analysis, computer simulation and GIS to evaluate or compare management techniques. The book is See ers not only for science or engineering students, but also for any reader who wishes to gain a better understanding of natural resources management techniques an rationale.—Lee Luckeydoo, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-40060, U.S.A. SIDA 21(2): 780. 2004 PSEUDOGNAPHALIUM CANESCENS (ASTERACEAE: GNAPHALIEAE) AND PUTATIVE RELATIVES IN WESTERN NORTH AMERICA Guy L.Nesom Botanical Research Institute of Texas can Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT ens,and = Pseud phalium canescens, Pseud halium microcephalum, Pseudognaphalium beneo Pscuilognaplia alium thermale (E.E. Ne eon) Nesom, comb. nov, are treated as separate species. cent treatment of the California flora combined these taxa as subspecies within a single species © canescens) but earlier floristicians have viewed them as distinct species. The current study also con- cludes that they are distinct and appropriately treated at specific rank. A nomenclatural and eco- logical summary and a distribution map are provided for each species and the four are morphologi- cally distinguished in a key. A lectotype is designated for Gnaphalium wrightii. RESUMEN Pse aa a dee canescens, PSeucee nap eau microcephalum, Pseudognaphalium beneolens, y e (E.E. Nelson) N comb. nov, se tratan como Caio gaan }: Un reciente tratamiento ee a fl ora ee Calitornia cout pipals estos taxa c 5 ie (Pc El presente x c [ bi a ne ] | ivel especilico. Se ofrece = t y qu un resumen nomenclatural y ecoldgico, asi como un mapa de Gistabicion de cada Seals y se g por wrightit. For the forthcoming treatment of Pseudognaphalium in the Flora of North America, | will recognize as species four taxa that were recently treated (Stebbins & Keil 1992; Stebbins 1993) as subspecies of a single species: P canescens (DC.) A. Anderb., P. microcephalum (Nutt.) A. Anderb., P beneolens (Davidson) A. Anderb., and (as validated below) P thermale (E.E. Nelson) Nesom. Pseudognaphalium thermale (E.E. Nelson) Nesom, comb. nov. Gnaphalium thermale E.E. Nelson, Bot. Gaz. 30:121. 1900. Gnaphalium microcephalum var. thermale (E.E. Nelson) Cronq., Leall. West. Bot. 6:47. 1950. Gnaphalium mic ae m subsp. thermale (E.E. Nel- son) G.W. Douglas, Canad. J. Bot. 64:2726. 1986. Gnaphalium canescens subsp. thermale (E.E. Nelson) Stebbins & Keil, Novon 2:437. 1992. ie m canescens subsp. thermale (E.E. Nelson) Kartesz, Syn. Checklist & Atlas . Innov. 1999. Pseudognaphalium eee (Nutt.) Anderb. var. thermale (E. a Dorn. Vasc. Pl. Wyoming, ed. 3, 375. 2001. TyPE: U.S.A. WYOMING. [Park Co. eens Park, geyser formations of Norris geyser basin, see 1899, A. Nelson and E. Nelson 6139 (HOLOTYPE: RM; ISOTYPES: GH!, US). Gnaphalium williamsii Rydb., Bull. Torrey Bot. Club 37:324. 1910. TYPE: U.S.A. MONTANA. [Flat- Is, LL Aug 1894, R.S. Williams s.n. (HOLOTYPE: NY). head Co. Columbia Fa SIDA 21(2): 781-789. 2004 782 BRIT.ORG/SIDA 21(2) Gnaphalium johnstonii G.N. Jones, Univ. Wash. Publ. Bot. 7:159, 176. 1938. Type: U.S.A. WASHING- TON. [Thurston Co. upper valley of the Nesqually, open, barren ground, 15 Sep 1896, O.D. Allen = 223 (ISOTYPE [as so annotated]?: WTU, a caagel Flowering Jun-Sep(-Oct). Dry, sandy roadbanks, roadside ditches, river beds and banks, lakeshores, granitic sand, open woods of yellow pine, Jeffrey pine, red fir, Douglas fir, mixed conifer, and mixed evergreen; (50-)300-2300(-2500) m. British Columbia; California, Idaho, Montana, Nevada, Oregon, Utah, Wash- ington, Wyoming. Pseudognaphalium beneolens (Davidson) A. Anderb., Opera Bot. 104:147. 1991. Gnaphalium canescens subsp. beneolens aco Stebbins & ms Novon 2:437. 1992. ole es sees ee subsp. | son) Kartesz, Syn. Checklist & Atlas, Innov. 1999. oe beneolens er Bull S. Calif | Sci. 17:17. 1918. TYPE: Wy U a CALIFORNIA. [Los Angeles Co.| Crescenta, 1 Sep 1917, FE. Burlew 3275 (ISOTYPE: GH)). Flowering (Apr-)Jun-Oct. Dry, open slopes and ridges, river beds, roadbanks and other disturbed sites, sandy flats, dunes, coastal sage scrub, chaparral, yel- low pine, foothill pine, blue oak woodland; (1-)50-850(-1950) m. California; Mexico (Baja California). Pseud hali i halum (Nutt.) A. Anderb., Opera Bot. 104:147. 1991. Guiphatinwn naicmcephalum Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:404. 1841. Gnaphalium canescens subsp. i ae Steepins & Keil, Novon 2:437. 1992. Pseudogna- Kartesz, Syn. Checklist &@ Atlas, Nom. Innov. 1999, Type: U.S.A.CAL IFORNIA, in Diego Co.|: San Diego, aie or early May], 1836, T. Nuttall s.n. (BM, photo-GH)). After examining a photo of the type of Gnaphalium ae Johnston (1924) noted that G, microcephalum was correctly applied to the species usually called Gnaphalium bicolor Bioletti (= Pseudognaphalium biolettii A. ee ), he pro- posed Gnaphalium albidum to account for the species left wattiont a name. ine piesent in- NJ ] terpretation, however, of ree and of I'sd re lum in the concept here (see i below). Ferris (1960 , p. 469) scam ie shotoaraok of the type and her interpretation of its identity was similar to that here. She commented that “The specimen probably was collected in the spring (see Madrono 2:146-47. 1934) [Jepson 1934]. This would account for the weather-beaten appearance (darkened olaee and rubbed tomentum) of this plant which nor mally flowers in summer and earl y la fa Gnaphalium albidum I.M. Johnston, Contr. Gray Herb. 70:84. 1924. Type: U.S.A. CALIFORNIA. San Diego Co. Granite, in chaparral, 1850 ft, 11 Jul 1916, M.F Spencer 69 (HOLOTYPE: GHI; ISOTYPE: Us!) Flowering (Apr-)Jun-Aug(-Nov). Grassy hillsides, gravelly canyon bottoms, coastal sage scrub, chaparral; 50-900(-1800) m. California; Mexico (Baja California). Pseudognaphalium canescens (DC.) A. Anderb., Opera Bot. 104:147. 1991 (non W. A. Weber 1991). Gnaphalium canescens DC., Prodr. 6:228. 1838. TYPE: MEXICO. GUANAJUATO: Leon, Méndez s.n. (HOLOTYPE: G-DC, fiche!, photos Fl and TEX!). Gnaphalium wrightii A. Gray, Proc. Amer. Acad. Arts 17:214. 1882. T ype: US.A. TEXAS. [E] Paso or Hudspeth Co: valley between El Paso and the Guadalupe Mts., Oct [1849], C. Wright NESOM 394 ( LECTOTYPE, designated here: GH!; ISOLECTOTYPES: GH, US!). Gray’s protologue referred to three collections: he see) cited (1) Parry & Palmer 419 eat from San Luis Potosi, Mexico, collected in 1878; he cited “G. microcephalum, Gray, Pl. Wright, i. & ii, non Nutt.” in relerence to his earlier ee ene as Gnaphalium wenden of (2) a ee by Wright i Texas from the “Valley between El Paso and the Guadalupe Mts,; Oct.” (PI. W aves one Knowl. 3(5) 1852] 1:124. 1852) and (3) another collection by ae 1851 from New xico, “from mountains near the copper mines” (PL. Wright. [Smithsonian ae Knowl. 5(6) on 2:99, 1853). Gnaphalium we Osterh., Muhlenbergia 1:141. 1906. Type: U.S.A. COLORADO. Larimer Co:: mpson River, 16 Aug 1905, G.E. Osterhout 3158 (ISOTYPE [as annotated by C. Anderson, a NY). Gnaphalium sonorae LM. Johnston, Contr. Gray Herb. 68:99. 1923. TypE: MEXICO. SONORA: Hermosillo, 1888, M.A. Crawford s.n. (HOLOTYPE: GH; ISOTYPE: US). Gnaphalium viridulum LM. Johnston, Contr. Gray Herb. 70:86. 1924. TYPE: U.S.A. NEW MEXICO. [Grant Co. Bear Mts. near Silver City, 2400 m, 19 Sep 1903, O.B. Metcalfe 742 (HOLOTYPE: GH}; ISOTYPE: US)). Gnaphalium texanum LM. Johnston, Contr. Gray Herb. 70:86. 1924. TYPE: U.S.A. TEXAS. [Brewster Co.| mouth of “Tarlingua” [Terlingua Creek], Sep 1883, V. Havard 26 (HOLOTYPE: GHh IsoTYPE: USI). Flowering Aug-Nov(-Jan). Lava beds, grasslands, oak, pine-oak, and pine wood- lands, 1150-2450(-2700) m; Arizona, California, Colorado, New Mexico, Okla- homa, Texas, Utah; Mexico (Baja California, Chihuahua, Coahuila, Durango, Nuevo Leon, San Luis Potosi, Sinaloa, Sonora, and other states to the south). KEY TO PSEUDOGNAPHALIUM CANESCENS AND PUTATIVE RELATIVES IN CALIFORNIA 1. Basal and lower cauline 1.5-6 mm wide, similarly colored on abaxial and adaxial sur- faces, cauline decurrent 5— i mm, the decurrent portion appearing as a thin line, ee ge prem rently sessile peculelel ese the rommen te pale aoMleu cs ally or es reduced in size and becoming linear upwards, not coiling upon wilting; capitulescence corymbiform; involucres 4-5 mm long; phyllaries in 3- 4(-5) series, usually slightly hyaline and shiny; bisexual florets (2-)4—-7; habitats at (600-)900—2500 m Pseudo gnaphalium thermale 2. Stems 35-100 cm tall land| | ar-oblanceolate, relatively even in size and shape upwards, usually ea or ea coiling upon wilting; capitulescence usually onda nah involucres 5-6 mm long; a in (4 -)5- 6(- 7) series, usually op ; bisexual florets (3-) 5-10(-13); habitats at 5—800(-1550) m Psetidog naphalium beneolens . Basal and lower cauline leaves (2-)5-10(-15) mm wide, often ae bicolored, cauline not decurrent, eglandular (2 microcephalum) or eglandular to minutely and Hea eect sessile- genguet Wasa ccap ht plants not aromatic ms 50-100 cm tall, usually 3 near the base; leaves eglandular; See in 4-5 series, outer ovate; bisexual florets 4-9 Ps cudographalim cephalum ot i) gS —~ — 3. Stems 20-70 cm tall, usually 2-3 mm diameter near the base; leaves ieee to minutely and inconspicuously sessile-glandular beneath the adaxial tomen- tum; phyllaries in 3-4 series, outer ovate-lanceolate; bisexual florets (1-)2—5(-6), 4-6 more common in the USA Pseudognaphalium canescens 784 BRIT.ORG/SIDA 21(2) DISCUSSION All four of the taxa considered here occur in California (Figs. 1-4). Pseudognaphalium microcephalum and P. beneolens are primarily Californian. Pseudognaphalium canescens ranges eastward to Colorado, Oklahoma, and Texas and southward in Mexico as far as Chiapas, although it appears to be rare south of the state of Mexico. Pseudognaphalium thermale occurs northward into southern British Columbia, Idaho, and Montana and barely reaches Utah and Wyoming. Stebbins and Keil (1992) observed that in southern California, where all four occur, they “intergrade to such a degree with respect to charac- ters used in current keys (Ferris 1960; Munz 1959, 1968, 1974) to differentiate them — decurrent versus nondecurrent leaf bases, nature of tomentum, char- acter of capitulescence, size and shape of heads, and acute versus obtuse phyl- lary tips — that o cannot be consistently be separated from each other.” Cronquist (1955) earlier treated Gnaphalium thermale and G. microcephalum at ie rank within a single species, but his concept of the latter see y included at least G. beneolens. In contrast, other botanists have regarded ea of the ae as a separate species (ie., Ferris 1960; Munz 1959, 1968), and | +e find that consistent separation is possible. Local and regional floras in California have treated Pseudognaphalium microcephalum and P. beneolens as separate species where they occur together: San Luis Obispo Co. (Hoover 1970), “southern California” (Munz 1974), the Santa Monica Mountains of Los Angeles Co. (Raven et al. 1986), and Santa Cruz Island of Santa Barbara Co. Junak et al. 1995). These two species are sympatric in the southern coastal counties (Figs. 2 and 3) and I have not seen collections that might be indicative of hybridization. = — ae Pseudognaphalium beneolens and P thermale are similar in their aromatic character and their relatively narrow, decurrent, concolored leaves prominently sessile-glandular beneath the other indument. The twoare largely allopatric in geographic range (Figs. 3 and 4) and P beneolens generally occurs at lower el- evations. Ferris 1960, p. +70) noted that “some plants of the lower western slopes of the Sierra Nevada are intermediate between [Gnaphalium beneolens| and G. thermale, having the opaque phyllary-tips and larger heads of the former and the growth habit and shorter inflorescence of the latter.” My observation is that these plants (e.g., Tuolumne, Mariposa, and Fresno cos.), which I identify as P beneolens, occur at higher elevations than coastal populations and they tend to have slightly wider basal leaves, but otherwise they fit within the species. In any case, even if they are somewhat differentiated from coastal populations, it is not clear that genes from P.thermale are the influencing factor. Further, there does not appear to be evidence of intergradation even in Mariposa and Fresno counties, where the two species apparently occur in relatively close proximity. Pseudognaphalium beneolens was reported for Oregon by Ferris (1960) and NESOM, 785 e| nb eh ae | rt =a a + + + 4 | i | ar) ell mee Je! SST 5 ) pik : a | i @ } 5 Pseudognaphalium Pe ee eee oe ie 4 escens ee hone | a A WW tribution of Pseud hali in the United States. The distribution continues into south-central Fic. 1 Mexico (see text). Munz (1974) and the current study records its occurrence in immediately adja- cent California (Fig. 3), considerably north of the main range of the species. Both records are documented here: Catirornia. Del Norte Co.: Hazelview Summit on Crescent City-Grant’s Pass road, 2800 ft, 24 Jul 1928, Kildale 5813 (LL); OREGON. Josephine Co: Illinois River between McGuire Gulch and Oak Flat, 1300 ft, 8 Aug 1929, Kildale 8881 (LL). Pseudognaphalium canescens and P microcephalum are mostly odorless and have relatively broader, non-decurrent, and weakly bicolored leaves with- out glands or with minute and weakly developed glands. These two species differ in geography and ecology and it is unlikely that they hybridize. Variation in Pseudognaphalium canescens Variability in phyllary morphology apparently was the primary basis for LM. Johnston’s descriptions of the several different taxa now treated within Pseudognaphalium canescens. Plants of P. canescens cl istically produce strongly white-opaque phyllaries with a filiform keel and apiculum, but in the southern portion of its range JJalisco southeastward) and scattered localities elsewhere, the phyllaries may be more hyaline and lack a pronounced keel and apiculum. Similar plants from New Mexico were the basis for Johnston's recog- nition of Gnaphalium viridulum, but these are not distinct on the basis of other 786 BRIT.ORG/SIDA 21(2) Pseudognaphalium microcephalum characters. The widely disjunct poy s of P. canescens in Texas, Oklahoma, and Colorado (Fig. 1) do not appear to be significantly differentiated from those in areas where the species is more continuously distributed. NESOM 787 Pseudognaphalium C) beneolens | ® Fic. 3. Distribution of Pseudognaphalium beneolens. BRIT.ORG/SIDA 21(2) 788 Pseudognaphaliu m thermale | A\AITHTY AIIBC) Winch? INCIT\ } ‘y ? NESOM, 789 ACKNOWLEDGMENTS David Giblin (WTU) provided data and a digital image of the type of Gnaphalium johnstonii, Scott Sundberg (OSU) helped in accessing the Oregon State University database of vascular plants, and Walter Kittredge (GH) pro- vided information on the syntypes and typification of Gnaphalium wrightii. Observations and distribution maps are based on specimens from ARIZ, GH, HSC, MO, TEX/LL, and SMU/BRIT, except where noted for Fig. 4. Review com- ments from David Keil are much appreciated. REFERENCES Cronauist, A. 1955. Vascular plants of the Pacific Northwest, Part 5: Compositae. Univ. of Washington Press, Seattle. Ferris, RM. 1960. Compositae. In: L. Abrams and R:S. Ferris. Illustated flora of the Pacific states, Vol.4. Stanford Univ. Presss, Stanford, California. Pp. 98-613. Hoover, R.F.1970.The vascular plants of San Luis Obispo County, California. Univ. of Califor- nia Press, Berkeley. Jepson, W.L. 1934. The overland journey of Thomas Nuttall. Madrono 2:143-147. JOHNSTON, |.M.1924. Taxonomic records concerning American spermatophytes. Contr. Gray Herb. 70:61-92. JUNAK, S., T. Ayers, R. Scott, D. WitKeN, and D. Youne. 1995. A flora of Santa Cruz Island. Santa Barbara Botanic Garden in collaboration with the Calif. Native Plant Society. Munz, PA. 1959. A California flora. Univ. California Press, Berkeley. Munz, PA. 1968. Supplement to a California flora. Univ. California Press, Berkeley. Munz, PA. 1974.A flora of southern California. Univ. California Press, Berkeley. Raven, PH., H.J. THomeson, and B.A. Pricce. 1986. Flora of the Santa Monica Mountains, Cali- fornia (ed. 2). Southern California Botanists Special Publ. No. 2. Steseins, G.L. and D.J. Kei. 1992. New combinations in Gnaphalium (Asteraceae: Inuleae). Novon 2:437. Steseins, G.L. 1993. Pseudognaphalium.|n:J.C. Hickman, ed. The Jepson manual: higher plants of California. Univ. California Press, Berkeley. Pp. 270-271. > 790 BRIT.ORG/SIDA 21(2) BOOK NOTICES/BOOKS RECEIVED Timber Press book on Lavender Tim Upson and Susyn ANpDREWS. 2004. The Genus Lavandula. (ISBN 0-88192-642- 6, hbk.) Timber Press Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204- 3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503- 227-2878, 1-800-327-5680, 503-227-3070 fax). $49.95, 442 pp., 30 color plates by Georita Harriott, Christabel King, and Joanna Langhorne), 4 b/w pho- tos, 126 color photos, 28 maps, 41 line drawings, 12 tables, 7 3/8" x 95/8". According to the authors, “The Genus Lavandula is the most comprehensive and authoritative ac- ount of lavenders to be published to date.” After spending some time thumbing through it, | am inclined to agree. Besides describing some 39 species and their hybrids, it encompasses the taxonomy, distribution, oo and nines with additional chapters on propagation, products, chemistry, essential oils, and p es, ina relatively small book. Besides making a lovely coffee table book, it is ore useful when ane species, with the 31 full page color Ea 45 line drawings and numerous maps. A part of the book that I found unique was the 50 on the History of Lavender Cultivation; an A-Z aun of all the people, places and ne a? involved in the world of Lavender. Among the nine appendices, some that | found to be useful were a highly detailed one on how to photograph oa giving advice on lighting, camera, film and backgrounds; Plant Breeders’ Rights and Plant Patents; and National Collections and Nurseries, which lists up-to- date information including, telephone and email addresses. With all that’s included in this book, | think it is well worth the $49.95 (U.S) it costs.— diol McElfish, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.: Editor’s note.—Appendix | list new taxa a ainidecd of which there are 18 including two new species (Lavandula samhanensis and Lavandula qishnensis). SIDA 21(2): 790. 2004 A TAXONOMIC REVIEW OF THE ERIOPHORUM RUSSEOLUM-—E. SCHEUCHZERI COMPLEX (CYPERACEAE) IN NORTH AMERICA icone Cayouette roe | lth Ri DI Natior val P. f Agr ie se ade ee a Bitildin — re menta ae Ottawa, Ontario, CANADA, K1A 0C6 email: cayouettej/@agr.gc.ca ABSTRACT The taxonomy of Eriophorum russeolum— B: serene, oho in North America is reviewed, including the northwestern North A E. chamissonis. A key to the taxa includ- new characters of the medial fertile scales and achenes i is Glee The new nothosubspecies Eriophorum Xmedium subsp. album J. Cayouette is described; it represents the hybrid between E. russeolum subsp. leiocarpum and E. scheuchzeri subsp. scheuchzeri from northeastern Canada (northern Quebec and Nunavut), based on es characters observed in herbarium speci- mens. The typical hybrid subspecies, E. xmedium subsp. medium (E. russeolum subsp. russeolum x . scheuchzeri subsp. scheuchzeri) is also characterized and ann to the flora of North America ae and ree = ne two hybrid subs pared to their parental subspecies. An- other related t chzeri subsp. arcticum, poorly oe in North America, is compared to the typical peu and its range is established in northeastern North America. Lectotypes of E. russeolum Fries and E. russeolum var. majus Sommier are here designated. RESUME Une révision taxonomique du groupe des Eriophorum russeolum—E. scheuchzeri en Amérique du Nord est proposée, incluant l’espéce voisine E. chamissonis ou Bole ones de Vamenice Une oe des 1 : rr ‘ PAS es iG 4 4 -}] ¢, +] Une nouvelle notho-sous-espéce, | Eriophorum b bum J. Cayouette, est decuite représentant Vhybride entre Eriophorum russeolum subsp. arena et TE. scheuchzeri ed sc clieuchver. et ee dans le ne est du eee nord ca Québec et Nunavut), a partir d’observations de | erbier. La sous-espéce typique de hybride,| "E.xm dium subsp. medium CE. fused subsp. deta x E. scheuchzeri subsp. scheuchzeri), est également caractérisée et rapportée comme nouvelle pour Amérique du Nord (Québec et Labra- dor). Les deux sous- ae del’ bypilees sont comparées a leurs sous- ba paces ag respectives. v( arcticum peu connuen mpa 1a a la sous-espéce typique, et son aire de répartition est précisée dans le nerd: -est de l’Amérique. Les lectotypes de ’E. russeolum Fries et de VE. russeolum var. majus Sommier sont désignés. Key Worps: Eriophorum, Cyperaceae, hybrid, Canada, Quebec, Labrador, Nunavut, arctic In the course of the preparation of the collaborative project “la Flore du Québec- Labrador nordique,” spearheaded by “le Centre d’Etudes nordiques” and “Herbier Louis-Marie” (QFA), both of Laval University, Quebec City, Canada, SIDA 21(2): 791-814. 2004 792 BRIT.ORG/SIDA 21(2 — the genus Eriophorum (Cyperaceae) was found to be very difficult taxonomi- cally, especially within the E. russeolum Fries in Hartman and E. scheuchzeri Hoppe groups of rhizomatous species with solitary spikelets, in subgenus Eriophorum. Approximately 700 specimens from the region of the projected flora were examined, covering the Quebec-Labrador peninsula north of 54° N, including adjacent islands in Hudson Bay and Ungava Bay that belong admin- istratively to Canada’s Nunavut Territory. Specimens from outside the region were also studied. Material was examined from the following herbaria: CAN, DAO, FI, GH, MICH, MINN, MT, MTMG, QFA, QFBE, QUE, SFS, TRTE, UPS, and WIS (abbreviations according to Holmgren et al. 1990). Marcel Raymond (1954) was one of the first to circumscribe the taxa re- lated to Eriophorum chamissonis C.A. Meyer and E. russeolum. He recognized five different species complexes worldwide based on the color and the shape of the spikelets, the pubescence of the achenes, the color pattern of the medial fertile scales, and in using some distributional characteristics. He also proposed three varieties of E. russeolum based on the color of the bristles and the pubes- cence of the achenes, two colored-bristle forms for E. chamissonis, and three colored-bristle forms for E. medium Andersson which he presumed to repre- sent the hybrid between E. russeolumand E. scheuchzeri. His key to these taxa was very brief and does not work well. He believed that both E. chamissonis and E. russeolum occurred in eastern and western North America. In north- eastern North America, he separated EF. russeolum into two varieties based on achene pubescence. Specimens determined by Raymond as E. chamissonis in northeastern North America did not match the characters he gave in his publi- cation for that species (first proximal scale length, stem width). It is almost impossible to clearly separate E. chamissonis from E. russeolum in northeastern North America based on his work. Later, Novoselova (1993) proposed an alternative circumscription of these taxa, and made other changes in the E. scheuchzeri group (Novoselova 1994a, 1994b). She believed that in subgenus Eriophorum two rhizomatous species with orange-brown spikelets occur in North America, E. chamissonis and E. russeolum subsp. russeolum. She restricted the range of E. chamissonisto North America (both western and eastern North America), considering the Russian material to be referable to other species of the group. She also considered E. russeolum subsp. russeolum to be an amphi-Atlantic taxon. Ball and Wujek (2002) included E. russeolum within E. chamissonis, and considered E. scheuchzeri to be monotypic. This investigation supports the circumscriptional concepts of Novoselova (1993, 1994a, 1994b), with the addition of a new taxon to the E. russeolum—E. scheuchzeri group. Moreover, based on examination of North American rhi- zomatous taxa of subgenus Eriophorum with orange-brown spikelets, it ap- pears that E. chamissonis is present only in Alaska and British Columbia, while CAYOUETTE, TAAVUINUINIT UE 793 Eriophorum russeolum subsp. russeolum occurs only in the northeastern North America. Examination of material from northeastern North America, revealed that rhizomatous specimens with orange-brown spikelets are highly variable. The variation includes typical and atypical E. russeolum subsp. russeolum, and what is known in western Russia and northwestern Europe as E. xmedium, the hy- brid between E. russeolum subsp. russeolum and E. scheuchzeri subsp. scheuchzeri (Novoselova 1993, 1994a), a taxon not previously reported in North America. A few specimens seem to be atypical E. russeolum subsp. russeolum, or perhaps backcrosses of E. xmedium with E. russeolum subsp. russeolum. Variation is even greater among rhizomatous taxa with white spikelets in the Eriophorum russeolum and E. scheuchzeri groups. Eriophorum russeolum subsp. leiocarpum Novoselova can be considered the white phase of E. russeolum. Two subspecies can be discerned in the E. scheuchzeri group, the common boreal and arctic-alpine subsp. scheuchzeri, and the recently recog- nized high-Arctic subsp. arcticum Novoselova. Novoselova (1994b) reported the latter subspecies for Arctic North America, without adequately establishing its distribution in northeastern North America. It should be noted that despite the high variability of E. russeolum subsp. leiocarpum (Novoselova 1993), some specimens with white spikelets match neither the description of that subspe- cies nor that of any subspecies in the E. scheuchzeri group. Since these speci- mens have many intermediate characters between E. russeolum subsp. leiocarpum and E. scheuchzeri subsp. scheuchzeri, | conclude that they repre- sent a hybrid of these two subspecies. To account for this hybrid, I propose and describe below a new nothosubspecies of E. x medium. With the naming and description of a new taxon, the discovery of a sec- ond as new to North America (E. xmedium subsp. medium), and the realiza- tion that a third (E. scheuchzeri subsp. arcticum), while recognized as present, is poorly known on the continent, it has been necessary to identify new charac- ters that help to distinguish these taxa from the commoner E. russeolum subsp. russeolum, subsp. leiocarpum and E. scheuchzeri subsp. scheuchzeri. The char- acters are based on various features of the medial fertile scales (size, shape, apex, color pattern), on features of the achenes (size, shape, surface, beak shape and size), and on the lengths of hypogynous perianth bristles and stigmatic branches. Complete descriptions of all six taxa are provided below. The main differences among these taxa are illustrated in Figs. 1-20, summarized in Tables land 2, and employed in the key to the taxa. As the northwestern North American E. chamissonis has been previously reported in northeastern North America, and confused with E. russeolum, a complete description is also provided and some of its main differences are in- cluded in Table 1 and employed in the key. General ranges are given for the widespread E. russeolum subsp. russeolum Spikelets Proximal fertile scales Medial fertile scales BRIT.ORG/SIDA 21(2) Achenes CAYOUETTE, TAXONOMY OF 795 and E. scheuchzeri subsp. scheuchzeri, while selected specimens or paratypes are cited for the other five taxa. Eriophorum chamissonis Eriophorum chamissonis C.A. Meyer in Ledeb. Fl. Alt. ane. ae Type: U.S.A. ALASKA: (LECTOTYPE: “Legit Eschsch. in Unalaska,” by N Herbs perennial with short to elongate rhizomes. coe shoots 1-3, 21-61 cm high, leaf margins glabrous. Stems erect, glabrous, terete in cross section, 27-95 cm high, 1.0-2.2(-2.5) mm in diameter below the inflorescence. Leaves basal and cauline 2-5. Proximal sheaths pale brown, pale reddish-brown to red- dish-brown, with orange-brown spots on distal membranous parts, ligules ob- tuse or truncate. Highest distal sheath situated above or below the medial part of the stem, 2.3-4.2 mm wide, without blades or with reduced blades. Blades of proximal sheaths flat to slightly cymbiform, 25-430 x 14-L5 mm, glabrous in distal parts, the apex rounded. Blades of distal sheaths 0 or 0.9-3.5 x 0.4-L0 mm. Spikelets solitary, typically spherical at maturity, sometimes widely obo- void, 2-6 x 2-7 cm, with 100-200 florets. Proximal scales 3-7, without florets. First proximal scale olive to pale gray-olive, becoming pale beige with reddish- brown dots in marginal and distal parts, triangular-lanceolate to elliptic, 12- 23(-30) x 2.6-6.5 mm, with 4-10 pale orange, pale brown to blackish nerves converging below the ae acute, acuminate or inely short-awned. Medial fer- tile scales with a non demarcated, or demarcated short to slightly extended proximal part, 1.0-2.5 mm long, averaging 19-42% of total scale length, whit- ish, greenish, pale beige-brown to pale orange-brown, with small reddish-brown longitudinal spots, with medial part frequently grayish or forming a + extended blackish triangle, with narrow-hyaline or whitish marginal and distal parts, very often covered with small dark reddish longitudinal spots, mostly lanceolate, sometimes elliptic, 4.2-6.3 x 1.2-1.5(-L8) mm, the widest part near the middle Fics. 1-20. Spikelets, proxi 1 medial fertil ( | f six Erioph taxa. 1-3.Friophorum eolum subsp. russeolum. 1. Spikelet (Gauthier & Roy 83-45 QFA). 2a. Mfs. Payee ge LM-148 QFA). 2b. Mfs. Ds 90-3090 QUE). 3a. Achene (Gauthier & Roy 83-47 QFA). 3b. Achene eae ie 4-6. Eriophorum medium. 4, Spikelet (Blondeau 501 QFA). 5a. Mfs. mono bibubo (Blond ne ee conan 6a—b. Achenes (Robinson 74b GH).7-10 i subsp. scheuchzeri.7. Spike- es anee oe ee Mian oa: aoe et al 7991 DAO). 8b. Pfs, eelondeat ove a) ae me, alae ante (Deshaye 80 QFA).10a.Achene (Calder 2308 D 87313 an N- 14. Eriophorum one subsp. arcticum. 11. Spikelet (Blondeau 85060 “ae 12. Pfs. (Dutilly et al 87516 SFS). 13a. Mfs. (Forbes 70 DAO). 13b. Mfs. (Dutilly et al 87562 QFA).14a—b. Achenes (Dutilly et al 87516 SFS). 15- 17. Eriophorum x medium subsp. album. 15. Spikelet (Blondeau 84435 QFA). 16a. Mfs. (Polunin 2599 CAN). 16b. Mfs. (Garneau 91-405M QFA). 16. Mfs. (Cayouette J82-212 DAO). 17a. Achene (Soper s.n. 1925 CAN—Holotype). 17b Achene (Cayouette J82-212 DAO). 18-20. Eriophorum russeolum subsp. ea 18. Spikelet (Dutilly et al 20806 QFA). 19a. s. (Brisson & Forest 20502 QUE). 19b. Mfs. (Dignard 98-159 QUE). 20a. Achene (Deshaye 90-1593 QUE). 20b. Achene (Dignard 98-159 QUE). Scale bar for Fig. 1 and all the spikelets equals 2 cm; for Fig. 2 and all fertile scales equals 3 mm; for Fig.3 and all achenes equals 1 mm. Taste 1. Selected morphological differences of Eriophorum x medium hybrids and their respective parents, also including £. chamissonis. Characters (1) Characters (2) E.chamissonis E.russeolum subsp. £. xmediumsubsp. —_E.scheuchzerisubsp. £.xmediumsubsp. _ E.russeolum subsp. russeolum edium scheuchzeri album leiocarpum Spikelet shape (mature) — spherical or obovoid to hemispherical, hemispherical hemispherical, ellipsoid or widely obovoid — ellipsoid ovoid or ellipsoi ovoid or obovoid obovoid color of bristles pale beige-brown orange-brown pale orange- white or cream white or dull white or dull to dark orange- — brown to hit white white brown orange-brown length (cm) 2.0-6.0 1.5-4.0 1.4-4.0 1.0-3.0 1.6-4.0 2.0-4.0 First proximal length (mm) 12-23(-30) 7-14(-18) 7-11(-17) 5-12 7.1-11.5 (5.7-)7-16.5 scale Medial fertile proximal part 1.0-2.5 1.5-3.5 0.9-1.5 0.1-0.9 0.5-1.7 0.8-3.2 scales length (mm) % of proximal 19-42 30-49 17-27(-37) 2-25 9-34 (11-)18-57 part to maximum length proximal part whitish, greenish, orange-brown whitish, pale whitish or whitish, pale orange-brown, color pattern pale beige-brown to beige-brown — green or pale pale green green or pale pale brown, pale beige beige reen or whitish color pattern of — grayish, or dark wide medial blackish with dark grey or dark medial usually dark wide other parts blackish medial triangle with usuall blackish triangle with medial triangle triangle with wide-hyaline narrow-hyaline — with dark or reduced hyaline — with wide or narrow-hyaline margins margins narrow hyaline margins reduced hyaline margins, with margins argins small dark longitudinal spots greatest width 1.2-1.5(-1.8) 1.3-2.2 0.7-1.3 0.4-1.0 0.6-1.1 (0.8-)1.0-2.4 (mm) 962 (Z)L2 vaIs/o¥O'LINa Taste 1. continued Characters (1) Characters (2) E. chamissonis E.russeolum subsp. £.xmediumsubsp. E.scheuchzerisubsp. £.xmediumsubsp. _ E. russeolum subsp. russeolum medium scheuchzeri album leiocarpum 7 position of the below or near above or near below middle below middle below middle above, below or greatest width middle middle or near base near middle apex acute acute, rarely acuminate narrowly acuminate acute obtuse or acuminate acuminate width (mm) at 0.3-0.5 0.2-0.5(-0.9) 0.1-0.3 0.05-0.1(-0.2) 0.15-0.3(-0.4) 0.25-0.6 0.2mm below the apex Hypogynous length (mm) 25-40 25-32 15-20 10-25 (10-)22-32 12-30 bristles Anther length (mm) 0.7-1.6 1.5-3.] 0.8-1.5(-1.8) 0.35-0.8 0.9-1.6 (1.3-)1.5-3.1 Stigmatic len (mm) 1.5-2.6 1.2-1 0.7-2.0 0.5-1.3 1.0-2.2 3-3.2 branches Achene shape ellipsoid or obovoid or frequently narrowly narrowly obovoid obovoid or slightly ellipsoid narrowly obovoid or narrowly ellipsoid obovoid obovoid ellipsoid 7 width (mm) 0.8-1.1 0.75-1.3 0.6-0.9(-1.1) 0.5-0.85 .6-0.9 6-1.2 a surface glabrous or glabrous or glabrous glabrous glabrous glabrous or (pubescence) scabrous scabrous scabrous i beak width at 0.1-0.2 0.1-0.25 0.1-0.15 0.05-0.1 0.1 0.1-0.2 base (mm) beak shape straight, rarely straight, rarely straight or more often straight or more often oblique oblique oblique oblique than oblique straight than straight oblique ifn LIAQNOYYI “JLLINOAV) L6L 798 BRIT.ORG/SIDA 21(2) Taste 2. Selected pholog the two subspecies of Eriophorum scheuchzeri. Characters E. scheuchzeri subsp. scheuchzeri E. scheuchzeri subsp. arcticum Spikelet shape Proximal fertile scale pattern Medial fertile scale shape Medial fertile scale width at the middle (mm) Medial fertile scale apex Medial fertile scale width (mm) ca.0.2 mm below the apex Anther length (mm) hemispherical missing hyaline margins or reduced margins well- separated from darker body narrowly lanceolate 0.3-0.7(-0.9) narrowly acuminate mostly 0.1 0.35-0.8 spherical conspicuous hyaline margins passing to inner various tones of gray, to darker medial and basal parts lanceolate (0.5—)0.7—1.4(-1.6) acuminate mostly 0.2 0.6—1.0 Achene color brown to olive-brown orange-brown to dark red- dish-brown dul Achene surface glossy or below, with 1-3 incomplete nerves, acute, 0.3-0.5 mm wide at 0.2 mm below the apex. Perianth of 50-80 hypogynous bristles, pale beige-brown (or whitish inf. turneri Raymond), 25-40 mm long. Stamens with filaments about as wide as perianth bristles, anthers pale yellow, 0.7-1.6 mm long. Styles with 3(-4) stig- matic branches closed or sometimes spreading at maturity, branches 1.5-2.6 mm long. Achenes orange-brown, ellipsoid or slightly obovoid, trigonous to com- pressed-trigonous, glabrous or scabrous in the distal part, mostly dull, 1.9-2.6 x 0.8-1.1 mm, base cuneate, apex obtuse, with a straight beak, rarely oblique, slightly conical, 0.2-0.5 mm long, 0.1-0.2 mm wide at base. Distribution and habitat.—Eriophorum chamissonis is restricted to north- western North America, in Alaska and British Columbia. It is found in various kinds of sphagnum and minerotrophic bogs, marshy and beaver meadows, shal- low ponds, muskeg, and heat tundra. Discussion.—This taxon has a long history of various interpretations, of which the most important are those of Raymond (1954) and Novoselova (1993). As many morphological features of E. chamissonis overlap with those of E. russeolum, E. chamissonis has been frequently mistakenly reported in eastern North America (Raymond 1954; Novoselova 1993) and in eastern Russia. Mate- rial from Russia has been reassigned by Novoselova to other taxa such as E. mandshuricum Meinsch. subsp. mandshuricum or subsp. sibiricum Novoselova. Eriophorum chamissonis is best differentiated from E. russeolum subsp. russeolum by the following characters (see also Table 1): mostly spherical spike- lets with pale beige-brown bristles (typically obovoid spikelets with red-brown — CAYOUETTE, TAAUNUMT UF 799 to dark orange-brown bristles in E. russeolum), anthers 0.7-1.6 mm (1.5-3.1 mm in E. russeolum), and various color pattern and shape of medial fertile scales: grayish to blackish middle and distal parts, sometimes with a more defined blackish triangle, with usually narrow-hyaline or whitish marginal and distal parts, very often covered with small dark reddish longitudinal spots (see Novoselova 1993, Fig. la), mostly lanceolate, the largest width below or near the middle (usually dark wide medial triangle with wide-hyaline margins, mostly without dark longitudinal spots, typically obovate, sometimes lanceolate or elliptic, the largest width above or near the middle in E. russeolum). There are some individuals in northeastern North America with occasional pale orange-brown spikelets, shorter anthers (0.9-1.9 mm), darker, elliptic or lanceolate medial fertile scales with the largest width sometimes below the middle. At first glance, they could be considered within the variation of E. chamissonis, but they differ by two main characters. In E. chamissonis the first proximal sterile scale is 12-23(-30) mm long and the stem diameter below the inflorescence is 1.0-2.2 mm, whereas the odd northeastern material has the corresponding measurements of 8-11 mm long and 0.6-1.2 mm in diameter re- spectively. Those specimens are considered in this paper to be atypical E. russeolum subsp. russeolumor backcrosses of E.xmedium subsp. medium with E. russeolum subsp. russeolum. See the discussion below under E. russeolum subsp. russeolum. In addition, micromorphological differences in the achene surface of E. chamissonisand E. russeolum have been pointed out by Tucker and Miller 990). They also consider E. chamissonis to be a western North American species. Selected specimens: CANADA. British Columbia: Hart Highway N of Prince George, 8 mi N of Ft. cLeod, 3 Aug 1954, J.A. Calder et al. 13949 (DAO), near Hilliers between Parksville and Alberni, 49°16'N-124°46'W, 13 Jun 1961, J.A. Calder & K.T. MacKay 30390 (DAO); near Kispiox River, about 12 mi NNW of Kispiox, N of Hazelton, 19 Aug 1954, J.A. Calder et al. 14728 (DAO), along Kitsumkalum Lake road, about 8 mi N of Terrace, 22 Aug 1954, J.A. Calder et al. 14907 (DAO), Hope Island, off N end of Vancouver Island, between Roller Bay and Mexicana Point, 5 Jul 1961, JA. ial & K. - Macks) 31290 (DAO); Lake Beautiful, 30 Jul 1935, PP Henson s.n.(DAO),; between P G rapids, 18 Jul 1954, J.A. Calder et al. 13216 (DAO); Queen Charl Islands (QCD) Guhanticend about 3/4 mi SW of Jalun Lake and 9 mi W of head of Naden Harbour, | Jul 1964, J.A. Calder & R.L. Taylor 35663 (DAO); QCI, Graham Island, Masset Inlet, Mamin River delta at Juskatla, 15 Jun 1957, RL. Tay- lor 124 (DAO); QCI, Graham Island, about 8 mi on road from Port Clements to Tlell, 9 Jun 1957, J.A. Calder et al. 21358 (DAO); QCI, Graham Island, 4 mi W of Tlell on road to Port Clements, 26 Jun 1964, JA. Calder & R.L. Taylor 35457 (DAO); QCI, Graham Island, 2-3 mi E of Tow Hill, 20 Jul 1957, J.A. Calder et al. 22756 (DAO); QCI, Graham Island, about 1 1/2-2 mi W of Tow Hill and E of Masset, 19 . 1957 i Calder et al. 22726 (DAO); QCI, Moresby pees Cumshewa Inlet, a few mi N of Moresby Camp, 29 Jun 1957, J.A. Calder et al. 21938 (DAO), Seeley Lake, S of Hazelton, 24 Jun 1949, R. =e 191 (DAO); Sicamous-Revelstoke a E end of Victor Lake, 7 Jun 1953, J.A. Calder & D.B.O. Savile 8796 (DAO); Southern Cariboo Mountains, Wells Gray Provincial Park, E side of Battle Mt., L.5 mi NE of Stevens Lake, 25 Jul 1961, L.& T. Ahti 7092 (DAO); 1 mi NW of Trout Lake on road from Beaton to Kaslo, 10 Jun 1954, J.A. Calder & D.B. O. Savile 8964 (DAO); about 1 mi N of Trout Lake, 50°39'N-117°34'W, 25 Jun 1962, ].A. Calder & K. W. Spicer 33647 (DAO). UNITED STATES. Alaska: Attu BRIT.ORG/SIDA 21(2) Island, Peaceful Valley, near Navy Town, 52°50'N-173°LI'W, 18 Aug 1983, B.F Friedman (83-59) & ].A. Michaelson (DAO, 2 collections), Eagle River, near Juneau, 28 Jun 1940, |.P Anderson 6201 (DAO, 2 — collections). Eriophorum russeolum—Eriophorum scheuchzeri complex A) Taxa with orange-brown spikelets (Figs. 1-6) 1. Eriophorum russeolum Fries in Hartman subsp. russeolum, Handb. Scand. Fl. ed. 3:13. 1838. Type: SWEDEN. TORNE LAPPM lo, L.L. Laestadius s.n. (LEC- TOTYPE, designated here: “Lappon. Tornens. Karesuando, L.L. Laestadius s.n. Herbarium normale Fasc. 3,no 67” UPS V-108936!, DAO (photograph)! The sheet selected bears three speci- mens, the middle one clearly rhizomatous. E eee russeolum var. — Fl. Ob Infer. 103.1896. Type: RUSSIA. WESTERN SIBE- Ob River, E. a Sn. ae TOTYPE, designated here: “Siberia, ad {lumen vt cis laevae | loso, sphagnoso, +-VIII-1880, E. Sommier s.n.” FI, DAO (pho- achenes and medial fertile scales terra firma Muzhi ph)! The four ee on the sheet | have scabrous togray with wide-hyaline margins. lerbs perennial with short to elongate rhizomes. Vegetative shoots 1-3, 19-32 cm high, leaf margins mostly glabrous. Stems erect, glabrous, mostly terete in cross section, 15-55 cm high, 0.7-1.6 mm in diameter below the inflorescence Leaves basal and cauline 2-6. Proximal sheaths brown, pale brown to gray- brown, with orange-brown spots on distal membranous parts, ligules obtuse Highest distal sheath situated above, below or near the medial part of the stem, 1.6-3.5mm wide, with reduced blades. Blades of proximal sheaths flat to slightly cymbiform, 30-240 x 0.9-L6 mm, glabrous or rarely scabrous in distal parts the apex obtuse to rounded. Blades of distal sheaths 2-18 x 0.9-1.2 mm. Spike- lets solitary, typically obovoid at maturity (Fig. 1), but often ellipsoid, 1.5-4.0 » 1.5-5.0 cm, with 100-150 florets. Proximal scales 3-7, without florets. First proxi- mal scale olive-brown to dark olive-green, becoming pale beige to hyaline in distal parts, triangular-lanceolate, elliptic to ovate, 7-14(-18) x 3.4-4.4 mm, with 3-8 orange-brown nerves converging below the apex, acute, acuminate or rarely short-awned. Medial fertile scales with well demarcated and extended proxi- mal part (Fig. 2), 15-3.5 mm long, averaging 30-49% of total scale length, or- ange-brown to beige-brown, with medial part forming a + extended dark tri- angle, with wide-hyaline or whitish marginal and distal parts, typically obovate (Fig. 2a), sometimes lanceolate or elliptic, 4.0-7.5 x 1.3-2.2 mm, the widest part near the middle or above, rarely below, with | incomplete nerve, obtuse, acute or acuminate, 0.2-0.5(-0.9) mm wide at 0.2 mm below the apex. Perianth of 50- 70 hypogynous bristles, pale to dark orange-brown or red-brown (Fig. 1), 25-32 mm long. Stamens with filaments about as wide as perianth bristles, anthers yellow, 15-31 mm long. Styles with 3 stigmatic branches open to spreading at maturity, branches 1.2-1.8 mm long. Achenes pale olive-green, gray-olive, dark olive-green or brownish (Fig. 3), obovoid or ellipsoid, trigonous to compressed trigonous, glabrous (Fig. 3b) or scabrous (Fig. 3a) in the distal part, lustrous or —_ CAYOUETTE, TAAYUNYUINUT UF 801 slightly lustrous, 2.05-2.70 x 0.75-1.30 mm, base cuneate, apex obtuse to slightly rounded, with a straight beak (Fig. 3a), rarely oblique (Fig. 3b), conical, 0.2-0.6 mim long, 0.1-0.25 mm wide at base. Figs. 1-3. Distribution and habitat—This typical subspecies is amphi-Atlantic, rang- ing from central Russia westward to Northern Europe and eastern North America, from Newfoundland, Labrador, the Maritime provinces, Quebec, the islands of Nunavut in James Bay to Ontario. Its ecological affinities in North America are boreal and its range does not extend far beyond the treeline. It is found mostly in fens or minerotrophic bogs of various kinds, at the edge of pools, ponds or on lakeshores, the typical habitats for all six taxa in the E. russeolum and E. scheuchzeri groups. Discussion.—This is the most common taxon with orange-brown spikelets in northeastern North America. Its distinguishing features are the typically obovoid spikelets and their orange-brown to dark orange-brown color (Fig. 1). The characters of the medial fertile scales are important (Table 1, Fig. 2):a unique color pattern consisting of a long demarcated proximal part, usually orange- brown, that covers up to half the length of the scale (Fig. 2a), a central zone represented by a wide black triangle, and wide marginal and distal whitish or hyaline parts; a wide lanceolate, elliptical or obovate shape, the widest of all the taxa considered here (1.3-2.2 mm), the widest area situated above (Fig. 2a) or near the middle (Fig. 2b) of the scale, with a mostly acute apex (better indi- cated by measurements taken at 0.2 mm below the apex: 0.2-0.5(-0.9) mm) (Table 1). Anthers and achenes are the longest and largest of all orange-brown taxa. Achenes (Fig. 3) are about equally glabrous or scabrous (Table 1, Fig. 3), which isa bit different from data reported for material from Russia and north- western Europe (Berggren 1969; Novoselova 1993). Achene beaks are the long- est of all the taxa (0.2-0.6 mm) and the widest at the base (0.1-0.25 mm), they are more often straight than oblique. The achene beaks of E. xmedium subsp. medium differ in that they are shorter, narrower and more frequently oblique (Table 1, Fig, 6). Variation is also encountered ina group of specimens considered atypical because of darker and narrower medial scales, shorter anthers (mostly 13-L9 mm), more frequently glabrous achenes, and achenes with narrower beaks. When these atypical specimens are found outside the range of the hybrid E. x medium subsp. medium, they could be interpreted as expressions of the varia- tion of E. russeolum subsp. russeolum. When they occur within the range of that hybrid they could also represent backcrosses of E. xmedium subsp. me- dium with E. russeolum subsp. russcolum. Experimental and field studies will be needed to help solve the problem represented by these atypical specimens. 2. Eriophorum medium Andersson subsp. medium, Bot. Not. 1857:62. 1857. (Eriophorum russeolum subsp. russeolum x E. scheuchzeri subsp. scheuchzeri) TYPE: SWEDEN. LULE LAPPMARK: prope Quickjock, N_J Andersson $.n. (HOLO- TYPE: S, not seen). 802 BRIT.ORG/SIDA 21(2) Eriophorum Xgauthieri Boivin, Provancheria 25:43. 1992. Type: CANADA: LABRADOR, Grady and Cross Islands, 26 Jul 1933, G. Gardner 18 (HOLOTYPE: QFA!: ISOTYPE: QFA!). Herbs perennial with short to elongate rhizomes. Vegetative shoots 1-3, 8-24 cm high, leaf margins mostly glabrous. Stems erect, glabrous, mostly terete in cross section, 16-42 cm high, 0.7-1.5(-L7) mm in diameter below the inflores- cence. Leaves basal and cauline 3-7. Proximal sheaths green, olive-green, beige- brown, reddish brown to dark brown, with orange-brown spots on distal mem- branous parts, ligules obtuse. Highest distal sheath situated below the medial part of the stem, 1.8-3.4 mm wide, with reduced blades. Blades of proximal sheaths flat to slightly cymbiform, 50-190 x 0.7-1.6 mm, glabrous or rarely sca- brous in distal parts, the apex obtuse to rounded. Blades of distal sheaths 1-13 x 0.4-0.9 mm. Spikelets solitary, hemispherical (Fig. 4), sometimes ovoid or ellip- soid at maturity, 1.4-4.0 x 0.9-6.0 cm, with 100-150 florets. Proximal scales 3-5, without florets. First proximal scale dark olive-brown to blackish, becoming hyaline brown in marginal and distal parts, ovate, 7-11(-17) x 2.2-4.7 mm, with 5-10 orange-brown or pale brown nerves converging below the apex, acumi- nate. Medial fertile scales with reduced proximal part (Fig. 5), 0.9-1.5 mm long, averaging 17-27(-37) % of total scale length, whitish, pale green or pale beige, with medial and distal parts blackish (Fig. 5a), with marginal and distal parts reduced-hyaline (Figs. 5b-c), lanceolate, 3.6-7.3(-8.0) x 0.7-1.3 mm, the widest part mostly below the middle, with 1 incomplete nerve, acuminate, mostly 0.1- 0.3mm wide at 0.2 mm below the apex. Perianth of 30-50 hypogynous bristles, orange-brown to pale orange-brown (Fig. 4) or red-brown, 15-20 mm long. Sta- mens with filaments about as wide as perianth bristles, anthers yellow, 0.8- 1.5(-L8) mm long. Styles with 3 stigmatic branches mostly closed at maturity, branches 0.7-2.0 mm long. Achenes chestnut brown (Fig. 6), obovoid, mostly narrowly obovoid, rarely ellipsoid, compressed-trigonous, glabrous, slightly lustrous, 1.6-2.5 x 0.6-0.9(-1.1) mm, base cuneate. apex acute or obtuse, with a straight (Fig. 6b) or oblique beak (Fig. 6a), mostly cylindrical, 0.2-0.3 mm long, 0.1-0.15 mm wide at base. Figs. 4-6. Distribution.—Described from Scandinavian material, up until now this hybrid had been found only in north central Russia, westward to northern Scan- dinavia (Novoselova 1993, 1994a). Reports of E. medium from an almost con- tinuous range in the Russian Arctic (Tolmachev 1996) do not in every case rep- resent hybrids between E. russeolum subsp. russeolum and E. scheuchzeri, because they refer to a taxon bearing white or orange-brown spikelets. Eriophorum xmedium subsp. medium can now be added to the flora of North America based on collections in Labrador and northern Quebec (Nunavik), ranging from ca 51°N to 59°N. Previous reports of E. x medium in North America, mostly from the Northwest, did not represent hybrids between E. russeolum subsp. russeolum and E. scheuchzeri subsp. scheuchzeri. E. xmedium is a bo- real amphi-Atlantic taxon like one of its parents, E. russeolum subsp. russeolum. CAYOQUETTE 803 Discussion.—This hybrid is occasional in the sympatric range of its two parental species in eastern North America. It had not been previously detected despite the fact that many morphological characters are intermediate between those of the two parents (Table 1). The orange-brown spikelets are more often than not paler and smaller (Fig. +) than those of E. russeolum subsp. russeolum, and their shape is highly variable, more often than not hemispherical, like those of E. scheuchzeri subsp. scheuchzeri (Fig. 7). Anther lengths are intermediate (0.8-1.5 mm) as are the majority of achene characters. The main differences are illustrated by the medial fertile scales which are closer to those of E. scheuchzeri subsp. scheuchzeri in the preponderance of blackish color, in the frequent re- duction of marginal and distal hyaline parts, in the color variation and short- ness of proximal parts, in the narrow width (0.7-1.3 mm) and acuminate apex, and in the maximum width mostly being located below the middle (Table 1, Figs. 5, 9). This is in accordance with Novoselova’s observations (1993) of E. xmedium in Russia and northwestern Europe. Some individuals from North America have medial scales with more developed hyaline margins and a nar- row central blackish triangle (Fig. 5c), corresponding to material from north- ern Europe studied by Faegri (1958: Fig. 1|D-E) and reported to belong in part to E. xmedium.Lobserved th pattern in some individuals of the hybrid between E. russeolum subsp. leiocarpum and E. scheuchzeri subsp. scheuchzeri (Fig. 16c). The examination of type material of E. xgauthieri Boivin (1992), described from Labrador as the hybrid between E. chamissonisand E. scheuchzeri, shows that it is identical to E. xmedium subsp. medium. Boivin (1992) included both whitish and orange-brown taxa of EF. russeolum within E. chamissonis. The presence of well-formed achenes in many individuals seems to indicate that some specimens may have become stabilized enough to be considered ortho- species of hybrid origin, as has been hypothesized for plants in northern Europe (R. Elven, pers. comm.). Further studies are needed to confirm this hypothesis. Specimens examined. CANADA. Labrador: Belle Isle, South Point, 51°53'N-55°24'W, 25 Jul 1986, TA. Hedderson 4061 (CAN), nes Island, 17-19 Jul 1938, G. Gardner 38113 (QFA, 2 collections); Grady and Cross Islands, 26 Jul 19 G. Gardner 18 [B] (QFA, 2 collections, holotype & isotype of Eriophorum xgduthieri Boivi eee with E. scheuchzeri subsp. scheuchzeri); Indian Harbour & Fox Cove, 16 Jul 1892, C. Waghorne 32288-B (CAN); Knob Lake area, valley on Geren Hill, 23 Jul 1961, J. Sangster s.n. (MTMG); Lake on n, Northwest Bay, 54°59'N-66°41'W, 19 Jul 1953, F Harper 3630(CAN); Port Manvers, 10 Aug 1922, R. Robinson 74 (GH); Red Bay, on Strait of Belle Isle, 23 Jul 1996, M.J. Oldham 19156 (MICH). pre Nunavik: Abloviak Fjord, [59°27'N-65°10'W], 2 mi from head, 1 mi from shore, 20 Jul 1978, H. Ouellet 82 (CAN, MT, SFS); environs de Kuujjuaq, ouest de la riv. Koksoak, env. 30 km au nord de Fort-Chimo, 58°22'N-68°14'W, 17 Jul 1982, M. Blondeau 501 (Hb. Blondeau, QFA); Fort Chimo area, 58°07'N-68°23'W, 4 Aug 1948, J.A. Calder 2338 (MT); Kangiqsualujjuaq, estuaire de la riviére George, | km auNNOdu ves a 4) a 'N-65°58'05"W, 26 Jul 1984, R. Gauthier 84-161 (MICH, QFA), 84-162 (QFA); idem, em| ge, 58°42'N-65°54'W, 20 Jul 1988, M. Blondeau GR-88079 (QFA); Lac Ford, 59°13'N-70°08'W, 10 Jul 1975, H. Ouellet s.n. (CAN); Riviére Boniface, 57°45'N-76°09'W, 9 Aug 1987, A. St-Louis 104 (QFA); idem, 4 l’est de la Passe au Renard, 57°43'50"N- 76°07'20"W, 26 Jul 1991, M. Garneau 91-553-M (QFA); idem, 4 l’est ducamp, 17 Jul 1994, P Levasseur 76 804 BRIT.ORG/SIDA 21(2) (QFA); Riviére aux Feuilles, 18 km enamont du le rapide, ca 538°30'N-70°30'W, 18 Jul 1974, H. Ouellet sn. (CAN): Riviére George, environ 3 milles 4 louest du lac Indian House, ca 56°20'N-64°47'W, 29 Jul 1947, J. Rousseau 564 (DAO, MT); idem, Lac Indian House, ca 50°25'N, 30 Jul 1947, J. Rousseau 580 (DAO, MT); idem, prés de Hades Hills, ca 56°58'N, 5 Aug 1947. J. Rousseau 734 (MT). B) Taxa with white spikelets (Figs. 7-20) 3. Eriophorum russeolum Fries in Hartman subsp. leiocarpum Novoselova, Bot. Zurn. (St. Petersburg) 788) 86. 1993. Type: RUSSIA. FAR EAST: E Chukotka, in vicinus pagi Nutepelmen, vallis rivi in sinum Pyngo-pilchin influentes, lo Aug 1969, A.A. Neczaeva & LV. Plievad s.n. (HOLOTYPE: LE, not seen). Herbs perennial with short to elongate rhizomes. Vegetative shoots I-3, 15-21] cm high, leaf margins mostly glabrous. Stems erect, glabrous, mostly terete in cross section, 14-51 cm high, 0.7-1.5 mm in diameter below the inflorescence. Leaves basal and cauline 1-7. Proximal sheaths brown, pale brown, chestnut brown to dark brown, with orange-brown spots on distal membranous parts, ligules acute to obtuse. Highest distal sheath mostly situated below the medial part of the stem, 2.1-3.5 mm wide, with blades reduced or lacking. Blades of proximal sheaths flat to slightly cymbiform, 40-230 x 0.7-2.3 mm, mostly gla- brous, the apex obtuse. Blades of distal sheaths 0.2-21 x 0.2-L.1 mm, or lacking. Spikelets solitary, ellipsoid or obovoid at maturity (Fig. 18), 2.0-4.0 « 1.5-3.5¢cm, with 150 or more florets. Proximal scales 4-6, without florets. First proximal scale olive-brown, olive-green, dark gray to blackish, becoming pale beige to whitish hyaline in distal parts, lanceolate to ovate-lanceolate, (5.7-)7-16 x 3.0- 5.3 mm, with 1-5 orange-brown or blackish nerves converging below the apex, acute or acuminate. Medial fertile scales with moderate to extended proximal part (Fig. 19), 0.8-3.2 mm long, averaging (11-)18-57% of total scale length, or- ange-brown, pale brown, pale green or whitish, with the medial part forming a + extended dark triangle, with marginal and distal parts mostly wide-hyaline (Fig. 19a), obovate, lanceolate or elliptic, 3.7-8.4 « (0.8-)1.0-2.4 mm, the widest part near the middle or above, rarely below, with 1 incomplete nerve, acute, 0.25- 0.6 mm wide at 0.2 mm below the apex. Perianth of 15-50 hypogynous bristles, white to dull white (Fig. 18), 12-30 mm long. Stamens with filaments about as wide as perianth bristles, anthers yellow or dark yellow, (1.3-)1.5-3.1 mm long. Styles with 3(-4) stigmatic branches barely open at maturity, branches 1.3-3.2 mm long. Achenes pale or dark olive-green, brownish or black-brownish (Fig. 20), obovoid (Fig. 20b) or ellipsoid (Fig. 20a), trigonous to compressed-trigonous, glabrous (Fig. 20b) or scabrous (Fig. 20a) in the distal part, slightly lustrous, 2.0-2.7 x 0.6-1.2 mm, base cuneate, apex obtuse, witha beak more straight than oblique, conical, 0.2-0.5 mm long, 0.1-0.2 mm wide at base. Figs. 18-20 Distribution.—The group of E. russeolum with white spikelets has a very different range from the group with orange-brown spikelets and was found to be distinct enough to be considered a subspecies by Novoselova (1993). Its range is amphi-Beringian, discontinuous in northwestern Russia, continuous from a Y CAYOUETTE, TAAUNUMT UT 805 north central Russia eastward to Alaska, the Canadian Yukon and Northwest Territories, the islands and continental portion of Nunavut, the Prairie prov- inces as far east as Manitoba, with scattered sites in eastern North America: Ontario, Quebec, Labrador, New Brunswick and Nova Scotia, with an exten- sion in Minnesota and Wisconsin. Reports of E. chamissoniss.l. from the Rocky Mountains (Ball @ Wujek 2002) have not been searched for this study and | dont know if they could refer to E. russeolum subsp. leiocarpum or not. The present study brought collections from the northern part of Nunavik in Que- bec (ca 60°-61°N) to light for the first time. Discussion.—For a long period in the North American literature, the group of E. russeolum with white spikelets was called FE. russeolum var. albidum F. Nylander, and even E. chamissonis var. albidum (F. Nylander) Fernald or f. albidum(E Nylander) Fernald. In his protologue, Nylander (1846: 10) designated a specimen from Alaska (Kodiak Island) that has not yet been found at LE as the type of his variety. This is not in accordance with Novoselova (1993) who states that Nylander’s variety, a taxon she considers a synonym for E. xmedium [subsp. medium], was described from Scandinavian material. Even if one ac- cepts that E. russeolum var.albidum was described from within the actual range of E. russeolum subsp. leiocarpum, it is difficult to consider var. albiduma syn- onym for subsp. leiocarpum because of a major divergent character of the me- dial fertile scales in Nylander’s protologue. Nylander described var. albidum’s fertile scales as being “narrowly-lanceolate” (squamis lineari-lanceolatis), whereas subsp. leiocarpum’s are typically obovate, elliptic or lanceolate (Fig. 19). Novoselova’s name (subsp. leiocarpum) is retained here because the sub- specific level is more appropriate for separating the almost non-sympatric E. russeolum taxa, and because leiocarpum was the first name to be attributed at the subspecific level. Eriophorum russeolum subsp. leiocarpum is quite variable and some speci- mens can be considered merely as white-colored counterpart nge-brown E. russeolum. Nevertheless, the spikelets of this subspecies tend to be more of - ten ellipsoid than obovoid, the achenes more often obovoid than ellipsoid, and the achene beaks narrower on average than in subsp. russeolum (Table 1). This variation is encountered in all parts of subsp. leiocarpum’s North American range. Specimens from higher latitudes (e.g. Nunavut) tend to have more black- ish medial scales (Fig. 19b), but all other characters are within the variation of the subspecies. a 2 ane examined. CANADA. Yukon: Kluane National Park, Alder Creek, 60°18'15"N-137°21'45'W, 7 Aug 1974, R.D. Wickstrom 318.11 K-AC (DAO), Northern Richardson oe tains, 68°22'53"N- 137°07'31'W, 7 Jul 1993, V. Loewen & J. Staniforth 93-154 (DAO), Idem, 68°11'42"N-137°2527'W, 17 Jul 1993, V. Loewen & J. pee 93-252 (DAO): North , ae Park, British Mountains, Firth River delta, 69°30'N-139° a il 1988, C. Kennedy 174 ). Northwest Territories: The Enter- prise-Mackenzie River Hi oe way, mile 50, 16 Jul 1959, i ee & RJ. Reich 5392 (DAO), Porter Lake, 61°44'N, 26 Jul 1970, WJ aa 19107 (DAO), Small Tree Lake, 61°N-L05°W, 20 Jul 1961, J.S. Maini 806 BRIT.ORG/SIDA 21(2) — 478 & J.M. A. Swan (DAO); Tuktoyaktuk Peninsula, Hutchison Bay, 69°42'N-132°18'W, 29-30 Jul 1981, D.L. Allen & V. Stringer 7766 (DAO): ph by lake in front of hospital, 62°27'N-114°22'W, 8 Aug 1949, WJ. Cody & B. McCanse 3324 (DAO). Nunavut: Bylot Island, A. R. Camp, Site B-3, 73°24'N- 80°43'W, 30 Jul 1983, G. Scotter SoG hesterfield Inlet, 1/4 mi W of settlement, 63°21'N-90°42'W, 4 Aug 1950, D.B.O. Savile & CT. Watts 1277 (DAO), Rasmussen Lowlands, S of Murchison Lake, 68°05'43.8"N-92°39'22.8'W, 12 Jul 1994, V Johnston 1(DAO). Manitoba: Vicinity of Churchill, 58°46'N- 94°10'W, 14 Jul 1956, W.B. Schofield & H.A.Crum sen AN): ioe 18 Jul ee WB. Schofield & H.A. Crum 6825 (CAN), Reindeer Lake, Sawbill, 57°37'N-LO1°44'W, 4 Aug 1951, W.K.W. Baldwin 2327 (MICH), Wasagaming, South Onanole, 50°37'N- ee W, 8 Jun 1967, Wk. W. Ae n 10900 (MICH), York Factory, 22-26 Jul 1949, HJ. Scoggan 5993 (CAN); idem, 20 Jul 1949, HJ. Scoggan 5929 (CAN). Ontario: Cape Henrietta-Maria, Hudson Bay, 55°1L0'N-82°20'W, 12-18 Jul 1979, R. A. Sims 2699B (MICH); Cochrane district, 50°O00'N-83°42'W, 10 Jul 1979, [.L. i ey 10576(CAN), 5 km N of Kesagami River, 50°14'N-80°12'W, 22 Aug 1983, D.F Brunton 4595 (CAN); Hudson Bay lowlands, Attawapiskat River, 53°08'N-83°18'W, 12 Jun 1957, A.E. Porsild, W.K.W. ce H. & G. Sjérs 200060 (CAN); Thunder Bay District, Fort William, 48°24'N-89°16'W, 4 Jun 1972, W. Hartley 1509 (CAN), City of Thunder Bay, NW side of Expressway in Northwood area of Intercity fen, 5 Jul 1978, CE. Garton 18297 (DAO), Thunber Bay Districy, Little Postagoni Lake, 2 Aug 1960, C.F. eee 7938(DAO):S of oo 2 about 25 km W of Cape Henrietta- sere 55°LO'N-82°50'W, 19-20 Jul 1979, R.A. Sims 2736A Québec: Nunavik, Nouveau-Québec, Lac Elizabeth, 55°40'N- oe Aug 1977, G. Lemieux 21031 (QFA (© collections), CAN); ne CRE At 55°17'N-77°46'W, 22 Jul 1969, S. Brisson & P. Forest 20502 (QUE, QFA); idem, rive N, 31/2 mia VE du poste, Ll Aug 1970, S. Brisson & P. Forest 22380 (SFS), Territoire du Nouveau-Québec, 55°46'N-76°13'W, 26 Aug 1990, J. Deshaye 90-1593 (QUE); Riviére Chukotat, ca 2 km au N, 14 km au SSO du lac Hubert, 61°19'38"N-76°21'49"W, 8-9 Jul 2003, |-F Duchesne s.n. (QUE), Riviére Korak, 60°58'N-76°58'W, 29 Jul 1987, L. Dion K9-4 (QFBE), Riviére eee 61°26'04"N-73°50'06"W, 7 Aug 1998, N. Dignard 98-159 (QUE); Riviére Puvirnitug, nviron 4 km au SO du lac Vaillant, 61°25'30"N-73°51'35"W, 7 Aug 1998, R. ence 98-148 (QFA); ae Por ep ree aaa E. Pepage. lie (QFA, 2 collections); idem, 9 Aug 1950, E. Lepage 12638 (QFA 1] baie Hannah, 51° 33'N- i 32'W, 16 Aug 1958, A. Dutilly . E. Lepage 36739 (QFA); Gaspé Co.: Tabletop Mts., headwaters of the Magdalen River, 1000- 1050 m. alt.,9 Aug 1906 ss Fernald & J.F Collins 73180 (CAN, MT); Tabletop Mts., Mt. Auclair, ca.1200 m. alt, 10 Aug 1923, M.L. Fernald & L.B. Smith 25603 (CAN, MT). Labrador: Battle Harbour, Fie Nl, 24 Aug 1871, C. ciated ae 2 (CAN). New Brunswick: Kent Co.: near Rexton, 13 Jul 1957, EC. Smith et al. 16418 (CAN, DAO). Nova Scotia: Cumberland Co.: West Advocate, chan 1950, a Smith N). UNITED STATES. ALASKA: Nome River, | mi from the sea, 5 Jul 1947, A. Dutilly, E. Lepage & H. O'Neill 20806 Sake MINNESOTA. Becker Co.: Itasca Park, E side of road to Morison Lake, 4 Jul 1933, ).B. Moyle 727 (MINN). Beltrami Co.: 10.8 mi N of Waskish P.O., 13 Jul 1975, G.B. Ownbey 4979 (MINN). Blue Earth Co.: [1883], J. R. pawl n. ne Cass Co.: near Swamp Lake, 10 Jul 1995, J. Boe Mea (MINN). Clearwater Co.: along Co. Rt. 39, about | mi N of Rt. 113, 1] Jun 1991, V. E. Neth 91-406 (WIS). Hennepin Co.: about 1.5 mi SE oe sai 4 Aug 1992, W.R. Smith 21238 N). Hubbard Co.: Itasca Park, 10 Jul 1929, C.O. Rosendah! 5908 (MINN). Lake of the Woods Co.: a Lake area, Brown's Creek Trail, 21 Jun 1979, J.S. Boe 328 fe N); idem, SE of Mud Lake, 21 Jul 1980, PH. Glaser 1300(MINN). Morrison Co.: Camp Ripley Military Reservation, 4 Jun 1991, B. Delaney 91082 (MINN). Roseau Co.: Roseau River Peatland, about 18 mi NW of Roseau, 14 Jun nee WR. Smith 9181 (MINN). Stearns Co.: 6 mi S of St. Augusta, 21 May 1998, M.D. Lee MDL2056 (MINN). St. Louis Co.: Cruiser Lake Trail, 48°28'22"N-92°48'49"W, 4 Aug 1977, M.R. Smith 466 (MINN); ne Highway 53, S of Kabetogama, 12 Jun 1950, O. Lakela 10363 (MINN). Wright Co.: by Hwy 55 on the SE side of Maple Lake, 16 Jun 1998, M.D. Lee & D. Wovcha MDL2140 (MINN). WISCONSIN. Ashland Co.: Long Island, lake Superior, S of Madeline Island, 10 Jun 1972, R.G. Koch 7378 (WIS). Bayfield Co.: NW of Eagle Lake, 10 Jul 1996, EJ. Judziewicz 11958 (WIS). Douglas Co.: SW side of junction of Co. A and CAYOUETTE, TAAVINUIMIE Ui 807 Empire Wilderness Road, 24 Jul 1996, E,J. Judziewicz 11979 (WIS). Iron Co.: W side of old rail road grade at Sandrock, 17 Jul 1996, EJ. Judziewicz 11938 (WIS) 4. Eriophorum xmedium Andersson subsp. album J. Cayouette, subsp. nov. (E. russeolum subsp. leiocarpum x E. scheuchzeri subsp. scheuchzeri) Tyre CANADA. NUNAVUT: Baffin Island, Nettiling Lake, 66°40'N-70°W, 28 Jul 1925, J. Dewey Soper s.n. (HOLOTYPE: CAN 25686). A subspeciei typica setis albis vel subcremeis, (10-)22-32 mm longis differt. Verosimiliter hybrida inter Eropn mune musecoutin suse! cocdypum et Eriophorum scheuchzeri subsp. eee Plantae vel elongatis. Culmi erecti, laeves, teretes, 22-40 cm alti, 0.8-1.1 mm diamet b spi . Vagina superior culmi media parte inferiore inserta, raro ad me- dium, foliis 0.3-17 mm ae 0.3-0.8 mm latis, vel nullis. Spiculae unicae, fructificatione hemisphaericae, ovoideae vel ebevoidene, 16 _ cm altae, setis paNpis vel subcremeis. aie sterilis infima 7.1-11.5 mm longa.S 1 m longa, 9- 7 ’o squamae totae Tongitudinis delemitante, alpica, pallide viride vel pallide fergie cum li 6-11 mm latae, squaimae media pet inferiore maxima netudine, apice acuminato 0 15-0:3620: . mm lato ad 0.2 )22-32 ereleneae Antherae 0.9-1.6 mm longae. Stigmatum rami tres a quatuor, 1.0- 2 Ds mm longi. Achenia glabra, anguste obovoidea vel oe 0.6-0.9 15-1 mm lata, rostro recto vel curvato, praecipue cylindrico, basi 0.1 mm lato. Figs. Herbs perennial with short to elongate rhizomes. Vegetative shoots 1-2, 6-24 cm high, leaf margins mostly glabrous. Stems erect, glabrous, terete in cross section, 22-40 cm high, 0.8-L1 mm in diameter below the inflorescence. Leaves basal and cauline 2-6. Proximal sheaths beige-brown to reddish brown, with orange-brown spots on distal membranous parts, ligules acute. Highest distal sheath situated below the medial part of the stem, rarely near the middle, 2.2- 2.9mm wide, with blades reduced or lacking. Blades of proximal sheaths flat to slightly cymbiform, 120-150 x 0.8-0.9 mm, glabrous, the apex obtuse. Blades of distal sheaths 0.3-17 x 0.3-0.8 mm, or lacking. Spikelets solitary, hemispheri- cal, ovoid or obovoid at maturity (Fig. 15), 1.6-4.0 x 2.0-4.5 cm, with about 150 florets. Proximal scales 3-5, without florets. First proximal scale dark olive-green or blackish, becoming brown-hyaline or brown-beige in distal and marginal parts, ovate to ovate-lanceolate, 7.1-11.5 x 2.6-3.7 mm, with 3-5 brown to or- ange-brown nerves converging below the apex, acuminate. Medial fertile scales with a reduced proximal part (Fig. 16), 0.5-1.7 mm long, averaging 9-34% of total scale length, whitish, pale green or pale beige, with the medial part black- ish forming a narrow triangle (Fig. lob), with marginal and distal parts reduced- hyaline (Figs. 16a-b), lanceolate, 3.8-6.7 x 0.6-1.1 mm, the widest part mostly below the middle, with 1 incomplete nerve, acuminate, 0.15-0.3(-0.4) mm wide at 0.2 mm below the apex. Perianth of about 50 hypogynous bristles, white to dull white (Fig. 15), (10-)22-32 mm long. Stamens with fil tsabout as wide as perianth bristles, anthers yellow-green, 0.9-1.6 mm long. Styles with 3(-4) stigmatic branches mostly closed at maturity, branches 1.0-2.2 mm long. Achenes beige-brown to orange-brown (Fig. 17), narrowly obovoid or narrowly ellipsoid, 808 BRIT.ORG/SIDA 21(2) compressed-trigonous or slightly biconvex, glabrous, slightly lustrous, 1.9-2.5 x 0.6-0.9 mm, base cuneate, apex obtuse, with a straight (Fig. 17b) or oblique (Fig. 17a) beak, mostly cylindrical, 0.2-0.3 mm long, 0.1 mm wide at base. Figs. 15-17. Distribution.—This nothosubspecies is currently known only in northeast- ern Canada, from continental Nunavut (Chesterfield Inlet), the Nunavut part of the Arctic Archipelago (Baffin and Southampton Islands, from 63°N to 66°N), south to northern Quebec (Nunavik), in the northernmost part of the peninsula (61°N-62°N) and at treeline near Hudson Bay (57°N-58°N). Since the sympatric range of the two parental taxa covers large parts of the Nunavut and Northwest Territories, the western Canadian provinces, Alaska, and eastern to western parts psp. album is likely to be discovered in some of these —— of Russia, E. xmediumsu major areas. Nevertheless, no specimen from outside of northeastern Canada has yet been identified as this hybrid subspecies (Novoselova pers. comm.). There is a slight possibility that the taxon described from Alaska as E. russeolum var. albidum by Nylander (1846: 10) and bearing “narrowly lanceolate scales” could refer to that new nothosubspecies, but the type should be searched and examined. Discussion.—A grouping among specimens that did not fit the normal varia- tion of either E. russeolum subsp. leiocarpum or E. scheuchzeri subsp. scheuchzeri was perceived and formally given the taxonomic status of a new nothosub- species. Although different from these taxa, the specimens shared most of the characteristics of E. x medium subsp. medium except for the color of the spike- lets and the length of the stigmatic branches (Table 1). The two subspecies of E. xmedium share size, shape, apex, color pattern, and maximum width below the middle of the medial fertile scales (Figs. 5, 16); medium-sized anthers (0.9- 1.6mm), and important achene characters (shape, width, beak width and shape) (Table 1, Figs. 6, 17). The shared characters are intermediate between those of E. russeolum subsp. leiocarpum and E. scheuchzeri subsp. scheuchzeri. Even if some characters of the hybrid subspecies overlap with extremes of variation of E. russeolum subsp. leiocarpum, | prefer to consider this taxon as a hybrid be- cause many characters are intermediate between those of the two parental taxa (Table 1). A similar situation has been encountered and studied ina few Arctic Ranunculus hybrids (Cayouette et al. 1997). Some specimens cited as paratypes have been previously considered by other authors as hybrids or potential hybrids involving the whitish E. russeolum and E. scheuchzeri. Polunin so annotated several specimens from Nunavut (CAN) and later discussed the possibility of E. russeolum-E. scheuchzeri hy- bridization (Polunin 1940: 100). Boivin (1992) cited one Polunin collection of the then undescribed E. medium subsp. album from Nunavut as a paratype of E. xgauthieri, a taxon that Boivin believed to be an E. chamissonis x E. scheuchzeri combination. All these published remarks match the description of E. xmedium subsp. album. A form of E. xmedium subsp. medium with white spikelets known as f. — CAYOUETTE, TAAUNUNUT UF 809 candidum (Norman) Blomgren, has been described from Scandinavia (Hylander 1982). Although not yet known in North America, this form may be expected in the vicinity of the treeline in northern Quebec, where the two subspecies of E. russeolum coexist with EF. scheuchzeri. In the event of its discovery there, E. xmedium subsp. medium f.candidum might be mistaken for E. x medium subsp. album. To avoid any possible confusion, I have selected a type specimen for E. xmedium subsp. album from Nunavut (Baffin Island), where only E. russeolum subsp. leiocarpum is sympatric with E. scheuchzeri subsp. scheuchzeri. To date, all specimens of subsp. album collected at the treeline in northern Quebec have turned out to be similar to the type specimen selected from Baffin Island (66°N). As for the typical hybrid subspecies, some paratypes of E. x medium subsp. album have been seen with good mature achenes, indicating that subsp. album has perhaps become a stabilized orthospecies of hybrid origin in some areas. Further investigations are needed to clarify this interpretation. PararyPes. CANADA. Nunavut: cane Island, me 15 Sep 1936, N. Polunin 2599(CAN); idem, Frobisher Bay. head of Tarr Inlet, 24 Jul 1965, A. McLaren 34 (CAN); idem, Frobisher Bay, vicinity of Air Base, 18 Jul 1953, V.C. Wynne-Edwat ee AN); idem, Lake Harbour, 26-28 Jul 1936, N. Polunin 1172 (GH); oe d Inlet, 1/2 mi NW of settlement, 63°21'N-90°21'W, 15 Aug 1950, D.B.O. Savile & CT. Watts of (DAO); Southampton Island, Coral Harbour, 64°10'N-83°15'W, I Jul 1976, S. White 761093 (TRTE). ae Nunavik, Cratére du Nouveau-Québec, 61°22'18"N-74°10'30'W, 28 Jul 2000, N. Dignard & J. Gagnon 00-174 (QUE); Golfe de Richmond, 1971-1973, S. Payette et al. GR-121 A (QFA, mixed with E. ri); environs d'lvujivik, 62°24'N-77°55'W, 28 Jul 1984, M. Blondeau 84435 (Hb. Blondeau, Or “A ); lec Chavigny, entre le lac Chavigny et le lac au sud, 58°03'N-75°05'W, 29 Jul 1982, J. Cayouette ‘82-212 (DAO, QFA); riviére Boniface, 9.5 km a Pouest de la Passe du Loup, 57°45'10"N-76°20'25"W, 21 Jul 1991, M. Garneau 91-405-M (QFA); bss riviére Boniface, troncon oe A riviére a l’ouest de la Passe du Loup, 57°45'10"N-76°20'25"W, 25 Jul 1991, M. Garneau 91-496-M(Q | de la petite riviére see 61°26'N-75°15'W, 16 Jul 1985, L. Dion 1.1-4 (QFBE); territoire du Nouveau-Québec, 56E39'N- 4E51W', 5 Sep 1989, J. Deshaye FOR89-85 (QUE) 5. Eriophorum scheuchzeri mas subsp. scheuchzeri, Bot. Taschenb. 104, plate 7. 1800. Type: AUSTRIA (“.. am Tuscher Tauern”) (HOLOTYPE: W, monocots destroyed, see Holmgren et al. 1990). Herbs perennial with short to elongate rhizomes. Vegetative shoots 1-3, 5-30. cm high, leaf margins glabrous. Stems erect, glabrous, terete in cross section, 9-42 cm high, 0.6-1.5(-L8) mm in diameter below the inflorescence. Leaves basal and cauline 3-5. Proximal sheaths pale green at first, becoming pale or dark orange- brown, with orange-brown spots on distal membranous parts, ligules acute or obtuse. Highest distal sheath very often situated below the medial part of the stem, 2.1-3.6 mm wide, with blades reduced or lacking. Blades of proximal sheaths flat to slightly cymbiform, 25-130 x 0.5-1.4 mm, glabrous or rarely scabrous in distal parts, the apex mostly acute. Blades of distal sheaths shorter, 0.2-55 x 0.2- 1.0 mm, or lacking. Spikelets solitary, typically hemispherical at maturity (Fig. 7), 1.0-3.0 x 1.4-4.5 cm, with 150 or more florets. Proximal scales 5-6, without florets. First proximal scale olive-brown or blackish, becoming pale brown, yel- 810 BRIT.ORG/SIDA 21(2) lowish brown or pale beige in distal parts, and pale beige or hyaline on the mar- gins, widely lanceolate or ovate, 5-12 x 1.6-4.3 mm, with 4-9 beige or olive nerves converging below the apex, acuminate. Medial fertile scales with a short proxi- mal part (Fig. 9), 0.1-0.9 mm long, averaging 2-25% of total scale length, pale green or whitish, with the medial part blackish or dark gray, with marginal parts blackish (Fig. 9a) or narrowly hyaline (Figs. 9b-c), and the distal part dark or hyaline, narrowly lanceolate, 3.2-5.3 x 0.4-LO mm, the widest part below the middle or close to the base, with width at the middle 0.3-0.7(-0.9) mm, with 1 incom plete nerve, nar rowly ac uminate, 0.05-0.1(-0.2) mm wide at 0.2 mm below the apex. Proximal fertile scales very similar (Fig. 8), blackish, with well-delim- ited narrow hyaline margins (Fig. 8b) or with hyaline margins lacking (Fig. 8a). Perianth of about 30 hypogynous bristles, white or cream white (Fig. 7), 15-25 mm long. Stamens with filaments about as wide as perianth bristles, anthers yel- low or pale yellow, 0.35-0.8 mm long. Styles with 3(-4) stigmatic branches barely open at maturity, branches 0.5-1.3 mm long. Achenes beige-brown to olive-brown (Fig. 10), narrowly obovoid, obscurely trigonous, slightly biconvex or plano-con- vex, glabrous, slightly lustrous, 1.7-2.4 x 0.5-0.85 mm, base cuneate, apex acute, witha beak more often oblique or curved (Fig. 1Ob) than straight (Fig. 10a), mostly cylindrical, 0.15-0.4 mm long, 0.05-0.1 mm wide at base. Figs. 7-10. Distribution —Eriophorum scheuchzeri was recently subdivided into two subspecies by Novoselova (1994b), and the typical subspecies is very common and widespread in the southern parts of the Arctic zones. Eriophorum scheuchzeri subsp. scheuchzeri is an arctic-alpine circumpolar taxon (Novoselova 1994a, 1994b). In North America, it covers both Arctic and Boreal zones (Hultén and Fries 1986). In northeastern North America, it is sympatric with both subspecies of E. russeolum, ranging southward to Labrador, New- foundland, and the southern reaches of James Bay at about 51°N (Scoggan 1978). Discussion.—The typical subspecies of E. scheuchzeri differs from other rhizomatous taxa with solitary whitish spikelets, including their hybrids with the two subspecies of E. russeolum, by having the shortest anthers (0.35-0.8 mm) and the narrowest (0.4-1.0 mm) and the most narrowly acuminate medial fer- tile scales (Table 1, Fig. 9). These scales are dark gray or blackish with narrow hyaline margins or with hyaline margins absent. Achenes are narrowly obo- void, obscurely trigonous, their beaks narrowly cylindrical, and more often oblique than straight (Table 1, Fig. 10). The main differences between subsp. scheuchzeri and subsp. arcticum are given in Table 2. 6. Eriophorum scheuchzeri Hoppe subsp. arcticum Novoselova, Bot. a oe Petersburg) 79(4):112. 1994. Type: RUSSIA. JENISSEJSK et Stellingii, 23 Jul 1915, L Trzhemesky 35 HOLOTYPE: LE, not seen). Herbs perennial with short to elongate rhizomes. Vegetative shoots l-3, 4-7 cm high, leaf margins glabrous. Stems erect, glabrous, terete in cross section, 11-28 CAYOUET ] E, TRAVUINVINIT UE 811 cm high, 0.7-1.6(-1.8) mm in diameter below the inflorescence. Leaves basal and cauline 1-4. Proximal sheaths apple green first, becoming pale orange-brown, with or without orange-brown spots on distal membranous parts, ligules acute or obtuse. Highest distal sheath most often situated below the medial part of the stem or near the base, 2.2-3.4 mm wide, with blades reduced or mostly lack- ing. Blades of proximal sheaths flat to slightly cymbiform, 15-80 x 0.5-1.1 mm, glabrous, the apex mostly obtuse. Blades of distal sheaths shorter, 2-20 x 0.4- 0.8 mm, or mostly lacking. Spikelets solitary, typically spherical (Fig. 11) or slightly flattened at maturity, 1.5-2.5 x 1.5-4.0 cm, with 100 or more florets. Proximal scales 1-4, without florets. First proximal scale blackish, becoming pale brown or pale beige in distal parts, with well developed hyaline margins, ovate-lanceolate, 6-9 x 2.9-4.2 mm, with 3-7 brown or gray nerves converging below the apex, acute. Medial fertile scales with a short proximal part (Fig. 13), 0.5-1.0 mm long, averaging 8-21% of total scale length, pale green, pale brown or blackish, with the medial part grayish or dark gray-brown, with marginal parts dark (Fig. 13b) or narrowly hyaline (Fig. 13a), and the distal part dark or hyaline, lanceolate, 4.0-6.0 x 0.7-1.5(-L7) mm, the widest part below the middle, with width at the middle (0.5-) 0.7-1.4(-1.6) mm, with 1 incomplete nerve, acuminate, 0.1-0.25(-0.3) mm wide at 0.2 mm below the apex. Proximal fertile scales different (Fig. 12), bicolor, with lower and medial parts dark but gradu- ally passing into various tones of gray and conspicuous marginal and distal hyaline areas. Perianth of 25-40 hypogynous bristles, white (Fig. 11), 16-25 mm long. Stamens with filaments about as wide as perianth bristles, anthers yellow or pale yellow, 0.6-1.0 mm long. Styles with 3 stigmatic branches barely open at maturity, branches 0.7-1.5 mm long. Achenes orange-brown to dark reddish- brown (Fig. 14), narrowly obovoid, mostly biconvex or slightly plano-convex, glabrous, mostly dull, 1.5-2.2 x 0.5-0.7(-0.9) mm, base cuneate, apex acute, with a beak more often oblique or curved (Fig. 14a) than straight (Fig. 14b), mostly cylindrical, 0.15-0.3 mm long, 0.05-0.1 wide at base. Figs. 11-14. Distribution.—Like Eriophorum scheuchzeri subsp. scheuchzeri, subsp. arcticum is at least partially circumpolar, according to Novoselova (1994a, 1994b), but its range is more High Arctic than the typical subspecies, as is illus- trated by its distribution in northern Russia. Novoselova claims that subsp. arcticum ranges across Alaska, Arctic North America and Greenland. My re- sults confirm its presence in the Canadian High Arctic Nunavut (from at least 67°N to 81°N) and establish its southern limits on islands in Hudson Bay and in Arctic Quebec at latitude 59°N or 60°N (see specimens examined). Since one cited collection is from Port Burwell, Quebec, its presence in adjacent northern Labrador is expected, but no specimen has yet been positively identified. Discussion.—Differences between the two subspecies were pointed out by Novoselova (1994b). Since no North American specimens of subsp. arcticum were cited in her work, the attempt was made to uncover voucher collections of 812 BRIT.ORG/SIDA 21(2) the subspecies in order to discover the most useful characters to differentiate between the two E. scheuchzeri taxa. The results are highlighted in Table 2 and have been confirmed by Novoselova (pers. comm.). The best characteristics were noticed in the color pattern of the proximal fertile scales (Figs. 8, 12), and in the widths of the medial fertile scales if measured near the middle of the scales and at 0.2 mm below the apex (Figs. 9, 13). Scales were wider in subsp. arcticum and acuminate (Fig. 13), instead of being narrowly acuminate (Fig. 9) as in the typical subspecies. Mature spikelets tended to be spherical (Fig. LL) in subsp. arcticum, rather than hemispherical (Fig. 7) as in subsp. scheuchzeri. A color difference was observed in mature achenes (Table 2, Figs. 10, 14). At the south- ernmost limit of F. scheuchzeri subsp. arcticum and elsewhere in the sympatric range of the two subspecies, a few specimens tend to be of intermediate nature. Since only a few voucher specimens of subsp. arcticum were collected within the borders of Quebec, I have proposed that it be added to the provincial list of threatened and — vascular plant species. Specimens examined, CANADA, Nunavut: Axel Heiberg Island, 79°54’ N-87°43' W, 19 Jul 1980, G.W. Scotter & S.C. Zoltai aa Baffin Island, head of Clyde Fjord, Jul 1950, M.E. Hale Jr. 40 WIS); Devon Island, Truelove Lowland, 75°38' N-84°30' W, 24 Jul 1989, B.C. Forbes 70 (DAO); Cambridge Bay, 69°03' N-104°50' W, 7 Aug 1950, E-H.N. Smith & G.K. Sweatman 42 (DAO); Ellesmere Island, east coast, between Baird weaee none tae acier, 78°29' N-76°31' W, 20 Jul 1979, J. Bridgland 694 (DAO), idem, Eureka, 80°O1' N-86° V,19 Aug 1953, PE Bruggeman 697 (DAO), idem, Eureka, 79°59" N- 85°50! W, 16-18 Jul 1980, - Ww. ee & S.C. Zoltai 45292 (DAO), idem, Hazen Camp, 81°49' N-71°21' W, 9 Jul 1962, D.B.O. Savile 4583 (DAO), idem, Skraeling Island, 78°36.5' N-75°38.5' W, 20 Jul 1981, W. Blake Jr. 24-1(DAO), Ottawa Islands [wrongly a as North Sleeper Islands, see Morisset and sia (1980)], 2 Aug 1939, G. Gardner 39891 la] (MT, QFA); idem [not N. Sleeper Islands], 59°17 J°40' W, 2 Sep 1939, A. Dutilly, H. ONeill, & M. ne poe (QFA); Ottawa Island Archipelago, Pattee Island, 59°42’ N-80°09 W, 27 Aug 1939, A. Dutilly, H. O'Neill, & M. Duman 87516 (CAN, DAO, SFS); Prince Charles Island, 67°51'27" N-75°06'07.2" W, 7 Jul 1997, V Johnston 97-161 (DAO); Somerset Island, 72°49' N-92°56' W, 19 Jul 1975, S.C. Zoltai 751135(DAO); Southhampton Island, Coral Harbor, 64°09' N-83°18' W, 16 Jul 1948, WJ. Cod ed 1348(DAO, WIS). Québec: environs d’Akulivik, 60°48 N-78°12' W, 8 Jul 1985, M. Blondeau 85060 (QFA); environs d'Ivujivik, 62°24 N-77°55' W, 17 Jul 1984, M. Blondeau 84235B (QFA); aa 62°25' N-78°05' W, 23 Jul 1938, M. Duman 1874 (QFA), Port Burwell, 60°22' N-64°50' W, 30-31 Aug 1927, M. O. Malte 118677 (CAN). a KEY TO TAXA (INCLUDING ERIOPHORUM CHAMISSONIS AND ATYPICAL FE. RUSSEOLUM SUBSP. RUSSEOLUM) . Spikelets with dark to aS eke bristles. 2. Medial fertile scales 0.7 m wide, acuminate, 0.1-0.3 mm wide at below the apex; aa cae obovoid, glabrous; hypogynous fete 30- 0, 15-20 mm long Eriophorum medium subsp. medium ae ae scales 1.2-2.2 mm wide, acute, ee obtuse or acuminate, 0.2 5(-0.9) mm wide at 0.2 mm below the apex; achenes ob id or ellipsoid, gla- an or scabrous; hypogynous bristles ae 25 40 mm long. 3. Anthers 1.5-3.1 mm long; medial scales with conspicuous hyaline margins and apex, the widest area near the middle or above; spikelets pala obo- void, with dark to pale orange-brown bristles iophorum russeolum subsp, russeolum N CAYOUETTE, TAXONOMY UI 813 3. Anthers 0.7-1.6(-1.9) mm long; medial scales often with reduced hyaline margins and apex, the widest area not above the middle; spikelets various, spherical, obovoid, or hemispherical, with pale beige-brown to darker bristles 4. Spikelets spherical, with pale beige-brown bristles; first proximal scale 12— 23(-30) mm origi stem Saal y 1e sl ieaalcen, Ke 2.2mm ae medial scales covered pots in hyaline areas; achene beak rarely curved; western North America _ ophorum chamissonis 4. Spikelets obovoid cae aad with pale to dark orange- oe bristles first proximal scale 8-11 mm long; stem below the inflorescence 0.6-1 2 m wide; medial ae eneels without reddish-brown longitudinal spots; achene ania ently curved; amphi-Atlantic atypical Eriophorum eolum and/or intermediates between E. x medium and E.russeolum 1. Spikelets with ne to whitish bristles 5. Medial scales (0.8-)1.0-2.4 mm wali acute, 0.25-0.6 mm wide at 0.2 mm below the apex, widest mostly at the middle or above, with well developed hyaline margins; anthers (1.3-)1.5-3.1 mm long; achenes ellipsoid or obovoid, scabrous or glabrous, beak base 0.1-0.2 mm wide Eriophorum russeolum subsp. leiocarpum . Medial scales 0.3-1.5(-1.7) mm wide, acuminate to narrowly acuminate, 0.05- 0.3(—0.4) mm wide at 0.2 mm below the apex, widest below the middle or close to the base, with frequently reduced hyaline margins; anthers 0.35-1.6 mm long; achenes narrowly obovoid, always glabro 1s, beak base 0.05-0.1 m 6. Anthers 0.9-1.6 mm long; hypogynous oe (10-)22-32 mm see aod branches 1.0-2.2 mm long Eriophorum medium subsp. album 6. Anthers 0.35-1.0 mm long; hypogynous bristles 10-25 mm long; stigmatic branches 0.5-1.3(-1.5) mm long Eriophorum scheuchzeri s.|. 7. Spikelets hemispherical; proximal fertile scales dark, with dark margins o vee) reduced hyaline margins shat ply differentiated from the darker Be. Sal dial scales narrowly acuminate (usually 0.1 mm wide at 0.2 mm below the apex), 0.3-0.7(-0.9) mm wide near the middle; mature achenes beige brown to olive-brown, slightly lustrous Eriophorum scheuchzeri bsp. scheuchzeri 7. Spikelets spherical; proximal fertile scales bicolored, with lower and me- dial parts dark but gradually passing to various tones of gray and con- spicuous marginal and apical hyaline ieee scales acuminate (usu- ally 0.2 mm wide at 0.2 mm below the apex), (0.5-)0.7-1.4(-1.6) mm wide near the middle; mature achenes orange-brown to dar reddish-brow mostly dull Eriophorum scheuchzeri aie arcticum ACKNOWLEDGMENTS The author thanks the curators of the cited herbaria for access to their collec- tions, as well as the following individuals for their help: M. Garneau, R. Néron, and K. Damboise of the Northern Québec-Labrador flora project for the illus- trations; my colleagues Y. Dalpé and J. McCarthy for the production of the color plate; M.S. Novoselova for her comments on an earlier version of this paper, her translations from the Russian and for various other bibliographic information; R. Moberg (UPS) for his help in selecting a lectotype of E. russeolum; R. Elven, P. Morisset, and M. Dubé for information; P.W. Ball, E. Small, PM. Catling, and G. Hall for criticism of the manuscript. 814 BRIT.ORG/SIDA 21(2) REFRENCES Bait, PW. and D.E. Wek. 2002. Eriophorum Linnaeus. In: Flora of North America Editorial Committee (editors). Flora of North America North of Mexico, Volume 23: Magnoliophyta: Commelinidae (in part): Cyperaceae. Oxford University Press, New York and Oxford. Pp. 21-27. Beraaren, G. 1969. Atlas of seeds and small fruits of Northwest-European plant species with morphological descriptions. Part 2. Cyperaceae. The Swedish Natural Science Research Council, Stockholm. Bown, B. 1992. Les Cypéracées de l’est du Canada. Provancheria 25:1-230. CayouerTTe, J., M.Bionbeau, and P.M. Catuina. 1997. Pollen abortion in the Ranunculus gmelinii- hyperboreus group (R ulaceae, Section Hecatonia) and its taxonomic implications. Rhodora 99:263-274, Facarl, K. 1958. Zur Hybridbildung in der Gattung Eriophorum. Veroff. Geobot. Inst. Rubel Zurich 33:50-58. Houmeren, P.K., N.H. Houmaren, and L.C. Burnett. 1990. Index Herbariorum, Part I: The Herbaria of the World, 8th edition. New York Botanical Garden. Hutten, E.and M. Fries. 1986. Atlas of North European vascular plants north of the Tropic of Cancer. Vols.| and Ill. Koeltz Scientific Books, Konigstein HyLanper, N. 1982. Nordisk Karlvaxtflora. Volume 2, ed. 2. Almaqvist & Wiksell, Stockholm. Morisset, P.and S. Payette. 1980. La flore et la végétation des Iles Dormeuses (Baie d'Hudson, Territoires du Nord-Ouest). Naturaliste Canad. 107:63-86. Novosetova, M.S. 1993. The taxonomy of the Eriophorum (Cyperaceae) species allied to Eriophorum russeolum. Bot. Zurn. (St. Petersburg) 78(8):80-89. [In Russian]. Novosetova, M.S. 1994a. The system of the genus Eriophorum (Cyperaceae). Il. Subgenus Eriophorum. Bot. Zurn. (St. Petersburg) 79(12):66-75. [In Russian]. Novosetova, M.S.1994b. Critical notes on the species of the genus Eriophorum (Cyperaceae) allied to Eriophorum scheuchzeri, Bot. Zurn. (St. Petersburg) 79(4):111-119. [In Russian]. NyLanber, F. 1846. Eriophori monographia. Acta Soc. Sci. Fenn. 3:1-23. Potunin, N. 1940. Botany of the Canadian eastern Arctic. Part |: Pteridophyta and Spermatophyta. Natl. Mus. Canada, Bull. 92:1-408. Raymond, M. 1954.What is Eriophorum chamissonis C.A. Meyer? Svensk Bot. Tidskr.48:65-82. Scocean, H.J. 1978. The flora of Canada. Part 2. Pteridophyta, Gymnospermae, Monocotyledonae. National Museums of Canada, Ottawa. Pp. 93-545. Totmactey, A.1. 1996. Eriophorum L.Cotton grass.|In:A.l.Tolmachev and J.G. Packer, eds;trans- lator G.C.D. Griffiths. Flora of the Russian Arctic. Volume 2: Cyperaceae—Orchidaceae. The University of Alberta Press, Edmonton, Canada. Pp. 3-18. Tucker, G.C.and N.G.Mitter. 1990. Achene microstructure in Eriophorum (Cyperaceae):Taxo- nomic implications and paleobotanical applications. Bull. Torrey Bot. Club 117: 266-283. TAXONOMY OF THE LIATRIS PILOSA (GRAMINIFOLIA) COMPLEX (ASTERACEAE: EUPATORIEAE) Guy L.Nesom Jon M. Stucky Botanical Research Institute of Texas Department of Botany 509 Pecan Street North Carolina State University Fort Worth, Texas 76102-4060, U.S.A. Raleigh, North Carolina 27695 gnesom@brit.org ABSTRACT. Liatris graminifolia Willd. is the name generally used for the grass-leaved gayfeather of the south- eastern United States. Gray (1884), Gaiser (1946), and Wilbur (1962) observed that the name Liatris pilosa (Aiton) Willd. apparently applies to this species and has priority; Fernald and Griscom (1938) dissented, but the present study concurs that L. pilosa should replace L. graminifolia as the correct name. A lectotype from BM is designated for Liatris (Serratula) pilosa. This specimen probably was collected in New JetSey or Delaware and appare ently Represents! a perucuiaey hairy populations, 7 varian nto but that tic of the oe in the broader Atlantic coast region. Two! taxa that have been identified as varieties of L. graminifolia are here treated at specific rank: Liatris elegantula (Greene) K. Schum. occurs on the Gulf coastal si in Missi spe! (rare), Alabama, Blotics, and Cores Liatris virgata. Nate SG Liatris regimontts, eae lina ang Georg northward through western North Carolin into Virginia. iphese three taxa iff ly I g vivgat is geographically juxtaposed Ragees L. pilosa and L. elegantula. | 1i 1 I ved in areas of sympatry at the range margins. Liatris cokeri Pyne & Stucky is a fourth member of this group, possibly most closely related to L. virgata. A taxonomic summary is provided, including nomenclature, distribution maps, ecological summaries, and a key. RESUMEN Liatris graminifolia Willd. es el nombre que se usa genealnente para la planta del Sureste de los Estados ae Gray (1884), SC eaieasigs y Wilbur (1962) obse ue el nombre Liatris pilosa (Aiton) W a esta especie y tiene sriorideck Fernald and Griscom (1938) oo a en el presente esudio se concluye que L. pilosa debe remplazar a L. graminifolia mbre correcto. Se designa un lectotipo de BM para Liatris (Serratula) pilosa. Este espécimen P once fue colectado en Nueva Jersey o Delaware y aparentemente representa una variante poblacional particularmente pelosa de la especie que se encuentra en el area pero que se intergrada con plantas mas caracteristicas de la Specie enla agen Atlantica costera mas ane Dee taxa que Liatris han sido identificados como variedades de L.g elegantula (Greene) K. Schum. vive en la Ilanura costera del Golfo e en Mississippi Gara), Alabama, Florida, y Georgia; Liatris virgata Nutt. (= Liatris regimontis, Lacinaria smallii) ocupa una posicion geografica apteEmed as yenee seeds Carolina del Sur y Georgia por el Oeste de Carolina del Norte la morfologia involucral y el rango de L. han observado hasta Virginia. virgata esta yuxtapuesto ee entre L. pilosa y L. elegantula. No se intermedios en areas de simpatria en los extremos de area. Liatris cokeri Pyne & Stucky es un cuarto mie pore de este erHpo, post emenle mas Helacionsae’s con = virgata. = ofrece un resumen 3 y una clave. wh oO SIDA 21(2): 815-826. 2004 816 BRIT.ORG/SIDA 21(2) — Liatris graminifolia Willd. is the name generally applied to the grass-leaved gayfeather, a taxon of the southeastern U.S.A. (e.g., Radford et al. 1968; Cronquist 1980; Figs. | and 2). Gaiser (1946) recognized five infraspecific taxa: var. graminifolia, var. elegantula (Greene) K. Schum., var. lasia Fernald & Griscom, var. dubia (WPC. Barton) A. Gray, and var. smallii (Britton) Fernald & Griscom Of these five, var. dubia and var. lasia both are representative of the Atlantic coast species, as is var. graminifolia; var. elegantula is treated here at specific rank; and var. smallii is treated here as a synonym of another formally recog- nized species. Fernald (1950) recognized L. graminifolia var. graminifolia, var. racemosa (DC.) Venard (as a replacement name for var. dubia), var. lasia, var. smallii, and var. virgata (Nutt.) Fernald. We observe that Fernald’s concept of var. virgata (1949, 1950) was artificial and that var. racemosa represents the same taxon as the type of var. virgata. The only recent treatment of the genus in the area that includes all of these variants is Cronquist (1980), who reduced the formally recognized taxa to L. graminifolia vars. graminifolia and elegantula. In our assessment, these two and two more, L. graminifolia var. virgata sensu stricto and L. cokeri Pyne & Stucky, constitute the evolutionary entities of this complex. Liatris cokeri is a species of the fall-line sandhills of southern North Carolina and adjacent South Carolina (Stucky & Pyne 1990). Our treatment rec- ognizes four taxa, each at specific rank: L. pilosa (Aiton) Willd., L. elegantula (Greene) K. Schum., L. virgata Nutt., and L. cokeri. a Taxonomic rank It is clear that Liatris pilosa, L. elegantula, L. virgata, and L. cokeri are closely related among themselves. Morphological differences among them, mostly in involucral features, are relatively small but they are consistent and a series of principal components analyses (Stucky 1990, 1992) indicates that L. cokeri, L. pilosa, and L. virgata are distinct. Liatris elegantula was not included in the analyses by Stucky, and it has consistently been treated as a variety of L. graminifolia since Gaiser reduced it in rank. In addition to morphology, the decision regarding the rank of these taxa rests on biology. Liatris cokeri is com- pletely sympatric with L. pilosa but contiguous or nearly so with L. virgata (Figs. Land 2). Liatris virgata is geographically juxtaposed between L. pilosa and L. elegantula and probably forms a reproductive barrier between them. From the sample of specimens studied and mapped here, it appears that the degree of sympatry between L. virgata and L. elegantula may be greater than between L. virgata and L. pilosa; in neither instance, however, have we seen collections that would clearly indicate that hybridization, intermediacy, or introgression oc- curs in the areas of sympatry (see comments below). Each of these taxa has been treated at varietal rank, but the nomenclature for treating them as species is already established. — 817 NESOM AND STUCKY, TAXONOMY OF LIATRIS PILOSA COMPLEX © Liatris pilosa eae = ata © ®@ Liatris virg . Gs ) Liatris virgata if antulain Elyrid J ef , J J SMU/BRIT, TEX/LL, USCH, and VDB. Tagg V, NCU, J Fic. 1.G ied from DO (1946). Some records for L. virgata (half-filled circles) are ret AAS .£ yy (90NNA) in Alah \ d 1990). ( added from Stucky & Pyne 818 BRIT.ORG/SIDA 21(2) neutegs ras : \ : : = ce Liatris cokeri Fic.2.G graphic distributi f Liatris cokeri.R d fi I i lied f NCU, SMU/BRIT, USCH, and VDB Identification of the species With the maps and following key, we believe that identifications can be made with accuracy and consistency. All key contrasts are not mutually exclusive, but they contribute toward an understanding of distinctions among the taxa. Species descriptions are provided in a treatment of the genus for the forthcom- ing Flora of North America volumes of Asteraceae (Nesom in prep.). KEY TO THE SPECIES OF THE LIATRIS PILOSA COMPLEX 1. Phyllaries apically rounded, lamina relatively thin, eglandular or with superficial to shallowly inset punctate glands, completely bordered by a narrow, hyaline rim; in- volucres turbinate (obconic) to turbinate-campanulate. 2 Cr lakh i} | “| : + | | | + | f y of Moaeratel erwnoie stem;leal g p pilo or ov laminae glabrous to sparsely pilose on abaxial surface; heads relatively densely arranged, on internodes (1-)2-5(-7) mm long; peduncles 0-10(-17,-80 in proxi- mal region of capitulescence) mm long; involucres (7-)8-10 mm long, phyllaries in (3-)4-5(-6) series; florets (6-)7-12(-13), mostly 9-13 in NJ. and Del. Liatris pilosa 2. Stems glabrous; leaf laminae glabrous; heads relatively loosely arranged, on in- ternodes (2-)5-10(-14) mm long; peduncles 0-2(-7) mm long; involucres 6-8 mm long; phyllaries in 3—4(-5) series; florets (7-)8-11(-13) Liatris elegantula . Phyllaries apically angular, lamina relative thin or thick, with inset or superficial glands, bordered on the lateral margins e But not at the apex by a narrow, hyaline rim; in- volucres cylindric-campanulat 3. Heads densely 1,0n ee oe 1—2(-5) mm long, often secund; phyllary apex sharply acurninate: acute, distinctly involute, lamina relatively thin, glands consistently present and superficial at least on proximal portion; florets 4-7(-9 er head; basal and lower cauline leaves 2-5 mm wide, gradually reduced in length distally Liatris cokeri 3. Heads loosely arranged, on internodes 6-15(—20) mm long,not secund; phyllary apex sharply acute to obtuse-angled with a thickened apiculum, not markedly involute, lamina relatively thick, usually with evidently sunken punctate glands, without superficial glands; florets 7-10(-12) per head; basal and lower cauline leaves 4—9(-12) mm wide, quickly reduced in width and length distally__ Liatris virgata ~ Liatris pilosa: the oldest correct name for L. graminifolia Liatris pilosa (Aiton) Willd. 1803 (based on Serratula pilosa Aiton 1789) is the oldest name in the L. graminifolia complex but was treated by Gaiser as a syn- onym of L. graminifolia var. dubia. Fernald (1950) did not include the name L. pilosa in his account of the genus. Gray (1856) had treated L. pilosa asa distinct, montane species (“Mountains of Virginia and southward”), but he later (1884) regarded L. pilosa as a questionable synonym of L. graminifolia var. dubia. Fernald and Griscom (1935) examined the “fragments of a head from Aiton’s type” of Serratula pilosa (from GH, perhaps obtained by Gray from type mate- rial at BM), but they concluded that “Serratula pilosa has nothing to do with Liatris graminifolia” - emphasizing the “long pedunculate” heads (from the type description) and the “linear and acute ... involucral bracts” (from the GH frag- ments). Gray (1884) had observed these same features and noted that L. pilosa represents a ‘state’ of L.graminifolia with “unusually narrow involucral scales.” Gaiser’s lengthy discussion of the typification of L. pilosa (1946, pp. 257-258) indicated that she regarded it as conspecific with L. graminifolia, and her place- ment of it in the synonymy of var. dubia seems to have been more of a nomen- clatural error than reluctance to use the name because of uncertainty about its identity. Wilbur (1962) accepted L. pilosa as the correct name for the species, perhaps based on Gaiser’s comments, while acknowledging the different inter- pretation by Fernald and Griscom; he noted that it seemed undesirable to pro- vide new combinations for infraspecific taxa prior to critical study of infraspe- cific variation. After a detailed survey of variation within Liatris graminifolia in the con- text of a study of the whole genus, and with the opportunity to study type material of Liatris pilosa from BM, we also conclude that the name L. pilosa does indeed apply to the species and must replace L. graminifolia as the correct name. Gaiser (1946, pp. 257-258) quoted notes from E.G. Baker of the National Herbarium, who apparently also examined type material of Serratula pilosa and whose observations regarding its morphology agree with ours. Some plants 820 BRIT.ORG/SIDA 21(2) from New Jersey and Delaware, at the northern extremity of the range of the species, which have prominently pilose stems and leaf lamina and a tendency to produce long peduncles, are similar to the BM type. We have seen collections of this “morphotype” from Atlantic, Camden, and Cumberland cos., New Jersey, and Sussex and Kent cos., Delaware. The lanceolate, apically acute phyllaries of the Serratula pilosa type are unusual for the species over most of its range but this feature appears sporadically in plants of the New Jersey-Delaware region. Some plants in New Jersey corresponding to Liatris pilosa sensu stricto occur in populations of relatively uniform morphology (David Snyder, pers. comm.): these plants have “lower branches up to 11 cm long with up to 5 heads. The branching is most pronounced on the lower half of the stem but the heads of the upper are long peduncled (up to 4 cm long). The branches are strictly ascending. Stems, peduncles, and leaf bases are densely hirsute.” On the other hand, plants more similar to those in eastern Virginia and North Carolina ap- parently are at least as common or more so in New Jersey (Atlantic, Burlington, Cape May, Ocean cos!) and Delaware (Sussex and Kent cos.!) as the prominently hairy ones. Plants from this region with stems and leaf lamina glabrous or sparsely hairy but with slightly larger heads were identified as L. graminifolia var. dubia by Gaiser (1946), who cited collections from New Jersey, Delaware, Maryland, District of Columbia, Virginia, and Pennsylvania (Bucks Co.). Our study substantiates the observation that some populations of this region are distinct in their combination of characters, but the tendencies for relatively densely pilose stems and leaves, long-pedunculate heads, more florets per head, and inner phyllaries with subacute apices apparently are only loosely corre- lated among themselves. We have not been able to meaningfully sort the varia- tion, but this is an area that needs to be investigated more closely. Liatris elegantula Plants of Liatris elegantula have consistently glabrous stems and leaves, rela- tively short and distinctly turbinate (obtriangular) involucres with a reduced number of phyllaries (evidenced by fewer series), and the heads tend to be more widely spaced than in L. pilosa. Records for this taxon cited by Gaiser (1946) from southwestern Alabama (Baldwin Co.) and adjacent Mississippi Jackson Co.) have not been examined in this study. Treatment of Liatris elegantula at specific rank is perhaps the most diver- gent proposal of the current overview. It is most similar to L. pilosa in involu- cral morphology, but small differences between the two are consistent and the geographic hiatus is real. Liatris elegantula and L. pilosa might be treated as conspecific, as has been generally done, or L. elegantula, L. pilosa and L. virgata might all be considered as a single species, but this would not account for ap- parent reproductive isolation in areas of sympatry (comments above) or a pos- sible close relationship between L. virgata and L. cokeri. — — NESOM AND STUCKY, TAXONOMY OF LIATRIS PILOSA COMPLEX 821 Liatris virgata and L. cokeri Liatris virgata has mostly been identified within Liatris graminifolia sensu lato, and as observed by Stucky (1992), this name has not been included in most of the pertinent taxonomic liter pais for the See even asasynonym, although one of its synonyms (L. graminifolia var. smallii) has sometimes been correctly applied. A ounce components analysis (Stucky 1992) indicates that L. virgata and L. graminifolia (L. pilosa) are morphologically distinct. The name L. regimontis (Small) K. Schum., now understood to be a synonym of L. virgata, mostly had been applied to the species segregated by Stucky and Pyne (1990) as L. coheri. The range of Liatris virgata is essentially contiguous with L. elegantula on the southwestern margin and with L. pilosa on the northeastern margin, but some overlap occurs in both areas (Fig. 1). Although L. virgata has been col- lected in close proximity to both of its closest relatives and all three species flower in generally the same period of time, our observations indicate that the taxa are discrete even in areas of sympatry. For example, from York Co,, S.C., we have studied five collections of L. virgata (Nelson 4994, Kennemore 917, 997, 1046, 1486, all USCH) and three of L. pilosa (Nelson 4989, 4998, 5024, all USCH)—all eight of these were collected within Kings Mountain National Military Park, From Richland Co., 5.C., we have studied 20 collections of L. virgata (USCH, NCU, BRIT) and a single one of L. pilosa (Nelson 11244, USCH), south Of its pr= mary range. Field and herbarium studies are needed to further the understand- ing of the geographic and evolutionary relationship between L. virgata and its close relatives. If hybridization and intergradation prove to be more significant than observed in the current study, treatment of these three taxa as conspecific might be more appropriate. Spacing of the heads and phyllary morphology are features that provide the most immediate recognition of Liatris virgata. Contrasted with L. pilosa and L. elegantula, the phyllary lamina is thicker and the glands are distinctly sunken into the tissue, and the apex is generally angular (vs. rounded) and lacks the narrow hyaline rim that borders the lateral margins. In Richland Co, S.C, from which numerous collections are available, the apex shape varies from sharply acute to obtuse, but even the obtuse angle is distinct, as the tip charac- teristically ends ina thickened and slightly raised (keel-like) apiculum or mu- cro. Similar variation occurs over the range of the species, although a tendency for obtuse apices apparently is more common on the coastal plain. Variation in Liatris virgata also occurs in involucral size and configura- tion. Larger-headed plants (including the types of Lacinaria smallii and Lacinaria regimontis) are mostly and piedmont. Larger heads are more elongate-cylindric and have phyllaries in 5-6(-7) series with more consistently sharply acute apices, while smaller heads have 3-5(-6) series. Number of flo- rets tends to be slightly higher in larger heads. 822 BRIT.ORG/SIDA 21(2) Finally, we note that the distribution of Liatris virgata from outer coastal plain into montane habitats is unusual, but L. pilosa and L. elegantula both oc- cur on the piedmont as well as their primary coastal plain range, and other species of Liatris range widely across habitats and ecological zones (e.g., L. squarrulosa Michx. and L. aspera Michx.). Further study of L. virgata may dem- onstrate geographic patterns of differentiation that we have not been able to delimit. Stucky and Pyne (1990) observed that apparent intermediates between Liatris virgata and L. cokeri occur on the coastal plain of North Carolina and South Carolina. In the present study, however, we have identified some of those putative intermediates as L. cokeri, and we have not confirmed the occurrence of L. virgata where the putative intermediates occur in North Carolina (Stucky S Pyne 1990, Fig. 10). Liatris cokeri is characterized by phyllaries with acute- angled apices, and this is likely an indication of close relationship to L. virgata. Indeed, evolutionary relationships within the L. pilosa complex, as outlined here, may be that of two sister pairs—L. pilosa-elegantula and L. virgata-cokeri. NOMENCLATURE AND TYPOLOGY Liatris pilosa (Aiton) Willd. Sey PL. 3:1636. 1803. Serratula pilosa Aiton, Hort. Kew. 3:138 1789. Lacinaria graminifolia (Willd.) Kuntze var. pilosa (Aiton) Britton, Mem. Torrey Bot.Club 5.314. 1894. Lacinaria pilosa (Aiton) A. Heller, Muhlenbergia 1:6. 1900. LECTOTYPE, here desig- mate U.S.A. Cultivated plant, without collection data but the original stock probably from New Jersey or Delaware, probably collected 7 ce Young, Jr. prior to 1783 (BM-Banks Herbarium, photol, fragment of lectotype GH andwritten inscription on the back of the lectotype sheet reads “Hort Kew. 1785” and eee i: handwriting of Jonas Dryander (Mar- a shall 1978), who assumed the primary responsibilities of describing and naming plants for the Hortus ese ae the ae of Daniel Solander in 1782. The publication itself, how- ever, credite p solely to William Aiton (see Britten 1912). Photos of the lectotype have been deposited at BRIT, a nee NCU, and US. The eprotsloniee! Serratula pilosa described the plants as “foliis linearibus pilosis, floribus axillaribus longe pedunculatis” and noted “Nat. of North America. Introd. 1783, by Mr. Will- iam Young.” William Young, Jr. lived in Philadelphia and eae orays into “the ae as he collected horticultural stock for English gardeners (Harshberger 1917). It seems a reason- able surmise that the material of L. pilosa was collected by Young in the region of his home, probably see by in New Jersey or Delaware, where plants of this morphology are known to occur (as al for the type of L. dubia, see below; Keller and Brown [1905] noted records in New Jersey sae Delaware ae ‘Liatris eae pilosa”). The type specimen presumably was grown in cultivation at Kew Gardens, as Young was supported as “Botanist to their Majestys” in collecting horticultural possibilities. Young informally used the name Serratula pilosa for gayfeather material in his plant collection (Young 1985). Liatris graminifolia Willd., Sp. Pl. 3:1636. 1803. Lacinaria Sy as (Willd) Kuntze, Revis. Gen. Pl. 1:349. 1891. Type: Original not located. U.S.A. NORTH CAROLINA. New Hanover Co: edge of Wilmington, common in the open pine ieee ee ting the Cypress Tree Park, 24 Oct 1948, E.O. White s.n. (NEOTYPE (Gaiser 1950, p. 414): GH, internet image!; ISONFOTYPES: [MO] noted by Gaiser to have been deposited at NY and US). The collection date apparently was miscited by Gaiser as “25 Oct,” because the GH specimen reads “24 Oct” and corresponds in 823 all other details with the citation. Many authors, including Gaiser (1946), have interpreted Willde name as a new combination based on a name of Thomas Walter (Anonymos peri Walter, Fl. Carol. 197. 1788), but as nod by Wilbur eae and others, Walter’s names using “Anonymos’ as tl lid (ICBN 2000: Arts. 20.4, 43.1). Willdenow’s ee e cited Anonvni mos Grameen. . Walt. oni 197.” and “Habitat in Carolina. 4 ake dit fully quoted Walter's description. Despite Willdenow’s apparent sug- gestion that hes ea corresponding to Walter’s type has not been erie Gaiser (1946, p. 255) noted that Bbeervatione had been une ona BM speci- men labeled “Chrysosoma aints E a0? Stopes au) fo ane to Fraser) and with Nuttall’s annotation Liatris in pencil,” 1, but she later rejected this i in favor of a hesepae The only eeecmnes in the Willdenow her- barium identified as Liatris graminifolia (B-Willdenow fiche 14838!) is a plant of Liatris spicata (L.) Willd. var. spicata with a label that notes “Habitat in Pensylvania;” the label also cites “Anonymos graminifolia W. carol. 197,” but it seems unlikely that Willdenow would have intended this collection as the type for L. graminifolia, which he explicitly understood was from “Carolina.” acest WPC. Barton, Veg. Mater. Med. US. 2:223, t. 49.1819. Liatris graminifolia Willd. var. a (WPC. Barton) A. Gray, Manual, ed. 2, 185. 1856. TyPE: U.S.A. [perhaps NEW JERSEY or DELAWARE, in the region where plants of this morphology occur]. This is the only element of potential type material used by the author and presum aby st bans as one HOLOTYPE (ICBN 2000, Art a No collection data were cited ae Barton, a Phil I ident, apparently drew tl ion f live plant, j ic di ionality of the draw- ing, even though the sa Bee and slonsste involucres suggest that considerable artistic license was in play. The description and illustration portray a plant with hairy stems and peduncles, linear-lanceolate lower leaves, and loosely arranged, long- a enad heads with elongate, “subacute” ae es Sey) mentioned” var. aunigs angers gramintfol ia, but he did not provide a basis of ref 1856. Fernald and Griscom (1935) ae that L. dubia was “suggestive of Aiton’s plant” (ie., L. pilosa s. ce Liatris ae (Aiton) Willd. var. laevicaulis DC, Prodr. 5:131. 1836. TYPE: U.S.A. Nov. Caesar. [No Cae = NEw JERSEY]. 1835, [no other data,] Mr. Torrey (HOLOTYPE: G-DC, fiche!). This one on ong ee ae ntly glabrous stems, long, narrow phyllaries apparently with sub- acute apices, and (fide de Candolle) 7-8 florets per hea ee ua Hoo ck , Bot. Mag. 67 (n. ser. 14): t. 3829. 1840. TyPE:[U.S.A., pea ede England) rega ard to its origin, Hooker noted only “Sent from the Horticultural ae urgh in the autumn of 1839, under the name i i Parana ihe illustration shows a plant with ies turbinate-cylindric ly spiciform array, acute phyllaries, and ancihae pean Banca cans ean | ae on the proximal margins. The stems and le he common name a oker to a plant, “Sharp-scale spiked I jatris? ‘referred to the osa, especially in view of the = giv acute phyllaries. The eee seems reasonable as L. pi prominently cilate leaves, but the sparsely pubescent (or glabrous?) stems, acute phyllaries, and relatively — florets (‘subdecemfloro,” from the description) leave open the possibility that it might be L. virgata. The name (L. propinqua) is ambiguous in reference until a type s nis ae or designated. Pein a ia Willd. var. lasia Fernald & Griscom, Rhodora 37:183. 1935. TYPE: U.S.A. NEW Jersey. Camden Co. Lindenwold, dry sandy soil, 29 Sep 1923, J.M. Fogg, Jr 622 (HOLOTYPE: GH). Flowering (Aug-)Sep-Oct(-Nov). Old fields, pine barrens, scrub oak-pine sandhills, openings in pine, oak, and oak-hickory woods, tidal marsh edges, sandy fields, dune hollows, wet sand near beach, edge of tidal marsh sand to 824 BRIT.ORG/SIDA 21(2) sandy clay-loam; ca. (O-)10-500 m. Delaware, Maryland, New Jersey, North Caro- lina, Pennsylvania, South Carolina, Virginia. Liatris elegantula (Greene) K. Schum., Bot. Jahresber. Just) 29:569. 1903. Lacinaria elegantula Greene, Pittonia 4:316. 1901. Liatris graminifolia Willd. var. elegantula (Greene) Gaiser, Rhodora 48:254. 1946. TyPE: U.S.A. ALABAMA. Lee Co.: Auburn, 18 Oct 1896, FS. Earle (HOLOTYPE: ND-G) Flowering Aug-Oct(-Nov). Longleaf pine-scrub oak, pine, live oak-pine, decidu- ous oak-pine, deciduous flatwoods, sandhills, savanna edges, edge of cypress depressions, depression meadows, live oak-pine-palmetto hammocks, sandy clay or loam, rarely clay; 5-300(-450) m. Alabama, Florida, Georgia, Mississippi. Liatris virgata Nutt.,J. Acad. Nat. Sci. Philadelphia 7:72. 1834. Liatrisgraminifolia Willd. var. virgata (Nutt.) Fernald, Rhodora 51:104. 1949. TyPE: U.S.A. GEORGIA. [probably Nov 1815], T. Nuttall s.n. (LECTOTYPE (Stucky 1992, p. 179): PH!; ia type material, “Hb. Nuttall” s.n., NY! M). Nuttall noes ‘Hab. In Georgia and North Carolina” in the protologue; the PH speci- men ieee ‘Geo.” as the only collection data. ee later (1841) described the habitat as “In the pine forests ‘i Georgia, and near Newbern, N. Carolina’—the plants from b are almost certainly Liatris cokeri Stucky & Pyne (see Stucky & Pyne 1990). ae nee PP. te 101) noted mae in aud October, 1815, Nuttall traveled by boat to Savannah, Georgia, h River to Augusta and vicinity, through longleaf pine sandhill vegetation and north at least to “where hills of deciduous trees (oaks, hickories, &) and primitive soil commence.” Nuttall’s protologue noted that the arenes was a sean ors and branched raceme and referred to the “long leafy oo of ie wers.” Gaiser (1946) apparently saw the PH specimen (she referred to it as “isotype”) a anes L. virgata : synonymy of L. graminifolia var. dubia, but the latter is oe meee as a syn- nye of pi ngETs pilosa’ sensu stricto. Fernald’s concept of L. graminifolia var. es li £ Ae - 50 Atlantic coast region an strongly branched een though he surely was aware that the type was sive Georgia, having indicated that he saw the Nuttall collection or at least a photo of it. Details on morphology of the PH specimen are provided in Stucky 992). Liatris spicata L. var. racemosa DC.,, Prodr. 5:130. 1836. Liatris graminifolia Willd. var. racemosa ( Venard, Rhodora 51:35. 1949. Type: U.S.A. GEORGIA, Savannah, 1832, no collector in cated (HOLOTYPE: G-DC, fiche!). The saeaue ges by: de Candolle noted “capitulis antic pedicellatis...caule glabro... foliis ciliatis 1.8.” The G-DC sheet has 2 branches with heads of each, identical morphology: one was broken off and the heads are borne on peduncles up to 5cm long; the other is intact and the distal heads are sessile while the proxi- mal ones are on peduncles ca. | cm long. The involucres are campanulate-cylindric, and the payllanes are e apically thickened and subacute sso outer aie spreading, DK.Schum., Bot. Jal Small Bull. toute Bot. Club 25:473. 1898. TYPE: USA. NorTH CAROLINA. Cleveland fon King’s Mt., wooded slopes, 27-30 Aug 1894, J.K. Small s.n. (HOLOTYPE: NY! ISOTYPE: ee a Man. FI. N. States 927.1901. Liatrisgraminifolia Willd. var. smallii (Brit- Rhodora 37:182. 1935. Type: U.S.A. VIRGINIA. Smyth Co. along Dickey Creek on Iron Mtn., 2900 ft, 8 Aug 1892, J.K. Small s.n. (HOLOTYPE: NY! ISOTYPE: MON). —, Flowering Jul-)Aug-Oct(-Nov). Edge of swampy woods, creek margins, slopes, clearings, and edges of upland woods, rocky woods, pine-oak woods, mixed de- NESOM AND STUCKY, TAXONOMY OF LIATRIS PILOSA COMPLEX 825 ciduous woods, roadsides, Iredell soil, clay; ca. 50-1000 m. Georgia, North Caro- lina, South Carolina, Virginia. Liatris cae Pyne & Stucky, Sida 14:205. 1990. Type: U.S.A. NORTH CAROLINA. Harnett Co: 0.2 mi E jet NC Rte 27 and Co. Rd. 1243 along NC 27 on S side of road, sandy roadside and margin oF longleaf pine/turkey nee a association, 23 Sep 1989, J.M. Stucky 511 (HO- LOTYPE: NCU; ISOTYPES: GH, NCSC, NCU, NY, US, USCH). Flowering (Aug-)Sep-Oct. Sand ridges, sy fields and roadsides, turkey-oak, longleaf pine-oak; 50-150 m. North Carolina, South Carolina. ACKNOWLEDGMENTS The analysis and map for this study were based on specimens were studied from BRIT, NCU, TEX,LL, and VDB (personal visits) and DOV, PH, USCH, and BM (loans). We are grateful to staff at the University of Texas Life Sciences Library for help with access to literature and microfiche, David Snyder, Botanist for the New Jersey Natural Heritage Program, for his insights regarding variation in Liatris pilosa, Walter Kittredge and other GH staff for help with data and type images, Milo Pyne for his perspectives on the identity of Liatris pilosa, and to Alan Weakley and Bruce Hansen for their review comments. REFERENCES Britten, J. 1912. The history of Aiton’s ‘Hortus Kewensis.’ J. Bot. 50, No. 599 (suppl.):1-16 Cronauist, A. 1980.Vascular flora of the southeastern United States, Vol.|. Asteraceae. Univ. of North Carolina Press, Chapel Hill. Fernatp, M.L. 1949. Studies of eastern American plants. Rhodora 51:43-57;61-85; 93-104. Fernaco, M.L. 1950. Gray's manual of botany (ed.8). American Book Co., New York. FerNatb, M.L.and L.Griscom. 1935. Three days of botanizing in southeastern Virginia.Rhodora 37:181-183. Galser, L.O. 1946. The genus Liatris. Rhodora 48:165-183, 216-263, 273-326, 331-382, 393-41 2. Gaiser, L.O. 1950. Chromosome studies in Liatris. ll. Graminifoliae and Pauciflorae. Amer. J. Bot. 37:414-423. Graustein, J.E. 1967. Thomas Nuttall, Naturalist: Explorations in America 1808-1841. Har- vard Univ. Press, Cambridge, Mass Gray, A. 1848. A manual of the botany of the northern United States (ed. 1). James Munroe and Co., Boston & Cambridge. Gray, A. 1856. Manual of the botany of the northern United States (ed.2). Ivison & Phinney, New York. Gray, A. 1884. Synoptical flora of North America. Vol.|.— Part Il Ivison, Blakeman, Taylor,and Co., New York. HarsHBERGER, J.W. 1917. William Young, Jr, of Philadelphia, Queen's botanist. Torreya 17: 91-99 826 BRIT.ORG/SIDA 21(2 — KeLLer, |LA.and S. Brown. 1905. Handbook of the flora of Philadelphia and vicinity. Philadel- phia Botanical Club, Philadelphia, Pa. Marsuatt, J.B. 1978. The handwriting of Joseph Banks, his scientific staff and amanuenses. Bull. Brit. Mus. (Nat. Hist.), Bot. 6:1-85. Nurtatt, 1. 1841.Descriptions of new species and genera of plants in the natural order of the Compositae. Trans. Amer. Philos. Soc., ser. 2, 7:284. Raprorb, A.E., H.E. AHtes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill. Stucky, JM. 1991. Affinity of Liatris cokeri Pyne & Stucky (Asteraceae), a sandhills endemic of the Carolinas, and its widely distributed relative, L. graminifolia Willd. Amer. Mid. Naturalist 125:323-330. Stucky, JM. 1992. Liatris virgata (Asteraceae) in the southeastern United States. Sida 15: 177-183 Stucky, JM. and M. Pyne. 1990 A new species of Liatris (Asteraceae) from the Carolina sandhills. Sida 14: 189-208 Wieur, R.L. 1962. The identity of Walter's species of Anonymos. J. Elisha Mitchell Sci. Soc. 78:125-131. WUNDERLIN, R.P. and B.F. Hansen. 2004. Atlas of Florida Vascular Plants [S.M. Landry and K.N. Campbell (application development), Florida Center for Community Design and Re- search.]. Institute for Systematic Botany, Univ. of South Florida, Tampa. Youne, W. 1985, Herbarium specimens (microfiche). Early American herbaria and related drawings from the British Museum (Nat. Hist.). Meckler Publ., Westport, Conn. A NEW SPECIES OF SYMPHYOTRICHUM (ASTERACEAE: ASTEREAE) FROM A SERPENTINE BARREN IN WESTERN NORTH CAROLINA Gary L. Kauffman Guy L.Nesom U.S. ai ieee ae Botanical Research Institute of Texas (BRIT) 509 Pecan Street Asheville, ee ae 28802, U.S.A. Fort Worth, Texas 76102, U.S.A. Alan S.Weakley Thomas E.Govus University of North Carolina Herbarium (NCU) 3711 Big Creek Road North Carolina Botanical Eaiden Fllijay, Georgia 30540, U.S.A. Campus Box #3280 Chapel Hill, North Carolina 27599, U.S.A. Laura M.Cotterman North Carolina Botanical Garden R 5 amnus Box #33 i Chapel Hill, North Carolina 27599, U.S.A. ABSTRACT Symphyotrichum rhiannon Weakley & Govus, sp. nov, is described from the Buck Creek ultramafic barren (over serpentinized dunite and olivine) in the Blue Ridge province of southwestern North Carolina. It is similar to Symphyotrichum puniceum but different in its smaller stature, thinner rhi- zomes, subspatulate cauline leaves, narrower and fewer-headed capitulescence, phyllaries with shorter, rhombic-lanceolate apical green zones, and shorter ray corollas. Its geographic range is im- dded within that of typical S. puniceum but morphological intergrades have not been observed, despite the co-occurrence of the two taxa within several meters of one another at the type locality. The Buck Creek site is within the Nantahala National Forest; 42 hectares of the site are managed by the United States Forest Service as a Special Interest Area and registered as a heritage area with the North Carolina Natural Heritage Program RESUMEN ee ae rhiannon Weakley & Govus, sp. nov, se describe del Buck Creek ultrabasico ae a y olivino serpentinizados) en la provincia Blue Ridge del Suroeste de Carolina del N ae a Symphyotrichum puniceum pero diferente por su talla mas baja, rizomas mas hoe Sc eamui Sue pa aca Ca ee mas ce y con menos aes: filarias con zonas rea Ide S. puni ha ol 1 lacioé morfoldgica a a pesar de la co-ocurrencia de los dos taxa a unos ces metros en la tecalicad tipo. El Buck Creek esta en el Nantahala National Forest; 42 hectareas estan gestionadas por el Servicio Forestal de los Estados Unidos como un Area de Especial Interés y registrado como un area del Natural Heritage Program de Carolina del Norte. SIDA 21(2): 827-839. 2004 828 BRIT.ORG/SIDA 21(2) INTRODUCTION A new species of Symphyotrichum is described from the Buck Creek ultrama- fic barren in the Blue Ridge province of southwestern North Carolina (Clay County). The naming of this species ends more than twenty years of bewilder- ment regarding its identity, which has troubled researchers at the site since the late 1970's. Mansberg (1981) was apparently the first botanist to encounter this entity and collect it, a significant component of the ultramafic barren and ad- joining woodlands. Despite consulting with experts at the time, she labeled this entity as “unidentifiable aster” and suggested that it might be a hybrid. Collec- tions made during later studies of the vegetation at Buck Creek barren by the North Carolina Vegetation Survey in the 1990s yielded similar confusion, with specimens being identified as “Aster laevis var. concinnus?,” “Aster surculosus?,” or “Aster # 1” Even more recently, U.S. Forest Service botanists involved in the management of Buck Creek barren have been troubled by the identity of this aster and have made additional guesses as to its affinity. Serendipitous circum- stances have led toa revisiting of this persistent problem, and following addi- tional study we now decisively describe this species as new. A comparison of the putative new taxon to all other species of Symphyotrichum in eastern and central North America reveals that it has a unique set of characteristics. Furthermore, it does not appear to be a recent or stabilized Fl hybrid, intermediate between any other two species of Symphyotrichum, although it is possible (as noted below) that gene flow has been involved in its origin. In our assessment, it is a distinct taxon worthy of specific rank, possibly most closely related to Symphyotrichum puniceum (L.) A.& D. Love. Symphyotrichum rhiannon Weakley & Govus, sp. nov. (Fig. 1). Type: U.S.A. NorTH CAROLINA. Clay Co.: Buck Creek Ultramafic Barrens, Nantahala National Forest, 1.3 km N of S 64 on es Creek Rd, just NE from Glade Gap, below FS Rd 6269 (above bridge crossing Buck Creek), NW ca. 0.5 km from gate on steep slope above E side of Buck Creek; serpentine barrens, steep W aspect outcrop of dunite and olivine, open woodland dominated by pitch pine and prairie grasses, 3385 {t elev, UTM zone 17S, 261513mE, 3885663mN (WGS-84), 3 Nov 2003, TE. Govus and G. Kauffman 1 (HOLOTYPE: NCU; ISOTYPES: BRIT, US, WAT) i, similis sed differt statura minore, rhizomatibus tenuioribus, foliis caulinis see aR angustioribus paucicapitatis, phyllariis zonis apicalibus viridibus rhombi-lanceolatis brevioribus, et corollis radii brevioribus. Plants perennial herbs, apparently without a distinct caudex, arising from a system of slender, scale-leaved rhizomes 0.5-1 mm wide and up to 10 cm long, older rhizomes woody or lignescent, thickened to 2-4 mm wide. Stems 15-40 cm tall, erect from the base, hirsute to hispid-hirsute with spreading to spread- ing-ascending, uniseriate trichomes 0.2-0.6 mm long (Type A, sensu Nesom 1976), evenly distributed or concentrated in vertical lines, lines especially evi- dent immediately below nodes, hirsutulous below heads, also with closely ap- KAUFFMAN ET AL.,A NEW OF EVIL VE Studies in Aster G, Nesom 2003 University of North Carolina Herbarium Symphyotrichum species novum Weakiey et al. Asteraceae Buck Cre ates, RUbiannon's Aster Buck Creek Utramafic Barrens. Nantahata piomatpaltinis {3 km N of US $4.00 frrotigcbiag Just northoest From Giada Gap, preteen rset tte! northwest trom gate on steep slope Serperti Appalachians Characters: Ue sore ae ch Ser —— om in beagnt UTM zone 17S, 281513mE 26aSeR MN (vs Sev. 3388 6. } icf hi, A Holotvpe: B. Cl YRS, 830 BRIT.ORG/SIDA 21(2) pressed, uniseriate trichomes 0.1-0.2 mm long [Type B, sensu Nesom 1976), eglandular. Leaves: basal and lower cauline usually not persistent at flowering, subspatulate when present, 3-7 cm long, blades oblanceolate-elliptic, 10-15 mm wide, shallowly crenate to serrate-crenate, prominent midvein and reticulate secondary venation, gradually narrowed to a petiolar region 4-6 mm wide, clasping to subclasping and shallowly auriculate, midcauline usually longer than lower, 5-11 cm long, 8-22 mm wide, with petiolar region remaining evi- dent but becoming shorter and relatively broader or the upper leaves oblong- lanceolate, upper cauline (of capitulescence) 1-3 cm long, 4-6(-8) mm wide, all adaxially scabrous with stiff, ‘thick- based, ascending hairs 0.1-0.2 mm long (foreshortened Type A), abaxially i y strigose (use lens) with thin, closely appressed trichomes 0.05-0.2 mm long (Type B). Heads in a sparsely leafy-bracteate, broadly cylindric-paniculate to subcorymboid arrangement, clustered near branch tips on ultimate peduncles 5-15(-30) mm long; involu- cres turbinate-campanulate, 6-11 mm wide (pressed); phyllaries in 3-4 weakly graduate series, mostly narrowly oblong-lanceolate, inner 6-7 mm long, outer 1/2-3/4 the length of the inner, outer white-indurate on the basal 1/2-2/3, upper 1/2-1/3 witha rhombic-lanceolate green zone 1.8-2.5(-3.0) mm long from widest point to tip, glabrous or the upper part of the green zone minutely stri- gose-puberulent, margins usually irregularly ciliolate, eglandular. Ray florets 18-32, corollas 6-9 mm long, laminae 0.8-1.4 mm wide, blue to lavender, tightly coiling with wilting or maturity, tube and lower lamina sparsely appressed- puberulent with minute biseriate trichomes (Type C, sensu Nesom 1976). Dise florets: corollas (4.0-)4.5-5.5 mm long, yellow but at least the lobes usually turn- ing purple, sparsely appressed-puberulent with biseriate trichomes (Type C), tube 1.5-2.1 mm long, throat and limb cylindric, lobes 0.8-1.0 mm long, erect: style branches with narrowly triangular collecting appendages 0. a0 5mm long. Achenes oblong to narrowly obovate in outline, slightly compressed, 2.6-3.0 mm long, 0.7-0.9 mm wide, 4-6-nerved (with 1 nerve on each edge and 1-2 on each face), sparsely short-strigose, tan or often purplish at maturity; pappus l-seriate, of 36-46 barbellate bristles. Etymology.—Symphyotrichum rhiannon is named in honor of Rhiannon Weakley, whose desire to rest during a field excursion led the authors to further investigate and finally resolve this decades-old taxonomic conundrum, and also in honor of the original Rhiannon, a Welsh goddess figure associated with the underworld, and therefore particularly appropriate for a plant endemic toa ser- pentine substrate. = Additional collections examined. NORTH CAROLINA. Clay Co.: Buck Creek Ultramafic Barrens, Nantahala National Forest, 1.6 km N of US 64 on Buck Creek Rd, NE from Glade Gap, 0.5 km below bridge over Buck Creekand W upslope along old road to powerline ROW, E aspect slope of dunite and olivine, pitch pine woodland with abundant prairie grasses, 3280 ft elev, UTM zone 17S, 261347mE, 3885588mN (WGS-84), 3 Nov 2003, Govus and Kauffman 2 (NCU); Buck Creek Serpen- KAUFFMAN ET AL., 831 tine Pine Barrens; in grassy understory, west- mens slope, midslope; pH 6.5, 20 Sep 1981, Mansberg 421 (NCU 575098); Buck C kS uncesta ue wesl mcngs) ad lower slope; pH 6.0, 20 Sep 1981, Men ber 422 (NCU 575100); Back Creel understory; west-facing slope, midslope; pH 7.0, 20 Sep 1981, Mansberg 423 (NCU 575099). DISCUSSION Symphyotrichum rhiannon is closest in morphology to S. puniceum and S. prenanthoides. These three taxa share a set of features: plants eglandular, stoloniferous; leaves with serrate margins, scabrous upper surfaces, the basal petioled but not cordate, cauline clasping, those of capitulescence reduced in size and relatively few; phyllaries long-acuminate to attenuate, subequal to weakly graduate in length. The leaves of S. prenanthoides are distinctly spatu- late with dilate-auriculate bases, which contrast with both of the other spe- cies, and the new species keys to S. puniceum (Radford et al. 1968; Gleason & Cronquist 1991; Cronquist 1980) because of the leaf shape. Of features noted ere as characteristic of S. rhiannon, only its subspatulate leaf shape appar- ently is not found inat least a few populations of S. puniceum. Thus, S. rhiannon might be considered intermediate between S. puniceum and S. prenanthoides, either from common ancestry or perhaps a result of hybridization. Contrasts between these three taxa are summarized in the following couplet. . Basal and cauline leaves distinctly spatulate, abruptly constricted to a petiolar re- gion, auriculate-clasping, the auricles dilated from the petiole base and completely enveloping stem; lower 1/2-1/3 of stem generally glabrous Symphyotrichum prenanthoides lanceolate, gradu- ally narrowed or not toward the oe base aeeoing to subclasping but not di- lated-auriculate and not completely enveloping the stem; lower 1/2-1/3 of stem 1 Gauli | kK tr ilat KI + hI hI lat 2. Plants 1.5-4 dm tall; young rhizomes thin, 0.5-1.0 mm wide; midcauline leaves subspatulate, slightly narrowed toward the base to a petiolar region; green area of phyllary rhombic-lanceolate, 1.8-2.5(-3.0) mm long from widest point to apex; ray corollas zn mm long Symphyotrichum rhiannon 2: Plants mostly 4-20 t f ickene d, 2-4mm wide; midcauline leaves mostly oblong to oblong- oblanceolate, not narrowed to a petiolar re- gion; green area of phyllary apices narrowly lanceolate, 2.5—4.0 mm long from widest point to apex; ray corollas 10-15(-20) mm long _____ Symphyotrichum puniceum Apart from the distinction in leaf shape, differences between Symphyotrichum rhiannon and S. puniceum are largely quantitative and are primarily reduc- tions in size. Stems of the new species are relatively short and the capitulescence tends to be narrower with fewer heads. Phyllaries with short green regions are not generally characteristic of S. puniceum but do occasionally occur in the species, but the rhombic-lanceolate green zones and more graduate phyllaries of S. rhiannon give the headsa distinctive appearance. The cumulative effect of these and the more quantitative differences, however, coupled with the unique 832 BRIT.ORG/SIDA 21(2) habitat and restricted distribution of S. rhiannon, is that the latter can be rec- ognized as sharply distinct. Typical S. puniceum occurs in most if not all of the counties in the mountains and piedmont of North Carolina (Radford et al. 1968), where it characteristically grows in wet habitats, especially fens, seepages, swamp forests, wet meadows, and in altered wetlands such as ditches and other drainages. Fernald (1950) treated Symphyotrichum puniceum (as Aster puniceus L.) with a number of varietal taxa, but all of these are now recognized as popula- tional variants or intergrading populations except one — Warners and Laughlin (1999) have provided a convincing case for treatment of S.firmum (Nees) Nesom as a distinct species, in contrast to many earlier treatments of it as Aster puniceus var. firmus (Nees) Torr. & Gray or A. puniceus var. lucidulus Gray. Evidence has been presented for recognition of S. puniceum var. scabricaule (Shinners) Nesom (Nesom 1997), the only infraspecific taxon currently recognized within the spe- cies. It might be argued that S. rhiannon would be more appropriately treated at varietal rank, especially in view of its largely quantitative difference, but we infer that the new species is reproductively isolated because its geographic range is imbedded within that of typical S. puniceum and morphological intergrades have not been observed, despite the co-occurrence of the two taxa within sev- eral meters of one another at the type locality. It is possible and likely that adaptation to the relatively drier habitat of the serpentine barren play a significant part in the observed size reductions of Symphyotrichum rhiannon. Common garden experiments would be of interest to determine what component, if any, of the differences are phenotypic. The differences in leaf shape, however, suggest that other genomes might have been involved in the evolution of S. rhiannon. Symphyotrichum prenanthoides, which apparently is closely related and which is sympatric with S. puniceum over much of its range, has cauline leaf blades distinctly narrowed to a petiolar re- gion and Semple (pers. comm.) hypothesizes that S. rhiannon is most closely related to S. prenanthoides, apparently weighting the tendency in S. rhiannon to produce subspatulate leaves. Symphyotrichum rhiannon, S. prenanthoides, S. puniceum, and S. firmum have been treated as members of Symphyotrichum sect. Symphyotrichum (sensu Nesom 1994), but because hybrids in many pa- rental combinations have been observed in Symphyotrichum, hypotheses re- garding evolutionary ancestry could justifiably include species of putatively more distant relationship. It is perhaps notable that the stem leaves of S. rhiannon are less strongly clasping than those of either S. puniceum or S. prenanthoides. Symphyotrichum rhiannon has only been documented within a serpen- tine plant community endemic toa 120 ha area surrounding Buck Creek in the southern Nantahala Mountains of Clay County, North Carolina (Fig. 2). Ser- pentine soils in the Southern Appalachian Mountains are very rare, and only a KAUFFMAN ET AL.,A WEY OFEAILO VE 833 le ire Ee Buncombe f ASHEVILLE f ROBSINSVILLE Blue Ridge Parkway (J County a City Cx USFS 60 0 60 120 Kilometers Great Smoky Mountains few serpentine barrens have been located and studied, beginning with a pio- neering study by Albert E. Radford (1948). The Buck Creek Serpentine Barren was discovered in 1951, following Radford’s dissertation, by the late Robert K. Godfrey, then at N.C. State University, and has been periodically studied by bota- nists and ecologists ever since, as it is the largest and floristically and vegetationally most distinctive of the Southern Appalachian serpentine bar- rens. In the late 1970s and early 1980s, Laura Mansberg (now Cotterman) con- ducted the most detailed study of this unique plant community, describing it as a Pine-Savanna (Mansberg 1981; Mansberg & Wentworth 1984). The unusual aster here described as Symphyotrichum rhiannon was initially noted during this study by Mansberg, and was referred to in her thesis and subsequent paper as “an undescribed Aster.” Schafale and Weakley (1990) classified the Buck Creek site as an Ultramafic Outcrop Barren, and considered it to be the best-devel- oped and largest example of this critically imperiled natural community type. The hierarchical United States National Vegetation Classification classifies the community association type as a Quercus alba-Pinus rigida / Sporobolus heterolepis- Andropogon gerardii Woodland and assigns it a conservation rank of Gl (Critically Imperiled) (NatureServe 2004). A physiognomic patchwork of forest, dense grass patches and partially 834 BRIT.ORG/SIDA 21(2) open woodland occurs across the Buck Creek site (Fig. 3). The dominant rock types, serpentinized dunite and olivine, influence the striking vegetation present on this site (Hadley 1949; Pratt @ Lewis 1905). Soil depth is variable, ranging from 0 to 60 cm, although rock outcrops represent between 5 and 10% of the local landscape. Soil characteristics reveal higher base saturation, cation exchange capacity, pH, and magnesium relative to surrounding sites in the Nantahala Mountains (Mansberg & Wentworth 1984). The serpentine plant communities occupy both east and west-facing slopes extending from 975 meters elevation along Buck Creek to over 1220 meters elevation atop Corun- dum Knob. Symphyotrichum rhiannon occurs within the woodland and grass- dominated areas. Within the woodland the forest canopy varies from 20-60% cover depend- ing on the intensity and frequency of recent prescribed burns. The woodland is dominated in the tree canopy by older stunted Quercus alba and smaller denser Pinus rigida stems. Tsuga canadensis, Sassafras albidum, Acer rubrum, Oxydendrum arboreum, and Amelanchier laevis also occur within the canopy and subcanopy. Shrub cover is meager, typically occurring in clumps and pro- viding no more than 10% cover. Diagnostic shrub species include Rhododen- dron viscosum, Physocarpus opulifolius, Viburnum cassinoides, Kalmia latifolia, Lyonia ligustrina var. ligustrina, Vaccinium stamineum, and V.corymbosum. The grass dominance within the herb stratum is reminiscent of prairie veg- etation and presents a striking contrast to the regionally typical herbaceous layer consisting of mesophytic, broad-leaved forbs. Andropogon gerardii, Schizachyrium scoparium var. scoparium, and Sporobolus heterolepis are the most important grasses. Characteristic forb species include Hexastylis arifolia var. ruthii, Thalictrum macrostylum, Packera plattensis, Castilleja coccinea, Phlox ovata, Oxypolis rigidior, Sanguisorba canadensis, Polygala pauciflolia, Asplenium platyneuron, Solidago nemoralis, Symphyotrichum undulatum and S. phlogifolium. A striking contrast within the herb layer is the juxtaposition of both mesophytic and xerophytic species. It is not unusual to observe Sanguisorba canadensis and Oxypolis rigidior emerging from a grassy thicket of Sporobolus heterolepis and Schizachyrium scoparium. Mansberg noted a perched water table while surveying the site and suggested that there is a com- plex soil moisture gradient within the serpentine site (Mansberg & Wentworth 1984). Many of the minerals predominant in mafic and ultramafic rocks chemi- cally weather to clays, and the soils at Buck Creek have a substantial clay com- ponent, providing a perched water table and abundant seepage after rains, but drying to highly xeric conditions during droughts. In response to periodic mining threats, 42 hectares of the serpentine site at Buck Creek are now managed by the USFS as a Special Interest Area and regis- tered asa Natural Heritage Area with the North Carolina Natural Heritage Pro- gram. In 1995, the U.S. Forest Service initiated active conservation management 835 KAUFFMAN ET AL Fic. 3.B D - canopy layer. BRIT.ORG/SIDA 21(2) Taste 1. Status of North Carolina rare plant species documented at Buck Creek serpentine site. Species G-rank* S-rank? Relative Rarity within North Carolina Sporobolus heterolepis GS 51 1 of 2 occurrences documented in NC; disjunct eastwards from a primary distribution in the Great Plains Deschampsia cespit p. glauca G5 S] Only known occurrence in NC; disjunct from further west Elymus trachycaulus ssp. trachycaulus G5T5 S1 Only known occurrence in NC; disjunct from further west Calamagrostis porteri ssp. porteri G4T4 S1 1 of 8 extant occurrer ithin NC Poa saltuensis G5 S1 1 of 2 occurrences cccamenica in NC; disjunct from further west Muhlenbergia glomerata G5 51 1 of 2 occurrences documented in NC; disjunct from further west Carex woodii G4 S2 Dense in forested area within and surrounding serpentine site; northern species near its southern range extent Calystegia catesbiana var. sericata G3T3 $3 Locally common in Southwestern NC Mts; broad Southern Appalachian endemic Liparis loeselii G5 S1 Locally restricted to only a few individuals; sel aa near its southern range Gentianopsis crinita G5 S1 1 of ; high ie occurrences in N Oenothera perennis G5 $2 ot high quality occurrences in N Ranunculus fascicularis G5 S1 1 of 5 occurrences documented in NC Pedicularis lanceolata G5 S] 1 of 2 drainages with extant populations in Parnassia grandifolia G3 $2 1 of 3 high quality occurrences in NC Viola appalachiensis G3 $2 Largest population documented in NC; Central and Southern Appalachian endemic Brachyelytrum arist G4G5 532 Locally common in higher (= B. septentrionale) elevations in the southwestern Mountains of NC; northern species near its southern range extent Drepanolejuenea appalachiensis G2? S2 Remarkable occurrence for tiny liverwort that more typically occurs in mesic hardwood forest; Southern Applachian endem with one disjunct occurrence in West Indies KAUFFMAN ET AL.,A NEW OF EID UF 837 TasLe 1. continued Species G-rank? S-rank® Relative Rarity within North Carolina Celastrina nigra G4 $2? 1 or 6 occurrences documented in NC, near eastern range limit Speyeria aphrodite cullasaja G5T1 S1? Only known occurrence in NC; a Appalachian endemic Chlosyne gorgone GS Si? Only known occurrence in NC primarily Midwestern and western species near eastern range limit Phyciodes batesii maconensis G5 S2 Largest population documented in NC; southern Appalachian endemic 8G-rank indicates global rarity and threat status. See NatureServe (2004) for definitions. »S-rank indicates state rarity and threat status. See NatureServe (2004) for definitions. of the site, using prescribed fire as the primary tool, resulting in reduction of woody growth encroaching on the site and an increase in herbaceous cover. particularly the grasses once dominant at the site (Fig. 4). Within the last 10 years monitoring plots were established both by both the North Carolina Veg- etation Survey and the U.S. Forest Service to document current vegetation and provide a baseline for detecting change. A review of this plot data shows that S. rhiannon prefers more open conditions, occurring exclusively within plots (n= 27) with less than 50% tree cover and primarily in plots with less than 33% canopy cover. S. rhiannon grows sparsely; cover estimates in plots have not exceeded 2% cover. S. rhiannon has either maintained its coverage or slightly increased its den- sity within fire-maintained areas at the Buck Creek Serpentine Barren. In addition to the presence of the unusual plant community, the presence of rare species adds to the conservation importance of the Buck Creek Serpen- tine Barren. Seventeen state-listed rare plant species and four state-listed but- terfly species occur within the site (Franklin & Finnegan 2004; LeGrand et al. 2004; Gatrelle 1998; Table 1). Most of these species are primarily wide-ranging and globally secure (G4 or G5 rank), yet rare in NC; a few are restricted to only this site within the state. Although serpentine is well known for its tendency to generate locally endemic species (Brooks 1987), relatively few endemics have been described from the serpentine areas in eastern North America. A number of endemics have been described from the well known serpentine barrens of Maryland and Pennsylvania, ine maing Symphyotrichum depauperatum (Fernald) Nesom, though the taxonomic d tiveness of many remains con- troversial. The less well known serpentine areas in Virginia, North Carolina, and Georgia pais? not heretofore yielded itel ule Cee clsedleee However, a number of odd } ive taxa (Carex, Hexastylis, Rhod Symphyotrichum, 838 BRIT.ORG/SIDA 21(2) Thalictrum) from serpentines in Virginia and North Carolina are currently the subject of taxonomic investigation and may add to a growing list of eastern North American serpentine endemics. Current information suggests that S. rhiannon is a serpentine endemic re- stricted to the Gl-ranked woodland community at the Buck Creek Serpentine Barren. We hope that publication of this species will result in the discovery of other populations, at the few other Southern Appalachian serpentine barrens, though the potential for locating other undocumented occurrences within ul- tramafic rock influenced woodlands or barrens within the region is probably limited given the overall scarcity of this habitat. ACKNOWLEDGMENTS We thank Luc Brouillet (Université de Montréal, MT), co-author of the taxo- nomic treatment of Symphyotrichum for the forthcoming FNA volumes (Flora of North America), for his comments on the distinctiveness of the new species. We thank John Semple (University of Waterloo, WAT) for helpful discussions about the taxon and comments improving the manuscript. We thank the cura- tors of GMU and VPI for the loan of specimens of other Symphyotrichum enti- ties found in ultramafic and mafic sites in Virginia. REFERENCES Ba.taro, TJ. 1947. Report of investigations on Buck Creek Corundum, Clay County, North Carolina.U.S. Department of Interior, Bureau of Mines, R.1. 4052. Brooks, R.R. 1987.Serpentine and its vegetation: A multidisciplinary approach. Dioscorides Press, Portland, Oregon. 454 pp. Cronauist, A. 1980.Vascular flora of the southeastern United States. Vol.1.Asteraceae. | North Carolina Press, Chapel Hill. Fernato, M.L. 1950. Gray’s manual of botany (ed. 8). American Book Co., New York, NY. FRANKLIN, M.A.and J.T. FINNEGAN. 2004. Natural Heritage Program list of the rare plant species of North Carolina. North Carolina Natural Heritage Program, Raleigh, North Carolina. 112 pp. Garrete, R.R. 1998. Two new Nymphalidae from western North Carolina: new subspecies of Speyeria aphrodite and Phyciodes batesii. The Taxonomic Report of the International Lepidoptera Survey volume 1,number 3, 7pp. Gteason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada (ed. 2). The New York Botanical Garden, Bronx, NY. Habe, J.B. 1949. Preliminary report on corundum deposits in the Buck Creek peridotite, Clay County, North Carolina. Washington, D.C. U.S. Department of Interior, Strategic Minerals Investigations 1945, Bulletin 948-E. LeGRAND, H.E., S.E. McRae, S.P. Hatt and J.T. Finnecan. 2004. Natural Heritage Program List of the rare animal species of North Carolina. North Carolina Natural Heritage Program, Raleigh, North Carolina. 99 pp. niv KAUFFMAN ET AL.,A NEW OFELILOS VI 839 Manseerc, L. 1981. Vegetation, soils, and canopy age-structure of the Buck Creek Serpen- tine Pine Barren, Clay Co., North Carolina.M.S. thesis, Curriculum in Ecology, North Caro- lina State University. Manseerc, L., and T.R.WentTworth. 1984. Vegetation and soils of a serpentine barren in west- ern North Carolina. Bull. Torrey Bot. Club 111:273-286. Nature Serve. 2004. NatureServe Explorer: An online encyclopedia of life [web application]. Version 1.8. NatureServe, Arlington, Virginia. Available www.natureserve.org/explorer. Nesom, G.L. 1976. A new species of Erigeron (Asteraceae) and its relatives in southwestern Utah. Brittonia 28:263-272. Nesom, G.L. 1994. Taxonomic overview of Aster sensu lato (Asteraceae: Astereae), empha- sizing the New World species. Phytologia 77:141-297. Nesom, G.L.1997.The status of Aster scabricaulis (Asteraceae: Astereae),an endemic of the Gulf coastal plain. Phytologia 82:300-315. Raprord, A.E. 1948. The vascular flora of the olivine deposits of North Carolina and Geor- gia. Ph.D. dissertation, Department of Botany, University of North Caroina at Chapel Hill. Raproro, A.E., H.E. AHLEs, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. University of North Carolina Press, Chapel Hill. ScHarate, M.P, and A.S. Weaktey. 1990. Classification of the natural communities of North Carolina, third approximation. N.C. Natural Heritage Program, Raleigh, N.C. Warners, D.P. and D.C. LAUGHLIN. 1999. Evidence for a species-level distinction of two co- occurring asters: Aster puniceus L. and Aster firmus Nees. Michigan Bot. 38:19-31. 840 BRIT.ORG/SIDA 21(2) BOOK REVIEW Blueberries and Cranberries JENNIFER TREHANE. 2004. Blueberries, Cranberries and Other Vacciniums. (ISBN 0-88192-615-9, hbk.) Timber Press Inc. 133S.W Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@ timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). U.S. $29.95, CAN $39.95, 272 pp., 66 color photos, 7" x 9". 1 Other Vacciniums, she begins by In the new book by Jennifer Trehane, Blueberries, Cra tracing the story of the most widely grown species of Vacciniums, blueberries and cranberries. The book deals with their propagation, cultivation, harvest, and uses, including their health benefits: varieties are also ee, oe Other relatively unfamiliar, fruiting vacciniums are dealt with in Id gaa he and neo- or ieuaed sara wcenaieds ‘forms.” Tl eee should note, the bool i blueberries, cran- berries, and even ein onasmall scale commercial basis, but is not intended as a complete in manual for large scale commercial a In part one the author gives a semi- ae f overview of the genus, including the history, archaeol- ogy, cultivation, pests, diseases, ie eS I found it to be useful in that the information was di- vided up by the species, common name, and inside that chapter, she covered all the aforementioned topics, specific to that plant—Asha McElfish, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. BOOK NOTICE MA DELA Luz ARREGUIN-SANCHEZ, RAFAEL FERNANDEZ Nava, and VAvID LEONOR QUIROZ GarciA. 2004. Pteridoflora del Valle de México. (ISBN not given) Instituto Politécnico Nacional, Escuela Nacional de Ciencias Biologicas, Prolongacion de Carpio y Plan de Ayala, Santo Tomas, CP 11340, México DE (Orders: same) Price not given, approx. +00 pp., color photos. Table of contents: Introduccion; Material y métodos; Morfologia de pteridofitas; referencias bibliograficas citadas en los antecedents e introduccion; Clave ae paral las f familias beg Sees fitas del Valle de México; Descripcion de las familias espeices descritas, Glosario de términus utilizado: Indice de moms cientificos del material del Valle de México. SIDA 21(2): 840. 2004 ASYNOPSISOP THE GENUS OTOST Vs (ORCHIDACEAE: MAXILLARIEAE SUBTRIBE ZYGOPETALINAE) WITH A NEW RECORD FROM SOUTHERN PERU Miguel Chocce John P. Janovec San Ma Herbarium (USM) gas Research Institute of Texas ei eo de Historia Natural Fort Worth, Texas 76102-4 Aveni a oe 1256, Jesus Maria Lene OL ole Apdo. 14- Coe ai 14, PERU (C L r) igquel] Eric A. Christenson 1646 Oak Street Sarasota, Florida, 34236, U.S.A. orchideric@juno.com ABSTRACT The sie Sey! is HOseiaacese)s is one i a the caus ion genera in the Aysopea um Sean whic pane and qanidad. The vies of this paper is to provide a synopsis of the genus, with descrip- tions of the four species and keys for their identification. Otostylis paludosa is resurrected from pre- vious synonomy under O. lepida and reported as a new record for the genus in Peru, occurring as a dominant herb in bog wetlands of the Department of Madre de Dios. RESUMEN cl AWinctaliel(Meehid Jo] 5 id laalianza de Zveonetalum SU] 4 pes | 5 i z a is LI ge ()+t rey fs : : 1: A & L en América del sur y Trinidad. El objetivo este articulo es proveer de una sinopsis del género, con descripciones de las cuatro especies y claves para su identificacion. Otostylis paludosa dejo de ser sinonimo de O. lepida y esta reportado como un record nuevo para el Pert, siendo una hierba dominante en los pantanos del Departamento de Madre de Dios. INTRODUCTION The circumscription of orchid genera related to Zygopetalum WJ. Hook. has been problematic throughout the history of botany, and periodically all taxa have been united within an impossibly broadly defined Zygopetalum. Our modern understanding of generic limits in the group begins with a generally accepted review by Garay (1973). Subsequent authors have dealt with nomen- clatural problems (Christenson 1988), generic reviews (Christenson 2002), and the confounding generic issues of taxa lacking pseudobulbs (Dressler 2000). ne of the lesser-known genera in the Zygopetalum alliance is Otostylis SIDA 21(2): 841-852. 2004 842 BRIT.ORG/SIDA 21(2) Schltr. It was described by the German orchidologist Rudolf Schlechter in 1918 in a review of genera previously combined with Aganisia Lindl. The name is derived from the Greek otos, meaning ear, and stylis, meaning column, referring to the conspicuous subapical wings on the column. This character separates it from related genera including Aganisia, Warreella Schltr, and Warreopsis Garay. Otostylis has been generally accepted in floristic accounts since its original publication. Rather astonishingly, despite their showy flowers, plants of Otostylis have been rarely cultivated in modern times and are only now being reintroduced to horticulture. The purpose of this note is to provide a synopsis of Otostylis and report the discovery of the genus for the first time in southern Peru. Recent collecting of vouchers for botanical and ecological studies in Madre de Dios, Peru, under the auspices of the Botanical Research Institute of Texas (BRIT) and the San Marcos Herbarium (USM) has revealed a substantial population of O. paludosa (Cogn.) Schltr.a previously endemic Brazilian taxon incorrectly reduced to synonymy in recent times. TAXONOMY Otostylis Schltr, Orchis 12:38. 1918. Tyre: Otostylis lepida (Linden & Rchb£.) Schltr. Caespitose terrestrials. Pseudobulbs ovoid, small, typically concealed by bracts and leaf bases. Leaves several, plicate, lanceolate, petiolate, acuminate, some- times with subsimilar foliaceous bracts subtending the pseudobulbs. Inflores- cences axillary erect long-pedunculate racemes, the floral bracts inconspicu- ous. Flowers numerous, several open at once Snowy, long-lasting, predominately white, crystalline in texture. Sepals an il ing. Lip unlobed or obscurely three- lobed, subauriculate, with a basal trans- verse crested callus, sometimes biseriate. Column straight, conspicuously winged, with a short foot; pollinia 4, on a common linear viscidium. Otostylis comprises four superficially similar species native to South America and Trinidad. , subequal, free, spread- KEY TO THE SPECIES at . Callus minutely warty, W-shaped with two short continuous arms extending onto the blade of the lip O.lepida . Callus not warty, entire or biseriate without continuous extensi Callus biseriate comprising a transverse ridge with three oa eae keels in front par N O. brachystalix 2. Callus entire, a high crest. 3. Callus toothed, inflorescences less than 50 cm tall (northern South America) O. alba 3. Callus ridged but not forming distinct teeth, inflorescences commonly 1m tall (southern Brazil and Peru) O. paludosa 1. Otostylis alba (Ridl.) Summerhayes, Kew Bull. 6:293. 1951. Aganisia alba Ridl., CHOCCE ET AL OF THE GENUS OTOSTYLIS 843 Timehri 5:204. 1886; Zygopetalum venustum RidL, Trans. Linn. Soc. Bot, ser. 2, 2:283, t. 47.1887; at ie (Rid1.) Rolfe ex J.D. Hooker, Bot. Mag. 118: sub t. 7270. 1892; Warreella venusta (Ridl.) Se Die Orchideen, ed. 1:425. 1914; Koellensteinia alba (Ridl.) Schltr, Orchis 9:32. 1915; Otos ae (Ridl.) Schltr. Orchis 12:41.1918. TYPE: VENEZUELA: Bolivar. Mt. Roraima district, sae the Kookenam River, 1000 m, E.F Im Thurn 360 E: BMk ISOTYPE: K!). Pseudobulbs ovoid, to 1.5 cm long, ca. 1 cm wide and 0.5 cm thick, completely concealed by the leaf bases, apically unifoliate. Leaves 3-4, arching, linear-lan- ceolate, petiolate, acuminate, to 25 x 2 cm. Inflorescences erect long-peduncu- late racemes to 40 cm long, longer than the arching leaves, the floral bracts triangular-ovate, acute, to 0.6 cm long. Flowers 10-12, to 2.5 cm across, white, the callus and surrounding field yellow. Sepals and petals subsimilar, subequal, elliptic, obtuse, the dorsal sepal to 1.3 x 0.8 cm, the lateral sepals to 1.5 x lcm, the petals to 1.2 x 0.7 cm. Lip unlobed, elliptic, obtuse, minutely emarginate, to 13 x 13cm, the callus a high transverse semicircular crest divided into blunt teeth, ca. 7 mm wide. Column winged, to 1 cm long, width at wings 7 mm wide, width at base 2.5 mm wide, the wings obliquely ovate. Fruits not seen. Distribution.— Venezuela, the Guianas, and Brazil at elevations around 1000 m (Foldats 1970) Some authors reduce this species to the synonymy of O. lepida (e.g., Dunsterville &@ Garay 1965; Pabst & Dungs 1977; Senghas & Gerlach 1996; Romero-Gonzalez 2003) but we are following Foldats (1970) in keeping it sepa- rate. Otostylis lepida has been recorded from the Brazilian states of Amazonas, Matto Grosso and Para (Pabst @ Dungs 1977). The records for Amazonas are likely correct; the records for Matto Grosso are based on the previous inclusion of O. paludosa in synonymy; and the records for Para require reexamination of those vouchers, outside the scope of this paper. 2. Otostylis brachystalix (Rchb.f) Schltr, Orchis 12:39. 1918. (Figs. 1, 2). Zygopetalum brachystalix Rchbf., Ann. Bot. Syst. 6:660. 1863; Aganisia brachystalix (Rchbf.) Rolfe, Orchid Rey. 22:200. 1914; Koellensteinia brachystalix (Rchb.f) Schltr, Orchis 9:31. 1915. TYPE: TRIN- IDAD: leg. ignot. s.n. (HOLOTYPE: W, drawing seen). Pseudobulbs short, ovoid, to 2 cm long, completely concealed by the leaf bases. Leaves 3-4, erect, grass-like, acuminate, to 70 x 2.cm, often half that size. Inflo- rescences erect long-pedunculate racemes to 70cm long, longer than the leaves, the floral bracts inconspicuous, ovate, acute, 6 mm long. Flowers 5-30, to 2.5 cm across, sepals and petals white, the lip white with a yellow center. Sepals elliptic, obtuse, to 1.8 x 1.2. cm. Petals obovate, obtuse, to 1.7 x 1cm. Lip obscurely three-lobed, to 1.2 x 1.2 cm, the lateral lobes small, subauriculate, flanking the callus, the midlobe transversely kidney-shaped, obtuse, Be oe a low trans- verse ridge across the lateral lip lobes in a raised cresce hes to the sides of the lobes. Column stout with conspicuous age ‘eee wings, to 0.8 cm long, 6 mm wide across the wings, 2.5 mm wide at the base. Fruits cap- sules, ellipsoid, smooth. BRIT.ORG/SIDA 21(2) Fic. 1. Habit of ee brachystalixfrom the Aripo Savannah wetland of Trinidad-Tobago. (Photograph by Julian Kenny, Trinidad-Tobago, 2003) CHOCCE ET AL., A SYNOPSIS OF THE GENUS OTOSTYLIS 845 Fic. 2 Inf] Al £n Ii- f L Iie €, Tobago, 2003) Trinidad-Tobago. (Photograph by Julian Kenny, Trinidad- 846 BRIT.ORG/SIDA 21(2) Distribution.—Trinidad, Colombia (Ortiz 1995), Venezuela, the Guianas, Brazil, and Peru (Loreto) at elevations of 150-1675 m. Much early confusion surrounding this species was resolved by Ames (1922) who provided the first illustration of O. brachystalix based ona plant from Trin- idad, where it was collected for the first time in the Aripo Savannah. Described by Reichenbach in 1861 as Zygopetalum brachystalix, this species is character- ized by having a weakly three-lobed lip and a biseriate callus consisting of a low transverse ridge between the auriculate lip lobules supplemented by three small longitudinal keels in front. Figures 1 and 2 provide images of the habit, inflores- cence, and flowers of O. brachystalix, courtesty of Dr. Julian Kenny. Pabst and Dungs (1977: 285) provided an illustration of a flower dissection of O. brachystalix. According to Julian Kenny (2004, pers. comm.), O. brachystalix is known in eastern Trinidad from the Aripo Savannah, which is a 360-hectare bog-like grassland lying on a bed of quartz sand over a clay pan. The savannah is boggy during the rainy season and arid during the dry season. The dominant vegeta- tion consists of grasses and sedges, as well as bog indicators, Drosera and Sph- agnum species. In the Aripo Savannah, O. brachystalix is more often seen on well-drained, slightly elevated patches. It [lowers typically in the dry season from January to April (kenny 1988). A mature plant grows to about 70 cm tall. Populations have declined significantly in the last 20 years (Kenny 2004, pers. comm.), presum- ably due to overcollecting, as it is the easiest to grow of the savannah orchids. A pot of sand and exposure to sun and proper watering is all that is necessary to cultivate this species (Kenny 2004, pers. comm.). Several other orchids exist in this habitat, such as Cleistes tenuis (Rchb.f.) Schltr, Epistephium parviflorum LindL, Habenaria leprieurii Rchb£f., H. mesodactyla Griseb.,and Sarcoglottis sim- plex (Griseb.) Schltr. Cyrtopodium parviflorum Lindl. also exists in the same habi- tat but according to Kenny (2004, pers.comm.) it is now rare due to overcollecting. 3. Otostylis lepida (Linden and Rchbf.) Schltr, Orchis 12:40. 1918. Aganisia lepida Linden and Rchbf., Beitr. Orchideenkunde 15, t. 5. 1869. Type: BRAZIL. AMAZONAS: Rio Negro, G. Wallis s.n. (HOLOTYPE: W)). Pseudobulbs slender, fusiform, to 6 cm long, to | cm in diameter, subtended by nonfoliaceous bracts. Leaves 1-3, arching-erect, linear-lanceolate, petiolate, acuminate, to 65 x 6 cm, often half that size. Inflorescences erect long-pedun- culate racemes, shorter than the leaves, to 25 cm long, the floral bracts incon- spicuous, ovate, acute, to 0.4 cm long. Flowers 4-10, 2 mm across, white, the sepals and petals often suffused with pink toward the apices, the callus and surrounding field yellow. Sepals and petals subsimilar, elliptic, acute, the se- pals to 2 x lcm, the petals to 1.8 x 0.8 cm. Lip unlobed, elliptic-suborbicular, slightly constricted above the base (= obscurely pandurate), obtuse-truncate with a minutely apiculate apex formed by conduplicate folding, to 18 x 18cm, 847 CHOCCE ET AL., A SYNOPSIS OF THE GENUS OTOSTYLIS = sat ~ = the hybrid origin of the species and a new contemporary pyene, P.Xc Hann llii. Because Folsom’s work was not Ewe aaa case auerncersandins a) t identity and rank of Ived in thi | | all have contrib- 5 x ] 13 j 4] Ls 5 7 =f: fo) 1 1 here. uted to the douldciea Th RESUMEN Platanthera chapmanii (Small) Luer emend. Folsom es una orquidea rara endémica de la llanura costera del Sur de los Estados Unidos. Desde su cesonpeten ongipal por omialh en He como Blepharigltschapmans Plat tantherachapmanit| ii Bal 1ORAY) A ; ] Fae Kes I pruebas del origen hibrido di la Bebe’ y un nuevo hibrido doutemponnees P._channellii. Debidoa que el trabajo de Folsom pliat disponible, todavia persiste mucho malentendido para la correcta identidad y rango oe Platanthe rd ae nii. Varias especies e hibridos estan implicados en este complejo y todos han contribuido a la confusion. Se clarifican estos errores y relaciones. Although geographically restricted to the southern portion of the southeast- ern United States, Chapman's fringed orchis, Platanthera chapmanii (Small) Luer emend. Folsom, is an important component of the summer-flowering or- chid flora of this area. No other complex within the Orchidaceae in the south- eastern United States has the unique position of having a species with an an- cestral hybrid origin: P chapmanii, and also having a current, or contemporary, occurring hybrid: P. xchannellii, with the same parentage. Historically known from East Texas, much of northern Florida, and southern Georgia, today it can be best found in the Apalachicola and Osceola National Forests of Florida. A few other small sites in northern Florida persist. The species is absent from the eastern half of the Panhandle. The Marion and Polk County, Florida records appear to be Channell’s hybrid fringed orchis, P. xchannellii Folsom. Few sites remain in East Texas (Liggio @ Liggio 1999) and the Georgia locales are based upon historic collections. No collections have ever been made from the area between the Apalachicola National Forest and East Texas. SIDA 21(2): 853-859, 2004 854 BRIT.ORG/SIDA 21(2) Although Correll cites the range for Platanthera chapmanii (as Habenaria xchapmanii) from New Jersey to Georgia and Florida and west to Texas, he was basing his knowledge on plants of both P chapmanii and P. xchannellii. True Platanthera chapmanii has always been, and continues to be, one of the rarest orchids found in North America and is endemic to this lower portion of the southeastern Coastal Plain (Fig. 1) (Brown & Folsom 2002). Chapman's fringed orchis originally was described by Small (1903) as Blephariglottis chapmanii. Ames (1910), noting its intermediacy between Habenaria ciliaris and H. cristata, made a new combination as Habenaria x chapmanii. This hybrid status remained for many years, including the new com- bination of Platanthera xchapmanii (Small) Luer made by Luer (1972). It was not until Jim Folsom’s work in 1984 that the taxon was restored to its rightful status of full species and a new, contemporary, hybrid, P. xchannellii, was de- scribed (Folsom 1984). Understanding Platanthera chapmaniiand its relationships to the closely related orange fringed orchis, P. ciliaris, and orange crested orchis, P cristata, is greatly simplified if the observer can see all three taxa in one field session. This can only be accomplished in the Osceola National Forest, because P. ciliaris is historically and apparently currently absent from any of the other known lo- calities. Liggio and Liggio (1999) clearly state that P ciliaris has never been found within any of the Texas locales for P chapmanii. Although vouchered from the anhandle counti ing the Apalachicola National Forest, no records exist for P-ciliaris within the Apalachicola National Forest (Anglin, pers. comm.; Brown & Folsom 2002; Folsom 1984, 1985). Conversely, P. cristata is often found growing within or nearby many of the P. chapmanii sites, especially in eastern Florida. Folsom (1984) clearly demonstrated that the origins of Platanthera chapmanii were most likely an ancient hybridization of P.ciliarisand P cristata. Therefore P. chapmanii is intermediate in size and characters between the two ancestors. Over the years it has evolved intoa stable, reproducing species with a very distinctive bent column. This evolution of the column shape is critical in the pollination of the species. At the same time the contemporary hybrid of P. ciliaris and P. cristata, Channell’s hybrid fringed orchis, P. xchannellii, occurs in rare situations when both parents are present. It, too, is intermediate between the parents, but the column is unlike that of P. chapmanii. Folsom (1984) illus- trates all of these characters in great detail. Because Folsom’s original publica- tion in Orquidea (Mex.) was not readily available to many interested orchidists, the article, with minor revisions, was reprinted in the North American Native Orchid Journal in 1995, and included all of Folsom’s graphics (Folsom 1995). One of the best aids in the initial determination of plants in the field is observing what predominates in the area. If both Platanthera ciliaris and P cristata are present and only a few intermediates are to be found, then they in BROWN, 855 Ni; A 2 . {rl Ce Me 4 j Lita (pi L L one el. L Wl +f i ¢. Fic. 1 all probability would be the hybrid P. xchannellii. If the majority of plants ap- pear intermediate between P ciliaris and P. cristata and only a few of either of the latter species are present then the observer needs to look carefully at the shape of the column, and most likely the majority of plants will be P chapmanit. The rostellum lobes of the column in P. chapmanii have a prominent, distinc- tive, and characteristic hook that is clearly visible while the rostellum lobes of the columns of P ciliaris are triangular and the tips pointing straight forward and those of P.cristata,are much shorter, nearly truncated, and witha very slight hook. In addition, characters that help in determining which species are present may also include geographic location, diameter of raceme, size of flower, length and position of spur, and shape of orifice (Folsom 1984, 1995). To simply state that Platanthera ciliaris is larger, P chapmanii intermediate in size, and P. cristata smaller has led to much confusion. For many orchid enthusiasts this, although not explicitly stated, implied overall size, especially height. That is not accurate and height should never be taken into account. All three species can grow from 10 or 15cm to, in the case of P chapmanii and P ciliaris, over a meter in height! When size comparisons are made they refer to the diameter of the raceme and the measurements of the individual flowers. Even the overall height of the flowering raceme is not a good criterion for identification. Because of the ancestral parentage of P. chapmanii eles can easily favor the overall raceme shape of either parent, but the raceme d appears to remain con- stant. Spur length in the three species is helpful as well. Typically, in P ciliaris the spur is 20-25 mm long, in P. chapmanii 10-14 mm long, and in P.cristata 5— 856 BRIT.ORG/SIDA 21(2) 8 mm long. Figures 2 and 3 show the relationships of P cristata, P chapmanii, and P ciliaris and will assist in understanding this comparison. In addition to understanding the species orchid observers need to be aware of the hybrids that are involved in this complex include: Platanthera xapalachicola PM. Brown & S. Stewart—(P. chapmanii x P. cristata) Platanthera xchannellii Folsom—(P. ciliaris x P cristata) Platanthera xosceola PM. Brown & S. Stewart—(P. chapmanii x P. ciliaris) Relationships among this group are best summed up in Figure 4. Platanthera blephariglottis, P conspicua, and P integrilabia are included in this diagram for completeness in the group (Brown 2003; Brown & Folsom 2004). These rela- tionships and putative parentages are based upon morphological criteria. Arti- ficially created hybrids, cytological, and molecular work has yet to be done on this entire complex. Because hybrid swarms of some or all three species occur it may be diffi- cult to determine individual plants. Platanthera xapalachicola is locally com- mon in northern Florida where both parents frequently grow together. They usually occur as individuals and may appear within stands of P chapmanii as smaller flowered, more slender plants or within stands of P. cristata as larger flowered more robust individuals. The hooked column of P chapmanii is usu- ally dominant, but the spur length and position is intermediate (Brown & Stew- art 2003). Platanthera xchannellii and P. chapmanii can be difficult to tell apart. In the field one of the best ways is to look about and see which other species are growing nearby. If all the plants observed are the same, and within the range of P. chapmanii, it is most likely P chapmanii, whereas if it is a colony of mixed species and only a few intermediate plants are present it is more likely to be P. xchannellii. Platanthera xosceola is known only from Osceola National Forest where it is the only place documented that both parents are found growing together. Plants of the hybrid usually occur as individuals and may appear within stands of P chapmanii as larger flowered, more robust plants with decidedly longer spurs or within stands of P. ciliaris as smaller more compactly flowered indi- viduals. The hooked column of P. chapmanii is not as dominant as in P. xapalachicola (Brown & Stewart 2003). Understanding both the history of Platanthera chapmanii and the hybrid swarms that may accompany plants in the wild hopefully will help in clarify- ing some of the mystery around this rare and spectacular orchid. Photographs and details of all of the taxa mentioned above are found in Brown and Folsom (2002, 2003, and/or 2004). All orange-flowered plants throughout the over- BROWN, m 857 w Fic.2. Ancestral parentage of Platanthera chapmanii. From left to right in all photos: Platanthera cristata, chapmanii, and ciliaris. BRIT.ORG/SIDA 21(2) vera A ce SN pes ate Brine pe tN ( 4 r = \ at Diat. eh tek: flanfe\ Diet ek L ong Fig. 3 ( fF ( ), Platanthera ciliaris (right). y Platanthera - chapmanii *-. ; an a mx é = 7 = Z a . ra : 7 ‘ x z . xosceola P. xapalachicola ae 7 7 Ss 2 . . ~ % P. cristata P. xchannellii <—————— P. ciliaris P. xcanbyi P. xbicolor P. blephariglottis P. xbeckneri | P. xlueri P. — P. integrilabia Fic. 4. Rel 7 hit g | 1 Sf i. +. L g J | fring j Ld pl Di wt 7 blept ig P. conspicua i; aes g cy em wes +4 1 j iva 1 * +L BROWN | 859 lapping ranges of P ciliaris, P. cristata, and P. chapmanii should be carefully examined for the possibility of additional sites for Chapman’s fringed orchis. ACKNOWLEDGMENTS The author thanks Scott Stewart, Bill and Pamela Anderson, Gustavo Romero (AMES), and Guy Anglin for assistance in locating populations in the field. Scott Stewart, Joe Liggio, and Larry Magrath made several helpful suggestions. Jim Folsom provided initial information some years ago and Stan Folsom executed the line art. REFERENCES Ames,O.1910. Orchidaceae: illustrations and studies of the family Orchidaceae issued from the Ames Botanical Laboratory, North Easton, Massachusetts. The genus Habenaria in North America. The Merrymount Press, Boston, MA.4:155. Brown, P.M. 2002. Revalidation of Platanthera conspicua, the southern white fringed orchis. North Amer. Native Orchid J. 8:3-9. Brown, PM. and S.N. Fotsom. 2002. Wild orchids of Florida. University Press of Florida, Gainesville. Brown, PM. and S.N. Fotsom. 2003. The wild orchids of North America, North of Mexico. University Press of Florida, Gainesville. Brown, PM.and S.N.Fotsom. 2004. Wild orchids of the southeastern United States, north of peninsular Florida. University Press of Florida, Gainesville. Brown, P.M. and S. Stewart. 2003. Two new hybrids in the genus Platanthera.North Amer. Native Orchid J.9:35. Fotsom, J.P. 1984. Relationships in the yellow fringed orchid complex. Orquidea (Mex.) 9:321-45. Fotsom, J.P. 1995. The real identity of Platanthera chapmanii. North Amer. Native Orchid J. 1(3):211-238. (reprinted with minor changes from Folsom, 1984) Liccio, J.and A.O. Licaio. 1999.Wild orchids of Texas. University of Texas Press, Austin. Luer, C.A. 1972. The native orchids of Florida. New York Botanical Garden, Bronx. Sma t, J.K. 1903. Flora of the southeastern United States. Published by the author, New York. P. 314. 860 BRIT.ORG/SIDA 21(2) BOOK NOTICE Beautiful Peonies Joser J. HALDA and James W. W appick (Botanical Illustrations by Jarmila Haldova). 4. The Genus Paeonia. (ISBN 0-88192-612-4, hbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwttim berpress com, mail@timber press.com, 503-227-2878, 1-800-327- 5680, 503-227-3070 fax). $34.95, 228 pp., 36 watercolors and 43 line draw- ings, 8 x” « IL" The Genus Paconia by Josef J. Halda with James W. Waddick, botanical illustrations by Jarmila Haldova, presents 36 watercolor paintings depicti g the peony. Simply put, botanical art is a mix- g | P ture of science and art. The more walanecd he mix, he higher the chance of success. The 22 color portraits here by book illustrator Jarmila, succeed more as art nies science. T works are beautiful and paints The papery quality of the petals is convincing. The dark colored Grand flowers, dramatically lit, are posed against s backgrounds. As art, the peonies so saturated in color as to appear satin-like. Veining on leaves is c: Lin rich detail. Deli- cate hairs on stems a virtuoso handling of the brush. But an inconsistent hits source that cast bright to almost white coloring on flowers yet placed no shadow at all on some stems and leaves, disap- pointed. Given the rigid standards of botanical drawing, better to abandon dramatic lighting alto- gether and spend all available time on the accuracy of the live plant specimen placed in front of you. An uncomfortable shortening of leaves and twisting of some flower heads to fit the formatting of the paper resulted in perspective issues. Ideal positioning of flower placement on paper should be stud- ied betore the start of the drawing. While not strict botanical renderings of peonies, these are stun- ning portraits of peonies lovely enough to rame In addition to the color paintings in The Genus Paconia, are 43 stunning black and white line and stipple drawings. These are sure and confident. Cooly devoid of illustrative styling. The perfect mix of art and science.—Cynthia ocean poled! Arts —_— bad Mixon Dr, Dallas, Texas 75220, U.S.A. 214) 351-3447 ph/fx il Padilla popular instructor for universities, arboretums, garden and Alan societies, noes xtensiv ely rese ‘tand nature. Available for research she commissioned plant portraits, workshops and a travel (ad eronsketch- ing tours. Her art is available for exhibit. SIDA 21(2): 860. 2004 A PRELIMINARY ANALYSIS OF CLEMATIS (RANUNCULACEAE) IN SUB-SAHARAN AFRICA Frederick B. Essig Institute for Systematic Botany Department of Biology SCA 110 University of South Florida Tampa, Florida 33620 U.S.A. ABSTRACT Based on a survey of the literature and a representative sampling of herbarium material, it appears that the approximately 70 published binomials for Clematis in sub-Saharan Africa can be reduced to 10-13 distinct species or speci plexes. A key to these species, along with a commentary on each, is provided as a preliminary guide for further floristic and taxonomic studies. RESUMEN Basado en una revision de la literatura y una prueba representativa del material herbario, parece que l imad 70 bin blicad las clematides del Africa sub-Sahariana pueden are oh gm me | te [ A ser reducidos a 10-13 complejos de especies, 0 especies distintas. Se proporciona una clave p ecies, junt iop dl guia preliminar p lios adicional floristicos y taxonémicos. INTRODUCTION This report originated as part of an effort to develop tentative hypotheses about the number and distribution of the species of Clematis in various parts of the world. It was found, however, that the literature referencing Clematis in sub- Saharan Africa, in which approximately 70 species have been named, was con- fusing, patchy, inconsistent and often contradictory. The goal of this exercise was therefore to make the first continent-wide survey of sub-Saharan Clematis in Africa, combining study and synthesis of the diverse literature on the sub- ject with a limited but representative herbarium study, in order to provide a preliminary overview of the number, correct names, and distribution of the species. North African species were excluded because they are more naturally a part of the Mediterranean flora. It is hoped that this preliminary report will provide a framework, some meaningful hypotheses, and a stimulus for regional African botanists to take up the detailed work that needs to be done. A compre- hensive treatment of African Clematis will require extensive study in the her- baria of Europe and Africa, as well as a great deal of fieldwork in all parts of this large continent. Since the study was initiated, global surveys of Clematis, including the African species, have been completed by Johnson (1997) and Grey- Wilson SIDA 21(2): 861-877. 2004 862 BRIT.ORG/SIDA 21(2) (2000). Both authors have great familiarity with Clematis and each brings some interesting perspectives to the study of this genus, but neither addressed the many taxonomic and nomenclatural problems of the African Clematis in any depth. The relationship of the African species with those of other parts of the world is of great interest, but no definitive phylogenetic studies have been com- pleted. Grey- Wilson considers the African species, with the exception of C. gran- diflora and the species of section Pseudanemone, to constitute a subsection of the section Meclatis. This is an interesting and feasible hypothesis, but needs to be born out by formal phylogenetic studies. Other authors, such as Tamura (1967) and Johnson (1997) have considered the African species to be part of the subgenus or section Clematis, which includes similar white-flowered species in Eurasia and the Americas. Even under the best of circumstances Clematis is a taxonomically diffi- cult genus, due to great variability within species and minimal diagnostic dif- ferences in floral morphology between species, at least as can be seen in her- barium material. Previous work with North American Clematis (Essig 1990) had suggested that taxonomic problems in this genus might be largely resolved through careful analysis of existing herbarium material. In that study, the two widespread and frequently confused white-flowered species in eastern North America were found to be distinguishable by subtle morphological features plus differences in blooming season and substrate preferences. Such studies are fea- sible for taxa that are readily collected by generalists, and hence abundantly represented in herbaria. This study was initiated with the anticipation that this would be true of African Clematis as well. MATERIALS AND METHODS The survey of herbarium material was based on the collection of African Clema- tis at the Missouri Botanical Garden (MO), consisting of some 550 specimens. The Missouri Botanical Garden hasa strong history anda sizeable ongoing pro- gram of fieldwork in Africa, and probably the best herbarium collection of Af- rican plants in North America. The collection includes sets of duplicates from important collectors in all parts of Africa, including Exell & Mendonca in An- gola, de Wilde, Ash, Westphal and Pichi-Sermolli in Ethiopia, Pawek, Phillips and Brass in Malawi, Reekmans in Burundi, Richards in Zambia and Tanzania, and countless other collectors who have collectively made a huge contribution. Table 1 shows the origin, by country, of the collections at Missouri. The best represented areas in the Missouri collections are the African upland areas stretching from South Africa to Ethiopia. Weakly represented areas include tropical west Africa from Senegal to Angola, the northern tier of the savanna belt running from Senegal to the Sudan, and the east coastal regions from Mozambique through Tanzania. These results probably reflect both the abun ESSIG, CLEMATIS IN SUB-SAHARAN AFRICA 863 Table 1. Collections in the Missouri Botanical Garden Hebarium (MO) by country of origin. Ethiopia 45 Uganda 20 Kenya 20 Burundi /Rwanda 33 Tanzania 46 Malawi 77 Zambia /Zimbabwe 67 Zaire 31 Angola 30 Namibia/Botswana 12 — South Africa 101 Mozambique 13 Tropical West Africa 36 dance of Clematis and the intensity of botanical activity in these areas, but sug- gest a reasonably balanced coverage of the continent, sufficient for at least the preliminary survey undertaken here. RESULTS The collections of the Missouri Botanical Garden were readily sorted into 10 broadly distributed groups that can be interpreted as species or potentially as species complexes. These were identified using the earliest validly published name applicable to each group, as determined through examination of type materials, and/or original descriptions/diagnoses of the many species described in the literature. These widespread and variable taxa include C. villosa, C. uhenensis, C. chrysocarpa, C. grandiflora, C. longicauda, C. dolichopoda, C. hirsuta, C. simensis, C. brachiata, and C. welwitschii. This is a conservative in- terpretation, a hypothesis of what appears to be the minimum number of dis- tinct species in Africa. Support for an additional three narrowly distributed species recognized by some authorities (C. burgensis, C. sigensis, and C. viridiflora), was weak or lacking in the Missouri collections, but are worth con- sidering further and included in this treatment. In addition, the subspecies of Clematis villosa and C. chrysocarpa recognized by Brummitt (2000) are well- supported by this study, and may be nearly as distinct as the other species rec- ognized. The preliminary nature of this study is stressed. Additional species may very well emerge after more exhaustive study. The species have distinctive but overlapping geographic and altitudinal ranges. A number of intermediate specimens suggest hybridization or incom- plete separation of the taxa, but most specimens fall clearly into one taxon or another. The taxa are defined primarily on the number, size, and shape of the leaflets, and in some cases by significant differences in flower size. Otherwise, flowers in African Clematis are relatively uniform with respect to morphology. All are bisexual and of a nodding habit (different from white-flowered species elsewhere, in which flowers are erect). Sepals vary only in size and slightly in texture, vestiture or color. Stamens are numerous, and all have hairy filaments. Carpels are numerous, all developing into achenes with long, plumose tails. Flo- ral characters referred to by earlier authors (e.g. anther shape) have been found to vary as much within taxa as between taxa. Finally, ecological separation is suggested by some differences in blooming season and altitudinal range, but 864 BRIT.ORG/SIDA 21(2) data on habitat and soil type were meager and inconsistent. Other subtle bio- logical differences, such as differences in floral fragrance and flower color, are hinted at sporadically on labels, and need to be investigated in the field A key to these species, and discussion of each, follow. KEY TO THE SPECIES OF CLEMATIS IN SUB-SAHARAN AFRICA 1, Plants erect, stiff-stemmmed, perennial herbs; flowers with sepals imbricate in bud (species formerly segregated as the genus Clematopsis, key based on Brummitt 0). 2. Achenes with oe hairs; flowers solitary; leaves with 1-2 (very rarely 3) pairs of pinn 3. Leaves ee ees 32- 55mm long;achenes (including plum style) 55-70 mm long. Angola, ss ean C. chrysocarpa ied assess 3 rae mostly 5—7-foliolate; oe as 32 mm long;achenes 35-50 mm Malawi, Tanzania and Mozambique ong. C.c 2. Achenes with gray hairs; flowers 1-many per stem; leaves simple to tripinn bbe subsp. bijuga 4. Leaves simple; flowers solitary; sepals 32-60 mm long. Angola and i to Tanzania C. uhehensis 4. Leaves trifoliate to tripinnate; flowers 1-many per stem; sepals 17-33 mm long (C. villosa) 5. Leaves trifoliolate or eee, 5-foliolate, flowers 1-5 ae em. Cameroun and Nigeria to Tanz ee a oliveri 5. Leaves pinnate to tripinnate, A trifoliolate; flowers 1—-many per ste 6, Leaves tripinnae with segments 1—5(-8) mm broad;flowers 1—5 per Zambia to amibia and S. Africa C. villosa subsp. stanleyi 6, Leaves pinnate to bipinnate, with segments 5-40 mm broad; flowers 1 many per stem. 7. Leaves densely sericeous beneath, pinnae lobes up to 15 mm broad and usually rounded at the ee flowers 1-7(-13) per stem; W. Zai Angola, S. Uganda, W. Tanz re, C. vi 7. Leaves sparsely to ec ae pubescent beneath, a illosa subsp. villosa lobes up to 40 mm broad and acute at eae flowers 5-many p stem; S. Zaire, Tanzania mbique, Malaw mbia . Plants vining or trailing; flowers with yee valvate in bu Cc villosa subsp. kirkii t 8 Flowers large, solitary or in clusters of 3, with sepals 15-45 mm long; carpels 50- ae an oe nijlat | ribbed, greenish to yellowish or cream- eae mostly more than 2.5 cm long (but with some specimens as short as 1.8 cm); leaves mostly 3-5- ieee: with leaflets mostly more than 50 long. 0. Leaves mostly 5-foliolate, leaflets lanceolate-ovate, rarely lobed, finely and bee, toothed; west equatorial Africa, from Guinea to Angola, 300 C. grandiflora 10. ee mostly 3-foliolate, leaflets broadly 3-lobed, ie dened rai fo- liage conspicuously reddish-golden pubescent; Ethiopia, 1350- 9. Flowers with spreading to reflexed sepals, sepals mostly 15-25 mm long! lets mostly less than 5 cm lon c longicauda af ESSIG, CLEMATIS IN SUB-SAHARAN AFRICA 865 11. Leaves 3-5-foliolate, leaflets broadly to narrowly ovate, abaxially reddish- golden pubescent; flowers white, reddish-golden pubescent externally, the sepals somewhat ridged; occurring in evergreen montane forest, Tanzania, Burundi, 1000-2500 m C. dolichopoda 11. boave mest iMies pInDatS leallet Ove apices to linear-elliptic, or ay tinged with pink, sepals not Gon epieuouely fibbed: eecinine in open woodland, Angola to Zaire, Tanzania, Malawi, & Zimbabwe, 400- 1600 m a welwitschii 8 Flowers small, numerous, with sepals mostly less than 15 mm long (to 20 long in some populations of C. hirsuta), white, cream, greenish cee mostly 5- or more foliolate, of various shapes; carpels fewer than 50 per flower. 12. Leaves 3-5-foliolate with leaflets ovate-lanceolate, rarely lobed 13. Leaflets regularly and finely toothed. Ethiopia to Zaire and Nigeria C. simensis 13. Leaflets with margins essentially entire. 14. Achenes with style 50-70 mm long; sepals 12-15 mm long. Kenya, Tanzania C. sigensis 14. Achenes with style to 33 mm long; sepals 10 mm long. Ethiopia C. burgensis 12. Lea or more foliolate with leaflets | | t | freq ly deeply che regular or cen toothed. 15. Flowers yellowish-green, coastal Mozambique and Tanzania C. viridiflora 15. Flowers white to cream or greenis 6. Leaves mostly once-pinnate, 5—9-foliolate, leaflets broadly ovate, ir- regularly lobed and toothed; tropical woodland and savanna, An- gola to n.e. South Africa and Zimbabwe, northward to Ethiopia, and from there westward to Senegal, 220-2340 m C. hirsuta Leaves mostly twice-pinnate, leaflets narrow, deeply lobed and in- frequently toothed; iomesiete to oe grassland, South Africa to Zimbabwe, Botswana 00-2200 m C. brachiata oY TAXONOMIC SYNOPSIS AND COMMENTARY Note: Three species of African Clematis, C. chrysocarpa, C. uhehensis and C. villosa, were formerly segregated in the genus Clematopsis, and not originally included in this survey. The recent paper by Brummitt (2000) provides a thor- ough treatment of those species, while providing the rationale for including them in Clematis. | am in full agreement with that decision, and include the species in this synopsis, but refer the reader to Brummitt’s paper for more detail and discussion. 1. Clematis brachiata Thunb,, Prod. pl. cap. 94. 1800. Type: not cited, but photos of authentic material from Uppsala have been seen. Clematis oweniae Harv. in Harv. & Sond., Fl. cap. 1:2. 1860. Clematis stewartiae Burtt Davy, Man. Flowering Pl. Ferns Transvaal. 1:37, 111.1926. Clematis thunbergii Steud., Nom., ed 2, 1:380. 1841, Harv. in Harv. & Sond., Fl. Cap 1:2. 1860. Clematis triloba Thunb., Prod. pl. cap. 94. 1800 (non B. Heyne ex Roth, Nov. pl. sp. 251.1851). Tyre: 866 BRIT.ORG/SIDA 21(2) The first species of Clematis described from sub-Saharan Africa, C. brachiata is similar to C. hirsuta, as both are widespread, variable species of open, dis- turbed habitats, and both produce numerous small white flowers. Exell and Milne-Redhead (1960), along with Thulin (1993), considered that C. brachiata and C. hirsuta would probably have to be combined, although earlier, Exell (937) recognized C. hirsuta in Angola and C. brachiata in South Africa. White (1962) considered C. hirsuta and C. inciso-dentata as synonyms of C. brachiata. The two species can, however, be generally separated by the simply pinnate leaves of C. hirsuta compared with the doubly- (or more) compound leaves of C. brachiata. The geographic separation, with C. brachiata confined primarily to the subtropical grasslands of eastern South Africa and C. hirsuta occurring pri- marily in tropical savannas and woodlands, also suggests ecological differences. Confusion between C. hirsuta and C. brachiata may have arisen in part because of a discrepancy between Thunberg’s diagnosis of the latter species and his own specimens at the Uppsala Herbarium. Thunberg actually described two species from South Africa. One (C. brachiata) supposedly had simply pin- nate leaves, and the other (C. triloba) had doubly compound leaves. Thunberg did not designate types, but authentic specimens have been found at Uppsala, one for each species, and apparently annotated by Thunberg. In contradiction to his diagnoses, both specimens have doubly compound leaves, as do the great majority of South African specimens. The specimen annotated as C. triloba has larger {lowers (sepals 10 mm long vs 5 mm long) than the specimen annotated as C. brachiata. Both flower size and the dissection of the leaf are quite variable, and the differences between these two specimens of Thunberg become insig- nificant when a large number of specimens are examined. Because of its sim- ply pinnate leaves, C. hirsuta may have been equated with Thunberg’s diagno- sis of C. brachiata by some authors. For them the species with doubly-compound leaves was C. triloba or one of the later names discussed below, There was some additional confusion concerning the name C. triloba Thunb.. It was apparently assumed by some later authors that this name was anteceded by C. triloba Roth ex B. Heyne, which however was not published until 1821. The name C. thunbergii was published by Steudel in 1841, appar- ently as a new name for C. triloba, so the latter name has been little used. In 1860, Harvey published C.oweniae for some South African specimens W ith dou- bly-compound leaves, apparently unaware of C. triloba. Some authors have at- tempted to distinguish C. oweniae from C. brachiata on the basis of anther shape, with the former species possessing short ovoid anthers and the latter elongate anthers. These differences do not hold up when a broad range of speci- mens is examined. The same applies to the species C. ste wartide. Johnson (1997) and Grey-Wilson (2000) recognized three South African species, C. brachiata, C. triloba, and C. owenide. Grey- Wilson distinguished C, oweniae trom C. triloba by its smaller flowers (sepals 10-12 mm long vs. 12-18 — — ESSIG, CLEMATIS IN SUB-SAHARAN AFRICA 867 mm long). He also says that the latter has “deliciously scented” flowers, but does not mention scent for C. oweniae. Flowers of C. brachiata are simply described as “fragrant,” and he appears to have misinterpreted C. brachiata as similar in foliage to C. simensis. Once again, variation in leaf form and flower size in South African Clematis are such that these kinds of lines are hard to draw without extensive field studies. Particular morphological combinations may indeed prove to be consistent within populations occupying specialized habitats or geographical areas. The issue of fragrance also needs to be examined more rig- orously and consistently. Hybridization between Clematis brachiata and Clematis villosa subsp. stanle yi, both of which are common in the Johannesburg area, has been noted in the literature (Letty 1962) and on some herbarium specimens. One specimen that is clearly intermediate in character between these two species is Mogg 36528 This underscores the close relationship between the species formerly segregated into Clematopsis and other African species, as well as the possibility that many unusual specimens in Africa could be the result of hybridization. — 2. Clematis burgensis Engl., Bot. Jahrb. Syst. 45:272. 1910. A distinct species according to Demel (1987), native to Ethiopia. Three of the specimens cited under C. simensis (Ash 1294, J. deWilde 6224, Westphal & Westphal-Stevels 3062) have smooth leaflet margins as indicated for this spe- cies, but lack flowers so the other traits could not be verified. 1 was unable to draw any conclusions about this species from the MO material. 3. Clematis chrysocarpa Welw. ex Oliv, FL. trop. Afr. 1:5. 1865. Clematis villosa subsp. chrysocarpa (Welw. ex Oliv.) Kuntze, Clematopsis chrysocarpa (Welw. ex Oliv.) Hutch. (see Brummitt, 2000). a. Clematis chrysocarpa subsp. chrysocarpa Brummitt (2000) places the following species in synonymy here: Clematis chrysocarpoides DeWilde., Clematopsis speciosa Hutch. (Clematis angolana M. Johnson, 1997, Slaktet Klematis: 145, nom. nov: replaced synonym: Clematopsis speciosa Hutch.) b. Clematis chrysocarpa subsp. bijuga Brummitt, Kew Bull. 55:97-108. 2000. Brummitt (2000) placed the following species in synonymy here: Clematopsis lineariloba Hutch. & Summerh., Clematopsis oliveri forma lineariloba (Hutch. S& Summerh.) Staner & Leonard. (Clematis africolineariloba W.T.Wang, Acta Phytotax. Sin. 39:336. 2001, replaced synonym: Clematopsis lineariloba Hutch- inson & Summerhayes). 4. Clematis grandiflora DC, Syst. nat. 1:151. 1818. Type: not indicated. Clematis pseudograndiflora Kuntze, Verh. Bot. Vereins Prov. Brandenburg 26:128. 1885. TYPE: ANGOLA: Welwitsch 1218, 1219. 868 BRIT.ORG/SIDA 21(2) Cl iskakoul isSchnell, Bull. Soc. Bot. France 96:223. 1949. Type: GUINEA: Mt. Kakoulima, 2000 ft, Schnell 2461 (HOLOTYPE: n.v.) Clematis chlorantha Lindl, Bdwardst Bot. Reg. 16, t. 1234. 1829. Belongs here according to Oliver (1868) and Johnson (1997). A very distinctive species with very large, campanulate flowers. Leaves are 5- pinnate, with leaflets ovate, rarely lobed, and finely toothed, much like those of C. simensis. Material referred toas C. kakoulimensisand C. pseudograndiflora have somewhat smaller flowers, with sepals 18-20 mm long, and about 50 car- pels per flower, but otherwise similar to the large-flowered specimens. They appear to represent extremes in the range of variation. A remarkable photo- graph of this species appears in Johnson (1997), showing large, pendant, yel- lowish flowers, reminiscent of Asian species in the section Connatae. 5. Clematis dolichopoda Brenan, Kew Bull. 1949:71. 1949. Clematis hirsuta var. dolichopoda (Brenan) te & J. Leonard, Bull. Soc. Roy. Bot. Belgique 82:36, 1950 and Flore du Congo Belge 2:187. 195 Clematis longipes Snr 1910. Ilegiti cf.Cl is longipesFreyn, 1890, from Madagascar. Poorly represented in the MO collections, this species appears to be similar to C. longicauda, but with smaller flowers. the foliage has the same distinctive indumentum of golden hairs Crusty ferrugineous”). Sepals are 13-20 mm long, variously described as white, golden-white, or yellowish, the latter perhaps due to the heavy external covering of golden hairs. Leaflets are cana heart- shaped, with irregular teeth, or in some material from Burundi, the leaflets are narrow-elongate. It is possible that the latter material represents hybrids with C. welwitschii. The species occurs mostly in high mountains, up to 3500 m, but as low as 1000 m in some Burundi material. One specimen, Williams 35, is placed here questionably. It is a fruiting specimen, but the large number of fruiting heads in the inflorescence suggests that the flowers are much smaller than in the other specimens. The hairs are much sparser on the leaves, though still golden in color. 6. Clematis hirsuta Guill. & Perr. in Guill, A. Rich. & Perr, Fl. Seneg. tent. Ll. 31.Typk: not cited, collected by Perrottet in 1829 at Cape Verde, Senegal near Kounoun and Rufisk. lematis inciso-dentata A. Rich., Tent. fl. abyss. 1:2, fig 1. 1847. TyPE: not specified, but collected yA. Petit in a Province, Ethiopia, between 1838 and 1843; = C. hirsute fide Demel (1987) and Johnson (199 Clonal is wightta na auct non Wall. n Wall.) sensu Oliv., Fl. trop. Afr. 1:7. 1868. Glemutise epic eset Fresen., Beitr. Fl. Abyssin., in Mus. Senckenb. 2: 268. 1837. TYPE: Not cited; = iG, hirs uta fide ape (1987) ses ng (1997). Cl sdjalonen Cheval. Bull. Mus. Hist. Nat. (Paris), ser. 2. 4:1010. 1932. TyPE: FRENCH GUINEA: Fouta- ar 1300 m, Chevalier 34545(nv)( C. hirsuta fide Hutch. & Dalziel). ESSIG, CLEMATIS IN SUB-SAHARAN AFRICA 869 Clematis chariensis A. Cheval., Bull. Mus. Hist. Nat. (Paris), ser. 2. 4:1012. 1932. Type: CENTRAL AFRICAN REPUBLIC: Haut-Chari, between Dekoua and Nana, Chevalier 6192; said to be a cousin of C. hirsuta by Chevalier, described as a low-growing scrambler adapted to frequent bush fires, sprouting ie from the rootstock. Clematis petersiana Klotsch in Peters, Naturw. Reise h bique 6(1):170. 1861. From the am- biguous description, ae species appears to be ae from ordinary C. hirsuta. It is from the upland, interior province of Tete, and so not likely to be equated with C. viridiflora. In the conservative treatment presented here, this isa widespread, variable spe- cies found throughout the tropical savannas and open woodlands of central Africa. It’s distribution correlates roughly with that of Acacia sieberanain mid- elevation plains and plateaus in what is called the Sudano-Zambezian region (Brenan 1978). The correlated distribution of the two species includes disjunct populations of each in central Angola. The two other widespread species with numerous white flowers appear to be geographically and/or altitudinally sepa- rated: C. simensis at generally higher elevations, and C. brachiata further south in the warm-temperate to subtropical grasslands of South Africa. A fourth spe- cies that appears to be in this complex, te viridiflora isa lowland species found along the coast of Mozambique and possibly Tanzania. There are morphologi- cal differences as well, but definitely gray areas between these species, both geo- graphically and morphologically. Clematis hirsuta is distinguished from C. simensis by its lobed, irregularly toothed leaflets, as opposed to the unlobed, finely dentate leaflets of the latter, and from C. brachiata by its simply pinnate leaves, as opposed to the doubly compound leaves of C. brachiata. C. viridiflora is hard to distinguish from herbarium material, but its flowers are somewhat larger, with the sepals said to be thinner and more yellowish. Clematis species with numerous, small, white flowers are found through- out the world, including the C. virginiana/catesbyana/ ligustifolia group in North America, C. dioica and its relatives in tropical America, C. grata/ wightiana and similar species in Asia, C. vitalba in Europe, and other species in New Guinea, Australia and New Zealand. All of these species are abundant, variable and widespread, occupying open, disturbed habitats, and can gener- ally be described as “weedy.” Species with larger, fewer, and often more colorful and/or fragrant flowers, on the other hand, tend to be less common and more restricted in both their distribution and habitat preferences. This pattern holds for Africa as well. The species placed in synonymy here were based on one or a few variant specimens and appear to fall within the range of variation in the widespread species, although they do warrant further study. The vestiture of the leaves in C. hirsuta is variable. Some, as the name implies, are rather densely hirsute, while others are nearly glabrous. The greatest concentration of heavily hirsute speci- mens is in the Ethiopia/ Uganda region. Some populations in Ethiopia have sig- nificantly larger flowers with rather attenuate sepals, as opposed to the more 870 BRIT.ORG/SIDA 21(2) obtuse sepals in other specimens, suggesting some degree of subspecific differ- and C. inciso-dentata, neither of which were recognized by Demel Tekatay (1987) in his thorough study of Clematis in Ethiopia. The present study also suggests that there are numerous intermediate specimens, making such a distinction difficult. Grey-Wilson (2000) recognized C. djalonensis from southwest Mali and northern Guinea, on the basis of its smaller flowers that are more rounded in bud and said to be more fragrant, and with shorter pedicels. This again ap- pears to fall within the range of variation for C. hirsuta, but needs to be investi- gated further. The photograph of C. hirsuta in Grey-Wilson appears to be misidentified. The flowers appear to be much larger than those of the many specimens examined in this study, and the leaves appear to be more deeply di- vided. The plant looks more like C. welwitschii. One difficulty in verifying the proper application of the name C. hirsuta is the lack of authentic type material. The type should be at Paris, but it cannot be found, and the Missouri collection contains no specimens from Senegal at al The protologue for that species indicates that the foliage is ternate to biternate, —_— and densely villous. No dimensions were given for the flowers, which were said only to be in clusters of 1-3. Biternate leaves are rare in the species as currently understood, suggesting the possibility that the population at Cape Verde (near sea-level) is distinct at some level from the more widespread form. In his flora of Senegal, Berhaut (1967) describes the leaves as having two pairs of leaflets, and the white flowers produced in large panicles, which is consistent with C. hirsuta elsewhere in its range. It is not known whether Berhaut had material from the Cape Verde area, however. If the species still exists there, it needs to be studied in order to resolve any further doubts. 7. Clematis longicauda Steud. ex A. Rich, Tent. fl. abyss. 1:2. 1847. Typr: ETHIOPIA: Schimper 1284 (rw). This taxon has large flowers similar to those of C. grandiflora. The 3-foliolate leaves differ conspicuously in the broader, lobed leaflets and the fine reddish- |: ie) gold pubescence. It is found at higher elevations (1350-2100 m), and only in Ethiopia. Johnson (1997) and Grey- Wilson (2000) both misinterpret this very distinctive species as a synonym of C. hirsuta. 8. Clematis simensis F'resen., Beitr. Fl. Abyssin., in Mus. Senckenb. 2:267. 1837. TYPE: Ruppell (nv). Clematis altissima Hutch. belongs here according to Hutchinson and Dalziel (1954) and Johnson 997). This is a distinctive species occurring throughout central and eastern Africa, generally at higher elevations than C. hirsuta, with which it overlaps geographi- cally. Leaves are typically 5-foliolate, with the leaflets finely toothed and ESSIG, CLEMATIS IN SUB-SAHARAN AFRICA 871 unlobed. Flowers tend to be numerous in elongate inflorescences, well exserted above the foliage. One specimen from Kenya, Taylor 1455, has unusually large flowers, representing perhaps a taxonomic variant or hybrid witha large-flow- ered species. 9. Clematis sigensis Eng]. Bot. Jahrb. Syst. +5:271. 1910. A distinct species according to Beentje (1989), found in Kenya and Tanzania; no matching specimens at MO 10. Clematis uhehensis Eng]., Bot. Jahrb. Syst. 28:387. 1900. Clematopsis scabiosifolia subps. uhenensis (Engl.) Brummitt, Clematopsis villosa subsp. uhehensis (Engl.) J. Raynal & Brummitt (see Brummitt, 2000). Brummitt (2000) includes the following names in synonymy here: ?Clematis villosa var. teuszii Kuntze, eee teuszii (Kuntze) Hutch.; Clematis homblei De Wild., Clematopsis homblei (DeWild) Staner & J. Leonard, Clematopsis hatangensis Hutch., Clematis katangensis (Hutch.) M. Johnson, Clematopsis simplicifolia Hutch & Summerh., Clematopsis grandifolia Staner & J. Leonard, Clematis grandifolia (Staner & J. Leonard) M. Johnson. 11. Clematis villosa DC., Syst. nat. 1:154. 1818. a. Clematis villosa subsp. villosa Brummitt (2000) placed the following species in synonymy here: Clematis scabiosifolia DC., Clematis villosa var. scabiosifolia (DC.) Hiern., Clematopsis scabiosifolia (DC.) Hutch., Clematis villosa subsp. argentea Kuntze, Clematis argentea (Kuntze) Prantl, Clematopsis argentea (Kuntze) Hutch., ?Clematis mechowiana Kuntze, ? Clematis villosa var. angolensis, Clematis stuhlmannii Hieron. ex Engl, Clematopsis stuhlmannii (Hieron. ex Engl) Hutch., Clematis villosa forma obtusiloba Hiern, Clematis villosa forma stenophylla Hiern, Clema- tis sapinii De Wild, Clematopsis sapinii (De Wild.) Staner & J. Leonard. b. Clematis villosa subsp. stanleyi (Hook.) Kuntze, Verh. Bot. Vereins Prov. Brandenburg 26:172. 1885. Clematis stanleyi Hook., Clematopsis scabiosifolia subsp. stanle yi (Hook.) Brummitt, Clematopsis stanleyi ( look.) Hutch. (see Brummitt, 2000) Brummitt (2000) places the following species in synonymy here: Clematis villosa var. tomentosa (Kuntze) T. Durand & Schinz. This is a very distinctive subspecies, with finely dissected leaves. c. Clematis villosa subsp. kirkii (Oliv.) Brummitt, Kew Bull. 55:97-108. 2000. Clematis kirkii Oliv, Clematis villosa var. kirkii (Oliv) Kuntze, Clematopsis kirkii (Oliv) Hutch., Clematopsis scabiosifolia subsp. kirkii (Oliv.) Brummitt (see Brummitt 2000). a Brummitt (2000) places the following species in synonymy here: Clematis villosa var. pubescens Kuntze; Clematis stanleyi var. pubescens (Kuntze) T. Durand & Schinz; Clematis goetzei Engl; Clematis busseana Engl; Clematis lugnignu De Wild. 872 BRIT.ORG/SIDA 21(2) d. Clematis villosa subsp. oliveri (Hutch.) Brummitt, Kew Bull. 55:97-108. 2000. Clematopsis scabiosifolia subsp. oliveri (Hutch.) Brummitt. Brummitt (2000) included the following species in synonymy here: Clematopsis nigerica Hutch. 12. Clematis viridiflora Bertol., Misc. bot. 19:7, pl. 3. 1830. The number of specimens available for this species at MO are few, but it appears to be a distinct species ecologically as well as morphologically. The name sug- gests that the flowers are greenish. This is confirmed on one specimen (Lemos & Balsinhas 37) who describe the flower color as “amarelo-esverdeades.” Exell and Milne-Redhead (1960), describe the sepals as “thin and membranous” as opposed to the thicker sepals of C. brachiata, which they equate with C. hirsuta. Ecologically, it appears to be a coastal species, apparently growing on dunes—a most unusual habitat for African Clematis, although in North America there are populations of C. catesbyana ina similar habitat (Essig 1990). A photograph of a cultivated plant attributed to this species in Grey- Wilson (2000) shows a plant with yellowish flowers. If accurately identified, it confirms a distinctly different species, and also strengthens the argument of a relationship with sec- tion Meclatis. One specimen from Zanzibar Island is tentatively placed here, though it has smaller flowers, reported to be white, and leaflets less lobed than the material from Mozambique. It may be represent C. zanzibarensis Bojer ex Loud., who found it similar to C. vitalba and C. grata, the latter a name widely misapplied to C. hirsuta. It has to be remembered also that Zanzibar has beena center of trade for centuries and that exotic species may have been brought in. 13. Clematis welwitschii Hiern ex Kuntze, Verh. Bot. Vereins Prov. Brandenburg 26:171. 1885. Type: Welwitsch 1217 (photo seen, MO- Angola); see also Exell & Mendonca (1937). Clematis antu it Engl. Bot. Jahrb. Syst. 45:274. 1910. as ANGOLA: Antunes A56 (BD, n.v.) ongs here en Hs to Exell and Mendonca (193 Eee tar Kuntze, Verh. Bot. Vereins Prov. ed burg 26:128. 1885. Type: ANGOLA: Welwitsch 1215a (MO-photo seen). Clematis prostrata Hutch., cene southern Africa. 484. 1946. TyPE: Hutchinson ai v.), ap- ears to belong here from the description, placed here by Exell @Milne-Redhead, 1960. oe thalictrifolia Engl., Bot. Jahrb. Syst. 45:270. 1910. Zaire, Zambia, Tanzania; close to C. welwitschii fide Exell & Milne-Redhead (1960), distinguished by larger, solitary flowers. This taxon includes specimens with medium-large flowers (sepals 1.5-2 cm long) with spreading sepals. Flowers are reported to be white to cream-colored, but sometimes with a pink tinge on the outside. Foliage is extremely variable in this species. Leaves are pinnately to doubly pinnately compound, with leaflets coarsely toothed, deeply lobed, elongate, and sometimes very finely dissected. The types of both C. welwitschii and C. commutata were collected in Angola, and both names have been applied to specimens with moderately large flowers ESSIG, CLEMATIS IN SUB-SAHARAN AFRICA 873 in a broad area from Angola, Zaire and Cameroon to Tanzania. Differences be- tween them, primarily leaf shape and number of flowers, blur considerably when a large number of specimens are examined, and they appear at this point to be just forms of one variable species. Johnson (1997) and Grey- Wilson (2000) both recognize C. commutata and C. thalictrifolia as separate species, but the distinctions are not clear or convincing, and the ranges overlap. The Missouri collections did not contain any specimens with flowers as large as those re- ported for C. thalictrifolia. They are said to be up to 50 mm across, which put them in the range of C. grandiflora or one of the species of section Pseudanemone, suchas C. villosa. This potential species is certainly worth investigating. Anum- ber of the specimens cited below have broad, cordate leaves and/or somewhat smaller flowers, strongly suggesting hybridization with a species such as C. hirsuta. These include Exell & Mendonca 1076 from Angola, Lacroix 3020, Tawakali & Kaunda 190, and Pawek’s 6255, 12644, and 13658 from Malawi. ERRONEOUS AND UNRESOLVED NAMES IN AFRICAN CLEMATIS The following names are either erroneous, invalid or require further research to determine their status. (note: IPNI = International Plant Name Index at wwwipniorg) Clematis capensis Poir. Encyc. Suppl. 2, 298. (= Anemone capensis according to IPND. Clematopsis costata Weim.,, Bot. Not. 1936:28. Zim babwe/Zambia. aaa eee Ulbr., Notizbl. Bot. Gart. Berlin-Dahlem, 10:914. 1930. Kenya. (= C. hirsuta fide J n 1997) G ematis oe Chiov, Ann. Bot. (Roma) 9:51. 1911. hybr, Ethiopia; said to be a natural hy brid between C. simensis and “C. thunbergii” (C. hirsuta?). Clematis iringaensis Engl., Bot. Jahrb. Syst. 28:388. 1900. Zambia and Tanzania; resembles C. welwitschi, but flower buds covered with “yellow” indumentum, possibly as in C. dolichopoda Clematis kassneri Engl. Bot. Jahrb. Syst. 45:274. 1910; Zaire. (= C. welwitschii fide Grey- Wilson ( ) 2000). Clematis keilii Engl, Bot. Jahrb. Syst. +5:273. 1910; Burundi. (= C. welwitschii fide Grey- Wilson (2000; but said by Engler to be “ferrugineo-pilosus” and similar to C. longipes, ie. C. dolichopoda). Clematis kerrii Steud., Nom. ed. 2. 1:379. 1840. South Africa. Clematis kissenyensis Engl. in Wiss. Ergebn. Deutsch. Zentr. Afr-Exped. 1907-8, 2:207. 1911. Tropi- cal East Africa. (=C. simensis fide Johnson 1997) Clematis massoniana DC,, Syst. nat. 1:135. 1818. (= C. brachiata fide IPND). Ethiopia? South Africa. Clematopsis pulchra Weim., Bot. Not. 1936:27. Clematis schinziana Engl. & Gilg ex Engl., Pflanzenw. Ost.-Afrikas 3:1. 1895. (Engl. & Drude, Veg, Erde oe cae in obey Nemibis) Cc] 1. Bot. Jahrb. Syst. 9:258. 1888. in obs. Zaire (cannot be deter- aaned fide Brummitt 2000). Clematis a Engl. Bot. Jahrb. Sys. 45:272 (1910), Malawi. (= C. eee Hage 1997, but w, rather large flowers; could be a form of C. welwitschii or a hy Clematis teguuons id Poir., Encye. Suppl. 2, p. 298. 1786. (=Anemone t seal ND. Cl |, Pflanzenw. Ost.-Afrikas 3:1. 1895. (Engl. & Drude, Veg. Erde 9:170. 1915, in obs. (= C. villosa var. a tedeee fide IPND). Angola. 874 BRIT.ORG/SIDA 21(2) Clematis tibestica Quezel, Bull Soc. fae Nat. Afrique N. 48:86. 1957. Tibesti sista of eudan (treated as a separate species by Johnson (1997) and Grey- Wilson (2000), / its close affinity with C. simensis). Clematis zanzebarica Sweet, Hort. brit. ed. 2, L. 1832. = C. zanzibarensis Loudon, fide IPNI (=C. viridiflora?). Clematis zanzibarensis Bojer ex Loudon, Hort. brit., ed 2,228. 1832. (=C. viridiflora?, similar to C. vitalba or C. grata fide Exell @ Mendonca 1937). APPENDIX Abbreviated listing of specimens examined (all from MO) Clematis brachiata | hunb—Botswana: Skarpe 284. Namibia: Leisiner et al. 194; Muller & Tilson 917: Seydal 2150, 4123, 4291. South Africa: Allardice 1572; Arnold 205; Balsinhas 2857, 3416: Bayliss 1188, 1331, 4674; Boot 32; Borle 508, 1110, Brink 596, 628; Buitendag 812; Burtt-Davy 15148; Codd 9648: Dahlstrand 2387, 2542, 3515, 3588, Davidse 6776, Drege s.n. (6 specimens with meager label data): Ecklon s.n. (3 specimens with meager label data); Edwards 175; Edwards & Vahrmeyer 4281; Germishuizen 225, 3168, 3924, Gibbs et al. 234; Goldblatt 1688, Halliwell 5126; Hilliard & Burt 9836; Joffe 232; Kemp 864; Krause 1234; Leendertz 50, Liebenbere 7564, 8820, Medias wood 4733, s.n. (1900): McLean 331, 534, 829; Mogg 19347, 21053, 25 20% 25832; Morley 377, Pillans 10910; Phillipson 551; Ray- mond s.n. (1972); Reardon 17; Retief izen 286, Rodin 3685, 3874, Scheepers 1398, 1485, Sidey 683, 2432, Stinton 83; Strey 2542, 9745. 86411181, 11263, Vahrmeyer 2425; van Hoepen 1648; Welman 729, 801; Wild 5781; Zambatis 1182. Swaziland: Kemp. Zimbabwe: Davies 388; Neoni 372; Norrerann ae tis subsp. chrysocarpa Welw. ex Oliv—Angola: F:xvell & Mendonca 794, 1360, 2547: Homble a BR); Welwitsch 1222 (photo, BM). Zaire: Symoens 6060 Clematis chrysocarpa subsp. bijuga Brummitt—Malawi: Brummitt et al. 15601; Chapman & Chap- man 7510; Lacroix 4257, Pawek 6256, 8044, 8415, 8894, 10797, Phillips 799, 1310; Reekmans 5527. Mozambique: Jansen & Boane 7882; Pereira etal. 1827. Tanzania: Bidgood & Congdon 147; Kayombo & Kayombo 166; Gereau & Kayombo 4363; Richards 15564; Stolz 2385 (ty pe of C. lineariloba Hutch. & Summerhayes) Clematis dolichopoda Brenan—Tanzania: Williams 35, Verdcourt 275. Schlieben 3991. Burundi: Reekmans 2298, 2365, 8476. Clematis grandiflora DC—Angola: Gossweiler 10311, Welwitsch 1218 (photo, BM, type of C. ae udograndiflora), 1219. pees Bates 1257, Guile 10, Latilo & Daramola FHI 28900, de Wilde & Wilde-Duyfjes 1157. Cote d'ivoire: Hepper & Maley s.n. 1984). Ghana: Vigne 2681. Guinea: Adam 857, 7562. Liberia: Konnel 621. Nigeria: Dalziel s.n. (1912). Sierra Leone: Adam 22116, 22315, 22409, ae Zaire: Liben 3192. Clematis hirsuta Guill. & Perr—Angola: Giess et al. 6612: Gossweiler 10285, 11379. Botswana: Smith 616, 3520. Burundi: Le walle 4674; Reekmans 595, 3444, 5132, 5133, 5138, 6271, 8029, 8983, 9145, 10258. 10420. Cameroon: Thomas 3105; Baldwin 13852; deWilde & de Wilde-Duyfjes 4332, 4117, Latilo & Dasabika 28773, Leeuwenberg 7561, 10497. Central African Republic: fay 5562, 6084. Céte d'Ivoire: Gautier-Beguin 460. Ethiopia: Ash 652, 1279, 1343; de Wilde & de Wilde-Duyfjes 8693, 9277; J. de Wilde 5773, 5866, 6225, 7350; Mengeshu s.n. (1958); Sean & Nievergelt 1227, Pappi 4642, 507, Pichi- Sermolli 2372; Schimper 212, 1481; Westphal & Westphal-Ste vels 2476, 2876, 2951. Fernando Po: Guinea 1876. Guinea: Adam 2630, 7174, 7190. Kenya: Agnew et al. 10283; Mwangangi 77; Faden et al. 74/655: Harmsen & Agnew 6540; Paolo 541, Perdue & Kibuwa 8155; Williams 315. Malawi: Banda © Thera 2612; Brass 1712, 17119, Chapman & Chapman 7496; Kwatha & Balaka 139, LaCroix 4562; Pawek 5547, 5548, 7162, 11354; Phillips 1438, 2678, 2823; Salubeni 1536, 2799, 3156; Salubeni & lawakali 4960, 5000: ESSIG, CLEMATIS IN SUB-SAHARAN AFRICA 875 phy 7, bao} 7 ta] 3( ()] ife Stolz 206; LaCroix 4562, Willan 65.M bi le K et al. OBB94, OBB501, Guile 13. Rwanda: Rouse & Rida 2216 Sierra Leone: ee 22214, “22768, 22223, 23259, 22944; Morton SL2804. Somalia: Pichi-Sermolli 138. South Africa: Cooper 191; Galpin 14413, 14418: Hemm 596; Stalmans 441. Swaziland: Kemp 734, 847. Tanzania: Flock 423; Frame 149: Gereau & Mziray 1678; Gereau et al. 4600; Grant s.n. 1928); Greenway & Kanuri 15164; Jefford et al. 243: Kayombo 555; Lovett 2182; Lovett & Congdon 1852; Lovett & Kayambo 3458; Mathias et al. A86; Newbould & Harley 4220; Paget-Wilkes 4, Richards 27090, Sanane220, Schlieben 4393, Tanner 627, 4922: Williams 690. Togo: Breteler 7229, Ern et al. 776, Robertson 72; Schafer 7628. Uganda: Brown 63; Dummer 177, Elliot 6567, 7818; Katende 2601, 2219; Loveridge 190, 421; Ross 889; Rwaburindore 326, 998, 1474, 2035, 2676, es 2214, 2275, 2550, 3283. Zaire: Bamps 3003; Callens s.n. July 1958), s.n. (Aug 1958): de Craene 231; Evrard 6350; Jean Lebrun 5352, 8241, 9132; Jean Louis 20, 709, 4509, Michel 2922. Michel & Reed 92; Robyns 2296, Toussaint 2417. Zambia: Angus 208, 1279, 1610, Bees 782, 786, Best 328, 327, Cun 8605; Davies 1166, Kabisa 22; ey 66; Richards 9421, 9553,15136, 15148, 15195, aes White 3222. Zimbabwe: Bayliss 10087, 10488, Biegel 3126, Chase 4887; Daves tee Mshasha 66; Muller 2475, 3403, 3087; Noel 2436, Nyarini 168; Plowes 1622; Rushworth 689; Siu 2194: West 2161; Wild 5781 Clematis longicauda Steud. ex A. Rich —Ethiopia: Ash 2767, de Wilde & de Wilde-Duyfjes 9384, 10133, 10434: J. De Wilde 6118, 6720, Meyer 7718, Pichi-Sermolli 2352, Schimper 1284. Clematis simensis Fresen—Burundi: Reek mans 7905, 10671. Cameroon: Thomas 2637. Ethiopia: Ash 35, 679, 1294: J. de Wilde 3062, 4077, 6224. de Wilde & de Wilde-Duyfjes 8238, 8374, 8966, 8984, 10145; Pichi-Sermolli 135, 138; Schimper 1512; Westphal & Westphal-St Bibi cei 2350, 3062. Kenya: Agnew 7707, 9438, D’Arcy 7364, Greenway & Kanuri aes 14895, Maas 6049; Mabberley & McCall 12; Robertson 1567; Taylor 125361455.M i: Brass 16106, 16860, 168: 36; ( chapman & Chap- man 7807; LaCroix 3103; 3; Pawel 7014, 97 11, 10000; an llips 1704, 2825. Nigeria: Sanford 5494. Rwanda: Fossey B/7.T i & Abdallah 1745; lwarsson et al. 1065; Jefford et al. 1745; Mathias & Tay- lor A86; Mwasumbi 16316; Richards 16802. Uganda: Katende 3305, Robertson 1567. Zaire: Louis 4817. Cl hehensis Engl.—Malawi: Pawek 7910; Phillips 136, 337,452, 1360. Mozambique: Sousa en Tanzania: Brummitt Pe) 18141, Gereau et al. 2833: Goetze 579 (photo, B); Magogo 255; Stolz Suleman & Fundi 16. Zaire: Bambs & Malaisse 8407; Kassner 3347 (photo, BM); Shantz a et ea Clematis villosa subsp. stanleyi (Hook.) Kuntze—Angola: Exell & Weeoue S.A. eee Rodin 9349; aaa Namibia: Se ydel 2067. South Africa: Bayliss 3145, 4664, Bou 571: Burke 1853; Joffe 2: Liebenberg 8815, Mogg 18811, 21054, Rand 1282; Scheepers 1486; side) 156 Thode aah Welman 52: Zimbabwe: SeD Re: — 353; ae 1647, Gonde 386, Leach 8200, Miller 2087, 218 - Mshasha 162; Opperman s.n.(1969). Zambia: Grant 4518, Harder pe 4018; Nawa et al. 123, EF White 1918. > oy ral ill kirkii Oliv.) Brummitt—Angola: Exel lemat | & Mendonca 120, 1702, 1741, 160. Malawi: Banda 1509; Brass 17453: Chapman & Chapman 7326, Grosvenor & Renz 933; Patel et al. 1484; Pawek 5463, 6634, 8253: Phillips 768A, 1181, 1449, 3671; Salubeni et al. 2518. Mozambique: Correia 215, Gomes & Sousa 1662; Torre & Paiva 379. Tanzania: Bally & Carter 16461, Gribb et al. 11247, Kayombo & Kayombo 212; Lovett 1482; Lovett et al. 1899, Muumba DSM 3029, Paget- Wilkes 785, Prins-Lambert 392: Stolz 146; Thulin & Mhoro 3125. Zaire: Brookes et al. 191, Robyns 1587. Zambia: Lusaka Natural History Club 228: Richards 22153. Zimbabwe: Bayliss 10659; Chase 7926; Davies 47; Gote 198; Rhodin 4368; Rutherford-Smith 482, West 254 Clematis villosa subsp. oliveri (Hutch.) Brummitt—Burundi: Reekmans 2756, 3824, 5337, 5527, 6754. Cameroon: De Wilde & de Wilde-Duyfjes 2344, Leeuwenberg 7619, 7668; T Thomas 6072. Migens: Ekwuno etal. 291, Sanford 5162, 6176; Witet al. ee patina Mic pee Tanzania: Gered ayombo 4773; Haarer 2251, Lovett & Congdon 2912; & Mabira 2251: Shabani 973. prun 9519a; Malaisse 11214; Miciele Reed 2,232. Zambia: Harder et al. 2610. Zaire: Grant 4501a; Le 876 BRIT.ORG/SIDA 21(2) Clematis villosa ~ eas villosa—Angola: Exell & Me foie — 2503, 2967. Burundi: Reekmans 17 tal. 723; Haarer 2252; Paget- Wilke Clematis viridiflora Bertol —Mozambique: Correia & Marques 2189, Edwards & Vahrmeyer 4281; Lemos & Balsinhas 37, deKoning & Hiemstra 9022. Tanzania (Zanzibar): Haji DSM 4026. matis welwitschii Hiern ex Kuntze—Angola: Carisso & Mendonca, 575; Exell & Mendonca 358, oe 396, 1076; Welwitsch 1215a (photo of type of C.commutata at BM), Welwitsch oe of type of C. welwitschii at BM). Cameroon: Breteler et al. 2448. Malawi: Chapman & Chapman 9008, LeCroix 3020, Tawakali & Kaunda 190; Lovett & Congdon 2885, Pawek 6255, 7151, 8405, 12644, 12686, 13658, 14352. Mozambique: Edwards & Vahrmeyer 4281. Tanzania: Bidgood & Congdon 150, Gereau & Kayombo 4228, Mwangoka & Kayombo 872; Mwasumbi 16172, Richards 20438. Zaire: Quarre 5383. Zambia: Lumba Natural History Club 38; Schmidt et al. 1333; Vesey-Fitzgerald 1172, White 2643. Zimbabwe: 10086; Leech & Brunton 9841, Rodin 4369. Q. matis welwit nex Kuntze—Angola: Carisso & Mendonca, 575; Exell & Mendonca 358, 396, 1076; ae ‘ D154 (photo of type of C.commutata at BM); Welwitsch 1217 (photo of type of C. welwitschii at BM). Cameroon: Breteler et al. 2448. Malawi: Chapman & Chapman 9008; LeCroix 3020, ae iG nouns 190; Lovett & Congdon 2885, Pawek 6255, 7151, ae ali 12686, 13658, 14352.0 is& Vahrmeyer 4281) Tanzania: Bidgood & Congdon 1 Kayomb« 4228; Vivanco & Kayombo 872, Mwasumbi 16172; Richards 20438. Zaire: Quarre 5383. Zambia: Lumba Natural History Club 38; Schmidt et al. 1333, Vesey-Fitzgerald 1172, White 2643. Zimbabwe: Bayliss 10086, Leech & Brunton 9841; Rodin 4369. — ACKNOWLEDGMENTS lam grateful to the staff of the Missouri Botanical Garden for making their col- lection of African Clematis available to me for this study, and to Richard Brummitt of the Royal Botanic Gardens, Kew, and Gretchen Walters of the Mis- souri Botanical Garden, for comments on earlier versions of the manuscript. REFERENCES Beenie, HJ. 1989. The reinstatement of Clematis sigensis Engl. (Ranunculaceae). Utafiti 2(1):16. BerHaur, J. 1967. Flore du Senegal. 2". Ed. Clairafrique. Dakar Brenan, J.P.M.1978.Some aspects of the phytogeography of Tropical Africa. Ann. Missouri Bot. Gard. 65:437-478. Brummit, R.K. 2000. Inclusion of Clematopsis Hutch. in Clematis L. (Ranunculaceae). Kew Bull. 55:97-108. Crossy,M.R. 1978. Systematic studies in Africa: the twenty-fourth Systematics Symposium. Ann. Missouri Bot. Gard. 65:367-368. Durano, T. 1895. Conspectus florae Africae. Jardin botanique de |'état, Bruxelles. Pp.1-8. Essic, F.B. 1990. The Clematis virginiana (Ranunculaceae) complex in the southeastern United States. Sida 14:49-68 Exett, A.\W. and Menoonca 1937. Conspectus florae Angolensis. Lisboa, Portugal. 1:1- Exett, A.W. and E. Mitne-ReDHead. 1960. Clematis. In: A.W. Exell and H. Wild, eds. Flora Zambesiaca. Pp. 89-93 Grey-Witson, C. 2000. Clematis, the genus. Timber Press. Portland, OR. ESSIG, CLEMATIS IN SUB-SAHARAN AFRICA 877 JoHNnson, M. 1997. Slaektet Klematis. Magnus Johnsons Plantskola AB. Soedertaelje. Sweden. HutcHINsON, J. and J.M. Datziet. 1954. Flora of West Tropical Africa. London, Crown Agents for Oversea Governments and Administrations. Pp.64—-65. Letty, C. 1962. Wild flowers of the Transvaal. Trustees of the Wild Flowers of the Transvaal Book Fund. P. 140. Ouiver, D. 1868. Flora of Tropical Africa. London, L. Reeve and Co. 1:7. Tamura, M. 1967. Morphology, ecology and phylogeny of the Ranunculaceae VII. Sci. Rep. Osaka Univ. 16:21-43. Texatay, D. 1987. Revision of the genus Clematis L.in Ethiopia. Belmontia 19(108):1-42. THULIN, M. 1993. Flora of Somalia. Royal Botanic Gardens, Kew. 1:22. Wuite, F. 1962. The forest flora of northern Rhodesia. Oxford University Press. Pp. 46-47. BRIT.ORG/SIDA 21(2) BOOK NOTICES Southeastern Botany RONALD W. GiLMour. 2002. Foundations of Southeastern Botany: An Annotated Bibliography of Southeastern American Botanical Explorers Prior to 1824. (ISSN 0008-7475, pbk.). The Southern Appalachian Botanical Society, Dr. Patricia Cox, TVA Heritage Program, PO Box 1589, Norris, TN 37828, US.A (Orders: same). Price not given, 142 pp., 6 3/4" x 10" Abstract: “This paper lists published sources pertaining to the lives and work of seventy individuals who contributed to early botanical knowledge of the Southeastern United States. General sources, primarily biographical compilations and scientific bibliographies, are listed at the beginning of the paper, followed by entries for individual botanists. For each botanist, the listed information includes name (withany variants), place and year of birth and death, location of manuscripts and plant speci- mens, citations for published portraits and handwriting samples, lists of any plant or fungal genera named for the botanist, a brief summary statement about the person's significance to Southeastern botany, and entries for published sources — iining to the botanist. Works both by and about each nroughout the botanist are listed with annotations. All et are cross-referenced parenthetically t text.” Davin E. ALLEN and GapricLLe HATFIELD. 2004. Medicinal Plants in Folk Tradition: An Ethnobotany of Britain & Ireland. (ISBN 0-88192-638-8, hbk.) Timber Press Inc. 133 S.W Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@tim ber press.com, 503-227-2878, |- 800-327-5680, 503-227-3070 fax). $29.95, 432 pp., 31 color plates, 57 b/w illus., | map, 6" x 9". Review forthcoming in Sida 21(3). SIDA 21(2): 878. 2004 THE ACTIN I INTRON—A PHYLOGENETICALLY INFORMATIVE DNA REGION IN CLEMATIS (RANUNCULACEAE) Jonathan M.Slomba, James R. Garey, and Frederick B. Essig Department of Biology University of South Florida Tampa, Florida 33620, U.S.A. essig@chumal.cas.usfedu ABSTRACT As part of a search for DNA regions suitable for phylogenetic analysis in the genus Clematis (Ranunculaceae), the nuclear-encoded Actin I intron was employed in a preliminary sampling of a small number of species selected to represent the major subdivisions of the genus. This DNA region was found to be more informative and to provide a more robust phylogenetic tree than chloroplast DNA regions. Trees generated were consistent with phylogenetic hypotheses based on morphology and with published molecular analyses. Moreover, three species of the section Crispae (subgenus Viorna) native to Florida, considered closely related on morphological grounds, formed a well-sup- ported clade, and were distinguishable from one another. This hat this DNA region might be a useful tool for distinguishing groups of closely-related species, individual species, and possibly hybrids. RESUMEN @ dq 1 eal : JoDNA I j licic fil g Pee | género Cena @anuncalacesd) se eurnleo el intron Actin. I nuclear en un muestreo preliminar de un DEqUEND: numero ods especies seleccionadas eae representar las grandes subdivisiones del género. laba un arbol filogenético mAs consistente las senioues de DNA plastidial. ree, oe penetra eran consistentes con la hipotesis et onic basadaen 2 monoNoe yco i ee tres especies e laseccién Cri i le Florid id las f en aspectos miortalesices ann un clado muy coherente, y eran pea una de otra. ea P sugiere que esta region del DNA puede ser un instrumento util para diferenciar grup muy emparentadas, especies individuales, y posiblemente hibridos. INTRODUCTION The infrageneric classification of Clematis (Ranunculaceae), a genus of more than 300 species distributed worldwide, has been uncertain pending definitive phylogenetic studies. Traditional classifications have relied primarily on floral characters for the major divisions of the genus (as in Tamura 1967). However, characters of seedling and juvenile morphology have been cited in recent de- cades as supporting a fundamental division in the infrageneric classification of the genus (Tamura 1987; Essig 1991; Miikeda et al. 1999). A specialized syn- drome of seedling and vegetative characters, featuring a suppressed hypocotyl SIDA 21(2): 879-886. 2004 880 BRIT.ORG/SIDA 21(2) and opposite seedling leaves (“Type II,” Essig 1991, “opposite” in Miikeda et al. 1999; see also Appendix 1) appears to have arisen from the more general Ranunculaceous type featuring an elongate hypocotyl and alternate seedling leaves (“Type |” or “alternate”), but it has been uncertain whether this morpholo- gical complex has arisen just once or more than once, as it is found in species formerly placed in different subgenera. We have been seeking appropriate molecular tools with which to resolve these and other phylogenetic questions within Clematis. The use of DNA se- quencing techniques has thus far been of limited success. Miikeda et al. (1999) utilized several chloroplast genes, including matK (maturase-encoding gene), trnK (UUU) intron, trnL (UAA) intron, the intergenic spacer between trnL and trnF (GAA), and the intergenic spacer between rbeL and atpB. Employing ap- proximately 4,400 bp. of sequence for the eight taxa included in the study, the team produced a tree that was consistent with Essig’s proposal, but was weak resol ved. Our own efforts with chloroplast DNA also produced weak results. A search for alternative tools led us to consider some nuclear DNA regions, which are expected to be more informative than chloroplast non-coding regions in determining species level phylogenies because the nuclear genome hasa sub- stitution rate 5 to 10 times faster than the chloroplast genome (Li 1997). In par- ticular, we have focused on a non-coding intron of the Actin] gene. Actin is one of the components of the cellular cytoskeleton, and is produced through the activity of a large multigene family (Moniz de Sa & Drouin 1996). The location of the Actin l intron is highly conserved across all angiosperm families, mak- ing the development of primers suitable for the PCR amplification of the intron possible. A preliminary test of this DNA region asa phylogenetic tool in Clematis was conducted using a small sampling of species representing the major sub- divisions of the genus. — y o MATERIALS AND METHODS Materials were obtained from the Chicago Botanical Garden and the Univer- sity of South Florida Botanical Garden (Table L). Samples were selected to rep- resent the major subdivisions of the genus Clematis. They include several spe- cies with Type I vegetative morphology, and several with Type II vegetative morphology, while also representing the traditional sections Clematis and Viorna (as in Tamura 1967), and the rearranged sections (elevated to subgen- era) of Tamura (1987) (TableL). The traditional sections each contained subsec- tions with Type Land Type II morphologies, and in his revision, Tamura (1987) reorganized his classification to reflect those different morphologies. Note that two of the species included in this study were realigned in that taxonomic shift. Clematis terniflora in subsection Rectae was formerly included in section Clematis, while C. stans, in subsection Tubiflorae was formerly included in sec- tion Viorna. This analysis is thus a preliminary test of that taxonomic revision. — SLOMBA ET AL., ACTIN | GENE REGION IN CLEMATIS 881 Table 1.Taxa included in the analysis (with seedling morphology type indicated as | or Il). Species Classification (Tamura 1987) Voucher Clematis ae Walt. Viorna: Crispae Il) Arias 71 (USF) Clematis cri Viorna: Crispae Il) ae 011001-6 (USF) Clematis en & A. Gray Viorna: Crispae II) ssig 011001-7 (USF) Clematis terniflora DC Flammula: Rectae II) ssig 860904-1 (USF) Clematis virginiana L. (Clematis: Dioicae |) Be go B.G.acc. # 356-81 5si i a Clematis stans Sieb.& Zucc (Campanella: Tubulosae |) Essig 011001- 3 (USF) Anemone pulsatilla var. cuaais L. (outgroup |) Essig 020305-2 (USF) The samples for this study also includes three species native to Florida that on morphological grounds appear to be closely related. Their inclusion provides a test of the resolving power of the Actin | DNA sequence. A species of Anemone (A. pulsatilla var. vulgaris L) was chosen as the outgroup. Anemone has traditionally been identified as closely related to Clema- tis,and A. pulsatilla shares with Clematis the very distinctive elongate styles of the mature achenes. A number of recent phylogenetic studies Johansson & Jensen 1993; Hoot 1995, and Kosuge et al. 1995) have identified a clade that in- cludes Clematis along with Anemone, Pulsatilla (sometimes treated as a segre- gate of Anemone), Knowltonia, Hepatica, and sometimes Ranunculus and/or Trautvettaria. A more comprehensive study will include more of these genera as outgroups. An unnamed species of Anemone was also used as the outgroup in the study by Miikeda et al. (1999). Angiosperm Actin (acl) gene sequences from a broad range of taxa were obtained from Genbank (Arabidopsis, Zea, Oryza, and Glycine (accession #’s M20016, J01238, X15865, and JO1298, respectively). These sequences were aligned using Clustal X (Thompson et al. 1994; Higgins et al. 1996). The primer sequences were selected from the alignment by anchoring the forward primer ina highly conserved (relatively guanine and cytosine rich [48%]) coding region just down- stream of the intron. The reverse primer was anchored in a highly conserved coding region just upstream of the intron (Table 2). This primer set corresponds toa region of approximately 300 nucleotides in the taxa listed above. Some specimens were deep frozen at -80° C before use, others were pre- pared immediately for extraction. Total genomic DNA was extracted from leaf samples following the modified CTAB protocol developed by Doyle & Doyle (1987) Polymerase chain reaction (PCR) was carried out on all extracted DNA samples using primers for the gene regions shown in Table 2. PCR reactions (am- plification) were carried out in 100 uL volumes, using a taq polymerase kit from Enzypol (Boulder CO), following their instructions. Thermal cycling param- eters were the same for all species: 1 min. initial denature at 95°C, followed by 35 882 BRIT.ORG/SIDA 21(2) Taste 2.PCR primers for Actin genes. Actin | forward: CCC GAA TIC CTT GTT TGC GAC AAT GGA AC Actinl reverse: CCC GAA TTC ACA ATT CCA TGC TCA AT cycles of 15 sec. at 95°C, annealing at 48 °C for 30 sec., a 90 sec. extension at 72°C, followed by a 10 min. hold at 72°C, and then a final hold at 4°C. The 300 bp PCR product of the Actin I intron was gel purified and cloned into a pBluescript vector and transformed into DH5p Escherichia coli cells (Gibco, Carlsbad CA). White colonies were picked from bacterial plates, grown insmall cultures with ampicillin and sequencing template prepared using the alkaline lysis method (Ausubel 2000). After purification, cycle sequencing re- actions were carried out in 200pL thin-walled capped tubes using a Perkin- Elmer (Foster City, CA) DYEnamic ET terminator cycle sequencing kit with an ABI Model 310 genetic analyzer. Phylogenetic trees were constructed from DNA sequence alignments gen- erated by Clustal X (Thompson et al. 1994; Higgins et al. 1996) and modified by hand using Genedoc (Nicholas et al. 1997). Neighbor-joining trees from molecu- lar data were made using MEGA 2.1 software (Kumar et al. date) using Kimura 2-parameter distances with 1000 bootstrap replicates. For comparison, a brief morphological analysis incorporating vegetative and floral characters used in recent classifications was carried out (Fig. 1B, and Appendices 1 & 2). The mor- phological tree was recovered using PAUP 4.0b10 (Swofford 2001) using an heu- ristic search with default parameters and 200 bootstrap replicates. RESULTS AND DISCUSSION The Actin intron sequences yielded an alignment of 316 bp with 27 variable sites, 17 parsimony informative sites and 28 sites with gaps. The alignment was relatively unambiguous and resulted in a neighbor-joining tree (Fig. 1A) with two well-supported clades that coincide with Type | and Type II seedling mor- phology as described by Essig (1991). A maximum parsimony tree based on morphology (Fig. 1B) recovered the type II clade, and illustrates the plesiomorphic distribution of the Type | character syndrome. The molecular analysis produced a strongly supported (bootstrap value 100%) derived clade containing Clematis crispa, C. reticulata and C. baldwinii. These are morpho- logically similar species native to the southeastern U.S. belonging to the tradi- tional group Crispae (variously designated as a section or subsection) in sub- genus Viorna, and exhibiting type Il morphology. There is a strong sister group relationship (97%) between this group and C. terniflora,a Eurasian species also with Type II morphology, but with panicles of small whitish flowers—a repro- ductive morphology syndrome it shares with Type | members of the traditional subgenus Clematis. Clematis virginiana and C. stans form a separate well-sup- SLOMBA ET AL., ACTIN | GENE REGION IN CLEMATIS 883 A. Actin Intron 54 C. reticulata O 100 C.crispae |g | p00) ne) . oe. a) @ of C. baldwinii = C. terniflora C. virginiana 80 | a TT = C. stans o Anemone peel 65 C. baldwinii B. Morphology e) . nn 63 C.crispae | |a se) TT Na . o | v1 C. reticulata = q a] C. terniflora C. virginiana < C. stans o Anemone 4 the Typ ll 1 Crisp d but ith 90 199 | I t | Pv i ly B M t y tree from morphological data (see Appendices 1 and 2). Type | and Type Il are synd f morphological characters, pri- ily of seedlings, as described in Essig (1991). Tt B indicates the origin of the Type Il syndrome. Crispae is be J po J th ° n fn. LA A | . £ } Ww (T 1987). J 884 BRIT.ORG/SIDA 21(2) ported clade (80%) and both have Type | seedling morphology, despite having different floral morphologies. The results of this preliminary analysis are consistent with those obtained by Miikeda et al. (1999). The species used in the two studies were different, but representative of the same infrageneric taxa. The results are also supportive of Tamura’s (1987) revised classification, and Essig’s (1991) proposal that taxa with type Il seedling morphology represent a monophyletic clade and might be placed together in a major infrageneric division of the genus. The results also confirm the close relationship of the 3 species of the subsec- tion Crispde occurring in the southeastern U.S.A., and appear to resolve those species from one another. The relationship of the three species was slightly dif- ferentinthe DNA analysis from that in the morphological analysis, or from what one would expect through conventional taxonomic analysis. More extensive sam- pling within species is needed, along with analysis of additional DNA regions, to fully evaluate the resolving power of the Actin | intron region at this level. Another discrepancy between the two analyses is the sister group relation- ship between C. virginiana and C. stans found in the molecular tree but not in the morphological tree. Too few taxa were included in this study to draw any conclusions about the deeper branches in the genus, however. A great many more taxa with both Type | and Type 2 morphologies exist. A more complete analysis will include a great many more of the species of this large genus, and in particular, as many of the recognized infrageneric taxa (sections, subsections) as possible, along with a comprehensive morphological analysis, in order to fully understand the phylogeny of this genus and develop a definitive infrageneric classification. In conclusion, the results of this preliminary analysis are consistent with taxonomic concepts based on morphology and with other DNA-based analy- ses, and also appear to discriminate among fairly closely related species. There- fore, it appears that the Actin I gene region will be a very useful tool for the analysis of infrageneric relationships in Clematis, and likely in other an- giosperm genera. APPENDIX | Characters used in morphological analysis. Note: characters l-4 are the primary features distinguishing the Type I (0) from the Type II (1) syndrome; characters 6-8 are the floral characters traditionally cited in distinguishing subgenus Clematis from subgenus Viorna. 1.Seedlings with hypocotyl elongate (0) vs hypocotyl suppressed (1) 2. Seedling leaves alternate (0) vs leaves opposite (1) 3. Eophylls 3-lobed (0) vs eophylls elliptic (1 4, Leaves dentate (0) vs leaves entire (1) 5.Stems erect (0) vs stems vining (1) — SLOMBA ET AL., ACTIN | GENE REGION IN CLEMATIS 885 6. Flowers with sepals spreading to reflexed nom De base(Q) vs HOWEts tubular, urceolate or cam- panulate with sepals spreading at the tips p with strongly reflexed limbs 2 7.Flowers colored (0) vs flowers white to cream (1) 8. Stamens with filaments glabrous (0) vs filaments hirsute (1) 9. Achenes narrow, turgid (0) vs achenes broad, flattened (1) APPENDIX 2. SPECIES / MORPHOLOGICAL CHARACTER MATRIX 1 2 3 4 5 6 7 8 9 Anemone pulsatilla 0 Clematis stans 0 Clematis virginiana 0 Clematis terniflora ] Clematis reticulata ] Clematis crispa 1 Clematis baldwinii ] a ee a ao a> a) 0 0 0 ] 0 0 1 0 ] ] ] ] ] ] -OC0O000-0 MN =| OO-o aoo--a00 a a ao a a) ACKNOWLEDGMENTS This publication was derived from the Master’s thesis of the first author. We acknowledge the support of NSF grants CHE-0221834 and DEB-0344372. We thank the Chicago Botanic Garden for providing materials. We thank Nancy Moreno and John Syring for providing valuable review comments. REFERENCES Aususel, F.M., R. Brent, R.E. Kinston, D.D. Moore, J.G. SeioMANN, J.A. SmitH, and K. StRUHL. 2000. Current protocols in molecular biology. Greene Publishing Associates, NY. Doyte, J.J.ANbD J.E. Doyte. 1987.A rapid DNA isolation procedure for small quantities of fresh leaf material. Phytochemistry J.19:11-15. Essia, F.B. 1991. Seedling morphology in Clematis (Ranunculaceae) and its taxonomic im- plications. Sida 14:377-390. Hicains, D.G., J.D. THompson, and TJ. Gisson. 1996.Using CLUSTAL for multiple sequence align- ments. Meth. Enzymol. 266:383-402. Hoor, $.B. 1991. Phylogeny of the Ranunculaceae based on epidermal microcharacters and macromorphology. Systematic Bot. 16:741—755. Hoot, $.B. 1995. Phylgenetic relationships in Anemone (Ranunculaceae) based on DNA restriction site variation and morphology. PI. Syst. Evol., suppl. 9:195—200. JOHANSSON, J.T. and R.K. Jensen. Chloroplast DNA variation and phylogeny inof the Ranunculaceae. Pl. Syst. Evol. 187:29-49. Kosuce, K., K. Sawapa, T. Denna, and K. Watanabe. 1995. Phylogenetic relationships of some genera in the Ranunculaceae based on alcohol dehydrogenase genes. PI. Syst. Evol., Suppl. 9:263-271. Kumar, S., K. Tamura, |.B. JAKogsen, and M. Ne. 2001. MEGA2: Molecular evolutionary genetics analysis software, Arizona State University, Tempe. 886 BRIT.ORG/SIDA 21(2) Li, W.H. 1997. Molecular evolution. Sinauer Assoc.|Inc., Sunderland, MA. Mukepa, O.S. Koa, T. Hanpa, and T. Yukawa. 1999. Subgeneric relationships in Clematis (Ranunculaceae) by DNA sequences. In: S. Andrews, A.C. Leslie, and C. Alexander, eds. Taxonomy of cultivated plants. Third International Symposium. Royal Botanic Gardens. Kew. Pp. 355-358. Moniz de Sa, M.and G. Drouin, 1996 .Phylogeny and substitution rates of angiosperm actin genes. Molec. Bio. Evol. 13:1198-1212. NicHotas, K.B., H.B. NicHotas, and D.W. Deerrieto Il. 1997.GeneDoc: analysis and visualization of genetic variation, Embnew. News 4:14. Tamura, M. 1967. Morphology, ecology and phylogeny of the Ranunculaceae VII. Sci. Rep. Osaka Unv. 16-2:21-43. Tamura, M. 1987.A classification of the genus Clematis. Acta Phytotax. Geobot. 38:33-44. THomeson, J.D., D.G. Hicains, and TJ. Gisson. 1994, CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-spe- cific gap penalties and weight matrix choice. Nucleic Acids Res. 22:4673-4680. NEW COMBINATIONS IN NORTH AMERICAN CARYOPHYLLACEAE J.K. Morton Department of Biology University of Waterloo terloo, Ontario, CANADA N2L 3G] jkmorton@sciborg.uwaterloo.ca ABSTRACT The following I d.Cerastium velutinum Raf. var. villosissimum (Pennell) J.K. Morton, Silene drummondii Hook. subsp. striata (Rydb.) J.K. Morton, Sil soaiatied Cav. ae californica (Durand) J.K. Morton, Silene ostenfeldii (AE. Porsild) J.K. Morton and St Willd. ex Schlecht. subsp. (Baldw. ex El.) J.K. Morton. RESUMEN Se proponen las siguientes nuevas combinaciones. Cerastium velutinum Raf. var. villosissimum (Pennell) J.K. Morton, Silene drummondii Hook. subsp. striata (Rydb.) J.K. Morton, Silene laciniata Cav. subsp. californica (Durand) J.K. Morton, Silene ostenfeldii (A-E. Porsild) J.K. Morton y Stellaria cuspidata Willd. ex Schlecht. subsp. prostrata (Baldw. ex Ell.) J.K. Morton. During the preparation of treatments of Cerastium, Silene and Stellaria for the Flora of North America, the need for the following new combinations became apparent. NEW COMBINATIONS Cerastium velutinum Raf. var. villosissimum (Pennell) J.K. Morton, comb. nov. BASIONYM: Cerastium arvense L. var. villosissimum Pennell, Bartonia 12:11. 1931. TyPE: U.S.A: PENNSYLVANIA. Chester Co. rocky cliff, serpentine below Lees Mills by Octoraro Creek, 21 Sep 1920, Pennell 10767 (HOLOTYPE: PH; ISOTYPE: NY). Cerastium arvense L.in North America consists of at least three species. One of these, Cerastium velutinum Raf., includes the plant that Pennell (1931) described as Cerastium arvense L. var. villosissimum. The new combination is required to accommodate this change. Silene drummondii Hook. subsp. striata (Rydb.) J.K. Morton, comb. et stat. nov. BASIONYM: Lychnis striata Rydb., Bull. Torrey Bot. Club 31:408. 1904. TyPE: COLORADO: Cam- eron Pass, 1000 ft, 30 Jul 1896, ae s.n. (HOLOTYPE: NY; ISOTYPE: MO). Silene drummondii contains two taxa, subsp. drummondii which is character- istically a prairie taxon, and subsp. striata which is associated with the Rocky Mountains from near the Canada border southwards. Though the two subspe- cies are clearly distinct in their extreme forms they intergrade where they come into contact. SIDA 21(2): 887- 888. 8.2004 888 BRIT.ORG/SIDA 21 N ) Silene laciniata Cay. subsp. californica (Durand) J.K. Morton, comb. et stat. nov, BASIONYM: Silene californica Durand, J. Acad. Nat. Sci. Philadelphia, ns. 3:83. 1855. TYPE: CALI- FORNIA: Pratten s.n. (HOLOTYPE: P. nv. The Mexican Pinks (S. laciniata, S. californica and S. greggii A. Gray) have fre- quently been regarded as distinct species. In the context of North America this is probably appropriate, for all three have clearly recognizable morphological characters. However, in Mexico, where their distribution is centred, many col- lections show intergrading characters and are not readily identifiable. For this reason, subspecific status is more appropriate. Silene ostenfeldii (A.E. Porsild) J.K. Morton, comb. nov. Basionym: Melandrium ostenfeldii A.E. Porsild, Sargentia 4:37. 1943. Type: CANADA: Narakay Island, Dease Arm, Gre Bear Lake, Northwest Territories, | Aug 1928, A.E. & R.T. Porsild 4839 (HOLOTYPE: CAN). Silene taimyrensis (Tolm.) Bocquet is the name that has been used for this spe- cies since Bocquet created the combination in 1967. Unfortunately, Bocquet did not have access to the type specimen on which the name is based (Jolmatchew 762; holotype, L). Recently Petrovsky & Elven (in the on-line Pan Arctic Flora- www.mun.ca/biology/delta/arcticf) reported that they had examined Tolmatchew’s specimen and consider it to be a form of Silene involucrate (Cham. & Schlecht.) Bocquet. The combination Silene ostenfeldii has apparently not been validly published. Electronic publication is not acceptable under Article 29.1 of the International Code of Botanical Nomenclature (Greuter et al. 2000). Accordingly the combination is validated here. Stellaria ee ay Willd. ex Schlecht. subsp. prostrata (Baldw. ex EI.) J.K. Mor- mb. et stat. NOv. BASIONYM: Siellavia prostrata Baldw. ex EIL., Sketch Bot. S. Caro- lina I: ae 1821. Type: FLORIDA: on the island of Fort George, East-Florida (Apalachicola), without date, Baldwin sn. (HOLOTYPE: NY) Stellaria cuspidata and S. prostrata frequently intergrade and hence are better treated as subspecies. The former tends to be montane and the latter to be a lowland weed. REFERENCES BocaueT, G. 1967. Physolychnidum olim Gastrolychnidum nomenclaturae fundamentum includens combinationes taxaque nonnuila Silenes generis. Candollea 22:1—38. GreuTer, W., J. MCNeiLt, FR. Barrie, HM. Burbert, V. Demoutin, T.D. Fiticueiras, DH. Nicotson, PC. Sitva, J.E. Skoa, P. TreHane, NJ. Turtanp, and D.L. Hawe«swortH (eds.). 2000. International code of botanical nomenclature (St. Louis Code) Adopted by the Sixteenth International Bo- tanical Congress St. Louis, July-August 1999. Koeltz, Konigstein PENNELL, F.W.1931.On some critical species of the serpentine barrens. Bartonia 12:1—23. TONESTUS KINGII AND T. ABERRANS ARE RELATED TO EURYBIA AND THE MACHAERANTHERINAE (ASTERACEAE: ASTEREAE) BASED ON NRDNA (TS AND ETS) DATA: REINSTATEMENT OF HERRICKIA AND A NEW GENUS, TRINITEURYBIA Luc Brouillet L. Urbatsch rbier Marie-Victorin LSU Herbarium and Biological Sci. Dept. Institut de recherche en biologie végétale 202 Life Sciences Bldg. Université de Montréal Louisiana State University 4101 Sherbrooke St.E Baton Rouge, Louisiana 70803, U.S.A. Montreal, Québec, CANADA, H1X 2B2 leu@Isu.edu luc.brouillet@um (author for an R.P. Roberts SU Herbarit logical Sci. Department 202 Life Sciences Bldg. Louisiana State University Baton Rouge, Louisiana 70803, U.S.A. and Department of Biological Sciences Towson University, Towson, Maryland 21252-0001, U.S.A. ABSTRACT Phylogenetic analysis of nrDNA (ETS and ITS) sequence | for 50 species of Astereae (Asteraceae) belonging to the Eurybia-Machaerantherinae clade show that Tonestus kingii and T. aberrans be- long to this group. The genus Herrickia is reinstated and expanded to include H. kingii as well as members of Eurybia section Herrickia sensu Nesom (1994). 7 new genus, Triniteurybia, is estab- lished for T. aberrans, which is sister to the Machaerantherinae. RESUMEN El andlisis filogenético de secuencia de bases de nrDNA (ETS y ITS) de 50 especies de Astereae (Asteraceae) pertenecientes al clado Eurybia- Maen Gera rinae muestra que Tonestus kingii y T. jai aberrans pertenecen a este grupo. El género Herric para incluir a H. kingii ri suN (1994) Se establece un nuevo género, asicomoa miembros de Eurybiaseccién H Triniteurybia, para T. aberrans, que es “sister group’ sds Machaerantherinae.. INTRODUCTION Several species of tribe Astereae in North America have proven difficult to as- sign taxonomically to genera or groups. Molecular phylogenetic studies, nota- bly following the seminal paper of Noyes and Rieseberg (1999), have helped to solve the relationships of many of these taxa. Nevertheless, the disposition of SIDA 21(2): 889-900. 2004 = 890 BRIT.ORG/SIDA 21(2) some species still remain controversial, notably species recently included in Tonestus by Nesom and Morgan (1990), T. aberrans (A. Nelson) G.L. Nesom & Morgan (basionym Macronema aberrans A. Nelson), and by Nesom (1991), T. kingii (D.C. Eaton) G.L. Nesom (basionym Aster kingii D.C. Eaton). The basis for inclusion of these species in Tonestus were the woody caudices, leaf shape and anatomy, eglandular (or sparsely short-stipitate-glandular) herbage, mostly single-headed capitulescences and foliaceous out however). Nesom and Morgan (1990) did not discuss the specific reasons for the inclusion of Macronema aberrans in Tonestus, but pointed out its similarities to T-graniticus. Nesom (1991) reviewed the detailed taxonomic history of Aster kingii, which has also been named Machaeranthera kingii (D.C. Eaton) Cronquist & Keck, and he pointed out that various authors had seen relation- ships of this species with genus Asters.l, notably western members of Eurybia and Oreostemma. The similarities invoked to link this species to Tonestus are those cited above. The only discordant feature in the group would be its white rays ina yellow-rayed group. All species placed in Tonestus are n=9, a primitive and frequent number in the tribe. A recent molecular phylogeny of Chrysothamnus and related Solidagininae (Roberts & Urbatsch 2004) suggests, however, that Tonestus sensu Nesom is polyphyletic. Further investigations by Urbatsch and colleagues (unpublished) suggested that T. aberrans and T. kingii might better be placed near Eurybia. Likewise, a study of the position of the North American asters within tribe Astereae had suggested that T. kingii was better placed with Eurybia and Oreostemma, two North American segregates of Asters.l. (see summarized phy- logeny in Semple et al. 2002) that will be called henceforth the eurybioid lin- eage. Therefore, in preparation for the treatment of Eurybia and relatives : the Flora of North America, we are using ITS and 3'ETS molecular phylogenetic data to investigate the taxonomic position of T. aberrans and TI’ kingii within the Astereae. These molecular markers have proved useful in determining the position of taxa in tribe Astereae (e.g, Noyes & Rieseberg 1999; Roberts & Urbatsch 2004). We show that these species belong to the eurybioid grade within the North American clade and are presenting the nec- essary combinations to reflect this phylogenetic position. Genus Herrickia is reinstated and T: kingii transferred to it as H. kingii (along with congeners H. glauca and H. wasatchensis), and a new monotypic genus is described to ac- commodate T.aberrans, Triniteurybia. These names are subsequently used in the paper. phyllaries (not in T.aberrans (Asteraceae MATERIAL AND METHODS Samples were preserved either as frozen leaf material, in silica gel, or taken from herbarium specimens. Data for the Machaerantherinae were taken from Markos BROUILLET ET AL., THE EURYBIA 391 and Baldwin (2001). Extracti d molecular methods used h described in Roberts and Urbatsch (2004) (Urbatsch laboratory) and in Fougére-Danezan et al. (2003) (ITS, Brouillet laboratory). For ETS (Brouillet laboratory), the prim- ers Ast-8 (Markos & Baldwin 2001) and 18S-2L (Linder et al. 2000) were used, with PCR conditions similar to ITS (above); this resulted in longer ETS sequences than those produced in the Urbatsch laboratory (T. kingii and T. aberrans). Sequences were input into already aligned matrices of ITS and ETS for the Astereae and manually adjusted. All new sequences used in this study are de- posited in GenBank under the accession numbers provided in Table 1. Sources for already published sequences are provided also. Preliminary parsimony analyses were done on the full ITS (more than 500 taxa in Astereae) and ETS (106 taxa) matrices, using PAUP* 4.0b10 (Swofford 2002). Resulting trees (not shown) clearly indicate that T. kingiiand T. aberrans are members of the Eurybia complex (eurybioids) at the base of the Machaerantherinae (see Semple et al. 2002), but the trees were unresolved (polytomy) in the current region of interest. Given that the trees were not in conflict, data were combined for 50 taxa for which both sets were available, in an attempt to better define the position of these species within the Eurybia complex. The total number of characters in the combined ETS + ITS matrix is 1149, of which 195 are parsimony informative; few phylogenetically informa- tive indels are found in the reduced taxon matrix (mostly in the ITS portion) and they were not coded as distinct characters. The matrix was subjected to parsimony analysis using PAUP* 4.0b10 (TBR, characters unordered, unweighted, gaps as missing, random addition, Multrees in effect), and to Baye- sian analysis using MrBayes3 (Ronquist and Huelsenbeck, 2003). For the latter, the following parameters were applied: 4 chains, 1,000,000 generations, burn in 100,000, every 10,000th tree saved, model GTR+ gamma+gamma inv, nst=6. Analyses were run to completion. Bootstrap and jacknife supports were calcu- lated for the parsimony analysis (5000 replicates, TBR, random addition); for the jacknife, 50% of characters were deleted in each run. Strict and 50% major- ity rule consensus trees were drawn for the parsimony analysis, and posterior probabilities were input onto the 50% majority rule tree for the Bayesian analy- sis. Trees were rooted using Chloracantha spinosa and Canadanthus modestus. RESULTS Parsimony analysis of the ETS +1TS matrix yielded 6910 trees of length 687, Cl 0.662, RI 0.812 and RC 0.538. In the strict consensus tree (not shown) the eurybioids + Machaerantherinae form a clade, within which Oreostemma, Eurybia and Eurybia sect. Herrickia form a polytomy sister to a T. aberrans- Machaerantherinae clade. The parsimony 50% majority rule tree is similar to that shown for the Bayesian analysis (Fig. 1), which resulted in 9002 Taste 1. List of voucher specimens for the | for the current study, and sources of previously published data. Current species names (e.g., Morgan & Hartman 2003){ Sida 20:1403} are eid and original published names are provided in Saas when differing (as deposited in GenBank; Narkos & Baldwin 2001). For eurybioids, names proposed here are used and formerly used ones are in parenthese Species Source (collectors, number, GenBank accession number Herbarium or citation) Canadanthus modestus (Lindl.) G.L.Nesom Semple 10639 (WAT) AY772432 AY772446 pilose eG (Bentham) G.L.Nesom Spellenberg 13101 (MT) AY77243] AY772445 (Torr & A. a one Semple 10419 (WAT) AY772430 AY77/2444 var, hae ic POLY G.L.INesom Tri ar . Nelson) Brouillet, Urbatsch & R.PRoberts clone 1 Urbatsch 7812 (LSU) AY77 2426 AY772440 eeu ens (A. Nelson) GLN m & D.R. Morgan) Triniteurybia aberrans (A. Nelson) acne G.L.Nesom & D.R.Morgan) clone 2 Urbatsch 7812 (LSU) AY772427 AY77244] (Tonectiie aberr, He Nelo n) Herrickia kingii (D.C.Eaton) Brouillet, Urbatsch & R-PRoberts clone 1 Garrett 1576 (US) AY772428 AY772442 Tonestus kingii (D.C.Eaton) G.L.Nesom) Herrickia kingii (D.C.Eaton) Brouillet, Urbatsch & R.PRoberts clone 2 GS. Goodrich 16357 (UT) AY77242 AY77 2443 (Tonestus kingii (D.C.Eaton) G.L.Nesom) Herrickia horrida Wooten & Stand. Spellenberg & Fletcher AY772425 AY772439 7 (NMC) be glauca (Nutt.) Brouillet (Eurybia glauca (Nutt.) G.L.Nesom) Semple 5 . (WAT) AY772424 AY772438 ta(L.) GL.Nesom Semple 10710 (WAT) AY772423 AY77 2437 Ennio id Sie Aiega (Michx.) G.L.Nesom Semple i 7 (WAT) AY772422 AY772436 Eurybia sibirica (L.) G.L.Nesom Semple 10627 (WAT) AY77242)] AY772435 Eurybia eryngiifolia (Torr. & A.Gray) G.L.Nesom Semple 10557 (WAT) AY772420 AY772434 Dieteria bigelovii (A. Gray) D.R.Morgan & R.L. Hartman Semple 10468 (WAT) AY772419 AY772433 (Machaeranthera bigelovii (A.Gray) Greene) 768 (Z)L2 Vvals/O¥O'LIYa TABLE 1. continued Species Source (collectors, number, Herbarium or citation) ITS GenBank accession number ETS identalis (H.M.Hall) D.D.Keck (Lessingia occidentalis H M. Hall) M.Lane) Haplopappus foliosus DC. Haplopappus glutinosus Cass. Haplopappus macrocephalus (Less.) DC. Haplopappus marginalis Phil. Haplopappus paucidentatus Phil. reladia a detonsa (Greene) Greene guarrosa (Hook. & Arn.) Greene var. grindelioides Hosarai ia whytnei (A.Gray) Greene Isocoma acradenia tl Greene subsp. eremophila (Greene) G.L.Neso [socoma menziesii on & Arm.) G.L.Nesom var. vernonioides (Nu Me GL piesoin lesa Pe ia filagi no ia a (ook: & Arn.) M.A.Lane var. californica (D C)M A.Lan essingia filaginifolia (Hook. & Arn.) M.A.Lane var. filaginifolia lege ingia germanorum Cham. Lessingia glandulifera A.Gray var. glandulifera Lessingia glandulifera A.Gray var. pectinata (Greene) Jepson ee ingia glandulifera A. var. tomentosa (Greene) Ferris heseinota lem coal A, bays var. lemmonii Markos & Baldwin (2001 Ww Markos & Baldwin (2001) Markos & Baldwin (2001) Markos & Baldwin (2001) Markos & Baldwin (2001) Markos & Baldwin (2001) Markos & Balan Markos & Baldwin Markos & Baldwin ( Markos & Baldwin ( AF251585 AE2Z5157/ AF251578 AF251579 AF251580 AF251581 AF251582 AF 251583 AF251584 AF251572 AF251571 AF251587 AF251593 AF251589 AF251596 AF251599 AF251597 AF251603 AF 251604 AF251606 AF251643 AF251642 AF251630 AF251629 AF251645 AF251651 AF 251647 AF251664 VISAGNI IHL TV Ld LATINOYS £68 Taale 1. continued Species Source (collectors, number, GenBank accession number Herbarium or citation) ITS ETS Lessingia lemmonii A.Gray var. peirsonii (J.T. Howell) Ferris Markos & Baldwin (2001) AF251608 AF251666 Lessingia lemmonii A.Gray var. ramulosissima (Nelson) Ferris Markos & Baldwin (2001) AF251610 AF251668 Lessingia leptoclada A.Gray Markos & Baldwin (2001) AF251612 AF251670 Lessingia micradenia Greene var. glabrata . D.Keck) Ferris Markos & Baldwin (2001) AF251614 AF251672 Lessingia micradenia Greene var. micradeni Markos & Baldwin (2001) AF251615 AF251673 Lessingia nana A.Gra Markos & Baldwin (2001) AF251616 AF251674 ined nemaclada ile Markos & Baldwin (2001) AF251618 AF251676 Lessingia ra Markos & Baldwin (2001) AF251620 AF251678 a tenuis ay a oale Markos & Baldwin (2001) AF251622 AF251680 Lessingia virgata A.Gray Markos & Baldwin (2001) AF251624 AF251682 Arida parviflora (A. Gray) D.R.Morgan & R.L.Hartman Markos & Baldwin (2001) AF251568 AF251626 (Machaeranthera parviflora A.Gray) Machaeranthera tanacetifolia (Kunth) Nees Markos & Baldwin (2001) AF251567 AF251625 Pyrrocoma lan Markos & Baldwin (2001) AF251574 AF251632 Xanthisma spi ie osum - sh) D.R.Morgan & R.L.Hartman Markos & Baldwin (2001) AF251569 AF251627 (Machaeranthera pinnatifida (Hook.) Shinners) Xanthisma texanum DC subsp drummondii (Torr. & A. Gray) Semple Markos & Baldwin (2001) AF251575 AF251633 3mm <2 2-3 mm Involucre height >6mm <5mm 5-6 mm Number of disk florets > 25 < 22 22-25 Disk floret length >46mm <4.3mm 43-46 mm SbF <14 14-17 Cypsela length > 2.8mm < 2.4mm 2.4-2.8 mm Pappus length >45mm < 40mm 4.0-4.5 mm Taste 2. Representative specimens of Symphyotrichum subulatum vars. tenuifolium and aphyllum, arranged from north to south along the Gulf coast of peninsular Florida. *Morphological character- istics of var. tenuifolius (T), var.aphyllus (A), or intermediate (-); characters are listed in the order used el. Florida County Voucher Character states* Identification Gulf Sundberg 2241 (TEX) T-TTTT-T var, tenuifolius Franklin Sundberg 2253 (TEX) T-TTTTTT var. tenuifolius | odf I FSU) TAAAAAAA intermediate Taylor Sundberg 2271 (TEX) AT 1-T— intermediate Sundberg 2291 (TEX) AT-T—T intermediate Levy sunavEIg ee 2293 a TATTTT-T intermediate Levy Cooley & E (FSU) ATA—— intermediate Citrus Godfrey 651 1 (FSU) AATT—AA intermediate Citrus Barilotti s.n. Ae TATAT-AA intermediate Hernando Sundberg 2302 (TEX) TA-AA-AA intermediate Hernando Sundberg 2302 (TEX) TAT—TA intermediate Hernando Cooley 5460 (NY) AA-A—A- intermediate Hernando Cooley 5460 (GH) TAT-TTA- intermediate Pinellas Thorne 9401 (GH) TA-AT— intermediate Hillsborough Sundberg 2308 (TEX) AAAAAAAA var. aphyllus Charlotte plies ae (TEX) AAAAAAAA var.aphyllus Lee 26 (NY) AAAAAAAA var.aphyllus eae wae tenuifolium (L.) G.L. Nesom var. aphyllum (R.W. Long) S.D. undb., comb. nov. Basiony: Aster tenuifolius L. var. aphyllus R.W. Long, Rhodora 72: - 70 TYPE: oar FLORIDA. Hillsborough Co.. NW of Tampa, S of State Route 580 and W Rocky Creek, 24 Dec 1962, Lakela 25610 (HOLOTYPE: GH! ISOTYPES: GA!, RSA! USF). Aster bracei Britton ex Small, Fl. Miami 190, 200. 1913. Symphyotrichum bracei (Britton ex Small) om, Phytologia 77:276. 1994 (1995). Type: BAHAMAS. NEW PROVIDENCE: 31 Aug 1904 Britton & Brace 394 (HOLOTYPE: NY!). 906 BRIT.ORG/SIDA 21(2) Symphyotrichum subulatum Symphyotrichum subulatum is widely distributed in moist habitats in the Americas, from southern and eastern states of the United States, through the Caribbean islands and Central America, to South America. Variety ligulatum is a widespread weedy annual on disturbed soils from Nebraska south to Tamaulipas, Mexico, and from Alabama to New Mexico. Variety parviflorum occurs in North America, West Indies, Mexico, and northern South America and has been introduced in other parts of the world. Variety elongatum grows in Florida, coastal Georgia, and the Bahamas. Variety subulatum is common in salt marshes and brackish areas along the Atlantic coast from New Brunswick to northern Florida. Variety squamatum is an introduced taxon in the United States and elsewhere, with a native habitat of saline and freshwater regions, especially in the southern half of South America. The five varieties recognized here differ in chromosome number, ligule size, capitulescence morphology, number of disk and ray florets, head size, and pres- ence of basal rosettes. Within a single population plants may be tall and with numerous capitula, or short and monocephalous. Within a variety it is typical for individual diagnostic characters to vary. Symphyotrichum subulatum has a base chromosome number of x = 5. Vari- eties ligulatum (Texas, Travis Co., Sund berg 1375), parviflorum, (California, Kern Co, Sundberg 2094) and subulatum (Georgia, Glynn Co., Sundberg 2342) are dip- loids with n = 5, and varieties elongatum (Florida, Dade Co., Sundbe rg 2324) and squamatum (Argentina, Proy. Salta, Lavin & Lavin 5809) are tetraploids with n= 10 (lor 79 more chromosome count citations, see Sundberg 1986). Greenhouse stud- ies of the species (Sundberg 1986) showed that varieties elongatum, parviflorum, squamatum, and subulatum are self-compatible. This may facilitate the fixation of variant forms and result in greater infraspecific variability in these varieties. Variety ligulatum is not self compatible and is the least variable taxon. The varieties intergrade morphologically where their distributions ap- proach one another. This may be the result of past hybridization events and limited gene flow across reproductive barriers. Artificial hybrids produced in the greenhouse among the varieties are highly sterile, yet, for example, 2% of the pollen of the triploid hybrid, var. parviflorum x var. elongatum stains darkly with cytoplasmic stain, suggesting that a small fraction of the pollen may be viable (Sundberg 1986) Varieties ligulatum and parviflorum: Populations intermediate in ligule length and width occur in trans-Pecos Texas, parts of New Mexico (including the type of A. neomexicanus Wooton & Stand. collected in Chaves Co.), Ari- zona, and Chihuahua, Mexico. These are fertile plants in stable populations and produce plump, apparently viable cypselae. Varieties elongatum and subulatum: Intermediates between these varieties occur sporadically in northeastern Florida (Duval County) and along the coast =— SUNDBERG, NEW COMBINATIONS IN SYMPHYOTRICHUM 907 of the Florida panhandle. Intergradation is demonstrated in the compactness of the « apit ulescence and the number of disk and ray florets. Varieties elongatum and parviflorum: Although readily distinguishable over much of their ranges, similar forms of the two varieties are found in south- ern Florida, where their distributions approach. In this area individuals of var. parviflorum are more robust (to 1.5 m tall) than elsewhere and the ligules are often pink, instead of white. Variety parviflorum is usually more diffusely and more equally branched in the capitulescence than var. elongatum. The latter variety often has long branches in the capitulescence, with shorter peduncles that are often disposed toward the upper side of the branch. In addition, vari- ety parviflorum has shorter heads, narrower phyllaries, fewer and shorter ray florets, and fewer disk florets than var. elongatum. Symphyotrichum subulatum (Michx.) G.L. Nesom, Phytologia 77:293. 1994 (1995). Aster subulatus Michx., Fl. Bor-Amer. 2:111. 1803. Type: U.S.A. “Pensylvania:” Michaux s.n.(LEC- TOTYPE [Bosserdet 1970]: PR photograph TEX!) Symphyotrichum subulatum (Michx.) G.L. Nesom var. elongatum (Boss.) S.D. Sundb., nov. BASIONYM: Aster subulatus var. elongatus Boss., Taxon 19:250. 1970. Type: U.S.A. FLorIDA. Hillsborough Co. Tampa, 20 Aug 1895, Nash 2416 (LECTOTYPE [Jones & Lowry 1986} P; photograph: TEX). Aster bahamensis Britton, Bull. aes Bot. emer 14. 1914. Astersubulatus Michx. var. bahamensis (Britton) Boss., Taxon 19:249, 1970. Symphyotrichum bahamense (Britton) G.L. Nes Phytologia 77:276. 1994 (1995). Type: BAHAMAS Great Bahama: Barnett’s Point, 5- ee ets Britton & Millspaugh 2621 (HOLOTYPE: NY!, photograph TEX!, IsOTYPE: FI). a ee subulatum (Michx.) G.L. Nesom var. ligulatum (Shinners) S.D. Sund nov. BASIONYM: Aster subulatus Michx. var. ligulatus Shinners, Field & Lab. 21:159. is Type: U.S.A. TEXAS. Hill Co: 6.9 mi SW of Hillsboro, bottom of dried-up pond, sandy clay, rays light lavender, 6 Oct 1949, Shinners 12057 (HOLOTYPE: SMU; ISOTYPE: GH). +9. a Tripolium divaricatum Nutt., Trans. Amer. Philos. Soc. n.s. 7:296. 1841. Aster divaricatus (Nutt.) Torr. & A. Gray, Fl. N. Amer. 2:163. 1841. non L., Sp. Pl: 873.1753, Symphyotrichum divaricatum (Nutt.) G.L. Nesom, Phytologia 77:279. 1994 (1995). Type: Innundated [sic] banks of the Missis- sippi, Nuttall s.n. (HOLOTYPE: PHI, photograph TEX!) Distinctive, localized forms of variety ligulatum occur in some areas. Collec- tions from the Dallas-Fort Worth area of Texas have particularly small heads and florets. Along the coast of Texas and Tamaulipas, near the mouth of the Rio Grande and southward, plants are especially large, sometimes over two meters tall, and exhibit the largest capitula, with the most phyllaries, ray florets, and disk florets found in the variety. These forms intergrade gradually into more typical forms and are not deemed to be worthy of nomenclatural recognition. Symphyotrichum subulatum (Michx.) G.L. Nesom var. parviflorum (Nees) S.D. Sundb., comb. nov. BASIONYM: dee Sage ads aed ) DC. var. B parviflorum Nees, Gen. sp. Aster: 157, 286. 1833. TYPE: U.S.A. HAWaIL Oahu, 1816 or 1817, Chamisso s.n. (LECTO- Type, here designated: G-DC; microfichel, Be a 908 BRIT.ORG/SIDA 21(2) eer | ap) aa ks eee Poepp. ex Spreng., Syst. Veg. 3:518. 1826. $ eX Spreng.) G.L. Nesom, Phytologia 77:281. 1994 Seen TYPE: can pl. Cub. MSS, In siccis ca ee ee at (HOLOTYPE: W?; ISOTYPES: HAL!, MOI, NY, FD). Aster divaricatus (Nutt.) Torr. & A. Gray var. sandwicensis A Gray in H. Mann, Proc. Amer. Acad. Arts 7:173, 18607. Aster sandwi is (A. Gray in H. Mann) Hieron., Bot. Jahrb. Syst. 29:20. 1901. Aster subulatus eon var. sandwicensis (A. Gray ex H. Mann) A.G. Jones, Brittonia 36:465. 1984. TYPE: U.S.A. HAWAIL Oahu, 1816 or 1817, Chamisso s.n. (LECTOTYPE: Jones 1984}: G-DC; microfichel, ah Jones (1984) designated a specimen in G-DC as the lectotype of Aster divaricatus var. sandwicensis and listed homotypic and heterotypic synonyms of the vari- ety. She listed Tripolium subulatum (Michx.) DC. var. B Nees as “unnamed” in the list of homotypic taxa, but did not recognize, or explicitly lectotypify, the variety. Sundberg later examined high resolution photographs of the lectotype and identified the specimen as var. parviflorum, and not var. squamatum, as the name has been applied in earlier publications. Nees (1833) cited Tripolium subulatum(Michx.) DC. var. Bas “B. Parviflorus, caule supradecomposito, calathiis dimidio minoribus,” which could be inter- preted as a polynomial, or an informal description of a form. However, on page 286, in “synonyma addenda vel corrigenda” Nees writes, “Ad Tripolium subulatum var. B parviflorum p. 157, Aster inconspicuus Less. in Schlechtend. Lin. V. p. 143,” which changed the “parviflorus” to “parviflorum” to agree in gender with Tripolium. He indicated that “B parviflorum” was based on A. inconspicuus Less., and associated “Var.” with the varietal epithet. Thus, Nees’ correction was to list A. inconspicuus Less as a synonym of his new variety. The lectotype of var. parviflorum is further selected on the basis of Nees’ (1833) statements on p. 143 that “Var. B in O Wahu insula (Cham.).” and “Vidi exempla Americae borealis et O Wahu insulae,” indicating that he had seen a specimen of the variety col- lected in Oahu by Chamisso. Variety parviflorum varies in ligule length, ligule pigmentation (white or pink), head size, and vegetative characters. Populations with abnormally large heads and pigmented ligules occur sporadically in the states of Mexico and Veracruz, of central Mexico. laa hale es subulatum (Michx.) G.L. Nesom var.squamatum (Spreng.) S.D. Sundb., comb. nov. Basiony: Conyza squamata Spreng., Syst. Veg. 3:515. 1826; Aster (S g.) Hieron., Bot. Jahrb. Syst. 29:19. 1901. Conyzanthus squamatus (Spreng.) Tamamechian FL, ULR.S.S. 25:186. 1959. Symphyotrichum squamatum (Sprengel) G.L. Nesom, Phytologia 77:292. 1994 (1995). Type: URUGUAY: Montevideo, Sello[w] s.n. (HOLOTYPE: P). Sprengel’s Asteraceae specimens were sold to Schultz-Bipontinus, whose her- barium is now part of the Cosson herbarium at P (Stafleu @ Cowan 1985). Be- cause this is the only known collection by Friedrich Sellow from Montevideo in the Sprengel Herbarium, the P specimen (Sprengel Herb #1064) may be re- garded as a holotype. Stafleu and Cowan note that Sello crossed out the letter SUNDBERG, NEW COMBINATIONS IN SYMPHYOTRICHUM 909 “w” on many labels to reflect the original family name. Sprengel used the spell- ing, “Sello” in the original description. The branching of the capitulescence, head size, and cypsela pubescence vary greatly in var. squamatum. Populations with glabrous cypselae, and narrow phyllaries and heads occur in the vicinity of Tucuman Province in Argentina, as well as around Buenos Aires. In other respects these plants are typical of the variety. In Chile a form with large involucres (8.5 mm high), ligules that extend 1.5 mm beyond the pappus, and cypselae ca. 3.2 mm long exist. These occur in proximity to populations of typical var. squamatum, which has shorter involu- cres, cypselae, and ligules. Similar large-headed forms, but with short ligules are found in Peru. Collections of these variants of var. squamatum are few and additional study may reveal that some of the populations represent distinct varieties. Symphyotrichum subulatum (Michx.) G.L. Nesom var. subulatum Aster subulatus Michx. var. obtusifolius Fernald, Rhodora 16:61.1914. TYPE: CANADA. NEW BRUN- Gloucester Co.: Bathurst, brackish marsh along Middle River, 13 Aug 1913, Blake 5372 ne OTYPE ae ISOTYPES: CASI, LL}, ae | USD. ee chx. var. euroauste ld & Griscom, Rhodora 37:183 1935. TYPE: U.S.A. VIR- GINIA. Norfolk Co. border of gum p near North Landing, 22 Sep 1933, Fernald & Griscom 2919 (HOLOTYPE: GH)). Aster ensifer Boss., Taxon 19:250. 1970. nee U.S.A. MASSACHUSETTS: Cambridge, margin of salt marsh, 2 Oct 1901, Robinson & F: 65 (HOLOTYPE: P, photograph TEX! IsoTYPEs: CASI, DSI, GA, GHI, ILL, LL!) MICH!, MO! NY! POM! UC, USD. Plants of variety subulatum from the northern Atlantic coast are shorter, with fewer capitula, larger leaves in the capitulescence, and longer ligules than plants from the southern Atlantic coast of the United States. The form that occurs in New Brunswick has been treated as Aster subulatus Michx. var. obtusifolius Fernald. However, this represents an extreme form in a gradual cline, and one variable variety is recognized here. ACKNOWLEDGMENTS I thank Linda Hardison for assisting with the production of the manuscript, Laurent Gautier and Muriel Hecquet for sending high-resolution photographs of a type specimen at G-DC, Almut Jones for providing photographs of type specimens, Guy Nesom for his comments on the manuscript, and the curators of CAS, DS, EF FSU, GA, GH, HAL, K, LL, MICH, MO, NLU, NY, POM, RSA, TEX, and USF for specimen loans used for this study. REFERENCES BosseroeT, P. 1970. Deux acceptions d’Aster subulatus. Taxon 19:244—-250. Jones, A.G. 1980. A classification of the new world species of Aster (Asteraceae). Brittonia 32:230-239. 910 BRIT.ORG/SIDA 21(2) Jones, A.G. 1984. Nomenclatural notes on Aster (Asteraceae)—lll. The status of A. sandwicensis. Brittonia 36:463—466. Jones, A.G.and PP. Lowry. 1986. Types and selected historic specimens of Aster s.l. (Asteraceae) in the Herbarium, Laboratoire de Phanérogamie, Muséum national d'Histoire naturelle, Paris (P). Adansonia 4:393-41 2. Nees, C.G. 1833. Genera et species Asterearum, 2nd printing. Nesom,G.L. 1994 (1995).Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the New World species. Phytologia 77:141-297 Sempte, J.C.and L. Brouittet. 1980.A synopsis of North American asters: the subgenera, sec- tions, and subsections of Aster and Lasallea. Amer. J. Bot.67:1010-1026. Starteu, FA. and R.S. Cowan. 1985. Taxonomic literature: vol. v: Sal-Ste. Boston: W. Junk Pub- lishers, The Hague. SunbBERG, S.D. 1986. The systematics of Aster subgenus Oxytripolium (Compositae) and historically allied species. Ph.D. dissertation, University of Texas at Austin, Austin. SEEDAND CAPSULE GHARACTERSINARC YOR VELLUM, BOUVARDIA, AND MANETTIA (RUBIACEAE), WITEENO LES ON A. SERPYLLEACEUM Edward E. Terrell! Harold Robinson Research Collaborator é National Museum of Natural History National Museum of Natural History Smithsonian Institution Smithsonian Institution Washington, DC 20013-7012, U.S.A. Washington, DC 20013-7012, U.S.A. ABSTRACT Seed and a charectets of scied species of Arcytophyllum, Bouvardia, and Manettia (Hedyotideae; Rub 1 and compared. Seed characters are illustrated by scanning electron microscopy. Meehan similarities and differences phasized and outlined by a descriptive key. All characters of Arcytophyllum serpyllaceum and A. nani um were compared in tabular f — ormat, and a strong similarity was noted. These results support previous work that consid- ered A. serpyllaceum a member of the genus Arcytophyllum. RESUMEN Las semillas y frutos de algunas eee de ie en Nee! saataes a Manettia (Hedyotideae; Rubiaceae) son descritas y comparadas. L — resentados mediante ian Pogues: per un Tosco Rle electronico de barrido. Las semejanzas y diferencias ig | | medio de una clave descriptiva. Todos los teres de Arcyto] i ll Pp) Ila ye A.muticum fueron comers en una nae qd Ret lt A . slitiid I apoyan eenidios previos, de ante ae serpyl Arcytophyllum. aceum es en realidad un miembro del genero INTRODUCTION This study of seeds and capsules of three genera belonging to the tribe Hedyotideae (Rubiaceae), follows the pattern of recent work on this tribe (e.g., Terrell 1996; Terrell & Robinson 2003). These studies involved examination of surface features of seeds by dissecting microscope and scanning electron mi- croscopy (SEM). Revision of Houstonia (Terrell 1996) showed seed characters and chromosome numbers to be especially important in revealing relationships. The three genera in the present study, Arcytophyllum, Bouvardia, and Manettia, are from Mexico, and Central and South America. These genera have largely escaped the past tendency to include many members of the tribe in a broad concept of the genus Hedyotis, and Bouvardia and Manettia have some- times been placed in the tribe Cinchoneae because of the conspicuous papery winged seeds ‘Address for correspondence: 14001 Wildwood Drive, Silver Spring, Maryland 20905, U.S.A SIDA 21(2): 911-927. 2004 912 BRIT.ORG/SIDA 21(2 Bremer and Manen (2000) in a molecular study placed Manettia and Bouvardia in the combined Hedyotideae/Spermacoceae where the older name Spermacoceae was adopted. Terrell and Wunderlin (2003) questioned this ac- tion on the basis of fundamental morphological differences between the two tribes. Here and elsewhere (e.g., Terrell @ Robinson 2003) we continue to place the present genera in the tribe Hedyotideae sens. str All three of our studied genera have recorded chromosome numbers of x = 9 (Lewis 1965: 199), a number that is frequent in the Hedyotideae, but unusual in the family Rubiaceae that has mostly x = 11. The purpose of this study is to present for comparison illustrations and/or data on the seeds and capsules of the three genera and to re-examine seed data published by Terrell (1999) and Andersson et al. (2002) for Bouvardia and Arcytophyllum serpyllaceum. MATERIALS AND METHODS Seeds were obtained from the U.S. National Herbarium (US), Smithsonian In- stitution, and from other herbaria including CHAPA, GH, NY, TENN. Seeds were examined by dissecting microscope and mature, well-formed seeds were viewed by scanning electron microscopy (SEM) at the Smithsonian Institution, and prior to 1985 at the Electronics Lab, US. Department of Agriculture, Beltsville, Maryland. Seed descriptions for each species and inclusive descriptions for each spe- cies-group provide basic data. The nomenclatural authors and collection data are added to the seed descriptions for each species. Seed collections viewed by SEM are indicated by a designation such as B35, ase.g. for Arcytophyllum lavarum, Light microscopy was used to observe pollen and some additional struc- tures such as corollas, calyx lobes, and stipules in type species and other se- lected species of all three genera. Material was mounted on slides in Hoyer’s solution (Anderson 1954). RESULTS The light microscope study included material of Arcytophyllum nitidum, A. serpyllaceum, Bouvardia ternifolia, and Manettia alba and M. reclinata. The first, third, and fifth of these species are the types of their respective genera. Pollen in Arcytophyllum and Bouvardia show essentially identical spherical, tricolporate pollen with minor variation in the granulation of the exine. Only Manettia differs by a sometimes more oblate shape with projecting pores. The grains of M. reclindta are commonly lying on their more flattened poles and seem almost triangular. The light microscope also showed prominent raphide bundles in stipules, calyx and sometimes the corolla of Manettia that were not seen in Arcytophyllum or Bouvardia slides. A review of the genera and species for seed and capsule data is as follows: TERRELL AND ROBINSON, RUBIACEAE GENERA 913 Arcytophyllum Willd. ex Schult. in Roem.& Schult, Syst. Veg. Mant. 3:5. 1827. This Andean and Central American genus of 16 species of shrubs, subshrubs, and suffruticose herbs grows at higher elevations from Costa Rica to Bolivia (Mena 1990). Most species have rather restricted ranges. Mena found the most useful characters to be the shape of the stipules and the leaf and corolla shapes We examined seeds of nine of the sixteen species of Arcytophyllum, of which six species were treated by SEM. The seeds of the studied species fell into three groups, as outlined below. Group A This group includes the type of the genus. The species are A. filiforme, A. lavarum, A. macbridei, A. muticum, A. nitidum, and A. serpyllaceum. Arcytophyllum lavarum, A. muticum, and A. serpyllaceum have lenticular seeds, polygonal or suborbicular in outline, with centric punctiform or slightly raised hila (Fig. 1). Arcytophyllum lavarum may or may not be winged, but is shown in this figure as being winged. The remaining three species, A. filiforme, A. macbridei, and A. nitidum were not done by SEM, but their descriptions are included below. Descriptions of the seeds of the six species follow: Arcytophyllum filiforme (Ruiz & Pav.) Standl.—Seeds 0.7-1L0 mm diam., black, moderately to strongly compressed, suborbicular in outline, lenticular, thickly to thinly concavo-convex, hilar area rounded or slightly raised, areoles not seen in detail. Harling & Andersson 12638(US), Ecuador (det. Boom); Fosberg &» Giles 23135 (US), Ecuador (det. L.B.Smith). Arcytophyllum lavarum K. Schum.—Seeds 0.7-1.0 mm diam., black, strongly compressed, suborbicular or polygonal in outline, lenticular, biconvex, wing absent or partial, very narrow, fragile, hilum punctiform, areoles isodiametric or polygonal, their walls thick, testa smooth. Cuatrecasas & Leon 26524 (US), Costa Rica, R8a; Herrera & Robles 791 (US), Costa Rica, B35 (Fig. 2). Arcytophyllum macbridei Standl.—Seeds 1.4-L6 mm diam., black, strongly compressed, suborbicular in outline, lenticular, concavo-convex, wing partial, very narrow, fragile, hilum punctiform or slightly raised, areoles not seen in detail. Wurdack 1273 (US), Peru (dupl.det. Steyermark). Arcytophyllum muticum (Wedd.) Standl.—Seeds 1.0-1.4 mm diam., black, moderately to strongly compressed, suborbicular or polygonal in outline, len- ticular, rather thickly biconvex or ony convex dorsally, hilum punctiform, slightly raised, or a short ridge | tric or polygonal, small, their walls thick, testa smooth. Fosberg 19196 (US), Colombia, (det. Mena), B36; Haught 5803 (US), Colombia, R7; Martin & Plowman 93 (US), Colombia (det. Mena) (Fig. 1). Arcytophyllum nitidum (Kunth) Schltdl.—Seeds 0.7-1.0 mm diam., black, strongly compressed, polygonal to suborbicular in outline, lenticular, convex dorsally, flat or bent ventrally, hilum punctiform, areoles not seen in detail. Fosberg & St. John 21880 (US), Colombia. 914 BRIT.ORG/SIDA 21(2) ee LP ane on a8; ty , > 3 G3 anions: wing? P 2; *y .. Fic. 1. Seeds of A phyll peci i 1 by SEM. A,B, E. A ! Avil i Fosberg 19196 (US), Colombia. C, D, F, Arcytophyll pyll Williams et al. 22771 (US), Guatemala. A, C, ventral views; B, D, dorsal views; E, F, areoles. Arcytophyllum serpyllaceum (Schltdl) Terrell—Seeds 0.6-1.2 mm diam., black, moderately to strongly compressed, suborbicular or polygonal in out- line, lenticular, rather thickly biconvex or flat on one face, hilum punctiform, areoles isodiametric or polygonal, small, their walls thick, testa smooth. Sharp TERRELL AND ROBINSON, RUBIACEAE GENERA: SEED 915 Fic.2. Seeds of Arcytophyllum | ined by SEM.A_B, D, Cuat & | 26524 (US), Costa Rica; C, Herrera & ) 7b, U, Robles 791 (US), Costa Rica. A, C, ventral views; B, dorsal view; D, areoles. 45143 (TENN), Guatemala, R12; Williams et al. 22771(US), Guatemala, B38; Al- exander 1063 (NY), Chiapas, Mexico (Fig. D. An inclusive description follows: Seeds 0.6-1.6 mm diameter, black, moder- ately to strongly compressed, suborbicular or polygonal in outline, thickly to thinly lenticular, faces (sides) flat, concave, convex, or biconvex, wings absent or present, partial, very narrow, fragile, hilum punctiform, rounded, or slightly raised, areoles isodiametric or polygonal, small, walls thick, testa smooth. Group B This group includes only A. aristatum. Seeds are polygonal in outline, lumpy, not lenticular, and have a strongly raised or ridged ventral face. Arcytophyllum aristatum Standl.—Seeds 0.7-1.0 mm diam., black, moder- ately to slightly compressed, lumpy, polygonal in outline, dorsal face convex, ventral face with a strongly raised hilar area or a short to long hilar ridge, ar- eoles small, isodiametric or polygonal, walls thick, testa smooth. Cuatrecasas 916 BRIT.ORG/SIDA 21(2) 18976 (US), Colombia, B34; Luteyn & Luteyn 6758 (US), Ecuador, Steere 8034 (US), Ecuador (Fig. 3). Group C This group includes Arcytophyllum rivetii and A. thymifolium and has ellipti- cal or oblong seeds with a low hilar ridge. Arcytophyllum rivetii Danguy & Cherm.—Seeds 1.0-1.3 x 0.6-0.7 mm, black, strongly compressed, broadly elliptic or oblong in outline, biconvex, concavo- convex, or flat, wing none or very narrow at one end of seed, hilum a low linear ridge, areoles isodiametric or polygonal, their walls thick, testa smooth. Wurdack 1288 (US), Peru, B37 (Fig. 3). Arcytophyllum thymifolium (Ruiz & Pav.) Standl—Seeds 0.8-1.2 « 0.5-0.7 plong in mm, black or dark brown, strongly compressed, broadly elliptic or o outline, rather thin, concavo-convex or flat, hilum a low linear or oblong ridge, areoles isodiametric or polygonal, their walls thick, testa smooth. Firmin s.n., 5/12/27 (US), Ecuador, B39; Fosberg 21196 (US), Colombia; Schultes & Villarreal 7880 (US), Colombia (Fig. 3). An inclusive description is as follows: Seeds 0.8-1.3 x 0.5-0.7 mm, black or dark brown, strongly compressed, broadly elliptic or oblong in outline, bicon- vex, concavo-convex, or flat, wing none or very narrow at one end of seed, hi- lum a low, linear or oblong ridge, areoles isodiametric or polygonal, their walls thick, testa smooth. Summary of Arcytophyllum.—A general survey of Arcytophyllum seeds shows that they are strongly or moderately dorsiventrally compressed, with a ventral punctiform centric hilum or a hilar ridge. A few species may have very narrow, fragile, partial wings, but the more common state is wingless. The areoles (cells) are usually isodiametric or polygonal and rather thick-walled with smooth testa. There are three distinct kinds of seeds. Group A seeds are polygonal or suborbicular in outline. The seeds are len- ticular, varying thicker or thinner and biconvex or concavo-convex or flat ona face. The hilum is centric, i.e., on or near the center of the ventral face of the seed, and is often punctiform, appearing asa dot ora small round or raised area. (Figs. 1, 2). Group B has thicker, lumpy seeds with ventral face raised or a short to long hilar ridge. (Fig. 3). Group C has oblongoid or ellipsoid seeds which are slightly to strongly concave — A low narrow hilar ridge extends along part or most of the seed (Fig. Mena’s (1990) ne of Arcytophyll ds as irregularly patelliform (dish- or saucer-shaped) or cymbiform (boat-shaped) and coarsely alveolated (honeycombed) is somewhat misleading because it suggests that they have shal- low or deep ventral cavities or concavities such as are typical of Houstonia (Terrell 1996). In Arcytophyllum, however, the seeds are in Group A lenticular, in Group B lumpy, and in Group C are ellipsoidal or oblongoid and longitudi- — TERRELL AND ROBINSON, RUBIACEA E GENERA 917 100 pm Cc Fic. 3. Seeds of Arcytophyllum species examined by SEM. se aronenhylaty rivet, Wurdack 1288 (US), Peru. C, Arcytophyllum thymifolium, Firmin s.n., 5/12/27, (US), Ecuador fi , Cuatrecasas 18976 (US), Co- lombia. A, C, D, ventral views; B, areoles. nally concave or bent. The seed surfaces are never honey-combed, but are re- ticulate and like most seeds of Hedyotideae are made up of walled areoles or cells. Capsules of Arcytophyllum were described by Standley (1921) as bilocu- late, turbinate to globose, and usually septicidally dehiscent to the base. Mena’s description was “septicidal, sometimes only the beginning of the dehiscence loculicidal”, crowned by the persistent calyx lobes and intercalycine teeth. For eight of our studied species Mena said that the capsules varied 1-3 mm in di- ameter, and were mostly subglobose. He found 2-15 seeds per locule (4-30 per capsule). For A. thymifolium Mena found 17-2 x 2-2.5 mm, which suggests a dimension wider than long. Terrell for A. thymifolium recorded 2 x 1 (ellipsoid or oblongoid) in one collection; in a second collection 15-2 x 1.0-L7 (subglobose to oblongoid). Terrell found 5-12 seeds per capsule in A. lavarum and A. thymifolium. The capsules of A. serpyllaceum are 15-4 x 2-3.5 mm, turbinate, loculicidal, nerved, and have 8-29 seeds per capsule. This species is further dis- cussed below and compared with A. muticum. 918 BRIT.ORG/SIDA 21(2) Bouvardia Salisbury, Parad. Lond. Pl. 88. 1805. Blackwell (1968) recognized 31 species in three subgenera in Bouvardia. The species are primarily Mexican, with extensions into Central America and south- western United States, and are shrubs except [or four species in the third subge- nus. The first subgenus recognized by Blackwell was Bouvardiastrum Schltd with 15 species, of which we examined B. capitata, B. cordifolia, B. laevis, and B. multiflora. The second subgenus, Bouvardioides Schltdl. has 8 species, none of which are represented in our study. The third subgenus, Bouvardia Schltdl. has 8 species of which we examined B. ternifolia and two recently described or transferred perennial herbs, B. rzedowskii and B. xestosperm (Terrell and Koch 1994), which are similar to and apparently related to the perennial herbs, B. rosea Schltdl. and B. tenuifolia Stand|. Several additional Bou vardia species have been described in recent years, but our study utilizes mainly the species stud- ied by Blackwell (1968). Previous field work in Mexico provided a foundation for the present study. Bouvardia rzedowskii was collected and described and B. xestosperma was re- collected in Oaxaca (Terrell and Koch 1994). Other species of Bouvardia were seen in Mexico during this work. Individual descriptions of seeds are given as follows, with capsule data added from Blackwell (1968). Two groups of species are recognized for this genus. Group A The first group includes four shrubby species in subgenus Bouvardiastrum (B. capitata, B. cordifolia, B. laevis, B. multiflora) and one shrubby species in subg. Bouvardia (B. ternifolia). These species seem to be generally similar in seed morphology. Bouvardia capitata Bull.—Seeds 2.5-3.5 mm diam., brown with tan wing, papery thin, orbicular in outline, ventral face concave, seed body 1.0-1.5 mm wide, wing 0.3-1.0 mm wide, its margin entire, undulate, hilum punctiform, often a raised point, areoles radially elongated, their walls thin, testa smooth. Hinton 8600 (US), Tejupilco, Tepascaltepec (?), Mex. (det. Blackwell), B41 (Fig. 4). Bouvardia multiflora (Cav.) Schult. & Schult.f.—Seeds 2-3 mm diam., dark brown or black with brown wing, papery thin, orbicular in outline, ventral face concave or almost flat, seed body 0.8-L4 mm wide, wing 0.2-1.0 mm wide, its margin entire, undulate, hilum punctiform, sometimes a raised point, areoles radially elongated, their walls thin, testa smooth. H.S. Gentry 8388 (US), Durango, Mexico, (det. Blackwell), B42; E. Palmer 708 (US), Mexico (Fig. 4). Bouvardia ternifolia (Cav.) Schltdl —Seeds 2-3 mm diam., black with brown wing, papery thin, orbicular in outline, ventral face concave, seed body 1.0-2.2 mm wide, wing 0.2-0.6 mm wide, its margin entire, undulate, hilum punctiform, a slightly raised point or rounded area, areoles radially elongated, their walls thin, testa smooth. Dziekanowshi et al. 1860 (US), Hidalgo, Mexico, H52 (Fig. 4). An inclusive description is as follows: Seeds 2-3.5 mm diam., brown, black, ~u TERRELL AND ROBINSON, RUBIACEAE GENERA 919 Fic. 4. Seeds of Bouvardia species examined by SEM. A-B, Bouvardia capitata, Hinton 8600 (US), Mexico. C, D, Bouvardia ternifolia, Dziekanovski et al. 1860 (US), Mexico. E, F, Bouvardia multiflora, H.S. Gentry 8388 (US views; F, enlarged section showing areoles. — , Mexico. A—F, ventral or tan, papery thin, orbicular in outline, ventral face concave, seed body 0.8-2.2 mm wide, wing 0.2-10 mm wide, its margin entire, undulate, hilum puncti- form,a raised point or rounded area, areoles radially elongated, their walls thin, testa smooth. Capsules in group A have the following inclusive description: 3-9 x 3.5-1] 920 BRIT.ORG/SIDA 21(2) mm., subglobose or slightly oblate, dehiscing loculicidally then septicidally (Blackwell). Standley (1921) recorded capsules as globose and loculicidally dehiscent. Group B. The second group includes two recently described or transferred perennial herbs in subg. Bouvardia (B. rzedowskii and B. xestosperma). Bouvardia rzedowskii Terrell & S.D. Koch—Seeds 1-2 mm diam., dark brown with brown wing, papery thin, orbicular in outline, ventral face concave or flat, seed body 0.5-L.0 mm wide, wing 0.1-0.4 mm wide, its margin entire, undulate, hilum punctiform, sometimes a raised point, areoles radially elongated, their walls thin, testa smooth. Koch & Koch 8935 (CHAPA), Oaxaca, Mexico, H5] (Fig. 5). Bouvardia xestosperma (Rob. & Greenm.) Terrell & S.D. Koch—Seeds |-2 mm diam., dark brown or black, thin, orbicular or oval in outline, ventral face flat or slightly concave, seed body 0.3-L0 mm wide, wing partial or complete, 0.1-0.5 mm wide, entire, sometimes undulate, hilum punctiform, a slightly raised point, areoles radially elongated, their walls thin, testa smooth. Koch & Koch 8934 (CHAPA), Oaxaca, Mexico, B45; Conzatti & Gonzalez 248 (GH), Oaxaca, Mexico, H4 (Fig. 5). Inclusive description: Seeds 1-2 mm diam., dark brown or black, papery thin, orbicular or oval in outline, ventral face concave or flat, seed body 0.3-L0 mm wide, wing partial or complete, 0.1-0.5 mm wide, entire, undulate, hilum punctiform, sometimes a raised point, areoles radially elongated, their walls thin, testa smooth. Capsules 3-5 x 3.5-4.5 mm in B. rzedowskii, and 2.5-4 mm long in B. xestosperma, with loculicidal dehiscence.(Terrell & Koch 1994). Due to limited sampling summaries are not given for Bouvardia and Manettia; however, a comparison of Bouvardia groups A and B shows that group B of perennial herbs has smaller seeds and partial instead of complete wings. Bouvardia ternifolia, a shrubby species, seems more allied to Bouvardiastrum or Bouvardioides which have shrubby species. ced Manettia Mutis ex L., Mant. Pl. 553, 558.1771, nom. cons. The genus includes vines or herbs distributed in tropical America, principally in South America. Standley (1921) treated 10 species from West Indies, Mexico, and Central America. Chung (1967, 1968) studied five species in the section Heterochlora K. Schum. and six species in section Pyrrhanthos K. Schum. Steyermark (1974) estimated 140 species of Manettia altogether, and treated 19 species from Venezuela. Manettia has broadly winged seeds similar to those of Bouvardia. Our data on Manettia are limited, and our main purpose here is to present seed descriptions of five species and SEM illustrations of three species of Manettia for comparison with similarly-seeded Bouvardia. We do not reach any conclusions regarding possible relationships to Bouvardia. TERRELL AND ROBINSON, RUBIACEAE GENERA Pe t 7 rset all. £D ih * - JL CORA D 4: Ay Lit es a L ONOICICUADA\ + Fic.5 A-B, ( ), Mexico.¢, D, Dp 4; 248 (GH), Mexico. A, ventral view; B, C, D, areoles We examined the seeds of the following five species of Manettia: M. calycosa var. karsteniana; M. divaricata; M.flexilis, M. meridensis,and M. reclinata. Three of these were examined by SEM: M. calycosa var. karsteniana, M. flexilis,and M. reclinata. Manettia calycosa Griseb.var. karsteniana K. Schum.—Seeds 0.8-11 mm diam., black with brown wing, thin, orbicular in outline, flat or slightly con- cave ventrally, seed body ca. 0.7 mm wide, wing 0.1-0.3 mm wide, its margin erose, hilum punctiform, areoles radially elongated, walls thin, testa smooth, Steyermark & Wiehler 106576 (US), Venezuela, B47 (Fig. 6). Manettia divaricata Wernham—Seeds 2-3 mm diam., black with brown wing, thin, orbicular in outline, flat or slightly concave ventrally, seed body 1.0-1.5 mm wide, wing 0.4-10 mm wide, its margin ornately erose or toothed, hilum depressed to slightly raised, areoles not seen in detail. Schunke V.838 (US), Peru, (det. D.R. Simpson). Manettia flexilis Brand.—Seeds 2.0-3.5 mm diam., dark brown with light brown wing, thin, orbicular in outline, flat, seed body 0.6-1.5 mm wide, wing 0.4-1.2 mm wide, its margin erose, hilum punctiform or flat, areoles radially 922 BRIT.ORG/SIDA 21(2) Fic. 6. Seeds of M I i ined by SEM. A, Manetti ar. karsteniana, Steyermark & Wichler 106576 (US), Venezuela. B, Manetslaredtiniata: Hooverl6l (US), meee: C- F Moneta lexis, Smith P2360 (US), Costa Rica. A—B, ventral views; C, dorsal view; D, hilar elongated, walls thin, testa smooth. A. Smith P2360 (US), Costa Rica, H32, (det. S. F Blake) (Fig. 6). Manettia meridensis K. Schum.—Seeds 2-3 mm diam., black with dark brown wing, thin, orbicular in outline, flat or slightly concave ventrally, seed TERRELL AND ROBINSON, RUBIACEAE GENERA 923 body 0.5-L5 mm wide, wing 0.4-0.8 mm wide, its margin entire or minutely erose, hilum punctiform, flat or a slightly raised point, areoles not seen in de- tail. Bro. Daniel 1707 (US), Colombia. Manettia reclinata Mutis ex L.—Seeds 2.5-3.0 mm diam., black with brown wing, thin, orbicular in outline, flat or slightly concave ventrally, seed body 0.7-1.2 mm wide, wing 0.5-1.2 mm, its margin erose, hilum area flat or slightly depressed, areoles radially elongated, walls thin, testa smooth. Hoover 161 (US), Se Mexico, B46 (Fig. 6). Inclusive description of the five species is as follows: Seeds 0.8-3.5 mm diam., black or brown, thin, orbicular in outline, flat or slightly concave ventrally, seed body 0.5-1.5 mm wide, wing 0.1-1.2 mm wide, its margin erose, coarsely toothed, or entire, hilum punctiform, flat or depressed, or slightly raised, areoles radi- ally elongated, walls thin, testa smooth. Seeds of Manettia have wings with margins erose or coarsely and irregu- larly toothed, much more ornate than Bouvardia wings. Manettia capsules were described by Standley (1921) as septicidally dehis- cent, obovoid or turbinate, and coriaceus or chartaceous. Steyermark (1974) also considered the capsules as septicidally dehiscent and described M. calycosa as subglobose to ellipsoid or turbinate and 4-7 mm long; for M. reclinata obconic and 10-12 x 7-9 mm. In this study we found M. divaricata with capsules 10 x 5.5 and obovoid; M. meridensis 5 x 5and obovoid; and M. reclinata 8 x 6.5-8 mm wide and subglobose or broadly ellipsoid. All capsules seen had septicidal dehiscence. Capsules often had two separated halves with each half having a slit or longitudinal open- ing into the locule reminescent of the diplophragmous capsules of Hedyotis subgenus Hedyotis (Terrell @ Robinson 2003). DISCUSSION Capsule characters Capsules in Arcytophyllum and Manettia have septicidal dehiscence, whereas Bouvardia primarily has loculicidal dehiscence followed by septicidal dehiscence. Capsule thicknesses and sizes are much greater in Bouvardia and Manettia and more varied in shape, although a predominant shape in Arcytophyllum and Bouvardia is subglobose. Seed characters The following descriptive key is based on the the present sample of species. Bouvardia and Manettia seeds are so similar that they are here treated as one unit. . Seeds 0.6-1.6 mm long or diam, cereale or elliptical in outline, moderately or ey compressed, lenticular, f concavo-convex, or flat or else some- ey concavo-convex, wings none or very narrow and partial, hila centric, flat or ssed or slightly raised, or else a low ridge, areoles isodiametric or some ne Bolvaenel their walls thick AFeoramnyittin 1. Seeds 0.8-3.5 mm diam, orbicular in outline, strongly compressed, flat, thin, most 924 BRIT.ORG/SIDA 21(2) den broadly winged, wing papery, fragile, seed body and wing often incurved concave, wing margin entire and undulate (Bouvardia) or erose or toothed a ettia),or else wing narrow, hila centric, flat or sometimes raised slightly to form a point, areoles radially elongated, their walls thin Bouvardia, Manettia Relationships of Arcytophyllum serpyllaceum The species long known as Hedyotis serpyllacea Schltdl. (Linnaea 9:599. 1834) occursat higher elevations (usually 1500-3500 m) in southern Mexico (Veracruz, Chiapas, Oaxaca) and Guatemala. It is a prostrate, creeping, mat-forming, sulfruticose herb that is often abundant in open, grassy places or fields. It pro- duces small white flowers that tend to be purplish or greenish on their abaxial faces. During field work in Mexico (Terrell & Koch 1994) the species was ob- served and collected in Veracruz. This species was long retained in Hedyotis but differed from other Mexican species that had been placed in that genus. Terrell (1999) described and discussed it in detail, transferred the name Hedyotis serpyllacea to Arcytophyllum (1827) and noted that there is a precedent for its placement in Arcytophyllum. John Donnell Smith (1893) described the species from Guatemala, placing it in Mallostoma (Karst. 1862), a synonym of Arcytophyllum, as M. shannoni. Smith provided a rather complete description of the entire plants, including the follow- ing: “capsule obovate ... 8-costate, loculicidal at apex, septicidal at base; seeds 8- 20, compressed, roundish, ... finely punctate, black and shining.” The “punctate” seed description apparently refers to the polygonal areoles, visible at lower power magnification as reticulations, a common feature of hedyotoid seeds. Standley (1916) transferred M. shannoni (without comment) to Arcytophyllumas A. shannoni, but included only A. lavarum in his treatment of Rubiaceae for the North Ameri- can Flora (Standley 1921).(Apparently, A. muticum was not known from Panama and Costa Rica in 1921; see other comments below about A. muticum) Although Mena (1990) in his revision of Arcytophyllum had merely listed a avons serpyllacea in 1 his Canee On) olf Excluded and Dubious Names, none- il stothe prostrate, sulfruticose species of Arcytophyllum, eo that A. lia, m could be an extension of Arcytophyllum north- ward from Costa Rica into Guatemala and southern Mexico. The transfer of this species to Arcytophyllum was questioned by Andersson et al. (2002) based on their Analysis 1, data from the rpsl6 intron. A phylogenetic tree indicated a close relationship to Bouvardia rather than Arcytophyllum. Their study of A. serpyllaceum was based on a Mexican collection from the MO her- barium, Stafford et al. 203. Hoping to examine this collection we corresponded with Charlotte Taylor, curator of Rubiaceae at MO. She has been unable to find this herbarium sheet either under Hedyotis or related genera, although Andersson has written to Terrell stating that the loan of MO specimens to GB was returned to MO. Presently, the location of this specimen remains unknown. — Taste 1. Comparison of characters of Arcytophyllum muticum and A. serpyllaceum. Characters muticum serpyllaceum Roots and stems Stipules Leaves Flowers Calyces Corollas Corolla lobes Corolla tubes Stamens Stigmas Capsules Seeds Subshrubs’ prostrate, stoloniferous, mat-forming Genus: interpetiolar, entire or with toothed projections, glabrous or with pustuliform papillae Species: Decurrent, deltate, with acuminate projections 0.4-0.7 mm long, scattered short hairs Sessile or subsessile, 2.5-4.5 x 0.8-1.8 mm, ovate, transversely falcate, coriaceous, glabrous Solitary, sessile, apparently isostylous 5 mm long, trullate or triangular, glabrous, with intercalycine teet m long, salverform—funnelform, purple or bluish abaxially, white adaxially 2-45 x 2-3.5 mm, deltate, papillose, scattered hairs in margin and abaxially, scattered shiny scales adaxially 3-4.5 x ] -2 mm, glabrous Anthers 1 x 0.3 mm, slightly exserted 1 mm long, papillose adaxially 1-3 mm diam., globose, septicidal, 4-24 seeds/capsule oS Aatlne lenticular, often biconvex, hilum sl. raised, areoles often isodiametric, small. 4mm diam., often strongly compressed, often polygonal in Suffruticose herbs, prostrate, woody rhizomes, creeping, mat-forming pale to 2 mm long, ovate, with short to long na audae, marginal teeth with apical glands, glabrous to nee Sessile, 2-9 x ] glabrous -5 mm, ovate or elliptic, thickened, coriaceous, Solitary, pedicels to 7 mm long, heterostylous Lobes (1-)3-8 x 0.5-1.5 mm, lanceolate to ovate or obovate, glabrous, with intercalycine teeth ( m long, funnelform, greenish or purple abaxially, white adaxially, thickis 2.5-4.8 X 1.2-1.7 mm, ovate or elliptic, densely white pubescent adaxially 3-5 x 2-3 (at throat), glabrous abaxially, pubescent distally adaxially Anthers ca. 1-2 mm long, oblong, slightly exserted (thrum flowers), included (pin flowers) —2.3 mm long (pin), 1-1.4 mm long (thrum) 1.5-4 x 2-3.5 mm, broadly oblong or obovate, loculicidal 8-29 seeds/capsule 0.6-1.2 mm diam., other characters very similar to those of A.muticum VUINd9 AVIVIGNa NOSNIGOd GNV Tidaddl S26 926 BRIT.ORG/SIDA 21(2) In the absence of confirmation of the voucher for the study by Andersson et al. (2002), the present study follows the species placement based on struc- tural features of the plants. Two species of Arcytophyllum occur in Central America and are of particular interest here. These are A. lavarum, endemic to Costa Rica and Panama, and A. muticum, native to Costa Rica and Panama and extending south to Ecuador. (Mena 1990 does not list A. muticum from Costa Rica except as the type fora synonym, but there are nine collections from Costa Rica in herbarium US). The two similar species, A. muticumand A. serpyllaceum, are compared in Table 1, listing all of the main characters. The flowers of the two species differ in styly, but this is not considered a generic difference (e.g., Terrell 1996 found both stylies in Houstonia). The data on capsules suggest that A. muticum differs somewhat in size, shape, and number of seeds per locule (Table 1) from A. serpyllaceum. The two species differ also in A. muticum having septicidal dehiscence versus loculi- cidal dehiscence in A. serpyllaceum. Comparison of dehiscence in other genera of Hedyotideae shows considerable variation and the presence of septicidal (e.g., Hedyotis subg. Hedyotis) and loculicidal dehiscence (e.g., Houstonia and Oldenlandia). Contrary to statements by Andersson et al. (2002) that A. serpyllaceum seeds resemble those of Bouvardia, we find that their seeds are distinctly differ- ent, not only in the presence or absence of conspicuous or inconspicuous wings, but also in the appearance of the often raised pointed hila and in the presence of thick-walled isodiametric areoles versus thin-walled radially elongate ar- eoles. Comparison of seeds of A. muticum and A. serpyllaceum show them to be so similar that we did not detect any differences in shape, position of hilum, or sizes and shapes of the areoles, and without identity data it would be impos- sible to distinguish them (Fig. 1). Arcytophyllum lavarum was also similar to the other two species (Fig. 2). A more detailed study of additional species of Hedyotideae is needed in order to provide both molecular and morphological evidence concerning rela- tionships. It is hoped that time will permit such a survey to be carried out with assistance from a person knowledgeable in molecular studies. ACKNOWLEDGMENTS We thank the herbarium curators for loans of specimens, Susann Braden for the SEM work, and Marjorie Knowles for formatting the illustrations. We also thank Charlotte Taylor, Curator of the Rubiaceae at MO, for her assistance in searching for a missing specimen needed to confirm an identity (see text), and Pedro Acevedo for providing the Spanish Resumen. Piero Delprete and an anony- mous reviewer provided helpful comments on an earlier draft. TERRELL AND ROBINSON, RUBIACEAE GENERA Pe REFERENCES Anderson, L.E. 1954. Hoyer’s solution as a rapid permanent mounting medium for Bryo- phytes. The Bryologist 57:242-244. Andersson, L., J.H.E. Rova, and F. Atazata Guarin. 2002. Relationships, circumscription, and al biogeography of Arcytophyllum (Rubiaceae) based on evidence from cpDNA. Brittonia Biackwett, W.H., Jk. 1968. Revision of Bouvardia (Rubiaceae). Ann. Missouri Bot. Gard. 55:1- 30. Bremer, B.and J.-F. MANeN. 2000. Phylogeny and classification of the subfamily Rubioideae (Rubiaceae). Pl. Syst. Evol. 225:43-72. CHUNG, I.-C. 1967. Studies in Manettia (Rubiaceae) section Heterochlora Schum.Phytologia 15:272-288, CHUNG, |.-C.. 1968. Studies in Manettia (Rubiaceae) section Pyrrhanthos Schum. Phytologia 17:353-366. Lewis, W.H. 1965. Cytopalynological studies of African Hedyotideae (Rubiaceae). Ann. Mis- souri Bot. Gard. 52:182-211. Mena V., P. 1990. A revision of the genus Arcytophyllum (Rubiaceae: Hedyotideae). Mem New York Bot. Gard. 60:1-26. SmitH, J.D. 1893. Undescribed plants from Guatemala, XI. Bot. Gaz. 18:203-204. STANDLEY, PC. 1916. Studies of tropical American phanerogams. 2. Contrib. U.S. Natl. Herb. 18:128 STANDLEY, P.-C. 1921. Rubiaceae. N. Amer. Flora 32: 38-39, 96-111. STEYERMARK, J.A. 1974. Manettia. In: T. Lasser and J. Steyermark. Flora de Venezuela. Vol. 9. Edicion Especial del Instituto Botanico, Ministerio de Agricultura y Cria, Caracas. Pp. 121-158. TerrELL, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48:1— 118 Terrett, E.E. 1999. Morphology and taxonomy of Arcytophyllum serpyllaceum (Rubiaceae), a transfer from Hedyotis. Novon 9:263-264. TeRRELL, E.E. and S.D. Kocu. 1994. A new species of Bouvardia (Rubiaceae) fram Mexico, and transfer of Hedyotis xestosperma to Bouvardia. Novon 4:179-182. TerreLt, E.E.and H.Rosinson 2003. Survey of Asian and Pacific species of Hedyotis and Exallage (Rubiaceae) with nomenclatural notes on Hedyotis types. Taxon 52:775-782 Terrett, E.E. and R.P. Wunpertin. 2002. Seed and fruit characters in selected Soermacoceae and comparison with Hedyotideae (Rubiaceae). Sida 20:549-547. 928 BRIT.ORG/SIDA 21(2) THE 2004 DELZIE DEMAREE TRAVEL AWARD RECIPIENT Mr. Orlando Alvarez-Fuentes Michigan State University Mr. Orlando Alvarez-Fuentes was awarded the 16" Annual Delzie Demaree Travel Award at the 5lst Annual Systematics Symposium (8-10 Oct 2004) at the Missouri Botanical Garden. Applications for the 2005 Delzie Demaree Travel Award should include a letter from the applicant telling how symposium attendance will benefit his/ her graduate work and letter of recommendation sent by the major professor. Please send letters of application to: Dr. Donna M.E. Ware, P.O. Box 8795, Her- barium, Biology Department, The College of William and Mary, Williamsburg, VA 23185-8795, U.S.A. 1-757-221-2799; Email: ddmware@wm.edu. Applications need to be received 4 weeks ahead of the Symposium date. The Systematics Symposium dates for 2005 are 7-8 October. Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research Institute of Texas, to Barney Lipscomb, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 1-817- 332-7432; Email: barney@brit.org. Thank you. SIDA 21(2): 928. 2004 GENERIC PLACEMENT OF CHAPTALIA HINTONII (ASTERACEAE: MUTISIEAE) Guy L.Nesom Botanical Research Institute of Texas 09 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT t7] lly Chaptalia hintonii, o south- tential: Mexico, was recently transferred to the other g the basis of the occurrence of staminodes in the pistillate flowers. No clear position within Gerbera was pee for the species. In contrast, C. hintonii also has been World placed among the eight species of sect. Chaptalia, based on its similarity in a set of features includ- ing scape morphology, head behavior, ray corolla morphology, cypselar vestiture, flower biology, and geography. Other aes of south American and Mexican Chapane incl other from sect. Chaptalia, al 1] lesi either stigial as the result of ont eeeenediacy Bence bisexual disc florets and outer, pistillate ray florets. The hypothesis that C. hintonii belongs with Chaptalia is stronger than that for positioning C. hintonii in Gerbera. RESUMEN iq] ¢ fe hintonii, del centro-sur de México, fue transferida del Viejo Mu Gerbera en base a la presencia de estaminodios en las Bicres pistiladas. No se ae una posicion clara en Gerbera para la epiet pol contra, C. hintonii ha sido colocada entre las ocho especies de la sect. Chaptalia, en base a s litud en un conjunto de caracteristicas que incluyen la morfologia del escapo, comportamiento del Se morfologia de los radios de la corola, indumento de las cipselas, biologia floral, y geografia. Otras especies de Chaptalia de Sur América y México, incluida otra de la sect. Chaptalia, también se sabe que producen estaminodios, una casas inter peta eau como Sesionnen tanto como ocurrencia vestigial o como Sarr tre fldsculos pega del ase y flores radiadas pistiladas. La meen C. feeieee ce ptali jue F C.hintoniien Gerbera. In a recent discussion of the Mexican species Chaptalia hintonii Bullock and its relationship to other species groups of Mutisieae-Mutisiinae, Katinas (1998) concluded that C. hintonii should be placed in Gerbera rather than Chaptalia. In contrast, in a revision of the 12 North American and continental Central American species most recently considered to be Chaptalia, Nesom (1995) treated C. hintonii among the six species of sect. Chaptalia. Two additional spe- cies from Mexico were added to sect. Chaptalia by Cabrera and Nesom (2003). Indeed, the distinction between the New World Chaptalia Vent., 1802, and the Old World genus Gerbera L., 1758, is not sharply defined, as observed by Jeffrey (1967). Hansen (1990) provided additional perspective on taxonomic problems in the larger generic group of monocephalous, essentially scapose species of SIDA 21(2): 929-933. 2004 930 BRIT.ORG/SIDA 21(2) Mutisiinae (the “Gerbera-complex”) that includes these two genera and infor- mally suggested a return to earlier positions of Schultz-Bipontinus (1856) and Baillon (1882), who believed that the limits of Gerbera should include New World species now treated in Chaptalia. Nesom (1995) also noted that recogni- tion of the taxonomic boundary between Chaptalia and Gerbera is problem- atic but maintained Chaptalia as distinct. In the observations by Katinas (1998), she did not deal directly with sig- nificant considerations pertinent to a judgement of generic placement of Chaptalia hintonii—it is among the species that constitute sect. Chaptalia (Nesom 1995), and cypselar vestiture suggests that Chaptalia has evolved apart from Gerbera (Hansen 1990). Species of sect. Chaptalia are characterized by monocephalous, ebracteate or few-bracted scapes, heads nodding in bud, broad, cream-colored rays witha purple midstripe on the abaxial surface, and functionally staminate disc flow- ers. This set of features apparently does not consistently occur in any other group of Mutisiinae. Chaptalia tomentosa Vent., the type of the genus, is native to the Atlantic and Gulf coastal plains of the southeastern United States; the other seven species occur in montane habits in eastern and south-central Mexico. Within the section, three species groups can be recognized: (a) C. tomentosa and C. madrensis Nesom produce narrowly beaked cypselas and pistillate ray co- rollas without an inner lip; (b) C. pringlei Greene is alone in its heads without a zone of rayless pistillate flowers (all pistillate flowers produce conspicuous rays), and cypselas with a short neck; (c) C. lyratifolia Burkart, C. hidalgoensis Cabr. & Nesom, C. mexicana Burkart, C. hintonii, and C. estribensis Nesom are simi- lar in their colonial habit from rhizomes, leaves with cordate blades and mucronulate margins, pistillate flowers with relatively short style branches, and unbeaked cypselas. Chaptalia hintonii differs from C. lyratifolia C. hidalgoensis, C. mexicana, and C. estribensis primarily in its thicker, more eas- ily disconnected rhizomes, and thinner leaves, but the geographic and morpho- logical coherence of these five species within sect. Chaptalia seems apparent. The transfer of Chaptalia hintonii to Gerbera implies that the species belongs to neither the C. lyratifolia group (“group c” above) nor sect. Chaptalia, but Katinas’s observation on the implied non-cohesiveness of sect. Chaptalia is in- direct—only through her removal of C. hintonii. “The characteristic lack of staminodes in Chaptalia and their presence in Gerbera” led [Katinas] to place [Chaptalia] hintonii in the genus Gerbera” (Katinas 1998, p. 380). Also, “Chaptalia hintonii is much closer to Gerbera in its floral characteristics” (p. 382). She noted that for C. hintonii, “all florets bilabi- ate,” “outer ray florets conspicuously surpass the involucre,” and “ray florets possess staminodes” were features supporting its removal from Chaptalia. Nu- merous other species of Chaptalia, however, have all florets bilabiate, all spe- cies of sect. Chaptalia have large outer rays (e.g., see numerous published pho- —_— NESOM, GENERIC CHAPTALIA HINTONII 931 4 ] in the inner pistillate flow- ers of at least three other Mexican species: C. estribensis (sect. Chaptalia) from the locality in Oaxaca, but not from the localities in Hidalgo; C. texana Greene (sect. Leria (DC.) Burkart) in plants of Baja California Sur but apparently not from mainland localities; and C. hololeuca Greene (sect. Leria) from one local- ity in Coahuila but not from others. Similar staminodes also occur sporadically in both the outer and inner pistillate flowers of the South American species C. incana Cuatr., a species suggested by Cuatrecasas (1961) to belong in sect. Leria (DC.) Burkart. Finally, Burkart (1944) viewed a group of cordate-leaved South American Chaptalia species as better placed in Trichocline Cass. than Chaptalia, particularly in view of their production of staminodes in the outer pistillate florets. Jeffrey (1967) and Hansen (1990) found that pappus texture and cypselar vestiture refer these species to Chaptalia rather than to Trichocline or any group of the Gerbera-complex. Parallel variation in the production of staminodes is found in Old World Gerbera. Katinas, following Hansen (1990), noted that a reduction to vestigial staminodes has occurred in several groups of Gerbera and that they are com- pletely absent in Gerbera sect. Parva H.V. Hansen. Hansen (1990) and Nesom (1995) have observed that the inner pistillate florets in Chaptalia are intermediate between the disc florets and the outer ray florets. “The inner pistillate flowers of Chaptalia appear to be transitional be- tween the ligulate, outer, pistillate ones and the tubular, inner, bisexual flowers in two ways: 1) a) nee are bilabiate, even if the outermost are not, and 2) in some sf hey sporadically produce small, abortive stamens |[staminodes]” (Nesom 1995, p. 155). The occurrence of staminodes in Chaptalia and other en- tities of the Gerbera-complex is a plesiomorphic feature (Hansen 1990)—they may be interpreted as vestigial organs or as developmentally intermediate in origin (as above). A plesiomorphic similarity between Mexican C. hintonii and certain groups of Old World Gerbera does not imply closeness of relationship. tographs of C tomentosa).and st What genus? Evidence at hand indicates that Chaptalia hintonii belongs in the same genus as C. tomentosa, the type species of Chaptalia. If C. hintonii is to become a spe- cies of Gerbera, then so also must the type and at least the other species of sect. Chaptalia. Most of Chaptalia as a genus holds together well. Species of sects. Leria (DC.) Burkart (C. nutans Polak. the type), Archichaptalia Burkart (C. rotundifolia D. Don, the type), and Pseudotrichocline Burkart (C. isernina Cuatr, the type) are similar to those of sect. Chaptalia in their nodding buds and cypselar vestiture, which provides evidence for the phylogenetic coherence of Chaptalia apart from Old World members of the Gerbera-group. “Almost all taxonomic main groups lof the Gerbera-complex] can be cir- cumscribed by having a special type of achene (cypsela) hair” (Hansen 1990, p. 932 BRIT.ORG/SIDA 21(2) 471). Chaptalia has “achene hairs as in Gerbera sect. Piloselloides Less. but more narrow and either much smaller or much longer and then less inflated” (p. 476, Table 4). Katinas (p. 382) noted that “the species of section Piloselloides seem to be the most closely related [to Chaptalia] by virtue of the length of the outer ray florets and the type of pubescence on the cypselas...” but her Table | and related comments do not indicate that the cypselar pubescence in these two groups is identical. There is slight variation among cypselar hairs within sect. Chaptalia, but I find the vestiture of C. hintonii indistinguishable from that of other spe- cies of the C. lyratifolia group. Chaptalia sects. Chaptalia, Leria, and Archichaptalia are overlapping in morphological features—leaf shape, degree of elaboration of bracts on the scape, presence/absence of inner pistillate florets, style morphology, elaboration of the inner lip of pistillate florets, ray size, and degree of elaboration of cypselar beak. Especially in their long-petiolate, cordate leaves and broad rays, species of sect. Archichaptalia are similar to those of the Chaptalia lyratifolia group of sect. Chaptalia. Even the distinctive ovarian sterility of the central florets of sect. Chaptalia is not a totally exclusive feature. A Brazilian species (Dias de Moraes 1998) is said to be most closely similar to species of sect. Archichaptalia, but it apparently is anomalous there in its functionally staminate disc florets (as wellas lack of inner pistillate florets, ebracteate scapes, all characters found in some species of sect. Chaptalia, where the Brazilian species might techni- cally be placed). Cuatrecasas (1905) described variation in fertility between heads of different plants of the Peruvian C. malcabalensis Cuatr: one head with outer radiate pistillate florets and bisexual fertile central florets, lacking inner rayless pistillate florets, another head with outer radiate pistillate florets, in- ner rayless pistillate florets, and central florets with sterile ovaries, except for two florets with fertile ovaries and sterile anthers. Chaptalia sect. Lieberkuhna (Cass.) Burkart (C. piloselloides VahD Baker, the type) and sect. Loxodon (Cass.) Burkart (C. exscapa (Pers.) Baker, the type), as a combined group, stand apart from all the rest of Chaptalia in production of erect buds, potential for cleistogamous heads, and distinctive cypselar vestiture, and it is possible that this small group of species will be separated at generic rank. This group apparently is the one underlying Hansen’s notice (1990) that Chaptalia is separable into two groups, based on morphology of cypselar hairs. The Antillean group sect. Microchaptalia Burkart appears to be heterogeneous, but agree with Burkart (1969) that all of it belongs with true Chaptalia, rather than being di- vided between Chaptalia and Lieberkuhna, as suggested by Jeffrey (1967). Katinas found similarities between Chaptalia hintonii and Gerbera sects. Lasiopus (DC.) Schultz-Bip., Pseudoseris (Baillon) C. Jeffrey, and Piloselloides Less. (= Piloselloides (Less.) C. Jeffrey), although the species did not fit within any of them. In Hansen's strict consensus tree for the Gerbera-complex (1990), these three groups are most closely related to Chaptalia and phyletically sepa- ey NESOM, GENERIC PLACEMENT OF CHAPTALIA HINTONII 933 rate from Gerbera sensu stricto, which is the basal OTU in the whole Gerbera- complex and separated from Chaptalia by Leibnitzia Cass., Perdicium L., and others. Thus the transfer of C. hintonii to Gerbera appears equivalent to a step toward recognizing the whole Gerbera-complex asa single genus rather thana precise hypothesis of relationship. The ultimate fate of all of Chaptalia may be to join a much broadened Ger- bera, as suggested by Hansen and earlier botanists, or it may be maintained (perhaps disjoined from Lieberkuhna) among a group of smaller genera, as sug- gested by Jeffrey, but the taxonomic decision in either case should be made ina broad perspective. The taxonomic segregation of C. hintonii has little support compared to the geographic and morphological coherence of this species with Chaptalia sect. Chaptalia and the apparent phylogenetic unity of most of the whole genus Chaptalia. ACKNOWLEDGMENTS Lam particularly grateful to John Strother for various helpful comments on the manuscript. REFERENCES BaiLton, M.H. 1882. Histoire des plantes 8.Compositae. Paris. Burkart, A. 1944. Estudio del género de Compuestas Chaptalia con especial referencia a las especies argentinas. Darwiniana 6:505-594. Burkart, A. 1969. Commentario acerca de Chaptalia Vent. (Compositae) a raiz de las publicaciones de Baldwin and Speese (1947), Jeffrey (1967) y otras autores. Darwiniana 15:554-555. Caprera, L.and G.L. Nesom. 2003. A new species of Chaptalia (Asteraceae: Mutisieae) from Mexico and rediscovery of C. mexicana. Sida 20:1363-1370. Cuatrecasas, J. 1961. Studies on Andean Compositae V. Proc. Biol. Soc. Washington 74: Cuatrecasas, J. 1965. Miscelanea sobre flora Neotropica, Il. Ciencia (México) 23:121-124. Hansen, H.V. 1990. Phylogenetic studies in the Gerbera-complex (Compositae, tribe Mutisieae, subtribe Mutisiinae). Nordic J. Bot. 9:469-485. Jerrrey, C. 1967. Notes on Compositae: Il. The Mutisieae in east Tropical Africa. Kew Bull. ra eee KaTINAS, L. 1998. The Mexican Chaptalia hintoniiis a Gerbera (Asteraceae, Mutisieae). Novon 8:380-385. Moraes, M.D. 1998. Chaptalia hermogenis (Asteraceae: Mutisieae), a new species from the Brazilian Atlantic rain forest. Novon 8:173-175. Nesom, G.L. 1995. Revision of Chaptalia (Asteraceae: Mutisieae) from North America and continental Central America. Phytologia 78:153-188. ScHuLtz-Bipontinus, C.H. 1856. Flora of the north-western Mexico. Compositae. In: B.C. Seemann. The botany of the voyage of H.M.S. Herald. London. Pp. 297-315. 934 BRIT.ORG/SIDA 21(2) COMMEMORATIVE Raymond John Taylor Jr. 20 January 1930-9 November 2004 As this issue of Sida was going to press, | received the news that my good friend and colleague had just passed away. Dr. John Taylor retired from Southeastern Oklahoma State University, Durant in 1990 after a long and productive career in botany. | remember John as an excellent teacher and an accomplished bota- nist who enjoyed field work and collecting plant specimens. A full memorial is forthcoming in Sida.—Barney Lipscomb, Botanical Re- search Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 21(2): 934. 2004 THE GERBERA COMPLEX (ASTERACEAE: MUTISIEAE). T@ SPLIT OR NOT TO Seer Liliana Katinas Departamento Cientifico de Plantas Vasculares Museo de La Plata Paseo del Bosque 1900 La Plata, ARGENTINA ABSTRACT The transfer of Chaptalia s rejected by Nesom (this issue), who retains the species in Chaptalia sect. Chantal Chaptalia and Gerbera belong to the Gerbera-complex, a ouR of scapose genera whose ¢ encumisel plenel are still in flux and whose largest genera still lack Important f the corollas, staminodes, and c ypsel a hairs are aise ussed, and a key to genera of the ee bera-complex is provided. The short, filiform corollas of the inner ray florets and the lack of Sa oaee are eaaraciers coat all ow the circumscription of Chaptalia.C within it must be rede- Gned. Staminodes anc inner ray florets with bilabiate Eerollns that are eee than the style are two characters that confirm the transfer of Chaptalia hintonii into the g Key worps: Gerbera hintonii, Chaptalia, Gerbera-complex, corollas, staminodes, cypsela hairs RESUMEN La transterencia de Chaptalia hintonii al gé Gerbera fue rechazada por Nesom (este volumen), quien retiene esta especie en C Payton ae ieee Chaptalia y Gerbera j al complej t £. J vascular a bundle on leaf base taken on day 2. (PE = petiole epidermis; V = vascular tissue; C issue; zone of shoot emergence). 948 BRIT.ORG/SIDA 21(2) be J ar J t reg . . 1 ra Fin S_F 1] of J Fics. 4—5. Scanning elect i hs of Sed ightii | lesi | ling days. Fig. 4. Emerging } t$5] L J Q 3+h remains of callus tissue. (PL = primordial leaf; ZRE = zone of root emergence). GRAVATT AND TAYLOR, ASEXUAL REPRODUCTION IN SEDUM WRIGHTII 949 Fics. 6—7.S ing el i hs of Sed igh | ling days. Fiq. 6. Leaf pair and elongating root taken on day 10. Fig. 7. Root hair formation, taken on day 14. (C = callus tissue; RH = root hairs). 950 BRIT.ORG/SIDA 21(2) of S. stahlii, left on a dry surface, gave rise to roots first and then shoots. Yarbrough (1936) further observed that in a moist chamber, as used in this study, shoots were not formed and roots only appeared occasionally. Yarbrough (1936) concluded that production of roots and shoots from leaves of Sedum stahlii, which at the time of detachment clearly possess no residual meristem, clearly is an example of regeneration through the formation of ad- ventitious structures. He further stated that the formation of callus tissue by the thinned-walled parenchyma cells of the short petiole involves differentia- tion. In this study, it appears that this is the case for Sedum wrightii as well, and compares well with the findings of Yarbrough (1936). The continued mitotic activity of the cells surrounding the callus makes possible the development of two fundamental growing points, the root and shoot primordia, and the subse- quent development of a new plant from the detached leaf. ACKNOWLEDGMENTS We would like to thank Guy Nesom (BRIT) and Sherwin Carlquist (SBBG) for various helpful comments on the manuscript. REFERENCES CLAUSEN, R.T. 1975. Sedum of North America north of the Mexican Plateau. Cornell Univer- sity Press, Ithaca. Gravatt, D.A. 2003. Crassulacean acid metabolism and survival of asexual propagules of Sedum wrightii. Photosynthetica 41:449-452. Green, P.B.and K.E. Brooks. 1978.Stem formation from a succulent leaf:lts bearing on theo- ries of axiation. Amer. J. Bot. 65:13-26. HoutHe, PA. and S.R. Szarek. 1985. Physiological potential for survival of propagules of crassulacean acid metabolism species. Plant Physiol. 79:219-224, Jordan, PW. and PS. Noset. 1981. Seedling establishment of Ferocactus acanthodes in rela- tion to drought. Ecology 62:901-906. McVeich, |. 1938. Regeneration in Crassula multicava. Amer. J. Bot. 25:7-110. Mims, C.W. 1981. SEM of aeciospore formation in Puccinia bolleyana. Scan. Electron Microscop. 3:299-303. SHaritz, R.R.and J.F. McCormick. 1973.Population dynamics of two competing annual plant species. Ecology 54:723-740. Stouot, H.N. 1934. Gemmipary in Brynesia weinbergii. Amer. J. Bot. 21:562-572. Stouot, H.N. 1938.Gemmipary in Kalanchoe rotundifolia and other Crassulaceae. Amer. J. Bot. 25:106-110. YARBROUGH, J.A. 1934. History of leaf development in Bryophyllum calycinum. Amer. J. Bot. 21:46/7-484. YarBROuGH, J.A. 1936. Regeneration in the foliage leaf of Sedum. Amer. J. Bot. 23:303-307. TAXONOMY, DISTRIBUTION, AND MEDICINAL USES OF LEGUME TREES OF PAKISTAN Mohammad Athar' Zahoor Ahmad! rs fA : | ip L | California Department of Food and Agriculture j gy 2014 Capitol Avenue, Suite 109 and Gencie Resources Sacramento, California 95814, U.S.A. Pakistan Agricultural en Council atariq@cdfa.ca.gov Islamabad, PAKIS ABSTRACT A literature search was conducted to determine the medicinal legume trees found in Pakistan and used as remedies for various human disorders or diseases. The taxonomic position of these trees is discussed and their distribution is described to facilitate their collection. There are 40 medicinal, egume tree species belonging to 25 genera of Fabaceae (Leguminosae). Most of these medicinal le- gume trees belong to Caesalpinioideae (10 genera and 16 species) followed by Mimosoideae (7 genera and 14 species) and Papilionoideae (8 genera and 10 species). This constitutes about half of the Paki- stani legume trees possessing medicinal properties used for various ailments. The plant parts usec for medicinal purposes included bark, roots, stem, Teaves flowers PONS and Seed These edicinal | tT COMMMCI~ jam trees have good vegetative growth under cial exploitation. There is potential for growing medicinal plants in palaerana as an income cee nese medicinal legume trees. However, there is a need for activity and for sustainable utilization of t systematic cultivation, commercial extraction using scientific netiees and sustainable conserva- participation. tion of endemic medicinal legume trees for substantial utilization t RESUMEN Se realizo una ee bibhiogratica para eet icar los arbol es medion ce las leguminosas Se discute que se encuentran en Pakistan y a posicion one de estos arboles Hay 40 especies medicinales de leguminosas arboreas, pertenecientes a 25 géneros de Fabaceae (Leguminosae). La mayoria de estas leguminosas arboreas medicinales pertenecen a las Caesalpinioideae (10 géneros y 16 especies) seguidas de Minesoleeas a penehee y 14 ea y ; y se deseube: su distribucion para facilitar su recolecci6n. Papilionoideae (8 géneros y LO especies). F t pakistanis que tienen propiedades medicinales eee para varios remedi usadas con propositos medicinales incluyen la corteza, raices, tallos, hojas, flores, frutos y semillas. Estos arboles medicinales tienen buen c crecimiento vegetativo en condiciones de cultivo y tienen xplotacion comercial. En Pakistan hay potencial para el incremento ios. Las Sante: de la planta amplio abastectiuento pate l | ividad gener adora de recursos econdmicos y para uso medicinal s 4 de las p my By | i ] : 1% Pero oO mediante métodos cientificos y conservacion sostenible de leguminosas arboreas medicinales endémicas para una utilizacion importante mediante participacion de la comunidad. 'Disclaimer: The views expressed in tls article are those a age and do not necessarily ue ent those of f this endorse- their respective departments. The publication of | t herbal icine. The authors or their departments make no warranty, for medical purposes. ment of the use of tl expressed or ape ae assume no legal liability for the use of these legume trees SIDA 21(2): 951-962. 2004 952 BRIT.ORG/SIDA 21(2 INTRODUCTION Fabaceae (Leguminosae) is the third largest family of flowering plants com- prising of 750 genera and 20,000 species with worldwide distribution (Sprent 2001). In Pakistan legumes form a considerable portion of the flora and are the third largest family in order of abundance (Kirkbride 1986). Ali (1973a, b; 1977) reported 107 genera and 539 species of legumes from Pakistan of which 68 gen- era have one or more native species (Kirkbride 1986). There are about 82 legume trees in Pakistan of which 27 tree species belong to Caesalpinioideae, 36 to Mimosoideae and 19 to Papilionoideae (Ali 197 3a, b; 1977). These trees have con- siderable significance as source of timber, lumber and fire wood. They are also used as ornamental and shade trees in gardens, orchards and roadside planta- tions (Malik & Farooq 1984). Classification of medicinal plants is organized in different ways depend- ing on the criteria used. In general, medicinal plants are arranged according to their active principles in their storage organs, particularly roots, leaves, flow- ers, seeds and other plant parts (Athar & Nasir 2004; Athar & Siddiqi 2004; Foster & Duke 1990; Moerman 1986, 1991, 1996; Shaheen et al. 2003). These prin- ciples are valuable to human for treatment of different diseases (Borchers et al. 2000; Duke 1997; Shaheen et al. 2003; Shinwari @ Khan 1998). Reports on the classification of many Pakistani medicinal plants that could be used for rem- edies and medicinal preparations are sporadic or lacking (Athar & Nasir 2004; Athar & Siddiqi 2004; Durrani et al. 2003; Saeed et al. 2004). Athar and Siddiqi (2004) described the taxonomy, distribution and flowering period of 95 spe- cies used as medicinal flowers in Pakistan. In another study, Athar & Nasir (2004) described the taxonomy of 78 plant species yielding vegetable oil used in cosmetics and skin and body care products. This third publication in the series presents the taxonomy of medicinal legume trees of Pakistan and their regional distribution, and summarizes the utilized tree parts and their princi- pal method of application for various ailments. MATERIALS AND METHODS A literature search was conducted to determine the legume trees of Pakistan that are used as remedies for various disorders or diseases (Allen & Allen 1981: Athar & Siddiqi 2004; Bach 1994; Baquar 1989; Borchers et al. 2000; Duke 1997; Gunasena & Hughes 2000; Malik & Farooq 1984; Nasir & Rafiq 1995; Palaniswamy 2003; Rasool 1998; Rizvi 1998; Said 1969: Shaheen et al. 2003; Shinwari & Khan 1998; William @ Ahmad 1999; Zaman & Khan 1970), and their taxonomic position was determined. The genera were arranged alphabeti- cally within sub-families. The scientific and local names are provided that are commonly associated with these legume trees. Their distribution in various parts of Pakistan is also described to facilitate their collection. The nomencla- ATHAR AND AHMAD, MEDICINAL USES OF LEGUME TREES OF PAKISTAN 953 ture and classification followed Polhill and Raven (1994), and author citations followed Brummitt and Powell (1992). RESULTS AND DISCUSSION The taxonomic position of 40 species of medicinal legume trees is given in Table 1. These legume trees are distributed in 25 genera and all three sub-families of Fabaceae (Leguminosae). This constitutes about half of the Pakistani legume trees possessing medicinal properties used for various ailments. The list is com- prehensive, yet not complete as more tree species could be investigated and added to the list. Most of these medicinal legume trees belong to Caesalpinioideae (10 genera and 16 species) followed by Mimosoideae (7 genera and 14 species) and Papilionoideae (8 genera and 10 species). The plant parts used for medici- nal purposes included bark, roots, stem, leaves, flowers, fruits and seeds. Table 1 summarizes the ailments, tree parts and principal methods of application. Spe- cific prescriptions, such as dosage and frequency, are not given, as these are rarely specified in literature and can vary considerably among users according to the recipe. Moreover, the table does not rank the order of the species importance as a medicinal source for the ailments, though this can be found in Hamdard Phar- macopoeia of Eastern Medicine (Said 1969). Legume trees are amazing natural resources. These trees provide wood for home building, packaging that keeps food fresh during shipment, and natural products like cellulose that are found in everything from the instant soup that we eat to the fabrics that we wear. Legume trees give us food, fodder, fabric, fur- niture, fatty acids, oils, tannins, gums, medicines, even cosmetics (Allen & Allen 1981), and the list goes on and on expanding to their nitrogen-fixing properties (Allen & Allen 1981; Sprent 2001; Subba Rao @ Rodriguez-Barrueco 1993). We get food, clothing and shelter from legume trees all in one environmentally friendly package. Essential oils have been extracted from legume trees for many years for their healing and nurturing properties (Athar & Nasir 2004). Today we use these same essential oils in aromatherapy to promote health, well-being and balance in body, mind and spirit. These plants are ever lasting, easily avail- able and centuries old, tested sources for healing various ailments (Athar & Siddiqi 2004, Athar & Nasir 2004). An estimated 80% of the rural population in Pakistan depends on tradi- tional medicines for their primary healthcare needs using legumes and other plants or their active principles (William & Ahmad 1999). By trial and error, these people have learnt to employ certain plants, plant parts and extracts to cure diseases and ailments. In this context, certain legume trees also merit con- sideration for their therapeutic properties, which are being commercialized. It is worth mentioning that nomadic people and rural populations are much more familiar with the use of medicinal plants as compared to people living in the Taste 1. Taxonomy, distribution and medicinal uses of legume trees in Pakistan. Species Local Name English Name Distribution Medicinal Use Fabaceae (Leguminosae) Caesalpinioideae Bauhinia purpurea L. Kaliar Bauhinia semla Wunderlin Kandla Bauhinia variegata L. Kachnar Caesalpinia bonduc (L.) Fevernut Roxb. Caesalpinia pulcherrima Gul-e-mohur Cassia fistula L. Amaltas Ceratonia siliqua L. Kharnub Chamaecrista absus (L.) Chasku H.S. Irwin & Barneby Delonix regia (Bojer Gulmohar ex Hook.) Raf. Purple Bauhinia Bauhinia Mountain ebony Gray nicker bean Paradise flower Golden shower, Indian laburnum Carob Cassia Peacock flower Grown in Punjab, NWFP. Rawalpindi Jowlian Reserve, Rawalpindi, Hazara Grown in gardens of Punjab and Sindh Grown in gardens of Pakistan Grown in gardens of Pakistan Naturalized throughout Pakistan Introduced and naturalized in Punjab, Islamabad and NWFP Wild in open places in NWFP and Punjab Planted in Karachi, Hyderabad and Lahore Flowers are used as nuraative. Flowers and buds used for indigestion. Stem bark decoction used for body pain and fever. Paste from bark is given for cancerous growths in stomach. Bark is astri {in diarrh re Decoction of leaf is used for | Jache and malaria. Flowers are aperient. Leaf decocti ken f | Idi pulverized seed infusion taken for fever and in- testinal worms. Root decoction used for fevers; flowers used for asthma and bronchitis and as anti-pyretic and expectorant. Flowers used for cough, diphtheria, constipation and edema. The pods are astringent and are used for coughs. Seed husk is astringent and purgative. Seeds used for skin diseases. Leaf decoction taken for constipation. Flowers are anthelmintic. 9S6 (Z)LZ VdIS/DYO'LIYS Table 1.continued Species Local Name English Name Distribution Medicinal Use Parkinsonia aculeata L. Saraca indica L. Senna alata (L.) Roxb. Senna alexandrina Mill. Senna italica Mill. Senna siamea (Lam.) H.S. Irwin & Barneby Tamarindus indica L. Valaiti kikar Ashok Dadmurdan Sennahindi Kasordhi Kasood tree Imii Jerusalem-thorn Ashok tree Ringworm Cassia Indian senna Senna Siamese shower Tamarind Planted in Sindh, Karachi, Hyderabad, Punjab and Lahore Cultivated in gardens in Punjab and Sindh Sometimes cultivated in Pakistan Wild in Punjab and Sindh Found in Punjab and Sindh Cultivated in Karachi, Sindh Grown in Sindh, Punjab, Jehlum, Karachi Leaf, fruit and stem decoctions taken for fever and also are abortive. Flowers and leaf poultice used for rheumatism. Bark is astringent; used in menorrhoea and uterine infections; also used for scorpion-sting. Flowers are laxative and also useful for softening skin. Stem, pods and leaves are useful in habitual castiveness. These lower bowels, increases peri- staltic movements of the colon by its local ac- tion upon the intestinal wall. Plant is used as expectorant, wound dresser, antidysentric carminative and laxative. Plant useful for malaria and constipation. Pods used as laxative and seed for stomachace. Flowers used as anthelminti anti-hypertensive, laxative, tranquilizer and sedative and for asthma, dandruff and insomnia. Leaf decoction taken for throat infection, intes- tinal worms and liver ailments. Flowers used as cy: anti-viral against new castle disease virus and are astringent and sedative. Fruits used for loss of appetite, constipation and rheumatism. QVWHY ONY YVHLY 4 NVLSINWd 40 S391 JWND37 40 S3SN IWNIDIGIW S56 Tale 1. continued Species Local Name English Name Distribution Medicinal Use Fabaceae (Leguminosae) Mimosoideae Acacia catechu (L.f.) Willd. Khair, Katha Black cutch Distributed in the Bark is astringent. Root is made into paste and foothill zone of NWFP applied on joints for rheumatism. and Punjab Acacia farnesiana (L.) Willd. Kabuli kikar Sweet acacia Grows in Sindh, Karachi, Leaves used for fever, typhus, tuberculosis Punjab gangrene, bladder infection and wounds; flower tea used for stomachache; crushed fruit juice used for diarrhea and eye and throat infections. Acacia nilotica (L.) Delile Kikar Gum-arabic Found in Sindh, Punjab, Flowers used for jaundice and palpitation. Leaf Baluchistan, NWFP juice is taken with fennel for bloody dysentery. Powdered bark given for acute diarrhea. Extracts from root and bark are used as taenifuge, and gum is used as emollient. Acacia senegal (L.) Willd. Khor Gum-arabic acacia Karachi, Sindh, Gum demulcent and emollient used internally Adenanthera pavonina L. Albizia chinensis (Osbeck) Merr. Albizia julibrissin Durazz. J Barighumchi Ghulabii siris Bead tree Silk tree Balochsitan Grown in Sindh, Karachi Sub-Himalayan tract, Punjab Outer Himalayan zone for intestinal mucosa and externally applied on inflammations Wood extract used for migraine and headache; bark and leaf decoction for diarrhea, dysentery and tonsillitis. Infusion of the bark used as lotion for cuts, sca- bies and skin diseases, Flowers used for mild constipation, boils carbuncles and swelling. 956 (Z)LZ VaIS/¥O'LING Tale 1. continued Species Local Name English Name Distribution Medicinal Use Albizia lebbeck (L.) Benth. Siris Lebbek tree Found in Sindh, Punjab, Similar to Albizia julibrissin Bajaur, Malakand Albizia procera (Roxb.) Benth. Sufaid-Siris Lebbek tree Cultivated and Leaves used as poultice on ulcer. Plant has naturalized, Sindh insecticidal properties. Karachi, Hasanabdal, Mirpur Leucaena leucocephala Ipil-Ipil Ipil-lpil Introduced in Sindh, Root decoction taken for fever; leaf tea taken for (Lam.) de Wit Punjab, NWFP typhoid and digestive tract ailment; root and twig decoction used for severe back pain; root and bark are abortive. Pithecellobium dulce Jungle Jaleebi Blackbead Found in Sindh and Root bark decoction taken for dysentery; leaf (Roxb.) Benth. Karachi used for indigestion; leaf poultice used for con- vulsions, venereal lesions and pain; fruit pulp used for hemoptysis; seed used for nasal con- gestion and internal ulcer. Prosopis cineraria (L.) Druce Jhand Mesquite Found in Sindh, Flowers are beneficial against miscarriage. Baluchistan and Punjab Prospis juliflora (Sw.) DC. Jhand Mesquite Found in Sindh, Eye drops made from leaf juice or cooked leaves Baluchistan and Punjab oe applied or taken orally for eye infections. trunk used for colds and flu. Fresh root taken orally for diarrnea.Bark and fruit decoction taken for bronchial infection and si- nus congestion NWILSINVd 40 $3941 JWN937 40 S3SN TVNIDIGIW ‘OVWHY ONV YVHLY £56 Tale 1.continued Species Local Name English Name Distribution Medicinal Use Samanea saman (Jacq.) Merr. Siris Fabaceae (Leguminosae) Papilionoideae Butea monosperma (Lam.) Taub. Dhak albergia sissoo Roxb. ex DC. Shisham Erythrina stricta Roxb. var. Dhauldak suberosa (Roxb.) Niyomdham Erythrina variegata L. Pangar Lebbek tree Flame of the forest, Bengal Kino Sissoo Coral tree Variegated coral tree Grown in Sindh, Karachi Cultivated in Punjab and NWFP, sporadic in Rawalpindi district Widely distributed in Punjab, Sindh and NWFP Sadiqpur Hills, Murree Road, Mirpur Cultivated as road side tre Fruit decoction used for BEE Os nervousness; fruit ingested for d tea or infusion used ~~ eoneioonar seed chewed for throat infection. Anti-pyretic, appetizer, aphrodisiac, blood purifier diuretic tri + Ai rj AA - Leaves are tonic, astringent; flowers are astringent, diuretic, depurative and aphrodisiac. Bark and seeds are used for snakebites. Leaves are bitter and stimulant; decoction is used in gonorrhea;root iS astrinaent.Wood js also used in leprosy, boils, eruptions and to stop vomiting. The bark is laxative, diuretic, EEN goog ex- pectorant, anthelmintic ibili and the decoction is given for dysentery and as a vermifuge and is effective as an eye lotion in ophthalmia. Leaves are anthelmintic, cathartic, glalactagoque and discutient. Leaf juice used externally as a oe for ulcers and for killing cass in es contain a mixture of alkaloid, hyaphorine is present in leaves, seeds and bark. Apart from hyaphorine, betaine and choline are also found. 856 (Z)LZ VaIs/uO'LIYE Table 1. continued Species Local Name English Name Distribution Medicinal Use Gliricidia sepium (Jacq.) Kunth ex Walp. Millettia pinnata (L.) Panigrahi <—- Ougeinia oojeinensis (Roxb. Peltophorum pterocarpum (DC) Backer ex K. Heyne Sesbania grandiflora (L.) Pers. Sesbania sesban (L.) Merr. Sapium/Lal Tali Karanja Sandan Peela Gulmohar Bansa Janter Madre tree Pongam Sandan Copperpod tree Sesbania Sesbania Only at experimental stations at Karachi and Islamabad Cultivated in Sindh, Cultivated in Punjab Gardens; Mirpur east (Sub-Himalayan tract MEL Cultivated in Punjab as road side plantation Planted in Karachi, Kutch, Sindh, Punjab Cultivated in Sindh, Punjab Leaf bath used for fever pain; leaf tea used for sinus inflammation and gonorrhea; root tea used kidney ailments, edema and hepatitis; leaf poultice used for skin diseases and wounds. Flowers are used for diabetes. The bark is febrifuge. The exudation obtained after incision is used in dysentery and diarrhea. The decoction is given to patients having highly colored urine; also used as fish poison. In traditional medicine, the bark used in various preparations as a tonic or as a astringent to cure or relieve intestinal disorders like consti- pation and ringworms, afterpain at childbirth, sprains, bruises and swellings, or as a lotion of eye troubles, muscular pains ae sores. Leaves in form of decoction used to w althy skin. Root paste used for rheumatism; root resin used for phlegm; bark decoction used for fever and diabetes; flower juice improves vision (as eye drops). Leaf compress applied to infected areas for suppuration. Anti-fertility activity of flowers also reported NWLSIMWd 40 S39YL IWN5D37 40 S3SN TYNIDIGIW ‘QVWHY ONV YVHLY 656 960 BRIT.ORG/SIDA 21(2) cities (Durrani et al. 2003; Saeed et al. 2004; William & Ahmad 1999). It is inter- esting to note that elderly people, particularly women, were more aware of the value of the medicinal flora than younger generations. Nomadic people and rural populations depend upon legume trees not only as medicinal plants but also for fuel wood and fodder and forage for their livestock. Some of the medici- nally important plants are over-harvested due to their extensive use, which may lead to their extinction. Overgrazing and cutting of legume trees for fuel wood and commercial exploitation has also resulted in reduced vegetation cover, pro- moted soil erosion and deterioration of Pakistani rangelands. However, legume trees cultivated in the gardens, parks, and roadside and riparian plantations in Sindh and Punjab have good vegetative growth and are in ample supply for com- mercial exploitation. Pakistan has the potential for sustainable utilization of its medicinal legume trees, and mainly exports crude plant materials that have a low value in the in- ternational market (Shaheen et al. 2003). On the other hand countries like China and India export value-added plant extracts standardized on the basis of their active ingredients. There is enormous price difference between crude plant ma- terial and properly processed plant extracts. Moreover, developed countries do not allow the import of unprocessed raw plant material due to rigorous phytosanitary regulations (Crockett @ Khan 2003; FAH 2003). It is therefore im- portant that Pakistani producers and traders of medicinal plants should consider adding value to their exports by carrying out the initial processing and quality evaluation in Pakistan. However, attention needs to be paid to the systematic cul- tivation, scientific extraction and sustainable conservation of endemic medici- nal legume trees for substantial utilization through community participation. A team from the Pakistan Forestry Institute, Peshawar has carried out an ethnobotanical study in northern areas of Pakistan (Rasool 1998). This study identified economically valuable trees that can be collected from the wild and medicinal plants that can be grown commercially on common or agricultural land. Several species are no longer found in the area, including Taxus baccata, the source of the anti-cancer agent Taxol. There is potential for cultivating me- dicinal plants as an income-generating activity (Athar & Nasir 2004; Athar & Siddiqi 2004). Further research should be carried out on the floristic composi- tion of medicinal plants and the present status of medicinal cum economic plants, including their market potential within the country and in the interna- tional market. The study should also look at the prospects for conserving en- demic medicinal and economic species for sustainable use through commu- nity participation. ACKNOWLEDGMENTS Special gratitude is expressed to Joseph H. Kirkbride, USDA, Agricultural Re- search Service, Beltsville, MD, for his help in checking the nomenclature and ATHAR AND AHMAD, MEDICINAL USES OF LEGUME TREES OF PAKISTAN 961 taxonomy of the plants and M. Akmal Siddiqi, Marshfield Clinic Research Foun- dation, Marshfield, WI, USA for valuable suggestions and helpful criticism on the manuscript. The authors are also grateful to M. Iqbal Choudhary, H. E. J. Research Institute of Chemistry, University of Karachi, Karachi, Pakistan for providing many original reprints used in the preparation of this paper. REFERENCES Atten, O.N and E.K. ALten. 1981. The leguminosae. A source book of characteristics, uses, and nodulation. The Wisconsin University Press, Madison, WI. Au, S.l. 1973a.Mimosaceae. Flora West Pakistan. 36:1-41. Au, S.1. 1973b. Caesalpiniaceae. Flora West Pakistan. 54:1—47. Au, S.l.1977. Papilionaceae. Flora West Pakistan. 100:1—-389. AtHAR, M.and S.M.Nasir. 2004. Taxonomic perspectives of plant species yielding vegetable oils used in cosmetics and skin care products. Int. J. Biol. Biotechnol.1(4).In Press. AtHar, M.and M.A. Sippial.2004.Some reflections on the taxonomy of medicinal flowers of Pakistan. Sida 21:357-369. Bacn, E. 1994. Flower remedies. Alternat. Med. 232-243. Baguar, S.R. 1989. Medicinal and poisonous plants of Pakistan. Printas, Karachi, Pakistan. Borcuers, A.T., C.L. Keen, J.S. Stern, and M.E. GersHwin. 2000. Inflammation and Native Ameri- can medicine: the role of botanicals. Amer. J. Clinic. Nutr. 72:339-347. Brummitt, K.R.and C.E. Powet (eds.). 1992. Authors of plant names. Royal Botanic Gardens, oe Crockett, S.L. and |.A. KHAN. 2003. Challenges of standardization: Marker compounds in plant species related and unrelated to top-selling herbs. J. Herbs Spices Med. PI. 10(3).: 13-24 Duxe, J.A. 1997. The green pharmacy. Rodale Press, Emmaus, PA. Durrani, M.J., A.M. Tay and F. Hussain. 2003. Folk medicinal plants of Nushki, District Chagai, Pakistan. J. Sci. & Tech.; Univ. Peshawar. 27:45-51. FAH (ForsSCHUNGSVEREINIGUN DER ARZNEIMITTEL-HERsTELLER €. V.) [German Research Association of Medicine Manufacturers]. 2003. Standard operating procedures for inspecting culti- vated and wild crafted medicinal plants. J. Herbs Spices Med. PI. 10(3):109-125. Foster, S.and J.A. Duke. 1990. A field guide to medicinal plants: eastern and central North America. Houghton Mifflin Co., Boston, MA. GuNASENA H.PM.and A. HucHes. 2000. Fruits for the future 1. Tamarind. Jamarindus indica L. International Center for Underutilized Crops (ICUC), UK. Kirkerive, J. 1986. Phylogeny and classification of Pakistani legumes. Pakistan J. Bot. 18: 287-299. Mauik, S.and S. Farooa. 1984. Cultivated trees, shrub and climbers of gardens of Pakistan. Taxonomic studies. PCSIR, Peshawar Laboratory, Peshawar, Pakistan. Moerman, D.E. 1986.Medicinal plants of North America. Museum of Anthropology, Univer- sity of Michigan, Ann. Arbor, MI. 962 BRIT.ORG/SIDA 21(2) Moerman, D.E. 1991. The medicinal flora of native North America: an analysis. J. Ethnopharmacol. 31:1-42. Moerman, D.E.1996. An analysis of the food plants and drug plants of native North America. J.Ethnopharmacol. 52:1-22. Nasir, YJ. and R.A. Raria. 1995. Wild flowers of Pakistan. Elite Publishers Ltd., Islamabad, Pakistan. PALANiswamy, U.R.2003.A guide to medicinal plants of Asian origin and culture. CPL Scien- tific Publishing Services Ltd., Newbury, UK. PotHitt, RM. and PH. Raven. 1981. Advances in legume systematics, part 1. Royal Botanic Gardens, Kew, UK. Rasool, G. 1998. Medicinal plants of northern areas of Pakistan: Saving the plants that save us. Pakistan Forest Institute, Peshawar, Pakistan, Saeed, M., M. ArsHap, M. AHmab, E. AHMab, and M. IsHaque. 2004. Ethnophytotherapy for the treatment of various diseases by the local people of selected areas of NWFP (Pakistan). Pak. J. Biol. Sci. 7:1104-1108 Saib, H.M. (ed.). 1969. Hamdard pharmacopoeia of eastern medicine. Time Press, Karachi. SHAHEEN, F., M. ATTAUR RAHMAN, K.VasisHT and M.laBal CHOUDHARY. 2003, The status of medicinal and aromatic plants in Pakistan, In: Medicinal plants and their utilization, UNIDO. Pp. 77-87. SHinwael, M.|.and M.A. KHAN. 1998. Ethonobotany of Margalla Hills National Park of Islamabad. Department of Biological Sciences, Quaid-i-Azam University, Islamabad and PASTIC — Press, Pakistan. SpreNT, J. 2001. Nodulation in legumes. Royal Botanic Gardens, Kew, UK. SupBa Rao, N.S. and C. Ropricuez-Barrueco (eds.). 1993. Symbioses in nitrogen-fixing trees. International Science Publications, New York, NY. Wittiam, J.T.and Z. AHmab. 1999. Priorities for medicinal plants research and development in Pakistan. Published by MAPPA, India and IDRC, Canada. Zaman, M.B. and M.S. KHAN. 1970. Hundred drug plants of West Pakistan. Pakistan Forest Institute, Peshawar and Ferozsons Ltd., Lahore, Pakistan. ABOUT 450 MILES UP THE RED RIVER IN A LARGE PRAIRIE: PETER CUSTIS’ SURVIVING BOTANICAL INFORMATION FROM THE RED RIVER EXPEDITION OF 1806 Michael H.MacRoberts and Barbara R. MacRoberts Bog Research, 740 Columbia Shreveport, Louisiana 71104, U.S.A. Herbarium, Museum of Life Sciences and Red River Watershed Management Institute Louisiana State University-Shreveport, Shreveport, Louisiana 71115, U.S.A. ABSTRACT In 1806 Peter Custis, botanist on the Red River Expedition, collected 26 plants mainly in the prairies in what is today Caddo and Bossier pari shes, Louisiana. These were sent to Benjamin Smith Barton in Philadelphia. Two of the specimens are still in the Barton Herbarium. They are very unusual for the region today. We have looked for the other 24 specimens but without success. Those 24 specimens would certainly aid in understanding the ecology of the Red River area in 1806. RESUMEN En 1806 Peter Custis, botanico en | pedicion del Red River, colect6 26 plantas principalmente el lo p P P p que hoy son los municipios de Caddo y Bossier, Louisiana. Fueron enviadas a Benjamin Smith Barton en Filadelfia. Dos de los especimenes estan atin en el Barton Herbarium. Estas son muy raras en la + id ] region actualmente. Hemos buscado los otros 24 especimenes pero no I éxito. Estos 24 i larian ciertamente a comprender la ecologia del area del Red River en 1806. INTRODUCTION The first part of the title of this paper comes from an herbarium specimen label written by Benjamin Smith Barton for a plant collected by Peter Custis in 1806. The plant comes from “the forgotten expedition’—the ill-fated and suppressed Thomas Jefferson-sponsored 1806 Expedition to the Red River. This expedition was to have been a southern counterpart to the now famous Lewis and Clark expedition (Figs. 1, 2). The specimen, housed in the Barton Herbarium at the Academy of Natural Sciences, Philadelphia, is one of three Peter Custis plant specimens there. It is one of two known surviving plants of 26 that were col- lected by Custis on the expedition. This paper has a twofold purpose. First, it makes the Freeman and Custis Red River Expedition of 1806 and its botanical contribution more familiar. The year 2006 will mark the bicentennial of the first botanical expedition into what is now northwestern Louisiana, southwestern Arkansas, and northeastern Texas (Anon 1807). Second, we call attention to an additional 24 collections made by Peter Custis in 1806 on the Red River that are not housed in the Barton Herbarium. These specimens, the identity of which is not known, might be crucial to under- SIDA 21(2): 963-975. 2004 964 BRIT.ORG/SIDA 21(2) Fic. 1.V . + Tree pe A \E llartad hy D Cc g Custis in 1806“450 miles up the Red Rver ina large prairie.” 1 in the Barton Herbari ; Academy of Natural Sciences, Philadelphia. Photograph made at the Old Courthouse Museum, Natchitoches, Louisiana. PETER CUSTIS ne ? hte . L hw. é a on Fic_2. Label inf Pa Pees» Geeerupaian oad A Di | J LAnls } AA Natchitoches, Louisiana standing floristic changes that have occurred in the Red River region over the past two centuries. THE EXPEDITION Louisiana was purchased from France in 1803. Thomas Jefferson sent probes into the newly acquired territory, the most notable of which was the Lewis and Clark expedition begun in 1804. Other expeditions soon followed: Dunbar and Hunter up the Ouachita, Pike up the Arkansas, and Freeman and Custis up the Red (Jackson 1981). This latter expedition was to skirt the Spanish/United States border, follow the Red River to its source, and test the Spanish resolve to prevent American western expansion. The Red River expedition, unlike other expeditions, was clearly politically 966 BRIT.ORG/SIDA 21(2) motivated but was disguised as scientific by the inclusion of a naturalist in the company. The purpose was to probe, none too delicately, the resistance of the Spanish beyond the as yet unsettled boundary between Louisiana and Mexico. Peter Custis, a young naturalist and medical student trained at that time by the most knowledgeable American botanist Benjamin Smith Barton, accompanied Thomas Freeman, a surveyor and astronomer and leader of the expedition, dur- ing the spring and summer of 1806. They were accompanied by 45 soldiers, interpreters, and guides on the Red River some 615 miles in search of its head- waters, then thought to be in the vicinity of Santa Fe. The expedition entered the Red River on May 2 and left Natchitoches, the northernmost post on the Red River,a month later. Soon after leaving Natchitoches, they encountered the lowest logjam of the Red River raft and had to divert their course to the bayous, “raftlakes,” and sloughs that surrounded the River (Figs. 3, 4, 5) (Humphreys 1971; Flores 1984; Triska 1984, Bagur 2001). They reentered the main River just above present day Shreveport to find themselves unobstructed ina land of prai- ries, cedar forests, river bottomlands, and pine-oak-hickory bluffs and uplands. The Spanish reaction was immediate and in force far beyond the modest Freeman party. The expedition was stopped on July 28 by Captain Don Fran- cisco Viana and a force of between 200 and 300 mounted soldiers and infantry at a point near what is today Spanish Bluff in Bowie County, Texas, and turned back (Fig. 6). The Red River expedition was a political failure and an embar- rassment to President Jefferson, who quietly suppressed it. Knowledge of the expedition was lost to history for almost 200 years. The Red River, north of Bowie County, remained virtually unexplored until the Marcy and McClellan Expe- dition of 1852, almost fifty years later (Marcy & McClellan 1854). PETER CUSTIS AND THE BOTANICAL ASPECT OF THE EXPEDITION Since Jefferson had received some criticism for not including a naturalist on the Lewis and Clark expedition, the Red River expedition was to be the first American-sponsored exploring expedition to include a trained naturalist. Pre- sumably Barton, at the University of Pennsylvania at the time, selected the list of candidates, and Freeman made the final choice. The nod was given to the 25- year old medical student, Peter Custis of Virginia, who had entered the Univer- sity of Pennsylvania as a medical student in 1804 and was about a year away from his degree. His background for the position consisted in having attended Barton's inclusive lectures on natural history. He had no field experience and was not considered a “naturalist” in any real sense of the word. On this expedi- tion no provision was made for proper scientific preparation to preserve and send specimens to Barton. Consequently, Custis attempted to identify and de- scribe in the field the plants he encountered either by familiarity or by refer- ence to the few floras he took with him, one of which probably was Linnaeus’s Systema Vegetabilium. As might be expected, there were names of many for- PETER CUSTIS 967 Fic. 3. Raft with vegetation growing on it on Red River, 1873. Archives Department Cc + 5 ee A bs Ch + Sh L L ? t reveport, Louisiana. R.B. Talf grap I eign species contained in the list of about 190 plants that Custis described (Mor- ton 1967; MacRoberts et al. 1997). Because Custis could not identify all of the plants to his satisfaction, he collected 26 and sent or took them to Philadelphia where two of them are known to survive in the Barton collection (Flores 1984). Custis listed these 26 plants separately in his report, most of which were collected in the extensive prairies that then existed north of present day Shreveport (Fig. 7). Both Freeman and Custis sent regular reports to President Jefferson, the Secretary of War Henry Dearborn, and to Professor Barton. These were drawn together, and Nicholas King was given the task of redacting the accounts. His redaction was published in a very limited edition in 1807, of which only about a dozen copies survive as stated in an extensive account of this publication by Flores (1984). The King redaction has many errors and some omissions, which Flores (1984), using primary material, has corrected. The redaction ina partial facsimile edition has been reprinted (Adams 1985). While some historians were aware of the expedition (e.g., Jackson 1981), it did not get full attention until Flores (1984) published his detailed account. In anthropological circles, the expedition was known because of its descriptions of Native Americans encountered above the Great Red River Raft (Morton 1967). 968 BRIT.ORG/SIDA 21(2) “4 ai 4 a i 1, a Py (Ee te ‘ 3 / ile J + ; : . 7 ¢ 4 ® { Arp Ae 2 O. ee ja ey 2¢ Pc S wa KEL 5 Biteute ¢ Fic. 4. Jacques Nicholas a 1764 map i gi isi ing Riv ver rin the1 8th cen ntury.A I ] y isi j H y j I port, Louisiana. In botanical, zoological, and ecological circles, the expedition remained un- known (e.g, McKelvey 1955; Ewan 1967; Sundell 1979; Lowery 1974a, 1974b). The expedition was unknown to Joseph Ewan (1952, 1969, 1988), Louisiana bota- nist and eminent historian of Bartonian Philadelphia. Morton (1967), using the MACROBERTS AND MACROBERTS, PETER CUSTIS, BOTANIST ON THE RED RIVER EXPEDITION 969 Q TAR vs sind mi ad a Th, ; Scale of miles Castor Li S, yi 2 = = ~ Alex dria | Cypress\Lake< Se 94° 93° Fic. 5. Map of the “raft lakes” along the Red River created by overflow caused by the Raft. H D t 488, 59th Congress, 1st Session. King redaction, published the first account of the plants observed by Custis. His aim was to interpret the Custis plant names and notes and provide modern identifications and nomenclature. Flores (1984) made the same attempt with the aid of Morton's (1967) paper. With the help of James Mears at the Academy of Natural Sciences in Philadelphia, Flores also located two of the Custis speci- mensand made photographic reproductions (Flores 1984:246-247). MacRoberts et al. (1997) reexamined the Custis accounts using, by that time, much better botanical information. Gilmore (2002) in his very important “Foundations of Southeastern Botany: An Annotated Bibliography of Southeastern American Botanical Explorers Prior to 1824” brought together the major works concern- ing the botanical aspects of the Freeman and Custis expedition. As Morton 970 BRIT.ORG/SIDA 21(2) * O* Natchitoches Natchez Fort Adams fe] nules Fic. 6. Freeman-Custi Red Ri Modified from Jackson (1981, Fig 10) (1967) has rightly pointed out, the botanical part of the narrative is quite infor- mative, being the first published information concerning the native plants of northwestern Louisiana and southwestern Arkansas, as well as the first descrip- tions of some of the plant communities, several of which have ceased to exist in the area, e.g., canebrakes, prairies, and cedar forests. THE SPECIMENS While it is possible to make informed guesses about the identity of the plants that Custis listed and described in his catalogues (Morton 1967; Flores 1984; MacRoberts et al. 1997), it is never possible to be certain without a specimen: thus the importance of the 26 plants collected by Custis that were forwarded to Barton. Fic.7. Facsimile copy of th , PETER CUSTIS aie 1, 2,& 4,. at the Coashatta. . 3. In prairies. No. 5, every where in suey. The Coashutia Indians make a decoc. tion with this which they drink at their green corn dance, previous to taking the black ‘drink, I. pukes them violently immediately after drink- heth ing it. ether it is the emetic property of - the plant, orth great quantity cf warm water which they drink that causes it to operate so svon fe aes Very plentiful, particularly on the de. clivites of the hills, . The poor people are said to use the root asa magne lor soap. The leaves are what the people of Campeachy make her eotce ee of. No. 8. Abundant in the prairies. The root is a Caddo remedy for the convulniene of chil- dren, It atall useful it is most probably in cases arising from worms, by its antheimintic apenas Is at the Coashatta No. 10. A speceis of Mimosa abundant in aco No. 11. Pepe in the prairies. No. 2 A climber. No. 13. On ae banks of the river. The leaves feathered with an odd one. No. 14. Abundent in prairies. No. 15. At the Coashatta Village. No. 16. On Lake anid No. 17, 18. In the prair No. 19. Lolypodium, every plwiicre abundant, No. 20 One of the most abundant vegeta~ bles in the country, found in Sate situation. No. 21. Plentiful in the prairic No. 22. A small shrub Be wing near the head of the great raft, . 23. Found in the prairies. No. 24. Supposed to be a species of Loni. cera. It grows near the ae village. No. 25. Very abundan No. 26. At the Ree village. Nore. Ne above ale refer to {peci- of the pla man-Cuctic let nf) | + " Hf baa ry a +} ui darti ftho £ r CUSTIS I 972 BRIT.ORG/SIDA 21(2) The two known surviving specimens, which come from above the Great Raft in the prairies along the Red River, are unusual for the present flora. One, Veronicastrum virginicum (L.) Farw, is very rare, if extant, in Louisiana today, and the other, Eustoma russellianum (Hook.) G. Don, has never been found in the area since. What were the other “prairie” specimens collected by Custis; and, if they were extant, would it be possible to envision the type of prairie where they occurred? Unfortunately, the prairies from which these plants came have long ceased to exist and are now known only from old maps and place names (Fig. 8). THE SEARCH On June 12 and 13, 2003, we made a thorough search of all of the specimens in the Barton and Lambert Herbaria at the Academy of Natural Sciences in Phila- delphia. The Barton Herbarium, kept separate at the Academy of Natural Sci- ences, consists of 1674 specimens originally housed at the American Philosophi- cal Society but moved to the Academy in 1898 (Pennell 1926). It consists of many specimens only a few of which were collected by Barton himself. The majority were collected by Frederick Pursh (Barnhart 1926). Barton’s collection was pre- viously searched for the same material by James Mears in the early 1980's with the same idea in mind; he found both of the specimens mentioned above (Flores 1984). Our search located three Peter Custis specimens, the same two from the Red River and one apparently from Virginia. These three specimens may have been all that were there in 1926 when both Pennell (1926) and Barnhart (1926) examined the collections or they would have done more than very briefly men- tion Custis in their papers. We also examined the Aylmer Bourke Lambert col- lection, which is the remains of Lambert's large collection purchased by Ed- ward Tuckerman and given to the Academy in 1842 (Miller 1970). It has a good deal of American material but no Custis collections. This leaves 24 specimens that have not been located. We do not know what happened to the other 24 Custis specimens. We only know that two of them did get to Philadelphia, then the hub of American botany (Pennell 1950). If two did, then probably all of them did. If that is so, then, where are they now? Itis unlikely they were discarded; collectors and curators prized their herbaria. But, the fragmentation and neglect of plant collections (includ- ing minimal labeling) in the early years of American botany is well known (Pennell 1950). For example, Frederick Pursh, Barton’s part-time curator and collector between 1805 and 1809, took various specimens from the Barton col- lection, including a significant portion of the Lewis and Clark collection, first to New York and then to London, and many plants simply disappeared (Ewan 1952; McCourt & Spamer 2003). After Barton’s death in 1815, his collections were warehoused for over 80 years at the American Philosophical Society be- fore they were entrusted to the Academy of Natural Sciences (Mears 1981). Por- MACROBERTS AND MACROBERTS, PETER CUSTIS, BOTANIST ON THE RED RIVER EXPEDITION 973 f4h L a5e 1 . 4 f-ndanD Fic. 8. Detail of Land Plat from 1837 of i iri i y ish, Louisi- ana. Clerk of Court Office, Caddo Parish, Louisiana. 974 BRIT.ORG/SIDA 21(2) tions of Pursh’s collections that he took to England and left to Lambert were returned to the Academy in the mid-19th century (Miller 1970; Pennell 1950; McCourt & Spamer 2003). But as any browser of the Index Herbariorum learns, specimens collected by famous botanists are scattered worldwide. We would like to locate the missing specimens to better interpret the Red River ecology prior to the massive changes that occurred to the area during the subsequent two centuries. ACKNOWLEDGMENTS James Macklin, Lucinda McDade, Alfred Ernest Schuyler, and Earle Spamer, at the Academy of Natural Sciences, Philadelphia, aided with this study. Amanda Crnkovic, Jolene Long, Cran Lucas, Steve Lynch, and Glenda Sharbono, Louisi- ana State University in Shreveport, aided with the figures. Laura Conerly, Domenica Carriere, and Glenda Sharbono, Archives Department, Noel Memo- rial Library, Louisiana State University, aided with the maps. The research was partly supported by a J. Bennett Johnston Science Foundation Grant that Vir- ginia Simons Lincove, Associate Vice Chancellor, Louisiana State University in Shreveport, was instrumental in securing for the research. Further support and assistance was provided by Gary Hanson and Patricia Peyton of the Red River Watershed Management Institute, Louisiana State University in Shreveport. Carolyn Harrington, Old Courthouse Museum, Natchitoches, Louisiana, allowed us to photograph the Custis specimens then on loan from the Academy of Natu- ral Sciences, Philadelphia. Ron Looney, Clerk of Court Office, Caddo Parish, gave permission to photograph land plats. Reviews by Eric Sundell, University of Arkansas, Monticello, and Ronald L. Stuckey, Ohio State University, Columbus, are greatly appreciated. REFERENCES Apams, G. 1985, An account of the Red River in Louisiana, drawn up from the returns of Messrs. Freeman and Custis to the War Office of the United States who explored the same in the year 1806. Ye Galleon Press, Fairfield, Washington. Anonymous. 1807 (redacted by Nicholas King). An account of the Red River, in Louisiana, drawn from the returns of Messrs. Freeman and Custis, to the War Office of the United States, who explored the same, in the year 1806. Baur, J.D. 2001. The history of navigation on Cypress Bayou and the Lakes. Univ. North Texas Press. Denton. Barnuart, J.H. 1926. Brief sketch of some collectors of specimens in the Barton Herbarium. Bartonia 9:35-42. Ewan, J. 1952. Frederick Pursh, 1774-1820, and his botanical associates. Proc. Amer. Philos. Soc. 96:599-628. Ewan, J.1967.A bibliography of Louisiana botany. Southwestern Louisiana J. 7:1-83. Ewan, J. 1969. A short history of botany in the United States. Hafner, New York. PETER CUSTIS 975 Ewan, J. 1988. Benjamin Smith Barton's influence on trans-Allegheny natural history. Bartonia 54:28-38. Fiores, D.L. 1984. Jefferson and Southwestern exploration: The Freeman and Custis ac- counts of the Red River Expedition of 1806. Univ. Oklahoma Press, Norman. Gitmore, R.W. 2002. Foundations of Southeastern botany: an annotated bibliography of Southeastern American botanical explorers prior to 1824. Castanea: Occasional Pap. Eastern Bot. 1:1-142. Humeureys, H. 1971. Photographic views of Red River raft, 1873. Louisiana History 12: 101-108. Jackson, D. 1981.Thomas Jefferson and the Stoney Mountains: Exploring the West from Monticello. Urbana. Univ. Ill. Press. Lowery, G.H. 1974a. The mammals of Louisiana and its adjacent waters. Louisiana State University Press, Baton Rouge. Lowery, G.H. 1974b. Louisiana birds. Louisiana State University Press, Baton Rouge. MacRoserts, D.T., B.R. MacRoserts, and M.H. MacRoserts. 1997.A floristic and ecological inter- pretation of the Freeman and Custis Red River Expedition of 1806. Bull. Mus. Life Sci., Louisiana State University 12:1—26. Marcy, R.B. and G.B. McCvettan.1854. Adventures on Red River. Report on the exploration of the headwaters of the Red River by Captain Randolph Marcy and Captain G.B. Mc- Clellan. Grant Foreman (ed.) 1937. Univ. Oklahoma Press, Norman. McCourt, R.M. and E.E. Spamer. 2003. The botanical legacy of Lewis and Clark: the most famous collection never heard of. Plant Sci. Bull. 49:1 26-130. McKewvey, S.D. 1955. Botanical exploration of the Trans-Mississippi West. The Arnold Arbo- retum of Harvard University. Jamaica Plain, Mass. Mears, J.A. 1981.Guide to the plant collectors represented in the Herbarium of the Acad- emy of Natural Sciences of Philadelphia. Proc. Acad. Nat. Sci. Philadelphia 133:141-165. Miter, H.S. 1970. The herbarium of Aylmer Bourke Lambert: notes on its acquisition, dis- persal, and present whereabouts. Taxon 19:489-656. Morton, C.V. 1967. Freeman and Custis’ account of the Red River Expedition of 1806, an overlooked publication of botanical interest. J. Arnold Arbor. 48:431-459. Pennett, FW. 1926. The elder Barton—his plant collection—and the mystery of his floras. Bartonia 9:1 7-34. PenNett, FW. 1950. Historic botanical collections of the American Philosophical Society and the Academy of Natural Sciences of Philadelphia. Proc. Amer. Philos. Soc. 94: 3 Zealaale Sunoett, E. 1979. A bibliography of Louisiana botany 1951-1975. Tulane Stud. Zool. Bot. 21:1-66. Taiska, F.J. 1984. Role of wood debris in modifying channel geomorphology and riparian areas of a large lowland river under pristine conditions: a historical case study. Verh. Internat. Verein. Limnol. 22:1876-1892. REFLECTIONS ON WILLIAM CHAMBERS COKER, PASSIONATE BOTANIST! Mary Coker Joslin liam R. Burk, John N. Couch eae ibrary olin pots c/o Wi Go No Oo S XO 1e) ay) ~ | Chapel Hill, North Carolina 27599-3280, U.S.A. ABSTRACT The Coker Arboretum at the University of North Carolina in Chapel Hill celebrated its centennial anniversary in the spring of 2003. The festivities began with a talk on botanist William C. Coker, which highlighted his humanistic traits. This paper briefly describes the events of the centennial celebration and provides the full text of the biographical presentation on Professor Coker. RESUMEN ss Coker Ar uate en ie Une a de Carolina del Norte en cna Hill celebré su centenario en I ] botanico William C. Coker, describe brevemente los eventos de la celebracion del 1 “ | eat =k oO centenario y aporta el le la presentacion biografica del Profesor Coker. INTRODUCTION William Chambers Coker (1872-1953) was a renowned botanist. His legacy in- cludes important contributions in botany as well as in horticulture and land- scaping. He arrived at the University of North Carolina at Chapel Hill in 1902 as the newly appointed associate professor of botany in the Department of Bi- ology. When the Department of Botany was established in 1908, Coker became its first chair. He set forward two major objectives: to foster a knowledge and appreciation of nature among people and to advance the discipline of botany. Among Coker’s lifelong scientific endeavors, mycology would occupy center stage and bring him international acclaim. Couch and Matthews (1954) de- scribed his life and scientific accomplishments. Coker was regionally known for his expertise in planting trees in Chapel Hill and in landscaping school grounds. Among his efforts to beautify the UNC campus, he began in 1903 to develop a five-acre arboretum on the northeastern edge of campus (Fig. 1). Planting the site with trees, shrubs, and vines native in North Carolina, Coker envisioned the collection asa living laboratory for botany students. Later, he added Asian species of woody plants. Today, nearly 575 spe- cies of trees and shrubs provide an inviting refuge and a living tribute to Coker. ‘Edited with an introduction by William R. Burk ohn N. Couch Biology Library, University of North Carolina, CB#3280 Coker Hall, Chapel Hill, NC 27599-3280) to whom correspondence should be sent. SIDA 21(2): 977-991. 2004 978 BRIT.ORG/SIDA 21(2) nn G. 1. View of the northern side of the William C. Coker Arboretum, University of North Caroli Chapel Hill, showing th . Wal It : A \Io] \ p pine (Pinus glabra Walter)—in center. (Courtesy of Brian Nalley, 14 April 2003). JOSLIN, 979 On the occasion of the centennial of the Coker Arboretum, the North Caro- lina Botanical Garden, the UNC Chapel Hill Libraries, and the Chapel Hill Museum hosted a series of events in the spring of 2003. From mid-March until mid-April, the UNC Libraries created several exhibits, all witha William Coker theme. The signature display was “WC. Coker: Legacy of a Lifelong Botanist” (Fig. 2).On 11-12 April, a two-day celebration took place in Chapel Hill. On the morning of the first day, the staff of the UNC Herbarium presented a reception and gave tours of their facility. Noon-time tours of the arboretum were followed by a birthday party that drew a large group of attendees who watched several distinguished guests plant a seedling of Crataegus marshallii, one of Coker’s favorites. Moving to the Morehead Planetarium, the group then listened to a number of distinguished speakers. Later that day, the focus was the Chapel Hill Museum’s exhibit “Coker Ar- boretum: The First Hundred Years.” There, Mary Coker Joslin (Coker’s niece) and illustrator Sandra Brooks Mathers signed copies of their book, Essays on William Chambers Coker, Passionate Botanist Joslin 2003). Although Dr. Joslin had previously presented a talk on 20 March 2003 to mark the publication of the book, the signing was the official release date. Following this, Mary Jane and Woodrow Burns and Florence and James Peacock graciously opened their homes and gardens (formerly part of Coker’s estate in Chapel Hill) with a re- ception. The first day of celebrations concluded with a formal dinner anda lec- ture. The next day, noted landscape gardener Chip Callaway entertained a large audience with the Evelyn McNeill Sims Native Plant Lecture entitled “North Carolina Natives—Gardeners and Their Gardens.” To close the celebration, nearly 150 people enjoyed a luncheon at the North Carolina Botanical Garden. The fes- tivities aptly commemorated not only the founding of the Coker Arboretum but also Coker’s tangible record of service to the university and to botany. The following commentary presents a humanistic view of William Cham- bers Coker, as related by Dr. Joslin at her 20 March talk. Filled with warmth, humor, and personal recollections of her uncle, the account provides a glimpse of Coker as a person. WILLIAM CHAMBERS COKER, PASSIONATE BOTANIST In these heartbreaking times of world violence and domestic threat to our en- vironment, it is good to celebrate the contributions of someone who passion- ately loved our world, the real natural world, loved to learn of it, teach of it, and preserve it. It is also good for us to smile and laugh a little, which we may do in these next few minutes. Also, let’s celebrate the first day of spring. Thank you for allowing me to talk about William Chambers Coker (Fig. 3), lately one of my favorite subjects. My obsession with this topic has become so obvious that when | launch into an anecdote about him at a family meal, | hear comments like, “Oh, Oh! Uncle Will again.” What a delight to have a captive au- 980 BRIT.ORG/SIDA 21(2) sisi Thervht tit eee ee CL Le Lead a i * Fic. 2.1 ic D Fant lth lIni | ray he iH rh | Hill, sah al Ul y' VA IH C Coker exhibit banners. (Courtesy of Brian Nalley, 14 April 2003). JOSLIN, 981 Fic. 3. William Chambers Coker, ca. 1940s. (Courtesy of the John N. Couch Biology Library, Botany Section, University of North Carolina, Chapel Hill). 982 BRIT.ORG/SIDA 21(2) dience this afternoon who will listen, | hope willingly, to a few stories about WC. Coker, some of which are not recorded in our recently published Essays on Will- iam Chambers Coker, Passionate Botanist Joslin 2003), where other tidbits may be found. Incidentally, I shall be referring to our subject in various ways accord- ing to context, using names such as William Chambers Coker, Will, Uncle Will, Professor Coker, Dr. Coker, ete. | hope this will be understandable and appropriate. We South Carolinians have the reputation of idealizing selected ancestors or family members. clearly remember an incident that lends some credence to that popular view. Some forty years ago, we were visiting my mother in Hartsville, South Carolina. Two of our young sons were bumbling about to the peril of various objects in their path. The younger child stopped, fixed his eyes on a somber portrait of my grandfather that hung over a fireplace. The artist had clothed him ina dark suit and posed him before an even darker background. His kindly face was bedecked with a flowing white beard. Our son grabbed his brother and asked, “Who is that?” Glancing up, the elder brother gave immedi- ately what was for him a satisfactory answer before they dashed on: “Why that’s God, | think.” True, my feelings toward my uncle, whom I knew and loved well, are tinted by admiration and affection. But maybe, even so, | can cast a realistic light on him. There was a real WC. Coker: very human like ourselves. We need to re- member this. His earthy humanity wasa great part of his charm. One hundred and thirty years after his birth, Professor Coker has become an historic figure at the University of North Carolina, where he left his mark. What | hope to do today is to humanize him for you. First, let’s look at some of his character traits and then attempt to breathe life into them with anecdotes from family oral tradition and from my own memory’s store. William C. Coker possessed a sense of humor that was quietly whimsical. He was brilliant, energetic, determined, tenacious, and even pas- sionate in his work. He was a stickler for accuracy and he expected the same tenacity and accuracy of his students. In his determination to answer a botani- cal question, he went directly for it. He could ignore completely any inconve- nience his floral quest might cause his colleagues, friends, or allies in the fam- ily. He took little note of the worldly rank of individuals on a societal scale. He was modest, even shy, not waiting for or even caring about recognition for his work. Rather, he directed his energies toward getting the work at hand done quickly and quietly and moving on to something else. He had no time or incli- nation to concentrate exclusively on the fields of botany in which he early be- came world-renowned, the Saprolegniaceae and the fleshy fungi. His interests were broader than these two fields. Nor did he have time regularly to attend meetings, where he would have periodically met with distinguished scholars and scientists. Though he had no children of his own, he cared deeply about his greater JOSLIN, 983 family: his parents, aunts, uncles, cousins, siblings, nieces, and nephews. He was himself a poet and a collector of his favorite poet, Walt Whitman. He deeply appreciated natural beauty. He was a conservationist. He was interested in literature and the visual arts. He had a good head for business. He was quietly generous. He had a succession of dogs and to each dog he gave his devoted affection. Several anecdotes, which you may not yet have heard or read may serve to illustrate some of these character traits. Some are in our Essays; some are not. First, I will mention his whimsical sense of humor. Those of us who knew him remember his crooked smile and his chuckle, but no uproarious laughter. He could recognize his own foibles. In recounting a rebellious incident of his early years, Uncle Will smiled at himself and at the same time paid tribute to the wisdom of his father. John Nathaniel Couch and Velma Matthews (1954), two of his former doctoral stu- dents, described one of their teacher’s reminiscences. As there was no public school at the time, older children in Will's family were usually sent to live fora time with relatives in Society Hill, South Carolina, a community fifteen miles away. There, they prepared for a university education at Saint David’s Academy, a good classical school incorporated in 1777 by the Saint David’s Society of the Pee Dee River’s Welsh Neck. Younger children were taught at home. There was for a time in Will’s home in Hartsville, a live-in teacher from Virginia. Will re- called that in his early teens, he took a dislike to this inescapable lady under his roof. His reaction to her was so strong that he refused to pay attention to her instruction or even to do his assignments. The lady reported this rebellion to his father, who summoned Will and calmly offered hima choice: “Will, of course ou may continue to pursue your studies respectfully, or, if you prefer, you may report at 6 A.M. tomorrow morning to our farm manager who needs your help in an area essential to our family’s wellbeing.” Will remained silent, but the next morning he reported for his studies and afterwards behaved himself. Uncle Will had a way of softening a major personal loss with a whimsical comment. One summer, Professor Coker led a company of colleagues and stu- dents on a plant collecting excursion in western North Carolina. At a stop on their return to Chapel Hill, someone took from the expeditionary car a large box containing about one-third of the priceless botanical collection of their foray. Rather than bemoan an irreparable loss to the University Herbarium, Uncle Will chose to defuse the tragedy by understatement and by whimsically as- suming the role of the disappointed robber. In 1923, Coker (1923) reported in the introduction of his book on clavarias that about one-third of the collection was lost through the activity of a misguided thief, who fancied he had found for himself something of value. Professor Coker was a stickler for accuracy when plants were concerned. He had little patience with romantic ignorance. When asked to upon 984 BRIT.ORG/SIDA 21(2) an article in the New York Times in praise of the Japanese honeysuckle that perfumes late-spring moonlit nights of the South, he waxed indignant, calling the invasive creeper a “first class pest, the worst pest since the chestnut blight.” Noted botanist M. L. Fernald would later call it “the yellow peril of the South.” The vine, Coker said, reaches out to throttle, not only shrubs but sometimes a considerable area of woodland, if left to its own devices. By this forceful reac- tion, he effectively dispelled for his interviewer the fanciful daydream of an urban journalist. What would he have said about kudzu? But that plant prob- ably would have no romantic champion. The native courtesy of Uncle Will was sometimes sorely strained by the inaccuracies of the botanically uninitiated. | remember a campus walk with Uncle Will on a lovely spring day. A talkative lady with our small party re- marked several times on “funguses.” Each time she used that expression Uncle Will, always a stickler for the correct use of botanical Latin, would mutter ina tone audible to most of us, if not to the lady in question, two clear syllables, es maetelea| ae William Coker was a modest, even shy individual, avoiding attention to himself if at all possible. Family legends illustrate that this character trait was evident early in his life. Two incidences ring of truth. When Will was a small boy, someone at the family breakfast table singled him out for praise for some accomplishment or other. There was no means of escape. A child then could not leave the family table unless excused by a par- ent. So Will picked up the large pancake on his plate to cover his face. Another delicious family story deals with his arrival back home after re- ceiving his Ph.D. degree from The Johns Hopkins University in Baltimore. That the son of a South Carolina farming region could and did earn a Ph.D. in 1901 was ararity,a source of pride for the community. Young Dr. Will returned to Hartsville, his home village, by train on the railroad spur built by local citizens to carry manutactured products and supplies to and from the town. As the train slowed to stop, Will spied down the track, to his horror, a considerable crowd of friends and relatives gathered to greet him. He even caught the strains of the local brass band playing “Hail the Conquering Hero Comes.” Though seemingly trapped, he quickly devised an escape. Grabbing his baggage, he descended from the train on the side away from the welcoming committee and temporarily vanished. William Coker tried to ignore or to make light of honors that inevitably came his way and which he was obliged to acknowledge. In 1943, he received a letter informing him of his election as honorary curator of the Charleston (South Carolina) Museum. In his response, he avoided the traditional formula: an ex- pression of gratitude, accompanied by a modest protest of unworthiness. He rather sidestepped the whole issue by diverting attention from himself to speci- mens in the museum's collection. He vividly recalled his youthful visits there with his father during the years when the family lived for four winters in La JOSLIN, ; 985 Charleston. During these Charleston years, from 1878 to 1882, Will was between five and nine years of age. The honoree simply remarked that ona recent visit to the museum he was struck by the remarkable fact that the expressions on the faces of the great deer and the giant buffalo had not changed at all in over 60 years Coker took little note of personal rank in the eyes of the world. He tackled his work in the chronological order of his commitment to it. People had to wait their turn. He had agreed to submit landscape plans for the factory buildings of a prominent industrialist in Durham, North Carolina. On 21 October 1915, the gentleman in question wrote asking him to run over and talk with him about his industrial park at his earliest convenience. Nine days later, the botanist-land- scaper answered the letter telling the gentleman that, though he was able to leave the university only at irregular intervals, he hoped to see him on Thurs- day or Saturday of the following week “as he was going to La Grange for Arbor pay and may have a few minutes in Durham either coming or going.” Later, asa volunteer extension agent for the university, he was to design the school grounds of this east Carolina farming community of La Grange. While pursuing any current subject of his botanical research and writing, Coker left no stone unturned to collect all possible data. His eagle eye was sharp todetecta plant of particular interest. He recruited or pressed into service friends who could help him gather the specimens he needed. While causing these vic- tims no little inconvenience, strangely enough, he often infected them with his own enthusiasm. On family trips, when Uncle Will was along, we could plan for no rigid schedule, as we expected an abrupt halt along the way if he spotted something special. I well remember one particular occasion. We had traveled some dis- tance across a seven-mile causeway over a marshy area where the driver was forbidden to stop or turn around. The eagle-eyed botanist called a halt and asked us to drive on and return for him presently, as he needed to get something on the edge of the swamp. He left the car, climbed over the guard rail and let him- self down into the rich vegetation. On our return, he had not yet finished his collection. We caught sight of him at some distance waving us on. Obliged to continue, we anxiously attempted to note some landmark, such as an extra tall cypress tree, in the pristine vegetation, where we had last seen him. After we had made at least two seven-mile crossings, Coker, pleased with his collection and unapologetic, was ready to climb back over the guard rail and return to Hartsville with his trophies. He then pressed them for eventual addition to the herbarium in old Davie Hall at UNC. During World War II, when Coker (1944) was preparing his classic article on “The Woody Smilaxes of the United States,” gasoline rationing denied him ready access to the South Carolina low country, where he needed to round up some key specimens he lacked. He leaned heavily on good friends there to col- 986 BRIT.ORG/SIDA 21(2) lect for him. Repeatedly and relentlessly, he wrote his friends G. Robert Lunz of the Charleston Museum and Frank Tarbox, master horticulturalist at Brookgreen Gardens on Pawley’s Island, South Carolina, to seek, find, dig up, press, and send to him different species of smilax in various stages of development. This was no small favor toask. The assignment involved constant vigilance for growth stages of the vines: male and female flowering and subsequent fruiting. It also meant innumerable sorties into a tangled woodland. Any woodsman knows that an encounter with some smilax thickets can be a very prickly experience. After the specimens were gathered, the collector had the time-consuming work of pressing the fresh plants, packing, and mailing them to Chapel Hill. These tasks, strangely enough, did not seem to turn Lunz and Tarbox against him—rather the contrary. Lunz was, during this period, an acting director of the Charleston Museum when Coker was made honorary curator and Tarbox shortly after his ordeal recommended Coker to Mr. Archer Huntington for appointment toa term as a trustee of Brookgreen Gardens, where he served for a period. The Venus flytrap was another passion of Uncle Will's. He assumed a pro- prietary role in protecting this indigenous North Carolina carnivorous plant, whose habitat centers around Wilmington. In November of 1920, he sent to the Georgetown Times, Georgetown, South Carolina, an ad written in the urgent style of someone in pursuit of a criminal at large (Coker 1920b): Wanted: Information about Venus’ Flytrap. One of the most remarkable plants in the world, callec Venus’ Flytrap because it catches and digests living insects, was reported from near Georgetown many years ago by [Stephen] Elliott in his ‘Sketch of the Botany of South Carolina and Georgia.’ No specimen from South Carolina is now known. Information in regard to the present occurrence of this plant in South Carolina is greatly desired. Address WC. Coker, Professor of Botany, University of North Carolina, Chapel Hill, N.C. Coker (1928) subsequently published an article on the “Distribution of Venus’s Fly Trap” in 1928, and he was always investigating reports of an extended range. There was something alluring about the passionate botanist’s single- minded devotion to plants, something akin to the medieval Holy Grail spirit. One somehow felt honored to be invited to become a partner in an important quest that could involve discomfort, risk, or danger. On 23 April 1938, Coker wrote his friend, the South Carolina poet Archibald Rutledge, thanking him for his recent hospitality. Coker enclosed for Rutledge one of his own poems. His letter expressed concern at the news that Rutledge had been sick, but quickly moved to the main question. The intrepid botanist greatly needed to know whether the Venus flytrap was indeed to be found in the “big ocean,” a local term for a botanically rich area near Rutledge’s home, and was counting on his friend to find it. Rutledge wrote to Coker in Chapel Hillof his initial failure. He had indeed searched for the flytrap one day, but instead of his finding the plant, he himself had come face to face with a diamond-back rattler. He bravely as- sured Coker, though, that he was returning to the area presently, as he knew the JOSLIN 987 plant was there. Though the poet had been sick, and though he had encoun- tered a deadly serpent in his effort to help his friend, the tenacious botanist kept his friend focused on the most important matter at hand, the location and collection of specimens of this plant. In responding to Rutledge four days after the rattlesnake letter, Coker dutifully requested a word from Rutledge on whether or not he had been “bit.” He ended the letter with the remark that he was looking forward with much interest to receiving the plants from Rutledge and that he greatly appreciated his continued interest. In the effort to verify information that he needed for an article, Uncle Will at times demanded the well-nigh impossible of Miss Alma Holland, later Mrs. C. Dale Beers, for numerous years his able research assistant and coauthor. In 1919, he heard that a species of mountain rhododendron had been found grow- ing naturally in sandhill country bordering North and South Carolina. Dr. Coker was then at his research desk in the New York Botanical Garden, where he worked from time to time during university vacations. Checking the report could not be delayed, as Coker’s article on rhododendron had been thought to be complete and was already in the hands of the publisher. Hearing that Miss Alma had not yet succeeded in verifying the plant’s location by correspondence, he urgently instructed her in a letter to visit the area personally. She was to take the train south to Rockingham, leaving Raleigh at 5 A.M. She was to find the plant, take specimens, and return the same day. Imagine the horror of that trip. The mere prospect of getting to Raleigh to take a train leaving at dawn, combined with the automobile transport of 1919, would have tempted a less dedicated botanist than Miss Alma simply to say, “I quit.” The reflected August heat in the sandhills can be hard to take. You have to keep dumping your shoes as the hot sand sifts in. But our heroine succeeded in her quest and the article was published without error. Alan Weakley, our able herbarium curator at UNC, discovered the evidence. He reported two herbarium specimens of Rhododen- dron minus labeled “August 3, 1919, Richmond County, N.C., Collector, Alma Holland,” thus providing ample proof of her strict obedience to orders and her success in accomplishing her mission. Weakley also checked the issue of the Journal of the Elisha Mitchell Scientific Society, where the article in question was published and found that Miss Holland's proof of the sandhills Rhododen- dron minus did indeed make the article before it was too late (Coker 1919). Our magnificent herbarium isa priceless archival research tool for solving all sorts of botanical mysteries. Uncle Will was delighted to obtain a new plant and even to read about the wondrous plants of distant climes, epochs, and cultures. Dr. Paul Titman, a re- tired professor in Chicago who had studied with Coker in the late 1930s and had returned to earn a Master’s degree in botany after his service in World War Il, recorded for me in December 1998 his recollections of his old professor, which included the following incident illustrative of Coker’s enthusiasm for plants. 988 BRIT.ORG/SIDA 21(2) Titman had described to Coker a particular kind of porcelain vine that grew in his grandmother's garden near Gastonia, North Carolina. Upon request, the stu- dent asked his grandmother to send a plant of this coveted vine for the Arbore- tum. Dr. Titman reminisced: “I still remember Miss Alma and Dr. Coker run- ning through the Arboretum, where | was doing something, | don’t know what, calling for me as if they were children at Christmas, to come quickly, that there was a package for me. And the gleam in his eye when that package was opened and he saw the anticipated vine was unforgettable.” There is no more effective pedagogical tool than a teacher’s own enthusiasm. Titman went on to earn his doctorate in botany at Harvard. He had planned to remain at Chapel Hill for his doctoral studies, but his professor advised him to seek instead, as he ex- pressed it, “the cross-pollination of Harvard.” Dr. Coker then pulled strings to assure that Titman was admitted to the doctoral program. I myself shall never forget Uncle Will’s excitement about the present Wil- liam and | gave to him when he took the place of my late father at our wedding in May of 1946. By a stroke of good fortune, I found for him a beautifully illus- trated book on an ancient Aztec herbal, reprinted under the title The Badianus Manuscript (Cruz, Badiano, and Trueblood] 1940). The volume is America’s ear- liest known book on herbal medicine. Ardently hoping that it would please him, | watched with bated breath as he unwrapped his gift. His unfeigned delight with our gift surpassed my wildest dreams. Ignoring the family hurly-burly swirling around us, he sat down to study it with total concentration. | believe that of all the gifts I have ever chosen for an adult, this book was the most ap- propriate and the most happily received. Uncle Will had real affection for some of his colleagues on the faculty, and good relations with most of them. Dr. William De Berniére MacNider opened letters to Coker with “Dear Old Man.” He closed them with “Affectionately,” or “Devotedly,” or “Bless your heart.” Dr. Dey of Romance Languages and Coker addressed each other as “Colonel,” with a complimentary close of “Very sin- cerely” or “Most sincerely yours.” The more formal “Yours truly” was reserved for business-like letters. Among his close friends were Collier Cobb, John Booker, Archibald Henderson, George Coffin Taylor, and our great librarian, Louis Round Wilson. With the renowned Professor Horace Williams, relations were somewhat less cordial. Professor Williams, it appears, was openly skeptical to his students of the laboratory method of the sciences and of the rigid demands for scholarly documentation in historical and literary research. “Oh those footnotes that Pro- fessor Greenlaw requires,” he might casually remark. These methods of research he considered far less valuable as tools for learning than the Socratic method, which Professor Williams used so successfully to arouse the curiosity and stimu- late the thinking of his students. Naturally this attitude caused some of his col- leagues in science, history, and literature to bristle. JOSLIN 989 The farms of Professors Williams and Coker were adjacent. The Orange County (North Carolina) Register of Deeds records that Coker bought this land from an H.H. Williams in 1906. I feel sure that this is the very gentleman under consideration here, though | have not definitively nailed it down. It would be natural for Professor Williams to retain a proprietary feeling for his former acre- age. The following letter from WC. Coker (1920a) to Professor Horace Williams reveals quite a bit about their relationship. I'll read it for you. February 21, 1920 Professor Horace Williams, Chapel Hill, N.C. Dear Professor Williams: I find that two of your sheep have been grazing every day on my wheat field for at least a month, - in fact, spend nearly all of their time on my place. l ask that you have your sheep removed to some other pasture, where they will not do constant damage to my crops. Yours truly, [W.C. Coker] When asked by the editor of the Chapel Hill Weekly,a local newspaper of Chapel Hill, for advice as to how to reach the ripe old age of seventy, one of Coker’s answers was: “Marry the right woman and manage always to have around a congenial dog.” Dogs were very important in his life. To each of an almost un- broken succession of canine friends, he gave his utter devotion. And it was am- ply returned. At their dinner table at “The Rocks,” Aunt Louise sat at right angles to him, on his left. When she turned her head to address someone at the other end of the table, one could observe Uncle Will surreptitiously passing a tidbit from his plate to Tinkerbell lying at his feet. Could this dinner-sharing be one reason why Uncle Will remained so pencil slim? One of his more colorful canine friends, Mickey, strongly resembled the famous Victrola dog of “His Master's Voice,” according to our own Laurie Rad- ford, one of Coker’s former graduate students and coauthor of the history of the University of North Carolina Herbarium (L. Radford and A. E. Radford, [2000)). Mickey was as much a one-man dog, as his master was a one-dog man. Uncle Will proudly wrote a niece that Mickey was the only dog he had ever owned who would actually bite. This statement reminds me of James Thurber’s de- lightful story, “The Dog that Bit People.” Thurber captures Uncle Will’s feeling toward Mickey, his beloved friend who could do no wrong. In Thurber’s tale, a one-dog lady replied to each victim’s complaint with the remark “Yes, he does love to bite but remember this, he never holds a grudge.” There is not enough time to tell other cherished tales and memories of my Uncle Will. I just hope that I have been able to humanize for you this very vital 990 BRIT.ORG/SIDA 21(2) person who walked the paths of our University for more than fifty years and did what he could to make our campus more beautiful, to interest us ina great variety of plants, and to encourage us to understand and conserve the wonders of our native fields and forests. |never think of Uncle Will without a frisson of delight and gratitude for his infectious love for our natural world. He, of course, was but one of the considerable number of extraordinarily gifted faculty mem- bers at Chapel Hill during the first half of the twentieth century. If William Coker were magically to return to Chapel Hill this spring, he could scarcely avoid seeing himself prominently displayed. He would doubtless immediately descend from the wrong side of the train, so to speak, and disappear to botanize incommunicado in an unknown North Carolina field or forest, or to study in some remote herbarium until all of this nonsense blows over. Please do forgive us, dear Uncle Will. We simply cannot help ourselves. You are irresistible. ACKNOWLEDGMENTS Many people, too numerous to cite, assisted us in preparing this paper. We par- ticularly thank Laura Cotterman, Rogers McVaugh, and Peter S. White for read- ing the entire manuscript and providing useful comments for changes. We also acknowledge the expert advice and assistance of Brian Nalley and Susan Whitfield with the creation and preparation of the illustrations. The Southern Historical Collection, University of North Carolina at Chapel Hill kindly pro- vided permission to quote two letters from the William Chambers Coker Papers. The North Carolina Botanical Garden funded this publication. REFERENCES Coxer, W.C. 1919. The distribution of Rhododendron catawbiense, with remarks on a new form. J. Elisha Mitchell Sci. Soc. 35:76-82 and plates 19-22. Coker, W.C. 1920a. Letter, dated 21 February 1920, to Horace Williams. (From Box 2, Folder 154, in the William Chambers Coker Papers # 3220, Southern Historical Collection, Louis Round Wilson Library, University of North Carolina at Chapel Hill). Coker, W.C. 1920b. Letter, dated 5 November 1920, to the editor of the Georgetown Times, including an enclosure (1 p.). (From Box 2, Folder 181, in the William Chambers Coker Papers # 3220, Southern Historical Collection, Louis Round Wilson Library, University of North Carolina at Chapel Hill). Coker, W.C. 1923. The Clavarias of the United States and Canada. University of North Caro- lina Press, Chapel Hill. Coker, W.C. 1928. The distribution of Venus's fly trap (Dionaea muscipula).J. Elisha Mitchell Sci. Soc. 43:221-228 and plate 33. Coxer, W.C. 1944. The woody smilaxes of the United States. J. Elisha Mitchell Sci.Soc.60:27- 69 and plates 9-39. JOSLIN, 991 Couch, J.N.and V.D. MattHews. 1954. William Chambers Coker. Mycologia 46:372-383. [Cruz, M. be LA, J. BADIANO, and E.W. Emmert (Trues.oop)]. 1940. The Badianus manuscript (Co- dex Barberini, Latin 241).Vatican Library.An Aztec herbal of 1552. Introduction, transla- tion, and annotations by E.W. Emmart. With a foreword by H. E. Sigerist. The Johns Hopkins Press, Baltimore. Josuin, M.C. 2003. Essays on William Chambers Coker, passionate botanist. With drawings by S.B. Mathers. University of North Carolina at Chapel Hill Library and Botanical Gar- den Foundation, Inc., Chapel Hill. Raprorb, L.S. and A.E. Raprorb. [2000]. The herbarium of The University of North Carolina: 1908-2000; history and perspective. With contributions from PS. White, J.R. Massey, and J.F. Matthews. The Herbarium (NCU), The University of North Carolina at Chapel Hill. ALLOZYME DIVERSITY IN ELEUTHEROCOCCUS SENTICOSUS AND E. BRACHYPUS (ARALIACEAE) FROM CHINA AND ITS IMPLICATION FOR CONSERVATION Shi-Liang Zhou Jun Wen! Laboratory of Systematic & Department of Botan Evolutionary Botany Field Museum of Natural History Institute ey 1400 S. Lake Shore Dr. Chinese Academy of Sciences Chicago, Illinois 60605, U.S.A. ang Xiangshan, Beijing 100093, CHIN Laboratory of Systematic & Evolutionary slzhou@ibcas.ac.cn Institute of Botany, The Chinese Academy of Scie iences Beijing 100093, CHINA; wen@fieldmuseum.org De-Yuan Hong Laboratory of Systematic & Evolutionary Botany Institute of Botany The Chinese Academy of Sciences Xiangshan, Beijing 100093, CHINA ABSTRACT The widespread Eleut! CR et Maxim.) Maxim. is threatened because of over- Lr r papveccne of its root ae for medicin: al uses. The geographically restricted E. brachypus becomes Loess Plateau of China. To facilitate the development of conser vation strategies, genetic diversity of both species was measured at 26 isozyme loci (12 enzymes). Eleutherococcus brachypus had a low proportion of polymorphic loci (P.=19.2%; P =13.1%) and low genetic diversity (H, =0.063; A =0.031). The genetic variability of the widespread E. senticosus was higher (P.=26.9%; P =20.76%, H,=0.094; H,,=0.059) than that of the restricted E. brachypus. Popula- tion differentiation of E. ee. -0.531) was greater than that of E. senticosus (G,=0.383). Clus- ter analysis showed that populations of E. senticosus in Northeast or North China are genetically closer within each region than between une two Bens BOpuIAnS ns o : Sad in NenEne st China have a higher level of g poy l L 1 ] Dp! } } 9 ae eS Tle | £ priority. Population tana pile popu- lations of the species cil be protected to maximize its genetic haat CHINESE ABSTRACT kl) fin (Eleutherococcus senticosus (Rupr. et Maxim.) Maxim. ) is F BABA AR Ab %, HAS ah EA a (ER oh SS eR; Donn el V2 ti BS: J FF TAH SRM IAT ANTENA AN PO FRA HTT BY ND AP EE 2 PE PEGE a AE 57 eee JE AEISEE in, ARLOT HN: oP ay F i te hin : Ing YH. naib iekrtie stu. Hg eae Kove sch v8 RK UPA REO. 094, RACE YU Ae a EOE 00. 059. AW HN eet St, FL (G,.=0. aD REMC, =0. 383). 32H AKA, Flite Ade lCorrespondina author v SIDA 21(2): 993-1007. 2004 994 BRIT.ORG/SIDA 21(2) Teor LUN Aide se aa . kak A fe AG HX A TL arate be el Ht De yy Jae aE hf DLA CRP OME. AA EL ee Te RPE HS i 7 uses i aes ie iy ae AAR i ROE FAY Ta DE AR RE PE BE EH FE RPROA >, witR Se REPEAIR, OOCCRIP TAT ESATA LEE 2 ae SPEER. a INTRODUCTION Understanding of the levels of genetic diversity is important in designing con- servation strategies for rare and endangered species (Holsinger & Gottlieb 1991; Hamrick & Godt 1996; Rieseberg & Swensen 1996; Newton et al. 1999; Gitzendanner & Soltis 2001). It is estimated that approximately 3000-4000 species (15%) of the vascular plants in China are endangered or threatened (Fu & Jin 1992). Since the 1970's, major efforts have been made to establish nature reserves to protect rare and endangered species and their communities. At present, few studies have been conducted to examine the genetic diversity of endangered species in China. Genetic data are needed from model plant groups in China to help design conservation efforts when resources are limited. Eleutherococcus senticosus [= Acanthopanax senticosus (Rupr. et Maxim.) Harms] (Araliaceae) distributed in China, North Korea and Far East of Russia, is commonly known as Siberian ginseng (Soejarto & Farnsworth 1978; Farnsworth etal. 1985; Duke 1989) and is considered to be of high medicinal value. It has been used for hundreds of years in China as a tonic (Hu 1980). Studies have been con- ducted (e.g, Xu et al. 1983; Zhao et al. 1990, 1991, 1993) to analyze its chemical constituents. Polysaccharides extracted from its leaves and roots have been re- ported to inhibit tumor cell proliferation (Xie et al. 1989; Liang et al. 1994) and to have antiviral activity (Glatthaar-Saalmiiller et al. 2001). Glucosides (such as liriodendrin) extracted from root or stem bark have also been reported to have an effect similar to those of that of ginseng (Panax spp.) (Slacanin et al. 1991), or as an adaptogen that exerts effects on both sick and healthy people by “correcting” any dysfunctions with no or few side effects (Davydov & Krikorian 2000). In- dustries have been developed to manufacture herbal products using E. senticosus. Herbal and pharmaceutical producers have been using material of the species collected from natural habitats, which has led to rapid destruction of natural populations. Although the species has a wide distribution in China, ranging from the North (Shanxi and Hebei provinces) to the Northeast (Liaoning, Jilin and Heilongjiang provinces) (Fig. L), itis now listed in the “China Plant Red Data Book” as a vulnerable species (Fu & Jin 1992) for its economical importance. Eleutherococcus brachypus (Harms) Nakai is a rare endemic species re- stricted to the Loess Plateau (southeastern Gansu and central Shaanxi prov- inces) of Northwest China (Fig. 1), which is heavily populated and has highly fragmented vegetation. Eleutherococcus brachypus isa clonal species with small populations (Wang et al., 1997), but has suffered from habitat loss, and is thus rare. Seeds of E. brachypus are usually not well developed, requiring 1.5 years of — ZHOU ET AL PpAcUYDIIC 995 after-ripening (Tian et al. 1998). Wang et al. (1997) reported that insect visita- tion was necessary for seed set. Yan et al. (1997) investigated genetic diversity of this species using three populations from Yanan City, Shaanxi Province, and reported that the percentages of polymorphic RAPD bands were relatively low, 5.4%, 18.5%, and 27.7%, respectively. Both Eleutherococcus senticosus and E. brachypus are shrubs with similar ecological preferences of sunny habitats, but different in geographic ranges. Eleutherococcus senticosus has a wide distribution, whereas E. brachypus is a rare endemic confined to the Loess Plateau of Northwest China (Fig. 1). This paper assesses the genetic diversity of these two congeneric species with allozyme markers using starch gel electrophoresis and discusses the implica- tions for conservation. MATERIALS AND METHODS Sampling Five populations of E. senticosus from five provinces in China and five popula- tions of E. brachypus from North Shaanxi to South Gansu were sampled, cover- ing nearly the full range of distribution of both species (Fig. 1, Table 1). Three populations of three other congeners: E. giraldii (Harms) Nakai, E. gracilistylus (W.W. Smith) S.Y. Hu, and E. sessiliflorus (Rupr. et Maxim.) S.Y. Hu (Table 1) were included in the UPGMA cluster analysis as comparisons. One to two year old twigs were collected in the spring from about 20 indi- viduals in each population. Populations of E. brachypus were small and the boundaries were easily determined. Samples were collected throughout the populations. Populations of E. senticosus were usually large with hundreds of individuals, and samples were collected at an interval of at lest 50 meters to minimize the possibility of collecting two samples froma single clone. The twigs were then kept in sealed moist plastic bags. Electrophoresis In the laboratory, the cut surface of each twig was immersed in water and incu- bated in a humid environment. When the first leaf appeared, the bud was re- moved and ground on ice with grinding buffer after removing bud scales. The grinding buffer (Tris-malate grinding buffer-PVP solution) was prepared fol- lowing Soltis et al. (1983) with the substitution of sodium bisulfite for sodium metabisulfite. DMSO was added to the mixture to a final concentration of 10% before adjusting pH to 7.5. The enzyme solution was absorbed onto wicks, which were frozen (-80°C) until electrophoresis. Electrophoresis was carried out on horizontal starch (Sigma cat no. $4501) gels at 4°C. Four buffer systems (elec- trode buffer / gel buffer) were used to assayl2 enzymes (Wang 1998): (1) 0.4M Citric acid trisodium salt (pH adjusted to 7.0 using 10M HCl) / 0.02M HistidineeHCl (pH adjusted to 7.0 using 1.0M NaOH) for aconitate hydratase 996 BRIT.ORG/SIDA 21(2) ) .¢] A Ay ~ -* 85x 2 ¢ S-HBs ~ aa, e rt) B- YA OB-PLeOPB-HL a e e@ o e-ts BXY* Yellow River a C e Yangtze River 30 Fic. 1.D f populations sampled. Loh accth i I (aa\ Ic es {a \infhi fa\ 7 | a Pa | Fae Ree 9 \ } (ACO, F.C. 4.2.1.3), fructose-bisphosphate aldolase (FBA, E. C. 4.12.13) and glyc- eraldehydes-3-phosphate dehydrogenase (G3PD, E. C. 1.2.1.12); (2) 0.3M boric acid (pH adjusted to 8.6 using NaOH) / 0.015M Tris (pH adjusted to 7.8 using citric acid) for an linope ptidase (AMP. E.C. 3.4.11.D), hexokinase (HEX, E.C. 2.7.1.1), phosphoglucomutase (PGM, E. C. 5.4.2.2) and triose-phosphate isomerase (TPI, E. C. 5.3.1.1; G3) 0.374M boric acid (pH adjusted to 8.0 using LiOH) / 0.033M Tris+0.005M citric acid+0.004M LiOH +0.030M boric acid for aspartate ami- notransferase (AAT, E. C. 2.6.1.1), alcohol dehydrogenase (ADH, E. C. LLL) and ZHOU ET AL., BRACH 997 Taste 1. Population localities, symbols and voucher specimens of Eleutherococcus brachypus, E. senticosus and close congeners. Locality Symbol Voucher (PE) Eleutherococcus senticosus (Rupr. et Maxim.) Maxim. Mao’ershan, Shangzhi County, soe Prov S-HU Zhou 009 Changbaishan Nature Reserve of CAS, A eat ae Province S-JL Zhou 010 Baishilazi Nature Reserve, Kuandian ae Liaoning Province S-LN Zhou O11 Mount Wutaishan, Shanxi Province S-SX Zhou 013 East Lingshan, Hebei S-HB Zhou 014 Eleutherococcus rie (Harms) Nakai Nanniwang, Yan’an City, Shaanxi Province B-YA Zhu 950004 Nanshan, Huanglong County, Shaanxi Province B-HL Zhou 007 Yangjiadian, Xunyi County, Shaanxi Province B-XY Zhou 003 Mount Kongtongshan, Pingliang City, Gansu Province B-PL Zhu 95016 Caijiashan, Lu’ergou, Tianshui City, Gansu Province B-TS Zhou 002 sy cere dae alii (W.W.Sm.) S.Y. Hu nkou, Kanxian County, Gansu Province Zhou 006 aes: giraldii (Harms) Nakai Mount Lianhuashan, Kanle County, Gansu Province Zhou 005 Eleutherococcus sessiliflorus (Rupr. et Maxim.) S.-Y. Hu Baishilazi Nature Reserve, Kuandian County, Liaoning Province Zhou O12 NADH-diaphorase (DIA, E. C. 1.6.2.2); and (4) 0.04M citric acid [pH adjusted to 7.5 using N-(3-aminopropyl)-morpholine] / 1:19 eaaias of seco oan oi isocitrate dehydrogenase (IDH, E.C. 1.1.1.42) and shi E.C. 1.11.25). Enzymes were visualized using stains in agar overlays except * AAT and AMP which were stained in buffer solutions. Data analysis Stained gels were photographed and the banding patterns were then drawn. The alleles at each locus were designated with letters a, b, and c, from the long- est migration distance to the shortest. The resulting genetic data (genotypes) were analyzed with Biosys-1 (Swofford & Selander 1989) for each species. For each population, the allele frequencies, mean number of alleles per locus, per- centage of polymorphic loci, heterozygosity observed and expected under Hardy-Weinberg equilibrium, F-statistics and unbiased genetic similarities/ distances (Nei 1978) were computed. All the populations were analyzed to gen- erate a dendrogram using UPGMA RESULTS Twenty-six putative loci from 12 enzyme systems were interpretable on the ba- sis of simple Mendelian genetics (Table 2). Nineteen loci exhibited polymor- phisms in one or both species. The frequency of one allele often dominated over the others in a given population (Table 2). 998 BRIT.ORG/SIDA 21(2 Genetic diversity of E. senticosus There were 13 polymorpic loci in E. senticosus (Table 2). The two regions of ac- tivity for AAT were designated as Aat-land Aat-2. Aat-1 had 2 alleles, the rare one Aat-lb and the common one Aat-Ic. Aat-2 had 3 alleles, Aat-2a, Aat-2b, and Aat-2c with Aat-2b being the common one. Only one locus was detected for ADH with 2 alleles, Adh-a and Adh-b, which were present in heterozygos- ity with low frequencies. Both Amp-1 and Amp-2 had 2 alleles, with the allele frequencies varying among populations. Dia-I exhibited polymorphism only in the S-JL population. Two alleles were detected on G3pd-2 and Hex-l, respec- tively. The rare alleles on Idh-2, Skd-1, Skd-2, Skd-3, Tpi-l,and Tpi-2 were unique to this species. Locus duplications were observed on PGM and TPL Eleutherococcus senticosus maintained a higher level of genetic diversity. The percentages of polymorphic loci ranged from 11.5% to 30.8% with a mean of 26.9% (Table 3). The mean number of alleles averaged over populations was 1.26, and the expected heterozygosity under Hardy-Weinberg equilibrium av- eraged over populations was 0.059. The population S-JL from Jilin Province (A=1.3; P=30.8%; He=0.073) and the population S-LN from Liaoning Province (A=1.3; P=23.1%; He=0.077) exhibited the highest genetic diversity. The south- west peripheral population S-SX from Shanxi Province had the lowest (A=L.2; P=11.5%; He=0.025). At the species level, the mean number of alleles per locus was 1.7,and 26.9% of the loci were polymorphic. The expected heterozygosity was 0.094. The ge- netic diversity was maintained at 13 polymorphic loci, especially at Amp-1, Amp-2, G3pd-2, Idh-2, Skd-3 and Tpi-1(Ht>0.2, Table 4+). Among these loci only Amp-l and Idh-2 contributed more to interpopulational than to intrapopulational variation (Gs>0.5). Of the mean total genetic diversity (H;=0.094), only 38.3% was maintained within populations (H.=0.058, Dst=0.0306). Genetic diversity of E. brachypus There were 10 polymorphic loci detected in five populations of this species (Table 2). Aat-la was unique to B-XY population. The alleles of Aat-2a and Aat- 2c were infrequent. Aco-la was common in B-XY population but rare or absent in the other populations. The unique allele Adh-c was present in heterozygos- ity with low frequencies. Did-2b and Dia-2c were characteristic of this species, which exhibited intrapopulational variation. Nearly complete divergence of alternative allele fixation was found at the three loci of IDH between E. brachypus and E. senticosus. Locus duplication was not observed at PGM. In- stead, both loci Pem-l and Pgm-2 were polymorphic. The genetic diversity of E. brachypusat the populational level was relatively low compared to that of E. senticosus (Table 3). The percentage of polymorphic loci varied from 3.8% to 23.1% with a mean of 13.1%. The mean number of alleles ZHOU ET AL 1D E. BRACHYPUS CONSERVATION 999 Taste 2. Allele frequencies in 10 populations of Eleutherococcus brachypus and E. senticosus (N indi- cates sample size). Populations Locus Allele Eleutherococcus brachypus Eleutherococcus senticosus BTS BPL BXY B-YA BHL S-HU S-JL S-LN S-SX_—S-BJ ] 20 (N) 20 19 15 14 20 9 20 20 20 Aat-l a = - 0.033 - - = - - - ~ b - - - - - - 0.200 0175 - ~ la 1.000 1.000 0.967 1.000 1.000 1.000 0.800 0825 1.000 1.000 Aat-2 a = - = 0.036 - 0.132 - - - - b 0.950 1.000 1.000 0.964 0.900 0868 0.975 1.000 1.000 1.000 C 0.050 - = - 0.100 - 0.025 = - - Aco-l a = - 1.000 0.071 - - - - b 1.000 1.000 - 0.929 1.000 1.000 1.000 1.000 1.000 1.000 Aco-2 a 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 Adh a 0,050 0026 - O07) 0025. = 0.100 - - 0.075 b 0950 0.947 1.000 0857 0.975 1.000 0.900 1.000 1.000 0.925 G 0.026 - 0.071 - = - - - a b = Amp-2 a - - - 0. b 1.000 1.000 1.000 1.000 1.000 0.553 0.700 1.000 0.975 1.000 Dia-1 a 1.000 1.000 1.000 1.000 1.000 1.000 0.925 1,000 1.000 1.000 Diy = = 7 = 7 0075: = - - Dia-2 a 0850 0.789 0.933 0286 1.000 1.000 1.000 1.000 1.000 1.000 BE OO75. 0105s ]0.0338e 205545 = - - - - G 0.075 0.105 0.033 0357 - - - = - - Fboa a 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 G3pd-1 a 1000 1.000 1,000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 G3pd-2 a 0.150 1.000 1.000 - 0.050 - - 0.300 - 0.350. b 0.850 - 1.000 0.950 1.000 1.000 0.700 1.000 0.650 Hex-1 a 1.000 1.000 1.000 1.000 0.950 1.000 0.950 0.950 0.975 1.000 bo = - - 0.050 - 0.050 0.050 0025 - Hex-2 a 1000 1,000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 Idh-1 a - = - - 1.000 1.000 1.000 1.000 1.000 b 1.000 1.000 1.000 1.000 1.000 - = - - ldh-2 a - - - - - 1.000 1.000 1.000 0.050 0.175 b 1.000 1.000 1.000 1.000 1.000 - - 0.950 0.800 C - = - = - - = - - 0.025 Idh-3, a ~ - - - - 1.000 1.000 1.000 1.000 1.000 b 1.000 1.000 1.000 1.000 1.000 - - - ~ Pgm-1 a 0.300 = 7 0.100 - = = = = b 0700 0947 1.000 1.000 0750 1.000 1.000 1.000 1.000 1.000 a - 0.053 - - 0.150 = - - - ~ Pgm-2 a 0.950 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1,000 1.000 b 0.050 - - - - - - - - - 1000 BRIT.ORG/SIDA 21(2) Taste 2. continued Populations Locus Allele Eleutherococcus brachypus Eleutherococcus senticosus 1.000 1.000 1.000 1.000 1.000 ad 1.000 000 =1.000 1.000 7 053 1.000 1,000 1.000 1.000 1.000 os 1.000 1.000 1.000 1.000 = = = = 0.025 1.000 1.000 1,000 1.000 1.000 : = 1.000 1.000 0.900 0.500 = = - - 0.075 Tpi-1 - - - - - = is) 025 b 1.000 1.000 1.000 1.000 1.000 0684 0950 0400 1.000 0,950 Cc = - - — _ 0.316 0.050 0.300 - 0.025 Tpi-? a 1.000 1.000 1.000 1.000 1.000 1.000 0.900 0.950 1.000 1.000 b = - - 7 = = 0.100 0.05 - Tpi-3 a 1.000 1.000 1,000 1.000 1.000 1.000 1.000 1.000 1.000 000 Tpi-4 a 1,000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 1.000 000 Taste 3. Genetic variability at 26 loci in the populati f Eleutl bract 1 F. senticosus (standard errors in parentheses). N indicates the sample size per os is indicates the mean num ber of alleles per locus; P indicates the percentage of polymorphic loci*; Ho indicates the observed heterozygosity; and He indicates the expected heterozygosity under nee Weiberg equilibrium.** Population N A P Hy H. Eleutherococcus brachypus B-TS 20 1.3(0.1) 23.1 0.013(0.008) 0.048(0.021) B-PL 19 1.2(0.1) 11.5 0.014(0.009) 0.024(0.015) B-XY 15 1.1(0.1) 3.8 0.005(0.004) 0.008(0.006) B-YA 14 1.2(0.1) 11.5 0.047(0.029) 0.045(0.028) B-HL 20 1.2(0.1) 15.4 0.006(0.004) 0.032(0.018) an 12 13.06 0.017 0.031 Species level 88 1.5(0.1) 19.2 0.016(0.009) 0.063(0.026) Eleutherococcus senticosus S-HLJ 19 1.2(0.1) 19.2 ae 017) 0.059(0.027) S-JL 20 1.3(0.1) 30.8 12(0.007) 0.073(0.025) S-LN 20 1.3(0.1) 23.1 oe 027) 0.077(0.034) S-SX 20 1.2(0.1) 11.5 0.019(0.011) 0.025(0.013) S-BJ 20 1.3(0.1) 19.2 0.012(0.006) a ean 1.3 20.8 0.023 059 Species level 99 1.7(0.1) 26.9 0.023(0.007) i es Note. * A locus is considered polymorphic if the frequency of the most common allele does not exceed ** Unbiased estimate (see Nei 1978) ZHOU ET AL., BRACHYPUS CONSERVATION 1001 Taste 4. Genetic diversity across populations of £. brachypus and E. senticosus. H, indicates the total gene diversity; H, indicates gene diversity within populations; D,, indicates the gene diversity be- tween populations; and G.,.is the ratio of D../H, Locus Ht H, Ds Gt Eleutherococcus brachypus 0. Aat-1 0.013 013 0.000 0.000 Aat-2 0.072 0.069 0.003 0.042 Aco-] 0.337 0.027 0.310 0.920 Adh 0.105 0.101 0.004 0.038 Amp-1 0.010 0.010 0.000 0.000 Dia-2 0.378 0.282 0.096 0.254 G3pd-2 0.493 0.070 0.423 0.858 ex-] 0.020 0.019 0.001 0.050 Pgm-1 0.218 0.185 0.033 0.151 Pqm-2 0.020 0.019 0.001 0.050 Mean 0.064 0.030 0.034 0.531 Eleutherococcus senticosus Aat-1 0.139 122 O17 0.122 Aat-2 0.061 0.055 0.006 0.098 0.068 0.064 0.004 0.059 Amp-1 0.466 0.215 0.251 0.539 Amp-2 0.261 0.192 0.069 0.264 Dia-1 0.030 0.028 0.002 0.067 G3pd-? 0.226 0.175 0.051 0.226 Hex-1 0.049 0.048 0.001 0.020 Idh-2 0.461 0.085 0.376 0.816 Skd-2 0.021 0.020 0.001 0.048 Skd-3 0.254 182 0.072 0.283 Tpi-1 0.342 0.257 0.085 0.249 Tpi-2 0.058 0.055 0.003 0.052 Mean 0.094 0.058 0.036 0.383 averaged over populations was 1.2, and the expected heterozygosity under Hardy-Weinberg equilibrium averaged over populations was 0.031. The popu- lation B-TS from Tianshui, Gansu Province, showed the highest genetic diver- sity (A=1.3; P=23.1%; He=0.048), while the population B-XY from Xunyi, Shaanxi Province, showed the lowest diversity (A=1.1; P=3.8%; H,-=0.008). Eleutherococcus brachypus exhibited an average of 1.5alleles per locus, and 19.2% of loci were polymorphic (Tables 3 & +). The expected heterozygosity under Hardy-Weinberg equilibrium was 0.063. The genetic diversity was main- tained at 10 polymorphic loci, notably at Aco-1, Dia-2, G3pd-2 and Pgm-1. The mean total gene diversity (H;) was 0.064. Nearly half of the total genetic diver- sity occurred within populations (Hs=0.030). Among the loci with high H; val- ues, Dia-2 and Pgm-1 showed most genetic variation within populations (Gs 1002 BRIT.ORG/SIDA 21(2) was 0.254 and 0.151, respectively), while Aco-Il(Gs=0.858) and G3pd-2 (Gs=0.920) showed genetic variation primarily among populations. Relationships among populations Genetic identity between populations within each species was higher than 0.9, and the genetic distances were lower than 0.09 (Table 5). The average distances between the populations were 0.043 (+0.028) and 0.047 (+0.019) for E. brachypus and E. senticosus, respectively. The dendrogram generated from ge- netic identity data using UPGMA (Fig. 2) showed that populations of E. senticosus from Northeast China were more similar to one another than with those from North China. The population B-XY of E. brachypus differentiated most significantly from other populations of the species. DISCUSSION Overall genetic diversity within species The levels of allozyme variation in E. senticosus and E. brachypus were rela- tively low at both species and populational levels in comparison with those of other species with similar attributes (Hamrick & Godt 1990), For example, Ps (26.9%) and Hes (0.094) of E. senticosus were about half the averages (64.7% and 0.177, respectively) of woody plants at the species level. Hamrick & Godt (1996) reported an average within species heterozygosity (Hes) for widespread and endemic species of 0.202 and 0.096, respectively. The Hes for the widespread E. senticosus was 0.094 and that of the restricted endemic species E. brachypus was 0.063. The allozyme heterozygosity within each of the two Eleutherococcus is thus considerably lower than the averages reported in the literature. The percentages of polymorphic allozyme loci reported in this study cor- respond well to DNA-RAPD polymorphism in E. brachypus (3.8-23.1% vs 5.4- 27.2%, Yan et al. 1997), but differ sharply from those of E. senticosus (11.5-30.8% vs 91.3-97.6%, Dai et al. 1998). However, the report of highly polymorphic RAPD bands in E. senticosus likely represents an overestimate because amplification failure from some individuals was interpreted as a lack of bands by Dai et al. (1998). Our allozyme data seem to be more reliable in presenting the overall genetic diversity of the species investigated. Levels of genetic diversity between species There is significant disparity between E. brachypus and E. senticosus in overall genetic diversity (Tables 3 & 4), with E. senticosus maintaining a higher level of genetic diversity than E. brachypus. These two species appears to be closely re- lated, but not sister taxa. They differ in several characters that may influence genetic diversity. First, they have highly different distributional ranges. Eleutherococcus senticosus is widespread across several thousand kilometers, from North to Northeast China and adjacent countries (North Korea and Far East of Russia). By contrast, E. brachypus is restricted to the Loess Plateau of ZHOU ET Al ANTE RDACHUVDIIC 1003 Taste 5. Matrix of Nei’s (1978) unbiased genetic identity (below diagonal) and distance (above di- agonal) between populations of £. brachypus and E. senticosus. B-TS B-PL B-XY B-YA B-HL S-HLJ S-JL S-LN S-SX 5-BJ B-TS 0.032 0.074 0.013 0002 0199 0169 0154 0.090 0.147 B-PL 0.969 0.040 0.048 0.039 0.239 0210 0170 0128 0.157 B-XY 0.929 0,961 0.089 0.080 0.289 0.258 0.216 0171 0.204 B-YA 0.987 0.953 0.914 0.017. 0.211 0.181 0.170 0.101 0.163 B-HL 0.998 0.962 0.923 0.983 0.193 0.164 0152 0.086 0.146 S-HLI 0.820 0.788 0.749 0810 0.825 0.009 0.039 0.079 0.050 “JL 0.844 0811 0772 0834 0848 0.99] 0.025 0.057 0.050 S-LN 0.857 0844 0.806 0.843 0.859 0.962 0.975 0.053 0.070 S-SX 0.914 0.880 0843 0.904 0918 0924 0.944 0.948 0.041 S-BJ 0.863 0.855 0815 0849 0864 0.952 0.951 0.933 0.960 E. brachypus B-XY E. senticosu $-SX E.giraldii E.gracilistylus E.sessiliflorus ———| 0.20 0.15 0.10 0.05 PRU ie igceseaasacac ri | 1 flIIDCAAAY £ 1 ted gri L L J Bl. °/. | 4 J > ait Fic.2 f \ f lide 3 southeastern Gansu Province and central Shaanxi Province. Another major dif- ference is the breeding system. Eleutherococcus senticosus is reported to be trioecious and protandrous (Liu et al. 1997a, 1998). But because individuals with ermaphroditic flowers are very rare (Liu et al. 1997b), this species is function- ally dioecious with insect-mediated outcrossing (Liu et al. 1998a). Eleutherococcus brachypus has hermaphrodite flowers. Both selfing and insect- mediated outcrossing are important in its sexual reproduction. Similar to E. senticosus, anthers of E. brachypus start to shed pollen at least five days before 1004 BRIT.ORG/SIDA 21(2) the receptivity of stigmas of the same flower (Wang et al. 1997), suggesting out- crossing. Wang et al. (1997) proposed that “outcrossing” within local popula- tions may represent selfing in a broad sense because local populations may be ramets of a single clone. But our study has shown that there is genetic variation within local populations. Thus the interpretation of a local population to be a group of ramets froma single clone is not supported (also see Yan et al. 1997). The degree of human impact may also explain the difference in genetic diversity between these two species. The populations of E. senticosus assayed in this study have not suffered serious disturbance because they were mostly in nature reserves. In contrast, all populations of E. brachypus sampled were seri- ously disturbed, and they were finely fragmented due to land reclamation for farming. Genetic diversity among populations Considerable genetic variation was detected among populations (Table 3). The level of genetic diversity varies across localities in E. senticosus. The population S-LN in Liaoning Province is the center of the present distribution of this spe- cies, and the genetic diversity (He) is highest. The population S-SX in Shanxi Province is peripheral and its genetic diversity is the lowest. This pattern has also been reported in many other plant species (see Crawford 1990; Hamrick & Godt 1996). In E. brachypus, however, the Tianshui population (B-TS) near the westernmost range of the species has the highest genetic diversity, whereas the Xunyi population (B-XY) near the distributional center shows the lowest level of genetic diversity. This unusual pattern may be due to a greater impact of human disturbance on E. brachypus. Implications for conservation In Eleutherococcus senticosus in China, populations near the distributional cen- ter have a higher level of genetic variation. From this center to the periphery, polymorphism decreases. The low genetic diversity in the populations of North China is expected due to their peripheral positions. When the entire distribu- tional range (including the Far East of Russia) is considered, the population S- HL] is also central. The relatively low genetic diversity of this population sug- gests that it may have suffered from genetic loss. Population destruction of E. senticosus has been serious in Heilongjiang Province. Although the genetic structure of E. senticosus has not been seriously dam- aged, overharvesting should be prevented to maintain the sustainability of this species. Recent establishment of nature reserves in Northeast China (the center of genetic variation of this species in China) has been successful in protecting some populations of E. senticosus. Demographic investigation outside the re- serves every few years is needed to determine the reduction rate of populations. ZHOU ET AL., BRACHYPUS CONSERVATION 1005 If it is rapid, use of this species should be controlled, and ex situ conservation of special genetic resources may be adopted in nearby nature reserves. Conservation of E. brachypus is urgent because considerable genetic loss has occurred. No significant correlation has been found between genetic differ- entiation of populations and their geographical distances from the distribu- tional center, suggesting that genetic structure has been altered. The vegetation of the Loess Plateau has been seriously fragmented. In order to maximize the conservation of genetic diversity, all natural populations should be protected. If nature reserves are to be established to conserve populations of this impor- tant species, priorities should be given to those in southern Gansu Province, where genetic diversity is high and habitat loss is serious. ACKNOWLEDGMENTS We thank Zheng Hongchun for assistance in sample collection, and Pete Lowry and Greg Plunkett for most helpful comments. This study was supported by grants from the National Natural Science Foundation of China (grant 3939150 to D-Y. Hong), the Excellent Overseas Chinese Research Grant of the Chinese Academy of Sciences (toJ. Wen), and the US National Science Foundation (DEB 0108536 to J. Wen). REFERENCES Crawroro, DJ. 1990. Plant molecular systematics. J.Wiley & Sons, New York. Day, S.L,, L.H. Wane, and N.H.Wu. 1998. RAPD analysis of Acanthopanax senticosus genetic diversity. Prog. Nat. Sci. 8:465—468. Davypov, M. and A.D. Krikorian. 2000. Eleutherococcus senticosus (Rupr. & Maxim.) Maxim. (Araliaceae) as an adaptogen:a closer look. J. Ethnopharmacology 72:345-393. Duke, JA. 1989. Ginseng: a concise handbook. Reference Publications, Inc., Algonac, Michigan. FarNswort, N.R., A.D. KincHorn, D.D. Soparto, and D.P. Water. 1985. Siberian ginseng (Eleutherococcus senticosus): Current status as an adaptogen. In: H. Wagner, H.Z. Hikino, and N.R. Farnsworth, eds.Economic and medicinal plant research.Volume 1.Academic Press, London. Pp. 155-215. Fu, L.G.and J.M.Jin. 1992.China plant red data book, rare and endangered plants. Volume 1. Science Press, Beijing. GitzeNDANNer, M.A.and PS. Soutis.2001.Genetic variation in rare and widespread Lomatium species (Apiaceae): A comparison of AFLP and SSCP data. Edinb. J. Bot. 58:347—-356. GLATTHAAR-SAALMULLER, B., F. SACHER, and A. Esperester. 2001. Antiviral activity of an extract de- rived from roots of Eleutherococcus senticosus. Antiviral Res. 50:223-228. Hamarick, J.L.and M.J.W.Goor. 1990. Allozyme diversity in plant species. |n:A.H.D.Brown,M.1. Clegg, A.L. Kahler, and B.S. Weir, eds. Plant population genetics, breeding, and genetic resources. Sinauer Associates Inc., Sunderland, Massachusetts. Pp. 43-63. 1006 BRIT.ORG/SIDA 21(2 = Hamrick, J.L.and M.J.W.Goor. 1996. Conservation genetics of endemic plant species. |n:J.C. Avise and J.L. Hamrick, eds. Conservation genetics: case histories from nature. Chap- man & Hall, New York. Pp. 281-304. Hotsinaer, KE. and L.D. Gorturs. 1991. Conservation of rare and endangered plants: Prin- ciples and prospects. In: D.A. Fatk and K.E. Housincer, eds. Genetics and conservation of rare plants. Oxford University Press, Oxford. Pp. 195-223. Hu, S.-Y. 1980. Eleutherococcus vs.Acanthopanax. J. Arnold Arbor. 61: 107-111. LIANG, M., B. Wu, N.C. Liana, L. Tone, T.Y. HuAne, and J.L.Li. 1994, Antitumor action and mecha- nism of Acanthopanax senticosus polysaccharides. Chinese Pharmacol Bull. 10: 105-109. Liu, L.D,, Z.L. Wana, G.W. Tian, and J.H. SHenc. 1997a. Observation on floral morphology and heteranthery of Eleutherococcus senticosus (Araliaceae). Acta Phytotax. Sinica 35:16. Liu, L.D., Z.L. Wane, G.W. Tian, and J.H. SHenc. 1997b. Studies on sexual reproduction and vegetative propagation of Eleutherococcus senticosus (Araliaceae). Acta Phytotax. Sinica Liu, L.D., Z.L.Wano, G.W.Tian, and J.H.SHenc. 1998. The pollination biology of Eleutherococcus senticosus (Araliaceae). Acta Phytotax. Sinica 36:19-27, Nei, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89:583-590, Newron, A.C., T.R. Atunurt, A.C.M. Gituies, A. Lowe, and R.A. Ennos. 1999. Molecular phylo- geography, intraspecific variation and the conservation of tree species. Trends Ecol. Evol. 14:140-145. RiesberG, LH. and S.M. Swensen. 1996. Conservation genetics of endangered island plants. In: J.C. Avise AND J.L. Hamrick, eds. Conservation genetics: case histories from nature. Chap- man & Hall, New York. Pp. 305-334. SoesarTo, D. D.and N.R. Farnsworth. 1978. The correct name for Siberian ginseng, Bot. Mus Leafl. 26:339-343. Sous, D.E., CH. Haurter, D.C. Darrow, and G.J. Gastony. 1983. Starch gel electrophoresis of ferns: a compilation of grinding buffers, gel and electrode buffer and staining sched- ules. Am. Fern. J. 73:9-27. SworrorD, D.L. and R.B. SeLANDeR. 1989. BIOSYS-1, a computer program for the analysis of genetic variation in populational genetic and biochemical systematics. Version 1.7. Illinois Natural History Survey, Champaign, Illinois. Tian, G.W., Z.L. Wane, L.D. Liu, and J.H. SHENG. 1998. Studies on the structure, afterripening and cytochemistry of seeds in Eleutt us brachypus Harms. Acta Phytotax. Sinica Wane, 2. L,, L.D. Liu, G.W. Tian, and J.H. SHENG. 1997, Flowering and pollination biology of Eleutherococcus brachypus. Chin. Biodiv. 5:251-256. Wane, Z.R. 1998. Plant allozyme analysis. Science Press, Beijing. Xie, C.S., WAR. ZHANG, 2.2. Lone, and S.K. Xu. 1989. Immunoregulatory effect of polysaccha- ride of Acanthopanax senticosus (PAS) l:immunological mechanism of PAS against can- cer. Zhonghua Zhongliu Zazhi 11:338-340. ZHOU ET AL., PRRACHYDIIC Adan Xu, R.S., S.C. Fens, Z.Y. Fan, C.N. Ye, S.K. ZHAl, and M.L. SHEN. 1983. The study of polysaccharide composition of Acanthopanax senticosus. Chinese Sci. Bull. 28:185-187. Yan, HJ.,S.L.Dai,and N.H.Wu. 1997.RAPD analysis of natural populations of Acanthopanax brachypus. Cell Res. 7:99-106. ZHAO, Y.Q., LJ. Wu, and X. Li. 1990. Structure studies of liriodendrin, active principle in Acanthopanax senticosus. Chinese Trad. Herb. Drugs 21:140-141. ZHAO, Y.Q,, S.S. YANG, J.H. Liu, and G.R. ZHao. 1993. Studies on the chemical constituents of Acanthopanax senticosus (Rupr. et Maxim.) Harms. J. Chinese Materia Medica 18: 428-429. ZHAO, Y.Q,, S.S. YANG, Y.D. Sun, and Y. Hao. 1991. |solation and identification of isofraxidin and stilbene compound in Acanthopanax senticosus. Chinese Trad. Herb. Drugs 22: 5 1Gr525: POTENTIAL BIOLOGICAL CONTROL OF LANTANA CAMARA IN THE GALAPAGOS USING THE RUST PUCCINIA LANIANAE Jorge Luis Renteria B. Carol Ellison Introduced Plants Program Invasive Species Management any Department CABI Bioscience UK Center (Ascot) eer ie Darwin Research Station Silwood Park, Ascot Galapagos, ECUADOR Berkshire, SL5 7TA, UK jrenteria@fcdarwin.org.ec c.ellison@cabi.org ASBTRACT Laboratory experiments were carried out in England he specifici denvi lreq ments of a Peruvian isolate of the fungus Puccinial Farlow, | Kk tl Lantana camara L.a serious problem in Galapagos. Eight species al ae representing five families were inoculated with the fungus and kept in a dew chamber for 48 hours. Lantana peduncularis Andersson and Lantana camara were sourced from Galapagos, ies species related to Lantana were sourced from other places. Dew periods of 5, 8, 11,14, and 20 hours were tested to determine the period necessary for csnes aaa jonmiation and net uate enOn ee mamas camara from Galapagos and Peru develor i y 15 days sori were fully developed. No non-target species developed pi I Mostimporant he rust did not attack the closest | lative f lapagos, the endemic Lant is. Eight in the dew cham- ber h to induce basidi f d host infection, I up to 20 hours induced eee more sori. Although we have not somite yet the experiments to determine the hos range specificity, P lant p gent for Lantana camara in see Key worps: Lantana peduncularis, Lant camara, Puccinia lantanae, rust, biocontrol, Galapagos RESUMEN Se realizaron experimentos de laboratorio en Inglaterra para determinar el nivel de especificidad y requerimientos ambientales del hongo Puccinia lantanae Farlow como potencial agente de control biologico de la eopecie invasora La ntana camara L. en Galapages’ Ocho especies ge pa L.camara ongo inantenttae durante 48 horas en la camara de roci6. Lantana peduncularis Andersson y ee camara fueron colectadas en Galapagos, ak otras iespecls ke laelonadas ck con pena se oe), en otros lugares. Para determinar el L las plantas de L. camara muerOn inociladas Di Scees a diferentes ee de rocio (5, 8, 1, 14, 20 horas). Puccinia lant y t mara ees de Galapagos y Pera. Los sintomas aparecieron seis dias después de la ee y a los 15 dias las pustulas estuvieron completamente desarrolladas. NOS se detectaron sintomas macroscopicos en las otras especies, principalmente en la endémica L. peduncu laris que es es eae mas cercana. Puccinia lantanae fue Fabs a ssporulat e€ amie cea Be camara luego d ; el mayor ere: de ] In 1 j ore ONT } infeccién y | ] liad is a | a “| 13} ic i p lagente, l to, P lant p ial d trol bioldgico Oo oO qmara nara Galapagos de I L | ioe i SIDA 21(2): 1009-1017. 2004 1010 BRIT.ORG/SIDA 21(2) INTRODUCTION Lantana camara (Verbenaceae), an ornamental shrub, native to tropical America, is now found in most tropical and subtropical regions of the world. It is not only widespread but it is also generally considered to be a major pest of agri- cultural and natural areas (Thaman 1974). The L. camara complex will tolerate a wide range of climates. Itcan be found at altitudes between sea level and up to 2000 meters, and between 45° N to 45° S. It flourishes in both dry and wet re- gions, growing on mountain slopes, along coastal areas and in val leys. It is some- what shade tolerant and thus can become the dominant understory plant in open forest and tropical tree crops in its weedy range (Holm et al. 1977). Repro- duction is by seeds that are spread via frugivorous birds, and short distance spread is by the rooting of horizontal stems in contact with the soil. Lantana camara was first introduced as an ornamental into Floreana Is- land in the Galapagos Archipelago in 1938 (Cruzetal. 1986), and has since spread or been carried to other islands, including Santa Cruz. The dense thickets ere ated by this invasive weed impact not only the indigenous flora but also fauna (Hamann 1984: Cruz et al. 1986). There is evidence, for exam ple, that it is affect- ing the nesting habitats of the endangered dark-rumped petrel, Pterodroma phaeopygia (Cruz et al. 1986). Lantana camara was the first weed ever targeted for classical biological control at the turn of the century. The first attempt at the biological control of lantana began in 1902, when 23 insect species were imported into Hawaii from Mexico. Fight of these species were established (Perkins & Swezey 1924). A to- tal of 36 insect species has since been released in 33 countries Julien & Griffiths 1998), but control in Hawaii, as well as in other parts of the world, has only been partially successful (Taylor 1989). This has mainly been due to the genetic diversity, and hence environmental adaptability, of the weedy biotypes which outstrip those of its natural enemies. New biocontrol agents are still being evalu- ated and released, including pathogens. A broad range of pathogens has been recorded infecting L. camara in its native range (Barreto et al. 1995). Three fun- gal agents have been released so far: a species of Septoria was released in Ha- waii, originally from Ecuador (Trujillo 1995), a rust Prospodium tuberculatum (ex Brazil) was released in Australia in 2001, and a leaf spot pathogen, Mycovellosiella lantanae var. lantanae (ex Florida, USA) was released in South Africa, also in 2001. The impact of these agents is still pending. Puccinia tuberculatum was screened against the invasive and native species of Lantana [rom the Galapagos at CABI Bioscience, but it was found to infect, albeit mildly, the native lantana, and was discounted asa potential agent. A number of other candidate pathogens have been identi fied with excellent potential, including a stem and leaf rust, Puccinia lantanae (Barreto et al. 1995). Puccinia lantanae Farlow (Basidiomycotina, Uredinales) occurs in tropical RENTERIA AND ELLISON, BIOLOGICAL CONTROL OF LANTANA CAMARA IN THE GALAPAGOS 1011 and subtropical regions of America: From Mexico and Florida, through the Caribbean and as far South as Argentina. Puccinia lantanae has been recorded from a number of Lantana spp., but there is evidence of distinct races that are only capable of attacking single species, and are even specific to biotypes within that species. This rust is recorded as a microcyclic (only teliospores and basid- iospores in the life cycle) and autoecious (completes life cycle on one host spe- cies) species. The teliospores remain in the sorus on the host plant, and are not released. Under conditions of high humidity, teliospores germinate and pro- duce basidiospores that are released from the teliospores. These infect fresh plant material, from which more teliospores result, and hence complete the life cycle. METHODS Plant material and fungal inoculations Plants species used in the experiment were grown from stem cuttings from the CABI Bioscience stock plant collection. Lantana camaraand Lantana peduncu- laris were collected originally from the Galapagos Islands (Santa Cruz). Using rooting powder, stem cuttings were planted in pots containing substrate John Innes no. 2). Plants were kept ina quarantine glasshouse set at a minimum tem- perature of 20 °C with 12 hours of artificial light and watered everyday. The rust Puccinia lantanae used was taken from the CABI Bioscience speci- men collection held on living plants (isolate reference number W1914). The fun- gus, like all rusts isa biotroph and therefore culturing can only be done in vivo. To inoculate the experimental plants for host range testing and assessment of the minimum dew period requirement, sori of Puccinia lantanae were sus- pended over new shoots; between two to four shoots were targeted for each pot- ted plant. The small piece of plant tissue containing the sori of teliospores, was attached to small Petri dish using petroleum jelly (Vaseline). Care was taken that no Vaseline was deposited on the fungal material. Petri dishes were attached toasmall stick a distance of 2 cm above the young leaves, making sure that the teliospores were directly above the leaf, so the basidiospores are released onto the potentially susceptible part of the plant. (Koutsidou 2000). The target area was usually the four youngest leaves of any given stem. The inoculated stems were marked by tying a string to the stem. Target plants were watered and the leaves were wetted witha fine mist of sterile distilled water before putting them in the artificial dew chamber (Mercia Scientific, Birmingham, UK). All in vivo experiments were done in a quarantine greenhouse at CABI Bioscience, Ascot. United Kingdom. There were two experiments described below: Symptom Development and Host Range Specificity For the host range specificity experiments, eight species from five families were used (Table 1). All test plants species were inoculated with the rust Puccinia 1012 BRIT.ORG/SIDA 21(2) Taste 1. Host range test species list and results of host specificity testing of Puccinia lantanae. Family Species Provenance Native in Susceptibility rating Bignonaceae Tecomanthe hilli Australia Australia 0 Boraginaceae Cordia dichotoma Australia Australasia ) Lamiaceae Plectranthus parviflorus Australia Australasia 0 Lamiaceae Vitex triflora Australia South America 0 Lamiaceae Gmelina leichhardtii Australia Australia 0) Verbenaceae Lantana camara Galapagos South America 3 Verbenaceae Lantana camara Peru South America 3 Verbenaceae Lantana montevidensis Australia South America 0) Verbenaceae Lantana peduncularis Galapagos Galapagos 0 lantanae using the method described above. At least three replicate plants were inoculated per species. Plants were incubated at 20° C for 48 hours ina dew simu- lation chamber to induce teliospore germination, basidiospores formation, and provide an optimum environment for potential plant infection. Symptoms were recorded according to a rating system devised to assess the susceptibility of plant species to P lantanae based on the visible symptoms (Koutsidou 2000): 0 No macroscopic symptoms; 1 Chlorosis on the leaf sur- face; 2 Restricted sporulation (sorus diameter <2mm), and; 3 Abundant sporu- lation (Sorus diameter >2mm). Dew Period Requirements Lantana camara plants (from Galapagos) were inoculated with the rust Puccinia lantanae using the method described above, although one sorus per shoot was used as the standard inoculum, of 5mm diameter. Three or more shoots were inoculated per replicate plant and a mean number of sori per shoot taken for each plant. Dew period treatments of 5, 8, 11,14, and 20 hours were compared, with two replicate plants per treatment. The dew chamber was set at 20° C. After this treatment, the inoculum was removed and plants moved into the quarantine glasshouse and pots watered normally avoiding the wetting of leaves. Plants were checked regularly for macrosymptoms and sorus development. RESULTS Symptom Development and Host Range Specificity The first appearance of symptoms of P lantanae infection on the L. camara from Galapagos, occurred 6-7 days after the inoculation, as small chlorotic spots. These spots enlarged, and after approximately 13-15 days the first symptom of sporulation i.e. sori became apparent. The size of the sori on leaves differed from between 1 mm to6 mm in diameter. In general, the younger the leaves (approxi- mately <5mm diameter) were at inoculation the larger the sori that formed. Leaves that had already partially expanded before infection produced the RENTERIA AND ELLISON, BIOLOGICAL CONTROL OF LANTANA CAMARA IN THE GALAPAGOS 1013 smaller sori. However, high densities of sori on a leaf also resulted in smaller average sorus size. No sporulation was observed on leaves that were fully ex- panded at inoculation. Sporulation occurred mainly on the lower surface of the leaf. When the density of sori was high, very premature leaf abscission was observed (around 13 days after inoculation). If the density of sori was lower, a necrotic area formed around them, which increased until earlier than normal leaf fall, but after full rust symptom expression. Infection often occurred on stems and petioles also. Table 1 gives the results of the host specificity testing. Although this is a limited host range test, the results suggest that P. lantanae is host specific to L. camara. Puccinia lantanae was not able to infect nor sporulate on any of the other seven related species used in the experiments. Even the most closely re- lated species L. peduncularis (Galapagos) and L. montevidensis were resistant to P lantanae, suggesting strong host-specificity. Dew Period Requirements Puccinia lantanae was able to sporulate and infect L. camara plants after only 8 hours of dew. Maximum infection and sori development was obtained at or after 20 hours of dew (see Fig. 1). Figure 2 shows the different levels of infection by P. lantanae after differ- ent lengths of dew period. Clearly, more basidiospores are released over a longer period of time in humid conditions. DISCUSSION Puccinia lantanae isolate W1914 from Peru seems to be significantly more de- structive to Lantana camara than other pathotypes of P lantanae, that are fre- quently observed throughout the native range of the plant. Previous records of isolates of P lantanae report that the pathogen only infects leaves (Barreto et al. 1995). The fact that the isolate W1914 can also infect petioles and stem means that the rust is much more damaging to the weed and is therefore a better po- tential biological control agent than originally estimated. Whole branches may drop asa result of stem infection and infection of the leaves can be very severe. Disease symptoms start to appear 5-7 days after inoculation and sori can grow up to 6 mm in diameter suggesting a rapid and destructive infection of this rust. This rust is able to release the basidiospores that can infect fresh host tis- sue within the first 8 hours of a dew period however longer periods of humid- ity favor it. Puccinia lantanae seems to be a promising biological control agent to tar- get L. camara from the Galapagos Islands. Nevertheless it is necessary to con- tinue with the host range specificity test using the related native and endemic species from Galapagos to avoid doubts about non-target species effects. In addition, more collections of L. camara from Galapagos need to be made 1014 BRIT.ORG/SIDA 21(2) 269.7 N n jo) No Oo oO 1 1325 Average number of sori per shoot S a Co © i 1 e) 8 1] 14 20 Dew period time in hours Fic. 1.A L ra ° } 1 | Batak: haf | Ce ea to ensure that the rust is able to attack all the possible forms of the weed that may occur in Galapagos. There is evidence (from observations on flower color and plant growth form), that this weed has a narrow genetic base on the is- lands, and hence this rust is likely to infect all populations. Although only lim- ited host specificity testing has been undertaken, the results suggest that this isolate is specific to L. camara. Classical releases of rusts with the same type of life cycle as P lantanae (e.g. microcyclic), show that such short-cycled rusts spread swiftly through and between plant populations (Morin et al. 1996.). The impact of successful classi- cal biological control agents on woody weed species, such as L. camara, has tended to take a decade or more to be demonstrated, and often a suite of natu- ral enemies is required. It is anticipated that due to the damaging nature of the rust and the short generation time, the impact may be observed sooner than is normal, and that this single agent may provide effective control. In the subsequent glasshouse based tests of this agent, untreated (not in- oculated) plants of L. camara should be compared with treated individuals, in RENTERIA AND ELLISON, BIOLOGICAL CONTROL OF LANTANA CAMARA IN THE GALAPAGOS 1015 A Fic. 2. Levels of infection of Puccinia | It of the different d iod A. eight hours, B. four- teen hours, C y hours in the dew chamber. 1016 BRIT.ORG/SIDA 21(2) order to investigate the effect of the rust on plant growth and survival. This may help give an indication of the possible level of impact of the agent in the field, although this is known to be quite difficult to study with woody species in controlled conditions. Lantana camara is difficult to control and appears impossible to eradicate due to the wide range it occupies in the Galapagos. Biological control isa realis- tic management option. This weed has been studied for over a century asa clas- sical biological control target. Although success has been limited, the rust Puccinia lantanae constitutes a new method and a potentially effective agent to try in Galapagos Islands. ACKNOWLEDGMENTS We gratefully acknowledge the following organizations and people for their fi- nancial support and help: Global Environment Facility (GEF) Invasive species project, through UNDP-ECU/00/G3l1, CABI Bioscience International, the Botany Department of the Charles Darwin Research Station, Richard Shaw, Harry Evans, Sarah Thomas, Djamila Djeddour, Marion Seier, Keith Holmes, Rob Tan- ner and Lynn Hill. Alan Tye and Chris Buddenhagen provided detailed com- ments on the manuscript. Joe Hennen (BRIT) and an anonymous reviewer pro- vided valuable comments. REFERENCES Barreto, R.W., H.C. Evans, and C.A. ELuison. 1995. The mycobiota of the weed Lantana camara in Brazil, with particular reference to biological control.Mycological Research 99:769-782 Cruz, F.J.Cruz, and J. Laweson. 1986. Lantana camara L.,a threat to native plants and ani- mals. Noticias de Galapagos 43:10-11 HAMANN, O. 1984. Changes and threats to the vegetation. |n:R. Perry, ed. Key environments: Galapagos. Pergamon Press, Oxford, UK. Pp. 115-131 Hom, L.G., D.L. PLUCKNETT, J.V. PANCHO, and J.P. Hersercer. 1977. The worlds worst weeds. Distri- bution and biology. University Press of Hawaii, Honolulu, Hawaii. Juuien, M.H. and M.W. GrireitHs. 1998. Biological control of weeds: A world catalogue of agents and their target weeds (Fourth edition).CABI Publishing:Wallingford, Oxon, UK. Koutsipou C. 2000. Studies on Puccinia lantanae, a potential biocontrol agent for Lantana camara. Msc Thesis, Imperial College, London. Pp. 16-68. Morin, L., B.A. Auto, and H.E. SmitH. 1996 Rust epidemics, climate and control of Xanthium occidentale. In: V.C. Moran and J.H. Hoffmann, eds. Proceedings of the IX International symposium on biological control of weeds. University of Cape Town, Cape Town, South Africa. Pp. 385-391. Perkins, R.C.L. and O.H. Swezey. 1924. The introduction into Hawaii of insects that attack lantana. Bull. Exp. Sta. of the Hawaiian Sugar Planters’ Assoc. 16:1-83 Taytor, E.E. 1989. A history of biological control of Lantana camara in New South Wales. Plant Protection Quart. 4:61-65. RENTERIA AND ELLISON, BIOLOGICAL CONTROL OF LANTANA CAMARA IN THE GALAPAGOS 1017 THAMAN, R.R. 1974. Lantana camara: \ts introduction, dispersal and impact on islands of the tropical Pacific Ocean. Micronesica 10:17-39. THomas S.E.and C.A. ELuison 2000.A century of classical biological control of Lantana camara: can pathogens make a significant difference? In:N.R. Spencer, ed. Proceeding of the X International symposium on biological control of weeds. Montana State University, Bozeman. Pp. 97-104. TRUJILLO, E.E. 1995. Septoria leaf spot of lantana from Ecuador: A potential biological con- trol for bush lantana in forests of Hawaii. Plant Disease 79:819-821. ISOLATION AND IDENTIFICATION OF FUNGI ASSOCIATED WITH THE RHIZOSPHERE AND RHIZOPLANE OF WILD AND CULTIVATED PLANTS OF PAKISTAN Shamim Akhtar Qureshi Viqar Sultana Department of Biochemistry ee DEBIOCnenitst University of Karachi Univer ity of Karachi Karachi-75270, PAKISTAN Karachi- 75270 PAKISTAN Syed Ehteshamul-Haque Mohammad Athar cas eat California Departm of Fi d Agricultur University of Karact 2014 Gn itol Avenue, Suite 109 Gaon 75270, PAKI STAN Sacramento, California 95814, U.S.A. atariq@cdfa.ca.gov ABSTRACT Fifty-seven species of fungi belonging to 23 genera were isolated and identified from the rhizosphere and rhizoplane of 65 plant species, belonging to 58 genera and 19 families from Sindh and Baluchistan, (Pakistan). A greater number of fungi were isolated from the rhizosphere than from the rhizoplane. In the rhizosphere, Fusarium solani and Aspergillus oa were > dominant followed by Drechslera dustraliensis. In the rhizoplane, Fusarium ual | t. Biocontrol agents like Tricho- derma ee £ honingit, T. viride, V.c losporium, and Stac ne ‘ys atra were pees in low frequency heir poor in the rhi | M ] Zed mays and Sorghun um n bicolor and Stac hybotrys ahlsl ay from Zea mays were reported for the first time from Pakistan profound influences on the soil fertility status as w ll as on the suppression er soil- porns plant his mM ple microbial d biocontro interactions involvi in many cases in comparison with bigcen eal agents used singly. importantly a soil that is suppres- sive to one palacesir ls is not necessaril y UES ssive to another, and so specificity in soil-plant-mi- Modern methods for analyzing microbial commu- nity structures may prove particularly ae to nee define the key organisms or groups of organisms responsible for such natural suy well as for monitoring the spread and impact of introduction of specific biocontrol ae or other management practices on natural microbial populations. RESUMEN f, selad -- ee | Cincuenta y ci ies de hong 23 género la rizosfera y rizoplana de 65 ies vegetales, pertenecientes a 58 g anes ye 19 familias de Sindh y Baluchistan, (Pakistan) Fue ais ee un numero mayor de hongos en la ri leen oplana. En la rizosfera, Fusarium ai y Bere SDB: fueron los ap aes eae de eee ién. Agentes de biocontrol como australiensis. En el rizoplan ichoderma harzianum, I. ee Tvi fide V.chl ee um, | Stachybotryssea ce en baja a recuencia lo que sugiere una competencia pobre en la rizosfe ed mays y Sorghum bicolour, y Stachybotrys parvispora de Zea mays se citan por primera vez de Pakistan. Los microorganismos de los suelos agricolas ejercen profundas influencias en la fertilidad del suelo asi ou SIDA 21(2): 1019-1053. 2004 1020 BRIT.ORG/SIDA 21(2) como en la supresion de enfermedades de las plantas originadas en el suelo. Las interacciones Gcropiau> aoe que implican pacievias y peer en la rizosfera se ve que ejercen un biocontrol sires Or ] 4 cipal [eee | o importante, que un suelo ee es supresivo ae un | y asi la especificidad en las interacci icin microbio para la supresion de la enicemedad existe. Los métodos modernos aes analizar la estructuras de la cum microbiana peeven ser muy valine ce ay udar a Geant rgani grup g I tal Pa Coa a +2 1; 1 pea } ip ee | [ Zz r de bi trol especificos practicas de gestion de poblaciones microbianas naturales INTRODUCTION The rhizosphere has become an important area to test and evaluate new oppor- tunities being developed in biotechnology. The rhizosphere is the portion of soil directly influenced by substances issuing from roots into the soil solution, favoring certain microorganisms, harmful around roots of unthrifty plants and beneficial around roots of healthy plants (Atkinson & Watson 2000; Curl 1982). There is an exchange of materials between the plant root and the surrounding micro-population within the rhizosphere. These materials may inhibit or pro- mote growth of the plant or the microorganisms (Bazin et al. 1990; Filion et al. 2004; Katan 2002). Rhizosphere is therefore the site where biological control of soilborne pathogens takes place. Pathogen population (inoculum density), growth and survival and infection or pathogenesis are all influenced by the rhizosphere (A bawi & Widmer 2000; Curl & Truelove 1986; Manka & Kacprzak 1999). Infection of roots by a soilborne plant pathogen is influenced by the physi- cal and chemical properties of the rhizosphere environment and interaction of the pathogen with other microorganisms in that environment (Dix & Webster 1995; Tate 1995). Saprophytic fungi and bacteria in the rhizosphere and root surface create a competitive deterrent to the colonization of rhizoplane and in- vasion of the plant roots by pathogens (Abawi & Widmer 2000; Tate 1995). It has been reported that m itive fungal species are found in rhizosphere than soil away from roots (Tate 1995). Another special habitat or site of microbial activity is rhi or the root surface which supports relatively high biologic activity than rhizosphere (A bawi & Widmer 2000; Atkinson & Watson 2000). It has been reported that legumes support larger rhizosphere population than non-leguminous plants (Subba Rao 1977). Similarly, rhizosphere of resistant cultivars of pigeon-pea (Cajanus cajan) harbored more Streptomyces and Trichoderma antagonistic to Fusarium udum causal agent of pigeon pea wilt, than susceptible cultivars and Trichoderma viride in the rhizosphere of varieties of tomato resistant to Verticillium wilt Subba Rao 1977). Trichoderma spp. and Paecilomyces lilacinus are known as effective bio- control agents against root infecting fungi and have shown promising results in microplot experiments (Boland & Kuykendall 1998; Burges 1998; Lewis et al. 1998; Whipps 1997, 2001). The opportunity to improve crop productivity by introduc- QURESHI ET AL., ROOT FUNGI OF WILD AND CULTIVATED PLANTS OF PAKISTAN 1021 Taste 1. Some of the soil characteristics of the collection sites. Province/Location Soil Type Soil pH Sindh Province Darsano Chano Surface and sub-surface of soil sandy loam 8.0-8.2 Gharo Surface and sub-surface of soil clay loam 8.2-8.5 Karachi University Campus Surface and sub-surface of soil silty loam 8.0-8.1 athor Surface coarse sand and sub-surface sandy loam 8.0-8.05 Surface and sub-surface of soil silty-sandy loam 8.05-8.1 Shah Faisal Colony Surface and sub-surface of soil sandy loam 8.0-8.1 Thatt Surface and sub-surface of soil clay loam 8.3-8.5 atta Baluchistan Province Hub Surface and sub-surface of soil sandy loam 8.0-8.2 ing organisms t to the unleospucnes is highlighting a major need for the study of fungal 1 the roots of plants. The present report describes the occurrence of fungal species on rhizosphere and rhizoplane of different plant species collected from different parts of Sindh and Baluchistan (Pakistan). MATERIALS AND METHODS Sites and Collection of Samples Eight sites were chosen for the collection of samples; seven from Sindh and one from Baluchistan. Details of collection sites along with some of the soil charac- teristics are provided in Table 1. Overall rainfall of these areas is very scanty ranging from 25mm to 102 mm per year. Average temperature is 50° C (maximum) and 25°C (minimum), and average winter temperature 25° C (maxi- mum) and 8° C (minimum). Young healthy plants were carefully dug out up toa depth of 15cm and root samples with adhering soil (25-50 g depending upon root size) were collected in polyethylene bags. Roots of cultivate ps were collected from agricultural fields. Roots of wild plants were collected from adjacent uncultivated fellow fields. Five replicates of each plant species were collected from each location. Samples were kept under refrigeration at 4°C until the isolation of fungi made within 24 hours. Potato dextrose agar was used for the isolation of fungi in this study, since it supports the growth of most of the fungi from rhizoplane and rhizosphere (van Elsaset al. 2002) and also for endophytes (Halleen et al. 2003), except obligate parasites and those which have special growth requirements. Isolation of Fungi from Rhizosphere Volume displacement technique was used for the isolation of fungi from rhizo- sphere soil as described by Reyes and Mitchell (1962). Root pieces with adhering soil were placed in a graduated cylinder containing 18 mL sterilized distilled water and shaken vigorously. The roots were removed and the process was repeated Taste 2. Fungi isolated from rhizoplane and rhiosphere of wild and cultivated plants growing in Sindh and Baluchistan, Pakistan. No and Host Rhizoplane Location Rhizosphere Location AMARANTHACEAE 1. Amaranthus virides L. Alternaria alternata (Fr.) Pee 4 Alternaria alternata (Fr.) Keissler 46 Aspergillus flavus Link ex Gray 45 Aspergillus is Link ex Gray 45 A.fumigatus Fres. 4 A.fumigatus 2 A. niger van Tieghem 45 A. nidulans jae Winter 5 A. nidulans (Eidam) Winter 5 A. niger van Tieghem > A.terreus Thom 4 A. terreus Thom 2,4,5,6 Chaetomium indicum Corda 2,6 Chaetomium indicum Corda 2,6 Fusarium solani (Mart.) Appel & Wollenw. Drechslera australiensis (Bugni) Subram. emend. S Han 2,5,6 & Jain ex M.B. Ellis 2,4,6 Macrophomina phaseolina (Tassi) Goid. 2 Fusarium solani (Mart.) Appel & Wollenw. emend. Rhizopus stolonifer (Ehrenb.ex Link) Lind. 5 nyd. & Hans 2,5,6 Unidentified black sterile mycelium 24 F oxysporum Schlecht emend. Snyd & Hans. 4 Penicillium crysogenum Thom. 4 tga osum Thom 2 Trichoderma viride Pers. ex Gray 45 Unidentified black sterile mycelium 2,4 Unidentified yellow sterile mycelium 24 2. Aerva javanica (Burm. f.) Alternaria alternata (Fr.) Keissler 5 Aspergillus flavus Link ex Gray 5 errill Aspergillus flavus Link ex Gray 5 A.niger van Tieghem 5 A.ni ger van Tieghem 5 A. terreus Thom 5 Fi Schlecht Cunninghamella echinulata (Thaxt.) Thaxt. =) emend. Snyd& Hans 5 Fusarium solani (Mart.) Appel & Wollenw. emend. F. solani (Mart.) Appel & Wollenw. Snyd. & Hans 5 emend. Snyd. & Hans 5 Penicillium luteum Zukel 5 izoctonia solani Kuhn 5 3. Digera muricata (L.) Mart. Aspergillus flavus Link ex Gray 1 Alternaria alternata (Fr.) Keissler 1 A. niger van Tieghem 1 Aspergillus flavus Link ex Gray 1 eZOL (Z)L@ VOIS/DYO"LING TasLe 2. continued No and Host Rhizoplane Location Rhizosphere Location A. terreus Thom ] A.niger van Tieghem 1 Fusarium solani (Mart.) Appel & Wollenw. A. terreus Thom 1 mend. S Han ] Cladosporium sp. 1 Macrophomina phaseolina Goid | Fi I | n (W.G. Sm.) Sacc. ] Penicillium waksmanni Zales 1 Myrotheci inctum (Corda) Sacc. ] Boniciiun waksmanni Zaleski 1 ASCLEPIADACEAE 4, Calotropis procera (Ait.) Alternaria alternata (Fr.) Keissler 2 Alternaria alternata (Fr.) Keissler 2 Ait.f. (Rooster TREE) Aspergillus flavus Link 2 Aspergillus flavus Link 25 A. niger van Tieghem 2 A. niger van Tieghem 2,5 A. terreus Thom 2 A. terreus Thom 2,5 Cunninghamella echinulata (Thaxt.) Thaxt. 2 Cephalosporium sp. 2 Fusarium solani (Mart.) Appel & Wollenw. Chaetomium flavum Omvik 2 emend. Snyd. & Hans 2 Trichoderma viride Pers. ex Gray 2 BORAGINACEAE 5. Heliotropium europeam L. Alt iq alternata (Fr.) Keissler 2 Alternaria alternata (Fr.) Keissler 2 (EUROPEAN HELIOTROPE) Aspergillus flavus Link ex Gray 2,6 Aspergillus flavus Link ex Gray 2,6 A. nidulans (Eidam) Winter 2 es 2,6 A. niger van Tieghem 2,6 A. nidulans (Eidam) Winter 2 Chaetomium indicum Corda 2 A. terreus Th 2 Drechslera australiensis (Bugni) Subram. Cephalosporium sp 2 & Jain ex M.B.Ellis. 6 Chaetomium indicum Corda 2 D.hawaiiensis nee Subram. Cunninghamella echinulata (Thaxt.) Thaxt. 2 & Jain ex M.B. Ellis 2 Drechslera gee fensis (Bugni) Subram. D. halodes ceca Subram. Ellis 6 Fusarium solani (Mart.) Appel & Wollenw. emend. ae an Hans 2,6 & Jain ex M.B. Ellis 2 Fusarium solani (Mart.) Appel & Wollenw. NVLSINWd 40 SLNW1d G3LVAILIND ONY C1IM 40 ISNNJ LOOY “Tv 13 1HS3¥Nd €Z0L TABLE 2. continued No and Host Rhizoplane Location Rhizosphere Location emend. Snyd. & Hans 2,6 Penicillium luteum Zuka 2 Rhizopus stolonifer (Ehrenb. ex Link) Lind. 2 Unidentified sclerotial fungus 2 Unidentified sclerotial fungus 2 Unidentified white sterile mycelium 2,6 CANNACEAE 6. Canna indica L. Drechslera australiensis (Bugni) Subram. Alternaria alternata (Fr.) Keissler 1 (Common Lity) Jain ex M.B. Ellis 1 Aspergillus flavus Link 1 Fusari YSp Schlecht emend. A.niger van Tieghem 1 Snyd. & Hans. ] . Sumani (Fres.) Them & Church 1 F. solani (Mart.) Appel & Wollenw. traliensis (Bugni) Subram. em S Hans 1 gain ex MB. Ellis 1 Macrophomina phaseolina (Tassi) Goid 1 Fusarium solani (Mart.) Appel & Wollenw. mend. Snyd. & Hans 1 Macrophomina phaseolina (Tassi) Goid 1 Penicillium luteum Zukal 1 CARICACEAE 7. Carica papaya L. Aspergillus flavus Link ex Gray 2 Aspergillus flavus Link ex Gray 2 (Papaya A. niger Vv hem 2 A. glaucus Link 2 nee ee Winter 2 A.niger van Tieghem 2 A. terreus Thom 2 A. nidulans (Eidarm) Winter 2 Rhizopus stolonifer (Ehrenb. ex Link) Lind. 2 A. terreu 2 ol. cil lium Uren Zukal 2 hod arzianum Rifai 2 CHENOPODIACEAE 8. Beta vulgaris L. (SUGAR BEET) Drechslera australiensis (Bugni) Subram. Alternaria alt ta (Fr.) Keissler 6 Jain ex M.B.Ellis 3 Aspergillus flavus ex Gray 3,6 Fusarium ee (Mart) Appel & Wollenw. A.niger van Tieghem 3,6 ; emend. Snyd. & Hans 3,6 Drechslera australiensis (Bugni) Subram. Macrophomina phaseolina (Tassi) Goid. 3,6 & Jain ex M.B. Ellis. 6 (2)L@ VaIS/HO'LIS vz0L Taste 2. continued No and Host Rhizoplane Location Rhizosphere Location Rhizoctonia solani Kuhn 3,6 Fusarium solani (Mart) Appel & Wollenw. ane ae & Hans 6 9, Chenopodium album L. Alternaria alternata (Fr.) Keissler 7 Al ata (Fr.) Keissler 7 (PiGweeD) ston lera australiensis (Bugni) Subram. & Jain pete us flavus Link ex Gray 7: Ellis A.niger van Tieghem 7 pe ium ae ene Appel & Wollenw. emend. A. terreus Thom F Snyd. & H Chaetomium galas Kunze ex Steud. 7 . indicum 7: h Cee is (Bugni) Subram. & ~ jain ex M.B. Ellis qi Fusarium semitectum Berk. & Rav. ri F. solani (Mart.) Appel & Wollenw. emend. Snyd 7 & Hans pile Hiv jum Leena Tho 7 10. Spin [ L. Alternaria alternata (Fr.) Keissler 2 a (Fr.) ae 2,6 (SPINACH) ne una esanel echin ulata (Thaxt.) Thaxt. 2 sera _ van Tieghem 2,6 aliensis (Bugni) Subram. & us Tho 2 Jain ex M.B.Ellis 2,6 sae 2 Fi ysporum Schlecht.emend. Synd. Deieee ae ese (Bugni) Subram. & & Hans 2 ae nex MBE llis, 2,6 F. solani (Mart.) Appel & Wollenw. emend. n Schlecht. emend. ee & Hans 2 S S 2,6 sclan (Mart) Appel & Wollenw.eme Macrophomina phaseolina (Tassi) Goid. 2,6 Snyd.& Hans 2,6 Rhizoctonia solani Kuhn 2,6 Rhizoctonia solani Kuhn 2 Unidentified white sterile mycelium 2,6 COMPOSITAE 11. Conyza bonariensis (L.) Alternaria alternata (Fr.) Keissler 5 Aspergillus flavus Link ex Gray 5 Cronquist Aspergillus flavus Link ex Gray 5 A.niger van Tieghem 5 NWLSINVd 40 SLNV1d G3LVALIIND ONY G1IM 40 ISNNd LOOY “TW 13 IHSauND S7OL Tasle 2. continued No and Host Rhizoplane Location Rhizosphere Location A. niger van Tieghem 5 A. terreus Thom 5 A. terreus Thom 5 Cunninghamella echinulata (Thaxt.) Thaxt. 5 Rhizopus stolonifer (Ehrenb.ex Link) Lind 5 12. Helianthus annuus L. (SUNFLOWER) Alternaria alternata (Fr.) Keissler 1,8 Aspergillus flavus Link ex Gray 1,8 Drechslera australiensis (Bugni) Subram.& Jain ex A. niger van Tieghe 1,8 M.B.Ellis. 1,8 A. terreus Thom ] Fusari ySf n Schlecht emend. Fusarium solani (Mart.) Appel & Wollenw. emend. 1,8 Snyd.& Hans 1,8 F. solani (Mart) os & Wollenw.emend. Macrophomina phaseolina (Tassi) Goid 1,8 Snyd. & Han 1,8 Pencillium luteum Zukal 1 roan phaseol ina (Tassi) Goid 1,8 Unidentified white sterile mycelium 2 Rhizoctonia solani Kuh 1,8 rales ce sterile Heal 1 13. Lactuca sativa L. ta (Fr.) Keissler 2 Alternaria alternata (Fr.) Keissler 2 (LetTUcE) Fusariu m oxysporum Schlecht emend. Snyd. Aspergillus flavus Link ex Gray 2,6 Hans. 2 A. niger van Tieghem 2,6 F. solani pe — & Wollenw.emend. Fusarium solani (Mart.) Appel & Wollenw. emend. Snyd. & H 2,6 Snyd.& Hans 2,6 Macr ate eae ina (Tassi) Goid 2,6 Rhizoctonia solani Ku 2,6 14. Launea nudicaulis Hook.f. Aspergillus niger Van ae hem 1 Aspergillus flavus Link ex Gray 1 Fusarium solani (Mart.) Appel & Wollenw. emend. niger van Ti ] Snyd. & Hans 1 A. nidulans (Eidam) Winter ] Macrophomina Mee ina (Tassi) Goid. 1 A. terreus ue 1 Rhizoctonia solani Kuhn 1 Drechslera a is (Bugni) Subram. Unidentified black sterile mycelium 1,2 & Jain ex MB. Ellis 1 9701 (Z)LZ VaIs/D¥O'LINS TasLe 2. continued No and Host Rhizoplane Location Rhizosphere Location Fusarium semitectum Berk. & R 1 F. solani (Mart.) hs ae emend. Snyd. & Han ] acro ee phaseolina (Tassi) Goid. ] Unidentified black sterile mycelium 1,2 CONVOLVULACEAE lvulus arvensi Alt ia alt ta (Fr.) Keissler 2 Alternaria alternata (Fr.) Keissler i Aspergillus flavus Link ex Gray 1,7 Aapergillus flavus Link ex Gray ] A. niger van Tie 1,2,7 A. niger van Tieghem ] ee a inulata (Thaxt.) Thaxt. 2 A. nidulans (Eidan) Winter 2 Cladosporiu 1,2 A. en = ) Thom & Churh ] Fusarium ae (Mart) Appel & Wollenw. A. terre 1,2 emend. Snyd. & Hans 2 ey Sp. F oxysporum Schlecht. emend. Snyd.& Hans 2 Fusarium es (Mart.) Appel & Wollenw. emend Macrophomina phaseolina (Tassi) Goid. ] Snyd. & Hans 2 Rhizoctonia solani Kuhn 7 F oxysporum Schlecht. emend. Snyd. & Hans 2 Rhizopus stolonifer (Ehrenb.ex Link) Lind. 2 Macrophomina phaseolina (Tassi) Goid. ] Rhizoctonia solani 7 Unidentified sterile fungus 7 CRUCIFERAE ica j (L.) Czern. Fusarium solani (Mart) Appel & Wollenw. Alternaria alternata (Fr.) Keissler 7 & Coss (Mustaro) emend. Snyd.& Hans 7,8 ok niger van Tieghem 7 Macrophomina phaseolina (Tassi) Goid 8 chslera .australiensis (Bugni) Subram. Rhizoctonia solani Kuhn 7,8 . Jain ex M.B.Ellis 7 Fusarium solani (Mart) Appel & Wollenw. mend. Snyd. & Hans 7 17. Brassica oleracea L. var. Aspergillus flavus Link ex Gray 4 Aspergillus flavus Link ex Gray 4 capitata L. (CABBAGE) A. terreus Thom 4 A. terreus Thom 4 NVLSIWVd 40 SLNW1d GILVAILIND ONY G1IM 40 ISNN4 LOOY “TV 1] IHSIHND L701 TABLE 2. continued No and Host Rhizoplane Location Rhizosphere Location Chaetomium globosum Kunze ex Staud. 3 Chaetomium globosum Kunze ex Staud. 3 Cunninghamella echinulata (Thaxt.) Thaxt. 4 Cunninghamella echinulata (Thaxt.) Thaxt. Fusarium solani (Mart.) Appel & Wollenw. emend. Fusarium solani (Mart.) Appel & Wollenw. emend. Snyd. & Hans 3,4 Snyd. & Hans 4 Macrophomina phaseolina (Tassi) Goid. 3,4 MAC OPNOn ina phaseolina (Tassi) Goid. 3 Rhizoctonia solani Kuhn 3 Penicillium waksmanni Zaleski 4 18. Brassica rapa L. (Rapifera Curvalaria lunata (Wakker) Boedijn 7 ta (Fr.) Keissler 7 group) (Turnie) Fusarium. oxysporum Schlecht emend. 7 Fusarium solani (Mart) Appel & Wollenw. emend Hans. ans 7 ee Sa il & Wollenw.emend. lomyces varioti Bain 7 ses &H 7. Unidentified ie mycelium 7 Rhizoc on 7 Unidentified yeast 7 19. Raphanus sativus L. pes ea Fa Keissler 4 Alternaria ene (Fr.) Keissler 4 (RADISH) Aspergillus nidulans eel Winter 4 Aspergillus flavus Link ex Gray 6 ‘ ni Ge r ven Ti ie: 4 A.nidulans (Eidam) Winter 4 is (Bugni) Subram.& niger van Tieghem 6 Jain ex. M.B.Ellis 6 Cladosporium sp. 4 Fusarium solani (Mart.) Appel & Wollenw. Drechslera australiensis (Bugni) Subram.& emen yd.& H 46 Jain ex. M.B.Ellis 6 Macrophomina phaseolina (Tassi) Goid. 6 Fusarium oxysporum Schlecht. emend. Sn ans 4 Rhizopus stolonifer (Ehrenb. ex Link) Lind. 4 eblani (Mart.) aad & Wollenw.emend. Rhizoctonia solani Kuhn 6 Snyd. & Han 4 eee phaseolina (Tassi) Goid. 6 CUCURBITACEAE 20. Citrullus lanatus (Thunb,) Aspergillus nidulans (Eidam) Witner 5 Articulospora sp. 5 Matsum. & Nakai A. niger van Tieghem 5 Aspergillus flavus Link ex Gray 5 (WaTERMELON) Drechslera hawaniesis (Bugni) Subram. & Jain A. glaucus Link 5 8Z0L (2)L2 WaIs/D¥O'LINa Tase 2. continued No and Host Rhizoplane Location Rhizosphere Location ex M.B.Ellis. 5 A. nidulans (Eidam) Witner 5 Sp Schlecht.emend A. niger van Tieghem 5 .& Hans. 5 A. terreus Tho om 5 F solani (Mart.) Appel & Wollenw. emend. Ce ara um sp. 5 5 Dre ree ae iensis (Bugni) Subram. & Jain ex M.B. Ellis 5 cone solani (Mart.) Appel & Wollenw. emend. Snyd.& Han 5 Myrothecium cinctum Tode Paecilomyces lilacinus (Thom) Samson 5 Penicillium javanicum Van Beijma 5 Scopulariopsis brumptii Salvanet-Duval 5 ea viride Pers. ex Grey 5 21. Cucumis sativus L. Fusarium solani (Mart.) Appel & Wollenw. emend. nata (Fr.) Keissler 7 (CucumBer) Snyd. & Hans 7 Apert us niger van Tieghem 7 Rhizoctonia solani Kuhn 7 A. terreus Thom 7 Unidentified sterile fungus 7 Fusarium solani (Mart.) Appel & Wollenw. emend. : S 7 Unidentified sterile mycelium 7 2. Cucurbita moschata L. Chaetomium globosum Kunze ex. Staud. 7 Aspergillus flavus Link ex Gray 7 (PUMPKIN) Drechslera australiensis (Bugni) Subram. & Jain ex A. niger van Tieghem 7 M.B. Ellis 7 Chaetomium globosum Kunze ex. Staud. vs Fusarium solani (Mart.) Appel & Wollenw. emend Fusarium solani (Mart.) Appel & Wollenw. emend. Snyd. & Han 17 Snyd. & Hans Macrophomina phaseolina (Tassi) Goid. 1,7 Rhizoctonia solani Kuhn 1,7 23. Lagenaria siceraria (Mol.) Aspergillus flavus Link ex Gray 1,2,5 Aspergillus flavus Link ex Gray 125 Standl. (BoTTLe GourD) A. niger van Tieghem 12 A. niger van Tieghem 1,2,5 NVLSINWd 40 SLNWTd G3LVAILIND ONY Q1IM JO INN LOO “Tv 13 IHSaNND 6201 TABLE 2. continued No and Host Rhizoplane Location Rhizosphere Location wus mium indicum Corda 2 A. nidulans (Eidam) Winter 5 ta (Wakker) Boe ] A. sulphureus (Fres.) Thom. & Church 5 es la echinulata erties ) Thaxt. 5 A. terreus Tho 2,5 Fusarium solani (Mart.) Appel & Wollenw. emend. Fusarium moniliforme Sheld 2 Snyd. & Han 1,2,5 F. solani (Mart.) Appel & Wollenw.emend. Gera phaseolina (Tassi) Goid 3 nyd. & Hans 2:5 Rhizoctonia solani Kuhn 1 Myrothecium roridum Tode 5 Pa se eele mee ted (Thom) Samson 5 24. Luffa aegyptiaca Mill. Chaetomium indicum Corda 2,6 Aspergillus flavus Link ex Gray 2,5,6 (SPONGE ee See ieee (Wakker) Boedijn 2,6 A. fumigatus Fres. 5 alee australiensis (Bugni) Subram. & Jain A. nidulans sl Winter 2,5,6 Ellis 2,5,6 A. niger van Tie 2,5,6 a jum oe (Mart.) Appel & Wollenw. emend. A. terreus Thom 2,5,6 Snyd. & Hans 2,5,6 Chaetomium indicum Corda 5 Macrophomina phaseolina (Tassi) Goid. 2,6 ae la eine ata (Thaxt.) Thaxt. 5 Paecilomyces lilacinus (Thom) Samson 5 is (Bugni) Subram. & Jain Rhizopus stolonifer (Ehrenb. ex Link) Lind. 5 M.B. Ellis 5 Rhizoctonia solani Kuhn 2,6 Fusarium moniliforme Sheld 2 Unidentified white sterile mycelium 2,6 F. solani (Mart.) Appel & Wollenw. emend. Snyd. & Hans 2,5,6 ee ‘aecaaal ina (Tassi) Goid. 2,6 Pae inus (Thom) Samson o) Penicillium purpurogenum Stoll 2 Trichoderma viride Pers.ex Gray 2 Unidentified sclerotial fungus 2 Unidentified white sterile mycelium 6 Unidentified yeast 6 OfOL (2)L2 Ydls/94¥O'LIa TaB_e 2. continued No and Host Rhizoplane Location Rhizosphere Location 25. Momordica charantia L. Fusarium solani (Mart.) Appel & Wollenw. emend. Aspergillus candidus Link ex Link 2 (BITTER GOURD Snyd. & 2 A.niger van Tieghem 2 Macrophomina phaseolina (Tassi) Goid. 2 Chaetomium indicum Corda 2 Rhizoctonia solani Kuhn 2 Drechslera australiensis (Bugni) Subram & Jain ex M.B.Ellis 2 Fusarium solani (Mart.) Appel & Wollenw. emend. Sny S 2 Penicillium funiculosum Thom 2 CYPERACEAE 26. Cyperus rotundus L. Aspergillus flavus Link ex Gray 1,5 Alternaria alternata (Fr.) Keissler 5 A. niger van Tieghem 5 Aspergillus flavus bink ex Gray 1,5 Chaetomium globosum Kunze ex Staud i A.niger van Tieghem 5 Cunninghamella echinulata (Thaxt.) Thaxt. 5 A. terreus Thom 5 gas australiensis (Bugni) Subram. & Jain ex. Drechs| is (Bugni) Subram. & Jain ex. Ellis M.B. Ellis 5 aes solani (Mart.) Appel & Wollenw. emend. Fusarium solani (Mart.) Appel & Wollenw. emend & Hans. 5 Sn ans i) Trichoderma viride Pers. ex Gray 5 EUPHORBIACEAE 27. Euphorbia hirta L. Aspergillus niger van Tieghem 5 oe lus ue Link ex Gray 5 ally ng TO echinlat (Thaxt.) Thaxt. 5 A. niger m 5 liensis (Bugni) Subram. & Jain ex. A. terreus ain 5 B. Ellis 5 Fusarium solani (Mart.) Appel & Wollenw. emen d.& Hans. 5 Macrophomina oo na (Tassi) Goid. 5 Penicillium purpurogenum Stoll 5 NVLSINVd JO SLNW1d G3LVAILIND ONY TIM 40 INNS LOOY “TV 13 1HSIYND LEOL TasLe 2. continued No and Host Rhizoplane Location Rhizosphere Location FABACEAE 28. Alhaji murorum Medik Drechslera halodes (Drechlera) Subram. & Jain 7 Aspergillus flavus Link ex Gray 7,8 Fusarium solani (Mart) Appel & Wollenw. A. niger van Tieghem 7,8 emend. Snyd.& Hans 7,8 29, Arachis hypogaea L. Chaetomium indicum Corda 1 Aspergillus flavus Link ex Gray 1 (PEANUT) urvalaria clavata (Wakker) Boedijn 1 A. nidulans (Eidam) Winter ] Drechslera australi (Bugni) Subram. A.niger van Tieghem 1 & Jain ex M.B. Ells, 1 A. terreus Thom ] Fusarium oxysporum Schlecht emend. Snyd. Fusari Y5D 1 Schlecht emend.. Wollenw 1 Snyd.& Wollenw ] F. solani (Mart.) aan & Wollenw.emend. F.solani (Mart.) Appel & Wollenw.emend Snyd. & Han 1 Snyd. & Hans 1 a scrophemina iad ina (Tassi) Goid. 1 Penicillium rugulosum Thom 2 ae (Berk. & Br.) Petch 1 Unidentified yellow sterile mycelium 2 Rhi Yee onal solani h ] Unidentified white sterile mycelium ] Unidentified yellow sterile mycelium 2 Unidentified white sterile mycelium ] 30. Cicer arietinum L. nel ium we dasa sum Kunze ex Steud. 8 Aspergillus oo van Tieghem 8 (CHICKPEA) is (Bugni) Subram. & Jain A. terreus Tho 8 ex M.B. Ellis. 8 Sic um globes Kunze ex Steud. 8 D. halodes (Drechslera) Subram. & Jain 8 (Bugni) Subram. & Jain Fusarium solani (Mart.) Appel & Wollenw. emend. B. Ellis Snyd. & Hanse 8 meee solani (Mart.) Appel & Wollenw. emend. Macrophomina phaseolina (Tassi ) Goid. 8 n 8 31. Cyamopsis tetragonoloba aie giol Cae Kunze ex Staud. 1 Alt ia alt ta (Fr.) Keissler 6 (L.) Taub. (Guar) is (Bugni) Subram. & Jain ex i lus flavus Link ex Gray 6 M.B. Ellis 1 niger van Tieghe 6 ZE0L (712 VaIs/¥O'LINS Taste 2. continued No and Host Rhizoplane Location Rhizosphere Location Fusarium semitectum Berk. & Rav. 6 Chaetomium globosum Kunze ex Staud. 6 F. solani (Mart.) Appel & Wollenw. emend. Curvularia clavata Jai 6 Snyd. & Hans 1,6 Fusarium semitectum Cees & Rav. 6 Macrophomina mneeeenae (Tassi) Goid. ] F. solani (Mart.) Appel & Wollenw.emend Rhizoctonia solani n 1,4,6 Snyd. & Hans 6 Unidentified white sterile mycelium 6 Penicilli itri Thom. 6 Trichoderma viride Pers. ex Gra 6 Unidentified white sterile mycelium 6 Unidentified yea 6 32. Glycine max (L.) Merr. Alternaria alternata (Fr.) Keissler 1,8 Aspergillus flavus a ex Gray 1,8 (SOYBEAN) Aspergillus candidus Link ] A. niger van Tieghem 1,8 A. flavus Link ex Gray 1,8 A. terreus Thom 1,8 A.niger van Tieghem 1,8 Curvularia lunata (Wakker) Boedijn 8 A. terreus Thom 1,8 Drechslera australiensis (Bugni) Subram, & Jain Curvularia lunata (Wakker) Boedijn 8 ex M.B. Ellis 1,8 Drechslera australiensis (Bugni) Subram. & Jain Fusarium solani (Mart.) Appel & Wollenw. emend. M.B. Ellis 1,8 Snyd. & Hans 1,8 Fusarium solani (Mart.) Appel & Wollenw. emend. Macrophomina phaseolina (Tassi) Goid. 1,8 Snyd. & Hans 1,8 Monodictys putredinis (Wallr.) Hughes ] wea annlig Eee ee (Tassi) Goid., 1,8 Myrothecium roridum Tode 1 m Tod ] Nigrospora oryzae (Berk. & Br.) Petch ] Biepctonia solani Kuhn 1,8 Penicillium purpurogenum Stoll 1 Trichoderma harzianum Rifai 1 Unidentified Basidomycetes 1 33. Lens culinaris Medic Alternaria alternata (Fr.) Keissler 1,8 [ternaria alternata (Fr.) Keissler 2 (LENTIL) Curvularia lunata (Wakker) Boedijn 8 Aspergillus flavus Link ex Gray 1,8 Drechslera australiensis (Bugni) Subram. & Jain A.niger van Tieghem 1,8 ex M.B. Ellis. 8 A. terreus Thom 8 NWISINWd 40 SLNV1d GLVAILIND GNY GTIM JO IDNN4 LOOY “TV 13 IHS3NND eeOl TABLE 2. Continued No and Host Rhizoplane Location Rhizosphere Location D. halodes (Drechslera) Subram. & Jain ex Fusarium solani (Mart.) Appel & Wollenw. emend. M.B. Ellis 8 Snyd. & Hans 1,8 Fusarium oxysporum Schlecht emend. Unidentified Basidomycetes 1 Hans. 8 F solani Hine Appel & Wollenw.emend d.& Han 1,8 oo phaseolina (Tassi) Goid. 1,8 Rhizoctonia solani Kuhn Stachybotrys atra Cor 8 Unidentified Basidiomycetes ] 34. Leucaena leucocephala Aspergillus flavus Link ex Gray 5 Aspergillus flavus Link ex Gray 5 (Lam.) de Wit A. terreus Thom 5 A.niger van Tieghem 5 (IPiL-iPiL) Drechslera liensis (Bugni) Subram. & Jain A. nidulans (Eidam) Winter 5 Ellis A. SU ew (Fres.) Thom. & Church. 5 on solani (Mart.) Appel & Wollenw. emend. A. sales 5 Snyd. & Hans 5 D lera australiensis (Bugni) Subram. & Jain Helicocephalum sp. 5 ex in B. Ellis 5 35. Medicag tiva L. Alternaria alternata (Fr.) Keissler 5,6 Alternaria alternata (Fr.) Keissler 5,6 (ALFALFA) fins illus flavus Link ex Gray 5 Aspergillus flavus Link ex Gray 3,5,6 A, ue Tieghem 5 A.niger van Tieghem 3,5,6 A. ter ane 5 A. terreus Tho 5,6 cunalari ee (Wakker) Boedijn 6 Fusarium semitectum Berk & Rav. 6 li (Bugni) Subram. & Jain F. solani (Mart.) Appel & Wollenw. emend. ex M.B. Ellis 3,6 nyd. & Hans 3,6 Fusarium semitectum Berk & Rav. 6 Macrophomina phaseolina (Tassi) Goid. 3,6 F. solani (Mart.) Appel & Wollenw. emend. Unidentified sclerotial fungus 3,6 Snyd. & Hans 5,6 Unidentified white sterile mycelium 3 vEOL (2)LZ YaIs/9¥O'LIYS TasB_e 2. continued No and Host Rhizoplane Location Rhizosphere Location Macrophomina phaseolina (Tassi) Goid. 3,6 Rhizoctonia solani Kuhn 3,6 Unidentified sclerotial fungus 3,6 36. Melilotus alba Medik. Alternaria alternata (Fr.) Keissler 2 Alternaria alternata (Fr.) Keissler 2 (WHITE SWEET-CLOVER) Aspergillus flavus Link ex Gray 2 Aspergillus flavus Link ex Gray 5 A. fumigatus Fres 2 A. niger van a 25 A. niger van Tie 2 A. terreus Tho 2 A.nidulans (Eidam) Winter 2 cepholesparm 2 Chaetomium flavum Ornvik 2 Chaetomium Aa Omvik 2 Drechslera Genta dee (Bugni) Subram & ete hee ae (Bugni) Subram & Jaine Ellis. 25 Jain ex M.B.Ellis. 5 ees ee Schlecht.emend. Fusarium a (Mart.) Appel & Wollenw. emend. Snyd.& H 2 Snyd. & Hans 2,5 F. solani (Mart) Appel & Wollenw. emend. F oxysporum Schlecht. emend. Snyd. & Hans 2 Snyd. & Han 2,5 37. eae vulgaris L. Alternaria alternata (Fr.) Keissler 46 Alternaria alt ta (Fr.) Keissler 4,6 (COMMON BEAN) Sai niger van Tieghem 4 Aspergillus niger van Tieghem 46 A. terreus Thom 6 A. se ve ) Thom. & Church. 6 Drehe australiensis (Bugni) Subram. & Jain A. terreu 6 B. Ellis Clados Sp. 4 ie solani (Mart.) Appel & Wollenw. emend. Deca eae s (Bugni) Subram. & Jain Snyd. & Hans 4,6 B. Ellis 6 Macrophomina phaseolina (Tassi) Goid. es solani oe ) Appel & Wollenw. emend. Rhizoctonia solani Kuhn 46 Snyd. & H 46 acro seas phaseolina (Tassi) Goid. 6 Rhizopus stolonifer (Ehrenb. ex Link) Lind. 4 NVLSINWd 40 SLNW1d G31VAILIND ONY GTIM 40 ISNNd LOOY “TY 13 IHSIHND SEol TaB_e 2. continued No and Host Rhizoplane Location Rhizosphere Location Scopulariopsis Brumptii Salvanet-Duval 4 Trichoderma aid jianum Rifai 6 T. koningii Oudem 6 38. Pisum sativum L. (Pea) Alternaria alternata (Fr.) Keissler 8 Alt ia alt ta (Fr.) Keissler 8 Chaetomi lob aud 8 pide us ieee Link ex Gray 8 piele dts australiensis com ae & Jain A. niger Vv 8 Ellis. 8 een pean Kunze ex. Staud 8 Pike (Drechslera) Subram. & Jain 8 Fusarium solani (Mart.) Appel & Wollenw. emend. Fusarium solani (Mart.) Appel & Wollenw. emend. 8 Snyd. & Hans 8 Macrophomina phaseolina (Tassi) Goid. 8 39. Sesbania sesban (L.) Merr. Aspergillus niger van Tieghem 3 Aspergillus flavus Link ex Gray 3 (EUROPEAN RIVER ee) Fusarium solani (Mart.) Appel & Wollenw. emend. A. niger Vv. 3 3 A. terreus Thom 3 Unidentified yellow sterile mycelium 3 Fusarium monoliforme Sheld. 3 Unidentified white sterile mycelium 3 F. solani (Mart.) Appel & Wollenw. emend. Snyd. & Hans 3 Macrophomina phaseolina (Tassi) Goid 3 Unidentified yellow sterile mycelium 3 pgstaes white sterile mycelium 3 40. Trifolium alexandrium (L.) Alternaria alternata (Fr.) Keissler 8 .) Keissler 8 Spergue ex. Turrill Chaetomium globosum Kunze ex. Staud. 8 nes illus flavus onl ex Gray 5,8 (Clover) alle unane emul Boedijn 8 A. niger van Tieghem 5,8 (Bugni) Subram. & A. terreus Thom 5 Jain ex M.B.Ellis 5,8 Drechslera australiensis (Bugni) Subram. & Fusarium solani (Mart.) Appel & Wollenw. emend. Jain ex M.B.Ellis. 5 Snyd. & Hans 5,8 Fusarium solani (Mart.) Appel & Wollenw. emend. 9E0L (2)L@ vdls/9¥O'LINd Tase 2. continued No and Host Rhizoplane Location Rhizosphere Location Macrophomina phaseolina (Tassi) Goid. 5,8 Snyd. & Hans 5,8 Rhizoctonia solani Kuhn 5,8 Macrophomina phaseolina (Tassi) Goid. 5,8 41. Trigonella foenum-graecum _ Alternaria alternata (Fr.) Keissler 2,7,8 Alternaria alternata (Fr.) Keissler 2 L. (FENUGREEK) Fusarium solani (Mart.) Appel & Wollenw. emend. Aspergillus flavus Link ex Gray 2,8 nyd. & Hans 2,8 A. niger van Tieghem 2,8 Macrophomina phaseolina (Tassi) Goid. 2,8 A. terreus Thom 8 Rhizoctonia solani Kuhn 2 Fusarium solani (Mart.) Appel & Wollenw. emend. Snyd.& H 2,8 42. Vigna mungo (L.) Hepper Curvularia lunata (Wakker) Boedijn 1 Aspergillus in Link ex Gray 1 (UID BEAN) Drechslera australiensis (Bugni) Subram. & A. niger van Tieghe 1 Jain ex M.B.Ellis. ] . com 1 an solani (Mart.) Appel & Wollenw. emend. lera australiensis (Bugni) Subram. & Snyd. & Hans. ~ Sain ex M.B.Ellis. ] Macrophomina tee (Tassi) Goid. Fusarium solani (Mart.) Appel & Wollenw. emend. Snyd. & Hans. 1 1 a izoctonia solani ] ] 43. Vigna radiata (L.) Wilezek A| ia alternat i .) Keissler 1,8 Alternaria alternata (Fr.) Keissler ] (MUNG BEAN) mapeigiis mies van Tieghem ] Aspergillus flavus Link ex Gray 1,8 Cha m Kunze ex Staud. 1,8 A. fumigatus Fre 1 ee lunata (Wakker) cin 8 A. nidulans en Winter 1 Drechslera nsis (Bugni) Subram. & A. niger van Tieghem 1 Jain ex M.B.Ellis. 1,8 A. terreus Thom 1,8 D.halodes (Drechslera) Subram. & Jain 8 Chaetomium globosum Kunze ex Staud. 1,8 Fusarium moniliforme Sheld. 8 Fusarium solani (Mart.) Appel & Wollenw. emend. Fsolani (Mart.) Appel & Wollenw.emend. Snyd. & Hans 18 Snyd. & Hans 1,8 Penicillium purpurogenum Stoll ] Macrophomina phaseolina (Tassi) Goid. 18 Macrophomina phaseolina (Tassi) Goid. 18 Rhizoctonia solani Kuhn 1,8 Myrothecium roridum Tode ] NVISINVd JO SLNV1d GILVAILIND ONY G1IM 40 IDNNJ LOOY “TV 13 IHSIYND LEOL Tasle 2. continued No and Host Rhizoplane Rhizosphere Location Unidentified Ascomycetes Unidentified white sterile mycelium 2 Unidentified white sterile mycelium GRAMINEAE 44, Avena sativa L. (Oar) Drechslera australiensis (Bugni) Subram. & Jain — illus nie Link ex Gray 5 A. niger 5 Fusarium solani (Mart.) Appel & Wollenw. ne era epee iensis (Bugni) Subram. emend. Snyd. & Hans io Jain ex M.B.Ellis 5 Rhizoctonia solani Kuhn 5 Fusarium solani (Mart.) Appel & Wollenw. emend. Snyd. & Hans 5 45. Cenchrno setigerus Vahl Aspergillus flavus Link ex Gray 5 Aspergillus flavus Link ex Gray 5 A. niger van Tieghem S A. niger van Tieghem 5 A. terreus Thom 5 A. terreus Thom 5 Cunninghamella echinulata (Thaxt.) Thaxt. 5 Cunninghamella echinulata (Thaxt.) Thaxt. 5 Fusarium solani (Mart.) Appel & Wollenw. emend. Curvularia clavata Jain 5 Snyd. & Hans 2 Paecilomyces lilacinus oe Samson 5 46. Cynodon dactylon (L.) Pers. Aspergillus flavus Link ex Gray 2 Aspergillus candidus 5 (BERMUDA GRASS) hem 2 A. flavus Link ex — 1,2,5 A. terreus Thom 2 A.niger van Tieghem 1,2,5 ees lunata (Wakker) Boedijn ] A. nidulans (Eidam) Winter 2 Drechslera australiensis(Bugni) Subram. & Jain ex. A. terreus Thom 2,5 M.B.Ellis Cephalosporium s 2 Fusarium proliferatum (Matsushima) Nirenberg Chaetomium flavum Omvik 5 F. solani (Mart.) Appel & Wollenw. emend. Cunninghamella echinulata (Thaxt.) Thaxt. 2,5 Snyd. & Hanse 2 Curvularia lunata (Wakker) Boedijn ] ete ieee phaseolina (Tassi) Goid 2 Drechslera australiensis (Bugni) Subram. & (Berk.& Br.) Petch 1 Jain ex.M.B.Ellis 1 Biizonin olor fer (ENrenb. ex Link) Lind. 2 Fusarium proliferatum (Matsushima) Nirenberg 5 8E0L (Z7)LZ VaIS/D¥O'LINa Taste 2. continued No and Host Rhizoplane Location Rhizosphere Location F. solani (Mart.) Appel & Wollenw. emend Snyd. & Hans 5 He aliecede roridum Tode 5 Sona yee (Berk.& Br.) Petch 1 nus (Thom) Samson 5 izopus stolonifer (Ehrenb. ex Link) Lind. 5 47. Oryzae sativa L. (Rick) Chaetomium globosum Kunze ex. Staud, 7 Aspergillus flavus Link ex Gray 7 Curvularia lunata (Wakker) Boedijn 7 A. niger van Tieghem 7 rechslera hawaliensis (Bugni) Subram. & Jain ex Chaetomium globosum Kunze ex. Staud. 7 M.B. Ellis Fusarium solani (Mart.) Appel & Wollenw. emend. Fusarium solani (Mart.) Appel & Wollenw. Emend. Snyd. & Hans 7 Snyd. & H 7 uns na phaseolina (Tassi) Goid. 7 48. Pennisetum americanum (L.) Alt ata (Fr.) Keissler 6 mena alternata (Fr.) Keissler 6 Leeve (MILLET) Aspergillus niger van Tieghem 5 andidus Link ex Link 2 A. nidulans (Eidam) Winter 5 A. pvas Link ex Gray 2,5,6 A. terreus Thom 5 A.niger van Tieghem 2,5 Drechslera australiensis (Bugni) Subram. & Jain A. cae inet 5,6 B. Ellis D liensis (Bugni) Subram. & Jain D. hawaiiensis (Bugni) Subram. & Jain ex. M. B. Ellis 2 M.B. Ellis. 5 Fusarium iliform oe 2 Fusarium solani (Mart.) Appel & Wollenw. emend. F.semitectum Berk. & 6 Snyd. & Hans 2,6 F. solani (Mart.) = een emend Macrophomina A ina (Tassi) Goid. 1,2,6 Snyd. & Hans 1,2,6 Rhizoctonia solani 2,6 Macrophomina phaseolina (Tassi) Goid. 1,2,6 Unidentified yellow caus mycelium 1 Monodictys putredinis (Wallr.) Hughes 1 Penicillium rugulosum Thom 6 NVISINWd 40 SLNW1d Q3LVAILIND GNV G1IM 40 IONNd LOOY “TV 13 1HSIYND 6£01L TABLE 2. Continued OrOL No and Host Rhizoplane Location Rhizosphere Location Unidentified Ascomycetes 2 Unidentified sclerotial fungus 1,2,6 Unidentified yellow sterile mycelium 1 enlcenunles white sterile ae 1 49. Setaria verticilata Alternaria alternata (Fr.) Keissler 14 ta (Fr.) Keiss 1,4 (L.) Beauv. Aspergillus flavus Link ex Gray 4 sper flavus Link ex | 4 HOOKED BRISTLE GRASS) A.niger van Tieghem 4 A. terreu om 14 A. terreus Tham 14 A. nidulans am Winter 4 Cladosp ] Cladosporium sp. 1 cura ae Jain 4 Drechslera australiensis (Bugni) Subram. & Jain traliensis (Bugni) Subram. & Jain ex M.B. Ellis ex M.B. Ellis 4 Fusarium solani (Mart.) Appel & Wollenw. emend. Rhizopus stolonifer (Ehrenb. ex Link) Lind 1 Snyd. & Hans 4 Myroth m (Corda) Sacc. 1 50. Sorghum bicolor Se Hs ao Jain 3 Aspergil us feuns ie ex Gra 3 (L.) Moench. is (Bugni) Subram. & A. sulphureus (Fres.) Thom. & Church 3 (SORGHUM) Jain ex M.B. Ellis. 3 Curvularia clavata Jain 3 Fusarium lon: (Mart.) Appel & Wollenw. emend. Drechslera. australiensis (Bugni) Subram. & Jain Snyd. & Hans 3 M.B.Ellis. hese ain na se (Tassi) Goid. 3 Fusarium solani (Mart.) Appel & Wollenw. emend. ta (Riv.) Galloway 3 Snyd. & Hans 3 ee phaseoli na (Tassi) Goid. 3 ta (Riv.) Galloway 3 ices white sterile mycelium I 51. Triticum aestivum L. Alternaria alternata (Fr.) Keissler 5,6 ta (Fr.) Keissler 6 (WHEAT) Chaetomium globosum Kunze ex. Staud. 5,6 Aspergillus avis Link ex Gray 5,6 Curvularia lunata (Wakker) Boedijn 6 A. niger van Tieghem 5,6 (Z7)LZ VIS/D¥O'LIYE Rhizoplane Rhizosphere Location Drechslera australiensis (Bugni) Subram. & A. terreus Thom 5 Jain ex MB. ele See On giopenun Kunze ex. Staud. 6 Fusarium Sheld. Sheld. 6 F. solani (Mart.) Appel & Wollenw. emend. F solani (Mart.) oe & Wollenw. emend. S Hans Snyd.& Han 5,6 Unidentified white sterile mycelium Penicillium ie um Thom 6 Unidentified white sterile mycelium 5 at Alternaria alternata (Fr.) Keissler Alternaria alternata (Fr.) Keissler 2,4,6 Aspergillus flavus Link ex Gray Aspergillus flavus Link ex Gray 1,2,4,6 A. niger van Tie A. fumigatus Fr 1,2 A. terreus Thom A.nidulans (Eidam) Win 1 Cunninghamella echinulata (Thaxt.) Thaxt. A. ee (Fres.) see & Church ] Drechslera australiensis (Bugni) Subram. & Jain A. terre m 2,4,6 x MB. Ellis eae sp. 2,4 e halode (Drechslera) Subram. & Jain haetomium indicum Corda 2 isarium moniliforme Sheld. Curvularia lunata (Wakker) Boedijn 4 ee Schlecht. emend. Snyd. & Hans Drechslera australiensis (Bugni) Subram.& Jain F. solani (Mart.) Appel & Wollenw.emend. ex M.B. Ellis 2 yd. & Han Fusarium life Sheld. 2 Macrophomina phaseolina (Tassi) Goid. F. oxysporum Schlecht. emend. Snyd. & Hans 2,4 Rhizoctonia solani Kuhn F. solani (Mart.) Appel & Wollenw. emend. Rhizopus stolonifer (Enrenb. ex Link) Lind. S 1,2,4,6 Macrophomina phaseolina (Tassi) Goid. ] Penicillium purpurogenum Stoll ] Rhizopus stolonifer (Ehrenb. ex Link) Lind. 4 Stachybotry parvispora Hughes 6 Unidentified yeast 1,4 NVLSINWd 40 SLNW1d G3LVAILIND ONY GTIM 4O ISNNd LOOY “TV 13 1HS4YND LvOL Tas_e 2. continued No and Host Rhizoplane Location Rhizosphere Location MALVACEAE 53. Abelmosch lentus (L.) Alternaria alternata (Fr.) Keissler 8 Alt ia alt (Fr.) Keissler 7 Moench (Okra) Drechslera ce fensis (Bugni) Subram. Aspergillus flavus Link ex Gray 1,7,8 Jaine Ellis. 7 A. abit eghem 1,78 Fusarium a (wart) Appl & Wollenw. A. terreus 1,8 Snyd. & Han 78 Drechslera lene (Bugni) Subram. ophomina pee ind (Tassi) Goid. 1,7 M.B.Ellis. 7 ace solani Kuhn 7,8 Fusarium solani (Mart.)Appl & Wollenw. : . ans 1,7,8 54. Abutilon indicum (L.) Sweet — Alternaria alternata (Fr.) Keissler 2 Alternaria alternata (Fr.) Keissler 2 (MONKEY BUSH) Aspergillus flavus Link ex Gray 2 Aspergillus flavus Link ex Gray 2 ‘i niger van Tieghem 2 A. nidulans (Eidam) Winter 2 A. terreus Thom 2 A. niger van Tieghem 2 Fusarium solani (Mart.) Appel & Wollenw. emend. A. terreus Thom 2 Snyd. & Hans 2 Cephalosporium sp. 2 Rhizopus stolonifer (Enrenb. ex Link) Lind. 2 Fusarium solani ie Appel & Wollenw. emend. Snyd.& Han Macrophomina sheet lina (Tassi) Goid. 2 55. Gossypium arboreum L. Alternaria alternata (Fr.) Keissler 1,8 Aspergillus flavus Link ex Gray 1,8 (Corton) Chaetomium globosum Kunze ex. Staud. 1 A. niger van Tieghem 18 eee ae cote! Boedijn ] e terreus Thom 1,8 traliensis (Bugni) Subram.& Jain ex n Schlecht emend. Snyd.& Hans 1 M.B. Ellis, 1 F solani (Mart) Ape & Wollenw.emend. Fusarium solani (Mart.) Appel & Wollenw. emend. S 1,8 nyd. & Hans 1,8 Benicili ium eae Zukal ] Macrophomina phaseolina (Tassi) Goid 8 Trichoderma viride Pers. ex Gray 1 vee solani Kuhn 1,8 Unidentified sterile fungus 2 1 1 ichoderma viride Pers. ex Gray Unidentified yeast 7r0L (2)LZ VdIS/DYO'LIYS TABLE 2. continued No and Host Rhizoplane Location Rhizosphere Location PEDELIACEAE 56. Sesamum indicum L. Aspergillus niger van Tieghem 5 Aspergillus candidus Link a AME) Curvularia lunata (Wakker) Boedijn 1 A.flavus Link ex Gra | Fe ae echinulata (Thaxt.) Thaxt 5 A.nidulans (Eidam) Winter ] rechslera australiensis (Bugni) Subram. & A.niger van Tieghem 5 ae ex MB. Ellis. 1 A. eae one & Church. 5 Fusarium solani (Mart.) Appel & Wollenw. emend. ‘ ieee 5 Snyd. & Hans 1 traliensis (Bugni) Subram. & Rhizoctonia solani Kuhn 1 Jain ex M.B.Ellis. Fusarium solani (Mart.) Appel & Wollenw. emend. onyd. & Hans ] Paecilomyces lilacinus (Thom.) Samson 5 Penicillium purpurogenum Stoll ] Rhizoctonia solani Kuhn ] Unidentified white sterile mycelium 1,2 PIPERACEAE 57. Piper betel L. (BETEL) Alternaria alternata st Keissler 2 Alternaria alternata (Fr.) Keissler 2 Chaetomium globosum Kunze ex. Staud 2 pe us flavus Link ex Gray 2 ee lunata a (Wake Boedijn 2 pales, van Tie oy 2 is (Bugni) Subram. & Jain nze ex. Staud 2 ex M.B. Ellis 2 Drechslera wisualense ees Subram. & Jain Fusarium Pail iforme Sheld 2 M.B. Ellis. 2 F oxysporum Schlecht emend. Snyd & Hans. 2 nee eld 2 F. semitectum Berk. & 2 F oxysporum Schec Bead Snyd & Hans. 2 F. solani (Mart.) eee & lean emend. F. semitectum Ber 2 Snyd. & Han 2 F. solani (Mart.) ae aialeni emend. Macrophomina Cent (Tassi) Goid. 2 Snyd. & Hans 2 Rhizoctonia solani Kuhn 2 NVLSIN¥d 40 SINVId G3LVAILIND ONY G1IM 40 ISNN LOOY “Ty 13 IHSaNNd fp0L TABLE 2. Continued No and Host Rhizoplane Location Rhizosphere Location SOLANACEAE ~Capsicum annuum L. Alternaria alternata (Fr.) Keissler 1,2,6,7 Alt It ta (Fr.) Keissler 1,2,6,7 (PEPPER) Aspergillus flavus Link ex Gray 2 pec lus flavus Link ex Gray 2 A. niger van Tie 1,2,6,7 A. niger 1,2,6,7 A. nidulans es Wie 1,2,6 A. a ans ea cies 1,2,6 A. terreus Thom 2 A. terreus Thom 2 Chaetomium indicum Corda 2 Cephalosporium sp. 2 Curvularia clavata Jain 2 Chaetomium indicum Co 2 2 ners ee Boedijn 7 Drechslera australiensis (Bugni) Subram. & Jain is (Bugni) Subram. & Jain ex MB a 2 ex M.B. Ellis 2 Fu n (W.G.Sm.) Sacc. 2 Fusarium culmurum (W.G. Sm.) Sacc. 2 F. solani Nt 098 & Wollenw.emend Foxysporum Schlecht emend. Snyd. & Hans 2,7 Snyd.& H 1,2,6,7 F solani (Mart.) ee & Wollenw.emend. Macrophomina ieee ina (Tassi) Goid. 2 1,2,6,7 — Penicillium funiculosum Thom ] ieee phaseolina (Tassi) Goid. 1,2 é ae osum om 2,6 Rhizoctonia solani Kuhn 1,2 rzial Rifai 2 Trichoderma viride Pers. ex Gray 6 Unidentified bck storie mycelium 1,2 Unidentified black sterile mycelium 1,2 9. Capsicum annuum L. var. hates alternata (Fr.) Keissler 2 Alternariaa [ternata (Fr.) Keissler 7 shimla (Pepper) spergillus niger van Tieghem 1,2,6 Aspergillus flavus Link ex Gray 1,2 i‘ terreus Thom 1,2 A. niger van Tieghem 2,7 Fusarium moniliforme Sheld. 1,2,7 A. terreus Thom 1 F solani (Mart.) Appel & Wollenw. emend. Drechs| traliensis (Bugni) Subram. & Jain Snyd. & Hans 1,2,6,7 ex M.B. Ellis 2 Trichoderma viride Pers. ex Gray Fusarium ilif Sheld 2 Prol (2)L2 VaIS/¥O'LINa Taste 2. continued No and Host Rhizoplane Location Rhizosphere Location Unidentified scleotial fungus ] F. solani (Mart.) Appel & Wollenw.emend Snyd. & Hans 1,2:6,7 Penicillium funicul Thom 1; Tri eae viride Pers. ex Gray 2 Unidentified sclerotial fungus 1,2 60. 5 ye icon esculentum Alternaria alternata (Fr.) Keissler 12:5 Alternaria alternata (Fr.) Keissler 15 Mill. (Tomato) Aspergillus ce van Tieghem ] bce niger van Tieghem ] Cladosporium sp. ] _candidus Link ex Link ] cunnlria ena wae) Boedijn 1,2 flavus Link ex Gray 1,2:5;7 lera aust is (Bugni) Subram. & Jain A.niger van Tieghem 1,2,5 ex MB. Ellis 1 A. terreus Thom ] Fusarium moniliforme Sheld. 2 Cladosporium sp. ] Foxysporum Schlecht. emend. Snyd. & Hans. 1,2 Drechslera liensis (Bugni) Subram. & Jain F. solani (Mart.) Appel & Wollenw. emend. ex MB. Ellis 1 1 F Schlecht emend.snyd.&Hans 1 Humicola grisea Traaen 5 F. solani (Mart) Appel & Wollenw. emen Macrophomina ee ee (Tassi) Goid. ] Snyd.& Hans 1 Rhizoctonia solani 1,2,5,7. Macrophomina ete (Tassi) Goid. ] Unidentified sclerotial fu ngus 1,2 Stachybotrys atra Cor ] Unidentified sclerotial fran G2 61. Solanum melongena L. Alternaria alternata (Fr.) Keissler 4 Aspergillus flavus Link ex Gray 1,2,4,6 (EGG PLANT) Aspergillus flavus Link ex Gray 4 A. fumigatus Fres 1,4 A. niger van Tieghem 4 A.niger van Tieghem 1,2,6 A. terreus Thom 4 ‘a terreus Thom 4 Fusari YS Schlecht.emend. Schlecht.emend. Snyd.& Hans 1,2 .& Hans 1,26 F soln (Mart) eos & Wollenw.emend. F. solani (Mart.) Appel & Wollenw. emend. nyd. & Han 4 Snyd. & Hans 4 eee ina phaseolina (Tassi) Goid. 4 NVLSINVd 40 SLNV1d G3LVALLIND ONY G1IM 40 INN LOOY “TW 13 1HSIYND SPOL Table 2. continued No and Host Rhizoplane Location Rhizosphere Location MOOR RONING phaseolina (Tassi) Goid. 1,2,4,6 Myrothecium roridum 4 lani Kuhn 1,2,4,6 Rhizoctonia solani Kuhn 1,2 see am chlomydosporium Goddar 1 62. Solanum nigrum L. (Bugni) ae & Aspergillus flavus Link ex Gray 2 (BLACK NIGHTSHADE) Jain ex M.B.Ellis. 2 — van Tieghem 2 Fusarium solani ia Appel & Wollenw. emend. hsl traliensis (Bugni) Subram. & Be &H 2 Jain ex M.B.Ellis 1 Kuhn 2 Fusarium solani (Mart) Appel & Wollenw. emend Snyd. & Hans 2 Rhizoctonia solani Kuhn 2 63. Solanum surrenttense edd us flavus Link ex Gray 5 Alternaria alternata (Fr.) Keissler 5 Burm. f rum Schlecht, emend., Aspergillus flavus Link ex Gray 5 Snyd. & bane 5 A.nidulans (Eidam) Winter 5 F. solani (Mart.) Appel & Wollenw.emend. A. niger van Tieghem 5 Snyd. & Hans 5 A. sulphureus (Fres.) Thom. & Church 5 Macrophomina phaseolina (Tassi) Goid. 5 A. terreus Thom 5 Chaetomium flavum Omvik 5 Fusarium solani (Mart.) Appel & Wollenw. emend. Snyd. & Hans 5 Myrothecium roridum Tode 5 Penicillium raistrickii Gilman & Abbott 5 UMBELLIFERAE 64. Coriandrum sativum L. Aspergillus flavus Link ex Gray 2 Alternaria alternata (Fr.) Keissler 7 (CORIANDER) A. niger van Tie 2,7 Aspergillus flavus Link ex Gray 2 Curvularia ste (Wakken Boedijn 2 A. niger van Tieghem 2,7 Bugni) Subram. & Jain ex A. nidulans (Eidan) Winter 2 M.B. Ellis 2 A. terreus Thom 2 (2)LZ YaIS/D4O'LINA 9P0L Tasie 2. continued No and Host Rhizoplane Location Rhizosphere Location 65. Daucus carota L. ssp. sativus (Haffm.) Acrang (Carrot ) D.halodes (Drechslera) Subram. & Jain ex M.B. Ellis. Fusarium solani (Mart.) Appel & Wollenw. emend. Snyd. & Hans Macrophomina phaseolina (Tassi) Goid. Unidentified white sterile mycelium Fusarium solani (Mart.) Appel & Wollenw. emend. Snyd. & Hans Fusarium solani (Mart.) Appel & Wollenw. emend. Macrophomina phaseolina (Tassi) Goid. Stachybotrys parvispora Hughes Unidentified sclerotial fungus Alt ia alt ta (Fr.) Keissler Aspergillus flavus Link ex Gray A. niger van Tie A. nidulans (Eidam) Winter 2 a 5 Fusarium semitectum Berk. & Rav. F. solani (Mart.) Appel & Wollenw. Emend Snyd. & Hans F.oxysporum Schlecht. emend. Snyd. & Hans Macrophomina phaseolina (Tassi) Goid Ctarhyvhot — atra rda Unidentified sterile fungus Unidentified yeast NON OM ON RN 1 = Karachi University Campus, 2 = Memon Goth, 3 = Darsano Chano, 4 = Shah Faisal Colony, 5 = Hub, 6 = Kathor, 7 = Gharo, 8 = Thatta NVISINWd 40 SLNV1d G3LVAILIND ONY G1IM 40 INN LOOY “TV 13 IHSINND Lv0L 1048 BRIT.ORG/SIDA 21(2) with additional roots until the total volume of soil and water become 20 mL, assumed to be 1:10 dilution. From this other dilutions 1:100, 1:1000, 1:10,000, 1:100,000 were made. One mL aliquot from each of the last two highest dilu- tions were poured in sterilized petri plate containing potato dextrose agar (PDA) incorporated with penicillin (00,000 units/L) and streptomycin (0.2 g/L) to check bacterial growth. Plates were incubated for 5 days at room temperature (25-30° C) under 12 hours light and dark conditions. Fungi that grew and sporu- lated on plates were identified with reference to Barnett and Hunter (1998), Dix and Webster (1995), Domsch et al. (1993) and Nelson et al. (1983). Special atten- tion was paid towards the occurrence of root infecting and well known biocontrol agents like Trichoderma, Paecoilomyces lilacinus, Verticillium chlamydosporium. Fungi that did not show morphological characters were in- cluded as unidentified. Isolation of Fungi from Rhizoplane Roots were washed under running tap water. Tap and lateral roots were cut into lcm long pieces, washed with sterilized water and transferred onto PDA plates containing penicillin 100,000 units/L) and streptomycin (0.2 g/L). Plates were incubated for 5 days at 28°C. Fungi grown on plates were identified as described above. RESULTS AND DISCUSSION The ecological relationships between host, pathogen and population of soil microbes have been represented by a triangle of interactions (Subba Rao 1977). Root infection by a parasite must be affected and often decisively by the micro- bial activity of the root surface and rhizosphere microflora (Abawi @ Widmer 2000; Boland & Kuykendall 1998; Filion et al. 2004; Manka & Kacprzak 1999; Marschner et al. 2002; Subba Rao 1977; Whipp 2001). In the present study, 57 species of fungi belonging to 23 genera were isolated and identified from rhizoplane and rhizosphere of 65 plant species (Table 2). These comprised 58 genera within 19 families collected from from Sindh and Baluchistan (Table 2). Most of the fungi isolated from rhizosphere and rhizoplane belonged to ascomycota and fungi imperfecti. Few genera belonged to zygomycotina and mycelia sterilia. Host plants are arranged under their families and associated fungi. A greater number of fungi was isolated from the rhizosphere than from the rhizoplane. In rhizosphere, Aspergillus flavus and A. niger were predomi- nant along with Fusarium solani (Table 2). Population of aspergilli like A. flavus, A. niger and A. terrus was high as compared to other fungal species (data not presented). It is interesting to note that A. niger was found in the rhizosphere of the all the members of families Cucurbitaceae, Fabaceae, Solanaceae and Umbliferae. Various studies have also reported the occurrence of greater num- QURESHI ET AL., ROOT FUNG! OF WILD AND CULTIVATED PLANTS OF PAKISTAN 1049 ber of fungal species in rhizosphere than in rhizoplane (Abawi @ Widmer 2000; Curl & Truelove 1986; Dix & Webster 1995; Frey et al. at Motean & Huhta 2002)). It has been shown that microbial population ti din rhizosphere (Manka & Kacprzak 1999). Organic andi bst ded from roots and sloughed off root cells enhanced the microbial population in the region (Tate 1995). Microorganisms in agricultural soils are known to exert profound influences on the soil fertility status as well as on the suppression of soil-borne plant diseases (Kennedy & Smith 1995). In fact, the health of soil can be defined in terms of its microbiological capacity to counteract alates the activity of plant pathogenic or plant deleter an 2002; van Bruggen & Semenov 2000). It is well known that some e soils are pees suppressive to some soil-borne plant pathogens such as Fusarium oxysporum, Pythium and Phytophthora species and this suppression relates to both physicochemical and microbiological features of the soil (Whipps 1997; 2001). Importantly, a soil that is suppressive to one pathogen is not necessarily suppressive to another, and so specificity in soil-plant-microbe interactions for disease suppression exists (Filion et al. 2004; Katan 2002; Marschner et al. 2002; Shiomi et al. 1999). Modern methods for analyzing microbial community structures may prove particularly valuable to help define the key organisms or groups of organisms responsible for such natural suppression as well as for monitoring the spread and impact of introduction of specific biocontrol agents or other management practices on natural microbial populations (Gamo & Shoji 1999; Smit et al. 1999; Postma et al. 2000; Whipps 2001). In the rhizoplane, root-infecting fungi like Fusarium solani, Drechslera australiensis, Macrophomina phaseolina and Rhizoctonia solani were found to be predominant. Aspergillus flavus and A. niger, Alternaria alternata, Chaetomium Sropesins, Curvularia clavata and Fusarium oxysporum showed an inter while remaining isolates were found in low frequency (Fig. 1; Table 2). Among the species of Aspergillus encountered frequency of Aspergillus flavus, A. niger was higher than for other fungi except F. solani. This is presumably due to their high sporulating ability and tolerance for different physico-chemical conditions of soil (Domsch et al. 1993). It is interesting to note that Fusarium solani was found predominant in both the rhizosphere and rhizoplane of most of the plant species reflecting their high competence in both rhizosphere and rhizoplane of different plant species in different ecological con- ditions. Characteristics like resistant nature, tolerance to a wide range of mois- ture, pH and temperature, and parasitic as well as saprophytic mode of life make F. solani ubiquitous (Hussain et al. 1966). Fusarium solani produces many anti- biotics and mycotoxins (Whipps 1997, 2001). Fusarium solani is also reported to possess cytotoxic effect on root knot nematode Meloidogyne javanica, besides parasitizing its eggs (Hameed et al. 2001). These characteristics play a signifi- cant role in its establishment in the rhizosphere and rhizoplane. Chaetomium 1050 BRIT.ORG/SIDA 21(2) number of plants be t Fr J Fic. 1. Major f globosum,C. indicum and Macrophomina phaseolina had a moderate frequency while Aspergillus fumigatus, A. nidulans, Fusarium moniliforme, F.oxysporum, F. semitectum, Penicillium spp., showed low frequency (Table 2). Penicillium spp. are generally abundant in soil and are often among the readily isolated species (Qureshi & Khan 1971). There are reports that Penicillium, Trichoderma, and Cladosporium are most common genera associated with rhizosphere of re- sistant varieties of flex to Fusarium wilt, while Alternaria, Cephalosporium, Fusarium, Helminthosporium and Verticillium were relatively common in the rhizosphere of susceptible variety (Subba Rao 1977). Chesters and Parkinson (1959) reported that Penicillium spp. are abundant in the rhizosphere of very young roots while in older roots they are replaced by members of family Hypocreaceae (e.g. Fusarium) and family Dematiaceae (e.g. Alternaria and Drechslera).In this study six sterile mycelium and one ascomycetous fungi were not identified. Sterile mycelium did not produced reproductive structure on agar media. Might be they required special condition for reproduction. Oligo- nucleotide finger printing of rRNA genes for analysis of fungal community in soil (Valensky et al. 2002) would be helpful for their identification. It is inter- esting to note that, in general cultivated plants and common weeds of families Fabaceae and Compositae showed more fungi than wild plants. Presumably roots exudates of cultivated plants and common weeds are more attractive to microbes than exudates of wild plants. In the present study well known biocontrol agents like Trichoderma spp., and Paecilomyces lilacinus were found in very low frequencies. Species of Tri- QURESHI ET AL., ROOT FUNGI OF WILD AND CULTIVATED PLANTS OF PAKISTAN 1051 chodermaare known to produce antibiotics (Tate 1995; Domsch et al. 1993). Their fewer occurrences reflect their weak competence in the rhizosphere, presum- ably their sensitivity to metabolites of competeting microorganisms. There are reports that a bacterium with a high level of resistance toa range of antibiotics is more likely to be successful competitor in the rhizosphere than a bacterium producing large quantities of highly active antibiotics but has a marked sensi- tivity to even a single antibiotic molecules (Bazin et al. 1990). Multiple micro- bial interactions involving bacteria and fungi in the rhizosphere are shown to provide enhanced biocontrol in many cases in comparison with biocontrol agents used singly (Boland & Kuykendall 1998; Whipps 1997, 2000). A wide range of fungi with antagonistic activity against root pathogens are discovered each year. However, ecological success of the antagonist on the plant roots is governed by its ability to colonize and utilize substrates on plant root surface, allowing it to compete effectively with pathogens and other competitive mi- croorganisms. Otherwise the success of a biocontrol agent with poor rhizosphere competence seems very remote. ACKNOWLEDGMENTS This work was carried out under a research grant from the Pakistan Science Foundation, Islamabad, Pakistan. We are thankful to William Campbell, Utah State University, Logan, Utah; M. A. Bari Mallik, Langston University, Langston, Oklahoma and Riaz Ahmad, University of Central Oklahoma, Edmond, Okla- homa for their critical comments and helpful suggestions on the manuscript. REFERENCES Asawi,G.S.and T.L.Wiomer. 2000. |mpact of soil health management practices on soilborne pathogens, nematodes and root diseases of vegetable crops. Appl. Soil Ecol. 15:37-47. ATKINSON, D.and C.A. Watson. 2000. The beneficial rhizosphere: a dynamic study. Appl. Soil. Ecol. 15:99-104. Barnett, H.L.and B.B. Hunter. 1998. Illustrated genera of imperfect fungi. 4" ed. APS Press, St.Paul, MN. Bazin, M.J., P. MarkHAM, E.M. Scott, and J.M. LyncH. 1990. Population dynamics and rhizo- sphere interaction.|n:J.M.Lynch,ed.The rhizosphere. John Wiley & Sons, Inc., New York. Pp. 99-127. Botanp, G.J.and L.D. KuyKenbatt (eds.). 1998. Plant-microbe interactions and biological con- trol. Marcel Dekker, New York, NY. Buraes, H.D. (ed.). 1998. Formulation of microbial pesticides. Kluwer Acad. Publishers, Boston, MA, Cuesters, C.G.C. and D. Parkinson. 1959. On the distribution of fungi in the rhizosphere of oats. Pl. & Soil 11:145-156. Curl, E.A. 1982. The rhizosphere: relation to pathogen behavior and root disease. Plant Dis. 66:624-630. 1052 BRIT.ORG/SIDA 21(2) Curt, E.A.and B. Tructove. 1986. The rhizosphere. Springer-Verlag, Berlin. Dix, NJ.and J. Wesster.1995. Fungal ecology. Chapman and Hall, London. Domscu, K.H.,W.Gams and T-H. ANDERSON. 1993.Compendium of soil fungi. Vol. 1,IHW, Eching. Fition, M., R.C. HAMELIN, L. Bernier, and M. St-Arnaup. 2004. Molecular profiling of rhizosphere microbial communities associated with healthy and diseased black spruce (Picea mariana) seedlings grown in nursery. Appl. Envir. Microbiol. 70:3541—3551. Frey, S.D.,E.T. Euiort, and K. Paustian. 1999. Bacterial and fungal abundance and biomass in conventional and no-tillage agro-ecosystems along two climatic gradients. Soil Biol. Biochem. 31:573-585. Gamo, M.and T. SHou!.1999.A method of profiling microbial communities based on a most- probable-number assay that uses BIOLOG plates and multiple sole carbon sources. Appl. Envir. Microbiol. 65:4419-4424. Hacteen, F., PW. Crous, and O. Perini. 2003. Fungi associated with healthy grapevine cut- tings in nurseries, with special reference to pathogens involved in the decline of young vines. Austral. Pl. Path, 32:47-52. Hameo, S., R. Riaz, V. SULTANA, S. EHTESHAMUL-Hague, and J. Ara. 2001. Variation in nematicidal activity of Fusarium solani (Mart.) Appel and Wollenw emend. Snyd and Hans strains. Pak. J. Biol. Sci. 4:423-425, Hussain, S.S., 5.M. Hassany and S.1. AHmep. 1966. A study of fungus flora of Karachi soils. Pak. J. Sci.Ind. Res. 9:265-268. Katan, J. 2002. Interactions of soil borne pathogens with roots and above ground plant organs. In: Y. Waisel, A. Eshel and U. Kafkafi, eds. Plant roots: the hidden half. Marcel Dekker, New York. Kennepy, A.C. and K.L. Smitx. 1995. Soil microbial diversity and the sustainability of agricul- tural soils. Pl. Soil 170:75-86 Lewis, J.A.,R.P. Larkin, and D.L. Rocers. 1998.A formulation of Trichoderma and Gliocladium to reduce damping-off caused by Rhizoctonia solani and saprophytic growth of the patho- gen in soilless mix. Plant Dis. 85:501—506. Manka, M and M. Kacprzak. 1999. Evaluation of rhizoshere biocontrol agent efficacy and communities of rhizosphere fungi. Bull. Pol. Acad. Sci., Biol. Sci.47:105-112. MarscHner, P., W. Marino, and R. Liesere!. 2002. Seasonal effects on microorganisms in the rhizosphere of two tropical plants in apolyculture agroforestry system in central Amazonia, Brazil. Biol. Fertil. Soils 35:68-71. McLean, M. and V. Huxta 2002. Microfungal community structure in anthropogenic birch stands in central Finland. Biol Fertil. Soils 35:1-12. Netson, PE., T.A. Toussoum, and W.F.O. Marasas. 1983. Fusarium species. An illustrated manual for identification. The Pennsylvania State University Press, University Park. Postama, J.and A.J.G. LurtikHowt. 2000. Effect of the indigenous microflora on the develop- ment of root and crown rot caused by Pythium aphanidermatum in cucumber grown on rookwool. Phytopath. 90:125-133. QuresHi, M.A. and A.S. KHAN. 1971. Fungi from the rhizosphere of Pinus roxburghii Sarg. Pak. J.Forest. 21:75-384. QURESHI ET AL., ROOT FUNG! OF WILD AND CULTIVATED PLANTS OF PAKISTAN 1053 Reyes, A.A.and J.E.MitcHett. 1962.Growth response of several isolates of Fusarium in rhizo- sphere of host and non-host plants. Phytopathology 52:1196-1200 SHiomI, Y., M. NisHivama, T. Onizuka, and T. Margumoto. 1999. Comparison of bacterial commu- nity structures in rhizoplane of tomato plants grown in soils suppressive and condu- cive towards bacterial wilt. Appl. Envir. Biol.65:3996-4001. Smit, E.,P.LeEeFLANG, B.GLANDORF, J.D. VAN ELsas,and K.Wernars 1999. Analysis of fungal diversity in the wheat rhizosphere by sequencing of cloned PCR-amplified genes encoding 18S rRNA and temperature gradient gel electrophoresis. Appl. Environm. Microbiol. 65:2614-2621. SusBa Rao, N.S. 1977. Soil microorganisms and plant growth. Oxford and IBH Publishing Co., New Delhi. Tate, R.L. 1995. Soil microbiology. John Wiley & Sons Inc., New York. VaLensky, L., G.D. Vedova, T. Jianc, and J. Borneman. 2002. Oligonucleotide fingerprinting of rRNA genes for analysis of fungal community composition. Appl.Environm. Microbiol. 68:5999-6004. VAN BruGGeN, A.H.C.and A.M. Semenov. 2000. In search of biological indicators for soil health and disease suppression. Appl. Soil Ecol. 15:13-24. VAN Evsas, J.D., P. Garseva, and J. Sates. 2002. Effect of agronomical measures on the micro- bial diversity of soils as related to the suppression of soil-borne plant pathogens. Bio- degradation 13:29-40. Wuiees, J.M. 1997. Interactions between fungi and plant pathogens in soil and the rhizo- sphere. In: A.C. Gange and V.K. Brown, eds. Multitrophic interactions in terrestrial sys- tems. Blackwell Science, Oxford. Pp. 47-63. Wuiees, J.M. 2001. Microbial interactions and biocontrol in the rhizosphere. J. Expt. Bot. 52:487-511. THE FLORISTIC ECOLOGY OF XERIC LIMESTONE PRAIRIES IN KENTUCKY, AND A COMPARISON TO LIMESTONE CEDAR GLADES AND DEEP-SOIL BARRENS PJ. Lawless J.M. Baskin C.C. Baskin Department of Biology Department of Biology Department of Biology University of Kentucky University of f entucky University of Kentucky Lexington, Kentucky 40506, U.S.A. Lexington, Kentucky 40506, U.S.A. Lexington, Kentucky 40506, U.S.A. an Department of Agronomy University of Kentucky Lexington, Kentucky 40546, U.S.A. ABSTRACT The flora of 18 xeric limestone prairies in the Interior Low Plateaus physiographic eer in Ken- tucky was surveyed and a checklist of vascular plants prepared for each site. Nine of the 18 sites are in the Knobstone Escarpment and Knobs, four in ine ammo one Plateau, two in nth Pn i Plain and three in the Outer Blue Grass. Life form, p geogray servation status and presence were cua for each taxon. Community coefficients were calcu- lated from site floristic lists, and the physiography, geology and soils associated with each site were documented. The flora of xeric limestone prairies in Kentucky was compared to that of limestone cedar glades of ae southeastern United States and of deep-soil barrens of the southwestern Penny- royal Plain in Kentucky and Tennessee. Three hundred and een Te taxa were identified in this plant parton! type, of which 20 are nonnative and 24 state-listed. Families with the highest number of taxa were Asteraceae (70), Poaceae (32) and Fabaceae (29); genera with the highest num- er of taxa were Symphyotrichum (15), Panicums ato (=Panicum + Dichanthelium) (10), Carex - Solidago (7) and Hypericum (6). Intraneous C, hemicryptophytes make up the majority of the flora. Community coe aie indicate ce similarity among all sites except those in the eastern outer Blue Grass, whi type. Endemic species, a higher percentage of ae e.annuals, especially winter agus anda higher number of taxa with western and northwestern geographic affinities distinguish the limestone cedar glade flora from those of xeric limestone prairies and deep-soil barrens. RESUMEN La flora de 18 praderas calcareas xéricas en Ja provincia fisiografica Interior Low Plateaus en ee tucky fue cerca y se preparo un catalogo de plantas vasculares en cada lugar. Nueve de los puntos estan en el Knobstone Escarpment y Knobs, cuatro en el et: Cave Plateau, dos en ‘ oe ies y a en ue Outer Blue Grass. Se — min6 el tipo bioldgico, ruta fotosintética, gr ; n. Se calcularon los coeficientes de comunidad a partir de las listas eee del punto, y se documenté la fisiografia, geologia y suelos asociados con cada punto. La flora de las praderas calcareas xéricas en Kentucky se compara con la de los claros calcareos de cedro del Sureste de los Estados Unidos y con la de los eriales de suelos profundos del Suroeste de la eee Plain en Kentucky y Tennessee. Se identificaron trescientos treinta y cinco taxa en este tl idad vegetal, de los que 20 no son nativos y 24 estan listados en el estado. Las ae con el mayor numero de taxa fueron Asteraceae (70), Poaceae SIDA 21(2): 1055-1079. 2004 1056 BRIT.ORG/SIDA 21(2) ] ] (32) y F abaceae (29): los¢g generos con el mayor numero de taxa fueron S sensu Ce ( Ee + Dichant ial ue wanes (7), Solid ago (7) e ype (6). Los fi la mayor parte de la flora. Los coeficientes ), Panicum de enue indican alta similicud ent re todos los puntos excepto los de Northeastern Blue Grass, que pertenecen a una segunda asociacion o tipo de comunidad. Las especies endémicas, un alto porcentaje de terofitos (ej, anuales, especialmente anuales de invierno) y | ; le taxa con 1 m (Lawless, unpublished). Ten of the 13 soil series upon which xeric lime- stone prairies occur are Alfisols, and the remaining three series are Mollisols (Corydon), Ultisols (Gilpin) and Inceptisols (Garmon). Nine of the 18 sample sites are developed on the Caneyville series, a Typic Hapludalf (Fig. 4). In addi- tion, many of the soil mapping units in xeric limestone prairies are rock out- crop complexes (e.g. Caneyville-Rock Outcrop Complex, Rock Outcrop-Corydon Complex and Rock Outcrop-Fredonia-Colbert Complex). All soil mapping units associated with sample sites in the Knobstone Escarpment and Knobs are mem- bers of the Garmon-Caneyville-Lenberg Soil Association, which in this region also includes the Corydon, Cumberland, Hagerstown and Vertrees series (Arms et al. 1979). Three of the four sites in the Mammoth Cave Plateau (Grayson County Barren, Lapland Road Barrens and Knight's Barrens) occur on the Caneyville series (Arms et al. 1979; Haagen 2001, Whitaker et al. 1972), and the fourth (Lapland Barrens) is the only site on the Rosine-Gilpin-Lenberg com- plex (Haagen 2001). Both sites in the Pennyroyal Plain (Logan County Barrens and Logan County Glade)are restricted to the Rock Outcrop-Fredonia-Colbert complex (Dye et al. 1975). The sample sites in the Outer Blue Grass occur on the Caneyville series (Pine Creek Barrens) (Whitaker & Waters 1986) and Beasley and Shrouts series (Crooked Creek Barrens 1 and 2) (USDA, NRCS, Soil survey of Lewis County, Kentucky. unpublished). METHODS In 2002 and 2003, each of 18 sites was visited a minimum of once per season in spring, summer and autumn. All vascular plant species were recorded and a 1064 BRIT.ORG/SIDA 21(2) TABLE 2. Series, family,and great group of soils in the 18 xeric limestone prairies sampled in Kentucky. For site identification, see Ta ble 1 Soil Series Family Great Group Site(s) Beasley Fine, mixed, active, mesic Typic Hapludalfs 11,12 Caneyville Fine, mixed, active, mesic Typic Hapludalfs 2, 3, 7, 8, 9, 10, 13 14,16 Colbert Fine, smectitic, thermic Vertic Hapludalfs 17,18 Corydon Clayey, mixed, superactive, mesic Lithic Argiudolls 15 Cumberland Fine, mixed, semi-active, thermic Rhodic Paleudalfs 6 Fredonia Fine, mixed, active, mesic Typic Hapludalfs 17,18 Garmon Fine-loamy, mixed, semi-active, Dystric Eutrudepts 8 mesic Gilpin Fine-loamy, mixed, active, mesic Typic Hapludults 15 Hagerstown Fine, mixed, semi-active, mesic Typic Hapludalfs 3 Lenberg Fine, mixed, semi-active, mesic Ultic Hapludalfs 15 Rosine Fine-silty, mixed, semi-active, mesic Ultic Hapludalfs 15 Shrouts Fine, mixed, mesic Typic Hapludalfs 11,12 Vertrees Fine, mixed, semi-active mesic Typic Paleudalfs 24,6 species list prepared for each site. Gleason and Cronquist (1991) was used for field identification, and taxa not identified in the field were collected and de- termined in the University of Kentucky Herbarium (KY) with the aid of a ste- reomicroscope. Each taxon was assigned a presence value based on the percent- age of sites in which it was recorded and placed in one of the following five presence classes (Cain & Castro 1959): 1 1-20%), 2 (21-40%), 3 (41-60%), 4 (61- 80%) and 5 (81-100%) . Sporobolus vaginiflorus was treated as a single taxon, since the two varieties found in this study are primarily distinguished by mi- croscopic characters in both vegetative and reproductive states (FNEC 2003). Community coefficients (CC) were calculated for all possible pair-wise site com- parisons using PC-ORD (McCune and Melford 1999). Community coefficients [CC= 2W/(A+B)] are based on the number of taxa shared between sam ple sites (W) and the total number of taxa in site A and in site B. The life form (sensu Raunkiaer 1934) of each taxon was obtained from Gibson (1961), Hansen (1952), Ennis 1928) and Baskin and Baskin (1978) and the photosynthetic pathway rom Baskin and Baskin (2003) and Waller and Lewis (1978). Life form and pho- tosynthetic pathway also were determined for each of the 342 taxa reported by Chester et al. (1997) in the deep-soil barrens of the southwestern Pennyroyal Plain of Kentucky and Tennessee for comparisons with the floras of xeric lime- stone prairies in Kentucky and limestone cedar glades of the eu eee United States (Baskin & Baskin 1999). We reviewed the list of st threatened and special concern species Cee by the ea State Na- ture Preserves Commission (2002) and noted all state-listed taxa occurring in the xeric limestone prairies of Kentucky. —— ndang os LAWLESS ET AL., XERIC LIMESTONE PRAIRIES IN KENTUCKY 1065 oe ee ms, N G x j . ’ 3 i? 1 ality, Ee Ne fies f+ (hte Be RRE See 2, Als ye hen wi Sys ¥ C a tetite Wyre Rell Dag ir, alluvium Ahlin-L di £el } j | . . | va L Fic g pography i Low Plateaus physiographic province in Hardin and Larue counties Kentucky. Soil series: C(n=Caneyville, Cr=Crider, Gm=Garmon, Hn=Hagerstown, Lf=Lenberg-Frondorf, No=Nolin-N ewark-Melvin, V=Vertrees. The geographic relationships of all taxa were determined by examining distribution maps obtained from Plants (USDA, NRCS 2004). Taxa in which Kentucky is in the central region of their current distribution were defined as intraneous, and those in which Kentucky is at the edge of their current distri- bution were defined as extraneous. When considering the geographic relation- ships of taxa in which Kentucky is near the edge of their range, the conserva- tion status or rank and/or county distribution in the adjacent state was (were) considered, if available. If Kentucky is near the edge of the taxon’s current dis- tribution and the taxon is listed (endangered, threatened or special concern) in the adjacent state or only distributed in counties adjacent to the Kentucky bor- der, the taxon was considered extraneous. The extraneous species were divided further into extraneous northern, southern, eastern and western groups in ac- cordance with the position of Kentucky in relation to the center of distribution of the taxon of interest. 1066 BRIT.ORG/SIDA 21(2) RESULTS A total of 335 vascular plant taxa representing 333 species, 215 genera and 72 families were recorded in the 18 xeric limestone prairies surveyed (Appendix). The genera Crataegus and Rubus were each treated as a single taxon, because no flowering individuals were observed in either genus, both of which are char- acterized by intergradation of vegetative characters between/among species and high taxonomic diversity. Three hundred and fifteen (94.0%) of the 335 taxa are native to Kentucky, and the remaining 20 (6.0%) nonnative (Fig. 5). Asteraceae, Poaceae and Fabaceae had the highest richness values of all fami- lies, containing 70 (20.9%), 32 (9.6%) and 29 (8.7%) taxa, respectively. Genera with the largest number of taxa were Symphyotrichum (15), Panicum sensu lato (=Panicum + Dichanthelium) (10), Carex (7), Solidago (6) and Hypericum (6). The xeric limestone prairies in Kentucky support 24 state-listed taxa (9 en- dangered, 8 threatened and 7 special concern), including three species (Del- phinium carolinianum ssp. calciphilum, Leavenworthia exigua var. laciniata and Talinum calcaricum) listed by Baskin and Baskin (1999) as cedar glade endemics. However, none of these state-listed taxa is federally listed, and none of the taxa recorded in this study is endemic to xeric limestone prairies in Ken- tucky. Furthermore, the three cedar glade endemics occur in areas of xeric lime- stone prairies that fit the description of limestone cedar glades when catego- rized at small spatial scales (Fig. 1). The flora of xeric limestone prairies in Kentucky is composed primarily of native C3 hemicryptophytes with broad geographical distributions encom- passing Kentucky. Ninety-two and eight-tenths percent of the taxa are C3 plants, and with the exception of Manfreda virginica, Opuntia compressa (CAM plants) and Cuscuta cuspidata (nonphotosynthetic), the remaining taxa are C4 plants. The hemicryptophyte life form group contains more taxa (52.9% of native spe- cies) than all other life form groups combined (Fig. 5). Seventy-three and five- tenths percent of the native taxa in the xeric limestone prairie flora in Ken- tucky (230 taxa) is intraneous (Fig. 6), and taxa with southern geographic affinities (44 taxa, 14.1%) make up the largest extraneous component of the flora. Although species richness values varied considerably across the 18 xeric limestone prairie sites surveyed, species composition among the sites was quite similar. Average site richness was 113 taxa (CV=13%) and ranged from 88 (Th- ompson Creek Glade) to 151 (Pine Creek Barrens) (Table 1). Calculation of com- munity coefficients (CC's) for all possible pair-wise site comparisons determined that the majority of xeric limestone prairies surveyed forma single association or community type. Only Il of the 153 CC's calculated were less than 0.50, the generally accepted association criterion (Barbour et al. 1999). Nine of the 11 CC’s less than 0.50 included either site 1 or site 2 in Crooked Creek Barrens, and the CC resulting from comparison of Crooked Creek Barrens site | and site 2 is high LAWLESS ET AL., XERIC LIMESTONE PRAIRIES IN KENTUCKY 1067 Percent of Taxa Ww oO oO 20.0 4 10.0 4 0.0 4 1 Ln Phanerophytes Chamaephytes Hemicryptophytes Cryptophytes Therophytes Stem Succulents Epiphytes Life Form Fic 5 Lig. €£, eh pe eas Sees fin “a. p 4 + +1 fi £ PR 4 I ue 4 ite + ky (XLP), deep-soil bar- Lal L ote dD Lola? incD\ i +4 J 1 J £ eh +h + we y ( g (LCG-SE). Percent of Taxa Intraneous Extraneous N Extraneous S Extraneous E Extraneous W Nonnative Geographic Affinity Fic. 6. Geographic affinities of the fl f xeric li iries in Kentucky (XLP) , deep-soil barrens of the south- t Ly 7 I} Diain (NCR) 1 1 lndanc tn thet IRaci £T, (LCG-C TCTHHE SCE LL 1068 BRIT.ORG/SIDA 21(2 (0.71). These data suggest the two sites in Crooked Creek Barrens may belong to a second association or community type. Particularly noteworthy was the ab- sence of Echinacea simulata and Hypericum dolabriforme in both Crooked Creek Barrens sites, since these taxa had high frequency values in the majority of the remaining sites (Lawless, unpublished). CC’ ranged from 0.39 to 0.80, and the average CC (x=0.58) minus one standard deviation (s = 0.07) exceeds the generally accepted 0.50 association criterion. As expected, the presence diagram for the xeric limestone prairie flora in nas (Fig. 7) shows that the majority of species are in presence classes | and 2. However, the percentage of taxa in pee nce class four (15.3%) is similar to that in presence class two (18.0% 6), and thus class four approaches asecond maximum (Oosting 1956). Only 13 taxa were present in all 18 sites, and all of them are na- tive. Five of these 13 taxa are broadly distributed phanerophytes (Cercis canadensis, Diospyros virginiana, Fraxinus americana, Juniperus virginiana and Rhamnus caroliniana), two are dominant perennial (Schizachyrium scoparium) and annual (Sporobolus vaginiflorus) peciesin this community type (Baskin & Baskin 2000; Lawless, unpublished), and the remaining six species (Euphor- bia corollata, Helianthus hirsutus, Lobelia spicata, Physostegia virginiana, Ruellia humilis and Solidago nemoralis) are relatively abundant forbs in the xeric lime- stone prairies in Kentucky (Lawless, unpublished). DISCUSSION The taxonomic distribution of the flora of xeric limestone prairies in Kentucky is very similar to that in the floras of cedar glades of the southeastern United States (Baskin & Baskin 1999) and of the deep-soil barrens of the Southwestern Pennyroyal Plain in Kentucky and Tennessee (Chester et al. 1999). However, the percentage of nonnative taxa in the xeric limestone prairie flora of Kentucky (6.0%) is considerably less than the percentages of nonnative taxa in the floras of the state of Kentucky (21.8%, Jones in press) of cedar glades of the Central Basin of Tennessee (18.7%, Baskin & Baskin 2003) and of southeastern United States 17.9%, Baskin & Baskin 1999) and of deep-soil barrens of the southwest- ern Pennyroyal Plain in Kentucky and Tennessee (9.1%, Chester et al. 1997) (Fig, 6). Furthermore, the vast majority of nonnative species in the xeric limestone prairie flora in Kentucky have low presence values. Nine of 20 taxa were present in only one site, and 18 occurred in four sites or less. The low percentage of nonnative taxa in the flora of xeric limestone prairies in Kentucky may be due to erosion of the organic horizon in these sites, which presumably has reduced nutrient availability in the surface layer. This reduction in surface layer nutri- entavailability may significantly decrease survivorship of nonnative seedlings, as these taxa typically are adapted to nutrient-rich sites (Grime 1974). Asteraceae and Poaceae have the highest richness values of all families in the floras of xeric limestone prairies in Kentucky, cedar glades of the south- rt ~ LAWLESS ET AL., XERIC LIMESTONE PRAIRIES IN KENTUCKY 1069 Percent of Taxa on oS ° oO 30.00 20.00 51 10.00 0.00 1 3 4 2 (41-20%) (21-40%) (41-60%) (61-80%) (81-100%) Presence Class Fic 7.P + £¢ +4 kK Ve xd | 4 10 4 t bg 3 ae L 1 - I plate hysi in Kentucky. P | follows: 1=1-20%, 2=21-40%, 3=41-60%, 4=61—80% and 5=81-100%. Number of taxa in presence class above each bar. eastern United States and deep-soil barrens of the southwestern Pennyroyal Plain, collectively accounting for 30.4%, 20.9% and 33.3%, respectively, of taxa in these floras. The Fabaceae ranks third in species richness in the floras of xeric limestone prairies in Kentucky and deep-soil barrens, whereas species richness in the Cyperaceae exceeds that in the Fabaceae in the flora of cedar glades of the southeastern United States. The flora of cedar glades also is dis- tinguished by 19 endemic and two near-endemic taxa. The genus Panicum sensu lato contains the largest number of taxa in the floras of cedar glades (16 taxa) and deep-soil barrens (10 taxa). The flora of xeric limestone prairies in Ken- tucky also is rich in Panicum species sensu lato (10 taxa); however, Symphyotrichum had the highest richness of all genera (15 taxa). The geographic affinities, photosynthetic pathway and life form distribu- tions among the floras of xeric limestone prairies in Kentucky, limestone cedar glades and deep-soil barrens are similar. However, the flora of limestone cedar glades differs from that of the other two community types in three important aspects. The floras of xeric limestone prairies and deep-soil barrens contain a strong extraneous southern component (14.1 % and 24.0% of floras, respec- tively). In contrast, taxa with western or northwestern geographical affinities make up the largest percentage of extraneous taxa (12.6% of flora) in the lime- stone cedar glade flora (Baskin & Baskin 1999). The phanerophyte life form group contains a considerably smaller percentage of native taxa in the flora of 1070 BRIT.ORG/SIDA 21(2) limestone cedar glades (11.5%) in comparison with the floras of xeric limestone prairies in Kentucky (19.4%) and deep-soil barrens (18.6%) (Fig. 5). The flora of limestone cedar glades also is distinguished by a relatively high percentage of native therophytes (23.0%), many of which are winter annuals. The cedar glade flora contains eight winter annuals in the genus Leavenworthia alone, which partially accounts for the richness of the Brassicaceae (17 taxa) in this commu- nity type and for the near endemic status of the genus Leavenworthia in the southeastern United States (Baskin & Baskin 2003; Estill &@ Cruzan 1999), Con- versely, the xeric limestone prairie flora in Kentucky contains only two native winter annuals, Leavenworthia unifloraand Leavenworthia exigua vat. laciniata, both of which occur in cedar glade-like areas of the perennial grassland ma- trix (Fig. L), and this life cycle type is absent in the deep-soil barrens flora. The winter annual life cycle is an adaptive phenological strategy in lime- stone cedar glades. It assures completion of the life cycle prior to summer-dry conditions, which winter annuals pass in the drought-tolerant seed stage (Baskin & Baskin 1985). The paucity of winter annuals in the floras of xeric limestone prairies in Kentucky and deep-soil barrens of the southwestern Pennyroyal Plain suggest the edaphic conditions in these habitats are considerably less se- vere, and/or these communities have not existed fora sufficient period to allow for evolution of the annual life cycle in a comparable proportion of resident taxa. Although the xeric limestone prairies surveyed support no endemic or fed- erally listed taxa, this community type is the primary habitat in Kentucky for almost all of the 24 state-listed species recorded in the present study. Therefore, preservation of these species in Kentucky is dependent upon conservation of the xeric limestone prairie community type. Presently, nine of the 18 sites sur- veyed are owned by conservation organizations (Kentucky State Nature Pre- serves Commission and The Nature Conservancy) or by the federal government (Fort Knox Military Reservation). Three of the remaining nine sites (Hardin County Cedar Glade, Knight’s Barrens and Lapland Barrens) are under man- agement agreement with The Nature Conservancy, and the Logan County NRCS office is currently in the process of purchasing Logan County Barrens, one of the two sites surveyed in the Pennyroyal Plain. The most significant threat to xeric limestone prairie conservation is all terrain vehicle (ATV) usage, which damages or destroys vegetation, thus promoting further erosion of denuded soils Baskin et al. (1994) proposed the following sequence of events for the origin of xeric limestone prairies in Kentucky: clearing of marginal agricultural lands by European settlers > cultivation and/or grazing > significant erosion of the topsoil > abandonment > colonization of these degraded areas by the xeric limestone prairie flora > succession to hardwood forest in the absence of dis- turbance or maintenance of xeric limestone prairie with periodic management (ie. disclimax). Forest succession is retarded by the highly eroded soils of these abandoned hillsides and by large heat loads (sensu McCune and Grace 2002) as- LAWLESS ET AL., XERIC LIMESTONE PRAIRIES IN KENTUCKY 1071 sociated with the moderate to steep slopes with south to west aspects. However, succession to hardwood forest eventually occurs in the absence of management, which typically consists of mechanical removal of large trees, particularly Juniperus virginiana, and periodic prescribed fire (2-5 year burning interval, David Skinner, KSNPC Eastern Preserve Manager, personal communication). The species composition of this community type is relatively consistent despite 1) their relatively recent origin, 2) the rarity of xeric limestone prairies at the landscape scale and 3) the small area occupied by these communities in the matrix of deciduous forests and agricultural lands. The richness of Asteraceae and Poaceae in the xeric limestone prairie flora of Kentucky is pri- marily responsible for the relatively high community coefficients (x = 0.58) observed in this study, since both of these families are characterized by large regional species pools and high proportions of wind-dispersed taxa. Further- more, we have observed many taxa in the xeric limestone prairie flora (e.g. Schizachyrium scoparium, Panicum flexile, Liatris squarrulosa, Ratibida pinnata, Croton monanthogynous, Hypericum dolabriforme, etc.) in open for- ests with shallow rocky soils, on rock ledges and on rocky stream banks. These habitats, in addition to deep-soil barrens and limestone cedar glades, are the likely sources of the xeric limestone prairie flora in Kentucky, since they ex- isted prior to European settlement. CONCLUSIONS Xeric limestone prairies are broadly distributed throughout the Interior Low Plateaus physiographic province in Kentucky and are most frequent in the Knobstone Escarpment and Knobs, where they primarily occur on eroded Alfisols derived from the Upper Mississippian Salem Limestone. The flora is rich in species of Asteraceae and Poaceae and contains 24 state-listed species, many of which are restricted to xeric limestone prairies in this part of their geographic range. Community coefficients suggest that all sample sites except those in the eastern Outer Blue Grass (Crooked Creek Barrens l and 2) belong to a single community type. Xeric limestone prairies are relatively well protected in Kentucky; the majority of sites are owned and/or managed by conservation organ ations and the federal government. The majority of taxa in the floras of xeric limestone prairies in Kentucky, limestone cedar glades of the southeastern United States and deep-soil barrens of the southwestern Pennyroyal Plain in Kentucky and Tennessee are intraneous C3 hemicryptophytes. The very low percentage (6.0) of nonnative species in the xeric limestone prairie flora of Kentucky is noteworthy. Among xeric lime- stone prairies, limestone cedar glades and deep-soil barrens, the flora of lime- stone cedar glades is particularly unique due to 1) 21 endemic/near endemic taxa, 2) relatively high percentage of therophytes and 3) an extraneous compo- nent with strong western and northwestern affinities (Baskin & Baskin 1999). 1072 BRIT.ORG/SIDA 21(2) In contrast, the floras of xeric limestone prairies in Kentucky and deep-soil barrens of the southwestern Pennyroyal Plain 1) contain no endemic taxa, 2) have higher percentages of phanerophyte taxa and 3) have an extraneous com- ponent with strong southern geographic affinities. APPENDIX Taxonomy is in accordance with USDA, NRCS (2004), with the exception of Elymus glabriflorus var. australis, which is based on the treatment by Camp- bell (1995). The name and authority for each taxon is followed, in parentheses, by the photosynthetic pathway (C3, C4 or CAM), life form (Ph=phanerophyte, Ch=chamaephyte, H=hemicryptophyte, Cr=cryptophyte and Th=therophyte; HP=holoparasite), geographic affinity (=intraneous, EN=extraneous northern, ES=extraneous southern, EE=extraneous eastern, EW=extraneous western and X=introduced) and conservation status in the state of Kentucky (E=endangered, T=threatened and S=special concern) (KSNPC 2002) in bold-faced type. Num- ber in bold-faced type following the final parenthesis refers to number of sites in which species was recorded. DIVISION PTERIDOPHYTA, FERNS AND FERN ALLIES ae a ne L. ee Ph, Dies ADIANTACEAE Do|] t (L.) Link (C3,H,1) 5 1s (L.) Kuntze (C3, Ph, |) 9 ASPLENIACEAE Asplenium platyneuron (L.) B.S.P. (C3, H,1) 3 sed ticlr iu gl fi Prantl (C3, Cr, ES) 2 DIVISION PINOPHYTA, CONIFERS CUPRESSACEAE Juniperus virginiana It (C3, Ph, 1) 18 PINACEAE Pinus virginiana Mill. (C3, Ph, |) 5 DIVISION MAGNOLIOPHYTA, FLOWERING PLANTS CLASS MAGNOLIOPSIDA (DICOTS) ACANTHACEAE Ruellia humilis Nutt. (C3, H, 1) 18 ACERACEAE Acer rubrum L. (C3, Ph, |) 9 Acer saccharum Marshall (C3, Ph, |) 11 ANACARDIACEAE Rhus aromatica Aiton (C3, Ph, |) 7 Rhus copallinum L.(C3, Ph, |) 12 APIACEAE Daucus carota L. (C3, H, X) 1 Eryngium yuccifolium Michx. (C3, Cr, ES) 1 ree rigidior (L.) Raf.(C3,Cr,!) 1 anicula ee L.(C3,Cr,l) 7 de (Michx.) Nutt. (C3, Cr, 1) 1 Zizi aptera os Fernald (C3,H,1) 10 Zizia aurea (L.) W.DJ.Koch (C3, Cr, |) 6 APOCYNACEAE An \ - } aspium arhin L.(C3,H, |) 14 ARISTOLOCHIACEAE Aristolochia serpentaria L.(C3,H, |) 12 ASCLEPIADACEAE maces tuberosa L.(C3,H,1) 1 Asclepias verticillata L.(C3,H, |) 12 Asclepias viridiflora L.(C3, H,!) 17 Matelea obliqua (Jacq.) Woodson (C3, |, ES) 4 ASTERACEAE Achillea millefolium L.(C3,H,1) 1 Ageratina altissima (L.) King & H.E. Robins. (C3, H, 1) 1 Ambrosia artemisiifolia L.(C3, Th, |) 10 Ambrosia trifida L.(C3, Th, !) 2 LAWLESS ET AL., XERIC LIMESTONE PRAIRIES IN KENTUCKY Arnoglossum atriplicifolium (L.) H.E. Robins. (C3 Richards (C3, Ch, na Antennaria plantaginifolia (L. 1) 2 Brickellia eupatoriodies (L.) Shinners = H,|) 16 Centaurea biebersteinii DC. (C3,H Cirsium discolor (Muh. ea ce (C3,H,1) 6 C. rae elestinum (L.) DC. (C3, H, ES) 3 Col is (L eee (C3, Th, |) 8 Coleen ie L.(C3, H, EN) 1 Echinacea simulata McGregor a H, EW) 14 Egon) suigests Muhl. ex Willd. (C3, H, 1) 6 L.(C3,H, ES) 13 Eupatorum hyssoptum I. Si H,ES) 1 oliatum L. (C3,H, I) 2 (L ) Nutt (C3, H, EN) 4 e L.(C3,H,l) 7 ee di ee ip oe Cr, |) 4 IS 3,Cr,l) 18 Hel lianthus a. Torr. & Gray (C3, H, Euthamia ia rami lene hirsutys ES) 5 Helianthus mollis Lam. (C3, Cr, |) 4 Blanes sects Riddell oe Crl) 3 es (L.) Sweet (C3, Cr, |) 3 Leucanthemum ee Lam. CG H, X) 4 Liatris aspera Michx. (C3, Cr, EW) 10 Liatris cylindracea Michx. (C3, Cr, EW,T) 2 Liatris spicata (L.) Willd. (C3, Cr, |) 8 Liatris squarrosa (L.) Michx. var. squarrosa (C3, Cr, ES) 15 Liatris squarrulosa Michx. (C3, Cr, ES) 3 Lonactis linariifolius (L.) Greene (C3, H, |) 4 Oligoneuron rigidum (L.) Small var. epee (El Braun) Nesom (C3,H, EW) 1 Packera anonyma (W cc aWaunceana Love (C3, H, ES) 5 Parthenium integrifolium L.(C3,H, 1) 1 Prenanthes aspera Michx. (C3, H, EW, =F Ratibida pinnata (Vent.) Barnhart a H, EW) 10 Rudbeckia fulgida Aiton (C3,H,|) 1 Rudbeckia hirta L. (C3, H,!) 1 See io glabellus poh (C3, Th, ES) 1 ides (L.) B.S.P. (C3, H, EE) 1 Silphium Thain L. (C3, H, EW, E) 1 Si Iphi um pinnatifidum Elliott (C3, H, ES, $) 3 Silphium terebinthinaceum Jacq. (C3, H, EW) 5 Silphium trifoliatum var. trifoliatum L. (C3, H, 17 Solidago bicolor L.(C3,H,1) 1 1073 Solidago canadensis L.(C3,H, |) 6 Solidago juncea Aiton (C3, H,1) 1 Solidago nemoralis Aiton (C3,H,I) 1 Solidese speciosa Nutt. var. erecta (Pursh) MacMill. (C3, H, ES) 3 Solidago sphacelata Raf. (C3,H,ES) 1 Solidago ulmifolia Muhl. ex Willd. var. ulmifolia = HI) 11 Symphyotrichum cordifolium (L.) Nesom (C3, H, 2 Symphyotrichum dumosum (L.) Nesom (C3,H,1) 1 Symphyotrichum laeve (L.) A. & D. Love var. concinum (Willd.) Nesom (C3, ,!) 2 ee laeve (L.) A.& D. Love var. laeve (C3,H, 1) 1 Senn laterifolium (L.) A.& D.Love (C3, H,l) 1 ee novae-angliae (L.) Nesom (C3, H, 1) 4 Symohyotd ichum oblongifolium (Nutt.) Nesom H, |) 2 Symphyotrichum oolentangiense (Riddell) Nesom (C3, H, EW) 2 Symphyotrichum patens (Aiton) Nesom var. pat- ens (C3,H, 1) 12 Symphyotrichum pilosum (Willd.) Nesom var. pilosum (C3, H Symphyotrichum pilosum (Willd.) Nesom var. pringlei (Gray) Nesom (C3, H, EN) 12 Symphyotrichum sericeum (Vent.) Nesom (C3,H, 6 aS ymphyotrichum shortii (Lindl.) Nesom (C3,H,1) 1 Symphyotrichum undulatum (L.) Nesom (C3, H, EE) 1 Symphyotrichum urophyllum (Lindl.) Nesom (C3, Taraxacum officinale G.H.Weber ex Wiggers (C3, H, X) 2 Verbesina virgin ica L.(C3,H, 1) 4 Vernonia (Walter) Trel. (C3, H, |) 4 BETULA ee americana Walter (C3, Ph, EN) 2 Ostrya virginiana (Mill.) Koch (C3, Ph, |) 16 BIGNONIACEAE Bignonia capreolata L. (C3, Ph, |) 1 Campsis radicans (L.) Seem. ex Bureau (C3, Ph, ES) 1 BORAGINACEAE Heliotropium tene es lum (Nutt.) Torr. (C3, Th, ES) 9 1074 Lithospermum canescens (Michx.) Lehm. (C3, H, BRASSICACEAE Arabis laevigata (Muhl. ex Willd.) Poir. var. ic H,1) 1 Cardamine hirsuta L. (C3, Th, X) 1 Draba verna L.(C3, Th, X) 1 Leavenworthia exigua Rollins var. laciniata Rollins C3,Th, ES,T) 1 yh tL Leavenworthia uniflora (Michx.) Britton (C3, H, 1) 2 CACTACEAE Opuntia humifusa (Raf.) Raf. (CAM, S, ES) 3 CAMPANULACEAE Lobelia spicata Lam. (C3, H, |) 18 CAPRIFOLIACEAE Lonicera japonica Thunb. (C3, Ph, X Symphor ricarpos orbiculatus ne (C3, Ph, |) 6 Vihurny Im prunif fingy L.(C3,P 1 Viburnum auth dili nnn Raf. (ey Ph, ES) 6 CARYOPHYLLACEAE Dianthus armeria L.(C3, Th, X) 1 Minuartia patula (Michx.) Mattf. (C3, Th, !) 1 CELASTRACEAE Celastrus scandens L. (C3, Ph, |) 7 Naeaiea eh is (C3,H, EE) 2 Hypericum ndolabriforme a (C3, H, ES) 14 hypericoides (| ny ae H, ES) 2 NM percum pri um L. (C3, Ph, Hy Nerictim punctatiim L. (C3, H, . ieene um Sphicroeaipte L.(C3,H, EW) 2 CONVOLVULACEAE Calystegia spithamaea (L.) Pursh (C3,H, 1) 1 pomoea pandurata (L.) G.F. Mey. (C3, Cr, |) 12 CORNACEAE Cornus drummondii C. A. Mey. (C3, Ph, EW) 4 Cornus florida L. (C3, Ph, |) 10 CRASSULACEAE Sedum ternatum Michx. (C3, Th, |) 2 CUSCUTACEAE Cuscuta cuspidata Engelm. (HP Th, EW) 1 EBENACEAE Diospyr OS virginiana L. (C3, Ph, \) 18 BRIT.ORG/SIDA 21(2) ERICACEA ie nium eee Marshall (C3, Ph, ES) 4 allidum Aiton (C3,Ph,1) 1 ICINITEM pa EUPHORBIACEAE Acalypha gracilens Gray ant —a C3,Th, 1) 6 Acalypha aie L.(C3,Th, 1) 1 Chamaesy itans (Lag.) Small = Th.) 8 Croton capitatus Michx (C3, Th, 1) 1 lei enOnED OU NOT Michx. ie Th, ES) 7 rollata L. (C3, Cr, I) 18 Es dentata Michx. (C3, Th, l) 3 FABACEAE Cercis canadensis L.(C3, Ph, l) 18 Chamaecrista fasciculata (Michx.) Greene var. fasciculata (C3, Th, |) 11 Coronilla varia L. (C3, Th, Dalea candida Michx. ex Willd. var. candida (C3 W) 1 Dalea purpurea Vent. var. purpurea (C3, H, EW, S)5 a Desmodium c s (L.) DC. (C3,H,1) 2 Desmodium ciliare (Muhl. ex Willd.) DC. (C3, H, 1) 12 Desmodium glabellum (Michx.) DC. (C3, H, ES) 4 Desmodium paniculatum (L.) DC. var. panicula tum ( Desmodium rotundifolium DC. (C3,H, 1) 1 Desmodium sessilifolium (Torr) Torr. & Gray (C3, H,1) 1 Desmanthus illinoensis (Michx.) MacMill. ex B.L. Robins. & Fern. (C3,Ph,EW) 1 = —_ Galactia volubilis (L.) Britton (C3,H, 1) 15 seiuield triacanthos L.(C3,Ph,1) 3 l Michx. (C3 _H, |, S) 3 hespedeea ue (Dum.Cours.) G. Don (C3, H, X) 2 Lespedeza procumbens L.(C3,H, |) 3 aia repens (L.) W. Bartram (C3,H,|) 1 pees 7a violacea (L.) Pers, (C3, H,1) 2 pede a virginica (L.) Britton (C3, H, | Mel ilotus alba Medikus (C3, H, X) 8 Orbexilum pedunculatum (Mill.) Rydb. var. psoralioides (Walt.) Isley (C3, Cr, ES) 1 Robinia pseudoacacia L.(C3, Ph, |) 2 Senna marilandica (L.) Link (C3, H, |) 4 Strophostyles umbellata (Muhl. ex Willd.) Britton (C3, Cr, ES) 3 Stylosanthes biflora (L.) B.S.P.(C3, H, ES) 6 Ter hrosia virginiana (L.) Pers. (C3, H, |) 8 = —a LAWLESS ET AL., XERIC LIMESTONE PRAIRIES IN KENTUCKY 1075 Trifolium repens L.(C3,H, X) 1 LYTHRACEAE Vicia villosa Roth (C3, H, Xx) 2 Cuphea VISCOSISSIMA Jacq. (C3, Th, ES) 2 FAGACEAE MAGNOLIACEAE Fagus grandifolia Ehrh. (C3, Ph, |) 5 Liriodendron tulipifera L.(C3, Ph, |) 6 pice pe L.(C3, Ph, |) 2 MALVACEAE uer ricaria Michx. (C3, Ph, 1) 3 Malvastrum hispidum (Pursh) Hoch. (C3, Th, EW, ewes landica Muenchh. (C3, Ph, ES) 13 T)1 Quercus muehlenbergii Engelm. (C3, Ph, |) 13 Quercus prinus L.(C3,Ph,1) 1 OLEACEAE Quercus rubra L. (C3, Ph, !) 10 fey tiera li iguana (Michx.) Poir. Ph,ES,T) 1 Quercus stellata Wangenh. (C3, Ph, |) 15 o L.(C3, Ph, |) Quercus velutina Lam. (C3, Ph, |) 6 Li igustrum sinense Lour. (C3, Ph, oe GENTIANACEAE ONAGRACEAE Frasera liniensis Walters (C3,H, |) 2 Gaura biennis L.(C3,H, EE) 4 Gentiana flavida Muhl. ex Nutt. (C3, H, |, E) 1 Gaura filipes Spach (C3,H, ES) 13 Gentianella quinquefolia (L.) Small ssp. Genothera biennis L.(C3,H,1) 1 pails: Say) Gillett (C3, H, 1) 9 OX ALIDACEAE Sabatia angularis (L.) Pursh (C3,H, |) 15 Oxalis stricta L.(C3,H,l) 1 JUGLANDACEAE Oxallis violacea L.(C3,H,1) 1 Carya alba (L.) Nutt. ex Ellis (C3, Ph, |) 3 PAPAVERACEAE Carya tomentosa Nutt. (C3, Ph, |) 3 PM (Gcreala Juglans nigra L.(C3, Ph, |) 13 HOLA an) PASSIFLORACEAE LAMIACEAE Passiflora lutea L.(C3,H, |) 2 Blephilia ciliata (L.) Benth. (C3, H, |) 13 PLANTAGINACEAE lcanthriie brachingiye B.S.P. (C3, Th, I) 13 Pie ; Mi " C3, Th | Lycopus americanus Muhl. ex W. Bartram (C3, H, NGGO Hera el i 1 Plantago patagoni ica Jacq.(C3,Th, X) 1 M narda fi fis tulosa iy (C3, H, \)1 Plantago virginica L. iT 1 l) Biycoseaia virginiana (L.) ae (G3, H, l) 18 PLATANACEAE Prunella vulgaris L. var. lanceolata (Barton) — Platanus occidentalis L. (C3, Ph, |) 1 eee ae POLEMONIACEAE ee pycnanthemoides (Leavenw.) em ald (C3, HES, 2 Jy! TUE PCHEITUtdd tenuifolium Schrad. (C3, H, 1) 11 POLYGALACEA E Phlox pilosa L. ssp. pilosa (C3, H,1) 8 sa wie brat bs (C3, H, 5 Polygala verticillata lis (C3,Th, 1) 13 1 elliptica Muh. (C3, 4,1) 3 BORIUEAer CE. Scutellaria parvula Michx.var.australis Fassett (C3, Ul n Ware (C3,H, ES, E) 1 Cris esi iee laMi var. missouriensis (Torr.) PRIMULACEAE Anagallis a is L. (C3, Th, X) 3 man & Lawson (C3, Cr, |) 2 Trichostema dichotomum L. (C3,Th, |) 1 LAURACEAE Dodecatheon meadia L.(C3,H,1) 1 Lysimachia lanceolata Walter (C3, H, |) 6 safrass albidum (Nutt, Nees (C3, Ph, ) 11 Lilary haan diel 1emone virginiana L. (C3,H, |) 13 LINACEAE Clematis pitcheri Torr. & Gray var. pitcheri (C3, Ph, Linum medium (Planch.) Britton (C3, H, |) 1 EW) 1 Linum sulcatum Riddell (C3, Th, EW) 16 Delphinium carolinianum Walter ssp.calciphilum Linum virginianum L.(C3,H,!) 2 Warnock (C3,H, ES, T) 1 1076 Thalictrum revolutum DC. (C3,H, !) 3 Thalictrum tha all Cr,1) 2 RHAMNACEAE Ceanothus americanus (C3, Ph, |) 3 Rhamnus Nalter (C3,Ph, ES) 1 ROSACEAE Agrimonia rostellata Wallr. (C3, H, l) 6 Amelanchier arborea (Michx. f.) Fernald (C3, Ph, 1) 6 sea L. [sp(p).] (C3,Ph) 5 nd virgit niana Duchesne (C3, Eh )5 Porteant stipulatus (Muhl. ex Willd.) Britton (C3,H,1)1 Potentilla iriples Michx.(C3,H,1) 1 Prunus americana Marshall (C3, Ph, a Prunus serotina Ehrh. (C3, Ph, l) 7 Rosa carolina L.(C3, Ph, |) 16 Rosa multiflora Thunb. (C3, Ph, X) 4 Rubus L. [sp(p).] (C3, Ph) 9 RUBIACEAE peas teres Walter C3,Th,) 4 Michx. (C3,H, 1!) 1 Galium pilosum Aiton (C3, H,l) 7 Galium triflorum Michx. (C3,H, |) 1 Hedyotis nigricans (Lam.) Fosberg (C3,Th,EW) 11 = a = =a Houstonia canadensis Willd. ex Roemer & J.A. Schultes (C3, H, EN) 13 SALICACEAE Salix humilis Marshall (C3, Ph, |) 1 SANTALACEAE Comandra umbellata (L.) Nutt. (C3, Cr, |) 1 SCROPHULARIACEAE Agalinis auriculata (Michx.) Blake (C3, Th, EW, E)1 Agalinis gattingeri (Small) Small (C3, Th, EW) 4 Agalinis tenuifolia (Vahl) Raf. (C3, Th, |) 9 Aureolaria flava (L.) Farw. (C3, Cr, |) 2 R bh ; — Castilleja coccinea (L.) Spreng. (C3, Th, |, E) 3 iol is ea L.(C3,H,1) 3 is Nutt.ex Sims (C3,H, 1) 1 a sehal imi (L.) Willd. (C3, H, |) 2 SIMAROUBACEAE Ailanthus altissima (Mill) Swingle (C3, Ph, X) 1 SOLANACEAE Physalis virgit Viana Mill. (C3, Cr, ES) 12 ictroides (L.) Eames & Boivin (C3, BRIT.ORG/SIDA 21(2) ULMACEAE Celtis occidentalis L. (C3, Ph, 1) 1 Celtis tenuifolia Nutt. (C3, Ph, |) 17 Ulmus alata Michx. (C3, Ph, ES) 6 Ulmus rubra Muh. (C3, Ph, |) 3 VERBENACEAE Phryma leptostachya L. (C3,H, |) 1 Verbena simplex Lehm. (C3, H,l) 6 VIOLACEAE ont concolor (T. Forst.) Spreng. (C3,H, | ii Brainerd (C3,H, ES,S) 9 es pane L.(C3,H,1) 7 Viola pedata L.(C3,H, |) 12 Viola sagittata Aiton (C3,H, |) 2 — 3 VITACEAE Parthenocissus quinquefolia (L.) Planch. (C3, Ph, 1) 9 Vitis aestivalis Michx. (C3, Ph, 11 DIVISION FLOWERING PLANTS CLASS LILIOPSIDA (MONOCOTS) AGAVACEAE Manfreda virginica (L.) Salisb. ex Rose (CAM, H, MAGNOLIOPHYTA, CYPERACEAE Carex complanata Torr. & Hook. (C3, Cr, |) 9 Carex crawei Dewey (C3, Cr, 1,S) 14 Carex glaucodea Tuckerman ex Olney (C3, H,1) 8 Carex juniperorum Catling, Reznicek & Crins (C3, H, EN, E) 2 Carex meadii Dewey (C3, Cr, |) 16 Carex pensylvanica Lam. (C3, Cr, l) 2 Carex umbellata Schkuhr ex Willd. (C3, H, !) 12 Eleocharis compressa Sullivant (C3, Cr, ES) 3 Eleocharis tenuis (Willd.) J.A.Schultes (C3, Cr, 1) 3 Fimbristylis puberula (Michx.) Vahl var. puberula _Cr,|, T) Scirpus pendulus Muhl. (C3, H,1) 8 Scleria olgantha Michx. (C3, Cr,!) 13 Scleria pauciflora Muhl. (C3, Cr, |) 2 DIOSCOREACEAE — = ta J.F.Gmel (C3, Ph, I) 2 IRIDACEAE Hypoxis hirsuta (L.) Coville (C3,Cr, |) 2 LAWLESS ET AL., XERIC LIMESTONE PRAIRIES IN KENTUCKY Iris cristata Aiton (C3, Cr, |) 1 Sisyrinchium albidum Raf.(C3,H, |) 13 LILIACEAE Allium cernuum Roth (C3, Cr, EN) 8 Maianthemum stellatum (L.) Link (C3, Cr, |, E) 1 Nothoscordum bivalve (L.) Britton (C3, Cr, |) 1 Polygonatum biflorum (Walter) Elliott (C3, Cr, 1) 10 Uvularia perfoliata L.(C3,Cr,l) 1 ORCHIDACEAE Cypripedium candidum Muhl.ex Willd. (C3, Cr, EN, E) 2 Cypripedium pubescens Willd. var. pubescens (C3, 2 Spiranthes lacera (Raf.) Raf. var. gracilis (Bigelow) Leur (C3,H,l) 4 Spiranthes magnicamporum Sheviak (C3, H, EW, T)9 Tipularia discolor (Pursh) Nutt. (C3, Cr, 1) 1 POACEAE Andropogon gerardii Vitman (C4, H, |) 17 Aristida longispica Poir. (C4,Th, l) 1 Aristida oligantha Michx. (C4,Th, l) 2 Aristida purpurascens Poir. var. purpurascens (C4, Bouteloua curtipendula (Michx.) Torr. (C4, H, |, S)3 Brachyeletrum erectum (Schreb. ex Spreng.) Beauvis. (C3, H,1) 1 Bromus pubescens Muhl. ex Willd. (C3, H, |) 1 Danthonia spicata (L.) Beauvis.ex Roemer & J.A. Schultes (C3, H, |) 14 Dichanthelium acuminatum (Sw.) Gould & CA. Clark var. fasciculatum (Torr) Freckmann (C3, H,1) 15 Dichanthelium boscii (Poir.) Gould & C.A. Clark (C3,H,1) 12 1077 Dichanthelium commutatum (J.A. Schultes) Gould (C3,H, I) 1 Dichanthelium dichotomum (L.) Gould var. dichotomum (C3,H, |) 6 Dichanthelium sphaerocarpon (Ellis) Gould (C3 H, |) 3 Dichanthelium villosissimum (Nash) Freckman (C3,H, 1) 1 Elymus glabriflorus Scribn.& C.R. Ball var. australis (Scribn. & C.R. Ball) J.J.N.Campb. (C3,H, 1) 5 Lolium arundinaceum (Schreb.) J.J. Darbyshire (C3,H, X)3 Muhlenbergia capillaris (Lam.) Trin. (C4, H, !) 2 Muhlenbergia cuspidata (Torr. ex Hook.) Rydb. iMapicneean sylvatica Torr. ex Gray (C4, H, |) 2 anion angneee Michx. (C4, Cr, ES) 2 illare L.(C4 ih ) : atu flexile (Gattinger) Scribn. (C4, Th, l) 14 Panicum linearifolium (Scribn. ex Nash) Gould (C3,H,l) 13 Paspalum leave Michx. (C4,H, |) 1 Schi elu scoparium (Michx.) Nash (C4, H, I) 1 Setaria ae (L.) P. Beauvis. (C4, Th, X) 2 Sorghastrum nutans (L.) Nash (C4, Cr, 1) 16 Sporobolus compositus (Poir.) Merr. var. com- positus (C4,H, |) 14 Sporobolus heterolepis (Gray) Gray (C4,H, EW) 1 Sporobolus vaginiflorus (Torr. ex Gray) Wood, including varieties ozarkanus (Fernald) Shinners (C4, Th, 1) and vaginiflorus (C4, Th, 1) 18 Tridens flavus (L.) Hitchcock (C4, H, |) 12 SMILACACEAE Smilax ae nox L.(C3,Ph, ES) 1 ACKNOWLEDGMENTS The authors sincerely thank the Kentucky State Nature Preserves Commission, The Nature Conservancy and all private landowners for granting the first au- thor access to the field sites studied in this project. REFERENCES Amos, D.H. 1972. Geologic map of the New Amsterdam Quadrangle, Kentucky-Indiana and part of the Mauckport Quadrangle, Kentucky. USGS Geologic Quadrangle Map GQ-990. 1078 BRIT.ORG/SIDA 21(2) Arms, F.S., MJ. MircHet, F.C. Warts, and B.L. Witson. 1979. Soil survey of Hardin and Larue counties, Kentucky. USDA, Soil Conservation Service, Washington, D.C Barsour, M.G., J.H. Burk, W.D. Pitts, F.S. Gittiam, and M.W. ScHwarrz. 1999. Terrestrial plant ecol- ogy, third ed. Benjamin Cummings, Menlo Park, California. Baskin, JM.and C.C. Baskin. 1985. Life cycle ecology of annual plant species of cedar glades of southeastern United States. In: J. White, ed. The population structure of vegetation. Dr.W. Junk, Dordrecht. Pp. 371-398. Baskin, J.M. and C.C. Baskin. 1999. Cedar glades of the southeastern United States. In: R.C. Anderson, J.S. Fralish, and J.M. Baskin, eds. Savannas, barrens, and rock outcrop plant communities of eastern North America. Cambridge Univ. Press, Cambridge. Pp. 206-219. Baskin, JM. and C.C. Baskin. 2000.Vegetation of limestone and dolomite glades in the Ozarks and Midwest regions of the United States. Ann. Missouri Bot. Gard. 87: 286-294. Baskin, JM. and C.C. Baskin. 2003. The vascular flora of cedar glades of the southeastern United States and its phytogeographical relationships. J. Torrey Bot.Soc. 130: 101-118. Baskin, J.M., C.C. Baskin, and E.W. CHester. 1994. The Big Barrens Region of Kentucky and Tennessee: further observations and considerations. Castanea 59:226-254, Braun, E.L. 1950. Deciduous forests of eastern North America. Blakiston, Philadelphia. Cain, S.A.and G.M de Ouiveira Castro. 1959. Manual of vegetation analysis. Harper & Broth- ers, New York. Campesett, J.N.N. 1995. New combinations in eastern North American Elymus (Poaceae) Novon 5:128. Cuester, E.W., B.E. Worroro, J.M. Baskin, and C.C. Baskin. 1997. A floristic study of barrens on the southwestern Pennyroyal Plain, Kentucky and Tennessee. Castanea 62: 161-172. Dye, J.W., A.J. Barton, and R.D. Froepae. 1975. Soil survey of Logan County, Kentucky. USDA, Soil Conservation Service, Washington, D.C Ennis, B.E. 1928. The life forms of Connecticut plants and their significance in relation to climate. Connecticut State Geol. Nat. Hist. Surv. Bull. 43. Esti, J.C. and M.B. Cruzan. 2001. Phytogeography of rare plant species endemic to the southeastern United States. Castanea 66:3-23. FeNNEMAN, N.M. 1938. Physiography of eastern United States. McGraw-Hill, New York. Fora oF NortH America Epitoria Committee [FNEC]. 2003. Flora of North America North of Mexico, Volume 25, Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press. New York. Gigson, D. 1961. Life-forms of Kentucky flowering plants. Amer. Midl. Naturalist. 66:1—60. Gicversteeve, B. 1978.Geologic map of the Leitchfield Quadrangle, Grayson County, Kentucky. USGS Geologic Quadrangle Map GQ-1316. Gteason, H.A. and A. Cronauist. 1991, Manual of vascular plants of northeastern United States and adjacent Canada. Second edition. New York Botanical Garden, Bronx. Grime, J.P. 1974. Vegetation classification by reference to strategies. Nature 250:26-31. Haaaen, J.E. 2001. Soil Survey of Breckenridge and Meade Counties, Kentucky. USDA, Soil Conservation Service, Washington, D.C LAWLESS ET AL., XERIC LIMESTONE PRAIRIES IN KENTUCKY 1079 Hansen, C.E. 1952. The life-forms of the flowering plants of Illinois. M.S. thesis. Northwest- ern Univ., Evanston, Illinois Jones, R.L.In Press. Plant life of Kentucky, an illustrated guide to the vascular flora. Univer- sity Press of Kentucky, Lexington. [KSNPC] Kentucky State Nature Preserves Commission. 2002. Endangered, threatened, special concern, and historical biota of Kentucky. unpublished. Frankfort. Keperrete, R.C. 1966. Geologic map of the Elizabethtown Quadrangle, Hardin and Larue Counties, Kentucky. USGS Geologic Quadrangle Map GQ-559. Keperrete, R.C.1967.Geologic map of the Colesburg Quadrangle, Hardin and Bullitt Coun- ties, Kentucky. USGS Geologic Quadrangle Map GQ-602. Keperrete, R.C. 1968. Geologic map of the Shepherdsville Quadrangle, Bullitt County, Ken- tucky. USGS Geologic Quadrangle Map GQ-740. McCune, B.and J.B.Grace. 2002. Analysis of ecological communities. MJM Software Design, Gleneden Beach, Oregon. McCune, B.and M.J. Merrorp. 1999.PC-ORD. Multivariate analysis of ecological data, version 4.MJM Software Design, Gleneden Beach, Oregon. McDoweut, R.C. 1981. Correlation chart for units on the geologic map of Kentucky. USGS Miscellaneous Field Studies Map MF-21. Miter, R.C. 1968. Geologic map of the Russellville Quadrangle, Logan County, Kentucky. USGS Geologic Quadrangle Map GO-714. Moore, F.B. 1965.Geology of the Millerstown Quadrangle, Kentucky. USGS Geologic Quad- rangle Map GQ-417. Oostine, H.J. 1956. The study of plant communities. Second edition. Freeman, San Fran- CISCO. Peck, J.H.and K.L. Pierce. 1966.Geologic map of part of the Manchester Islands Quadrangle, Lewis County, Kentucky. USGS Geologic Quadrangle Map GQ-581. Peterson, W.L. 1966. Geologic map of the Nelsonville Quadrangle, central Kentucky. USGS Geologic Quadrangle Map GQ-564. QuartermaN, E.and R.L. Powett. 1978. Potential ecological/geological natural landmarks of the Interior Low Plateaus. United States Department of the Interior, Washington, D.C. Rainey, H.C. 1965. Geologic map of the Dennis Quadrangle, Logan County, Kentucky. USGS Geologic Quadrangle Map GQ-450. RAUNKIAER, C. 1934. The life forms of plants and statistical plant geography.Clarendon Press, Oxford. USDA, NRCS. 2004. The PLANTS Database, Version 3.5 (plants.usda.gov). National Plant Data Center, Baton Rouge. Water, S.S.and J.K.Lewss. 1979.Occurrence of C3 and C4 photosynthetic pathways in North American grasses. J. Range Manage. 32:12-28. Wuitaker, O.J., ER. Cox, H.T. Converse, J.L. TaYLor, J.V.Bentey, and E.H. Jacoss. 1972. Soil survey of Grayson County, Kentucky. USDA, Soil Conservation Service, Washington, D.C WHITAKER, O.J.and B.A. Waters. 1986. Soil survey of Bullitt and Spencer Counties, Kentucky. USDA, Soil Conservation Service, Washington, D.C A FLORISTIC SURVEY OF FORT MATANZAS NATIONAL MONUMENT, ST. JOHNS COUNTY, FLORIDA Wendy B. Zomlefer David E. Giannasi Department of Plant Biology Depar tment of Plant Biol ogy mei se rgia University of Geo orgia 1? Plant Sciences 2502 Plant Sciences Athens, a. coe: 7271,USA. Athens, Georgia 30602-7271, U.S.A. Walter S. Judd Lisa M. Kruse Department of Botany Department of Plant Biology University of Florida University of Georgia PO. Box 118526 2502 Plant Sciences Gainesville Florida 32611-8526, U.S.A. Athens, Georgia 30602-7271, U.S.A. Kelly A. Bettinger Department of Plant Biology University of Georgia 2502 Plant Sciences Athens, Georgia 30602-7271, U.S.A. ABSTRACT Fort Matanzas National Monument, administered by the National Park Service, U.S. Department of the Interior, comprises 127 ha (313 acres) that ae Roan one two Paes is eas) in St. Johns County, Florida: northern Rattlesnake Island, and south- em anaes a8 A floristic survey was conducted to provide Park Service personnel with a list of vascular plant species, supplemented with salient mlormation SUC is as rela- tive abundance, aa data, and SoS Four 2003- 2004 ee 230 ess of vascular ee in 189 baad 2 B families. The fone largest families are oaceae, Asteraceae, Fabac and A map, descriptions, and photo- ae of oe various a ee communities are also provided. RESUMEN El Monumento Nacional Fuerte Matanzas, administrado por el Servicio de Parques Nacionales del Departamento del Interior, tiene una extension de 127 ae e pe) we dos islas ee barrera localizadas en el condado de St. Johns, Florida: al Norte la Isla Ratt] Fuerte Matanzas, y al Sur la Isla Anastasia. Fue realizado un monitoreo floristico por el personal del Servicio de Parques Nacionales, utilizando una lista oe aa pouzO, ue Deen pad aes sobre la descripcién, abundancia relativa, localida ad y d pla antas vasculares de la zona. Se realizaron cuatro ies de colecta intensiva entre 2003 y 2004, de los cuales se obtuvieron un total de 237 especies de plantas vasculares, pertenecientes a 189 géneros y 75 familias. Las cinco familias mas representadas son Poaceae, Asteraceae, Fabaceae, Amaranthaceae y Euphorbiaceae. Se presentan mapas, descripciones y Pugh de varias de las comunidades de plantas muestreadas. SIDA 21(2): 1081-1106. 2004 1082 BRIT.ORG/SIDA 21(2) INTRODUCTION Study Area Fort Matanzas National Monument, administered by the National Park Service (NPS; U.S. Department of the Interior), is located 14 mi (22.5 km) south of St. Augustine along State Road (S. R.) AIA in St. Johns County, Florida (Fig. 1A). The park comprises 313 acres (127 ha; Fig. 1B): the southern tip of Anastasia Island (138 acres, 56 ha) and the northern third of Rattlesnake Island (175 acres, 71 ha; NPS 2004). Both islands are separated from mainland Florida by the Matanzas River and the Intracoastal Waterway. These linear shaped barrier is- lands of quartz sand parallel the gently sloping Atlantic coastline and occur on underlying coquina (the Anastasia formation), a soft whitish limestone formed primarily of coarsely broken shells naturally cemented together (Water- bury 1993). The historic Fort Matanzas (discussed below), built from quarried coquina stone, is situated on northeast Rattlesnake Island overlooking the Matanzas River. A comprehensive and vouchered survey of the vascular flora had not previously been conducted for this NPS controlled land. Anastasia Island.—The Anastasia Island portion of the park [1.0 mi (.6 km) long; minimum width 0.02 mi (0.03 km), maximum width 0.4 mi (0.6 km)] consists of stabilized beach dunes rising as muchas 27 ft (8.2 m) above sea level and is bisected by S.R. A1A that becomes a bridge over the southernmost tip of the island (Fig. 1B). Park property borders along A1A are 50 ft (15.2 m) from the center line of the highway on each side (east and west). The park is hemmed in by dense beachfront housing development to the north, and the Summer Ha- ven community across the S.R. AlA bridge, to the south. Most NPS land here is accessible to the public. Along southwest S. R. ALA are two entrance points to the Matanzas River (main visitor center and a dune boardwalk, numbered | and 4, respectively, in Fig. 1B), and across S. R. A1A to the east, two points of easy access to the Atlantic Ocean [beach ramp (2 in Fig. 1B) and dune boardwalk (5)]. The visitor center encompasses a parking lot/gift shop/picnic area, park headquarter offices/maintenance areas, boat dock, and a nature trail/boardwalk [through dense forest (3)]. A small parking lot for a boardwalk (4) through the dunes out to the Matanzas River is 0.35 mi (0.56 km) further south along west S. R. A1A. Directly across the highway to the east is another parking lot for the third, much longer boardwalk (5) over the fragile dune system, which terminates in an overlook of the Atlantic Ocean. Another parking area opposite the visitor center entrance (2) cuts through the dunes and allows vehicle access to the Atlantic Ocean. This popular beach even has traffic “lanes” along the shore (speed limit 10 mi/hr) policed by NPS personnel; however, only four-wheel drive vehicles are allowed along the southwest “hook” of the island bordering Matanzas Inlet where the sand is less compacted and where Least Terns nest for part of the year. ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT 1083 A @Tallahassee” ® Jacksonville 28° . Augustine Ft. Matan aytona Orlando Beach FLORIDA . i Tampa ~ 0 100 km 100 mi Q Miami 4 - 26° 86° 82° ut 1 J \ B A1A ANASTASIA ATLANTIC ISLAND OCEAN Be, ~~. SE N(3) Visitor Center (5) RATTLESNAKE ISLAND () Park Entrance @)Beach Ramp (car access) @G) Boardwalk (maritime hammock) @) Boardwalk (shrubby backdune) MATANZAS VINLET Summer (5) Boardwalk (dune system) Haven © Parking (access, s. Rattlesnake Island) Fic. 1. Location Ih 4 fc AA Net TMA AG WW +i £ Cart Mat ca 14 mi (22.5 km) south of St Augustine Florida. B. Fort M National M t k ty (shad 1), comprising the 7 4h: 4 £D 1 L | } 4 L f F t a7\ +h e fA WAT EH 1 Madi£fiad f Ipc (/9nNA)\ \ J 1084 BRIT.ORG/SIDA 21(2) Rattlesnake Island.—When Fort Matanzas was built in 1740, the island it oc- cupied totaled less than two acres, and a soldier on the gun deck could look directly out over the ocean. Erosion and shifting tidal deposits have altered the inlet, and the fort is now about 0.5 mi (0.8 km) farther from the Atlantic Ocean (Fig. 2). In the early 1900's the Army Corps of Engineers dug the channel for the Intracoastal Waterway west of the fort (Fig. 1B), and the little island was joined with other islets, creating a much larger isle now called Rattlesnake Island (Chandler 2002). The Rattlesnake Island park area [1.28 mi (2.1 km) long; minimum width 0.03 mi (0.05 km), maximum width 0.31 mi (0.50 km)] is bordered to the south by exclusive waterfront homes. The northern and southern ends of the park have been created mainly by dredged spoil from the Intracoastal Waterway, and tidal creeks anda labyrinth of mosquito control ditches fragment the low- lying central portion (Fig. 3). Most of the island is less than 5 ft (1.5 m) above sea level, although the central fill area at the northern end rises to ca. 17 ft (5.2 m). The majority of NPS land on Rattlesnake Island is closed to the public. Fort Matanzas on the northeast coast is accessible for brief (ca. 45 min.) guided tours via a forty-two passenger boat that crosses the river eight times daily from the visitor center on Anastasia island. Annually, over 50,000 tourists visit the fort (NPS 2004). A parking area at the southeastern park border (6 in Fig. 1B) allows access lor fishing, although we noted very little activity along the jetties, espe- cially in comparison to the popular southwestern beaches of Anastasia island along the river and inlet. oH Brief History of Fort Matanzas National Monument Spanish colonial history in Florida (mid-sixteenth to early nineteenth centuries) involved crucial events around the Matanzas Inlet (Manucy 1943; Schesventer et al. 1980; Chandler 2002). The name Matanzas, the Spanish word for slaugh- ters, reflects the violent history of the area - specifically, the massacre of 245 French soldiers in 1565 under the orders of Pedro Menéndez de Avilés. On two occasions, soldiers trying to reach Fort Caroline (near present-day Jacksonville) had shipwrecked in the inlet and surrendered to the Spanish. As “heretics” (Hu- guenots or Protestants) and a threat to Catholic Spanish colonization in Florida, they were subsequently executed behind the dunes on Anastasia Island. A sentry post at Matanzas with no armament was erected by 1569 as part of aseries of lookouts along the uninhabited Matanzas Inlet. The wooden watch- tower was not optimal for weathering the warm temperatures and high hu- midity. The well preserved stone-masonry fort standing on present-day Rattle- snake Island (see photograph in lower right of Fig. 3) was constructed in 1740-1742. Fort Matanzas, designed by engineer Pedro Ruiz de Olano, isa simple, two-story, square structure: 49.5 ft (15.0 m) on each side with a 12 ft (3.7 m) gundeck with two 4 ft (1.2 m) tall parapets (east and west sides) and 30 ft (9.1 ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT -~ ‘ bos is oN \ / a f ~~ \ { ‘ \ \ \ \ “ aN \ Sas ‘ x Fort ~% \ . Matanzas Ss Visitor \ ‘ \ Center \ av \ \ x : ‘ \ \ \ : ae \ \ \ ‘ \ ‘\ ae N \ RATTLESNAKE \ <4 \ \ ‘N x } \ | . ISLAND, ‘sl . ri Va v \ 9) / Lee r os Ke s ; j \ y J T qT 1 ; \ fe i: a 0.3 mi are i { \ / / 5. 2 XS | \ / 1740's boundaries \. XS \ 7 ~ > ! = / ——~- Current boundaries as \ ae \ ra ~ : ees ees x Fic. 2. Perimeter changes of Rattl I d Anast 250 Ei | (108N)\: nracao etal Mf 740’s (when years. Island boundaries in the 1 SS ee errors Tene ee Tan Pf ohns County m) tower (north side; Schesventer et al. 1980). The foundation originally com- prised a system of closely-set pine pilings, driven deeply into the marshy sub- strate. The walls are coquina blocks comprising native shellstone (mainly Donax variabilis) probably quarried from El Pefion (within present-day Sum- mer Haven) and originally were covered with mortar made of lime from burnt oyster shells, mixed with sand and water. Fort Matanzas was armed with five cast iron cannon and usually manned by seven to 10 soldiers but could accommodate a planned maximum of 50 sol- diers. The men were assigned to the fort for one-month duty tours as part of their regular rotation among the outposts and missions near St. Augustine. They had to bring supplies to last the whole month on small boats from St. August- ine; drinking water was either transported by boat or collected into a cistern as rainwater fell on the observation deck of the fort. The strategic location of Fort Matanzas at the mouth of Matanzas Inlet helped maintain Spanish control of the waterway and served asa sentry to warn garrisons at St. Augustine about potential enemies approaching from the south 1086 BRIT.ORG/SIDA 21(2 _| Open Beach Foredune Herbaceous Backdune Canals and Tidal Creeks Bordering Flats Disturbed Areas ) 1 £ Cart AAat Nati TRA rn J tallita ? t Jo hns County GIS 2002) and field sGsciatian: Lower right: photograph of east face of Fort Matanzas. Photo credit: Wendy B. mlefer. ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT 1087 via the Matanzas River. The fort also functioned as a rest stop, coast guard sta- tion, and a place where Spanish vessels headed to St. Augustine could be ad- vised on navigating the river. The inlet was cl nough to Fort Matanzas that ships trying to enter the inlet came within range of the fort’s guns. However, soldiers fired the fort cannons in battle only once, in 1742, thwarting two Brit- ish vessels attempting to gain the inlet. Over the next hundred years, Florida became a diplomatic pawn (Gold 1969). Fort Matanzas suffered periods of neglect during subsequent British (1763-1784) and second Spanish (1784-1821) occupations. The aging fort posed serious maintenance problems (Krakow 1986). By the late 1700's, the east foun- dation, facing the Matanzas River, was eroding. By the time Spain deeded Florida to the United States in 1819, the fort was so badly deteriorated that soldiers could no longer live inside. The United States formally took possession in 1821 but never occupied the fort. At this time Fort Matanzas became the administrative property of the War Department asa national military park. By the turn of the twentieth century, the fort was overgrown with vegetation, including mature shrubs and trees, such as Juniperus virginiana, Myricacerifera,and Quercus virginiana growing from the gundeck (Bryant 1872; Schesventer et al. 1980). The walls were cracked, and the south side leaned precariously on a weakened foundation Undetmined by the tides. In 1916, funds were granted by Congress for tand of the structure under a contract between the War Department and the St. Augustine Historical Society and Institute of Science (Krakow 1986). In 1924 President Calvin Coolidge proclaimed Fort Matanzas a national monument (Executive Proclama- tion No. 1713), and three years later the land around the fort was assigned to the Department of Agriculture as a bird refuge (Executive Order No. 4704). President Franklin D. Roosevelt issued Executive Order No. 6166 in 1933, transferring Fort Matanzas (and other military parks and cemeteries) to the National Park Service, Department of the Interior. Over the past seventy years, the NPS has overseen extensive restoration, including installation of stainless steel rods that hold the upper portion of the fort together (Schesventer et al. 1980; Chandler 2002). Asa result of these preservation efforts, Fort Matanzas, a unique relic of military uiclalliccioniee: in este United States, retains its basic his- toric structure. Shoreli vigilance. The origi- nal national monument site consisted of only the fort on Rattlesnake Island, but through the years, NPS has acquired additional land on both Rattlesnake and Anastasia Islands (see Krakow 1986), thereby preserving an intact (and imperiled) barrier island ecosystem. MATERIALS AND METHODS The first authors, Pl Giannasi and coPI Zomlefer, lead four intensive field trips in 2003 (28-30 May, 29-31 July, 26-27 September) and 2004 (12-13 April) to 1088 BRIT.ORG/SIDA 21(2) collect vascular plant specimen vouchers in triplicate using standard field and herbarium techniques (under NPS collecting permit #FOMA-2003-SCI-0005) with assistance of the coauthors and other personnel listed in the acknowledg- ments. Plant associations were also assessed. A complete set of vouchers is de- posited at GA, and one duplicate set at FLAS. The second duplicate set has not yet been distributed pending resolution of issues concerning NPS ownership of voucher specimens. The floras of Wunderlin & Hansen (2000, 2003) were pri- mary sources for plant identification, supplemented by Godfrey & Wooten (1979, 1981). The majority of plants were identified by WBZ and DEG; coauthor LMK identified most ferns, grasses, and sedges; and coauthor WS] confirmed other problematic determinations. RESULTS AND DISCUSSION Floristics The 485 numbered collections (194 from Rattlesnake Island, 291 from Anasta- sia Island) comprise 237 species (plus two varieties): 125 species from Rattle- snake Island and 197 from Anastasia Island (see ANNOTATED CHECKLIST OF SPECIES below). Included in the list are 14 species planted around park headquarter buildings on Anastasia Island (indicated as “cutt”); Zamia pumila, a native spe- cies, occurs naturally in the park and is also cultivated there, and Severinia buxifolia is cultivated and has also escaped to the nearby hammock. The larg- est families are Poaceae (32 spp.), Asteraceae (28 spp.), Fabaceae (14 spp.), Amaranthaceae s.|.(9 spp.), Euphorbiaceae (9 spp.),and Rubiaceae (8 spp.). With completion of this survey, we have vouchered 57 new county records (53 spe- cies and four varieties) for St. Johns County, Florida, according to Wunderlin and Hansen (2004). No Florida endemics (Wunderlin & Hansen 2003) nor any state/federally listed endangered /threatened plants (Coile & Garland 2003; US. Fish and Wildlife Service 2004) were found. However, Zamia pumila (Florida arrowroot, coontie), which occurs naturally in the backdune/maritime ham- mock habitats (also planted around park headquarters), is a state listed “com- mercially exploited plant” (Coile & Garland 2003). In addition, harvesting Uniola paniculata (seaoats) is prohibited by Florida Statute 370.041 (State of Florida 2004) because of this species’ crucial role as a dune builder and stabilizer. Excluding the 12 cultivated exotics, the remaining 46 introduced species represent 20.4% of the total and include five listed as invasive exotics (FLEPPC 2004). Four (Asparagus aethiopicus, Cinnamomum camphora, Nephrolepis cordifolia, Lantana camara )are ranked as Category | (invasive exotics altering native plant communities by displacing native species, changing community structures/ecological functions, or hybridizing with natives), and one, Pteris vittata, as Category II (invasive exotics increasing in abundance/frequency but not yet altered Florida plant communities to the extent shown by Category | ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT 1089 plants). In addition, Schinus terebinthifolius (Brazilian pepper), another Cat- egory I plant, had recently been extirpated by NPS personnel from the south- ern portion of Rattlesnake Island; reinvasion is possible, however, from fruiting plants observed by the authors near some homes adjacent to the park bound- ary. The one colony of Nephrolepis cordifolia (tuberous sword fern) growing on the edge of the forest near park headquarters (Anastasia Island) was sprayed by park personnel with herbicide the day after we collected vouchers, aggressive efforts to eradicate Asparagus aethiopicus (Sprenger’s asparagus-fern) from the boat dock area were also underway. The relatively few plants of Lantana camara (lantana) occur sporadically in disturbed areas and occasionally in the backdune community. The one cultivated tree of Cinnamomum camphora (camphortree) is near maintenance buildings adjacent to the hammock. The cliff fern Pteris vittata (Chinese ladder brake), a significant arsenic hyperaccumu- lator (Ma et al. 2001), grows on the outer walls of Fort Matanzas and woul require careful, probably mechanical, removal since the rhizomes deeply pen- etrate the precious coquina. Plant Communities The six major community types of the study area (open beach, foredune, backdune, maritime forest, salt marsh, and disturbed areas), discussed below and depicted in Figs. 3 to 5, are based upon satellite imagery (St. Johns County GIS Division 2002), our field observations, and classifications of similar areas by FNAI(1990), Johnson and Barbour (1990), Montague and Wiegert (1990), and Easley and Judd (1993). Barrier islands are dynamic habitats: zonation of spe- cies from the coast inland is controlled by the tolerance of plant growth habits to salt spray and sand burial, as well as wave-driven erosion and overwash (Leatherman 1988; Johnson & Barbour 1990). Overall species diversity is low, and several species may occur in more than one community. These maritime habitats are fairly uniform but may intergrade in the study area. We, therefore, have also designated several habitat subcategories to accommodate certain eco- tones (Fig. 3): herbaceous backdune, shrubby backdune/maritime forest, and bordering flats (of the salt marsh). Open Beach.—This area, also referred to as upper beach (e.g., Johnson & Bar- bour 1990), comprises exposed sandy beach up to the high tide line. High en- ergy waves pound the shoreline and deposit sand grains, forming the beach. Anastasia Island has a well-developed open beach along the Atlantic Ocean, Matanzas Inlet, and the southwestern tip bordering Matanzas River (Figs. 3, 4A). The open beach along Rattlesnake Island, however, is more limited and is best developed along the southeastern shore along the river. Coastal perimeter and sandbar development varies with the tid dsand deposition, especially along the southern hook of Anastasia Island (Fig. 3). This habitat is generally 1090 =~ BRIT.ORG/SIDA 21( Fic. 4. Pl ee eee A Aa Park. A. Open beach ( hte of Ipomoea pres-caprae and I. imp g perpendicul I hore. B. Foredune (eastern Anastasia Is- 1 A\ If iL . T fat 1 a? + | WH s ai. L TL Ht ,and Uniala I iculata (ak t I A tasia Island) at | iwalk west S.R. ATA: Gaillar- dia pulchella, Helianthus debilis, Muhl Mari 4 Dal Fi I€ ee ee a ee ee oe butt (kh L 4 ih) ch kk J f Dasel L lel Z -\ y a “ ic 7 (£, d); Junig gini 1 // (backg 1). Photo credits: A, David E Giannasi; B-D, Wendy B. Zomlefer. ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT 1091 unvegetated with the exception of ramets of Ipomoea imperatiand L. pes-caprae that may trail over the high tide line on Anastasia Island (Fig. +A) where ve- hicles are allowed on the beachfront but not near inland dune vegetation. Foredune.—This plant community, also called beach dune (FNAI 1990) or ocean beach (Easley & Judd 1993), includes terraces, overwash, and blowout sites ad- jacent to the beach itself Johnson & Barbour 1990) and is a mobile and harsh environment sparsely to densely vegetated with pioneer species, especially seaoats (“seaoats zone,” Uniola paniculata). The foredune forms as sand accu- mulates around plants. These sand particles, compared to those of the adjacent open beach, are smaller due to selective uplifting by wind. The vegetation must, therefore, tolerate burial by sand, as well as exposure to wind, salt spray, intense sunlight, and storms. Dune height is determined by wind strength and growth habits of certain dune-forming plants. Plants continuously recolonize the habi- tat due to periodic disturbance by waves om storms gue high tides (Oertel & Larsen 1976). Thiscommunity is p to human impact: large gaps or blowouts in the foredune from seen or off-road vehicle trails desta- bilize the substrate, thereby providing opportunities for erosion by wind and water. The foredune community is best developed as a distinct border between the open beach and much higher backdune (described below) along the Atlan- tic coast and southern hook of Anastasia Island (Figs. 3, 4B). The fragile system on Anastasia Island is protected from trampling by two boardwalks over the dunes (Fig. 1B); direct public access by foot or vehicle is forbidden. On Rattle- snake Island, this vegetation often intergrades with backdune and occurs mainly along the river beach (Fig. 3). Foredune habitats in the study area have been usually built by Uniola paniculata, as well as two other dune grasses, Panicum amarum and Spartina patens. The growth of these species is stimulated by sand burial, with vertical growth keeping pace with burial, and lateral growth via runners forming a con- tinuous dune ridge (Wagner 1964). The colonial and succulent morning-glo- ries, Ipomoea imperati and I. pes-caprae, are usually the first invaders of foredune and beach after storm erosion. The plants produce long stolons that creep across the barren sands at right angles to the coast (Fig. 4A), ensuring ramets both on the old and newest foredunes Johnson & Barbour 1990). Other pioneer species (often succulent), consistently found seaward of the foredune, include: Atriplex cristata, Cakile edulenta, Cakile lanceolata, Cenchrus tribuloides, Chamaesyce bombensis, Gaillardia pulchella, Helianthus debilis, va imbricata, Salsola kali, Sesuvium portulacastrum, and Sporobolus virginicus. Croton punctatus, Hydrocotyle bonariensis, Oenothera humifusa, Phyllanthus abnormis, and Physalis walteri are examples of wider-ranging species also characteristic of this zone. Several common weedy species, such as Chenopodium ambrosioides, 1092 BRIT.ORG/SIDA 21(2) Chamaesyce maculata, and Heterotheca subaxillaris also thrive on the foredunes; dense patches of Distichlis spicata occur in several wetter areas. Shrubby plants with lower salt tolerance, such as Ilex vomitoria and Serenoa repens, typically grow on the lee side of the foredunes where they are some- what protected from sand burial and salt spray. Backdune.—The backdune, also called the transitional zone johnson & Barbour 1990), coastal strand (FNAI 1990), open shrubby interior, maritime thicket, and coastal scrub (FNAI 1990; Easley & Judd 1993), is an ecotonal com- munity generally occurring between foredune and maritime hammock and shares many species of both. These deep, stabilized, wind-deposited coastal dunes are covered with variable, often patchy, vegetation. Backdune communi- ties are generally stable, and as prime beachfront real estate property, comprise one of the most rapidly disappearing community types in Florida (FNAI 1990). Originally a nearly continuous band along the Atlantic coast, backdune now occurs in isolated short stretches. Along eastern Anastasia and Rattlesnake Islands, backdune comprises herbs to low shrubs (Fig. +C, D) plus adjacent regions covered with low, dense, often impenetrable woody vegetation intergrading with maritime hammock community to the west (Fig. 5A). Therefore, the backdune habitat of the study area is here divided into two, more or less, well-demarcated sul s (discussed below), herbaceous backdune and shrubby backdune. Herbaceous backdune zone.—This backdune zone, immediately bordering the foredune (Fig. +C), is characteristically a broad flat area occupied by a mix- ture of herbs, often low-growing, including Chamaecrista fasciculata, Gaillar- dia pulchella, Helianthus debilis, Heterotheca subaxillaris, Hydrocotyle bonariensis, Ipomopsis rubra, lresine rhizomatosa, Opuntia pusilla, Opuntia stricta, Solanum chenopodioides, and Strophostyles helvola, as well as several predominant grasses: Andropogon glomeratus, Muhlenbergia capillaris, Spartina patens, and Uniola paniculata. Some low shrubby plants, such as Borrichia frutescensand Iva imbricata may also occur. Asin the foredune, many kd t, have thickened cuticles, root readily from frag- ments, produce floating seeds, and/or spread by runners. Shrubby backdune zone/maritime hammock.—Further inland, the backdune community commonly comprises a low, dense, often impenetrable thicket of salt-tolerant shrubs and small trees. The characteristic pruned and dwarfed form of these woody plants (see Juniperus in Fig. 4D) results from salt-spray aden winds that kill terminal buds on twigs facing the sea Johnson & Bar- bour 1990). In the study area, this shrubby backdune zone (Figs. 3, 4D) gradu- ally intergrades with maritime hammock, especially on the northwestern Anas- tasia Island portion of the park. Characteristic shrubby species (also often in forest understory) include: Baccharis halimifolia, Ilex vomitoria, Iva frutescens, Myrica cerifera, and Sideroxylon tenax; Serenod repens commonly favors the — _ bac — 1093 ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT k, continued. A Dp Nn ) Fic.5. Pl (overstory). B.S J (right).D.B { J ard ima J J 1). Photo credits: A Al B. Zomlefer. 1094 BRIT.ORG/SIDA 21(2) protected lee slopes and flats behind steeply eroded dunes. Juniperus virginiana, Persea borbonia, Prunus serotina, Sabal palmetto, and Zanthoxylum clava- herculis are common tree associates. On the northwestern side of the Anastasia Island es area, portions of shorter shrubby backdune are almost completely covered by dense mats of tangled woody vines of Ampelopsis arborea, Cissus trifoliata, Parthenocissus quinquefolia, Smilax auriculata, Vitis aestivalis, and Vitis rotundifolia, as well as the herbaceous vine, Mikania cordifolia. Along the west border of the disturbed area comprising south Rattlesnake Island (see Fig. 3), a narrow strip of shrubby backdune vegetation includes species such as Chiococca alba, Teucrium canadense, Vigna luteola,and Zamia pumila. Maritime Hammock.—This vegetation type, the terminal succession stage in these coastal areas, is defined as the impenetrable band of “hardwood” forest just inland of the dune community (Laessle & Monk 1961; Stalter & Dial 1984; FNAI 1990). The habitat is also referred to as coastal hammock (Easley & Judd 1993), stable dune zone, and maritime forest Johnson & Barbour 1990). The dense wind-pruned canopy over the old, stabilized, white sand-dunes combined with humus buildup contributes to some moisture retention, but soils gener- ally remain well-drained because of underlying deep sand. Many species over- lap with those characteristic of the shrubby backdune. As with backdune, maritime hammock is prime resort and residential property and originally was an almost continuous band (with the dune system) along the coast of Florida but is now fragmented by development into short segments (FNAI 1990). Typical mature maritime forest, best developed along the western coast of Anastasia Island (Figs. 3, 5A), forms a continuum with the shrubby backdune zone to the east (described above). The forest often covers relatively steep ter- rain, and sometimes the tops of large trees are near eye-level as one stands on the peak of an old dune. The dominant species are Quercus virginiana and Q. geminata, whose branches are characteristically covered with epiphytes Pleopeltis polypodioides, Tillandsia recurvata, and Tillandsia usneoides. Persea borbonia is a principal understory tree, along with Asimina parviflora, Juniperus virginiana, Quercus myrtifolia, Prunus serotina, and Sabal palmetto. Callicarpa americana, Ilex vomitoria, Myrica cerifera, Sideroxylon tenax, Ser- enoa repens, Rhus copallinum, and Zamia pumila are common understory shrubs (or shrubby trees). As in the backdune, woody vines are prevalent (ie., Parthenocissus quinquefolia, Smilax auriculata, Smilax bona-nox, Vitis aestivalis, Vitis rotundifolia), herbaceous understory plants include Galium hispidulum, Oplismenus hirtellus, Rivina humilis, Ruellia caroliniensis, and Teucrium canadense. The northern tip and western coastline of Rattlesnake Island, along the Intracoastal Waterway, support a much different maritime forest vegetation lacking oaks and dominated by Celtis laevigata, Juniperus virginiana, Persea ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT 1095 borbonia, Pi lliottii,and Sabal palmetto. Pinus elliottii 10-12 m tall) is more common on the northern portion of the island. Numerous snags of slash pine along the western coast indicate that this once dominant species has been de- clining and has been replaced by the more common juniper, cabbage palm, and large trees of Celtis laevigata that reach heights of over 12 m (0.5-1.0 m dbh). Common understory shrubs (to small trees) include Ilex vomitoria (4-5 m tall), Myrica cerifera, Serenoa repens, Sideroxylon tenax, Zamia pumila, and Zan- thoxylum clava-herculis. Smilax auriculata is also common throughout the hammock. The dark forest floor supports little herbaceous understory except for occasional plants of species such as Bacopa monnieri, Galium hispidulum, and Pilea microphylla in exposed damp areas. Salt Marsh.—The salt marsh system includes tidal marsh (FNAI 1990), tidal creeks, and bordering flats (Easley & Judd 1993) - distinguished as separate habitat subtypes in Fig. 3 for the study area. Salt marshes in Florida, most abun- dant north of the normal freeze line, are coastal communities of nonwoody salt- tolerant plants occupying intertidal zones at least occasionally inundated with salt water (Montague & Wiegert 1990). These plants must tolerate poorly aer- ated saline substrate, frequent submersion, and intense sunlight. Salt marshes develop at the land-marine water interface, especially in regions with low re- lief, high tidal range, and low wave energy. Therefore, the elevation varies from slightly below to slightly above sea level, with vegetation growing in intertidal and supertidal zones. Salt marsh ecotone functions in sediment stabilization and coastline storm protection. Dense stems and roots of colonizing plants trap sediments from upland runoff, and decaying marsh plant detritus accumulates to form anaerobic layered soils. Within the park, the salt marsh system is best developed in the north-cen- tral portion of Rattlesnake Island, as well as some smaller areas along the east- central and south-central coastlines (Figs. 3, 5B). The salt marsh system here also includes a distinct network of drainage gullies, tidal creeks, and pools (Fig. 5C). Smaller salt marshes also occur within the southwestern tip of Anastasia Island (Fig. 3),as well as the northwestern most strip of the park property along the Matanzas River. Salt marsh proper (tidal marsh).—Al|though salt marsh vegetation is often distinctively zoned in other locations (each zone dominated by a different spe- cies), large expanses of dense monotypic stands of Spartina alterniflora char- acterize the study area (Fig. 5B). Juncus roemerianus, another important indi- cator species in salt marshes elsewhere in Florida (Montague & Wiegert 1990), only occurs occasionally in a few small patches. Spartina alterniflora tends to grow along the deepest portions, grading subtly to other salt-tolerant plants, such as Batis maritima (Fig. 5C), Distichlis spicata, Limonium carolinianum, Sarcocornia perennis (Salicornia perennis), Sesuvium portulacastrum, and 1096 BRIT.ORG/SIDA 21(2) Suaeda linearis along the edges. Small shrubby trees of Avicennia germinans (black mangrove) to 3 m tall (see background in Fig. 5D) also are well estab- lished within Spartina stands on both Rattlesnake and Anastasia Island. We observed little flowering and fruit set, however: our collections represent the northernmost limit of this tropical species along the east coast of Florida (Wunderlin & Hansen 2004). Bordering flats.—A distinct flora also characterizes the slightly elevated ridges bordering the salt marsh proper and associated tidal waterways in the study area (Fig. 3; Easley & Judd 1993). These flat sandy meadows (Fig. 5D) are vegetated with scattered herbs (mainly grasses and sedges), including: Chamaecrista fasciculata, Cynanchum angustifolium, Cyperus esculentus, Cyperus polystachyos, Cyperus retrorsus, Distichlis spicata, Eragrostis elliottii, Fimbristylis spadicea, Juncus dichotomus, Limonium carolinianum, Muhlenbergia capillaris, Opuntia pusilla, Paronychia herniarioides, Pluchea odorata, Portulacca pilosa, Scleria triglomerata, Setaria parviflora, Solidago stricta, Sporobolus virginicus, and Triplasis purpurea, as well as occasional woody species, such as Borrichia frutescens, Iva frutescens, Juniperus virginiana, Pinus palustris, Prunus serotina, and Zanthoxylum clava-herculis. Disturbed areas (ruderal community).—On Anastasia Island, disturbed habi- tats have developed around public-access areas, and on Rattlesnake Island, con- sist of large Intracoastal Waterway dredge fill areas, land within the network of mosquito control ditches, and Fort Matanzas itself (see Fig. 3). Disturbed ar- eas associated with construction and heavy human use on Anastasia Island (ie., land bordering S. R. AIA, visitor center/picnic tables, parking lots, park head- quarters/roads) have few species in common with the habitats discussed in the preceding sections. Common weedy plants occurring primarily in these dis- turbed areas include: Acalypha graciliens, Andropogon glomeratus, Cenchrus spinifex, Chamaesyce hirta, Chamaesyce hyssopifolia, Conyza canadensis, Cro- ton glandulosus, Cynodon dactylon, Dactyloctenium aegyptium, Indigofera spicata, Lepidium virginicum, Malvastrum corchorifolium, Paspalum setaceum, Phyla nodiflora, Pteridium aquilinum, Salvia lyrata, Sida rhombifolia, Sonchus asper, Spermacoce assurgens, Triodanis perfoliata, Verbena bonariensis, and Verbena officinalis. The sandy dune-like fill area comprising the northern tip of Rattlesnake island (Fig. 3) hasa distinct ring of shrubby backdune/maritime hammock veg- etation (e.g., Celtis laevigata, Iva frutescens, Juniperus virginiana, Pinus elliottii, Sabal palmetto; discussed above under maritime hammock) enclosing an open center of patchy, herbaceous to shrubby, backdune plants, including Myrica cerifera, Baccharis halimifolia, Opuntia pusilla, Phyllanthus abnormis, Prunus serotina, Sideroxylon tenax, Uniola paniculata,and Zanthoxylum clava-herculis. Small specimens of Oxalis corniculata and Pteris vittata were the only plants ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT 1097 growing on the coquina walls of Fort Matanzas (northeastern coast) - a stark comparison to the varied and lush flora of 56 species we found covering the walls of Castillo de San Marcos in St. Augustine during the same study period (Zomlefer & Giannasi 2005). The small mowed lawn surrounding the fort comprises Cynodon dactylon, Hydrocotyle bonariensis, and Stenotaphrum secundatum. The labyrinth of mosquito control ditches in the central southwest por- tion of the Rattlesnake Island park area (Fig. 3), excavated circa 1950-1960’s (D. Parker, pers. comm.), drained large expanses of salt marsh, allowing growth of shrubby backdune and maritime forest (discussed above). The narrow fill area [ca 0.13 mi (0.21 km) long, 0.06 mi (0.10 km) wide, tapering to 0.03 mi (0.05 km); see Fig. 3] forming the southernmost portion Rattlesnake Island park property isan exposed flat ridge of very compact sand bordered by the Matanzas Inlet to the east and the Intracoastal Waterway to the west. This harsh, severely wind- blown habitat supports an odd flora of stunted plants (e.g., Opuntia pusilla, O. stricta) and compressed forms of normally upright plants (Cnidoscolus stimulosus, Gaillardia pulchella, Oenothera humifusa, Phyllanthus abnormis), as well as sand-hugging rosettes of Chamaesyce bombensis and C. maculata, large cushions of Stenaria nigricans, depauperate strings of Galactia volubilis, and hardy scattered tufts of grasses, such as Cenchrus echinatus and Eragrostis secundifolia. — ANNOTATED CHECKLIST OF VASCULAR PLANT TAXA A list of 237 vascular plant species representing 189 genera in 73 families is here compiled from Giannasi & Zomlefer specimens (collection numbers in italic) in alphabetical order by family within three major groups (ferns, gym- nosperms, and angiosperms). Genera, species, and infraspecific taxa are alpha- betical within each family. Scientific nomenclature and common names follow Wunderlin & Hansen (2003); exceptions are vernacular names of a few horti- cultural plants (not included in their flora) that conform to Huxley (1992). Fam- ily circumscriptions for ferns and gymnosperms follow FNA (1993), and for the angiosperms, APG (2003). Non-boldface collection number = collection from Anastasia Island; bold- face collection number = collection from Rattlesnake Island; underlined taxa = new vouchered St. Johns County records according to on-line species list by Wunderlin & Hansen (2004); * = exotic (Wunderlin & Hansen 2003); invasive exotics (FLEPPC 2004): [car 1]= Category I [cat 1] = Category II; cuit = cultivated, ie., planted on park grounds. Habitat data: pa = disturbed areas; Fp = foredune; HB = herbaceous backdune; MH = maritime hammock; 08 = open beach; s8 = shrubby backdune; sB/mH = shrubby backdune/maritime hammock ecotone; sm = salt marsh; sm/sr =salt marsh/bordering flats. Relative abundance: c = common (gen- erally abundant throughout a particular habitat; species easily found), 0 = oc- casional (locally common and/or several individuals distributed within a habi- 1098 BRIT.ORG/SIDA 21(2) tat; species not too difficult to locate); i = infrequent (sporadic occurrence of a small number of individuals; species relatively scarce and not easily found); r= rare (very few individuals encountered) FERNS DENNSTAEDTIACEAE Pteridiu uilin (L.) Kuhn Se in aes ann Clute ex A, baller Bracken fern, Da; 0; 127 NEPHROLEPIDACEAE *Nephrolepis cordifolia (L.) C. Presl, Tuberous sword fern, [car |], 0A; 0; 92 POLYPODIACEAE Alebodium aureum (L.) J.Sm.,Golden polypody, Fo; 1; 547 Pleopeltis polypodioides (L.) E.G. Andrews & Windham var. michauxiana (Weath.) E. G. Andrews & Windham, Resurrection fern, MH; c; 28] PTERIDACEAE *Pteris vittata L., Chinese ladder c 55 brake, [cat i], DA; GYMNOSPERMS CUPRESSACEAE Juniperus virginiana L ., Red cedar, MH, 58/MH;C: 77, PINACEAE Pinus elliottiiE Im.,Slash pine, MH, s8/MH;c: 427, 544,670 ZAMIACEAE Zamia pumila L., Florida arrowroot, MH, $8/MH; 0; 284 [cuit], 397, 647 ANGIOSPERMS ACANTHACEAE Avicennia germinans (L.) L., Black mangrove, smu; 0;51, 247, 392, 4/7/ “Justicia brandegeana Wassh. & L. B. Sm., rimpplant, cut; 529 Ruellia caroliniensis (J. F.Gmel.) Steud., Carolina wild petunia, Mu; i; 276 aun bucus nigra L. subsp. canadensis (L.) R. Bolli, American elder, ba; 0; 130 “Viburnum odoratissimum Ker Gawl., Sweet viburnum, cult; 593 AGAVACEAE *Yucca pela : Spanish bayonet, He, MH, SM/BF; 1,268, 5 AIZOACEAE Sesuvium portulacastrum (L.) L., Shoreline seapurslane, Fb, sM/BF; 0; 16,409, 439 AMARANTHACEAE riplex cristata Humb.& Bonpl.ex Willd., Crested saltbush, Fb, sM/er;0; 244,402,413, 440 Blutaparon vermiculare (L.) Mears, Samp 6 > Vire, FD: I: 59 “Chenopodium album L.,,Lamb'squarters, 0a; i;602 *Cheno podium ambrosioides L., Mexican tea, DA; 0; 128 “Gomphrena serrata iL, Globe amaranth, pa;1;603 |. Rootstock bloodleaf FD, HB; 0; 437,468 *Salsola kali L. subsp. pontica (Pall.) Mosyakin, rickly Russian thistle, Fp; 0; 64, 72, 115, 452 Sarcocornia perennis (Mill) A. J. Scott, Perennial ort, SM, SM/BF; C; 27, 480, 545 [= Salicornia perennis viletre segregation of the perennial from the annual species of laxy I Salicc PEtrca Hhlay oa der Sarcoc ornia eee. etic, and Salicornia s.\. (ine nial and annual species) is likely monophyl etic on the basis of the truncate perianth apices, pubescent nonperispermous seeds, and membranous testa Judd & Ferguson 1999) Suaeda linearis (Elliott) Moq., Sea blite, sm, sm/BF; 0; 475,476 Aare deanetnaa _, Poisonbulb, curt; 280 ANACARDIACEAE Rhus copallinum L.,Winged sumac, mH;0; 327,519 ANNONACEAE Asimina parviflora (Michx.) Dunal, Smallflower pawpaw, Md; 0; 3,319,518 ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT APIACEAE Ptilimnium capillaceum (Michx.) Raf., Mock bishopsweed, 0a; i; 600 APOCYNACEAE Cynanchum angustifolium Pers., Gulf coast swallowwort, SM, SM/BF; i; 26, 70,404 *Nerium oleander L., Oleander, cuit; 410 AQUIFOLIACEAE lex \ penis 13,59, omitoria Aiton Yaupon MH, SB, SB/MH:C: 110, 323,621,624,648 ARALIACEAE i cite bonariensis Com e Largeleaf ma ee DA, FO; C; 52, 90, 113,122 ARECACEAE Sabal palmetto (Walter) Lodd.ex Schult.& Schult f,, Cabbage palm, MH, 58/MH Serenoa a . a sll on palmetto, 4, SB, SB/M x Lams, ASPARAGA *Aspara ie sL., Sprenger's asparagus- fern, DA; 0; [caT 1]; 7 ASTERACEAE Ageratina jucunda (Greene) Clewell & Wooten, Hammock snakeroot, Mb; i; Ambrosia artemisiifolia L, Common ragweed, ba; score ee Michx., willow, st ee es ia cae tree, DA, MH, $8; 0; 494,682 Saltwater Bidens alba (L.) DC. var. radiata (Sch. Bip.) R. E. Ballard ex Melchert, Begaenien: DA; 0; 133 id 1099 Gaillardia pulchella Foug., Firewheel, Fo, 48; c; 80, 102,454 Gamochaeta antillana (Urban) A. Anderberg, Narrowleaf purpl lasting, DA; 0;608,651 Vv [Gamochaeta falcata (Lam.) Cabrera, misap- plied (R. Wunderlin, pers. comm.), as in Wunderlin & Hansen (2003): The correct name for the species in the southeastern ree States is clarified by Nesom (2004).] debilis Nutt. subsp. debilis, East coast dune sunflower, FO, He; ¢; 79, Heterotheca subaxillaris (Lam.) Bae & Rusby, rweed, DA, FD, HB; C; 389, 497, 503 Iva frutescens L., Bigleaf sumpweed, s8, s8/MH; C; 267 lva imbricata Walter, Seacoast marshelder, FO, HB; 0; 246, 398 Krigia virginica (L.) Willd., Virginia dwarfdandelion, HB; 1; aes Lactuca as nifolia Michx., Grassleaf lettuce, Da; ia cordifolia (L. f.) Willd., Florida Keys hempvine, $8; 0; 470 Pluchea odorata (L.) Cass., Sweetscent, sm/BF; i; 397 Solidago odora Aiton var. chapmanii (A. Gray) Cronquist, Chapman’s goldenrod, pa; 1; 502 Solidago stricta Aiton, Wand goldenrod, ba, sm/ ar; 1; 530, 560 meee | *Sonchus asper (L.) Hill, Spiny sowthistl 607 dA; 1510, *Sonchus oleraceus L., Common sowthistle, Da; i; 635 *Taraxacum officinale ee ex “ H.Wigg.,Com- mon dandelion, 0A; r; 663, *Youngia japonica (L.) ae te false Borrichia frutescens (L.) DC., Bushy seaside FD, SB, SM/BF; 0; 54, 407, 626 *Calyptocarpus vialis Less., Straggler daisy, DA; r; CYC, 509 Cirsium _horridulum Michx., Purple thistle, DA, $8; ; 500, Conyza canadensis (L.) Cronquist var.canadensis adian horseweed, pa, He; 0; 251, 490, 535 Erechtites hieracifolius (L.) Raf. ex DC., American bur aie! quercifolius Poir., Oakleaf fleabane, DA, HB; 611 at orium capillifolium (Lam.) Small ex Porter Britton, Dogfennel, sm/sr; r; 538 hawksbeard, pa; i; 644 BATACEAE Batis maritima L., Saltwort, sm;c; 20,248, 479 BORAGINACEAE FD; 1; 245 BRASSICACEAE Cakile edulenta (Bigelow) Hook. subsp. harperi (Small) Rodman, American searocket, FD, HB, sM/8F; C; 73, 114,618,627, 66 Cakile lanceolata (Willd.) O. E. Schultz, Coastal searocket, F0; i; 98 1100 *C pus didymus (L.) Sm., Lesser swinecress, A; 1; 60 Desai Belsiee ge Britton, Western mustard, b ay / i? LEPICiurit Vil ginicum L. die pepperweed, Da, HB, SM/B8F; 1; 24,35, 548,620 BROMELIACEAE *Neoregelia aaa is (Moore) L.B. Sm., Painted ingernail, CULT Tillandsia recurvata (L.) L., Ballmoss, mH; 0; 522 Tillandsia usneoides (L.) L, Spanish moss, MH; Cc; 309 CACTACEAE Opuntia pusill Haw., tle ait ears DA, FD eaves -¢: 48 266 Opuntia stricta (Haw.) Haw., Erect ee DA, HB, $8, SM/BF; C; 77,75, 116 CAMPANULACEAE Triodanis perfoliata (L.) Nieuwl., Clasping Venus’ lookingglass, Da; 1; 605 NNABACEAE Celtis ines Willd., Sugarberry, mH; c; 660 CARYOPHYLLACEAE ee al (Torr. & A. Gray) Fenzl ex Baldwin's nailwort, sm/sr; 1; 537 = ov Nutt., *Stellaria media (L) Vill, Commer chickweed, Da; 0; 637, 643, 665 COMMELINACEAE f.var.diffusa, Common apilniere Whitemoiith dayfl wer, DA i;61, 96,243 a eae Raf, Bluejacket, Da; 0; 1, 458 CONVOLVULACEAE Dichondra caroliniensis onysfoot, HB; i; 654, 671 nee ena (L) Lam., Sweetpotato, DA; Fr; Michx., Carolina | ear eee hey pea Dennst.,, Fievine, 04; 1;505 Ipomoea imperati (Vahl) Griseb., Beach morning- glory, FD, 08; C; L.) G. Mey., Man-of-the- Ipomoea pes-caprae (L.) R. Br., Railroad vine, Fo, BRIT.ORG/SIDA 21(2) *Merremia dissecta (Jacq.) Hallier f, Noyau vine, oasr; 107,289 CYPERACEAE *Cyperus esculentus L., Yellow nutgrass, sM/8F; 0; 29,69 Cyperus polystachyos Rottb., Manyspike flatsedge, Da, su/BF;0;28, 419,455 Cyperus retrorsus Chapm., ee. flatsedge, HB, SM/BF; O; 36, 37, 38, 260,261,557 Cyperus tetragonus Elliott, a a flatsedge, DA; 310,521 Fimbristylis spadicea (L.) Vahl, Marsh fimbry, 0a, sM/BF; 0; 31, 264, 424,540 Scleria triglomerata Michx., Tall nutgrass, Da, sM/ BF; 1; 263, 312 EBENACEAE Diospyros virginiana L.,cCommon persimmon, MH; 01 mS ERICACEAE “Rhododendron simsii Planch., Indian azalea, CULT EUPHORBIACEAE Acalypha gracilens A. Gray, Slender threeseed mercury, DA; 1; 526 Chamaesyce bombensis Jacq.) Dugand, Dixie sandmiat, DA, 08, FD; C; 62, 4 Chamaesyce hirta (L.) Millsp., Pillpod andmat, Da: 1,301 a ae eer (L.) Small, Hyssopleaf at 0;491,5 chomoesyce fie He (L.) Small, Spotted sandm cox e fae ee Engelm.& A. Gray, softly, DA, FD 106 oe eee * var. glandulosus, Vente migo, DA, HB; 0; 3 428, 482,527 Croton punctatus Jacq., Gulf croton, Fo; ¢; 45, 112, 44] al cyathophora (Murray) Bartl., Paintedleaf, da, He; i; 17, 94, 135,406, 447, 493 FABACEAE Centrosema virginianum (L.) Benth. Spurred but- terfly pea, DA, HB; 0; 291, 4 Chamaecrista fasciculata et Greene, Par- tridge pea, FD, HB; C; 65, *Desmodium tortuosum ei ticktrefoil, ba; 1; 478 DC., Dixie ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT 1101 Erythrina herbacea L., Coralbean, DA, HB; 1;430, 616, lactia volubilis (L.) Britton, Downy milkpea, Da; 0; 81, 288, 396 * Indigofera hirsuta L., Hairy indigo, pa; r; 487 *Indigofera spicata Forssk., Trailing indigo, pa; i; 303,425 *Medicago lupulina L., Black medick, ba; 0; 607, 631 * ; . . Medicaan polymorpha L., Burrclover, DA; O; 630, MEA Aiea Ih KA Pm id | VA Loe Faas 82,617 *Meliotus bs icus (L.) All, Indian sweetclover, pa; 1,614,633 *Senna obtusifolia (L.) H.S. Irwin & Barneby, ffeeweed, Hp, 1: 59 Strophostyles helvola (L.) Elliott, Trailing fuzzybean, DA, FD, HB, SM/BF;C; 32, 86, 119,390, 437,471,487 Vigna luteola Jacq.) Benth., Hairypod cowpea, DA; O; ; FAGACEAE Quercus chapmanii Sarg., Chapman’s oak; mH; i; Quercus geminata Small, Sand live oak, MH, SB/MH; 318,644 Quercus myrtifolia Willd., Myrtle oak, MH, 5B/MH; O; Quercus virginiana Mill, Live oak, MH SB/MH; C; 317 GERANIACEAE Geranium carolinianum L., Carolina cranesbill, pa; i619 HYPERICACEAE Hypericum gentianoides (L.) Britton et al., Pineweed, sw/se; 1; 534 Hypericum hypericoides (L.) Crantz, St. Andrew’s- CrOSs, SB, MH, SM/BF; i; 277,436, 536 JUGLANDACEAE *Carya illinoinensis (Wangenh.) K. Koch, Pecan, CULT; 2 JUNCACEAE Juncus dichotomus Elliott, Forked rush, sm/Br; 0; Scheele, Black rush, sm, SM/BF; 0; 27,394 LAMIACEAE eae americana L., American beautyberry, —= re cue Rich.) Briq., Tropical bushmint, DAL 29 Monarda fetes L., Spotted beebalm, Da, HB; 0; 283,297, Salvia lyrata 7 yi pa; 0; 5, 495, 5 Stachys floridana Shuttlew. ex Benth., Ges edgenettle, DA; i; 598 m canadense L., Wood sage, $8, MH; 0; 371 Trichostema dichotomum L.,Forked bluecurls, He, sM/BF; 1; 249,427,485 LAURACEAE *Cinnamomum camphora (L.) J. Camphortree, [car i], cuLT; 638 Persea borbonia (L.) Spreng., Red bay, MH, $8/MH; C; 7,89, 107,255, 322,467, 486 MAGNOLIACEAE Magnolia grandiflora L., Southern magnolia, mH; 1,677 MALVACEAE Malvastrum corchorifolium (Desr.) Britton ex Small, False mallow, ba; i; 737 *Malvastrum coromandelianum (L.) Garcke, Threelobe false mallow, pa; i; 294, *Malvaviscus penduliflorus DC.,Turkscap mallow, ULT; (Persisting ae cultivation near ence road en ce.) Sider a ifolia L. crane 0a; 0; 295,571 Pres|, ORACEAE Morus rubra L.,Red mulberry, Mu; 1; 552 MYRICACEAE Myrica cerifera L., Southern bayberry, MH, $B, s6/ MH; C; 279, 448, 514,656 NYCTAGINACAE Gg, DA; 0; 304 OLEACEAE Forestiera segregata (Jacq.) Krug & Urb., Florida amppris vet, 5B; 1; 476 ee mesnyi Hance, Japanese jasmine, Cutt; ONAGRACEAE Gaura angustifolia Michx., Southern beeblossom, DA, FD; C; 37 1102 Oenothera humifusa Nutt., Seabeach evening- primrose, DA, Fb; C; 18, 87, 124 Oenothera laciniata Hill, Cutleaf eveningprim- rose, DAT; 6 *Oenothera speciosa Nutt., Pinkladies, ba; i; 595 ppisaeie ate alis corniculata L. (incl. O. stricta 2 ee ellow woodsorrel, DA, H8; C; 56, 4 *Oxalis rubra A. St.-Hil., Windowbox woodsorrel, ba; 1; 596, 674 PHYLLANTHACEAE Phyllanthus abnormis Baill, Drummond's leafflower, Da, FD, H8; C; 39b, 88, 104,253, 293, 434 a es tenellus Roxb., Mascarene Island eafflower, DA, HB; 1; 97, 462 PHYTOLACCACEAE ne americana L. var. rigida (Small) Cal ins& tt, American pokeweed, Da, HE; 15,429 [This distinct geographical race, previously included in the flora by adeeb ee but not in the recent edi- & Hansen 2003), merits rec- scan pa Caulkins & Wyatt 1990).] Rivina humilis L., Rougeplant, mx; i; 282 PLANTAGINACEAE Bacopa monnieri (L.) Pennell, Herb-of-grace, sm/ BF; 0; 400, 546 Linaria canadensis (L.) Chaz., Canada toadflax, Hp; V; Plantago virginica L., Virginia plantain, ba; 1; 675 649 Sweetbroom, HB Scoparia dulcis | sM/eF; 1; 408, 498 PLUMBAGINACEAE Carolina sealavender, sm, sM/8F; 0; 252, 539 POACEAE Andropogon glomeratus (Walter) Britton et al.var. hirsutior (Hack.) C. Mohr, Bushy bluestem, pa; C457 sauce glomeratus var. pumilus (Vasey) .Dewey, Bushy bluestem, DA, #8; ep 8 Andropogon virginicus L. vat. Broomsedge bluestem, a; i; 507 Cenchrus echinatus L., Southern sandbur, DA, FO; 1; 411 vIrginicus, ' BRIT.ORG/SIDA 21(2) Cenchrus gracillimus Nash, Slender sandbur, DA; 0; 53,84, 117 Cenchrus spinifex Cav., Coastal sandbur, ba; 0; 305 Cenchrus oo ides L., Sanddune sandbur, 123, *Cynodon en on (L.) Pers., Bermudagrass, DA; c; 307 *“Dactyl octenium aegyptium (L.) Willd.ex Asch. & Schweinf., Durban crowfootarass, DA; 0; 420, 466 Dicanthelium scabriusculum (Elliott) Gould & C. Clark, Woolly witchgrass, DA; 0; 520, 646 *Digitaria bicornis (Lam.) Roem. & Schult., Asia $5, DA; 1; 126 Digitaria filiformis (L.) Koeler var. filiformis, Slen- der crabgrass, 0a; 0; 417 Distichlis spicata (L.) Greene, Saltgrass, H8; 0; 556 *Fleusine indica (L.) Gaertn., Indian gooseqrass, dA; O; 286 Eragrostis elliottii S.Watson, Elliott's lovegrass, sm/ BF; O; 562 Eragrostis secundiflora J. Presl subsp. oxylepis (Torr) S. DB. Koch, Red lovegrass, DA, H8; 0; 77, 12] Eustachys petraea (Sw.) Desv., Pinewoods fingergrass, DA, H8; 0; 41, 83, 499 ‘Lolium perenne L.,, Italian ryegrass, ba; i; 613 Muhlenbergia capillaris (Lam.) Trin. var. filipes (M. A.Curtis) Chapm.ex Beal, Gulf hairawn mubhly, HB; C; 485, 489, 549 Muhlenbergia capillaris var. trichopodes (Elliott) Vasey, Cutover muhly, sm/rF; 0; Oplismenus hirtellus (L.) P. Beauv., Woodsgrass, MH; 0; 525 Panicum amarum Elliott, Bitter panicgrass, Fb, HB; 5464,515,551 *Paspalum notatum Fliiggé var. saurae Parodi, ahiagrass, DA; 0; 765 Paspalum setaceum Michx., Thin paspalum, pa; 0; 287, 308, 528 “Poa annua L., Annual bluegrass, 0a; 0; 650, 668 Setaria parviflora (Poir.) De Yellow bristlegrass, sm/BF; 0; 22 Spartina alterniflora Loisel., Sie cordgrass, ieisy Spartina nahn (Aiton) ao cordgrass, fF; 0;23, 393, Sphenopholis obtusata ih sea Prairie wedgq escale, DA;0; 622 omy ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT 1103 * Sporobolus indicus (L.) R. Br. var. pyramidalis (P. . eauv.) Veldkamp, West Indian dropseed, 0a; obo virginicus (L.) Kunth, Seashore dropseed, Fb, SM/8F; C; 256,271,412, 443,558 Prenat secundatum (Walter) Kuntze, St. Augustinegrass, s8;1; 30 [This common law grass is listed in Wunderlin & Hansen Hes as native, and our collections in remote ar- eas are likely not escapes from cultivation. Triplasis purpurea (Walter) Chapm., Purple sandgrass, DA, SM/BF; 1; 532, 559 Uniola paniculata L., Seacats, Fb, H8;C; 67,68, 125 262 —— POLEMONIACEAE [pomopsis rubra (L.) Wherry, Standingcypress, H8; 1,432 POLYGALACEAE Polygala incarnata L., Procession flower, sm/BF; i; 533 POLYGONACEAE Rumex hastatulus Baldwin, Heartwing dock, ba, 1,610 PORTULACACEAE Portulaca oleracea L., Little hogweed, Da; 1, 298 Portulaca pilosa L., Pink purslane, DA, sM/BF; 1; 273, 299, 415, 444 ROSACEAE *Eriobotrya japonica (Thunb.) Lindl., Loquat, mx; 1; 274 (A colony of several saplings natural- ized in the hammock.) Prunus caroliniana (Mill.) Aiton, Carolina laurelcherry, DA; 1; 640 Prunus serotina Ehrh. var. serotina, Black cherry, MH; 0; 44, 5417, 641,655 *Rhaphiolepis indica (L.) Lindl. Indian hawthorn, CULT Rubus trivialis eae Southern dewberry, DA, HB, SB/MH; 1; 612, 6 RUBIACEAE Chiococca alba (L.) Hitche., Snowberry, Hs; 1; 407, 531 Diodia teres Walter, Poor joe, 0a; 1; 666 Galium hispidulum Michx., Coastal bedstraw, DA, HB, MH; 0; 58,414, 433,524 Houstonia procumbens (J.F.Gmel.) Standl., Inno- cence, H8;i;442,473,653,679 ill ne Flattop mille grain A; 1; 300 sae ak brasiliensis Gomes, Tropical Mexican clover, Da; 1; 306 Spermacoce assurgens Ruiz & Pav., Woodland false buttonweed, parr; 292, 513 Stenaria nigricans (Lam.) Terell var. nigricans, Diamondflowers, DA; 0; 57 TACEAE *Severinia buxifolia (Poir.) Ten., Chinese boxorange, mH; 278,678 [cut] (Cultivated near park maintenance buildings and also es- caped and apparently established in the ne ck.) Zanthoxylum clava-herculis L., Hercules-club, mx, sp/MH; C; 47,257, 403, 453,632 SANTALACEAE Phoradendron leucarpum (Raf.) Reveal & M.C. Johnst., Oak mistletoe, mx; 1; 478 SAPOTACEAE Sideroxylon tenax L., Tough bully, Mu, s8/MH; C; 9, 42,108,270, 435,469 SMILACACEAE Smilax auriculata Walter, Earleaf greenbrier, MH, sp, SB/MH; C; 4,43, 109, 426, 4 Smilax bona-nox L., Saw greenbrier, 0; MH, SB/MH; 8 ViAe 14 Walter, Christmasberry, sm SOLANACEAE F317; 553,625 bye 5 walteri Nutt., Walter's groundcherry, Fo, HB 111 HB; 1, 445 TETRACHONDRACEAE Polypremum procumbens L. Rustweed, DA, HB; O; 118, 134, 254 URTICACEAE Parietaria praetermissa Hinton, Clustered pelli- tory, DA; 1; 93 Pilea mi ‘cophulla (L.) Liebm., Artillery plant, MH; 1; Lam., Black nightshade VERBENACEAE *Lantana camara L., Lantana, [CAT 1], DA, HB; 0; 85, 100, 599 Phyla nodiflora (L.) Greene, Turkey tangle fogfruit, DA, HB; 0; 120, 136, 285 1104 *Verbena bonariensis L., Purpletop vervain, da; i; 132 Verbena officinalis L.subsp. hale/ oo S.C.Bar- ber, Texas vervain, Da; 0; 99, | BRIT.ORG/SIDA 21(2) Parthenocissus quinquefolia (L.) Planch., Virginia creeper, DA, SB, MH; 0; 12, 78, 449 Vitis aestivalis Michx., Summer grape, MH, $8; Cc; 40, 315 Vitis rotundifolia Michx., Muscadine, mH, $8; 0; 70, alinente pelopsis orh v rea (L.) Koehne, Peppervine, ba, H, 58; 0; 14,34, 250 ZINGIBERACEAE Cissus trifoliata (L.) | “ zerumbet (Pers.) B.L. Burtt & RM. Sm., ellflower, cutT; 639 Sorrelvine, sb;i: 374 ACKNOWLEDGMENTS We thank William Carromero and Alexander Reynolds for enthusiastic assis- tance in the field. We greatly appreciate the invaluable support and coopera- tion of National Park Service personnel: Joe De Vivo, Network Coordinator; Gor- don (Gordie) Wilson, Superintendent; David Parker, Site Supervisor and Chief of Resource Management; Andrew Rich, Park Ranger; and Ronnie Dean, Bio- logical Science Technician. Park Guides Ray Hamel and Thomas Birmingham expertly commandeered the Matanzas Queen II around Rattlesnake Island, al- lowing our amphibious landings to otherwise inaccessible shores. Kent D. Per- kins and Richard P Wunderlin provided essential constructive criticisms of the manuscript, Kerik D. Cox created the electronic version of our vegetation map and assisted with the other figures; J. Steve Davis at the University of Florida generously lent us NPS structure study documents on Fort Matanzas; and Fric Fuchs translated the abstract into Spanish. We are also grateful to the Depart- ment of Plant Biology, University of Georgia, for the use of a a: armen ve- hicle for three trips to transport | | field suppli | personnel. This survey was funded by National Park Service contract agreement J2114-03-0006 (PID. E. Giannasi & coPI WB. Zomlefer). REFERENCES Anciosperm PHyLoceny Group [APG]. 2003. An update of the Angiosperm Phylogeny Group Classification for the orders and families of flowering plants: APG Il. Bot. J. Linn. Soc. 141:399-436, Bryant, W.C. (ed). 1872. Picturesque America, Vol. |.D. Appleton and Company, New York, New York. Cau kins, D.B. and R. Wyatt. 1990. Variation and taxonomy of Phytolacca americana and P. rigida in the southeastern United States. Bull. Torrey Bot. Club 17:357-367. CHaNoLER, L. (ed.). 2002. Fort Matanzas National Monument park handbook. Historic Map and Print Co., St. Augustine, Florida. Corte, N.C.and M.A. Gar.anb. 2003. Notes on Florida's endangered and threatened plants. Botany Contribution 38, ed. 4. Florida Department of Agriculture and Consumer Ser- vices, Division of Plant Industry, Gainesville, Florida. ZOMLEFER ET AL., FLORISTICS OF FORT MATANZAS NATIONAL MONUMENT 1105 Eastey, M.C.and WS. Juop. 1993. Vascular flora of Little Talbot Island, Duval County, Florida. Castanea 58:162-177. Flora or NortH America Eviroriat Committee [FNA]. 1993. Flora of North America, vol. 2, Pteri- dophytes and gymnosperms. Oxford University Press, New York, New York. Froripa Exoric Pest PLant Councit [FLEPPC]. 2004. Florida Exotic Pest Plant Council's 2003 list of invasive species. URL: www.fleppc.org/Plantlist/O3list.htm/. FLorIDA Naturat Areas Inventory [FNAI]. 1990. Guide to the natural communities of Florida. Tallahassee, Florida. Goorrey, R.K.and J.W.Wooten. 1979. Aquatic and wetland plants of the southeastern United States. Monocotyledons. University of Georgia Press, Athens, Georgia. Goorrey, R.K.and J.W.Wooten. 1981.Aquatic and wetland plants of the southeastern United States. Dicotyledons. University of Georgia Press, Athens, Georgia. Goto, R.L.1969. Borderline empires in transition:the triple-nation transfer of Florida. South- ern Illinois University Press, Carbondale, Illinois Huxtey, A. (ed). 1992. The new Royal Horticultural Society dictionary of Gardening. Stock- ton Press, New York, New York. JOHNSON, A.F.and M.G. Barsour. 1990. Dunes and maritime forests.In: Ecosystems of Florida, R.L. Myers and John J.Ewel, eds. University of Central Florida Press, Orlando, Florida. Pp. 429-480. Juop, WS.and |.K. Fercuson. 1999. The genera of Chenopodiaceae in the southeastern United States. Harvard Pap. Bot. 4:365-416. Krakow, J.L. 1986. Administrative history of Castillo de San Marcos National Monument and Fort Matanzas National Monument. United States Department of the Interior, National Park Service, Washington, D.C. Laesste, A.M. and C.D. Monk. 1961. Some live oak forests of northeastern Florida. Quart. J. Florida Acad. Sci. 24:39-55. LeaTHERMAN, S.P. 1988. Barrier island handbook, 34 ed. Coastal Publication Series, Labora- tory for Coastal Research, University of Maryland, College Park, Maryland. Ma, L.Q., KM. Komar, C. Tu, W. ZHANG, Y. Cal, and E.D. Kennettey. 2001. A fern that hyperaccu- mulates arsenic. Nature 409:579. Manucy, A.C. (ed). 1943. The history of Castillo de San Marcos & Fort Matanzas, from con- temporary narratives and letters. Source Book Series, No. 3. United States Department of the Interior, National Park Service, Washington, D.C. Montacue, C.L.and R.G. WiecerT. 1990. Salt marshes. In:Ecosystems of Florida, R.L. Myers and John J. Ewel, eds. University of Central Florida Press, Orlando, Florida. Pp. 481-516. National Park Service [NPS]. 2004. Fort Matanzas online. URL: www.nps.gov/foma/home/ home.htm/. Nesom, G.L. 2004. New distribution records for Gamochaeta (Asteraceae: Gnaphalieae) in the United States. Sida 21:1175-1185. Otrret, G.F.and M. Larsen. 1976. Developmental sequences in Georgia coastal dunes and distribution of dune plants. Bull. Georgia Acad. Sci. 34:35-48. 1106 BRIT.ORG/SIDA 21(2) SCHESVENTER, G., C.C. Frazier, L.A. ARANA, E.C. Bearss, R.W. CopeLanp, G. CUMMINS, J.C. Paice, and TLL. Wonc [Fort Matanzas Stabilization Team]. 1980. Historic structure report for Fort Matanzas National Monument, St. Johns County, Florida. Historic Preservation Branch, National Park Service, United States Department of the Interior, Denver, Colorado. ST. JOHNS County GIS Division. 2002. Fort Matanzas National Monument 2002 aerial imagery [map]. St. Johns County Geographic Information Systems, St. Augustine, Florida. Stacter, R.and S.C. Dia. 1984. Hammock vegetation of Little Talbot Island State Park, Florida. Bull. Torrey Bot. Club 111:494-497, STATE OF FLoripa. 2004. Online sunshine: official internet site of the Florida legislature. URL: www.leg.state.fl.us/. U.S. FisH AND WILDLIFE Service. 2004, Species information: threatened and endangered ani- mals and plants. URL: endangered.fws.gov/wildlife. html#Species/. Waaner, R.H. 1964. The ecology of Uniola paniculata L.in the dune-strand habitat of North Carolina. Ecol. Monogr. 34:79-96. Watersury, J.P. 1993. Coquina. Townet Press, Inc., St. Augustine, Florida. WUNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, Gainesville, Florida. WUNDERLIN, R.P.and B.F. Hansen. 2000. Flora of Florida, vol. 1,Pteridophytes and gymnosperms. University Press of Florida, Gainesville, Florida. Wunber tin, R.P.and B.F. HANSEN. 2003. Guide to the vascular plants of Florida, ed. 2. University Press of Florida, Gainesville, Florida. Wunpertin, R.P. and B.F. Hansen. 2004. Atlas of Florida vascular plants. URL: www.plantatlas.usfedu/. Zomerer, W.B. and D.E. Giannasi. 2005 [in press]. Floristic survey of Castillo de San Marcos National Monument, St. Augustine, Florida. Castanea 70. JUNIPERUS ASHEI (CUPRESSACEAE): PHYSIOGNOMY AND AGE STRUCTURE IN THREE MATURE TEXAS STANDS Caren McLemore University of North Texas Department of Biological Sciences Denton, Texas 76203, U.S.A. otanical Research Institute of Texas 509 Pecan Street, Fort Worth, Texas 76102-4060, U.S.A. Glenn C.Kroh John E. Pinder III IOUS Christian Uni iversi . Colorado ogee tant MEESIO partment of B Biolc Department of f E t Ith Fort worth Texas 761] 39, U SA. Fort Collins, Colorado 30523, U 6 7. ABSTRACT Despite the abundance of Juniper hei Buchholz in Texas, mature, intact | lativel This study compares structural patterns and growth dynamics among three mature stands on the Edwards Plateau and documents He AUOHSAIDS between pur uetura) changes ane rempor develop- ment of these forests. Each stand has varied By identifying and compan. these properties this study provides iaioamanon relevant to conser- vation an g g tof. ashei. RESUMEN A pesar de la abundancia de Juniperus ashei Buchholz en Texas, las agrupaciones maduras e intactas ] i um Pier ] J] ] ] i 4 7 J 7 j tr ya entre tres agrupaciones maduras en el altiplano Edwards y se docu tan las relaciones entre los cambios esa: y el desarrollo oa de —— Sane ee tat aeee tiene diversas ificar y comparar estas propiedades, este estudio oe informacion pertinente para la conservacion y decisiones de supervision relacionadas con J. ashei. INTRODUCTION Background Juniperus ashei Buchholz (Ashe juniper), one of the nine Texas species of the genus Juniperus (Correll & Johnston 1970; Simpson 1999), has dense popula- tions from the Ozark Mountains in Missouri and Arkansas, to the Arbuckle Mountains of northeastern Oklahoma, and is found throughout central Texas particularly on southern and eastern portions of the Edwards Plateau where it is the dominant woody species and formsa significant component of the state’s vegetation (Van Auken 1988; Diamond et al. 1995; Jackson & Van Auken 1997; Smeins et al. 1997). It also occurs in northeastern Mexico (Little 1992). Although SIDA 21(2): 1107-1120. 2004 1108 BRIT.ORG/SIDA 21(2 J. ashei has an overlapping distribution with both J. virginiana L. (eastern red cedar) and J. pinchotii Sudw. (redberry juniper), chemical analyses suggest that hybridization does not occur (Adams 1972, 1975; Kelley 1976; Flake et al. 1978). Juniperus ashei is typically found on thin, calcareous limestone- or dolo- mite-derived soils (Vines 1960) and also grows in deeper, sandier soils often in association with Quercus fusiformis Small (plateau live oak), Diospyros texana Scheele (Texas persimmon), Q. stellata Wang. (post oak), Q. sinuata var. breviloba (Torr) CH. Mull. (scaly-bark oak) and Q. buckleyi Nixon & Dorr (Texas oak) (Van Auken et al. 1978; Riskind @ Diamond 1986; Diggs et al. 1999). Co-occur- rence of J. ashei with broadleaf trees constitutes prime habitat for Dendroica crysoparia (golden-cheeked warbler), an end: d species which nests solely in juniper/oak woodlands and uses the bark from mature (> 30 yrs old) J. ashei trees as nesting material (Doughty & Parmenter 1989; Beardmore et al. 1995). Mature, second-growth J. ashei stands are rapidly disappearing due to high rates of urban and suburban expansion (Doughty & Parmenter 1989; Diamond et al. 1995; Patoski 1999). Effective land and endangered species management must include an understanding and appreciation of J. ashei’s role in establish- ing and maintaining stable, mature communities (Diamond et al. 1995) and its importance to the endangered golden-cheeked warbler. This study investigates the structure and dynamics of three mature J. ashei stands and provides information regarding the establishment and persistence of these stands. By identifying and comparing several structural and age-re- lated characteristics, it provides information relevant to conservation and man- agement decisions. Structural patterns and growth dynamics are compared among stands to document relationships between structural changes and tem- poral development of these forests. METHODS Study Areas The three study aus are on the Edwards Plateau of central Texas (Fig. 1) where eroded marine s , limestones, shales, and dolomites are covered by thin soil deposits (Riskind & Diamond 1988) on upland areas deeply dissected by streams. Precipitation, which averages 85 cm per year in the region of the study sites (Riskind & Diamond 1988), percolates downward to the water table, ex- pands fissures in the limestone, and forms the sinkholes, caves, and underground drainages characteristic of ‘karst’ topography (Spearing 1991). Two study sites are in Guadalupe River State Park, a 769 ha park in Comal and Kendall counties. The first site (hereafter Guadalupe South) is located south of the Guadalupe River on a 35 ha ‘karst dome’ (elevation = 385 m; N 29° 51' W 98° 30'). The second site (hereafter Guadalupe North) is north of the Guada- lupe River atop the river’s escarpment (elevation = 342 m; N 29° 52' W 98° 28)). MCLEMORE ET Al 1109 cc 4 200 miles Fic. 1. Map of T Both stands are on undulating, well drained, cherty clay loam soils where chert and limestone cobbles cover >20 percent of the surface and subsoil layers be- low 15-20 cm are 75%, by volume, limestone fragments (U.S. Department of Agriculture 1984). Soils at both locations are typical of the region. The third study site is on moderately deep, well-drained, loamy soil in Meridian State Park, a 204 ha reserve in Bosque County (elevation = 326 m; N 31° 53' W 97° 41’). Surface soil and subsoil layers are up to 38 and 94 cm deep, respectively, over a karst bedrock (U.S. Department of Agriculture 1980). Sampling Procedures Circular, 100 m2 plots were established at 20 m intervals along transects in each stand. Transects differed in length according to stand dimensions. All trees were identified to species, mapped by their distance and bearing to the plot center, and their diameter breast height (dbh) recorded. Trees with dbh >8 cm were marked with individually numbered aluminum tags and cored at the base us- 1110 BRIT.ORG/SIDA 21(2) ing a 4.3mm diameter increment borer. Dead or unhealthy trees were not cored. Cores were glued to wooden mounting boards and sanded toa flat surface. Rings were counted under 37.5x magnification. Because J.ashei forms false rings in response to environmental changes, ring number does not accurately reflect tree age. Therefore a formula for estimating age using ring counts was developed with cores from a separate set of trees of approximate known age from Meridian State Park. Photographs dating back to the park’s development in 1933-34 were analyzed to isolate specific areas de- void of J. ashei. Trees now present in these areas were assumed to have germi- nated immediately after the park’s establishment, giving them a maximum age (at the time of the study) of sixty-seven years. Cores from these trees were ana- lyzed and a formula was derived by () counting rings of each tree, (2) dividing approximate age by ring count, and (3) pooling results and computing a mean. Approximate age of each J. ashei in this study was then calculated using the resulting formula: ring count x 0.67. Large rays and the diffuse porous nature of the deciduous hardwoods made it impossible to accurately determine ages of those trees. Ring count information was used to determine forest age structure. Tree numbers, dbh, and height were used to determine mean height, mean basal area, size distribution, relative density (number of J. asheias a proportion of the total number of individuals of all species), relative frequency (frequency of J. asheiasa proportion of the sum of the frequencies for all species), and rela- tive basal area of each tree species. Importance values (Brower et al. 1998) were calculated. Measurements for height, basal area, and age were tested for normality and homogeneity of variance (Sokal & Rohlf 1973) in order to determine the appro- priate method of statistical analysis. All variables were normally distributed but displayed heterogeneity of variance, therefore non-parametric analysis of variance (ANOVA) was chosen to test for significant differences between stands. RESULTS Tree Species Identified and Importance Values Table | provides numbers of each tree species found at each study site. Only Juniperus ashei was common to all three sites. Relative density, frequency, and basal area of species may be summed to produce importance values (ranging from 0-3). Importance values integrate these separate measures to provide an indication of species influence in the community (Smith 1974). High importance values occurred for J. ashei at all three sites, with Guadalupe South at 2.70 and Meridian and Guadalupe North at 2.36 and 1.93, respectively (Table 2). These values indicate the dominance of this species in these communities MCLEMORE ET AL., PHYSIOGNOMY AND 1111 Taste 1. Summary counts of trees sampled. Scientific Name Common Name Guadalupe South Guadalupe North Meridian Juniperus ashei Ashe juniper 138 13] 86 Diospyros texana Texas persimmon = 3 38 0 Celtis laevigata hackberry 0 5 0) Ulmus crassifolia cedar elm | 6 0) Quercus texana Texas oak | 0 2 Quercus fusiformis plateau live oak 0 4 12 Quercus stellata 2 1 0 Quercus sinuata scaly-bark oak | 6 0) Fraxinus texensis exas ash 0 0 4 Sideroxylon lanuginosum gum bumelia 0 0 1 Total Sampled 146 19] 105 Taste 2. Relative density, relative frequency, relative basal area and importance values. Relative Density Relative Frequency Relative Basal Area Importance Values Guadalupe South Juniperus ashei 95 9 96 2.70 Diospyros texana 02 07 01 10 Ulmus crassifolia 01 03 01 05 Quercus texana 01 03 02 06 Quercus stellata 01 03 01 06 Quercus sinuata 01 03 .003 04 Guadalupe North Juniperus ashei 69 39 85 1.93 Diospyros texana 20 ny, 01 A8 Ulmus crassifolia 03 06 04 ls Quercus sinuata 03 04 003 07 Celtis laevigata 03 10 01 14 Quercus fusiformis 02 2. 07 21 Quercus stellata 01 02 01 04 ridian Juniperus ashei 82 62 92 2.36 Quercus fusiformis 11 19 O7 37, Fraxinus texensis 04 10 003 14 Quercus texana 02 05 01 08 lanuginosum 01 05 .001 06 Basal Area and Size Class Distribution Mean basal area of J. ashei was calculated for each site. Data indicate 29.58 m*ha™ (+114), 33.89 mZha! (=12.13), and 39.30 m2ha™! (=10.63) for Guadalupe South, Guadalupe North, and Meridian, respectively. Analysis of variance (Table 3) showed no significant differences between basal area of the three stands. 1112 BRIT.ORG/SIDA 21(2) Taste 3. Kruskal-Wallace one-way multisample non-parametric ANOVA with ties correction and x’ cantina oT tree bded area, height, and age at three sites. Mean sums of ranked scores are t differences at p<0.05 via Student-Newman-Keuls Multiple Range ae Guadalupe South Guadalupe North Meridian (n=84) (n=77) (n=66) Basal Area (cm?) 111.42 (a) 108.73 (a) 123.43 (a) r= ee ered Age (yrs) 128.2 (a) 134.5 (a) 72.0 (b) x’ = 38.53, p<0.0001 Height (m) 71.89 (a) 154.03 (b) 120.89 (c) ie Oar p<0.0001 Highest percentages of J. ashei at each site were in the smallest size class category (>30-300 cm?) with Guadalupe South at 45.3%, Guadalupe North at 48.5%, and Meridian at 36.5% (Fig. 2). Fewer than 5% of J. asheiat each site were in each of the four largest size class categories. Age Structure Ages of cored J. ashei were calculated and divided into five equal groups, 27-56 years, 5/-86 years, 87-116 years, 117-146, and 147-177 years. At all sites most trees were younger than 86 years and few were over 147 years (Fig. 3). Mean ages for the stands ranged from 80.4 years at Guadalupe North to 55.5 years at Meridian (Table +), and these differences were statistically significant (Table 3). There was no significant difference in mean ages between Guadalupe North and Guadalupe South. However, this result is believed to be due to the inability to determine ages of the many dead trees at Guadalupe North. Pattern for mean ages was reflected in the pattern for oldest trees (Table 4). The oldest trees at Guadalupe North and Guadalupe South sites were >150 years old. The oldest tree at Meridian was about the same age as the mean trees at both Guada- lupe sites and was less than half the age of Guadalupe North’s oldest tree. Height Mean heights were calculated for each tree species comprising >3 percent of each community. Juniperus ashei occupied the canopy at all three sites. At Guadalupe South and Guadalupe North, where the canopy was shared with other species, only Ulmus crassifolia (cedar elm) at Guadalupe North was taller than J. ashei (Table 5). Mean heights of J. ashei differed significantly among all sites with the greatest heights at Guadalupe North and the least at Guadalupe South (Table 3). Meridian had the highest rate of height increase (cm yr’!), grow- ing approximately 35% faster than Guadalupe South over the lifetime of the two stands (Table 6). DISCUSSION Mature Juniperus ashei dominated all three sites in this study; however, each stand had varied physiognomic characteristics and age-related structure. MCLEMORE ET Al 1113 Guadalupe South @ Guadalupe North < O Meridian =] o 5 Oo >30-300 >300-600 >600-900 >900-1200 >1200- >1500- >1800- >2100 150 1800 Size (Basal Area cm*) Fic. 7.Si | Pera ee £ If = L i] L 1 b o J. ashei (“%) w 5 Guadalupe South @ Guadalupe North O Meridian 20 10 re} ‘ 27-56 57-86 7-116 117-146 147-177 Estimated age (yrs) Fic 2 fj i Lat 4° tad g 1 dAictrih Tasle 4. Mean Juniperus ashei ages and oldest trees. Location Mean Age J. ashei (yrs) 5.D. Oldest J. ashei (yrs) Guadalupe South 773 *29.6 158 Guadalupe North 80.4 yy: 177 Meridian 55:5 *124 82 1114 BRIT.ORG/SIDA 21(2) Taste 5.Mean heights (m) of trees comprising >3 percent of each stand. Mean Height (m) Standard Deviation Guadalupe South Juniperus ashei 6.19 “1.37 Gu rth Juniperus ashei 7.93 *2.10 Diospyros texana 371 *0.50 Ulmus crassifolia 10.47 *3.45 Quercus sinuata 4.97 50.53 ridian Juniperus ashei 19 “1.11 Quercus fusiformis 79 *2.36 6.23 *1.92 Fraxinus texensis Taste 6. Juniperus ashei mean height increase (cm yr‘). Location Mean Increase (cm yr'') S.D. Guadalupe South 9.05 4.1/ Guadalupe North 11.57 * 4.99 Meridian 13.8] £3.23 Tree Species Identified and Importance Values Importance values were determined at all three locations (Table 2). The lowest J.ashei value occurs at Guadalupe North. Also present at this site is Diospyros texana (Texas persimmon), a small tree usually less than 12 m tall (Little 1992). Diospyros texana is exclusively an understory tree at this location, with the tall- est individual measuring 4.7 m. At Guadalupe South, with the highest J. ashei importance value, D. texand is rare (Table 1). Shading conditions make no sig- nificant difference in germination rates of D. texana (Everitt 1984). However, Diospyros species are reported to require full sun for optimum growth (Crock- ett 1972). At Guadalupe North these trees are often found clustered near dead J. ashei. These gaps in the canopy appear to provide ideal locations for the con- tinued growth of this species. Few D. texana occur under the canopy at Guada- lupe South, but may become more abundant as the stand ages and gaps are opened by tree death. Basal Area and Size Class Distribution Van Auken (1988) reported J. ashei mean basal areas of 38.6, 21.4, 43.2, and 18.4 m?ha'in four mature, undisturbed, woodlands computed from diameters mea- sured at 0.1 m above ground surface. These results are consistent with those for this study where mean basal areas of 29.6 (“11.4), 33.9 (=12.1), and 39.3 (+106) MCLEMORE ET AL 1115 m2 ha for J. ashei were measured at the Guadalupe South, Guadalupe North, and Meridian sites, respectively. Analysis of variance (Table 3) showed no significant differences between basal areas of the three stands. However, since Meridian is a significantly younger stand (Table 4), this indicates a faster basal growth rate at that loca- tion. Some of this difference may be accounted for by the faster growth rate of younger trees. However, deeper surface soil with its associated greater moisture- holding capacity is probably the most important factor (Bockheim 1982) influ- encing tree growth and may have led toa faster grown rate at Meridian than at the two Guadalupe River State Park locations wl thi and rockier. Highest percentages of J. dshei were in the smallest size class category (>30- 300 cm)at all three locations with Guadalupe South at 45.3%, Guadalupe North at 48.5%, and Meridian at 36.5% (Fig. 2). Less than 5% of J. ashei at each site were in each of the four largest size class categories. This arrangement results in a negative exponential size distribution, usually representing relatively early successional establishment (Van Auken 1993). However, size distribution does not necessarily reflect age distribution in forest age class studies. A generaliza- tion may be made that larger trees are likely to be old. However, it cannot be assumed that a small tree is young (Harper 1977). Many of the smaller J. ashei in this study were older than expected and size class distributions, in this case, do not indicate early successional stages, expanding populations, or relative youth of the majority of trees. However, size class distribution is useful in describing the condition of a population in terms of its future and may offer insights into reproductive per- formance. While reproduction is often analyzed in terms of age structure, quite often it is a function of size and can best be studied using size distributions (Harper 1977). Juniperus ashei cone production is partially determined by en- vironmental conditions, particularly rainfall, but is also dependent on tree size with trees reaching reproductive maturity at about 1.5 m height and about 50 cm? basal area. Results from this study indicate the majority of trees are repro- ductively mature and playing an important role in the reproductive dynamics of the population. — Age Structure False rings formed by many species of Juniperus (Panshin &@ Dezeeuw 1964) cause considerable difficulty in age determination. Van Auken (1993) believes it is impossible to accurately determine ages of junipers from growth rings due to formation of several rings each year in response to fluctuating rainfall. Fuhlendorf (1992; pers. comm.) reported an inability to differentiate true and false annual rings while determining J. ashei ages from ring counts. Adams (pers. comm.) expressed doubt concerning dating method accuracy for J. ashei (Adams et al. 1998). 1116 BRIT.ORG/SIDA 21(2) Methodology devised for age determination of J. ashei in this study is a novel approach based on ring counts of trees of known age. Although some conifers have a propensity to produce relatively more false rings when young and fewer when old, Grissino-Mayer has found no indication that Juniperus species have a tendency to do this (pers. comm.). Therefore, although the trees from Merid- ian were only 67 years old, the rate of false ring production should be similar to that of even the oldest trees in Guadalupe North. Although the trees were sampled at Meridian State Park, similar precipitation patterns at both parks also help validate use of the same formula constant for all three sites. Forest populations often progress as a sequence of even-aged cohorts initi- ated by disturbance. However, the mixed-aged structure characterized in this study (Fig. 3) indicates rarity of disturbance and infers continuous recruitment over the life of the stands (Kelly &@ Larson 1997). These stands appear to have escaped the relatively frequent fires that historically occurred in Texas at the time of their establishment (Smeins et al. 1997) and apparently have been fire- free throughout their existence. Interpreting age structure is complicated by the fact that there is no way to determine past mortality rates of a population. Age structure determination usually considers only survivors (as in this study) and does not utilize recruit- ment and mortality data (Harper 1977). However, accurate determination of stand age is dependent on mortality, as the oldest trees may be dead. This diffi- culty played a major role in determining the true age of Guadalupe North, where much of the forest was composed of dead trees. Despite these limitations, gen- eralizations can be made concerning age structure of these stands. Analysis of variance (Table 3) results indicated no significant differences between the ages of the two stands at Guadalupe River State Park. However, the stand at Meridian was significantly younger with a mean tree age of 55.5 years and no tree sampled older than 82 years (Table 4). All J. ashei at Meridian were in the two youngest age categories (Fig. 3). Stand age broadly corresponds to the establishment of the park in 1934. Much of the area now occupied by this stand was historically midgrass prairie (Riskind, pers. comm.), and the woodland’s presence demonstrates the ability of J. ashei to colonize many ter- rain types in the absence of fire. Mean ages for J. ashei were similar for Guadalupe South and Guadalupe North, with Guadalupe North results indicating a slightly (but not significantly) older stand (Table 4). Guadalupe South’s age distribution is typical of an aging population, with trees in age categories of 27-56 yr, 57-86 yr, and 87-116 yr al- most equally distributed (Fig. 3). Guadalupe North is the oldest stand with its establishment dating back toat least 170 years ago. Its greater age is reflected in the shift toward older trees (Fig. 3). It appears to be a declining population with relatively few individuals in the youngest 27-56 yr age category. Field observa- tions indicated many large, old, dead trees for which ages could not be deter- MCLEMORE ET AL 1117 mined. This difficulty caused an underestimate of the stand’s true age. There- fore, despite ANOVA results, Guadalupe North is believed to be older than Guadalupe South. Both Guadalupe North and Guadalupe South met some cri- teria for old-growth J. ashei stands as proposed by Diamond (1997) and perhaps could serve in refining the definition. Height Although J. ashei was the most abundant tree and dominated the canopy struc- ture, other species, notably Ulmus crassifolia, Quercus fusiformis, and Fraxinus texensis were present in the canopy. At Guadalupe North the J. ashei canopy, at 7.93 m, is overtopped by a number of U. crassifolia (Table 5). Whether the taller U. crassifolia are older or whether they grow more quickly than J. ashei could not be determined. Presence of Diospyros texana is also significant at Guada- lupe North. This species is primarily an understory tree as indicated by its mean height that is approximately half that of J. ashei. Quercus fusiformis and Fraxinus texensis have mean heights shorter than J. ashei but still share the canopy at Meridian (Table 5). This stand developed in a grassland or savanna environment and records indicate that the area was a cotton field prior to establishment of the park (Riskind, pers. comm.). There- fore, trees Snanne the canoe alee Sa) probably established concurrently. ANOVA different canopy heights between sites (Table 3). Although younger, Meridian had a greater mean height (7.19 m) than Guadalupe South (6.19 m). Reasons for this pattern are not known. However, deeper soils with greater moisture-holding capacity at Meridian could be one expla- nation for the observed height differences (Table 6). Tallest mean J. ashei height (7.93 m) is at Guadalupe North and differences between that site and Guada- lupe South may be due to Guadalupe North’s greater proportion of older trees. Information is lacking concerning height growth rates of J. ashei but it is historically considered slow-growing (Blomquist 1990). Based on tree ring analy- sis, J. pinchotii grows in height an average of 6.01 cm yr? for the first thirty years (McPherson & Wright 1989; Ueckert 1997). Juniperus ashei height growth rates may be expected to be similar to those of J. pinchotii but data from this study indicated greater mean height increases at 9.05, 11.57, and 13.81 cm yr! for Guadalupe South, Guadalupe North, and Meridian, respectively (Table 6). If J.ashei height growth rates are similar to those of J. pinchotii, discrepancies may be due, in part, to previous studies overestimating tree age from faulty interpre- tation of annual ring counts. Jackson and Van Auken (1997) recorded that J. ashei seedlings in edge habi- tats grow an average of 13.98 cm yr!. Their data are similar to height growth rates reported for Meridian. Deeper soils and the high light environment of open grassland during stand establishment may have resulted in relatively high rates of increase at this location. f 1118 BRIT.ORG/SIDA 21(2) Guadalupe South and Guadalupe North receive similar amounts of pre- cipitation and have similar soil depths. Growth rate discrepancies between the two sites are probably due to underestimating the true age of Guadalupe North. If the Guadalupe North stand is older than data from this study indicate, an adjustment downward in height growth rate would result, giving Guadalupe North a growth rate more similar to that obtained for Guadalupe South. ACKNOWLEDGMENTS We would like to thank David Riskind for suggesting this project and Texas Parks and Wildlife Department funding the research. Two anonymous review- ers provided helpful reviews. REFERENCES Apams, R.P. 1972. Chemosystematic and numerical studies of natural populations of Juniperus pinchotii Sudw. Taxon 21:407-425. Apams, R.P. 1975.Gene flow versus selection pressure and ancestral differentiation in the composition of species: analysis of populational variation of Juniperus ashei Buch. us- ing terpenoid data. J. Molec. Evol. 5:177-185. Apams, R.P., L.E. Flournoy, R.L. SINGH, H. JOHNSON, and H. Mayeux.1998. Invasion of grasslands by Juniperus ashei:a new theory based on random amplified polymorphic DNAs (RAPDs). Biochem. System. & Ecol. 26:371-377. Bearomorge, C., J. Hatrieto, and J. Lewis. 1995, Golden-cheeked warbler population and habi- tat viability assessment report. Grant Rept. U.S. Nat. Biol. Serv., Grant No. 80333-1423. Biomauist, K.W. 1990. Selected life history and synecological characteristics of Ashe juni- per on the Edwards Plateau of Texas. Unpublished M.S. Thesis, Texas A&M Univ., Col- lege Station. BockHeim, J.G. 1982. Forest soils.In:R.A. Young, ed. Introduction to forest science. John Wiley & Sons, New York. Pp. 93-111. Brower, J.E., JH. Zar, and C.N. von Enve. 1998. Field and laboratory methods for general ecology (fourth edition). McGraw-Hill Companies, Inc Corrett, D.S. and M.C. JoHNsToN. 1970. Manual of the vascular plants of Texas. Texas Res. Found., Renner. Crockett, J.U. 1972. Trees. Time-Life Books, New York. Diamonp, D.D. 1997. An old-growth definition for western juniper woodlands: Texas Ashe juniper dominated or codominated communities. Forest Service, U.S. Department of Agriculture. Gen. Tech. Rep. SRS—15. Diamonp, D.D., G.A. Rowett, and D.P. Keppy-Hector. 1995. Conservation of Ashe juniper Uuniperus ashei Buchholz) woodlands of the central Texas hill country. Nat. Areas J. 15:189-197, Dicas, G.M., Jr, B.-L. Liescome, and RJ. O'Kennon. 1999. Shinners & Mahler's illustrated flora of north central Texas. Sida, Bot. Misc. 16. MCLEMORE ET Al NOMY AND 1119 Doucuty, R.W. and B.M. Parmenter. 1989. Endangered species. Texas Monthly Press, Austin, Texas. Everitt, JH. 1984. Germination of Texas persimmon seed. J. Range Manag. 37: 189-192. Fiake, R.H., L. Urgatscu, and BL. Turner. 1978. Chemical documentation of allopatric intro- gression in Juniperus Syst. Bot. 3:129-144. FuHtenporr, S.D. 1992. Influence of age/size and grazing history on understory relation- ships of Ashe juniper. Unpublished M.S. Thesis, Texas A&M Univ., College Station. Harper, J.L. 1977. Population biology of plants. Academic Press, Inc., New York. Keitey, WA. 1976.|soenzyme investigations in Juniperus L.Unpublished Ph.D. Thesis, Colo- rado State Univ., Fort Collins. Ketty, RE. and D.W. Larson. 1997. Dendroecological analysis of the population dynamics of an old-growth forest on cliff-faces of the Niagara Escarpment, Canada. J. Ecol. 85: 467-478. Jackson, J.T.and O.W. Van Auken. 1997. Seedling survival, growth and mortality of Juniperus ashei (Cupressacae) in the Edwards Plateau region of central Texas.Texas J. Sci. 49: 267-278. Lirtue, E.L. 1992. The Audubon Society field guide to North American trees. Chanticleer Press, Inc., New York. McPuerson, G.R. and H.A. WricHT. 1989.Direct effects of competition on individual juniper plants: a field study. J. Appl. Ecol. 26:979-988. Pansuin, AJ. and C. Dezeeuw. 1964. Textbook of woad technology, volume 1 (second edi- tion). McGraw-Hill, New York. Paroski, JN. 1999. The war on cedar. Texas Month. 25(12):114-117. Riskino, D.H.and D.D. Diamonp. 1986. Plant communities of the Edwards Plateau of Texas: an overview emphasizing the Balcones Escarpment zone between San Antonio and Austin with special attention to landscape contrasts and natural diversity. In:P.L.Abbott and C.M.Woodruff, Jr, eds. The Balcones Escarpment, central Texas: geological society of America. Pp. 21-32. Riskino, D.H. and D.D. Diamonb. 1988. An introduction to environments and vegetation. In: B.B. Amos and FR. Gehlbach, eds. Edwards Plateau vegetation: plant ecological studies in central Texas. Baylor Univ. Press, Waco, Texas. Pp. 1-15. Simpson, B.J.1999.A field guide to Texas trees. Gulf Publishing Company, Houston, Texas. Smelns, F., S. FUHLENDORE, and C. Taytor, Jr. 1997. Environmental and land use changes: a long- term perspective. In: 1997 Juniper Symposium (Chapter 1). Texas A&M Univ. Press, San Angelo. Pp. 3-21. Sokat, R.R. and FJ. Rouir. 1973. Introduction to biostatistics. W.H. Freeman & Company, San Francisco. SpEARING, D.1991. Roadside geology of Montana. Ueckert, D.N. 1997. Biology and ecology of redberry juniper. In: 1997 Juniper Symposium (Chapter 3). Texas A&M Univ. Press, San Angelo, Texas. Pp. 3-10. Texas. Mountain Press Publishing Company, Missoula, 1120 BRIT.ORG/SIDA 21(2) Uniteo States DEPARTMENT OF Acricutture. 1980. Soil survey of Bosque County, Texas. United States DEPARTMENT OF AGRICULTURE. 1984. Soil survey of Comal and Hays Counties, Texas. Van Auken, O.W. 1988. Woody vegetation of the southeastern escarpment and plateau. In: B.B. Amos and FR. Gehlbach, eds. Edwards Plateau vegetation: plant ecological studies in central Texas. Baylor Univ. Press, Waco, Texas. Pp. 43-55. Van AukeNn, O.W.1993. Size distribution patterns and potential population change of some dominant woody species of the Edwards Plateau region of Texas. Texas J. Sci. 45: 199-210. VAN AuKeNn, O.W., A.L. Foro, A. Stein and A.G. Stein. 1978. Woody vegetation of upland plant communities in the southern Edwards Plateau. Texas J. Sci. 32:23-35. Vines, R.A. 1960. Trees, shrubs, and woody vines of the southwest. Univ. of Texas Press, Austin. HIGH RESOLUTION GIS MAPPING AND CURRENT STATUS OF THE TEN VIABLE POPULATIONS OF SHORT’S GOLDENROD (SOLIDAGO SHORTII-ASTERACEAE) IN KENTUCKY Brent D. Smith 2 0d ee aiOateal Sciences a aster Nc ky University Bt th , entucky 40475, U.S.A James B. Beck peparninen of Biological | Sciences Ea Kentucky 40475, U.S.A. Andrew 1. Denham Pepatemens er Siciogicd) Sciences Eastern University Richmond, Kentucky 40475, U.S.A. Cunentagal 2266 Ard Ridge Rd. ada ancy, Kentucky 42544 U.S.A. Patrick J. Calie' pu ered: Sciences University Easter Pepe entucky 40475, U.S.A. Richmo eee Department of Biology pat.calie@eku.e Washington University St. Louis, Missouri 63130-4899 U.S.A. ABSTRACT Short’s goldenrod (Solidago shortii Torr. & A. Gray) is an endemic species with a highly restricted distribution, the Kentucky populations occurring in and around the vicinity of Blue Licks in the northeastern ee of the state. The general occurrence of the species was first mapped in 1987, with several similar maps being published from 1989-2000. Due to changes in local land use prac- tices the status of the popwauogs ee rapidly oases Pane aS al obsolete. A census of 1] ] Ubcuncanies of etch Ee eee e) 1 c Fey £2 | - | : | Ic extant population Was Mapp! Between 1989 and 2003 four of gel populations were extirpated, eight dedineda in number cr stems present, and one increased in both number of stems and area coverage RESUMEN | ‘a oe 4 yd ron is 4” Sc lidc J s} i Torr & A \ = g a | . ] t sa enyalosa ees ae Blue Licks, al norteste de Renin, La digmabue ion general de las poblaciones de esta eSpeclen os eras pes aoe vez en 1987 y varios mapas han sido publicados entre 1989-2000. para cultivo, la distribucion original de las poblaciones de la especie se han modificado en gran medida, y en consecuencia los mapas custentes oa ebseleioy Un censo eee Weeks las Bepatones ob eee de campo, y * oe lest =) [ y la delimitacion fisica de cada nope De sie a 2003 eudue ee las pobl ginales han sepa pesciceis o, ocho disminuyeron I de tallos hertiura bs ‘author for correspondence: Department of Biological Sciences, 235 Moore Science ag 521 Lancaster Ave., Richn hmond, | Kentucky 40475-3124, U.S.A. Ph: 859-622-1505; fax: 859-622-1399; SIDA21(2): q21- 1130. 2004 1122 BRIT.ORG/SIDA 21(2) INTRODUCTION —_— Solidago shortii (Asteraceae) is listed as an Endangered Species in the Federal Register (Anonymous 1985). Charles Wilkens Short originally discovered speci- mens of the species growing on boulders at the Falls of the Ohio River in Jeffer- son County, Kentucky, in 1837. All remnants of those populations were either destroyed by inundation resulting from the construction of the McAlpine locks and dam to facilitate navigation on the Ohio River in 1925 (Buchele et al. 1989) or were extirpated in the latter half of the 19th century (Baskin et al. 2000). The species was “rediscovered” by E.L. Braun in 1939, in the vicinity of Blue Licks, KY (Braun 1941) (Fig. 1). The first map showing the spatial distribution of the Blue Licks populations was constructed in 1986 (Evans 1987). Several subse- quent reports contained maps of similar resolution (e.g. Buchele et al. 1989), with a new population being noted in Baskin et al. (2000 During the course of our field investigations Gai 1995-2003, we observed marked changes in the spatial size and occurrence of specific populations, due in part to local land-use practices and to local successional changes in several habitats. It became very clear that updated maps were needed to facilitate man- agement practices involving this species. This study was undertaken with two objectives in mind: |) to develop higher resolution maps of each known Kentucky population of Short’s goldenrod using Geographic Information Systems (GIS) cartographic technology; and 2) to up- date the status of each population first demarcated by Evans in 1987. Given the demonstrated utility of Geographic Information Systems technology in land use management (Longley et al. 1999) and species inventory applications (DeMers 1996), this application was a logical choice for developing accurate maps. MATERIALS AND METHODS Field Work-Population Census.—During the 2000 field season the boundary of each population first documented by Evans (1987) and Buchele et al. (1989) was defined cneues field surveys. Multiple transects through each population were established, dividing the population into parallel 3-meter-wide strips. The space between successive transects was then traversed, each individual S. shortii stem being counted. The majority of populations exhibit a linear rather thana polygonal distribution, which made this direct count approach technically fea- sible. This represents at best a minimal estimate, as undoubtedly some stems within the population boundaries escaped our detection, and some scattered plants do exist outside the demarcated boundaries. Field Work-GPS Coordinates.—GPS Lat/Long coordinates, taken in the degrees/minutes/seconds format, were determined with a Magellan 2000 hand- held GPS device at 50-meter intervals along the established perimeter of each population. Reference maps for each population were drawn in the field, using SMITH ET AL., CURRENT STATUS OF SOLIDAGO SHORTII IN KENTUCKY 1123 Fic. 1. Location of the populations of Short’s goldenrod in K ky. The “star” indicated the approximate location of the ec ¢ Inti . . f Fleming (F). Robertson (R) d Nicholas (N) ti measurements taken from local landmarks to orient field maps with aerial photographs and topographic maps (described below). Area estimates for each population were calculated from field measurements and combined with stem counts taken in 2001 to obtain density estimates for each population. GIS Mapping.—An Event Theme using the GPS data points was created in ArcView 3.0. The X coordinate was set to Longitude, the Y coordinate to Lati- tude, and the Projection was set to Lambert Conformal Conic. Aerial and topo- graphic images of the field research area were downloaded from the Kentucky Office of Geographic Information Systems (KYOGIS) website (ogisstate.ky.us/). The KYOGIS download included a file containing georeferencing information. A Line Theme was created to show the plants’ distribution. The plants’ locations and their proximity to landmarks visible on the aerial photo were verified and corrected based on our field observations. Other data sets were com- bined with the images and population lines. For example, a State Highways theme and Counties theme from Environmental Systems Research Institute (ESRD were added to show the locations of highways and county boundaries. Finally, we created the included maps (Fig. 2) using the ArcView Layout tool. All data and map files are available to appropriate scientific investigators and state and federal agencies upon request. RESULTS The ArcView system allows us to superimpose the GPS-derived data points for each population onto a number of high resolution cartographic interfaces, e.g,, an aerial photograph (Fig. 2A) or a topographic map (Fig. 2B). Comparisons with earlier maps are not quantitative due to differences in cartographic meth- odologies. Likewise, exact comparisons of surface area coverage for each popu- 1124 BRIT.ORG/SIDA 21(2) r rt be Naan tied | Fic. 2A. Example of a GlS-generated map showing the location of the known extant Kentucky populations of Short’s teh +b - Fo | 4 Had and #16 RQ yi L = £ I *: £1] re (41n07)\ Solid goldenrod " } pace ee CBSA et " + ey ae Th +h ££ tl is the Licking River, and lation t dari hown in red. Only those populations found on properties SMITH ET AL., CURRENT STATUS OF SOLIDAGO SHORTII IN KENTUCKY 1125 tere Fic 2B The id . I shar fin in ff JA 1126 BRIT.ORG/SIDA 21(2) lation relative to an earlier study (Buchele et al. 1989) are difficult due to differ- ences in sampling techniques. However, we are confident that both sampling lologies are of sufficient accuracy to allow for general comparisons. Two populations found on lands in private ownership (#11 and #15) are not shown on our maps to provide those populations with some measure of protection. The census results, rather dramatic when com pared to earlier stem counts, are summarized in Table 1. Based upon comparison with earlier stem counts and surface area estimates (Buchele et al. 1989) the populations cluster into the following categories. Decline.—This is the status of populations #1-4, #7, #11 and #12, all hav- ing declined in stem numbers ranging from a 1.25-fold to 33-fold. Population #6 exhibits a larger fold decrease, now consisting of only one stem. We con- sider this population to be extirpated. A concomitant decrease in surface area isalso seen in population #1-4, #7 and #8. While experiencing a slight decline in stem number (1.25-fold), population #12 has expanded its surface area by 13- fold. Census data for population #11 are not available for 2002, as we were un- able to obtain permission from the private property owner to examine this site. GIS data were obtained for this population along a bordering road, and mapped from prior field observations. Extirpated.—Population #6 occurs along a highway right-of-way, and isa remnant of a once larger population that was intentionally destroyed by a lo- cal landowner. There is now a single stem remaining. The original population #10 (Evans 1987) was destroyed by a local landowner (Mr. Allison, pers.comm.). We discovered a remnant set of eight plants in 1998, ca. 100 meters east of the original locality. This set is now extirpated due to local successional changes, e.g., increased canopy cover from arborescent species (primarily Quercus spp.) Increase.—Population #5 is the only population that has increased in stem number, from 530 in 1989 to an estimated 3,488 in 200L This is accompanied by an increase in coverage area from 870 m2 in 1989 to 5,380 m2 in 2001. This is now the largest Kentucky population of Short’s goldenrod in terms of both area and stem number. Doubtful reports.—Population #14, located on private property (the Kingsolver farm near Blue Licks Battlefield State Park) has been of doubtful determination since its “discovery”. There are no voucher specimens from this site, and repeated efforts by the authors in 1998 and 1999 and by D. White (Ken- tucky State Nature Preserves Commission, pers.comm.)to locate S. shortii plants met with failure. This population, if ever extant, is now extirpated Recently discovered population.—Population #15 was discovered by Mr. Nick Drozda of the KSNPC during a survey of a bison trace (trail) in 1998. This population was revisited in 2003 and found to have increased in number of stems and distribution. 1 Tee Taste 1.Summary of population census and area of coverage of all known verified populations of Short’s goldenrod in Kentucky. Data from 1989 are those of Buchele et al. (1989), from 2001-2003 are from this study. The numbering of populations #1-13 is that of Evans (1987). Population #14 was designated and first mapped by Baskin et al. (2001), and #15 is the recent discovery by N. Drozda. The “First Report” entry refers to the earliest notation in status ous maintained by the USFWS and provided by D.White of the KSNPC. KSNPC = Kentucky State Nature Preserves Commission; P = private ownership; KSPC = Kentucky State Parks Commission; KHC = Kentucky ey Commission; ROW = Right-of-Way; USFWS = United ae Fish and Wildlife Service. *extirpated = population consisting of 1 plant, but in essence extirpated; **14 = dubious report; ***15 = stem number estimate provided by N. Drozda (pers.comm.). Population Stem # Fold increase/ Area m2 Fold increase/ Density Ownership First Report 1989/2001 1989/2002 (# stems/m2) 1 42,000 / 2,549 16-fold decrease 4,600 / 3,027 1.5-fold decrease 9.13 /0.842 KSNPC/KSPC 1936 2 10,150/ 573 18-fold decrease 12,840 / 1,367 9.4-fold decrease 0.790 / 0.419 KSPC 1983 3 3,500 / 193 18-fold decrease 4,500 / 300 15-fold decrease 0.778 / 0.642 P 1987 4 1,400 / 42 33-fold decrease 1,290 / 193 7-fold decrease 1.08 /0.217 KSNPC / KSPC 1936 (?) 5 530 / 3,488 7-fold increase 870 / 5,380 6-fold increase 0.609 / 0.648 KSNPC 1983 6 2,100 / 10 (00), 1 (02) “extirpated Sills al NA NA P / KDOT (ROW) 1986 7 6,300 / 1,000 6-fold decrease 6,230 / 524 12-fold decrease 1.01/1.91 KDOT (ROW) 1957 8 1,780 / 672 3-fold decrease 2,570 / 766 3-fold decrease 0.692 / 0.877 KSNPC 1934 9 640/<25 (00), 3 (02) extirpated 2,485 / 1 NA NA P (ROW) 1987 10 240 / 13 (00), 0 (02) extirpated 15/0 NA 16.0/0 P / KDOT (ROW) 1985 11 500 / 800 3-fold decrease 265 /NA NA 943 /NA P 1985 12 2,300 / 1,846 1.25-fold decrease 390/4,877 13-fold increase 5.89 / 0.378 USFWS (ROW) 1939 13 180/0 extirpated Feb-00 NA 90.0 /0 P 1987 mA 15-20(?) / 0 extirpated(?) NA NA NA P 1989 mS NA / 100 (03) NA NA/120 NA NA /0.83 P 1998 AMDNLNAY NI ILLYOHS OOVGITOS 4O SNLWLS LNIYHND “TV 13 HLIWS Z7LL 1128 BRIT.ORG/SIDA 21(2) DISCUSSION Given the state of mapping technology in 1987 (and the passage of time) it is not surprising that maps derived through GIS applications are different from the original maps. What is of particular note is the general numeric decline in all but one population (#5) of S. shortii. Populations #1, 2, and 4 have been within the jurisdiction of the Kentucky State Fans Commission during this period of comparison, yet all three have suff 1assive decline. An unequivo- cal cause for the decline of these three and/or any wohet populations under pro- tection (i.e, populations #5 and 8, under the jurisdiction of the KSNPC, and population #12, under the USFWS) is not clear. The spatial distribution patterns of the populations fall into two catego- ries. The first we term “linear” (populations #1, 3-4, 6-9, 11 and 12). Several of these populations occur along either power line or highway rights-of-way (Table 1), but others occur in what appears to be uninterrupted habitat (e.g. #4) con- forming toa linear pattern. This could be due to localized edaphic conditions, as the preferred habitat is one with shallow soils. The second category we refer to as “polygonal” (populations #2 and #5). These habitats are continuous areas uninterrupted by roads. The plants, however, are distributed discontinuously throughout the mapped area, the perimeter of the populations assuming an irregular geometric outline. The precise reasons for this difference in spatial distribution patterns is at present unknown but could be due to subterranean factors, e.g., rock shields underlying shallow soils and providing an unsuitable habitat for competing species. Alternatively, the plants’ distribution could be due to seed dispersal patterns of those specific populations. At present both of these postulates remain untested. The results of the stem count comparison are dramatic. Numbers of stems has decreased since 1989 for all observed populations except #5. This popula- tion exists in an old field that is currently under the management and owner- ship of the Kentucky State Nature Preserves Commission. The results of the area estimates are equally dramatic, since all of the observed populations ex- cept for #5 have decreased in area since 1989. Population #1 has experienced the most marked decline, most likely due to improper management and in- creased development and use of park recreation facilities by visitors. This popu- lation is now dissected i into several distinct groups, each following either a road or a power line right-of-way or a former bison trace remnant. The glade area within the park where the plants were previously observed in abundance is now populated primarily by S. nemoralis. A similar situation exists in popula- tion #2. The impact of S. nemoralis on the long-term persistence of S. shortii is not known, although we have observed a steady decline in these two popula- tions (Ta Three populations have been extirpated, one by natural means and two by SMITH ET AL., CURRENT STATUS OF SOLIDAGO SHORTII IN KENTUCKY 1129 human intervention. Population #13 occurred in an open field and consisted of 180 stems in 1989 (Buchele et al. 1989). The field was subjected to mowing in the early years of the 1990s, but in the latter half of the decade the field was not cultivated. Competition from non-native grasses (e.g, Festuca) and forbs (eg., Lespedeza) have contributed to the loss of this population. Population #6 grew in an actively grazed pasture and was removed by the landowner in 1988 by bulldozing the habitat (USFWS records and D. White, pers. comm.). In discus- sions with the local landowner it was revealed that population #10 was elimi- nated through extensive mowing of caulescent stems and the deposition of con- crete debris on the persistent rosettes, beginning prior to 1995. Population #11 occurs along a woodland edge and appeared rather stable over the period of this study. The status of a questionable population has also been resolved. Population #14 was first cited in Baskin et al. (2000). We were unable to locate plants in the field during 1998 and 1999, and there was confusion regarding the accuracy of the original species determination (D. White, pers. comm.). Records maintained by the USFWS indicated ca.15-20 stems of “Short’s goldenrod” in 1989, but none was found in either 1997 or 1998. Independent efforts by D. White (pers. comm.) were also unsuccessful. If this population of Short’s goldenrod ever existed, it is clear that it is now extirpated. In 2000, a new population was discovered by N. Drozda of the KSNPC along a former bison trace (trail) in Fleming County. This population consisted of ca.25 stems, and the plants were described as “depauperate” (USFWS records). The population, designated as #15 in Table 1, persisted into 2003 and now con- sists of ca. 100 stems concentrated along the bison trace with a few individuals scattered in the adjacent woods. In 1995 seven “clumps” of cultivated S. shortii (originally obtained froma Blue Licks population) were planted on the Indiana shoreline of the Ohio River, across from the type locality at the Falls of the Ohio River in an effort to rein- troduce the species into suitable riparian habitat (Homoya 1996). These plants were lost in the following year due to increased water flow from winter runoff (D. White, pers. comm.). A recent report describes a population of Short’s gold- enrod in Indiana (wwwin.gov/dnr/public/novdecOl/newsl.htm). This is the first verified record of Short’s goldenrod outside of Kentucky (a voucher speci- men has been deposited at MOBOT), the site occurring within a former migra- tion pattern of the extinct eastern woodland bison (Bison bison L.). This could represent a very old population that has been genetically isolated from the Ken- tucky populations for perhaps several centuries. Alternatively, this population could consist of escaped colonizers from the 1995 effort to reintroduce the plants in Indiana, as the newly discovered population is ca. 50 nautical miles down- stream from the attempted reintroduction. Genetic analyses of samples from 1130 BRIT.ORG/SIDA 21(2) both the Kentucky and Indiana populations are now underway in an effort to resolve this issue. ACKNOWLEDGMENTS Appreciation is extended to the Wilderness Road Girl Scout Council Troop #1205 for their mapping of population #4, and to Bruce Davis and the GIS Map Laboratory of Eastern Kentucky University for assistance with GIS software and map generation. Funding in support of field and computational activities was obtained through a Sigma Xi Grant-in-Aid of Research JBB), through con- tracts #20379 and #99161405 from the Kentucky State Nature Preserves Com- mission and the US Fish and Wildlife Service, two EKU Faculty Research Awards and NIH/NCRR Award #P20 RR16481-SI (PJC). Discussions with Marc Evans and Deborah White, and the field asians of Tiffany Carpenter and Chris- tina Shackleford a tefully acknowledged. Nick Drozda escorted PC to popu- lation #15 and oe critical background information. The comments and reviews of Moriah Beck and Rob Naczi on the manuscript, and assistance from Sara Fuentes and Lupita Sanchez with Spanish translation are appreciated. REFERENCES Anonymous. 1985. Listing of threatened and endangered species. Fed. Reg. 50:36085-36089. Baskin, J.M, J.L. Watck, C.C. Baskin, and D.E. Buchete. 2000. Ecology and conservation of the endangered plant species Solidago shortii (Asteraceae). Native Pl. J. 1:35-41. Beck, J.B., R.C Naczi,and PJ. Catt. 2001. Insights into the species delineation and popula- tion structure of Solidago shortii (Asteraceae) through morphometric analysis. Rhodora 103:151- BucHete, D.E., J.M. Baskin, C.C. Baskin. 1989, Ecology of the endangered species Solidago shortii |. Geography, populations and physical habitat. Bull. Torrey Bot. Club 116:344—355. Braun, E.L.1941.A new jecallty for Solidago shortii,. Rhodora 43:484. DeMers, T.1996.Remote sensing and geographical information systems: Spatial technolo- gies for preserving phytodiversity. In: T.F. Stuessy and S.H. Sohmer, eds. Sampling the green world. Innovative concepts of collection, preservation, and storage of plant di- versity. Pp. 125-139. Evans, M. 1987. Short's goldenrod recovery plan. Unpublished Technical Agency Draft. USFWS, Southeast Region, Atlanta, GA Homoya, M. 1996. The return of Short’s goldenrod. Endang. Spec. Tech. Bull. 21:24—25. Lonctey, P.A., M.F. GoopcHitp, DJ. MaAcuire, and D.W. RHino (eds.). 1999. Geographic Informa- tion Systems: Principles, Techniques, Applications and Management. Second ed. John Wiley and Sons, New York. AN ANNOTATED, PRELIMINARY CHECKLIST OF THE VASCULAR FLORA OF CAMP BUTNER, NORTH CAROLINA Alexander Krings Carlyle Franklin Herbarium, Department of Botany Department of Forestry North Carolina State University North Carolina State University Raleigh, North Carolina 27695-7612, U.S.A. Raleigh, North Carolina 27695-8006, U.S.A. Alexander_Krings@ncsu.edu Carlyle_Franklin@ncsu.edu ABSTRACT For the past five years, the Woodlot Forestry Research and Development Program at North Carolina State University has been assisting with the implementation of the Land Condition Trend Analysis (LCTA) program at Camp Butner. A es ant inventory has been an ongoing aspect of associated re- search. This checklist i inventory of the flora of the site, comprising 78 fami- c t lies, 178 genera, and 241 species. RESUMEN En los cinco anos anteriores, el Woodlot Forestry Research and Development Program de la Univer- sidad del Estado de Carolina del Norte ha estado ayudando con la umipleientacion del programa Land Condition Trend Analysis (LCTA) en Camp Butner. El inventario vegetal ha sido un aspecto de a investigacion asociada. Este listado representa un inventario preliminar de la flora local, que comprende 78 familias, 178 géneros, y 241 espec The Camp Butner National Guard Training Site (Camp Butner’) in Durham and Granville counties, North Carolina comprises 1975 hectares and includes pine plantations, mixed pine-hardwood forests, and bottomland hardwood forests. For the past five years, the Woodlot Forestry Research and Development Pro- gram at North Carolina State University has been assisting with the implemen- tation of the Land Condition Trend Analysis (LCTA) program at Camp Butner (WEFRDP 2003). The Land Condition Trend Analysis (LCTA) inventory was instituted by the US Army Construction Engineering Research Laboratory USACERL) in order to monitor the natural resources on military installations and to provide information for making sound training and land management decisions (Tazik et al. 1992). In association with annual vegetation monitoring, land use assessment, and wildlife monitoring, a plant inventory has also been an ongoing aspect of research at Camp Butner since 2001. This checklist repre- sents a preliminary inventory of the flora of the site. METHODS To inventory the vascular plants of Camp Butner, the site was visited numer- ous times between 2001-2003. Principal collectors of the flora associated with our project include: C. Wiecek, V. Miller, C. Sheats, and K. Summitt. Batson (1952) SIDA 21(2): 1131-1139. 2004 1132 BRIT.ORG/SIDA 21(2) also collected in the area. Collected specimens were pressed and dried using standard herbarium techniques and identified using the collections of the North Carolina State University Herbarium (NCSC). RESULTS AND DISCUSSION Seventy-eight families, 178 genera, and 241 species are currently known from Camp Butner (Table 1). Ruellia purshiana Fern. (Acanthaceae) is known from the site (M. Franklin, pers.comm.), but has not been collected. Taxa are arranged alphabetically within the major subgroups of ferns, gymnosperms, dicots, and monocots. The number of genera, then species follows each family name. Brief habitat descriptions follow each species entry when available. Landscape fea- tures for the site are discussed by Hall 1995). Taxonomy follows APGII (APG 2003). Nomenclature primarily follows USDA, NRCS (2002). Specimens are deposited at NCSC, unless otherwise indicated. The composition of the flora is typical for central North Carolina—the site hosts mixed second growth woodlands, pine plantations, and clearcuts (see also Palmer 1990; Hall 1995). However, compared to other near sites, the flora of Camp Butner is noticeably depauperate (Table 2). The flora of Umstead State Park, which is somewhat larger than Camp Butner, includes about three times as many species (Sawyer 1968). Even floras of sites smaller by an order of magni- tude comprise 1.5 to 2 times as many species (Table 2). We suspect two main reasons to explain the relative species poorness of the Camp Butner flora. The majority of the site was not forested as late as 1950 based on black and white (1 to 60,000) photography flown by the US Army in November, 1950. The site now hosts forests that are under higher disturbance regimes from military land use than “nature preserves” such as Durant Park (Skean 1982) or Umstead State Park Sauls 1968). Prior to its current ownership, Camp Butner lands were in to- ultivation and pastures, as well as host toa WWII prisoner of war camp ana artillery range. ampling intensity could also be an important factor. Based on comparisons with other floras (Table 3), it appears evident that several large families, includ- ing Asteraceae, Fabaceae, Cyperaceae, and Poaceae, remain undersampled in our study and that future efforts must concentrate on increasing representation FERNS AND ALLIES ASPLENIACEAE 1/1 DENNSTAEDTIACEAE 2/2 Asplenium platyneuron (L.) B.S.P. (Sheats 106; = Dennstaedtia punctilobula (Michx.) T. Moore Wiecek 68) Pine-hardwood mix. (Sheats 92) BLECHNACEAE 1/1 eae areolata (L.) T. Moore (Sheats 107) ne-hardwood mix DRYOPTERIDACEAE 3/3 Athyrium filix-femina (L.) Roth ssp. asplenioides (Michx.) Hultén (Summitt 148) Near stream. Pteridium aquilinum (L.) Kuhn (Sheats 138) Pine orest CAMP BUTNER, NORTH CAROLINA 1133 Taste 1. Summary of numbers of families, genera, and species of Camp Butner. Ferns and allies Gymnosperms Dicotyledons Monocotyledons Total Families 8 2 58 10 78 1] 2 139 26 178 Species 11 4 192 34 241 Taste 2. Comparative floristics of Camp Butner to other sites (F = families; G = genera; S = species). Camp Butner’ __Yates Mill Durant Nature = White Pines Umstead Pond? Park? Natural Area* State Park® (ca. 4880 ac) (ca. 180 ac) (ca. 237 ac) (ca. 242 ac) (ca. 5439 ac) F Gr <5; «JF G § F “~G s& F Gan seri G ‘Ss Ferns andallies 8 lyr bh. 28 11 15 8 13 18 8 14 19 9 14 15 Gymnosperms 2 2 4 2 2 4 2 2 4 2 2 6 Dicotyledons 5 139 192 72 164 24783 206 337 86 241 39892 290 529 Monocotyledons 10 26 34 11 56 96 11 75 12814 73 13915 90 186 Total 78 178 241 93 233 362 104 296 487 110 330 562 118 396 734 'Present study; Jones (1971); *Skean (1982); *Swab (1990); “Sawyer (1968). Taste 3. Comparison of species richness of four large families at Camp Butner and other sites. Camp Butner’ — Yates Mill Pond? ~— Durant Nature Park? White Pines Natural Area* Asteraceae 36 35 54 66 Fabaceae 22 2] 30 30 Cyperaceae 3 16 24 28 20 47 62 64 'Present study; Jones (1971); *Skean (1982); “Swab (1990). Onoclea sensibilis L.(Summitt 153) Near stream. POLYPODIACEAE 1/1 Polystichum acrostichoides (Michx.) Schott — Pleopeltis polypodioides (L.) Andrews & Windham (Summitt 163; Wiecek 63) Near stream; Pine (Sheats 203) Bottomland hardwoods. plantation. THELYPTERIDACEAE 1/1 LYCOPODIACEAE 1/1 Thelypteris noveboracensis (L.) Nieuwl. (Summitt Sheats 223) Near stream. — Lycopodium digitatum Dill. ex A. Braun 95) Pine-hardwood mix. OSMUNDACEAE 1/1 Osmunda cinnamomea L. (Summitt 222) Near CUPRESSACEAE 1/1 stream. Juniperus virginiana L. (Sheats 98) Pine-hardwood MIX. GYMNOSPERMS 1134 PINACEAE 1/3 Pinus echinata i a 94) Pine-hardwood upland mix. Pinus taeda L. Gun 136) Pine forest. Pinus virginiana P. Mill. (Sheats 127) Pine forest. ANGIOSPERMS BASAL ANGIOSPERMS AND EUDICOTS ACANTHACEAE 1/1 Ruellia caroliniensis 218) Roadside. J.F. Gmel.) Steud. (Summitt jeter ue im L.(Summitt 213) Near lake. Vieuiatm create ium L.(Batson 1274, DUKE Viburnum rafinesquianum J.A. Schultes (Sheats 147) Bot tomland hardwoods. ALTINGIACEAE 1/1 Liquidambar styraciflua L. (Sheats 77) Upland hardwood mix ANACARDIACEAE 2/2 Rhus copallina L.(Sheats 115) Upland hardwoods. oxicodendron radicans (L.) Kuntze (Sheats 101) Pine-hardwood mix. ANNONACEAE 1/1 Asimina parviflora (Michx.) Dunal (Wiecek 15 Bottomland hardwood forest. APOCYNACEAE 2/2 Apocynum cannabinum L. (Summitt 210) Road- side. Asclepias tuberosa L. (Sheats mix. AQUIFOLIACEAE 1/2 llex decidua Walt. (Batson 1029, DUKE llex opaca Sol. (Sheats 113) Upland hardwoods. ARISTOLOCHIACEAE 1/4 Hexastylis lewisii (Fern.) H.L. Blomaq. & Oost. (Batson s.n.) Bluff. — minor (Ashe) HL. cle (Wiecek 14) miand hardwood fores ei 5 shuttleworthii (Britten : Baer f.) Small Summitt 131) Lake edge Hexastylis virginica (L.) Small (Sheats 75) ASTERACEAE 27/36 Achillea mi jemi L. (Sheats Summitt 125) Roadside. WS wae 175) Roadside/pine 125, Sheats 176; BRIT.ORG/SIDA 21(2) Ambrosia artemisiifolia L. (Wiecek 39) Upland hardwoods. Antennaria plantaginifolia (L.) Richards (Batson 1314, DUKE) Anthemis arvensis L Leen Roadside. istosa Mi .) Britt. (Miller 27) Xeric site, near erosion ) monitoring plot. Chrys uM virginianum L. (Sheats 81;Summitt 81) | Roadade: ins near stream. Chrysopsis mariana (L.) Ell. (Miller 26; Wiecek 47) she = hardwoods; Xeric site, near erosion monitoring ae ing. Cirsium ee ee (Summitt 204; Sheats 126) Roadside. vanes — ae Cronq. (Sheats 221) C a L.(Sheats 86) Alluvial forest. ies) ay — Eleph ee tomentosus L.(Batson 1290, DUKE) Erechtites hieracifolia (L.) Raf.ex DC. (Sheats 222) Erigeron annuus (L.) Pers. (Sheats 108) Upland — ardwoods. Eupatonium el ifolium (Lam.) Small (Wiecek 71) Pine-hardwood mix Eupatorium nner ifolium L. (Batson 655, DUKE) mee cole rotundifolium L. ine 183; Wiecek ine plantation/road sea amarum (Raf.) ae (Sheats 16/7, Sheats 179; Summitt 220) Roadside; pine mix. ree autumnale L. a 78) Roadside/ pine-hardwood m Hieracilum gronovil > (Wiecek 5, Wiecek 48) Loblolly pine plantation and clearings; Up land hardwoods Hieracium venosum L. (Sheats 73) Pine-hardwood mix. 3 Krigia virginica (L.) Willd. (Summitt 206) Roadside. leucanthemum vulgare Lam.(Summitt 60; Wiecek 60) Roadside; Pine plantation Liatris squarrosa (L.) Michx. (Sheats hardwood mix/roadside. Packera anonyma (Wood) W.A.Weber & A. Love (Summitt 166) Roadside. Pluchea foetida (L.) DC. (Sheats 140) Pine forest. Pseudognaphalium obtusifolium (L.) Hilliard & tt (Summitt 117; Wiecek 7) Roadside; loblolly pine plantation Seriocarpus asteroides (L.) B.S.P. (Sheats 1 37, Sheats 72; Su 199) Pine- ~ mmitt 221) Roadside pil LA sa mi . Solidago caesia L.var.curtisii (Torr.& Gray) Wood (Wiecek 52) Upland hardwoods. F CAMP BUTNER, NORTH CAROLINA 1135 Colida qAnadenc g L. var. scabra Torr. & A. Gray (Batson 640, DUKE) Solidago nemoralis Ait. (Batson 625, DUKE) Solidago odora Ait. (Sheats 207) Roadside. Solidago pinetorum Small (Sheats 163) Roadside. Solidago roanensis Porter (Miller 23) Xeric site, near erosion monitoring plot. Solidago speciosa Nutt. var.erecta (Pursh) MacM. (Wiecek 58) Slope near creek Symphyotrichum dumosum (L.) Nesom ( 44) Upland hardwoods. Taraxacum officinale G.H. Weber ex Wiggers (Summitt 183) Roadside. BALSAMINACEAE 1/1 Wiecek Impatiens capensis Meerb. (Sheats 223) Creekside BETULACEAE 5/5 Alnus | a Aj .) Willd (Summitt 207) Stream- side. Betula nigra L.(Sheats 193; Summitt 211) Bottom- land hardwoods; streamside. Carpinus caroliniana Walt. (Sheats 91) Pine-hard- Woo mix. Corylus americana Walt. (Miller 20) Xeric site, near erosion monitoring plot Ostrya virginiana (P. Mill.) K. Koch (Summitt 214) BIGNONIACEAE 1/1 Campsis radicans (L.) Seem. ex Bureau (Sheats 103, Sheats 194) Pine-mix. BRASSICACEAE 4/4 Barbarea verna (P. Mill.) Aschers. (Summitt 129) Roadside Cardamine hirsuta L. (Summitt 187) Roadside near stream-crossing. Draba verna L.(Summitt 185) Roadsi Senin (L.) Ait.f.(Summitt a Road- side CAMPANULACEAE 1/4 iecek 35) Knap of Reeds Lobelia cardinalis L. (Wi Creek. Lobelia inflata L.(Wiecek 4) Loblolly pine planta- tion. Lobelia puberula Michx. (Wiecek 38) Knap of Reeds Cre Lobelia spicata Lam. (Sheats 109) Upland hard- woods sialic, 6 1/1 nicera hes Thunb. (Sheats 96) Pine-hard- ood m CARYOPHYLLACEAE 2/2 Cerastium fontanum Baumg. ssp. vulgare (Hart- Greuter & Burdet (Summitt 179) Road- side. Scleranthus annuus L. (Sheats 214) Roadside/ pine-mix. CISTAC 1/1 Lechea pan: a Raf. (Sheats 198) CLUSIACEAE 1/4 Hypericum gentianoides (L. Quarry. mon B.S.P. (Sheats 197) Hypericum eee (L.) Crantz (Sheats 144, Shea 6; Wiecek 6, Wiecek 19) Loblolly pine ee bottomland hardwood forests/ roadside. Hypericum nudiflorum Michx. ex Willd. (Sheats 201) Bottomland hardw Hypericum punctatum ene 189-Summitt 217) Bottomland hardwoods; roadside CONVOLVULACEAE 1/1 Ipomoea pandurata (L.) G.FW. Mey. (Miller 32; Sheats 173) Xeric site, near erosion monitor- ing plot; roadside/pine m CORNACEAE 1/1 Cornus florida L. (Sheats 80; Summitt 156) Pine- hardwood mix; roadside. EBENACEAE 1/1 Diospyros virginiana L. (Sheats 218) Pine mix. ERICACEAE 6/8 Chimaphila maculata (L.) Pursh (Sheats 111; Summitt 82) Upland hardwoods. Gaylussacia frondosa (L.) Torr. & Gray ex Torr. Summitt 215) Leucothoe racemosa (L.) Gray (Wiecek 16) Bot- omland Bet ies forest ydend arboreum (L.) DC. (Sheats 88, Sheats 142, Sheats 220) Pine: hardwood upland mix; bottomland hardwoods. Rhododendron Ber ao: Shinners (Sheats 89) Vaccinium fuscatum Ait. (Summitt 84) Upland hardwoods. Vaccinium a idum Ait. (Sheats 119, Sheats 209) orest — Michx.) 1136 Vaccinium stamineum L.(Summitt 145; Wiecek 62) Roadside; Pine plantation. EUPHORBIACEAE 2/3 Bete maculata (L.) Small (Sheats 165) Uplan ee er cb : ee 188) Bottomland hardw Euphorbia ie Pursh (Wiecek 37) Knap of Reeds Creek FABACEAE 16/22 Albizia julibrissin Durazz. (Sheats 177) Roadside. Amphicarpaea bracteata (L.) Fern.(Wiecek 40) Upland ha rdwoods. Centrosema vi rgit vanum (L ) Benth. (Sheats 211 eh is Canageris L. neat ] — <5 reg: 157) Pine forest. Chamaecrista nictitans (L.) Moench (M Xeric site, near erosion monitoring plot. Cytisus scoparius (L.) Link (Sheats 168) Pine mix. Desmodium nudiflorum (L.) DC. (Summitt 216) Roadside near lake Desmodium paniculatum (L.) DC. (Batson 929, Miller 21) UKE) Desmodium rotundifolium DC. (Wiecek 43) Up ods. (L.) Britt. (Batson 980, DUKE) 3) Pine-hard- land hardw Galactia volubilis Gleditsia triacanthos L. (Sheats 12 wood mi ix. ss ne bicolor Turcz. (Sheats 174, Sheats 210; cek 8) Clearcut; roadside/pine- ee cuneata (Dum.-Cours,) G. ae (Miller 22; Sheats 161) Roadside; xeric site, near ero- sion monitoring plot Lespedeza procumbens Michx. (Miller 31) Xeric site, near erosion monitoring plot Lespedeza virginica (L.) Britt. (Miler 30) Xeric site, near erosion monitoring plot. Pueraria montana (Lour.) Merr. (Sheats 164) Up- land hardwoods. Robinia pseudoacacia L.(Sheats 192) Bottomland ardwoo S. Strophostyles umbellata (Muhl. ex Willd.) Britt. (Miller 28; Sheats 187) Xeric site, near erosion monitoring plot; bottomland hardwoods/ roadside. Stylosanthes biflora (L.) B.S.P.(Sheats 180) Roadside. Tephrosia virginiana (L.) Pers.(Summitt 172) Road- side BRIT.ORG/SIDA 21(2) Sims) DC. (Summitt 89) Roadside. — Wisteria sinensis FAGACEAE 2/9 Fagus grandifolia Ehrh.(Sheats 78) Upland slopes. Quercus alba L.(Sheats 76) Upland hardwood mix. Quercus falcata Michx. (Sheats 134) Pine forest. Quercus marilandica Muenchh.(Sheats 181) Pine- = hardwood mix. Quercus montana Willd. (Sheats 87) Dry oak- hickory mix, Quercus phellos L.(Sheats 128) Pine forest. Quercus rubra L. (Sheats 79) Upland hardwood Mix. Q lata Wangenh.(Sheats 135) Pine forest. Quercus velutina Lam. (Sheats 2 GENTIANACEAE 1/1 Sabatia angularis (L.) Pursh (Sheats 190; Wiecek 1) Bottomland hardwoods; loblolly pine plantation. HAMAMELIDACEAE 1/1 Hamamelis virginiana L.(Wiecek 17) Bottomland hardwood forest JUGLANDACEAE 2/3 Carya alba (L.) Nutt.ex Ell. (Sheats 97, Sheats 149, Sheats 225) Upland lak bottomland hard- d forest; pine fore ee g ee (P.Mill.) ses caches 2) Road- side near lake. Juglans nigra L.(Sheats 122, Sheats 151) Pine for- est; pine-hardwood m LAMIACEAE 4/7 Prunella vulgaris L. (Miller 24; Summitt 24) Xeric site, near erosion monitoring plot; roadside. Pycnanthemum muticum (Michx.) Pers. (Sheats 185) Bottomland hardwoods Pycnanthemum pycnanthemoides (Leaven- worth) Fern. (Wiecek 70) Bottomland hard- wodd forest Pycnanthemum tenuifolium Schrad. (Sheats 184) Pine plantation/roadside Salvia ae . (Summitt 209) Rgadsida: S Hlaria tica Muhl. ex Spreng. (Sheats 170) = pland hawood ea aria integrifolia : (Wiecek 69) Pine-hard- wood mix LAURACEAE 1/1 Sassafras albidum (Nutt.) Nees (Sheats 112, Sheats 146) Bottomland hardwoods; upland hardwoods RA OF CAMP BUTNER, NORTH CAROLINA 1137 LINACEAE 1/1 Linum sulcatum Riddell var. sulcatum (Batson 1013, DUKE) AGNOLIACEAE 1/1 Liriodendron tulipifera L. (Sheats 83) Upland oak- hickory mix. MELASTOMATACEAE 1/1 Rhexia mariana L.(Sheats 171;Summitt 171) Pine mix; roadside MORACEAE 2/2 Maclura pomifera (Raf.) Schneid. (Sheats 102) Pine-hardwood mix. Morus rubra L. aoe 124) Pine-hardwood mix. NYSSACEAE 1/1 bes es ca Marsh. (Sheats 226; Summitt 279) Pine hardwood mix; near stream. OLEACEAE 1/1 Fraxinus americana L. (Batson 114 ONAGRACEAE 2/3 Ludwigia decurrens Walt. (Sheats 224) Oenothera biennis L. (Summitt 132) Ditch. Oenothera fruticosa L. (Sheats 93) Roadsides. OROBANCHACEAE 3/3 Agalinis purpurea (L.) Pennell (Wiecek 46) Upland hardwoods Aureolaria virginica (L.) Pennell (Sheats 191) Bot- omland hardwoods. Epifagus virginiana (L.) W. Bart. (Wiecek 56) Slope near creek 1, DUKE) OXALIDACEAE 1/3 Oxalis corniculata L. (Wi Upland hardwoods. Oxalis stricta L.(Sheats 205. Summitt 170) Roadside. Oxalis violacea L. (Summitt 191; Wiecek 36) Knap of Reeds Creek; near stream PAULOWNIACEAE 1/1 Paulownia tomentosa (Thunb.) Sieb. & Zucc. ex Steud. (Sheats 150) Pine forest. Wiecek 54) PHRYMACEAE 1/1 Mimulus alatus Ait. (Sheats 200) Bottomland ardwoods PHYTOLACCACEAE 1/1 Phytolacca americana L.(Sheats 139) Pine forest. PLANTAGINACEAE 2/4 Nuttallanthus canadensis (L.) D.A. Sutton (Summitt 195) In clearcut. Pi aristata Michx.(Sheats 152) Pine forest. Plantago rugelii Dcne. (Sheats 206) Roadside. P. lantago virginica [is (Summi tt 208) Roadside. PLATANACEAE 1/1 ai yeas Occi identalis L.(Sheats 90) Alluvial forest (streamside). POLYGALACEAE 1/1 Polygala curtissii Gray saan 169; Summitt 9; Wiecek 9) Clearcut; roadsi PORTULACACEAE 1/1 Claytonia virginica L.(Summitt 193) Along creek. RANUNCULACEAE 4/4 epatica nobilis Schreb. var. obtusa (Pursh) Steyermark (Sheats 114; Summitt 188) Upland hardwoods; along streambank. eo. abortivus L.(Summitt 182) Grass/pine lr. e ee thalictroides a. .) Eames & Boivin (Summitt 189) Streamban Xanthorhiza simplicissima Veal (Sheats 202) Creekside. ROSACEAE 10/12 Amelanchier arborea (Michx. f.) Fern. (Summitt 114, Summitt 224) Forest edge Crataegus flava Ait. (Sheats 133) Pine forest. Fragaria virginiana Duchesne (Batson 744, DUKE) ae oe (Ait.) Michx. (Sheats 208) Pine- mix. ee trifoliatus (L.) Britt. (Sheats 74) Pine- hardwood oe ses L. (Sheats 84, Summitt 85) Roadside. one simplex Michx. (Batson 738, DUKE) Prunus serotina Ehrh. (Sheats 162) Roadside. Pyrus communis L. (Sheats 154) Pine forest. Rosa carolina L. (Batson 878, DUKE Rubus argutus Link (Summitt 137) Roadside. Rubus trivialis Michx. (Sheats 82) RUBIACEAE 4/6 Diodia teres Walt. (Miller 29; Sheats 204) Roadside. Xeric site, near erosion monitoring plot Galium circaezans Michx. (Wiecek 11, Wiecek 41) Bottomland hardwood forest; Upland hard- woods. Galium pilosum Ait. (Sheats 130) Bottomland hardwoods. Galium triflorum Michx.(Wiecek 42) Upland hard- woods 1138 Houstonia caerulea L.(Summitt 180, Summitt 187) Roadside Mitchella repens L. ( SALICACEAE 1/1 Salix nigra ane (Sheats 141) Bottomland hard- woods. SAPINDACEAE 1/1 Acer rubrum L. (Sheats 72) Pine-hardwood mix. SAXIFRAGACEAE 1/1 Heuchera americana L. (Summitt 159) Near streambank Wiecek 64) Pine plantation. SI MARQUEACENE 1/1 P.Mill.) Swingle (Sheats 160) ere Roadside. SOLANACEAE 1/1 Solanum carolinense L. (Sheats 116) Pine forest. SYMPLOCACEAE 1/1 Symplocos tinctoria (L.) U'Hér. (Sheats 196) Pine- hardwood mix/roadside. ULMACEAE 1/1 Ulmus alata Michx. (Batson 797, DUKE) VIOLACE AE 1/2 mulifolia L. (pro sp.) (lanceolata x ea (Summitt 143) Roadside. Viola tricolor L.(Summitt 184) Roadside. VITACEAE 2/3 Parthenocissus quinquefolia (L.) Planch. (Batson 43, DUKE) Vitis aestivalis Michx. (Batson 1044, DUKE Vitis rotundifolia Michx. (Wiecek 65) Pine-hard wood mix — MONOCOTS ALLIACEAE 1/2 Allium ampeloprasum L.(Sheats 159) Allium vineale L.(Summitt 158) Near streambank. AMARYLIDACEAE 1/1 Narcissus sp. (Summitt 75) Roadside. COMMELINACEAE 1/1 Commelina communis L. (Wiecek 34) Knap of eeds Cree CYPERACEAE 3/3 Carex crinita Lam. (Wiecek 18) Bottomland hard- ood forest Cyperus retrorsus Chapman (Sheats 219) BRIT.ORG/SIDA 21(2) Scirpus cyperinus (L.) Kunth (Wiecek 55) Upland hardwoods. IRIDACEAE 2/2 Iris verna L.(Sheats 156, Sheats 182) Pine planta- tion/roadside. Sisyrinchium angustifolium P.Mill. (Summitt 205) Roadside. LILIACEAE 1/1 Erythronium americanum Ker-Gawl. (Summitt 190) Lake edge. MELANTHIACEAE 1/1 Trillium catesbaei Ell. (Sheats 85) ORCHIDACEAE 2/3 Goodyera pubescens (Willd.) R. Br. ex Ait. f. (Summitt 192) Along creek. Spiranthes cernua (L.) L.C. Rich. (Wiecek 59) Pine plantation. Spiranthes praecox ( Slope near creek POACEAE 13/20 ste de ternarius Michx. (Wiecek 70) Pine- ardwood mix chosnantn lanolin (Michx.) Yates (Wiecek 12 mland hardwood forest. Rane ai laxum (L.) Yates (Wiecek 49) Up- land hardwoods. Chasmanthium sessiliflorum (Poir.) Yates (Wiecek 53) Upl hardwoods Danthonia sericea Nutt. (Sheats 178) Pine forest/ roadside Walt.) S. Wats. (Wiecek 57) Danthonia spicata (L.) Beauv, ex Roemer & J.A. Schultes (Batson 338, DUKE) Dichanthelium laxiflorum (Lam.) Gould (Sheats “Batson 475, DUKE) Dichanthelium sphaerocarpon Sea ) Gould (Wiecek 2) Loblolly pine plant Echinochloa crus-galli (L.) Beauv ae 227) Elymus hystrix L.(Sheats 120) Pine forest. Holcus lanatus L. (Sheats 153) a um vimineum (Trin.) A.Camus (Sheats 100; cek 67) Pine-hardwood mix; Pine- oo Mix. Panicum anceps Michx. (Sheats 217; Wiecek 51) Upland hardwoods; roadside. Panicum dichotomiflorum Michx. (Wiecek 67) e plantation. eee floridanum Michx. (Miller 33) Xeric site, near erosion monitoring plot. CAMP BUTNER, NORTH CAROLINA 1139 Paspalum notatum Flueggé (Sheats 2716) Road- — Schizachyrium scoparium (Michx.) Nash (Wiecek side. 45; Batson 406, DUKE) Upland hardwoods. Pennisetum glaucum (L.) R. Br. (Wiecek 66) Pine- SMILACACEAE 1/1 hardwood mix. Saccharum brevibarbe (Michx.) Pers. var. contor- tum (EIL) R.Webster (Wiecek 50) Upland hard- ds. Smilax rotundifolia L. (Sheats 99) Pine-hardwood MIX. ACKNOWLEDGMENTS We thank C. Wiecek, V. Miller, C. Sheats, and K. Summitt for their collecting efforts and, in part, identification assistance; H. Davis and M. Kruger for mount- ing and organizing the specimens; Sherri Herndon for pulling respective speci- mens at DUKE; and Misty Franklin (NC Natural Heritage Program) for supply- ing information on rare species. REFERENCES ANGIOsPERM PHYLOGENY Group. 2003. An update of the Angiosperm Phylogeny Group classi- fication for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399-436 Batson, W., Jk. 1952. Floristics of the Iredell soil series in the central Piedmont of North Carolina. Ph.D. dissertation, Duke University, Durham Hatt, S.1995. Inventory of the wildlife habitats, movement corridors, and rare animal popu- lation of Durham County, North Carolina. North Carolina Natural Heritage Program, Raleigh. Jones, S..1971. Floristic survey and vegetational analysis of Yates Pond Biological Area in Wake County, North Carolina. M.S. Thesis, North Carolina State University, Raleigh Parmer, M.W.1990. Vascular flora of the Duke Forest, North Carolina.Castanea 55:229-244. Sawyer, G.P, Jr. 1968. The vascular flora of William B. Umstead State Park, Wake County, North Carolina. M.A. Thesis, University of North Carolina-Chapel Hill. SkeAN, J.D., Jr. 1982. The vascular flora and plant community types of Durant Nature Park, Wake County, North Carolina. North Carolina State University, Raleigh.M.S. Thesis, North Carolina State University, Raleigh Swas, E.C. 1990. The flora and vegetation of White Pines Natural Area, Chatham County, North Carolina. North Carolina State University, Raleigh.M.S. Thesis, North Carolina State University, Raleigh Tazik, D.J., S.D. Warren, V.E. Diersina, R.B. SHAW, R.J. Brozka, C.F. Bactey, and W.R. WHitwort. 1992. U.S.Army land condition-trend analysis (LCTA) plot inventory field methods. USACERL Tech. Rep. N-92/93. U.S Army Corps of Engineers. USDA, NRCS. 2002. The PLANTS Database, Version 3.5 (plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. Woootot Forestry RESEARCH AND DEVELOPMENT Procram. 2003. Camp Butner Training Site Land Condition Trend Analysis (LCTA) Program: Progress Report for the 2002 Field Season. NC State University, Raleigh A VASCULAR FLORA SURVEY OF EMERGENT CREEK BED MICROHABITATS OF KISATCHIE BAYOU TRIBUTARIES IN NATCHITOCHES PARISH, LOUISIANA Ray Neyland and Melissa Hennigan Lowell E. Urbatsch Department of Biological and Department of Plant Biology Environmental Sciences Louisiana State University McNeese State University Baton Rouge, Louisiana, 70803, Lake Charles, Louisiana 70609, U.S.A. reyland@mail.mcneese.edu ABSTRACT Tributaries flowing into Kisatchie Baye in southwestern Natchitoches Parish, pomsuna are see acterized by clear flov er and white sand bottoms. In places d emerge during nie Oa g levele but quickly become peri during periods of rain. These emergent creek Be eee a uence microhabitat that hosts a distinctive flora. A total of 70 spe- cies representing 30 152 genera were discovered in this base-line vascular plant survey of these emergent ae oe With few exceptions, the plant species here appear to be well-adapted for life in this challenging microhabitat. ce Key Worps: Kisatchie Bayou, plant microhabitat, vascular flora survey RESUMEN Los afluentes que di por Kisatchie Bayou en el sudoeste de Natchitoches Parish, Luisiana se caracterizan por el agua clara que flu y oe fondos nba blanca. En eee eestes sae bancos d : ] ‘ : oOo los periodos de lluvia. Estos bancos imespeuadios proporcionan un microhabitat unico que alberga una flora deans oe fore de #0 especies ae representan a Sd familias y 52 generos se descubrieron tas especies parecen estar bien adaptadas para la vida en este iaieenabieat dectane Tributaries flowing into Kisatchie Bayou in southwestern Natchitoches Parish, Louisiana are characterized by clear {lowing water and white sand bottoms. In places, sandstone or siltstone creek beds emerge during normal water levels. Their surfaces are devoid of sand and soil except in small crevices and fissures. The emergent creek beds remain perpetually damp because water permeates these fissures. hysiographic expression of the sector where Kisatchie Bayou and its tributaries reside reflects the lithologies of the Miocene Epoch (Andersen 1993). Andersen (1993) designates the physiography of the creeks draining the area as recent alluvium (undifferentiated) of the Holocene. Martin et al. (1990) designated the soil along the stream banks, which ulti- mately washes into these tributaries as “Kisatchie-Oula.” This soil type is broadly SIDA 21(2): 1141-1147. 2004 1142 BRIT.ORG/SIDA 21(2) defined as a very strongly acidic fine sandy loam occurring on 5-40% slopes. Additionally, this soil is low in fertility and runoff is rapid (Martin Jr. et al. 1990). Some emergent creek beds are inhabited by a small number of vascular plants, bryophytes (mosses and liverworts) and lichens. Plants are rooted in the crevices and fissures. However, lichens are directly attached to the rock sub- strate. Only those plants that manage to remain rooted during periods of inun- dation survive in this unique and challenging microhabitat. The purpose of this study is to survey the vascular flora of the emergent creek beds of Kisatchie Bayou tributaries. This survey provides a base-line in- ventory of this distinctive plant community that can be used for comparison to monitor changes that may occur due to natural or human perturbance. METHODS Several tributaries to Kisatchie Bayou were located by examining Natchitoches Parish aerial soil survey maps (Martin Jr. et al. 1990). Study sites were discov- ered by exploring these tributaries on foot. Each study site was surveyed periodically throughout the entire year of 2002. At least one voucher specimen was collected for each species; voucher numbers are indicated in Table 1. All vouchers are housed at McNeese State University Herbarium (MCN). Nomenclature follows Kartesz (1999) with the exception of Aletris lutea (Narthecicaceae), where nomenclature follows An- giosperm Phylogeny Group (1998). RESULTS Vascular plants are not present in all emergent creek beds. However, they typi- cally occur in areas where the creeks are broadened and the water levels are comparatively low. Additionally, these areas are often associated with small watertalls and rippled currents. The forest canopy is more open in these broad- ened areas with consequent increased light levels. A detailed description of each study site follows. The location of each site is indicated in Figure 1. Site L occurs in Little Bayou Pierre. This area is characterized by sandstone islets and peninsulas that lie just above the normal water level. Small water- falls are present. The site is about 85 m long and about 18m at its widest point. The site is just south of the bridge along Hwy. 118 near Mink, 31° 23' 38" north and 93° 03' 52" west. Site 2 occurs in Little Sandy. This area is characterized by islets and a few small peninsulas. The substrate here appears to be siltstone which is softer and darker than the sandstone found in the three other study sites. There are no watertalls here; however, the creek bed topology produces turbulence and rip- pling. The area is about 73 m long and about 19m at its widest point. The site is about | km north of the bridge on Hwy. 118 and about 2.5 km east of the town of Kisatchie, 31° 24' 37" north and 93° 09' 7" west. NEYLAND ET AL., FLORA OF KISATCHIE BAYOU TRIBUTARIES IN LOUISIANA 1143 31° 27' 117 Kisatchie T5N ee Kile \ t t 3 , I ° Lite fl es 31 21' a 93° 14! R8W R7W nam +—{ 1 mile O Study Site K+ 1km @ Town ARKANSAS ( 4 MISSISSIPPI Natchitoches e TEXAS BATON ROUGE e GULF OF MEXICO Fic. 1. Detailed map of th gent creek bed study sites is depicted in the upper illustration; Louisiana State High ways 117 and 118 are indicated. TI | ight ill ionst the stud inrelati Natchitoches Parish. The u 1144 BRIT.ORG/SIDA 21(2) Site 3 occurs in Rocky Branch. This area is characterized by a mixture of islets and peninsulas. A small waterfall is present. The area is about 38 m long and about 14 m at its widest point. The site is about 100 m south of the bridge on Hwy. 118 and about 3.6 km east of the town of Kisatchie, 31° 23' 50" north and 93° 05' 40" west. Site 4 occurs in Little Kisatchie Bayou. The area is characterized by a mix- ture of peninsulas and islets. Rippled water flow occurs here but no waterfall is present. The area is about 49m long and about 9m at its widest point. The site is about 100 meters south of the bridge on Hwy. 118 about 3km east of the town of Kisatchie, 31° 23' 49" north and 93° 06' 29" west. A total of 70 vascular plant species representing 30 families and 52 genera were discovered in this survey (Table 1). Of these, 32 species were found in two or more study sites. With a total of 57 species discovered, study site 1 was the most diverse. In study sites 2, 3 and 4, a total of 23, 20 and 21 species were dis- covered, respectively (Table 1). DISCUSSION Emergent creek beds within the Kisatchie Bayou tributaries provide a distinc- tive plant microhabitat. Observations made during this study suggest that this flora is stable. Periods of submergence had little impact on the overall health of the flora. For example, the effects of heavy rainfall from the remnants of hurri- canes Lili and Kenna in October 2002 were minimal. Although larger plants were lodged by the strong force of rapidly moving water, they appeared to suf- fer no irreparable damage and recovered rapidly. Although most species appeared to be thriving on the emergent creek beds, there were two notable exceptions. Of the several individuals of Pinus taeda that were discovered, all were seedlings or juveniles. It appears that the small fissures in which they were rooted had insufficient soil or space for plants to reach maturity. The one individual of Baccharis halimifolia appeared to have been repeatedly damaged by high water. Although individuals of these two species had managed to germinate and survive fora time, they do not appear to be adapted well for survival on the emergent creek beds. Additionally, observations made during this study suggest that this flora is fertile. Specifically, all discovered species, with the exception of Pinus taeda and Baccharis halimifolia, produced spores or seeds during 2002. ACKNOWLEDGMENTS We thank Mark Paulissen for his review of an earlier version of this paper and the Kisatchie District of the U.S. Forestry Service. NEYLAND ET AL., FLORA OF KISATCHIE BAYOU TRIBUTARIES IN LOUISIANA 1145 Taste 1. Species found listed by family and division with voucher numbers indicated. Presence of a species at each study site is indicated by an"X.” Specimens curated at McMeese State University (MCN). Taxon Study Site Voucher DIVISION LYCOPODIOPHYTA ae cane = a (Chapm.) Cranfill X 1964 Ly i DIVISION FILICOPHYTA Dryopteridaceae Onoclea sensibilis L. X xX 1987 Fygoclacene dium japonicum (Thunb.) Sw x 1996 Osmundacea Osmunda er L. var. spectabilis (Willd.) A. Gray X x 1997 DIVISION CONIFEROPHYTA Pinaceae Pinus taeda L. xX Xx 1968 DIVISION MAGNOLIOPHY TA Crass Macnotiopsipa (Dicots) Apiaceae peda aod ium Walt. Xx xX Xx 2035 aceum (Michx.) Raf. x X 1993 Re Baccharis halimifolia L. X 2060 Coreopsis linifolia Nutt. x 2053 Coreopsis tripteris L xX xX 2038 Elephantopus carolinianus Raeusch X X xX 1988 Helianthus hirsutus Raf. X x 2055 Mie ae graminifolia (Michx.) Nutt. X 2057 P ata (L. X X X 2063 Solidago fgota P.Mill. X X X xX 2041 Symphyotrichum lateriflorum (L.) A.& D. Love xX X x 2065 Betulaceae Alnus serrulata (Ait.) Willd. Xx X X X 1787 Buddlejaceae SEH, oot L. X Campanulac Lobelia ae Michx. var. pauciflora Bush x Xx 2054 Beers itilum L. Xx X X X 1973 pec a lum (Spach.) Steud. X X X 2019 Droser. ee ee folia Pursh x 1998 1146 Tase 1.continued BRIT.ORG/SIDA 21(2) Taxon Study Site Voucher 1 2 3 Ericaceae Rhododendron canes (Michx.) Sweet x x 1999 Vaccinium elliottli Chapm. Xx x 1994 Fabaceae Desmodium lineatum DC. x 2057 Desmodium paniculatum (L.) DC. x Xx 2058 Lespede ad virginica (L.) Britt. X 2045 Lamiaceae Lyc irgit nicus Lb xX X xX X 2039 sein ae L. x X 2007 Lentibulariaceae eae ula pumila Michx. X 2000 ta Mic Xx 1956 Utriculari fa juncea Vahl X 1955 ho gaiila eae lifolia (J.F.Gmel.) G. Don x 2023 ivtht ae jue (DC.) Wood. x 2028 Melastomataceae Rhexia virginica L. X 1965 Myricaceae Myrica cerifera L. X X 2001 Narthicaceae Aletris lutea Small x 2002 sa ea Ludwigia alternifolia L. X X X 2016 Rubia ens repens L. X x 2003 Scrophulariaceae Mecardonia procumbens (P. Mill.) Small x 1990 Gratiola pilosa Michx. X xX 2005 Violaceae Viola _ primulifolia L. (pro. sp.) Xx xX X 2059 Ctass Litiopsipa (Monocots) urmanniaceae annia capitata (Walt.) Mart. x 958 Cyperaceae Carex amphibola Steud. X 1960 Carex tribuloides Wahlen x 2026 Cyperus ha X xX 2011 Eleocharis microcarpa Tor xX 2015 Fimbristylis autumnalis ie Roem. & Schult. x 2044 Fuirena simplex Vahl x 2043 Fuirena squarrosa Michx. X 2025 NEYLAND ET AL., FLORA OF KISATCHIE BAYOU TRIBUTARIES IN LOUISIANA Tas_e 1. continued 1147 Taxon Study Site Voucher 1 3 Rhynchosp iculata (Lam.) A. Gray x 1985 Rhynchospora glomerata (L.) Vah| x 2017 hynchospora inexpansa (Michx.) Vahl x 2018 Juncaceae Juncus coriaceus Mackenzie X xX 1961 Juncus nodatus Coville X 2027 Juncus scirpoides Lam. xX 1959 Juncus tenuis Willd. X 2020 Poaceae Chasmanthium laxum (L.) Yates x x 2030 Dichanthelium dichot (L.) Gould var. ensifolium (Baldw. ex Ell.) Gould & C.A. Clark X X X 1958 Dichanthelium sphaerocarpon (Ell.) Gould var. isophyllum (Scribn.) Gould & C. A. Clark x 4 1957 Dichanthelium scoparium (Lam.) Gould x 2010 Panicut iT virgatun rL. 2042 Paspalum setaceum Michx. xX 2032 Paspalum urvillei Steud. Xx 2009 Steinchisma hians (Ell.) Nash x 2008 Xyridaceae Xyris ambigua Bey. ex Kunth X 2031 Xyris baldwiniana Schult. xX 1962 Xyris difformis Chapm. var. curtisii (Malme) Kral x 1967 Xyris laxifolia Mart. var. iridifolia (Chapm.) Kral x 1963 Xyris torta Sm.in Rees X 2006 REFERENCES ANDERSEN, H.V. 1993. Geology of Natchitoches Parish. Geological Bulletin No. 44 Louisiana Geological Survey, Baton Rouge. ANaiosPeRM PHYLOGENY Grour. 1998. An ordinal classification for the families of flowering plants. Ann. Missouri Bot. Gard. 85:531-553. Kartesz,J.1 .1999.A synonomized checklist of the vascular flora of the United States, Canada and Greenland. 1st.ed.In:Kartesz, J.T.and C.A. Meacham, eds. Synthesis of North Ameri- can floraVersion 1.0. North Carolina Botanical Garden, Chapel Hill. Martin Jr., PG., C.L. Butter, E. Scott, J.-E. Lytes, M. MARIANO, J. RAGU s, P Mason, and L. SCHOELERMAN. 1990. Soil Survey of Natchitoches Parish, Louisiana. United States Department of Agri- culture, Soil Conservation Service. U.S. Government Printing Office, Washington, DC. SARRACENIA PURPUREA (SARRACENIACEAE) IN LOUISIANA Michael H. MacRoberts and Barbara R. MacRoberts Bog Research, 740 Columbia Shreveport, Louisiana 71104, U.S.A. and Herbarium, Museum of Life Sciences Louisiana State University in Shreveport, Shreveport, LA 71115, U.S.A. Sarracenia purpurea L. was once thought to extend from British Columbia along the Canadian-United States border to the Atlantic coast, down the coast to Geor- gia and Florida, and west along the Gulf Coastal Plain to Louisiana. However, Nacziet al. (1999) have recently separated the Gulf Coastal Plain population as a distinct species, S. rosea Naczi, FW. Case & R.B. Case, which does not reach Louisiana but occurs in southwest Georgia, north Florida, south Alabama, and southeast Mississippi. However, two specimens—one collected in 1842 by Will- iam Marbury Carpenter (s.n. U.S.) and the other collected in 1870 by Americus Featherman (s.n. LSU)—both of which have been identified as S. purpurea by Naczi et al.(1999) and both reported from southeast Louisiana west of the range of S. rosea, are anomalous; they leapfrog S. rosea and are disjunct from the near- est S. purpurea population in eastern Georgia (see MacRoberts and MacRoberts [1988] for a detailed review of S. purpurea in Louisiana). Naczi et al. 1999) sug- gest possible labeling error to account for these anomalous specimens. The label of the Featherman Sarracenia purpurea specimen, along with other Featherman collections from the same locality, is shown in Figure 1. MacRoberts and MacRoberts (1988) originally interpreted the script just below “Covington” on the Sarracenia specimen to read “AF” the collector’s initials. Naczi et al. (1999:1200) question this interpretation: “The script leaves determi- nation of the final pair of letters equivocal. Instead of ... Covington/AF [Americus Featherman] ... the label may actually read ‘.. Covington/AL. [Covington, Ala- bamal.” According to this assessment, the specimen is from Covington Co., Ala- bama, not Covington, St. Tammany Parish, Louisiana. On reexamination, this interpretation fails for two reasons. First, the script following “Covington” is neither “AF” nor “AL,” but “St T.” Of the Featherman specimens with “Coving- ton” on the label, some read “Covington” only, but others have something writ- ten below “Covington.” We (and Diane Ferguson, pers. comm.) examined these and found that what was written was either “St Tam.” or “St T.”, abbreviations for “St. Tammany.” Second, corroborative evidence that this specimen comes from Louisiana is to be found in Featherman’s (1871) published description of SIDA 21(2): 1149-1152. 2004 1150 BRIT.ORG/SIDA 21(2) ih Lyi MACE hw sabres Yate Leil. do , bd ay LoL 1 Lozern peeling fp GLEE hie, . ae hy A A COAL : i. Se a i ee f eg Hae Bigot MOP)? ODO Ll ve SP eras go * @ Fic. 1 1 L ee . £ Cc . | 1 ie al r L I . £ ° ocd 2 gt St Tammany Parish, Louisiana. 1151 its provenance. In 1869 the Louisiana State Legislature passed Act. No. 72 in which professors of the Louisiana State Seminary of Learning and the Military College (later Louisiana State University) were to spend four months each year in survey work in their respective fields. Americus Featherman traveled exten- sively throughout Louisiana and produced three reports on the botany (Brown 1944). In his (Featherman 1871:78) list of species collected during his botanical survey of southern and central Louisiana in 1870, Featherman gives the follow- ing entry: “Sarracenia rubra Walt, [this is what Featherman called the species] Red Flowered, Trumpet-Leaf, Covington, St. Tammany,” which, except for the abbreviation of St. Tammany, is exactly what is written on the specimen label. Reading Featherman’s (1870, 1871, 1872) accounts of his extensive botanical surveys of Louisiana leaves no doubt of his competence as a careful botanist and researcher (see brief account of his Louisiana botanical career in MacRoberts [1984] and of his Louisiana botanical publications in Ewan [1967)). The provenance of the Carpenter specimen is less certain. The original la- bel reads “Sarracenia purpurea, wet pine woods, St. Helen, La.” This was anno- tated by Charles Mohr who purchased Carpenter's collection some years after his death: “Sarracenia purpurea L., St Helena Parish, Carpenter 1842.” Because in 1842 St. Helena Parish encompassed not only the present parish but also the northwestern quarter of Tangipahoa Parish, it is impossible to know from which present-day parish the specimen may have come. Carpenter was a prominent Louisiana botanist during the first half of the nineteenth century who collabo- rated with the leading botanists and naturalists of his time, including Charles Lyell and John James Audubon (Cocks 1914). He collaborated with J.L. Riddell and Josiah Hale on a work entitled “Plants of Louisiana,” which, unfortunately, was never published and is now lost. However, J.L. Riddell in 1852 published an abridged version “Catalogus Florae Ludovicianae,” and in his introductory re- marks, Riddell writes: “The following systematic list, embodying the results of a great many years of observation, by Dr. Josiah Hale, by the late Professor W.M. Carpenter, and by the author, has been abridged from a manuscript work, con- tributed by the author, in 1851, to the Smithsonian Institution. The MS. work alluded to is entitled, ‘Plants of Louisiana’ It comprises the technical and the vulgar names of the flowering and filicoid species of plants, well ascertained as growing within the limits of the State of Louisiana, [nearly all of which are rep- resented by specimens in the author's herbarium|],—with special localities, times of flowering, and full descriptions of the new species.” Unfortunately, this manu- script no longer exists and we are simply left with the bare list, which contains about 1800 taxa and which remained the most extensive compilation for Loui- siana until 1982 (MacRoberts 1984). As MacRoberts (1984:13) says: “an exami- nation of Riddell’s list leaves one with a feeling of confidence in his reports.” Interestingly, S. purpurea is on the list but, of course, without provenance or col- lector identification. But it seems likely that this was the Carpenter specimen. 1152 BRIT.ORG/SIDA 21(2) While labeling error, as Naczi et al. (1999) suggest, is always possible, there is no evidence of such in these cases, and there is strong corroborative evidence that the Featherman specimen came from Louisiana. As to the Carpenter speci- men, there is corroborative evidence of its provenance, not strong but sugges- tive. Thus, there seems no reason to remove Sarracenia purpurea from the Loui- siana flora. ACKNOWLEDGMENTS Diane Ferguson, Collections Manager, LSU Herbarium was instrumental in in- terpreting the Featherman labels and in providing scanned copies for our use. Amanda Crnkovic aided with the figure. wo anonymous reviewers provided useful comments. REFERENCES Brown, C.A. 1944. Historical commentary on the distribution of vegetation in Louisiana and some recent observations. Proc. Louisiana Acad. Sci, 8:35-47, Cocks, R.S. 1914. William M. Carpenter, a pioneer scientist of Louisiana. Tulane Graduates Mag. 3:122-127. Ewan, J. 1967. Bibliography of the botany of Louisiana. SouthW. Louisiana J. 7:1-83. FEATHERMAN, A. 1870. Report on the botany of Louisiana. |In: Annual report of the board of supervisors of the Louisiana State Seminary of Learning and Military Academy for the year ending December 31, 1869 to the governor. New Orleans. Pp. 111-122. FEATHERMAN, A. 1871. Report on the botanical survey of southern and central Louisiana made during the year 1870. In: Annual report of the board of supervisors of the Loui- Ssiana State University. The Republican Press, New Orleans. Pp. 1-131. FEATHERMAN, A. 1872. Third annual report of botanical survey of southwest and northwest Louisiana made during the year 1871. In Annual report of Prof. D.F. Boyd, superinten- dent, Louisiana State University, for the year 1871, to the governor of the state of Loui- siana. Pp. 101-161. MacRoserts, D.T. 1984. The vascular plants of Louisiana. An annotated checklist and bibli- ography of the vascular plants reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci. 6:1-165. MacRoserts, M.H. and B.R. MacRoserts. 1988. A note on Sarracenia purpurea L. in Louisiana. Phytologia 65:191-194. Naczi, R.F.C., E.M. Soper, FW. Case, and R.B. Case. 1999. Sarracenia rosea (Sarraceniaceae), a new species of pitcher plant from the southeastern United States. Sida 18:1183-1206. Riovett, J.L. 1852. Catalogus florae Ludovicianae. New Orleans Med. Surg. J. 8:743-763. ELEOCHARIS MUTATA (CYPERACEAE) NEW TO THE FLORA OF NORTH AMERICA NORTH OF MEXICO David J.Rosen Stanley D. Jones S.M. Tracy Herbarium Herbarium, B | Research Center Department of Rangeland Ecology & Management PO. Box 6717 Texas A&M University Bryan, Texas 77805-6717, U.S.A. College Station, Texas 77843-2126, U.S.A. ABSTRACT Eleocharis mutata(L.) Roem. & Schult. is reported new to the flora of North America north of México based on recent collections from Brazoria Co., Texas, U.S.A. A key to separate E. mutata from other species of Eleocharis subg. Limnochloa in North America north of México as well as a technical de- scription and ecological notes are provided. RESUMEN Se cita Eleocharis mutata (L.) Roem. & Schult. Nueva para la flora de Norte América al norte de México en base a recientes colecciones de Brazoria Co., Texas, U.S.A. Se ofrece una clave para separar my tytn . As Til l : bg. Limnochloa en Norte América al norte de México asi : f como una descripcion técnica y notas ecologicas. Recent field work in Texas produced collections of a member of Eleocharis R. Br. subg. Limnochloa (P. Beauv. ex T. Lestib.) Torr. (= Eleocharis ser. Mutatae Svenson) with triquetrous culms. These specimens keyed to E. fistulosa Schult. (= F.acutangula[Roxb.] Schult.) in Manual of the Vascular Plants of Texas (Correll & Johnston 1970) and Aquatic and Wetland Plants of Southwestern United States (Correll & Correll 1975). Correll and Johnston (1970) report E. fistulosa from the Rio Grande plains of Texas without mentioning any specific county. Gonzalez-Elizondo et al. (2002) discussed the misapplication of E. fistulosa in North America, referring to Texas specimens as E. obtusetrigona (Lindl. & Nees) Steud. A review of pertinent manuals and treatments of subg. Limnochloa (Svenson 1929; Adams 1972; Hooper 1972; Kern 1974; Haines & Lye 1983; Koyama 1985; Wilson 1993; Gonzalez-Elizondo 1994; Browning et al. 1997; Gonzalez- Elizondoet al. 2002) and a critical examination of numerous herbarium speci- mens confirm our collections are of Eleocharis mutata (L.) Roem. & Schult. Eleocharis subg. Limnochloa comprises over 20 species of perennial, rhi- zomatous, aquatic herbs distributed in tropical, subtropical, and warm tem- perate areas worldwide, and recognized by indurate and prominently to ob- scurely longitudinally many-veined scales and cylindrical spikelets (Gonzalez-Elizondo & Peterson 1997; Gonzalez-Elizondo 2002). Despite the usually coarse and conspicuous habit of this group, new species have recently been described from Venezuela (Gonzalez-Elizondo & Reznicek 1996) and SIDA 21(2): 1153-1160. 2004 1154 BRIT.ORG/SIDA 21(2) México (Roalson 1999). Eleocharis mutata has not been previously reported for Texas (Hatch et al. 1990; Jones et al. 1997). In Flora of North America, Gonzalez- Elizondo (2002) reports six species belonging to subg. Limnochloa: E. cellulosa Torr., E. elongata Chapm., E. equisetoides (Elliott) Torr, E. interstincta (Vahl) Roem. & Schult. E. obtusetrigona (Lindl. & Nees) Steud., E. quadrangulata (Michx.) Roem. & Schult. and E. robbinsii Oakes. A dichotomous key modified from Svenson (1929) and Gonzalez-Elizondo (2002) and a technical descrip- tion of E. mutata follow. KEY TO ELEOCHARIS SUBG. LIMNOCHLOA IN NORTH AMERICA, NORTH OF MEXICO 1. Culms without septa 2. Culms triquetrous to terete, not distinctly quadrangular. 3. Culms relatively coarse, (2—)3-5(-8.5) mm thick at leaf sheath summit; spike- mm thick; achene 1.4-1.8 mm w z Culms triquetrous (rarely distally secre 3 angled);achene apex slightly constricted at the summit into a hard annular thickening (see Fig. 1d); pe- rianth bristles irregularly retrorsely spinulose Eleocharis mutata 4, Culms more or less terete or distally obscurely 3—5-angled, never trique- trous; achene apex daaualy: io maiseayy SOnSTIEIEG, but WIMIOUE hard annular thict — | retrorsely SpInulose. of ly pi D: ae apex gradually narrowed into a stout spongy region; perianth bristles usually smooth or sometimes finely retrorsely spinulose (see ig. le) Eleocharis cellulosa 5. Achene apex markedly constricted to a short neck; perianth bristles coarsely feed spinulos Eleocharis es Culms slender, 0.5-1.5 mm thick can leaf sheath summit; spikelets 3 mmo less thick; achene 0.5— a mm wide. 6 Achene 0.65-1.4 long x 0.5-0.8 wide; floral scales 3.5—4.5 mm long; tubers absent Eleocharis elongata 6. Achene 1.9-2.6 ae x 1-1.4mm wide; floral scales 5—7.8 mm long; tuber eae robbinsii 2. Culms a Feat (4-angled) Eleocharis quadrangulata . Culms septate. 7. Perianth bristles one than the achene, coarsely retrorsely spinulose; achenes — uously sculptured at 10-15; culm septa extending up the culm to im diately below spikelet Eleocharis interstincta 7. ent bristles much shorter than the achene, thin and soft, without teeth; achenes not conspicuously sculptured at 10-15 x,appearing nearly noe culm septa extending up the culm to well short of spikelet isetoid q Ww Eleocharis mutata (L.) ROCH: & — (Fig. 1 a-d). Scirpus mutatus L., Syst. Nat 10) 2:867.1759. Eleoc! Roem. & Schult., Sys. Veg. 2:155. 1817. Limnochloa mutata (L.) Nees, FI. Bras. 2(1):101. 1842. ae (LECTOTYPE, typified by Browning et al. [19 LINN photo!). Eleocharis scariosa Steud., Syn. Pl. Glumac. 80. 1855. TYPE: BRASIL: Martius Herb. Fl. Bras. 229 (ISOTYPES: GH], M, NY). 1155 ROSEN AND JONES, Ue ER LER (L.) Roem. & Schult.a. Habit. b. Det | end of culm. d. Detail of achene. Eleocharis cellulosa Torr. e. Detail of achene. a—d drawn from Rosen 2614 (MICH drawn from Rosen 2698 (SBSC) by Neva Mikulicz. Fic. 1.E/ ande ) 1156 BRIT.ORG/SIDA 21(2) Plants perennial, rhizomes long, 2-5 mm thick, scales to 8 mm, tubers absent; roots coarse fibrous, gray-brown to maroon. Culms triquetrous, usually con- spicuously so distally (rarely in the field Texas plants on dryer sites obscurely 3 angled), sometimes twisted, (31-)53.8-93(-116) cm x (2.2-)2.6-5.1(-8.5) mm, soft to hard, internally spongy, with incomplete transverse septa, longitudinally striate when dry, shinny and smooth when fresh, dark green. Leaf sheaths 2, apically notched, apex acute to acuminate, membranous, loose, friable, maroon- chestnut to cinnamon brown; blade reduced toa mucro to 5mm long. Spikelets cylindric, obtuse (acute), proximal 2-3 scales empty, the first amplexicaul and appearing asa continuation of the culm, (12-)23-44(-66) x 3-)3.8-5.4(-8) mm; floral scales appressed to weakly spreading upon drying, ovate to broadly ovate, apex broadly rounded, the distal 0.2-0.3 mm hyaline-erose, central area broadly keeled from base for ca. 1/3-1/2 scale length, (2.8-)3.2-4(-4.8) x (1.9-)2.5-3.4(- 4.8) mm, finely many veined, mid-vein evident only in adaxial view, cartilagi- nous, stramineous, abaxially red-maculate or more frequently witha dark band near apex, adaxially red-maculate. Flowers with (5-)6-8 perianth bristles, ir- regularly oriented, narrow to somewhat broad and strap-shaped proximally, irregularly retrorsely spinulose nearly to the base, mostly exceeding the achene, stramineous, the margins and spinules sometimes dark reddish; stamens 3; an- thers 1.3-2.0 mm, reddish-brown; style trifid. Achene biconvex, more or less obpyriform, obovate, or sometimes broadly elliptic, the apex constricted to about 0.6 the width of the achene, broadening again into a hard annulus of the same texture and color as the achene, (1.2-)1.3-1.6(-1.9) mm (not including annulus or tubercle) x (-)LI-L4(-L8) mm, with ca. 20 longitudinal rows of deeply pit- ted horizontally rectangular cells visible through transparent periclinal layer oneach achene face, dull, cream colored, maturing to lustrous olive-yellow (am- ber); annulus oblong or tapering apically, (0.05-)0.09-0.18(-0.3) mm high; tu- bercle dorsoventrally compressed, triangular, well formed to withered, distinct or sometimes appearing to merge with the annulus or shouldered by it, (0.15-) 0.3-0.5(-0.9) mm x 0.4-0.8 mm, dark brown. Phenology and Ecology.—In Texas, flowering from early June through early November. Eleocharis mutata forms large monotypic colonies in dark gray, clay- loam soils of a shallow, semi-permanently flooded freshwater marsh near the coast, associated in the dryer fringes of the marsh with Cyperus elegans L., C. oxylepis Steud., C. polystachyos Rottb., Eleocharis cellulosa, E. olivacea Torr, E. quadrangulata, Juncus roemerianusG. Scheele, Paspalum floridanum Michx. P. vaginatum Sw., and Spartina patens (Ait.) Muhl. (Fig. 2). The nativity of Eleocharis mutata in Texas is uncertain; as we encountered it in Brazoria County, it was ecologically dominant, forming pure stands in a habitat usually more diverse. Distribution—In North America north of México, currently known from Brazoria County, Texas, on the eastern edge of the Gulf Prairies and Marshes. ROSEN AND JONES 1157 Fic 2. F } hotlabac K INN? a+ UncL: AA J L D -. Ales LULA, Daf, bp ae al * ith Eleochari: (L.) Roem. & Schult. in foreground Expected also in similar habitat from Jefferson County, SW to Goliad County, $ through the Rio Grande Valley. There is an immature specimen at TAES with triquetrous culms (Glazener s.n., collected 16 miles W of Goliad in 1941) anno- tated as Eleocharis acutangula. Unlike E. acutangula, however, this specimen has broadly ovate, finely many veined floral scales which fits our concept of E. mutata. It is possible that E. mutata is more widespread in southern Texas, and has been overlooked because of its affinity for aquatic habitats and its superfi- cial similarity to E. cellulosa and E. quadrangulata with which it occurs. Coastal habitats in United States, México, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Cuba, Jamaica, Haiti, Dominican Republic, throughout the West Indies, tropical South America, and tropical Africa. Vernacular name.—scallion grass (Adams 1972). 1: NORTH AMERICA. U.S.A.. TEXAS. Brazoria Co.: Hoskin’s Mound within the Brazoria National Wildlife Refuge, 3.8 km SE of the intersection of FM 2004 and Co. Rd 277, locally abundant in shallow water along roadside ditch and marsh S of perimeter oil-field ser- vice road, N of the mound, 21 Oct 2002, Rosen & Jones 2382 (MO, TAES, TEX, VSC), 29 Aug 2003, Rosen 2606 (GH, SBSC, NY, US, WIS, Z), Rosen 2614 (CIIDIR, K, MICH). Goliad Co.: 16 mi. W of Goliad, in water, 23 Aug 1941, Glazener s.n., (TAES). BARBADOS. Christ Church: G Hill Swamp, domi- 1158 BRIT.ORG/SIDA 21(2) nant understory» peut CREME a eva ebougneut much of the swamp, 09 Nov 1996, Rogers 96-128 MICH). BELIZE p, Ll. Mar 1933, Lundell 1816 (MICH); Belize City, vacant lots, roadsides, guns sites about town, | m elev. swamp on S side of town, 09 Aug 1992, Worthington 21439 (MO). COSTA RICA. Limon: Limon airport, beach at mouth of Rio Banano, near sand dunes and shallow oon 35 m from shore, 07 Jul 1966, Denton 1139a (MICH). Guanacasta: Palo Verde National Park, swamp ca. 2 km from park headquarters at Catalina, along transect to Laguna Nicaragua, abundant, 07 Sep 1984, Crow & Rivera 5981 (MO). DOMINICAN RE- PUBLIC. Proy. Maria: Llanura de Nagua, Trinidad Sanchez, Las Gordas, 18.7 km, Llanuras plantadas de arroz, zona pantanosa, con muchas malezas despues de la ultima cosecha (en el camino de Las Gordas a Mata Bonita), 19°25N, 70°O0W, 05 Oct 1982, Mejia & Pimentel 23601 (MO). Prov. Peravia: very common in muddy sites at roadside, Galeon, Bani, 29 Oct 1976, Cicero 8276 (NY, TAES). HAITI shallow pond and swamp area between Terrier Rouge and Fort Libertr, northeastern alluvial plain, 26 Jun 1941, Bartlett 17480 (MICH, NY, US). HONDURAS. Atlantida: in boggy area near the seashore vicinity of Ceiba, 06 Jul 1938, Yuncker et al. 8243 (MICH, NY); Sibun River, 28 Nov 1934, Gentle 1429 (MICH, NY); Hector Creek, Sibun River, 28 Nov 1934, Gentle ats MICH, NY, US). Toledo: in wet area on river bank, Monkey River, 18 Oct 1941, Gentle 3708 (MICH, NY, US); Monte Redondo lake, vicinity of Yeguare river, El Eamorano, 23 Jan 1970, Molina pe (MO, NY); commonly found along the edge of brackish lagoons also thrives where it receives continuous sea spray, All Pines, 23 Jan 1970, Schipp 786 (MICH, NY, Z). JAMAICA. St. Thomas, just N of Grant Pen, off of 10 ad . 4, a little above sea level, at edge of and growing in open water at roadside, 21 Jul 1963, Crosby et al. 822 (MICH, NY, US): Port Antonio, Dec 1890, Hitchcock s.n. (MO). MEXICO. Jalisco: about 2 km N of Puerto Vallarta in cultivated areas west of the airport, near sea level, very common in wet Scape beside road,13 Nov 1963, Feddema 2533 (MICH); La Huerta, Rancho Cuixmala, Gargollo farm, on E side of Cerro de la Alborada, Lat. 19°24°N, pene 104°59'W, elev. below 50 m, highly ime remnants oF tropical semi- ae sl st now used for cattle, locally | with feet in water, n g 4,04 Nov 1991, Lott et al. 4116 Ux, MICH, NY WIs)3km al N de Puerto Vallarta, obi el camino al aeropuerto, terrenos planos, salobres, en suelo humedo, escaso, 16 Nov 1963, oo she (MICH). Pemauass pequenta zona inundada a 8 kms al SE de Sisal, 11 Aug 1978, Lot 2582 (MO). M t sedge in center of Chance’s pond, alt. 2700 ft, 14-18 Jun 1950, Howard i pies NICARAGUA. Zelaya: in wet sand, Fl Bluff near Bluefields, 14 Dec 1968, Seymour 642 (BRIT); dense stands covering many acres, brackish inlet, Corn Island, 06 Mar 1971, Svenson 4317 (BRIT). Managua: near the mouth of Rio El Carmen, 30 km NW of Masachapa, freshwater marsh | km from shore, 16 Nov 1976, Neill 1276 (MO). PANAMA. San Blas: Comarca de San a ys cai oi lata) elev. 0-30 m, in marsh by airport, 27 Jun ieee de ae ver Herrera 8105 (M Beach road, growing in water, 03 Aug 1967, Kir} ias69 (MO, NY). Bocas del Toro: out along road W Almirante, 17 Oct 1965, Blum 1415 (MO). PUERTO RICO: 1.3 mi Son Rte 687 from junction with Rte 686, elev. i m, ie end of eee Tortuguero, en soil in marsh, 31 Dec 1980, Solomon 5749 (MO); Yabucoa, su | J guayj O Feb 1886, Sintenis 4942 (GH, NY, Z-2 aoe ). SOUTH RICA. BRAZIL. Ceara: brackish Le Barra do Ceara, eee de ee 25 Sep 1935, iniaeree oa NY). COLOMBIA 1 km W of Turbo, probable lat. ca. 8° 5' N, long ca. 76° 43' W clevation sea level, common growing in mud at edge of brackish slough in sun, 31 Mar 1962, Feddema 2111 (MICH); Santa Marta, 100 ft, Sep 1898-1901, Smith 245 (MICH, NY). E DOR. Costa: Prov. Esmeraldas, km 102, Esmeraldas-La Tola, a lado de la carretera, en pantano en un ts grande, 27 Jul 1984, Dodson et al. 14564 (MO). PARA- GUAY: no date, Reugger s.n. (ZT). TOBAGO: Buccoo Bay, 20 Apr 1939, Elmore 510 (MICH). TRINIDAD: ca. 2-3 mi E of Trinidad along the Beetham Hwy. low lying sedge marshes, ey Aug 1970, Davidse 2546 (MO, NY). VENEZUELA. Sucre: Laguna La Bodega, inmediatamente al Este de Santa Fe, 17 Sep 1973, Steyermark et al. 108552 (MO). LE oa ROSEN AND JONES 1159 AFRICA. ANGOLS: Money: Ikula bot ees by R. Zambezi, plentiful in rock pools in 1-15 ft of water, tufted er sheaths pale brown; upper and stem green, trian- gular, glumes green, edged red- en turning «pale brown, 17 Jan 1938, Milne-Redhead 4195 (Kk). LIBERIA: Crew Town, Monrovia, in water, 27 Jun 1909, Massey 82 (NY). SENEGAL: Lyndiane, 30 Sep 1956, Jacques-Georges 12460 (MO); Ziguinchor, km 54-57 Cap Skiring-Bignona, vicinity of Nyassia, oist area in mangrove-area, ll Sep 1994, Laegaard et al. 16891 (K, US); Basse-Casamance, Fegroum, eS Nov 1990, Berghen 9301 (MO); 10 Apr 1946, Roberty 6209 (Z). SIERRA LEONE: emplodered me swamp, Rokupr, 03 Jul 1947, Jordan 42 (K). TANZANIA. Pemba: Makongwe Is, in shallow water on th edge of a pond, leaves and stems (hand drawn triangle symbol) and flower spikes sessile at the apex of the stem, 16 Dec 1930, Greenway 2730 (K). TOGO: Agwegan, entre route et lagune, depression sale, savane herbense inonde eau sale 9 gr Natt/litre, 26 Jun 1985, Schafer 8629 (MO, US). > ACKNOWLEDGMENTS We thank the curatorial staff at the herbaria listed. We thank Mark Spencer (LINN) for making excellent images of the type specimen available for study, Hajo Esser (M) for assisting with nomenclatural uncertainties of E. scariosa,and Mary Stiffler and Linda Oestry with the Missouri Botanical Garden Library for assistance locating certain literature items. We thank Richard Carter for pro- viding many helpful suggestions and critical review of this manuscript, and for alerting the first author to certain important literature items. Thanks to Socorro Gonzalez-Elizondo and Galen Smith for their critical review and many helpful comments. The first author is grateful to Brian Cain and Ron Brinkley (USFWS) for presenting specimens they discovered during their survey of Hoskins Mound, and to Mr. Tommy Thompson and Mr. Nathan Gutowsky with ChevronTexaco for facilitating access to the site which led to this discovery. Special thanks to Neva Mikulicz for preparing the illustration. REFERENCES Avams, C.D. 1972. Flowering plants of Jamaica. University of the West Indies, Mona. Brownine, J., K.D. GorDon-Gray, and C.J. Warp. 1997. Studies in Cyperaceae in Southern Af- rica 32: Eleocharis subgenus Limnochloa section Limnochloa.S. Afr. J. Bot.63:1 72-184. Corrett, D.S.and H.B. Corrett. 1975. Aquatic and wetland plants of southwestern United States. Vol. 1. Stanford University Press. Stanford. Corrett, D.S. and M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner. GonzALez -ELIZONDo, M.S. 1994. Eleocharis. In: G. Davidse, M.S. Sousa, and A.O. Chater, eds. Flora Mesoamericana, Vol. 6: Alismataceae a Cyperaceae. Universidad Nacional Autonoma de México, Ciudad Universitaria. Pp. 458-464. Gonzatez -Euizonvo, M.S. 2002. Eleocharis subg. Limnochloa. In: Flora of North America Edi- torial Committee, eds. Flora of North America North of Mexico, Vol. 23. Oxford Univ. Press, New York. Pp. 116-120. GONZALEZ-ELIZONDO, M.S., M. GonzALez-ELizondo, and S.G. Smith. 2002. Eleocharis obtusetrigona (Cyperaceae) new to North and Central America. Acta Bot. Mexicana 60:/-11] 1160 BRIT.ORG/SIDA 21(2) GONZALEZ -ELIZoNDO, M.S. and P.M. Peterson. 1997. A classification of and key to the supraspecific taxa in Eleocharis (Cyperaceae). Taxon 46:433-449. GOnzALez -ELizonpo, M.S.and A.A. Reznicek. 1996. New Eleocharis (Cyperaceae) from Venezu- ela. Novon 6:356-365. Haines, R.W. and KA. Lye. 1983. The sedges and rushes of East Africa. East African Natural History Society. Nairobi. Hatch, S.L., KN. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Bull. MP-1655. Hooper, 8.8. 1972. Eleocharis (Cyperaceae). In: J. Hutchinson and J.M. Dalziel, eds. Flora of West Tropical Africa, 2nd ed. Whitefriars Press, Ltd., London. Pp. 311-314. Jones, S.D., J.K. Wiprr, and P.M. Montcomery. 1997. Vascular plants of Texas:a comprehensive checklist including synonymy, bibliography, and index. University of Texas Press, Aus- Kern, J.H. 1974. Cyperaceae.In:C.G.G.J.van Steenis, ed. Flora Malesiana, Ser. 1, 7(3). Noordhoff, Leyden. Pp. 435-753. Koyama, T.1985.Cyperaceae.|In: M.D. Dassanayake and FR. Fosberg, eds. Flora of Ceylon, Vol.5. Amerind Publishing Co. Pvt. Ltd., New Delhi. Pp. 125-405 (Eleocharis Pp. 255-268). Roatson, E.H. 1999. Eleocharis yecorensis (Cyperaceae), a new species of spike-sedge from Mexico. Aliso 18:57-60. Svenson, H.K. 1929. Monographic studies in the genus Eleocharis. Rhodora 31:121-135; 152-163 (plate 188). Wivson, K.L. 1993. Cyperaceae. In: G.J. Harden, ed. Flora of New South Wales, Vol. 4. New South Wales University Press, Kensington. Pp. 293-396, COMMENTS ON THE REDISCOVERY AND DISTRIBUTION OF CUNILA ORIGANOIDES (LAMIACEAE) IN TEXAS Jason R. Singhurst Walter C. Holmes Wildlife Diversity Program Department of Biology Texas Parks and Wildlife Department Baylor University th IH-35, Suite 100 Waco, texas 76798-7388 U.S.A. Austin, Texas 78704 U.S.A. ABSTRACT Cunila origanoides has been rediscovered in Texas based on recent herbarium and field studies. RESUMEN Cunila origanoides ha sido redescubierta en Te li Li i de herbario y campo. Herbarium and field studies on the Texas flora have yielded the following records pertaining to the occurrence and distribution of Cunila origanoides within the state. Cunila origanoides (L.) Britt. (Lamiaceae)—Small (1933), Cory and Parks (1937), Correll and Johnston (1970), Hatch et al. (1990), and Jones et al. (1997) attributed this species to Texas. Correll and Johnston (1970) cited the distribu- tion as north central Texas, while Hatch et al. (1990) gave the distribution as the Post Oak Savannah, Blackland Prairies, and Cross Timbers and Prairies veg- etational areas. The latter mentioned distribution was cited by Diggs et al. (1999) who included Cunila in the Flora of North Central Texas. Gould (1962, 1969) and Turner et al. (2003) did not include the species in their respective works. The exclusion from the latter work seems to have effectively removed Cunila from being known asa part of the state’s flora. Most likely, the ambiguity in the literature is a consequence of no cited voucher specimen. Recently, we have uncovered the two specimens cited below, thus are able to resolve the matter. Specimens examined: Kaufman Co.: vicinity of Terrell, 6 Sep 1904, Tyler s.n. (BAYLU (photo), US). Lamar Co.: 1.4 mi E of jet of St. Rt. 906 and Hwy 271 pees) on St. Rt. 906, slope forest below Pat Mayse Lake Dam, 18 Oct 2002, Singhurst G Harris cay (BAYLU). It may be assumed that Small (1933) had reference toa specimen, thus its inclu- sion in his manual, but under the genus Mappia House. The Kaufman County specimen was annotated by Donovan S. Correll in 1967, hence its inclusion in Correll and Johnston (1970). However, neither the Small (1933) nor Correll and Johnston (1970) reports give precise location data or indicate the source of their information. The other literature citations concerning the occurrence of the species in Texas seem to have originated from these two reports. Without loca- SIDA 21(2): 1161-1163. 2004 1162 BRIT.ORG/SIDA 21(2) tion data at least to county, it is understandable, and certainly correct, for Turner etal. (2003) to not include the species in the Atlas of the Vascular Plants of Texas. Cunilais a New World genus of about 15 species native from eastern North America to Uruguay (Diggs et al. 1999). Cunila origanoides isa native to eastern North America that is distributed from New York, south to Georgia, west to Kansas and Texas (USDA, NRCS 2004). The occurrence of C.origanoides in Texas is very significant. In eastern Texas, calcareous disjuncts, such as C. origanoides, have been receding in distribution and are evidence of calciphile relics of an earlier cooler climate (Kral 1963). The real significance of the occurrence of C. origanoides is in the disjunct nature of the sites. It is not known what the 1904 location of C. origanoides in Kaufman County was like and whether such habi- tat still exists there. There are still a few remnant hardwood sites, such as Cedar Creek Island (the big island) in Cedar Creek Lake, in Kaufman County that contain old growth oak stands with eastern taxa like Carollorhiza wisteriana, Erythronium albidum, Phryma leptostachya and Verbesina helianthoides. In Lamar County, C. origanoides occurs in rich hardwoods on slopes along a spring fed creek where the Woodbine and Eagleford shale formations merge. This is cretaceous geology, which is of rather restricted occurrence in northeast Texas. It does, however, support other disjunct and peripheral species such as the rare [in Texas] Prenanthes barbata and the more eastern Heuchera americana. There- fore, the presence of C. origanoides in northeast Texas is significant as this na- tive species is persisting in remnant mature hardwood habitat. This habitat type is dwindling from land use practices and the resulting fragmentation. Poten- tial habitat for C. origanoides also occurs in Fannin, Hunt, Delta, Hopkins, Fran- klin, Titus, Red River, and Bowie counties. This information is valuable as an indicator of calcareous slope hardwood forests that may contain other periph- eral and disjunct taxa and its discovery aids the understanding regional plant communities and their floristic make-up. — ACKNOWLEDGMENTS We wish to thank the curators of GH, NY, TEX, and US for their search for and use of specimens that made this study possible. During the study, the junior author visited US and is grateful for the assistance and kindnesses extended. We are grateful to Margie Knowles of US who arranged to have a digital photo of the Kaufman Co. specimen sent to the authors. We are very grateful to Barney Lipscomb for his comments and suggestions in producing this manuscript. REFERENCES Corrett, D.S. and M.C. Jounston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner Cory, V.L. and H.B. Parks. 1937. Catalogue of the flora of Texas. Texas Agricultural Experi- mental Station Bulletin 550, College Station. 1163 Diacs, G.M., B.L. Liescoms, and R.J. O’KenNon. 1999. Shinners & Mahler's illustrated flora of north central Texas. Sida, Bot. Misc. 16, Botanical Research Institute of Texas, Fort Worth. GouLo, FW. 1962. Texas plants—a checklist and ecological summary. Texas Agricultural Experimental Station publication MP-585, College Station. Gouto, FW. 1969. Texas plants—a checklist and ecological summary. Texas Agricultural Experimental Station publication MP-585/ revised, College Station. Hatcu, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agricultural Experimental Station publication MP-1655, College Station. Jones, S.D., J.K. Wire, and PM. Montcomery. 1997. Vascular plants of Texas.A comprehensive checklist including synonymy, bibliography, and index. University of Texas Press, Austin. Krat, R. 1966, Observations of the flora southeastern United States with special reference to northern Louisiana. Sida 2:395—408. Smatt, J.K. 1933. Manual of the southeastern flora. Published by the author. New York. Turner, B.L., H. NicHois, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Vol.1. Sida, Bot. Misc. 24. Botanical Research Institute of Texas, Fort Worth. USDA, NRCS. 2004. The PLANTS database (plants.usda.gov/cgi_bin/plant_search.cgi). National Plant Data Center, Baton Rouge, LA. GEOCARPON MINIMUM (CARYOPHYLLACEAE), NEW TO TEXAS Eric L.Keith Jason R. Singhurst Raven Environmental Services Texas Parks and Wildlife Dept. PO. Box 6482 4200 Smith School Road Huntsville, Texas 77342, U.S.A. Austin, Texas 77342, U.S.A. ek7275@cox-internet.com Stan Cook Temple-Inland Forest Products 700 N.Temple Dr. Diboll, Texas 7594, U.S.A. ABSTRACT Geocarpon minimum, a federally threatened species, is documented for the flora of Texas for the first time. Approximately 200 plants were found in an extensive saline prairie near the floodplain of the Neches River in Anderson County, Texas. Included is a general description of the habitat, known distribution of the species, and associated flora in Texas. RESUMEN ‘dala lictatedoealad gta Neeeaty yy ; lanura A la How a dee por primera vez. ae eee cee rapide 200 enese enuna N c A nderson, Texas. Se incluye una see general oT habitat i: distribucién conocida de la especie, y la flora asociada en Tex Geocarpon minimum MacKenzie (geocarpon, tiny Tim, earth fruit) is listed asa federally threatened species by the U.S. Fish and Wildlife Service and is cur- rently known from Arkansas, northwestern Louisiana, and southwestern Mis- souri (MacKenzie 1914: Moore 1958; McInnis et. al. 1993; USDA, NRCS 2004; USFWS 1993). In Missouri, Geocarpon minimum occurs in glades and other open, sparsely vegetated areas on shallow soils over outcrops, often in shal- low depressions (Morgan 1980; Palmer & Steyermark 1950; Steyermark, et. al 1959; Thurman 1989; USFWS 1993). In Arkansas it is typically found in sparsely vegetated areas on saline prairies (Moore 1958; Orzell & Bridges 1987; Pittman 1988; Rettig 1983; Shepherd 1987; USFWS 1993). Soils in both habitat types are high in magnesium or sodium (Rettig 1983). This species was brought to the attention of the former Texas Natural Heritage Program in 1993 by Peggy Horner of the Missouri Department of Conservation as potentially occurring in north- east Texas. Several surveys were then conducted on saline prairies in East Texas, SIDA21(2): 1165— 1169. 2004 1166 BRIT.ORG/SIDA 21(2) including the Davy Crockett National Forest, Temple-Inland Forest Products Corp. (Temple) property, and other private lands. In summer of 2003, we discovered a large saline prairie complex occurring on Temple property in Anderson County, Texas, just above the floodplain of the Neches River. This prairie was surveyed in the summer and fall of 2003 and found to contain several halophytes typically associated with Geocarpon mini- mum. In surveys conducted in early March 2004, approximately 200 plants of Geocarpon minimum were discovered at eight stations throughout the 75 acre saline barren complex. All of the plants occurred on the edge of sparsely veg- etated areas commonly referred to a “slick spots” (USFWS 1993). These slick spots vary in size from one to approximately thirty square meters. They have very high sodium content and are typically devoid of any vegetation, with the exception of Sibara virginica, Talinum parviflorum, Plantago pusilla,and Cleo- mella angustifolia. This vegetative edge around the slicks is referred to as a “cryp- togamic lip,” where a spongy, leathery crust of moss protonemata, lichens, liv- erwort thalli, Nostoc sp., and the associated micro-flora interact to cement the lip in place (Shepherd pers. comm.; USFWS 1993). Woody species associated with Geocarpon adjacent to these slicks includes, Celtis laevigata spp. laevigata, Crataegus spp.,Juniperus virginiana, Opuntia macrorhiza, Pinus taeda, Quercus similis, Sabal minor, Smilax bona-nox, Vaccinium arboreum, and Ulmus crassi- folia. Associated herbac ies include Anagallis minima, Aphanostephus skirrhobasis, Astranthium integrifolium, Chaetopappa asteroides, Cleomella angustifolia, Coreopsis tinctoria, Crassula aquatica, Draba brachycarpa, Evolvu- lus sericeus, Gratiola flava, Houstonia rosea, Houstonia micrantha, Houstonia pusilla, Hypericum drummondii, Isolepis carinata, Krigia occidentalis, Lepuro- petalon spathulatum, Ophioglossum crotalophoroides, Plantago pusilla, Portu- laca spp., Rumex hastatulus, Sagina decumbens, Schoenolirion wrightii, Sedum nuttallianum, Sibara virginica, Sporobolus vaginiflorus,and Talinum parviflorum. The surrounding topography also includes mima mounds with micro highs and lows. The Brimstone Silt Loam described in McInnis et. al. (1993) may be represented in this saline prairie. However, in Coffee (1975), this area is not dif- ferentiated from the surrounding bottomland soil type, and saline soils are not mentioned. The mapped soil series for the prairie area is classified as Nahatche- Wehadke Soil Series with typically loamy bottomland soils. However, in an older soil survey of Anderson County from 1890, the Anderson County saline areas are briefly described (Department of Agriculture, Insurance, Statistics, and History 1890). Collection data for Geocarpon minimum follow: UNITED STATES. Texas. Anderson Co.: 120 m NE of a roadside park, 0.1 mi E of the jet. of U.S. Hwy 84 and Neches River; saline barren complex, 8 Mar 2004, J. Singhurst 12921, E. Keith, S. Cook, & B. Shepherd (BAYLU, TEX) KEITH ET AL., GEOCARPON MINIMUM NEW TO TEXAS 1167 Hi To kat £ Pes reli ak. 4 I A £415 Fig. 2. t G J J L + ia SHCK Spots, 1168 BRIT.ORG/SIDA 21(2) TAXONOMIC DESCRIPTION Glabrous winter annual, stems simple or branched at the base, the branches few, erect or spreading ascending, mostly 3-4 cm high and less than 0.5 mm thick, often a bright reddish or pale purplish color. 1 imple, opposite, green or reddish in color, 3-4 mm long, narrowly oblong or ovate- oblong, the mar- gins entire, and the apex acute. Flowers usually axillary, regular, funnelform- campanulate; sepals 5, 3-4 mm long, reddish or reddish-green; petals absent; stamens 5; staminodes 5; ovary superior, lance-ovoid, somewhat trigonous, about the length of the sepals. Fruit isa capsule containing numerous, funicular seeds, 0.5 mm long (NatureServe 2004; Steyermark 1963). ACKNOWLEDGMENTS Appreciation is given to Temple-Inland Forest Products Corp., which currently owns the entire saline prairie complex and is conserving the habitat. We would also like to thank Bill Shepherd, retired biologist from the Arkansas Natural Heritage Commission, for accompanying us on the collecting trip and sharing his expertise on Geocarpon minimum and its habitats. We would also like to thank Guy Nesom of the Botanical Research Institute of Texas and Monique Reed, Herbarium Botanist at Texas AQM University, for critical review of the paper. REFERENCES Corree, D.R. 1975. Soil Survey of Anderson County, Texas. United States Department of Agriculture Soil Conservation Service In Cooperation with Texas Agricultural Experi- ment Station. DEPARTMENT OF AGRICULTURE, INSURANCE, STATISTICS, AND History. 1890. Papers Accompanying the Annual Report of the Geological Survey of Texas 1890. www.lib.utexas.edu/books/ dumble/publications/08/08b315a.html Mackenzie, K.K. 1914. A new genus from Missouri. Torreya 14:67-68. McInnis, N.C.,L.M. Smith, and A.B. Pittman 1993. Geocarpon minimum (Caryophyllaceae), new to Louisiana. Phytologia 75:159-162. Moore, D. 1958. New records for the Arkansas flora. IV. Proc. Arkansas Acad. Sci. 12:12. Morcan, S. 1980. Status report on Geocarpon minimum in Missouri. Missouri Department of Conservation, Jefferson City, Missouri. NatureServe. 2004. NatureServe Explorer: An online encyclopedia of life [web application]. Version 4.0. NatureServe, Arlington, Virginia. Available www.natureserve.org/explorer. (Accessed: August 22, 2004) Orzett, S.L.and E.L. Brioces. 1987. Further additions and noteworthy collections in the flora of Arkansas, with historical, ecological, and phytogeographical notes. Phytologia 64: 81- Pacmer, E.J. and J. Stevermark. 1950. Notes on Geocarpon minimum Mackenzie. Bull. Torrey Bot. Club 77:266-273. KEITH ET AL., GEOCARPON MINIMUM NEW TO TEXAS 1169 Pittman, A.B. 1988. Identification, survey and evaluation of potential habitats of Geocarpon minimum MacKenzie in Arkansas. Arkansas Natural Heritage Commission, Little Rock, Arkansas. Rettic, J.H. 1983. A new Arkansas station for Geocarpon minimum MacKenzie (Caryophyllaceae). Bull. Torrey Bot.Club 110:213. SHEPHERD, W. 1987. Monitoring of Geocarpon minimum at Warren Prairie Natural Area in the spring of 1987. Arkansas Natural Heritage Commission, Little Rock, Arkansas. SHEPHERD, W. 2004. Personal communication on the ecology of Geocarpon minimum. STEYERMARK, J., J.W.Voict, and R.H. MoHLENBRock. 1959. Present biological status of Geocarpon minimum MacKenzie. Bull. Torrey Bot. Club 86:228-235. STEYERMARK, J.A. 1963. Flora of Missouri. lowa State Press. Ames, lowa. THurMAN, C.M. 1989. A Missouri survey of six species of federal concern. Final report. Mis- souri Department of Conservation. USDA, NRCS. 2004. The PLANTS Database, Version 3.5 (plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. U.S. Fish AnD Witotire Service (USFWS). 1993. Recovery plan for Geocarpon minimum MacKenzie. Atlanta, Georgia. SOLANUM VIARUM (SOLANACEAE)—TROPICAL SODA-APPLE—CONFIRMED FROM TEXAS Monique Dubrule Reed Mary Ketchersid Biology Department Herbarium 115 Agronomy Field Lab TAMU 3258 TAMU 2488 5 A&M University Texas A&M University College Station, Texas 77845-3258, U.S.A. College Station, Texas 77845-2488, U.S.A. Richard L. Thompson Jasper County 2/1 East Lamar Jasper, Texas 75951, U.S.A. ABSTRACT (Sol i d f South America. Since Solanum viarum, Tropical Soda-A it is known from the southern US, including Louisiana, it las been watched for in Texas. A population has now been confirmed and vouchered from Jasper County, Texas. Other populations probably exist. RESUMEN Solanum viarum le sod ),es una mala hierba agresiva de Sur América. sde que es conocida del Sur de Estados Unidos, incluyendo Louisiana, ha sido b 1a en Texas. Se ha confirmado ahora una poblacion y se han tomado testigos del condado de Jasper, Texas. Solanum viarum Dunal, Tropical Soda-Apple, is a perennial native to South America. It was first found in the U.S. in Florida in 1988 (Wunderlin et al. 1993), though it may have been present as early as 1981 (Coile 1993.) It can be aggres- sively weedy in warmer areas and is now also known from Alabama, Georgia, Mississippi, North Carolina, South Carolina, Pennsylvania, and Tennessee USDA, NRCS 2004). It is also present in Louisiana and Arkansas (Medal et al. 2003). Medal et al. (2003) also listed the plant for Texas, but we have been un- able to locate any voucher specimens or specific collection site information. (It is possible they were referring to the Jasper County population documented here.) The species was not listed for Texas by Correll and Johnston (1970) or by Jones et al.(1997).This is the first vouchered report of its occurrence in the state. In 2002, plants were found growing in a cattle pasture in Jasper County. The land owner cleared as many of the plants as he could but left the dead plants (with mature fruit) on the property. By 2004, the infestation was much worse (hundreds of individuals on ca. 40 to 60 acres), and he sought identifica- tion of the weed. We determined that it was indeed Solanum viarum. The lan owner suspects the seeds may have been introduced ina shipment of hay from Louisiana. SIDA 21(2): 1171-1174. 2004 1172 BRIT.ORG/SIDA 21(2) Voucher specimen: U.S.A. TEXAS. Jasper Co: Ranch, entrance on TX Hwy 63, ca. | mi N jet. TX Hwy 190, pasture. Mary Ketchersid 5-2004-1, 24 May, 2004 (TAMU). Images of the Jasper County plants may be seen at wwwecsdl.tamu.edu/FLORA / cgi/gallery_query?q=solanum+viarum In October, 2004, a live specimen was sent to Texas A@M from Edna in Jackson Co. This represents a second confirmed infestation and underscores the possibility that S. viarum is already more widespread in Texas than is cur- rently known. We have also seen photographs of what appears to be another population S. viarum in Texas, but the property owner has declined to disclose the location of the plants. Solanum viarum (Fig. 1) somewhat resembles S. carolinense L. or S. dimi- diatum Raf, both native to North America, but it has unbranched rather than stellate hairs on the upper leaf surfaces. The plant may reach | m or more tall, with a branched, prickly stem. The leaves are ovate to ovate-triangular, to 15 cm long and 12 cm wide, sinuate, and with sharp prickles to 1 cm or longer on both the upper and lower surfaces. The flowers are white (vs. purple to white in S.carolinense or S. dimidiatum), with the corolla lobes propor tionately narrow and strongly recurved. The fruit are globose, 1.5-2(-3) cm in diameter, striped light and dark green when young, becoming yellow at maturity. A full descrip- tion can be found in Wunderlin et al. 1993) and Bryson and Byrd (1994.) This plant has the potential to be a serious weed in Texas, and it has al- ready been proposed for the Texas Noxious Weed List, as per Texas Agricul- tural Code rule 4TAC § 19.300(a.)(Texas Register 2004). Cattle avoid eating the foliage, allowing the plant to persist in grazed pastures. The fruit, however, is eaten by cattle and small mammals, which distribute the seeds in their feces. Small infestations may be controlled by herbicides, but this may be prohibi- tively expensive over large areas, where mowing before flowering stage is the first line of defense. Trials are underway in Florida to determine the efficacy of the beetle Gratiana boliviana as a biocontrol agent (Medal et al. 2003). Anyone finding what they know or believe to be Solanum viarum in Texas should contact their county agent and/or Mary Ketchersid at the above address for confirmation of identification and specific control instructions. ACKNOWLEDGMENTS Thanks go to Rodney Young, National Plant Identifier for USDA-APHIS-PPQ, for his confirmation of the identification. We thank two anonymous reviewers for helpful comments. REFERENCES Bryson, C.T. and J.D. Byrp. 1994. Solanum viarum (Solanaceae) new to Mississippi. Sida 16:382-385. REED ET AL., SOLANUM VIARUM CONFIRMED FOR TEXAS 1173 Fic. 1. Solanum viarum.A. prickly | 1 fl B. immature fruit, C. flowers I y Cotte, N.C. 1993. Tropical soda apple, Solanum viarum Dunal: the plant from hell. Florida Dept. Agric. & Consumer Services, Div. Pl. Industr. Bot. Circ. No. 27. Corrett, D.S. and M.C. JoHnston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner. Jones, S.D., J.K. Wire, and P.M. Montcomery. 1997. Vascular plants of Texas:a comprehensive checklist including synonymy, bibliography, and index. University of Texas Press, Austin. Meoat, J.C.,D.Ganpotro, and J.P. Cupa. 2003. Biology of Gratiana boliviana, the first biocontrol agent released to control tropical soda apple in the USA. Document ENY-826, IFAS Extension, University of Florida. Texas Recister. May 21, 2004. 29(21):5003-5150. www.sos.state.tx.us/texreg/archive/ May212004/index.htm| 1174 BRIT.ORG/SIDA 21(2) USDA, NRCS. 2004. The PLANTS Database, Version 3.5 (plants.usda.gov Data Center, Baton Rouge, LA. WUNDERLIN, R.P,, K.R. DeLAney, M.Nee,and J.J. MuLLAHey. 1993. Solanum viarum and S. tampicense (Solanaceae):two weedy species new to Florida and the United States. Sida 15:605-611. .National Plant SS NEW DISTRIBUTION RECORDS FOR GAMOCHAETA (ASTERACEAE: GNAPHALIEAE) IN THE UNITED STATES Guy L.Nesom Botanical Research Institute of Texas 509 Pecan Stre Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Gamochaeta stagnalis is reported for the first time for the United States (Arizona and New Mexico). In Arizona these plants have previously been identified as Gamochaeta purpurea and Gamochaeta falcata. The species is common in Mexico and ene is at the northern limit of its distribution in Cochise, ae and Santa Cruz cos., Arizona, and Hidalgo Co., New Mexico. Gamochaeta purpurea Arizona, Ae . its main range in the eastern USS.A. Plants of the southeastern U.S.A. previously identified in some treatments as Gamochaeta falcata are separated into two species: Gamochaeta calviceps and Gamochaeta antillana. A lectotype is chosen for G. antillana. poles haeta placie is eee n eae rom the southeastern U.S.A. but is reported from t ti California. G ‘tata is first reported for Arkansas and Varese and further documented for California. Gamochaeta stachydifolia, a South American native, is re- ported from two counties in California. For each of the 12 Gamochaeta species recorded for the US.A., a hypothesis of nativity is given, with a brief rationale. RESUMEN S it } g li e primera Ve para los E tados Unidos (Arizona y Nuevo México). En ] | a x ro | ] F od i t b + Ff t y CTU PTLOCTIQCLE falcata. La especi ' México y aparentemente tiene su limite Norte de distribucion en los condados de Cochise, Pima, y Santa Cruz, en aa y condado de Hidalgo, en Nuevo México. Gamochaeta pur] Arizona, disyunta de su area prince ae enel oc ot e US. A. Las plantas del Sureste de US jaleat se separan en dos especies: Gamochaet calviceps y fa mochacta cna Se cs un po par G.antillana ! | US A. pero e citan dos col nes en California. Gamochaeta coarctata se cita por primera vez de Arkansas . Virginia, y se edocunente s otras citas para Sa eee SeaG NO nativa de S América, -alifornia d para US.A., se of rece una hipotesis de la posibilidad de ser nativas, con un informe razonado. In connection with preparation of a taxonomic treatment of the genus Gamochaeta Wedd. for the developing Flora of North America volumes, vari- ous range extensions and new records have come to light. Gamochaeta purpurea sensu stricto in Arizona Plants of Gamochaetain Arizona have been identified as G. purpurea (L.) Cabr. (as Gnaphalium purpureum L.: Kearney & Peebles 1960; Lehr 1978) and G. falcata (Lam.) Cabr. (Nesom 1990). Gamochaeta purpurea sensu stricto does SIDA 21(2): 1175-1185. 2004 1176 BRIT.ORG/SIDA 21(2) indeed occur in Arizona, but further study shows that the more common plants are instead a species widespread in Mexico but previously unreported for the United States (see below). The native range of G. purpurea sensu stricto appar- ently is the eastern U.S.A, including eastern Texas, but it occurs as an adventive in many parts of the world. In the U.S.A., Arizona is the only other state in which G. purpurea is known to occur. =, Specimens examined. ARIZONA. Cochise Co.: Chiricahua Mts. Rucker Canyon, streambank near upper road crossing, scarce, ca. 6300 ft, 8 Jun 1980, Toolin 797 (ARIZ). Pima Co.: Rincon Mts., Speed Ranch, 7400 ft, 23 Aug 1909, Blumer 3301 (GH, MO); Spud Rock Spring, wet meadow, 7200 ft, 17 Sep 1982, Bowers R828( ARIZ): Rincon Mts., 7500 ft, 1891, Neally 230(TEX, US); Santa Catalina Mts., Bear Canyon, near Tucson, 27 Apr 1930, Cory s.n.(ARIZ, GH); Mt. Lemmon, control road, 23 Aug 1931, Har- rison and Kearney 8128 (ARIZ, LL), Sabino Canyon, 26 Mar 1926, Loomis 1410 (ARIZ); 10 mi NE of Tucson in Sabino Canyon, frequent along banks of small stream, 15 May 1965, Matthews 387 (ARIZ, ASU), Sabino Canyon near Tucson, 26 March 1926, Peebles et al. 1410 (US); old Sawmill on Apache Camp Trail, 6800 ft, 24 Jul 1914, Shreve s.n. (ARIZ); Sabino Canyon, moist alluvial soil, 2800 ft, 2 May 1903, Thornber 416 (US), Sabino Canyon, moist alluvial soil, 3000 ft, 2 May 1903, Thornber 468 (ARIZ, MO, NMC, US); Sabino Canyon, 5 May 1903, Thornber s.n. (ARIZ); Saw Mill, 24 Jul 1914, Thornber 7303 (ARIZ). The plant of Harrison and Kearney 8128 (ARIZ; the LL specimen has a single stem with drawing of the habit) produced numerous decumbent, rhizome-like or caudex-like branches arising from a central axis and apparently was dis- tinctly perennial. At least one of the plants collected by Shreve (s.n., ARIZ) also appears to have been perennial. It will be interesting to investigate whether the highly unusual modification of Harrison and Kearney 8128 is phenotypic or whether a distinct genetic race might be present. Gamochaeta purpurea probably is native to eastern North America (see below), where it is the least weedy of its congeners, but the species apparently occurs widely through the world as a adventive. Plants of G. purpurea in south- ern Arizona (Pima Co.) occur along sandy banks of perennial streams in Sabino Canyon and Bear Canyon at the base of the Santa Catalina Mountains east of Tucson. The first known collections were made in these long-popular recreation areas in 1903, perhaps accidentally established there through heavy visitation, as the same sites are heavily infested by other, more aggressive non-native spe- cies. On the other hand, collections of G. purpurea also have been made in the Rincon Mts. (Pima Co.) and the Chiricahua Mts. (Cochise Co.), where the spe- cies is less likely to have been introduced by human activity. It also seems un- likely that plants of G. purpurea sensu stricto in scattered Mexican localities were introduced there by human activity. A record of Gamochaeta purpurea from New Mexico (Allred 2003 ) was attributed to documentation in McIntosh (1996), which instead reported records for Pseudognaphalium leucocephalum (A. Gray) Anderb. The voucher for the Gamochaeta record in New Mexico is identified here as Gamochaeta stagnalis (see citation below). Documentation for a report of Gamochaeta purpurea from NESOM, 1177 Montana (Dorn 1984, as Gnaphalium purpureum) has not ben verified. Re- ports of Gamochaeta purpurea from California, Oregon, Was ton, and Brit- ish Columbia are primarily based on G. ustulata (Nutt.) Holub (a native and relatively common species), although three other species (non-native, relatively uncommon) of the genus are now known from California: G. calviceps, G. stachydifolia, and G. coarctata. The status of Gamochaeta stagnalis The present report documents the occurrence of Gamochaeta stagnalis in the US.A.,, where it occurs in Arizona and New Mexico. These plants have previ- ously been identified in Arizona mostly as Gamochaeta purpurea. In the US.A., G. stagnalis does not geographically overlap with any other species and its iden- tity should now be easily ascertained. A full description is given here, since one apparently does not exist elsewhere. Gamochaeta stagnalis (I.M. Johnst.) Anderb., Opera Bot. 104:157. 1991. Gnaphalium stagnale I.M. Johnston, Contr. Gray Herb. ser. 2, 68:99. 1923. TypE: MEXICO. SAN LUIS POTOSI: marshes about San Luis Potosi (‘in palustris circa urbem”), Aug 1876, J.G. Schaffner 225 (HO- LOTYPE: GH!). A specimen at US, Sianereoe. ith “225” and “212” also written on the label), collected in Sep 1879, is the same species but apparently not a duplicate of the type. Plants annual froma short, very slender to filiform taproot, less commonly from very shallow fibrous roots. Stems single and erect or 2-8 and decumbent-as cending, 2.5-20(-35) cm long, densely and loosely arachnoid-tomentose. Leaves mostly cauline, oblanceolate-spatulate to narrowly oblanceolate or nearly lin- ear, 1-4cm long, 2-6(-10) mm wide, basal usually not persistent, cauline oblan- ceolate, slightly reduced upward in size, equally loosely tomentose above and beneath or the adaxial surface glabrescent and greener. Capitulescence a capi- tate cluster (in smallest plants) of heads or an interrupted series of small glom- erules subtended by divergent-ascending bracts similar to the upper cauline leaves, sometimes branching at lower nodes. Involucres campanulate, 2.5-3 mm high, conspicuously imbedded in loose tomentum, the outer bracts basally hairy; inner phyllaries narrowly oblong-lanceolate, with rounded-obtuse, whit- ish lamina, usually purple above the stereome and along proximal margins of the lamina, outer phyllaries ovate-triangular, translucent; receptacles deeply concave to crateriform. Florets: bisexual (2-)3(-4); all corollas purplish-tipped. Flowering (Mar-)Apr(-May). Sandy, often moist soil, washes and permanent streams, canyon bottoms, flower beds, riparian, desert grassland, juniper-grass- land, creosote bush-mesquite-cholla, oak woodland; 900-1750 m; Ariz., N.Mex.; Mexico (Sonora, Chihuahua, Coahuila, Nuevo Leon, Baja California Sur, Sinaloa, Durango, Zacatecas, San Luis Potosi, and other states to the south). — Specimens examined: ARIZONA. Cochise Co.: floodplain of Miller Canyon, 0.8 mi by road W of Hwy 92, under Quercus emoryi in open woodland, 14 Apr 1991, Bowers 3426 (ARIZ); Dragoon Mts., Noonan Canyon, SE slope of S-facing saddle, 5080 ft, with Fouquieria, 29 Apr 1983, Caffey-Moquin 396(UNM) 1178 BRIT.ORG/SIDA 21(2) pr 1952, Goodding 58-52 (ARIZ); Huachuca and 399 (UNM); Whetstone Mountains, rocky draw, 27 A Mountains, Coronado National Memorial, rocky bench in wash, FE end of park, 5000 ft, 20 Apr 199] McLaughlin 6233 (ARIZ); San Bernardino Ranch, 18 mi E of Douglas, moist area around Middle Well ay 1981, Marrs-Smith 561 (ASU): SW corner of Chiricahua National Monument, grass- land, juniper, 5400 ft, 19 Apr 1975, Reeves R2595 (ASU). Pima Co.: Rincon Mountains, along the Man- ning Camp trail, moist draws in desert grassland, 4500 ft, 27 Apr 1983, Bowers R124 (ARIZ): Rincon Mts., unnamed canyon at W base of Tanque Verde Ridge, 3200 ft, moist soil in streambed, 29 April 1984, Bowers R1433 (ARIZ, LL): Altar Valley, 8 airline mi SSW of Robles Junction (Three Points), creo- sote bush-mesquite-cholla woodland, ae in disturbed areas, 2900 [t, 6 Apr 1973, Holmgren 6668 (NY); Baboquivari Canyon, ll Mar 1926, Ledingand Thackery 1123 (ARIZ); Forestry Cabin at W Papago a ee vation, Pe manent stream in oak-Sonoran desert zone, grazed, 6-7 Apr 1973, Lehto etal. 1 JS). Pag vation, 0.3 mi E of Topawa, mes- quite-cactus, bur-sage desert, ae meches 6 Apr 1973, tape al 10749( ARIZ); 22.0 mi Sof Rob- les Jet.at Las Delicias Ranch Road along Hwy 286, desert grassland _ yaa ae burroweed, Opun- tid spp., 3200 ft, 14 May 1988, McLaughlin 4568 (ARIZ): Buenos Ai onal Wildlife Refuge, along Brown wash, with hackberry and mesquite, 3600 ft, 25 Apr 1991, een 6253 (ARIZ), Lower Bear Canyon, moist sand along stream, 3200 [t, 13 Apr 1961, Mason 1908 (ARIZ); along Santa Cruz River at Cortaro Road, NW of Tucson, 5 Apr 1976, Mason 3171a (ARIZ); Little Tucson, Ascencio flood- water field section last plowed summer 1978, 17 Apr 1979, pede Hate ie Pima Co. Coyote Mts., 5, Parker 5797a (ARIZ, BRIT, LL, Santa Catalina Mountains, Ventana 3800 ft, 25M base of Baboquivari Peak, P = TC) = =a) ay £6) Mendoza Canyon, 3200 ft, abundant on canyon floor, 22 Apri NY); wet banks of the Rillita, 14 Apr 1881, Pringle 13744 (GH, MO); Canyon, 1908, Shreve s.n. (ARIZ): Fort Lowell, Rillito, 23 Apr 1903, Thornbers.n.(ARIZ): Tucson, First Avenue at Roger Road, irrigated flower bed around parking lot, 14 Aug 1994, Van Devender 94-444 (TEX). Santa Cruz Co.: Nogales-Lochiel Road, 6 mi from Ariz. Hwy 82,9 airline mi E of downtown Nogales, sandy soil around oak tree, 4200 ft, 18 Apr 1973, Holmgren 6866 (ARIZ, NY); Agua Caliente ng stream near road crossing, riparian vegetation with Celtis, Baccharis, Fraxinus, 3800 Canyon, alot {t, 2 Apr 1978, Reeves 6640 (ASU); Santa Rita Mountains, Gardner Canyon, 5700-5800 ft, 8 May 1975, Van Devenders.n. (ARIZ). NEW MEXICO. Hidalgo Co.: Peloncillo Mts,, Granite Gap, occasional on W- facing granitic slope with Ericameria Vaciiolin Fouquieria splendens, Agave palmeri, Opuntia phaeacantha var. discata, 21 Apr 1993, McIntosh 2665 (NMC). Localities for Gamochaeta stagnalisin Arizonaand New Mexico areat the north- western extremity of its overall range, where flowering is restricted to the end of the cool season. The species is common and widespread in Mexico, from Baja California Sur, Sonora, and Chihuahua southward and eastward to Jalisco and Colima, Nuevo Leon, San Luis Potosi, and Veracruz, where it occurs at eleva- tions of 200-1800(-2600) meters in rocky or gravelly soil, including stream beds and other periodically wet sites, in areas of thorn-scrub, tropical deciduous, or oak woodland, usually in open or disturbed sites. In Mexico it [lowers Decem- ber through May but sometimes continues longer in wet seasons Plants of Gamochaeta stagnalis are recognized by their annual duration, usually from a filiform taproot, oblanceolate leaves equally tomentose on the lower and upper surfaces, interrupted capitulescence, small, basally tomentose heads, and phyllaries conspicuously purplish at the stereome/lamina junction and along the proximal margins of the lamina. It is similar to and probably closely related to G. antillana (below). NESOM, 1179 The status of Gamochaeta antillana Gamochaeta antillana (Urb.) Anderb., a common species in the southeastern US.A., was combined in concept with Gamochaeta calviceps (Fern.) Cabr. and identified as Gamochaeta falcata (Lam.) Cabr. by Nesom (1990). Godfrey (1958) separated G. calviceps and identified the other species as G. falcata. It is now clear that two taxa occur in this region, and they are now known to be widely sympatric, countering Godfrey's notion that they might be treated as geographic varieties. Gamochaeta antillana (Urb.) Anderb., Opera Bot. 104:157. 1991. Gnaphalium antillanum Urban, Repert. Spec. Nov. Regni Veg. 13:482.1915. Tyre: CUBA. “In insula Saba ad Great Hill et Gumbeygut, m. April, fruct. delapsis, Suringar” (holotype: B, apparently de- stroyed). Two paratype collections cited by Urban (Britton 10009; Britton 9619) are at NY, with internet- ae Heo on the NY type database. Peter see in mh 1982, annotated Llo n designated a Suringar s| ed 2 additional speci- my aan mens— ee ae and Britton, Britton and Cowell 9619. Thes Suringar specimen no longer exists; | believe that the fruiting aes Britton, Britton and Cowell 9619 should be desig- nated lectotype.” It obviously is somewhat arbitrary, but | were that the features of the species are better paiia sy ants of 10009 and designate it here as the LECTOTYPE: Cuba. Prov. Pinar del R afre, hillside, 15 Mar 1911, N.L. are ee Gnaphalium subfalcatum Cabr., Rev. Mus. La Plata (ns.) Bot. 4:174. 1941. Gamochaeta subfalcata (Cabr.) Cabr, Bol. Soc. Argent. Bot. 9:383. 1961. Type: ARGENTINA. PROV. BUENOS AIRES: Pdo Avellaneda, Isla Maciel, 12 Oct 1920, Cabrera 944 (HOLOTYPE: LP). Cabrera (1961) cited collections of Gamochaeta subfalcata from Texas and Florida, extending the range far from northeastern Argentina, as circumscribed by the original citations (Cabrera 1941). Freire and Iharlegui (1997) also identi- fied this species in the U.S.A. as G. subfalcata, and it seems inescapable that C. antillana and G. subfalcata are synonyms. Gamochaeta antillana is known to occur in Alabama, Arkansas, Florida, Georgia, Louisiana, North Carolina, Okla- homa, Mississippi, South Carolina, Tennessee, Texas, and Virginia; also in South America, Europe, and New Zealan Gamochaeta antillana is closely similar to G. stagnalis but the plants often are generally taller (6-40 cm vs. 2.5-20(-35) cm in G. stagnalis) and the basal leaves are oblanceolate with the cauline quickly becoming linear (in G. stagnalis, the cauline leaves and those subtending the clusters of heads are oblanceolate). Gamochaeta antillana occurs in humid climates and habitats while G. stagnalis isa species of arid climate and habitat. Further study of the distinction between these two taxa is needed. a. Involucres 3-3.5 mm, lightly arachnose only at the base or not at all; capitulescence interrupted at least distally, main axis visible to terminal heads; phyllaries in 5-7 series, outer and middle ovate-triangular with sharply acute-acuminate apices, 1/ 3-1/2 as long as the inner, none with purplish color; a oie Jul mochaeta calviceps Involucres 2.5-3 mm, seated in tomentum; capitulescence initially ae an uninterrupted, at least distally, main axis obscured by clustered heads; phyllaries in @ 1180 BRIT.ORG/SIDA 21(2) ~4(—5) series, outer and middle ovate-lanceolate with narrowly to broadly acute apices, outer 1/2-2/3 as long as inner, at least innermost commonly tinged with ce he at stereome-lamina junction; fl (Feb-)Mar—May, sometimes later with moistu Gamochaeta antillana Gamochaeta calviceps in California Gamochaeta calviceps occurs widely in the heastern U.S.A. The first known North American records outside of that region are reported here. It is known to occur in Alabama, Arkansas, California, Florida, Louisiana, Mississippi, North Carolina, Oklahoma, South Carolina, Texas, Virginia, as well as South America, Europe, and New Zealand. As noted above, G. calviceps and G. antillana consti- tute the plants in the eastern U.S.A. most commonly identified in the past as Gamochaeta falcata. CALIFORNIA. Fontes Costa Co.: Tilden | Eeoiouay Park Botanic Garden, Wildcat Canyon; a rapidly ading ly came with plant material from Delano, Kern Co., 27 Jun 1975, True 7872A (TEX) San Diego Co.: : Peninsular Ranges, near Riverside Co. line, E of I-15 off Pala Road along Rancho Heights Rd, 1364 ft, chaparral, mostly past flower, 24 Jun 2003, Spjut and Marin 15384 (BRIT, UCR). Gamochaeta stachydifolia in California This species is known to me by two collections from central California, the lo- calities separated by about 200 kilometers. The 1990 collection suggests that it has probably is naturalized in that region and should be expected at more lo- calities. The plants are recognized by their slender-taprooted habit (probably annual), oblanceolate and concolorous leaves, cylindric capitulescence, acute to acute-acuminate outer and middle phyllaries, inner phyllaries with brown- ish-hyaline, rounded-apiculate lamina, and yellowish-tipped florets. The GH collection (Mason 6991) was annotated by Peter Michael in 1990 as Gamochaeta berteriana (DC.) Cabr., but this identification is problematic, as G. berteriana apparently is perennial, thicker stemmed, and bears heads in capitate clusters; it is native to high elevation habitats in Chile (type: Chile, 1833, Bertero 8222, B, photo-TEX!). Freire and Iharlegui (1997) noted that the range of G. stachydifolia includes Argentina, Brasil, and Uruguay. — Gamochaeta stachydifolia (Lam.) Cabr., Bol. Soc. Argent. Bot. 9:382. 1961. Gnaphalium stachydifolium Lam., Encycl. 2:757. 1786. Gnaphalium purpureum L. var. stachydifolium (Lam.) Baker in Martius, Fl. Bras. 6(3):125. 1882. Probable HOLOTYPE (see com- ments by Cabrera 1961): Uruguay or Argentina. “Des environs de Montevideo et de Buenos Aires,” without date, Commerson s.n. ie F photo-LL ). Lamarck noted that “Commerson a trouve cette espeéce ¢ a Monte- Video. Plants annual, slender taprooted. Stems 4-15cm high, erect, single from the base, densely and loosely gray-white tomentose-arachnoid. Leaves basal and cauline, basal mostly withered and withering by flowering, oblanceolate, 1-2 cm long, 2-4 mm wide, cauline similar to basal, oblanceolate, commonly folded, subclasping but not auriculate, 2-3 cm long, 3-6 mm wide, continuing nearly unreduced into lower inflorescence but none longer than heads, dark apical NESOM, 1181 mucro often evident, ey Buy white tomentose-arachnoid on both surfaces. conti der 2-3(-4) cm long, lO-12 mm wide (pressed). involacies campanulate, 3.5- 4 mm high; phyllaries in 4-5 gradate series, outer ca. 1/3 as long as innermost, outer and middle narrowly ovate-triangular, apically acute to acute-acuminate, lightly tomentose at the very base, inner- most oblong, stereome ca. 1/2 length, lamina brownish-hyaline, apically rounded-apiculate; receptacles shallowly concave. Florets: bisexual 2-4; all co- rollas yellowish. Cypselae: mature fruits not seen. Collections examined. CALIFORNIA. Amador Co.: Sierra Nevada foothills, hill above lone, 25 Apr 1932, Mason 6991 (GH, LL). Butte Co.: ca. 1/4 miS of the Feather River, ca. 0.4 mi W of Pacific Heights Road, ca. 4.5 mi SW of Oroville, T18, R3E, ne/S3, riparian woodland (destroy od), 100 ft, uncommon, inconspicuous, growing on dry, bare disturbed, sandy soil in the borrow area, 28 Apr 1990, Ahart 6466 (MO) Gamochaeta coarctata in Arkansas, California, and Virginia Gamochaeta coarctata (Willd.) Kerg. was previously noted to occur in Califor- nia (Nesom 1990), but it was identified as Gamochaeta americana. In view of the rapid spread and pervasive occurrence of this species in the southeastern US.A.,, it seems likely that it also is becoming increasingly common in Califor- nia. It is common throughout Louisiana, and its representation in southern Ar- kansas probably is already significantly greater in herbaria than the single record reported here. I have seen the following specimens. ARKANSAS. Bradley Co.: “Southern Bluff” ca. 2.3 mi NW (by air) of the center of Warren, 26 Jun 1976, Locke 2002 (BRIT). CALIFORNIA. Humboldt Co.: Canyon Creek, 6 mi SE of Blue Lake, hillside pas- ture in logged area, local and scarce, 1200 ft, 1 Aug 1936, Tracy 15057 (NCU, TEX). Sacramento Co.: weed in irrigated alfalfa field, Aschwanden farm, 3 mi W of Galt, 10 Aug 1953, Tucker 2674 (SMU). Stanislaus Co.: San Joaquin Valley, near Ceres and Turlock, 2 mi WSW of Keyes, uncommon annual weed in almond orchard, 80 ft, 8 Jul 2000, Sanders 23532 (BRIT). VIRGINIA. N end of Hog Island, inner dune, | Jul 1996, McAvoy 1603 (DOV) Further comments on biology and nomenclature of Gamochaeta coarctata are given in two other papers in this issue (Nesom 2004; Pruski &@ Nesom 2004). Nativity of North American Gamochaeta species Assessment of the nativity of North American species of Gamochaeta is prob- lematic. Most Gamochaeta species are native to South America, and most of the North American species cl istically occur in ruderal habitats, commonly in company of known non-native species of various families. Some, if not all, of the North American Gamochaet as weeds in parts of the world other than South America (although. inconsistencies in identification and ap- plication of names make it difficult to accurately evaluate overall distributions of the widespread species). Thus by behavior and association, all rudera Gamochaeta species in North America might also be expected to be non-na- tive. The mode of introduction of those clearly non-native is not known. 1182 BRIT.ORG/SIDA 21(2) Circumstantial evidence, however, suggests that some of the North Ameri- can Gamochaetd species are native. Gamochaeta purpurea and G. ustulata were described from collections made early in the history of the U.S.A., presumably efore non-native colonizers became abundant, others are known only from more recent collections. Several species are distributed over broad latitudinal and ecological range, suggesting that geographic differentiation may have oc- curred; the geographic range (and presumed genetic variability) of others is more restricted. Four of the species suggested as native on a geographic-eco- logical basis form two species pairs (the two of each pair with strong morpho- logical similarities: G. purpurea and G. sphacilata, G.argyrinea and G. ustulata), suggesting that the evolutionary differentiation was autochthonous. Gamochaeta pensylvanica, G. antillana, and G. stagnalis are similar among themselves and possibly closely related; their nativity is uncertain, but at least itseems likely that G. stagnalis is native. For those non-native, evidence is strong that they are naturalized (sensu Nesom 2000a). For each of the Gamochaeta species recorded for North America (north of Mexico), a hypothesis of nativity is given, with a brief rationale. Distribution maps for G. purpurea, G.argyrinea, G. ustulata, G. chionesthes, G. simplicicaulis, and G. coarctata are provided in Nesom (2004). Gamochaeta purpurea (L.) Cabr—Native: widespread in the eastern U.S.A. over a broad latitudinal and ecological range; early collections from known range in the U.S.A; possibly closely related to G. sphacilata, which apparently occurs natively over a wide area, including South America and Mexico, into southwest Texas. Gamochaeta purpurea sensu stricto is found over a wide area of peninsular Florida, but G. argyrined and G. chionesthes, both segregated from the concept of G. purpurea in the US.A., are restricted to the northern counties of the state (a loan of specimens from USF was extremely helpful in establish- ing this). ~— Gamochaeta sphacilata (Kunth) Cabr—Native: widespread from South America to the US.A., occurs in essentially undisturbed habitats at mid and relatively high-elevation in Mexico and the northern extension of its range in trans-Pecos Texas; possibly closely related to G. purpurea, which apparently is native to the eastern U.S.A. Gamochaeta argyrinea Nesom—Native(?): widespread in the eastern U.S.A. over a considerable latitudinal and ecological range and also known from Pu- erto Rico; early collections from known range in the U.S.A. probably closely related to G. ustulata, which apparently is native to the western U.S.A. Gamochaeta ustulata (Nutt.) Holub—Native: distinctive habitat and geo- graphic range in Pacilic coast states, over a wide latitude, mostly in coastal and near-coastal habitats; early collections from known range in the western U.S.A.; closely similar and probably closely related to G. argyrinea, which perhaps is native to the eastern US.A. a NESOM, 1183 Gamochaeta chionesthes Nesom—Non-native: relatively scattered and re- cent collections in the southeastern U.S.A. (see Nesom 2004). It possibly has been identified in South America by a misapplied name (G. americana?); possi- bly closely related to G. simplicicaulis, a native of South America. Gamochaeta simplicicaulis (Willd. ex Spreng.) Cabr—Non-native: scattered and recent collections in the southeastern U.S.A., the earliest known in 1957- 1959, when it was discovered in nine counties of North Carolina and South Caro- lina (Nesom 1999, 2000b). Widely distributed in South America and known from early collections there; recorded as adventive in other parts of the world before its discovery in North America. Gamochaeta coarctata (Willd.) Kerg.—Non-native: collections from the US.A. before about 1970 are rare. Small (1933) did not include this distinctive species in his treatment of the Southeastern flora. Goalie (1958) noted that he knew the species (as Gnaphalium spicatum Lam.) from from around Wilmington, North Carolina, and from Florida, “in and around Tallahassee, thence westward to Pensacola.” Perhaps the earliest collection or one of the ear- liest was made in 1949 in Wilmington (Godfrey 49341, originally identified as Gnaphalium purpureum, SMU, NCU), where it was “abundant in vacant lots and weedy places”, it was collected again in Wilmington in 1950 (Godfrey 50362, SMU), identified as an “unusual form” of Gnaphalium purpureum. The range of G. coarctata in the southeastern U.S.A. is now apparently much more con- tinuous than indicated by existing collections (personal observation), suggest- ing a recent and rapid expansion. The type of Gnaphalium coarctatum and its synonym Gnaphalium spicatum was described from Uruguay from a collec- tion made in the 1700s (Pruski & Nesom 2004), and it seems likely that the species is native to South America. It is documented as adventive in other parts of the world. Gamochaeta pensylvanica (Willd.) Cabr. (synonyms: Gnaphalium spathulatum Lam. [non Burm. f.|, Gnaphalium peregrinum Fernald)—Non-na- tive(?): common in the southeastern U.S.A. (nearly restricted to Atlantic Coast and Gulf Coast states); common in eastern South America and throughout the world as a weed. Similar and perhaps related to G. antillana, for which the na- tivity also is uncertain but suggested to be North American and Antillean. On balance, however, it seems likely that G. pensylvanica arrived early as an ad- ventive in North America, especially in view of its apparent complete fidelity toruderal habitats on this continent and its near-cosmopolitan occurrence as a weed. In Willdenow’s proposal of the name Gnaphalium pensylvanicum, he noted that the ‘habitat’ was in Virginia and in Pennsylvania, north of its char- acteristic range in the U.S.A. My guess is that it was collected as a ballast waif in Philadelphia prior to 1809, the year of Willdenow’s proposal. The species is shown only in Pennsylvania County by Wherry et al. (1979), who noted that it is “introduced” in the state; it was not included at all in a later summary of the — 1184 BRIT.ORG/SIDA 21(2 a= Pennsylvania flora (e.g., Rhoads & Block 2000). The type of Gnaphalium spathulatum, described by Lamarck in 1788, was from plants cultivated at the “Jardin du Roi’ in Paris; Lamarck noted that he did not know the origin of the garden plants but that Commerson had found a similar form near Buenos Aires. In his description of Gnaphalium peregrinum, Fernald (943) noted that G. spathulatum was a later homonym (non Burm. f. 1768) and probably the same species as G. peregrinum, but because of his uncertainty regarding the identity of the plant in the type photo (G. spathulatum Lam.), he chose a new type for the new name. Burman’s name (Prodromus florae capensis 25. 1768) is validated by citation of a figure in Breyne’s Prodromi (tab. 18, fig. 3.1738) and accompany- ing legend (p. 29)—it apparently is a species of Helichrysum. Gamochaeta antillana (Urb.) Anderb.—Native(?): common in the southeast- ern U.S.A., most in coastal states; described from Cuba in 1915, known from most islands of the Antilles, South America, and apparently weedy in other parts of the world. Closely similar to and possibly closely related to G. stagnalis. Gamochaeta stagnalis (1.M. Johnst.) Anderb.—Native(?): common in north- ern Mexico and reaching southern Arizona and southwestern New Mexico, where it flowers in early spring in desert habitats. Probably closely related to G. antillana but different in geography and ecology. Gamochaeta calviceps (Fern.) Cabr—Native(?): described in 1935 from Vir- ginia and known mostly from recent collections in eastern North America (states of the Atlantic Coast and Gulf Coast); apparently widespread in South America other parts of the world. Gamochaeta stachydifolia (lam.) Cabr—Non-native: known in North America only from two counties in central California. Native to South America. — ACKNOWLEDGMENTS lam grateful to Alan Weakley and Carol McCormick for checking locality data of NCU collections and to John Pruski and Susana Freire for reviews. Loans of specimens were studied from ARIZ, ASU, DOV, MO, NMC, and USE Data also were obtained on visits to MO, NCU, NY, GH, TEX, and US. REFERENCES Autrep, K. 2003. A working index of New Mexico vascular plant names. Accessed 1 Mar 2004. Casrera, A.L. 1941. Gnaphalium.|In: Compuestas Bonaerenses. Rev. Mus.La Plata (n.s.), Secc. Bot., 4:1-450. Pp. 161-180 CasrerA, A.L. 1961.Observaciones sobre las Inuleae-Gnaphalinae (Compositae) de América del Sur. Bol. Soc. Arg. Bot. 9:359-386. Dorn, R.D. 1984. Vascular plants of Montana. Mountain West Publishing, Cheyenne, Wyoming. Fernato, M.L.1943.Virginian botanizing under restrictions. Rhodora 45:357-413,445-480. NESOM 1185 Freire, S.E.and L. IHartecul. 1997. Sinopsis preliminar del género Gamochaeta (Asteraceae, Gnaphalieae). Bol. Soc. Argent. Bot. 33:23-35. Goorrey, R.K. 1958. A synopsis of Gnaphalium (Compositae) in the southeastern United States. Quart. J. Florida Acad. Sci. 21: 177-184. Kearney, T.H.and R.H. Peestes. 1960. Arizona flora and supplement. Univ. of California Press, Berkeley. Lene, JH. 1978. A catalogue of the flora of Arizona. Desert Botanical Garden, Phoenix, Arizona. McIntosh, L. 1996. Seven additions to the flora of New Mexico. Phytologia 81:365-368. Nesom, G.L. 1990.The taxonomic status of Gamochaeta (Asteraceae: Inuleae) and the spe- cies of the United States. Phytologia 68:186-198. Nesom, G.L. 1999. Gamochaeta simplicicaulis (Asteraceae: Gnaphalieae) in four southeast- ern states and new for North America. Sida 18:1259-1 264. Nesom, G.L. 2000a. Which non-native plants are included in floristic accounts? Sida 19: 189-193. Nesom, G.L. 2000b. Gamochaeta simplicicaulis (Asteraceae: Gnaphalieae) in Georgia. Sida 19:413. Nesom, G.L. 2004. New species of Gamochaeta (Asteraceae: Gnaphalieae) from the east- ern United States and comments on similar species. Sida 21:717-741. Pruski, J. and G.L. Nesom. 2004. Gamochaeta coarctata, the correct name for Gamochaeta spicata (Asteraceae: Gnaphalieae). Sida 21:711-715. Ryoaps, A.F.and T.A. Bock. 2000.The plants of Pennsylvania: An illustrated manual. Univ. of Pennsylvania Press, Philadelphia. Smatt, J.K. 1933. Manual of the southeastern flora. Univ. of North Carolina Press, Chapel Hill. Wuerry, E.T., J.M. Foas, Jr. and H.A. Want. 1979. Atlas of the flora of Pennsylvania. Morris Arboretum of the Univ. of Pennsylvania, Philadelphia. pase VASCULAR FLORA OF WASHITA BATTLEFIELD NATIONAL HISTORIC SITE, ROGER MILLS COUNTY, OKLAHOMA Bruce W. Hoagland Amy Buthod Oklahoma Biological Survey Oklahoma Biological Survey and Department oes University of Oklah University of Oklahom Norman, Oklahoma 73019, U.S.A Norman, Oklahoma 73019, sh bhoagland@ou.edu Wayne Elisens Oklahoma Biological Survey and Department of Botany and Microbiology University of Oklahoma Norman, Oklahoma 73019, U.S.A. ABSTRACT This article reports the results of a vascular plant in pene of the Washita Battlefield National His- toric Site in western Oklahoma. Two hundred and seventy-two species of vascular plants were col- lected from 201 genera and 62 families. T ne most specious families were the Poaceae (53), Asteraceae (48), Fabaceae (22) and Euphorbiaceae (22). One hundred and seventy-five species were perennials, ninety-five annuals, and 2 biennials. Twenty-eight woody plant species were present. Twenty-one species exotic to North pon a were collected representing 7.7% of the flora. Five species tracked by J the Oklahoma Natural I y were found. This study reports 205 species previously not } documented in Roger Mills County: ABSTRACT Este articulo presenta los resultados de un inventario de rota vascular del Washita Battlenesd Na- d | tional Historic Site en nel ee de Oklahoma. $ S] le plantas vasculares | 201 géneros y 62 familias. Las familias con mas epecus es Poaceae (53), ena ae (48), me (22) y Euphorbiaceae (22). Ciento setenta y cinco pepe eran perennes, noventa y cinco anuales, y 2 bianuales. Estaban presentes s lenosas. Se colectaron veintiuna especies exoticas para Norte América que representan el 77% de la flora. Se encontraron cinco especies Segeicas por el seuss a Natural Heritage Inventory. Este estudio cita | i nel condado de Roger Mills. INTRODUCTION The objectives of this study were twofold; to fill a gap in floristic data for west- ern Oklahoma and provide resource managers at the Washita Battlefield Na- tional Historic Site WBNHS)a comprehensive species list. Prior to 2002, when collecting began for this study, 446 specific and intraspecific taxa were reported from Roger Mills County (Hoagland 2004). Erigeron bellidiastrum Nutt., col- lected by J. Engleman on 3 July 1919, was the first botanical specimen gathered in Roger Mills County. There are no subsequent collection records until 1929. SIDA 21(2): 1187- 1197. 2004 1188 BRIT.ORG/SIDA 21(2) Peak collecting years in Roger Mills County were 1939 (261 specimens), with the return of J. Engleman, and 1976 by Susan Barber and Rahmona Thompson (124 specimens) on behalf of the Robert Bebb Herbarium at the University of Oklahoma (Hoagland 2004). During the course of this research, Freeman et al. (2003) published a floristic list from the Thurman Ranch in Roger Mills County, located south of WBNHS, which documented 470 species from 85 families. Study Area The WBNHS was established on 12 November 1996 and encompasses 136 hect- ares in Roger Mills County (Fig. 1). Latitudinal extent ranges from 35.63°N to 35.62°N and longitudinal extent from 99.70°W to 99.71°W. The WBNHIS is lo- cated within the subtropical humid (Cf) climate zone (Trewartha 1968). Sum- mers are warm (mean July temperature = 27.7°C) and humid, whereas winters are relatively short and mild (mean January temperature = 1.9°C). Mean annual precipitation is 105.6 cm, with periodic severe droughts (Oklahoma Climato- logical Survey 2004). Physiographically, the study area is located in the Osage Plains section of the Central Lowlands province (Hunt 1974) and within the High Plains province of Oklahoma (Curtis @ Ham 1979). Elevation in the study area ranges from 588 m along the Washita River to 610 m. The surface geology is primarily Permian red sandstone in the uplands to the south of the Washita River, and Quaternary silt, sand and clay adjacent to and north of the river (Bran- son & Johnson 1979). The primary soil association at WBFNHS is the Yahola- Port, which is composed of alluvial soils on bottomlands and terraces. The Woodward-Quinlan association occurs on uplands and is level to very steep loamy soils underlain by red sandstone (Burgess et al. 1959). The predominant potential vegetation types are mixedgrass prairie with a smaller component of bottomland forests and stabilized dunes (Duck & Fletcher 1943). Much of the Washita River bottomlands were cleared for agriculture and pasturage. a METHODS Fight collection sites were established at WBNHIS for intensive floristic sam- pling. Sites were selected following a review of US Geological Survey 1:24,000 topographic maps and field reconnaissance. The predominant vegetation asso- ciation at these sites was classified according to Hoagland (2000). Collection effort was not limited to established sites. Collections were made on a monthly basis from March through October 2002. Vouchers for species exotic to North America were made from naturalized populations only, thus excluding culti- vated and ornamental plants. Specimens were processed at the Robert Bebb Herbarium of the University of Oklahoma (OKL) following standard herbarium techniques. Manuals used for specimen identification included Correll and Johnston (1970), Gould (1975), Waterfall 1969) and Great Plains Flora Associa- tion (1986). Origin, whether native to introduced to North America, was deter- HOAGLAND ET AL., FLORA OF HISTORICAL SITE 1189 Observation ®CS1 @cs3 Tower ree Cyr ges Washita Battlefield National Historic Site Fic. 1. L ati nofW. eee sf lafialA Al 4 Tut tf Davnyw AASII fa y Oklahoma mined using Taylor and Taylor (1991) and the United States Department of Ag- riculture-Natural Resources Conservation Service (USDA-NRCS 2004). The nomenclature used is in concordance with USDA-NRCS (2004). Voucher speci- mens were deposited at OKL. RESULTS AND DISCUSSION A total of 272 vascular plants in 201 genera and 62 families were collected (Table 1). Among the angiosperms, 66 were monocots and 205 were dicots. One gym- nosperm was found. The most species were collected from the families Poaceae (53), Asteraceae (48), Fabaceae (22), and Euphorbiaceae (14). The genera Chamaesyce (5), Eragrostis (4), Dalea (4), and Solanum (4) had the most spe- cies. Ninety-seven species were annual or biennials, and 178 perennial. Twenty-one exotic species from 14 families were collected, representing 7.7% of the flora. The greatest number of exotic species were in the families Poaceae (11) and Asteraceae (4). This is higher than the 10% exotic flora reported from the Thurman Ranch (Freeman et al. 2002), but is comparable to recent floristic inventories from other areas in Oklahoma. For example, a flora of the Chickasaw National Recreation Area reported 12% exotic species (Hoagland & Johnson 1190 BRIT.ORG/SIDA 21(2 Taste 1.Summary of floristic collections at the Washita — National Historic Site, Roger Mills County, Oklahoma. Table format follows Palmer et al. (19 Taxonomic Group Species Native spp. Introduced spp. Equisetophyta 0 @ 0 Pteridophyta 0 0 0 Coniferophyta ] | 0 Magnoliophyta Magnoliopsida 205 184 21 Lillopsida 66 56 10 Total 271 240 3] 2001), 9% at Oologah Wildlife Management Area (Hoagland & Wallick 2003), 15% at Keystone Wildlife Management Area, and 11% for an inventory of Tillman County (Hoagland et al., in press). However, the percentage was lower, 6.6%, at Red Slough and Grassy Slough in southeastern Oklahoma (Hoagland & Johnson, in press). Five species tracked by the Oklahoma Natural Heritage Inventory were found: Argythamnia humilis (G5S253), Gaura brachycarpa (G4G5 SIS2), Solanum triflorum (G5S182), Sporobolus giganteus (G5S1S3), and Zinnia gran- diflora (G5S?2). Species are ranked according to level of imperilment at the state (S) and global (G) levels on a scale of 1-5; 1 representing a species that is imper- iled and 5 one that is secure (Groves et al, 1995). Asaresult of this study, 651 species are now known to occur in Roger Mills County. Of the 361 species reported in this study, 156 had been previously col- lected in the county (Hoagland 2004). This study documented 205 species not previously reported from Roger Mills County. When compared with the Demp- sey Divide site (Freeman et al. 2002), there were 219 species found at both sites. However, 53 species were documented at WBNHS that were not reported at Dempsey Divide. Two hundred and fifty-one additional species were found on the Dempsey Divide that were not found at the WBNHS. The difference in total area of the two sites may account for this discrepancy; the Thurman Ranch is 3,755 hecatres in area and contains 19 vegetation types whereas WABA is only 136 hectares with four vegetation types. (Freeman et al. 2002). The 8 collection sites occurred within four vegetation associations. A brief description of each follows: Sapindus saponaria woodland association This association was limited to large sand dunes located on the northside of the Washita River. Although S. saponarida was abundant, dominance was locally vari- able. Celtis laevigata var. reticulata was a common woody species in this vegeta- tion association, as was UImus pumila,a species introduced to western Oklahoma for shelterbelt plantings. Other common woody species included Forestiera pubescens, Gymnocladus dioicus, Prunus angustifolia, Ribes aureum, and Sideroxylon lanuginosum. Associated herbaceous species included Andropogon hallii, Argemone polyanthemos, Asclepias arenaria, Cyclanthera dissecta, Dimorphocarpa candicans, Froelichia gracilis,and Funastrum cynanchoides. Schizachyrium scoparium—Bouteloua hirsuta herbaceous association Occurred on Permian red sandstone in the uplands overlain by the Woodward- Quinlan soil association. Associated species included Aristida oligantha, Am- brosia psiolstachya, Bouteloua curtipendula, Eriogonum annuum, Penstemon albidus, Sphaeralcea coccinea, Thelesperma megapotamicum,and Yucca glauca. Disturbed areas and old-field vegetation This includes much of the Washita River floodplain, which had been under cultivation for many years. It also includes roadsides and areas visited by WBNHS visitors and other areas exhibiting signs of physical disruption. Com- mon plants in disturbed areas and old fields included Ambrosia trifida, Bothriochloa ischaemum, Chenopodium simplex, Cynodon dactylon, Digitaria ciliaris, Melilotus officinalis, and Sorghum halepense. APPENDIX 1 Annotated species list for the Washita Battlefield National Historic Site. The first entry indicates life history (A=annual, P=perennial), species not native to North America (designated with an asterisk), habitat (DAOF=disturbed area/ old-field; MGP=mixed grass prairie, RA=riparian area; SW=sandy woodland), and collection number. Voucher specimens were deposited at the Robert Bebb Herbarium at the University of Oklahoma (OKL). PINOPHYTA Apocynaceae ADOcy anna mL-P; SW;WAS192 Cupressaceae Jur niperus virginiana L.—P; SW; WAS193 Asclepiadaceae Asclepias arenaria Torr.—P; MGP, SW; WASO MAGNOLIOPHYTA Asclepias asperula (Dcne.) Schlechter— p ee MAGNOLIOPSIDA AS195 Asclepias stenophylla A. Gray-P; MGP; WASO5O Amaranthaceae Asclepias syriaca L.-P; MGP; WAS280 Amaranthus albus L.A; MGP;WAS322 Funastrum cynanchoides (Dcne.) Schlechter-P; Amaranthus palmeri S.Wats.—A; DAOF; WASO093 : 100 Froelichia gracilis (Hook.) Moq.-A; SW; WAS103 Asteraceae Anacardiaceae Ambrosia psilostachya DC.-P; MGP; WAS148 Rhus aromatica L.—P; MGP; WAS230 Ambrosia trifida L—A:M 95 Rhus glabra LP; MGP; WAS044 Amphiachyris dracunculoides (DC.) Nutt-A;MGP; Toxicodendron radicans (L.) Kuntze-P; RA; WAS283 Aphanostephus skirrhobasis (DC.) Trel.—A; MGP; Apiaceae WAS323 Chaerophy. Cymopterus macrorhizus Buckl—P;MGP;WAS171 lum tainturieri Hook. —A:MGP:WAS222 Artemisia dracunculus L-P; MGP; WAS293 Artemisia filifolia Torr.—P; MGP; WAS116 as 1192 Artemisia ludoviciana Nutt.-P; MGP; WAS289 Baccharis salicina Torr. & A. Gray—P; RA;WAS118 Brickellia eupatorioides (L.) Shinners—P; MGP; AS294 Chaetopappa ericoides (Torr.) Nesom—P; MGP; O15 Cirsium undulatum (Nutt.) Spreng.—P; MGP; WAS248 Cirsium vulgare (Savi) Ten.-B; MGP;WAS247 Conyza canadensis (L.) Crong.-A; DAOF; WAS 141 Eclipta prostrata (L.) L.-A; RA; WAS133 Engelmannia peristenia (Raf.) Goodman & Law: a 4 Erigeron bellidiastrum Nutt.-A; MGP. SW;WAS188 Eupatorium serotinum Michx.-P; RA; WAS129 Euthamia gymnospermoides Greene-P; DAOF; WAS314 Gaillardia pulchella eee MGP; WAS228 so suavis (A. Gray & Engelm.) Britt. & usby—P; MGP; WAS229 es papposa Nesom & Suh-A; MGP; WAS153 liantt 5 L-A; DAOF; WASO8 Helianthus maximiliani Schrad.—P; cee Helianthus petiolaris Nutt.-A; DAOF; WASO84 Heterotheca subaxillaris (Lam.) Britt. & Rusby-A; WAS 144 Heterotheca villosa (Pursh) Shinners—P; MGP; WASO28 Hymenopappus flavescens A. Gray-B; MGP; WAS258 /va annua L.-P; DAOF, RA; WAS317 Lactuca serriola L.*~A; MGP; WASO22 Liatris pycnostachya Michx.-P; MGP; WAS282 ygodesmia juncea (Pursh) D. Don ex Hook.-P; R; M AS285 Machaeranthera pinnatifida (Hook.) Shinners—P; MGP; WAS2 Pluchea odorata (L.) Cass. var. odorata—A; RA: WAS108 Ratibida columnifera (Nutt.) Woot. & Standl.—P MGP; $276 ’ densis L—P; DAOF;WAS113 Sélidago gigantea Ait.-P; DAOF; Len Solidago petiolaris Ait.—P; MGP; WAS30 Symphyotrichum ericoides (L.) Nesom— is DAOF; Symphyotrichum oblongifolium (Nutt.) Nesom— P; MGP;WAS303 BRIT.ORG/SIDA 21(2) —_ Symphyotrichum subulatum (Michx.) Nesom-A; RA; WAS 132 Taraxacum officinale G.H.Weber ex Wiggers*-P; DAOF; WAS302 Tetraneuris scaposa (DC.) Greene-P; MGP; WAS226 Thelesperma megapotamicum (Spreng.) Kuntze- P; MGP;WASO51 AON Aihyipie *_ A: DAOF; WAS182 ebesna encatoiaes ca Benth. & Hook. f.ex A. Gray—A; DAOF; esis baldwinii Torr.— : ome 105 m strumarium L—A; RA; WAS135 se ia en ie Nutt.—P; MGP; WAS271 Boraginaceae Helioitropium convolvulaceum (Nutt.) A. Gray-A; SW; WASO95 Lithospermum incisum Lehm.—P; MGP; WAS172 Brassicacea Camelina Cd ica Velen.-A; MGP;WAS231 Capsella bursa-pastoris (L.) Medik.*-A; DAOF; 6 Descurainia pinnata (Walt.) Britt.-A; MGP; AS177 Dimorphocarpa candicans (Raf.) Rollins—A; SW; eva oe (Lam.) Fern.-A; MGP; WAS16 | 7 Small-A; MGP; WAS] vA Lesquerel gordon (A. Gray) S. Wats.-A; MGP; Cactaceae ae vivipara (Nutt.) Buxbaum-—P; MGP; WAS315 Opuntia macrorhiza Engelm.-P; MGP; WAS159 iia abate aii holzingeri McCVaugh-A; MGP; WAS 266 Trindanic aryophylla pentane ia L.*-A; DAOF; WAS163 Paronychia jamesii Torr. & A. Gray-P; MGP; WASO55 Stellaria media (L.) Vill*—A; DAOF; WAS 174 Senopodiaceae Ch L.*-A; MGP; WAS287 henenod) um simplex (Torr.) Raf.-A; MGP; WAS150 Cycloloma atriplicifolium (Spreng.) Coult.-A; MGP; WAS264 Kochia scoparia (L.) Schrad.*-A; MGP; WASO09 HOAGLAND ET AL., FLORA OF HISTORICAL SITE 1193 Sonvelvulaceae Col is L.*-P: MGP; WAS 196 Evolvalg nuttallianus J. A. Schultes—P; MGP; WAS215 [pomoea leptophylla Torr.—P; MGP; WAS260 Cucurbita Cucurbita pee Kunth-P; MGP; WASO18 Cyclanthera dissecta (Torr.& A. Gray) Arn.—A; SW; WAS140 Euphorbiaceae aan esol aaa A; MGP; WAS031 is (Engelm. & A. Gray) Muell.- oC —P; MGP; WASO mies ie eri ae & A. Gray) Small—P; — see (Engelm.) Small-A; MGP, SW; WASO90 Chamaesyce maculata (L.) Small-A; DAOF; WAS122 Chamaesyce missurica (Raf.) Shinners—A; MGP, DAOF;WAS304 Chamaesyce stictospora (Engelm.) Small—-A; DAOF; WAS069 Croton glandulosus L.—A; MGP;WASO37 Croton texensis (Klotzsch) Muell.-Arg.-A; MGP; O11 phorbia dentata Michx.-A; MGP; WASO1 2 Euphorbia ‘eae Nutt. ex Spreng.—A; MGP; eis ongicruris Scheele—A; MGP; WAS 160 Euphorbia marginata Pursh—A; DAOF; WAS 142 Tragia ramosa Torr.—P; MGP;WAS058 Fabace Amerpha ittcoa L—P; RA; WASO86 tiflorus Hook.-P; MGP; WAS180 oe Nutt.-P; MGP;WAS181 Baptisia australis (L.) R. Br. ex Ait. f-P; MGP; WAS191 Actraqgalise« Caesa W, >> pinia jamesii (Torr.& A. Gray) Fisher—P; SW; $102 Cercis canadensis L.-P; DAOF; WAS065 Chamaecrista fasciculata (Michx.) Greene-A; GP: WAS Dalea aurea Nutt. ex Pursh-P; MGP; WAS274 Dalea candida Michx. ex Willd.-P; MGP; WAS267 Dalea enneandra Nutt.-P; MGP; WASO57 Dalea purpurea Vent.-P; MGP; WAS250 Desmodi um illinoense A. Gray—P; MGP; WASO032 W;WAS300 Gleditsia triacanthos Li- P:S oi ladus dioicus (L.) K. Koch—P; SW; WASO16 jata Ortega—P; SW; WASO91 Medicago minima (L.) L.*-A; DAOF; WAS224 eliotue offici nalis (L.) Lam.*-A; DAOF; WAS246 imo A. Gray—P; MGP; WAS199 Mimosa eat (DC.) B.L. Turner-P; MGP; WAS201 Pedi jomelum linearifolium (Torr. & A. Gray) J es—P; MGP; WAS048 Pees nuttalliana B.L.Turner—P; MGP;WAS243 Strophostyles leiosperma (Torr. & A. Gray) Piper- A; MGP; WAS042 Fumariaceae Corydalis micrantha (Engelm.ex A.Gray) A.Gray— A; MGP;WAS178 Geraniaceae Frodium cicutarium (L.) LU’Her. ex Ait.*-A; DAOF; WAS169 Geranium pusillum L.*-A; MGP; WAS218 Grossulariaceae Ribes aureum Pursh—P; SW;WAS167 Juglandaceae Jugla Berl.—P; SW; WAS101 Krameri Krameria eee Torr.—P; MGP; WAS039 Lanuacese ium amplexicaule L.*-A; DAOF; WAS168 Lycopus americanus Muhl. ex W. Bart.—P; RA; WAS 128 ees clinopodioides A.Gray—A; MGP;WAS25 1 onarda punctata L—A; MGP;WAS254 ae azurea Michx. ex Lam.—P; MGP;WAS301 Scutellaria resinosa Torr—P; MGP; WASO40 Scutellaria wrightii i A. Gray —P: MGP;WAS214 im canadense L.—P; RA; WASO75 Teurcium la ‘icin Tons P: MGP;WAS221 Linaceae Linum pratense (J.B.S. Norton) Small—A; MGP; WAS? 12 Linum rigidum Pursh—A; MGP; WAS204 Loasaceae Mentzelia nuda (Pursh) Torr. & A. Gray-P; MGP; Lythraceae A Rottb.—A; RA; WAS143 1194 Malvaceae Callirhoe involucrata (Torr. & A. Gray) A. Gray-P; MGP; WAS279 Hibiscus trionum L.*—P; MGP; WAS269 Sphaeralcea coccinea (Nutt.) Rydb.-P; MGP; WAS219 Menispermaceae Cocculus carolinus (L.) DC.-P; SW; WASO78 Hira e erticillata L—A; DAOF, SW; WAS024 IVIQTT Moraceae Morus alba L.*-P; DAOF; WAS061 Nyctaginaceae Mirabilis albida (Walt.) Heimer|—P; MGP: WAS 298 Mirabilis linearis (Pursh) Heimerl|—-P; MGP;WASO30 Mirabilis nyctaginea (Michx.) MacM.—P; MGP, SW; WAS189 Oleaceae Forestiera pubescens Nutt.-P; SW; WAS036 ¢ herlandier al ae lylopht i Spach—P; MGP; WAS207 ae Rae hartwegii (Benth.) Raven—P; MGP; WAS237 Calylophus serrulatus (Nutt.) Raven—P; MGP; WASO049 Gaura brachycarpa Small—A; MGP; WAS202 Gaura parvi iflora Dougl. ex Lehm.-A; MGP Oenothera jamesii Torr.& A. Gray—P; RA; WAS125 Qenothera laciniata Hill-P; MGP; WAS206 Oenothera rhombipetala Nutt. ex Torr.& A. Gray— P; MGP; WAS265 Oxalidaceae Oxalis stricta L.-P; SW; WAS110 Papaveraceae Argemone polyanthemos (Fedde) G.B. Ownbey- A; SW; Pedaliaceae Proboscidea louisianica (P. Mill.) Thellung—A; MGP; Plantaginaceae Plantago patagonica Jacg.-A; MGP; WAS 197 P. rhodosperma Dcne.-A; MGP; WAS225 Polygonaceae Eriogonum annuum Nutt.-A; DAOF, MGP; 0) BRIT.ORG/SIDA 21(2) folium Nutt.-P; MGP; WAS054 L.*-A; MGP; WASO14 Polygonum lapathifolium L.—A; RA; WAS 136 Rumex crispus L.*-P; MGP; WAS185 Por] ; | i Y Portulacaceae Portulaca oleracea .-A; OF; WAS066 Primulaceae Androsace occidentalis Pursh-A; MGP; WAS 162 Ranunculaceae Delphinium carolinianum Walt. ssp. virescens (Nutt.) Brooks—P; MGP; WAS240 Rosaceae Prunus angustifolia Mars.—P; SW; WAS291 Rubiaceae Cephalanthus occidentalis L.—P; RA; WAS 106 Galium pilosum Ait.-P; DAOF; WAS089 edyotis nigricans (Lam.) Fosberq—P; MGP; $273 Salicaceae Populus deltoides Bartr.ex Marsh.—P:RA;WAS117 Salix exigua Nutt.-P; RA; WAS111 Salix nigra Marsh.-P; RA; WASO85 Sapindaceae Sapindus saponaria L.—P; SW; WASO70 Sapotaceae Sideroxylon lanuginosum Michx.-P; SW; WAS046 Scrophulariaceae a purpurea (Nutt.) G. Don var. citrina (Pennell) Shinners—P; MGP; WAS23? Penstemon albidus Nutt.-P; MGP;WAS213 Veronica arvensis L.*-A; DAOF; WAS165 Solanaceae Chamaesaracha conioides (Moric. ex Dunal) Britt. —P; MGP:; WAS238 Physalis cinerascens (Dunal) A.S. Hitchc.—P; MGP; WAS205 Ele Phy Nutt.-P; MGP; WASO19 as sri ie or Par P: DAOF; WAS203 P; DAOF: WAS194 Solanum ead Cav.-P; DAOF, MGP; WAS 234 Solanum rostratum Dunal-A; DAOF, MGP; atic | ifr |i a WASO25 Solanum triflorum Nutt.-A; MGP; WAS029 HOAGLAND ET AL., HISTORICAL SITE 1195 Tamaricaceae Tamarix ramosissima Ledeb.*—P: RA; WAS115 Ulmaceae Celtis atc. Willd. var. reticulata (Torr.) L.Ben- on-P; SW; WASO04 Ten pumila ic *_P; ae WAS 268 Ulmus rubra Muhl.—P; SW; WAS154 Urticaceae Parietaria pensylvanica Muhl. ex Willd.-A; DAOF, Verbenac Glandularia pumila P;WAS198 Phyla lanceolata (Michx.) Greene-P; RA; WASO096 (Rydb.) Umber-A; DAOF, Violaceae Hybanthus verticillatus (Ortega) Baill.—P; MGP; WAS242 VICAEESE. data Michx.—P; RA; WAS12 Coss incisa auct. non Des Moulins— 3 SW; WASO Vitis bac Raf—P; RA; WAS109 Zygophyllaceae Kallstroemia parviflora J.B.S. Norton—A; DAOF; WASO004 Tribulus terrestris L.*-A; DAOF; WAS308 LILIOPSIDA Agavaceae Yucca glauca Nutt.-P; MGP; WAS200 Commelinaceae Commelina erecta L—P; MGP;WAS052 Tradescantia occidentalis (Britt.) Symth—P; MGP; Cyperaceae Carex gravida Bailey—P; MGP; WAS241 Cyperus odoratus L.-A; MGP; WAS145 Cyperus schweinitzii Torr—P; MGP; WASO23 Cyperus setigerus Torr. & Hook.—P; RA; WAS126 Fimbristylis vahlii (Lam.) Link-A; RA; WAS137 Schoenoplectus pungens (Vahl) Pall-P; RA; WASO77 lridaceae Sisyrinchium angustifolium P. Mill.—P; MGP; WAS236 Juncaceae Juncus torreyi Coville-P; RA; WAS083 prise im canadense L.-P; MGP; WAS227 Allium drummondii Regel-P; MGP; WAS173 Poaceae Andropogon hallii Hack.—P; MGP, SW; WASO73 Aristida oligantha Michx.-A; MGP;WAS325 Aristida purpurea Nutt—P; MGP; WAS053 Bothriochloa ischaemum (L.) Keng*—P; MGP; WAS299 Bothriochloa laguroides (DC.) Herter—P; MGP; WAS257 Bouteloua curtipendula (Michx.) Torr.—P; MGP; WASO020 Bouteloua gracilis (Willd. ex Kunth) Lag. ex Griffiths—P; DAOF, MGP; WAS272 Bouteloua hirsuta Lag.-P; MGP; WASO05 Bromus japonicus Thunb.ex Murr.*-A; DAOF, SW; 239 Buchloe dactyloides (Nutt.) Engelm.—P; DAOF, Cenchrus spinifex Cav.—P; SW; WAS097 cele PuatY Nutt.-P; mie MGP;WAS263 Cyn n (L.) Pers.*-P; DAOF; WAS255 ium ceo (Nash) Gould-P; WAS261 = ty pee ie oligosanthes (J.A. Schultes) Gould—P; MGP; WAS253 Digitaria ciliaris (Retz.) Koel—P; DAOF; WASOO1 Digitaria cognata (J.A. Schultes) Pilger—P; MGP; WAS256 —= Distichlis spicata (L.) Greene-P; DAOF; WAS062 Echinochloa crus- gay (L.) Beauv.*—A; RA;WAS127 Elymus canadensis L.—P; MGP; WAS297 Eymus virginicus LP; RA; WASO76 Eragrostis barrelieri Daveau*-A; DAOF; WAS307 Eragrostis cilianensis (All.) Vign. ex Janchen*—A; OF; WASO07 Eragrostis curvula (Schrad.) Nees*-P; MGP; Eragrostis spectabilis (Pursh) Steud.-P; SW; WASO99 Erioneuron pilosum (Buckl.) Nash-P; MGP; WAS318 Hordeum pusillum Nutt—A; DAOF; WAS208 Leptochloa fusca (L.) Kunth ssp. fasicularis (Lam.) now-—A; RA; WAS134 ium perenne L.*=P; MGP; WAS252 Q 1196 Muhlenbergia asperifolia (Nees & Meyen ex Trin.) Parodi-P; SW; WAS158 Muhlenbergia racemosa (Michx.) B.S.P—P; MGP; AS152 Muhlenbergia sobolifera (Muhl.ex Willd.) Trin.-P; MGP;WAS151 Panic pillare L.-A; MGP; WASO26 Panicum obt Kunth—P; MGP, RA; WAS124 LP; MGP;WAS319 +d ‘ . PULNCUTTT VIFGQALUTTT Pascopyrum smithit (Rydb.) A. Love-P; MGP; WAS064 Paspalum setaceum Michx.-P; DAOF; WASO8 1 Poa arachnifera Torr—P; RA; WAS244 ae giganteum (Walt.) Pers.—P; RA; WAS14 sheen paniculatus (Nutt.) Trel-P; DAOF; BRIT.ORG/SIDA 21(2) Setaria parviflora (Poir.) Kerguelen—P; DAOF; WAS114 oe idis G .) Beauv.*-A; MGP; WAS02 1 ghastrum nutans (L.) Nash—-P; MGP; WAS130 Sorghum h ies nse (L.) Pers.*-P; DAOF; WAS270 eat na pectinata Bosc ex Link-P; RA; WASO74 Sporobolus cryptandrus (Torr) A.Gray—P; MGP, SW; WASO79 Spor ao0! US Gi paniens Nash—P; MGP; WAS321 Sporobolus Va aginiflorus (Torr rex A, Gray) Wood- Tridens flavus (L.) A.S. Hitchc.—P; DAOF; WAS119 eee purpurea (Walt.) Chapman—-A; MGP; WAS288 Triticum aestivum L.*-A; DAOF; WAS Vulpia octoflora (Walt.) Rydb.-A; ve on WAS063 Schizachyrium scoparium (Michx.) Nash—P; MGP; WAS156 ACKNOWLEDGMENTS This project was funded by a grant from the National Park Service. We thank Adam K. Ryburn and anonymous review for their helpful comments on an ear- lier draft. REFERENCES Branson C.C.and K.S. JOHNSON. 1979. Generalized logic map of Oklahoma. |n:K.S.Johnson, C.C. Branson, N.M. Curtis, W.E. Ham, W.E. Harrison, M.V. Marcher, and J.F. Roberts, eds. Geology and earth resources of Oklahoma.Oklahoma Geological Survey, Norman. P.4. Burcess, D.L., J.D. NicHots, and O.G. Henson. 1959. Soil survey of Love County, Oklahoma. United States Department of Agriculture, Washington D.C. Correll, D. S. and M.C. Jounston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner Curtis N.M. and W.E. Ham. 1979. Geomorphic provinces of Oklahoma. In: K.S. Johnson, C.C. Branson,N.M.Curtis, W.E.Ham,W.E. Harrison, M.V. Marcher, and J.F. Roberts, eds. Geology and earth resources of Oklahoma. Oklahoma Geological Survey, Norman. P.45. Duck L.G. and J.B. FuercHer. 1943.A game type map of Oklahoma. Oklahoma Department of Wildlife Conservation, Oklahoma City. Freeman, C.C.,C.A. Morse, and J.P. THurmonp. 2003. The vascular flora of the Ogallala eco- tone on the Dempsey Divide, Roger Mills County, Oklahoma. Sida 20:1217-1245. Great Plains Flora Association. 1986. Flora of the Great Plains. University Press of Kansas, Lawrence. Groves C.R., M.L. Kcein, and T.F. Breoen. 1995. Natural heritage programs: public-private part- nerships for biodiversity conservation. Wildlife Soc. Bull. 23:784-790. Goutp, FW. 1975.The grasses of Texas. Texas A&M University Press, College Station. HOAGLAND ET AL., FLORA OF HISTORICAL SITE 1197 Hoactano BW. 2000. The vegetation of Oklahoma: a classification of landscape mapping and conservation planning. Southwest Nat. 45:385-420. Hoaclanb, BW. 2004. Atlas of the flora of Oklahoma [online]. Available: www.biosurvey.ou.edu. (Accessed on 14 January 2004). Hoactano B.W. and A. ButHop. 2003. Vascular flora of the Keystone Wildlife Management Area, Creek, Pawnee, and Osage counties, Oklahoma. Oklahoma Native Pl. Record 3: 23-37. Hoaatanp B.W.and F.L. Jonnson.In press. The vascular flora of Red Slough and Grassy Slough Wildlife Management Areas, Gulf Coastal Plain, McCurtain County, Oklahoma. Casta- nea 69. Hoactanbd B.W. and F.L. JoHNson. 2001.Vascular flora of the Chickasaw National Recreation Area, Murray County, Oklahoma. Castanea 66:383—400. Hoactanp B.W. and K. Wattick. 2003. Vascular flora of Oologah Wildlife Management Area, Nowata County, Oklahoma. Proceedings of the Oklahoma Academy of Science 83: 47-62. HoacLaNo B.W., P.CRawrorD-CALLAHAN, P.CRawForb, and F.L. JOHNSON. 2004. Vascular flora of Hack- berry Flat, Frederick Lake, and Suttle Creek, Tillman County, Oklahoma. Sida 21: 429-445, Hunt C.B. 1974. Natural Regions of the United States and Canada. W.H. Freeman, San Francisco. OKLAHOMA CLIMATOLOGICAL SuRveY. 2004. Oklahoma Climatological Data [online]. Available: www.ocs.ou.edu/. (Accessed on 1 March 2004). OktAaHoma NaturAt HeriTace INveNToRY. 2004. ONHI working list of rare Oklahoma plants [on- line]. Available: www. biosurvey.ou.edu/publicat.html. (Accessed on 1 March 2004). Paumer M.W., G.L. Wave, and P. Neat. 1995. Standards for the writing of floras. Bioscience 45:339-345, Taytor RJ.and CS. Tayior. 1991. An annotated list of the ferns, fern allies, gymnosperms, and flowering plants of Oklahoma. Southeastern Oklahoma State University, Durant. TrewartHa G.T. 1968. An introduction to Climate. McGraw-Hill, New York. USDA-NRCS 2004. The PLANTS database [online]. Available: plants.usda.gov/plants. Na- tional Plant Data Center, Baton Rouge, LA. (Accessed on 14 January 2004). WarerrAatt, U.T. 1969. Keys to the flora of Oklahoma. 4th edition. Published by the author, New York. ALTERNANTHERA PARONICHYOIDES (AMARANTHACEAE) AND RUMEX MARITIMUS (POLYGONACEAE) NEW TO OKLAHOMA Bruce W. Hoagland Amy Buthod Oklahoma Biological Survey Oklahoma Biological Survey and Department of Geography University of Oklahoma University of Oklahoma Norman, Oklahoma 73019, U.S.A. Norman, Oklahoma 73019, U.S. Oe ee fac ABSTRACT This paper reports the occurrence of two species previously unknown to the flora of Oklahoma. Alternanthera paronichyoides St. Hil. which occurs in Texas, was discovered in Pushmataha County in southeastern Oklahoma. Rumex maritimus L., a species found in all states adjoining Oklahoma was collected in Washita County in western Oklahoma. RESUMEN Este articulo cita la ocurrencia de d iz descouesicn de la lore de Oklahoma. Alternanthera paronichyoidesSt. Hil. ee fue d le Pushmataha enel Sureste de Oklahoma. Rumex maritimus L., una ncuentra en todos los estados adyacentes de Oklahoma, se colecté en el condado de shite en el Oeste de Oklahoma. Alternanthera paronichyoides St. Hil(Amaranthaceae) is a perennial plant spe- cies native to the old world tropics. It has been reported from Texas and several Gulf Coastal Plains and Atlantic Seaboard states (USDA-NRCS 2004). The popu- lation reported below was robust and growing in a ditch which drains into the Kiamichi River in southeastern Oklahoma. Associated species included Arundinaria gigantea (Walter) Muhl.,, Eleocharis obtusa (Willd.) Schult, and Symphyotrichum subulatum (Michx.) Nesom. Voucher specimen: OKLAHOMA. Pushmataha Co.: 4.0 mi W of Rattan on Hwy. 3 at Rattan Landing on the Kiamichi River, T4S, RI7E, sec 14, 5 May 2001 Hoagland and But hod hugo509 (OKL) Rumex maritimus L. (Polygonaceae) is a native annual species that has been reported from all states adjoining Oklahoma (USDA-NRCS 2004). It was grow- ing in the drawdown zone of a municipal reservoir. Associated species included Ammannia coccinea Rottb., Cyperus setigerus Torr. @ Hook., C. squarrosus L., Echinochloa crus-galli (L.) Beauv., Leptochloa fusca (L.) Kunth ssp. fascicularis (Lam.) N. Snow, Symphyotrichum subulatum (Michx.) Nesom. Voucher specimen: OKLAHOMA. Washita Co.: Clinton Lake, ca. 4.4 mi NE of Canute, shoreline habi- tat, TIN, R1IOW sec. 16, 23 Aug 2002, Hoagland and Buthod AB-2867 (OKL). SIDA 21(2): 1199-1200. 2004 1200 BRIT.ORG/SIDA 21(2) ACKNOWLEDGMENTS We appreciate the comments of Adam K. Ryburn and an anonymous reviewer. REFERENCE USDA-NRCS 2004. The PLANTS database [online]. Available: plants.usda.gov/plants. Na- tional Plant Data Center, Baton Rouge, LA. (Accessed on 1 March 2004). SCHOENOPLECTUS HALLII (CYPERACEAE), A GLOBALLY THREATENED SPECIES NEW FOR TEXAS Robert J.O’Kennon Caren McLemore Botanical Research ol of Texas Botanical Research Institute of Texas and 509 Pecan Stre University of North Fort Worth, TX 76102, a A. Denton, Texas 76203, a okennon@brit.org clm0093@unt.edu ABSTRACT Schoenoplectus hallii (A. Gray) S. Galen Smith, Hall’s bulrush, has been discovered in the Lyndon B. Iehncon N anona) Grasslands, Wise County, Texas. Previous Epo ts of this Species in Texas were based I PI record for the st ale. RESUMEN Schoenoplectus hallii (A. Gray) S. Galen Smith, ha sido descubierto en los Lyndon B. Johnson Na- tional Grasslands, Wise County, Texas. Las citas Previas de esta especie en Texas estaban basadas en especimenes mal identificados, y la presente cita es aparentemente la primera para el estado. Schoenoplectus hallii (A. Gray) S. Galen Smith, Hall’s bulrush, has been discov- ered during an extensive floristic survey of the Lyndon B. Johnson National Grasslands (LBJGL), 15.6 km north of Decatur, Wise County, Texas. The LBJGL are managed by the U.S. National Forest Service and comprise numbered “units” scattered throughout much of north-central Wise County. Schoenoplectus hallii grows in Unit 66 primarily along the moist sandy-clay margins of three small ponds that merge during periods of high water. The populations are approxi- mately 100 meters apart and comprise about two hundred individuals each. Specimens of Schoenoplectus hallii were collected in May 2003 through April 2004 and their identity was confirmed by Galen Smith of the University of Wisconsin. Earlier reports of the species in Texas (Correll & Johnston 1970, Hatch et al. 1990) and various herbarium none a (later correctly annotated) were based on misidentifications of S.sax inus (Fern.) Raynaland S. erectus (Poir) Palla ex Raynal (Schuyler 1969, Smith 1995). Because S. hallii had not been confirmed for Texas, it was not included in the recently published Illustrated Flora of North Central Texas (Diggs et al. 1999). Smith (2002) reported the oc- currence of the species in Georgia, Illinois, Indiana, Kansas, Kentucky, Massa- chusetts, Michigan, Missouri, Oklahoma, and Wisconsin, Schoenoplectus hallii (A. Gray) S. Galen Smith, Novon 5:101. 1995. Scirpus hallii A. Gray; Scirpus supinus L. var. hallti (A. Gray) A. Gray Voucher specimens. TEXAS. Wise Co.: Along margin of pond in Unit 66 of the Lyndon B. Johnson National Grasslands, N33° 22' 12" -woo7e 32! 28", 17 May 2003, O’Kennon and McLemore 18344 (BRIT, SIDA 21(2):1201-1204. 2004 1202 BRIT.ORG/SIDA 21(2) WIS), Along margin of pond in Unit 66 of the Lyndon B. Johnson National Grasslands, N33° 22' 12", W097° 32° 28", 23 August 2003, O’Kennon and McLemore, 18853(BRIT); Along margin of pond in Unit 66 of the Lyndon B. Johnson National Grasslands, N33° 22' 15", W097° 32' 26", 29 August 2003, O’Kennon and McLemore, 18892 (BRIT, WIS), Along margin of pond in Unit 66 of the Lyndon B Johnson National Grasslands, N33° 22' 12", WO97° 32' 27", 3 Oct 2003, O’Kennon and McLemore, 19070 and ee (BRIT) Along margin of pond in Unit 66 of the Lyndon B. Johnson National Grasslands, 2'12", W097? 32' 28", | Nov 2003, O’Kennon and McLemore “Oidtand 19142 (BRIT), seer pants at bottom of pond in Unit 66 of the Lyndon B. Johnson National Grasslands, N33° 22'1 19 Dec 2003, O’Kennon and McLemore 19180 (BRIT); oe Sao al eee s pone! in ae ve of the Lyndon B. Johnson National Grasslands, N33° 22 O’Kennon and McLemore 19191 (BRIT); Submerged plants at bottom 7 pond in Se ol “A ai B. Johnson National Grasslands, N33° 22' 15", W097? 32' 26", 28 Feb 2004, O’Kennon and McLemore 19197 (BRIT), Submerged plants at bottom of pond in Unit 66 of the Lyndon B. Johnson National Grasslands, N33° 22' 15", W097° 32' 26", 17 Mar 2004, O’Kennon and McLemore ieee (BRIT); Emer- gent plants at bottom of pond in Unit 66 of the Lyndon B. Johnson Nation ands, pees a = ; W097° 32' 26", 9 Apr 2004, O’Kennon and McLemore ue banca Mus i otha submerged) in Unit 66 of the Lyndon B. Johnson Nati 3°22 1 woor" 32 26. I Apr 2004, O’Kennon and McLemore 19279 (BRIT): Madey: | Sept. 2003) in Unit 66 of the Lyndon B. Johnson National erin N33° 22' 15", WO97° 32 26", 21 Apr 2004, O’Kennon and McLemore 19290 (BRIT); Beets: along margin of pond in Unit 66 of the Lyndon B. Johnson National Grasslands, N33° 22' 15", W097? 32' 26", 21 Apr 2004, O’Kennon and McLemore 1929] (BRIT). Duplicates to be distibucd Identification of the Texas plants.—The following key to species of Schoenoplectus sect. Supiniin Texas is modified from Smith (2002). Schoenoplectus hallii found elsewhere are usually annual, but in Texas many perennial plants have been observed. 1. Achenes in spikelet ly equilaterally sharply tri all 3-fid; from pan- handle to central and far south Texas Schoenoplectus saximontanus . Achenes in spikelets penta to plano-convex or obscurely compressed- trigonous; styles 2-fid or a ia -fid. , ; = chenes Ikebe 1] | It \orizontally JY un) vex center; scikalet aay eae anh distally orange- or nee nia in Tes known only from south coastal counties inland to Atascosa Co. Schoenoplectus 2. Ache dl ae) rsometimes nearly plane; spikelet scale flanks pale orange to nee ie hacun from one site in north central Texas __ Schoenoplectus hallii Habitat of the Texas plants.—Schoenoplectus hallii has a narrow habitat toler- ance (Beatty et al. 2004). Al though itis found in various soil types in other states, in Texas it is found along widely fluctuating margins of small sandy clay ponds r tions apparently are more suitable for S. hallii than for many of its associates (Schuyler 1969). Many plant species occur with S. hallii in Texas (Table 1). These associates appear and disappear throughout the year as water levels fluctuate, but S. hallii can be found along pond margins from April to Decem- ber. It is most closely associated nearly year-round with the rare Pilularia americana (Marsiliaceae). During periods of normal water levels P americana 1203 Tasle 1. Species associated with Schoenoplectus hallii in Texas, ranked by abundance. Pilulari icana A.Braun Eleocharis obtusa (Willd.) J.A. Schultes Rotala ramosior (L.) Koehne Cyperus acuminatus Torr. & Hook. ex Torr. Fimbristylis vahiti (Lam.) L Ludwigia peploides nae Raven \mmannia inea Rottb. Eleocharis palustris (L.) Roem. & Schult. Fuirena simplex Vahl Eleocharis quadrangulata (Michx.) Roem. Lindernia dubia (L.) Pennell var. anagallidea (Michx.) Cooperrider Eleocharis coloradoensis (Britt.) Gilly Callitriche heterophylla Pursh Ammannia robusta Heer & Regel Eleocharis engelmannii Steud Najas guadalupensis (Spreng.) Magnus Veronica peregrina L.var.xalapensis (Kunth) Pennell Fuawigie ia glandulosa Walter ranthera limosa (Sw.) Willd. Scalia is GtOpUIUicd (Retz.) J.& K. Presl Raf. ee onde anion var. cylindricus (Ell.) Fern. & Grisc. Cyperus squarrosus L. Panicum rigidulum Nees Juncus texanus (Engelm.) Coville Echinodorus berteroi (Spreng.) Fassett /) idtallisataiadiaaln Buckl Spirodela polyrhiza (L.) Scheid. Limnosciadium pinnatum (DC.) Mathias & Constance Anagallis minima (L.) Krause Marsilea vestita Hook. & Grev is a submergent and can be conspicuous just below the surface around the en- tire margin of the pond. However, during periods of extended drawdown it be- comes a marginal plant. The ponds occur in the West Cross Timbers of north central Texas at an elevation of 254 meters. During the first year of our LBJGL survey, twelve other species undocu- mented for Texas, in addition to S. hallii, were discovered (O’Kennon et al. 2003; O’Kennon & McLemore in prep.). We believe that this probably reflects previ- ous undercollecting rather than particularly unique habitats in this area. Overall distribution.—Schoenoplectus hallii is a globally threatened spe- cies (Nature Conservancy ranking of G-2) known recently from only 8 states and fewer than 100 sites in the U'S.A. Texas Parks and Wildlife Department has assigned a ranking of S-1 (critically imperiled) for the species Jackie Poole, pers. comm.), based on the discovery in Wise County. The closest populations of S. hallii to the Wise Co. populations are in Comanche Co., Oklahoma, about 190 kilometers northwest. Prior to 1973, Schoenoplectus hallii was reported in 9 states, 15 counties, and 29 sites. During 1973-1997, it was reported in 8 states, 17 counties, and 46 sites. During 1993-1997, it was reported in only 6 states, 11 counties and 37 sites (McKenzie 1998). Since not all sites are being monitored each year, it is difficult to ascertain the actual number of existing populations. Plants in Massachusetts have not been seen since 1931 and are considered extirpated. Schoenoplectus hallii has not been collected in Georgia since 1966. It has not been confirmed in lowa since 1890. There is a record from lowa from 1960, but that specimen has not been seen nor confirmed (McKenzie 1998). Popu- lations in these states are considered “possibly extirpated.” 1204 BRIT.ORG/SIDA 21(2) ACKNOWLEDGMENTS We thank Guy Nesom of BRIT for assistance in identifying the original speci- mens and reviewing the manuscript. We gratefully acknowledge Galen Smith of the University of Wisconsin for assistance in confirming the identity of this plant. We thank James Crooks, Jimmy Dickerson, and Alfred Sanchez of the Lyndon B. Johnson National Grasslands and the U.S. Forest Service for assistance and coop- eration in our work on this threatened species and for preserving the site. REFERENCES Beatty, L.B., W.F. Jenninas, and R.C. Rawtinson. 2004. Schoenoplectus hallii (Hall's bulrush): a technical conservation assesment. USDA, Forest Service, Rocky Mountain Region, Spe- cies Conservation Project. Denver, Colorado. Corrett, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner, Texas. Diccs, G.M. Jr., B.L. Liescome, and R.J. O’Kennon. 1999, Shinners & Mahler's illustrated flora of North Central Texas. Sida, Bot. Misc. 16. Hatch, S.L., K.N.GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Misc. Publ. No. 1655, McKenzie, PM. 1998. Hall's bulrush (Schoenoplectus hallii) status assessment. U.S. Fish and Wildlife Service. Columbia, Missouri. O’Kennon, RJ.,C. McLemore, and A. Nei. 2003. Fagopyrum esculentum, (Polygonaceae), new for Texas. Sida 20:1717-1720. ScHuYLer, A.E. 1969. Three new species of Scirpus (Cyperaceae) in the southern United States. Notulae Naturae 423:1-12. Smith, S.G. 1995. New combinations in North American Schoenoplectus, Bulboshoenus, Isolepis and Trichophorum (Cyperaceae). Novon 5:97-102. Smith, S.G. 2002. Schoenoplectus sect. Supini, In:Vol.23 (Cyperaceae) Flora of North America North of Mexico, Flora of North America Association, Oxford Univ. Press, N.Y. Pp. 58-60. NEW RECORDS IN PSEUDOGNAPHALIUM (ASTERACEAE: GNAPHALIEAE) FROM NEBRASKA AND CALIFORNIA Steven B. Rolfsmeier Guy L.Nesom High Plains Herbarium Botanical Research Institute of Texas Chadron State College 509 Pecan Street Chadron, Nebraska 69337, U.S.A. Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Pseudognaphalium jaliscense is first reported from Nebraska, documented here from twelve coun- ties; an earlier report of this species from California was based on collections of a form of P. stramineum. Pseudognaphalium roseum is first documented as occurring in the U.S.A., where it has been collected in southern California. RESUMEN Pse udoenae eee Jamsconses se cita igs puea vez oe NepEa se documents aqui de doce =] ] Ulla for Hida ae P. stramineum. Pse ae na yhalium roseum se documenta por primera vez de U.S.A., donde ha sido 8 ae colectado en el Sur de Califo In connection with taxonomic studies of Gnaphalieae, preparation of treat- ments for the Flora of North America volumes of Asteraceae, and floristic stud- ies in Nebraska, we report new distribution records for two species of Pseudognaphalium Kirp. Pseudognaphalium jaliscense in Nebraska Pseudognaphalium jaliscense (Greenm.) Anderb. was recently reported for the US.A., based on collections from Arizona, New Mexico, Texas, and Colorado (Nesom 2001, including details of typification and description). The current report documents its occurrence in twelve counties of Nebraska. It was first collected in the state in 1968 by Steve Stephens (Custer Co.) but was not recol- lected until 1991 by Herb Karcher (Hayes Co.). It has since been recollected in both counties in addition to numerous other sites in the southwest and the Ne- braska Sandhills in the north central part of the state. The recency of the col- lections and the common roadside habitat suggest recent introduction, but other collections are from apparently natural habitats, and the nativity of Pjaliscense in Nebraska is not clear. Distribution of the species is shown in Figure | for northwestern Mexico and the U.S.A., based on the Nebraska citations below and those in Nesom (2001). Additional records for New Mexico also are cited below and suggest that the species may be even more common in that state, where habitats do not suggest that the species is a recent colonizer. If the distribution of Pseudognaphalium SIDA 21(2):1205—1208. 2004 1206 BRIT.ORG/SIDA 21(2 am ie \ \ ae, | jaliscense PROC \ ko x aha Wen a Be ace £D, 4 fee a +. eh WcA Ad } Movi Th H ; } ~s Fr JF li Navarit.andA li {localities i t IN Leo tst The records ) £ Movi £ llart tidiad hy M £, * L L jaliscense is native in the U.S.A, itapparently should be expected in counties of northeastern Colorado and northwestern Kansas. With further perspective, collections reported as Pseudognaphalium jaliscense from Mendocino, Marin, and Del Norte counties, California (Nesom 2001), prove to be the form of Pseudognaphalium stramineum (Kunth) A. Anderb. sometimes identified as Gnaphalium stramineum var. confertifolium Greene. These plants have larger heads with more numerous, more opaque phyllaries and often more florets than typical P stramineum, which is abundant in California. — ROLFSMEIER AND NESOM, NEW RECORDS OF PSEUDOGNAPHALIUM 1207 U.S.A. NEBRASKA. Blaine Co.: T24N R25W Sec 26, | Aug 1998, Morris 899 (CSCN). Brown Co.: Long Pine, 12 mi ye 6 mi W, a in interdunal Sandhills fee 2 Aug 2000, Rolfsmeier i GR. ee NEB). C .: 2.5 mi N of Hooker Co. line, sandy road ditches along Nebr. 97, 13 Sep et aap ae CSCN, KANU, NEB): S of ene Reservoir, sandy Poadeide: - Sep 1995, oyna 20% (BRIT, SOE KANU ae near turnoff to Marsh Ranch on Nebr. 97, ca. 3.5 mi N of ee lin sandy 22 Aug 1996, I Rolfsmeier 12911 (BRIT, CSCN, KANU, ee caster Co.: Anselmo, 4 mi NW, eae of prairie slough, moist, sandy soil, several plants, 28 Aug 1968, Stephens 28200 (KANU); Oconto, 0.25 mi N, 3.5 mi E, ravine bottom in mixed- grass prairie, 22 Jul 1999, Rolfsmeier 14903 & se (BRIT, NEB). Frontier Co.: T7N R30W Sec 18, heavily-grazed sand prairie near Hayes Co. line, 6 Sep 1992, Rolfsmeier 11117 (BRIT, KANU, NEB, CSCN); S of Curtis, T7N R28W Sec 11, prairie pasture, 29 Sep 1992, John s.n. (CSCN). Grant Co.: 0.1 mi W of Hooker Co. line, scattered at t base of of railroad ceutaanneee 22 Aug 1996, Rolfsmeier 12913 (BRIT, CSCN, ul KANU). Hayes Co.: 13.5 mi S of junction of Nebr. s 23 & 25, roadbank, 21 Jul 1991, Karcher 820 (CSCN, KANU); Hayes oe Wildlife Nonagenen ce ee oie on east side of lake, 6 Sep 1992, Rolfsmeier 11109 (BRIT, KANU, NEB, CSCN). Hooker Co.: 6.25 mi E of Grant Co. line, base of railroad embankment, 22 Aug 1996, RaVemeier 12912 ete CSCN, ANU: NEB). Keith Co.: Paxton, 3.3 mi W, sandy road ditch between US. 30 and railroad, 25 Sep 1992, Rolfsmeier 11282 (CSCN, KANU, NEB). Lincoln Co.: several miles north of — Co. line, sandy prairie along Nebr. 25, 6 Sep 1992, ee (CSCN, NEB); Dickens, 9 mi E, sandy roadbank along Nebr. 23, 1] Sep 1992, trad T, CSCN, KANU, NEB); T12N R34W Sec 26, sandy bank of ditch, W side of hwy 25, 1997, G. see 253 (NEB), Wallace, 6 mi N, sandy slope along highway 23, 20 Sep 1998; aa 51 (CSCN). Loup Co.: Taylor, 7.5 mi ESE, sandy foadbanls elu 199 9, Rolfsmeier 14974 & Parker (CSCN NEB). eee Co.: Elsie, 0.25 mi E, as prai h, 11 Sep 1992 Rolfsmeier 11193 (BRIT, CSCN, KANU). New Mexico. Grant Bayard, open woodland, 63 00 ft, 18 bale 1967, Gierisch ee Hidalgo Co.: ieee grassy ville 22 Aug 1955, Castetter 9959 ( y Ranch, Howe Camp, cienega, 20 Aug 1993, Ivey s.n. (UNM). Lincoln Co.: Sacramento fore Me Gavilan Ridge at trail to Blue Lake, burned area that was originally mostly ponderosa pine, 7300 ft, 30 Sep 2001, Worthington 30728 (UNM). Pseudognaphalium roseum in California Pseudognaphalium roseum (Kunth) A. Anderb. is widespread in Central America and Mexico, from Panama northward through Chiapas to Chihuahua, Veracruz, Nuevo Leon, Tamaulipas, and southeastern Coahuila. It ly above 1000 m elevation in a variety of habitats. The closest collections of P. roseum southward from California are from Sinaloa and southern Chihuahua, but the species is abundant only in eastern and southern Mexico. It is recog- nized by its persistently tose stems and leaves, the leaves subclasping and non-decurrent, weakly bicolored and sessile-glandular beneath the tomentum, often thick stems, large capitula with numerous, opaque-white or sometimes pink phyllaries, numerous florets, and smooth-surfaced cypselae. It has been confused with the smaller-headed P canescens (DC.) A. Anderb., but plants of P. roseum with relatively few bisexual florets can be distinguished from P. canescens by their subclasping leaves commonly with closely wavy margins, broader and more numerous phyllaries, and smooth-surfaced cypselae. A key with diagnostic characters for these taxa is provided below. Collections of P. roseum cited from San Mateo Co. and Santa Barbara Co. may be atypical in their slightly smaller capitula. 1208 BRIT.ORG/SIDA 21(2) U.S.A. CALIFORNIA. Orange Co.: Catalina Island, 16 Jun 1920, Fisher s.n. (SMU). San Diego Co.: Bor- der Field State Park, southwesternmost acre of continental U.S.A., 16 Mar 1989, Turner 15925 (TEX); Tia Juana, 24 Apr 1913, — 2918 oy San Mateo Co.: Pebble Beach, Pescadero, 29 June 1927, Wolf 563 (TEX). Santa Barb . Purissima Canyon, 26 Mar 1925, Munz 4737 (TEX). Ventura Co.: North Signal Street, Ojai, Boe oe Pollard 20331 (SMU) a. Mid-cauline leaves mostly 3-7 cm long, (3-)6-15(-20) mm wide, subclasping, the margins often closely sinuate pistillate florets 45—90(-110); bisexual florets (5-)6-12(-18); cypsela ies stnisoen (non papillate) Pseudognaphalium roseum . Mid-cauline leaves mostly 1-4 cm long, 2-8(—15) mm wide, not clasping, the mar- gins usually straight; pistillate a (16-)24—44; bisexual florets (1-)2—5(-6);cypsela surfaces densely and minutely papillate Pseudognaphalium canescens Pseudognaphalium roseum (Kunth) A. Anderb., Opera Bot. 104:148. 1991, Gnaphalium roseum Kunth in H.B.K., Nov. Gen. & Sp. 4 [folio|:63. 1818; 4 [quarto}81. 1820. TYPE: MEXICO. GUANAJUATO: Humboldt and Bonpland s.n. (HOLOTYPE: P fichel). Ae a8) Taprooted annual to short-lived perennial herbs, the stems 5-20 dm tall, per- sistently woolly-tomentose. Leaves oblong-lanceolate to oblanceolate, the mid- cauline 3-7 cm long, G-)6-15(-20) mm wide, subclasping to clasping but not basally ampliate, not decurrent, the margins often closely sinuate-wavy, per- sistently woolly-tomentose or sometimes tardily glabrescent above, with short- stipitate to sessile glands beneath the tomentum. Involucres 4-4.5 mm high; phyllaries ovate, in 5-6 graduate series, usually opaque-white but sometimes pink. Pistillate florets 45-90(-110). Bisexual florets (5-)6-12(-18), the corollas often pink on the lower half. Cypselae shallowly longitudinally ridged, smooth. ACKNOWLEDGMENTS We gratefully acknowledge loans from UNM, the staff at TEX for helps in visits there, staff at NEB and KANU for confirming collection data for specimens de- posited there, and Caleb Morse for a review of the manuscript. REFERENCE Nesom, G.L. 2001. New records in Pseudognaphalium (Gnaphalieae: Asteraceae) for the United States. Sida 19:1185-1190. THE REDISCOVERY OF THE SOUTH AMERICAN HYBANTHUS PARVIFLORUS (VIOLACEAE) IN NORTH AMERICA B. Eugene Wofford Juliana de Paula-Souza Dept. of Botany Universidade de Sdo Paulo- adi ESA The University of Tennessee v. Pddua Dias 11, Knoxville Tennessee 37996, U.S.A. 418-900 Piracicaba-Sdo Pe: BRASIL Jupsouza@esalq.usp.br Alan S.Weakley Thomas E.Govus University of North Carolina Herbarium (NCU) 3711 Big Creek Road North Sais Botanical Garden Ellijay, Georgia 30536, U.S.A. ey Box #3280, University of North Carolina Chapel Hill, North Carolina 27599-3280, U.S.A. weakley@unc.edu ABSTRACT An unknown plant collected in 1998 at Fort Pulaski, Chatham County, Georgia has been identified as Hybanthus parviflorus (Violaceae), a South American native. The Georgia collection represents only the third report of this species in North America, and the first since the 1880s, when it was collected on eee ballast at two port cities in New Jersey. It is possible that the introduction of the was also via ship’s ballast, as Fort Pulaski would have been a port of call for ships Sein to and fon Savannah, a major seaport in the ie and ase centuries; if the Georgia plant was introduced on ballast, it is apparently established. It i hat iti introduc- tion; if so, however, the means of introduction of this sspecis is obscure, as ae is not typically an agri- eee nor of horticultural interest. The u lig il facilitated the rapid and accurate identification of this wee species, which fould now be considered an estab- lished, though rare, component of the North American flora. RESUMEN Una planta desconocida colectada en 1998 en Fort Pulaski, Chatham SS ee aes sido identificada como Hybanthus parviflorus (Violaceae), nativa de Sur Am .La gia es la tercera cita as esta Ee cie en geese Nata sTet y la primera ne los 1880s, ee nie i eee Es posible que la introduccion Georgia | id bié li l ya que Fort Pulaski podria haber J KT coe adaen sido un iene de asda para barcos que van o regresan de Savannah, un gran puerto en los siglos XVII y XIX; si la planta de Georgia fue introducida en el lastre, esta aparentemente establecida. Es también posible que sea una introduccion més reciente; en este caso, sin embargo, el sistema de Eee ees esta epaicn no esta gre ye que BBEAmente noes una ar meee agricola, ni de interés ho a. El y 5 bapice y 2 o o 7 5 ae ei 4 ey Ct C L pe aunque raro, de la flora de Norte América. The University of Tennessee Herbarium (TENN) recently received a loan of se- lected Gratiola (Scrophulariaceae) from the University of Georgia Herbarium SIDA 21(2): 1209-1214. 2004 1210 BRIT.ORG/SIDA 21(2) (GA). Included within the loan were a few unidentified specimens of either unrequested Gratiola or the closely related Lindernia, and an undetermined specimen (Figs. 1, 2) collected by Govus (1998) from Ft. Pulaski National Monu- ment, Chatham Co., Georgia, as part of a catalog of the vascular flora of that USS. National Park Service unit (Govus 1998). At GA, this last specimen had been tentatively identified by an unknown individual as Gratiola virginiana, and then that identification had been crossed through, but the specimen was ap- parently left in a Gratiola folder and sent out with the Gratiola loan to TENN. Upon receipt of the loan at TENN, Wofford immediately eliminated this speci- men asa member of Scrophulariaceae based on overall morphology. A few flow- ers were rehydrated and upon dissection its floral morphology unequivocally placed it in Violaceae, ie, flowers zygomorphic, sepals 5, subequal, petals 5, unequal, the lower one slightly spurred at the base and wider than the others; stamens 5, the lower two provided with nectariferous appendages, the connec- tive prolonged into a membranous appendage; ovary superior, unilocular, 3- valved, placentation parietal. TENN houses only two genera of Violaceae (Viola and Hybanthus), this speci- men obviously was not a Viola but the flowers were strikingly similar to the common eastern North American Hybanthus concolor (TE Forst.) Spreng. and the ee North American H. verticillatus (Ortega) Baill. Wofford de- veloped the h hat this might be an introduction of an extracontinental species of Hybanthus, a ene genus (extending into warm temperate areas, as in eastern North America) of about 85 species (Ballard, in press), though sometimes credited with as many as 150 species (Mabberley 1997). Additional specimens from Mexico and Cuba available at TENN were examined, but none remotely fit the overall morphology of this specimen. Digital images of the unknown plant were then sent to Weakley at the University of North Carolina Herbarium (NCU). Two NCU specimens of Hybanthus (from Paraguay and Argentina) closely resembled the unknown from Georgia; unfortunately, both specimens were labeled only “Hybanthus” and lacked an identification to the species level. Finally, an electronic image of the Georgia specimen was sent to Paula-Souza, who has hinterests in Violaceae, especially Hybanthus. She immediately responded that “this plant is Hybanthus parviflorus (Mutis ex Lf.) Baill. This is a very common plant here in South America, and although it is considered a weed in some places, | have never seen a record of it from the United States.” Further investigation revealed that H. parviflorus had indeed been reported previously from North America, by Dowell (1906), as Calceolaria glutinosa (Vent.) Kuntze, based on several specimens collected “on ballast at Communi- paw Ferry,” New Jersey, in July and September 1880 (see specimens cited be- low). H. parviflorus is additionally represented by a specimen - a Academy of Natural Sciences in Philadelphia (PH), identified as Ionidi Vent. WOFFORD ET AL., HEVISCUVERT VE 1211 Fic. 1. Image of Govus coleeual of Aypantnas uu Fic. 2.€ from Fort Pulaski, Georgia g y parviflorus from Fort Pulaski, Georgia. Photograph by Gene Wofford. 3 and labeled “Herbarium of Isaac Burk, Philadelphia; Pa., on ballast, Kaigns Pt., N,J., October 1885.” However, catalogs of the flora of New Jersey (Britton 1889), and the various manuals covering the Northeastern United States (Fernald 1950, Gleason 1952; Gleason & Cronquist 1991) do not include this taxon, presum- ably considering it a ballast waif, and it is also not included in the recent check- list me Boe of North America (Kartesz 1999). viflorus is widely distributed over South America, from Ven- ae through southern Brazil (Figs. 3, 4), Chile, and Argentina, occurring pref- erentially in cold regions and at higher altitude places in warmer regions. The species is commonly found in open sites, frequently behaving as a weed in pas- 1212 BRIT.ORG/SIDA 21(2) tures and roadsides, though not asa weed of cultivated crops. The 1880 New Jersey specimens came from ship’s ballast, at the ferry landing of the Communipaw Ferry, one of sev- eral busy ferries between Jersey City and New York City. The 1885 speci- men also came from ballast, at Kaighn’s Point on the Delaware River in Camden, alsoa busy ferry landing serving Camden, New Jersey and Philadelphia, Pennsylvania, in opera- tion from 1806 until at least 1926. It seems plausible that H. parviflorus was introduced to Fort Pulaski, Geor- gia, via ballast as well. Fort Pulaski has been actively used by ships since at least the mid 1700s, and would have been a regular docking area for sailing ships through much of the late 18th and early 19th centuries, including use by ships going to and from the large and active port city of Savannah, Georgia (ca. 20 miles away) to destinations around the world. If H. parviflorus were intro- duced to Georgia on ballast, it is ap- parently established, as the 1998 col- lection is long after the use of solid ballast material was discontinued. It is also possible that this represents a more recent introduction, but how and why this species would have ar- rived at Fort Pulaskiisobscure,asthe = ——— eee species in neither typically a weed of _ Fis.3.Habit of Hybantt iflorus. Photograph by Juli agricultural crops nor an object of horticultural interest. Given this more recent occurrence of H. parviflorus, found 118 years after its initial collection in North America, it appears that H. parviflorus should be considered a rare, alien component of the North American flora, and collectors should be aware of its potential occurrence, especially in the vicinity of old seaports. It does not appear, at least at this time, that it is likely to be an aggres- om de Paula-Souza, from live material in Itararé, S40 Paulo, Brazil. WOFFORD ET AL., REDISCOVERY OF 1213 be fared 7 Fic. 4. Detail of flower of Hybanthus parviflorus. Pt Paulo, Brazil. sive alien weed. Below, we present a description to aid in its recognition, should it occur elsewhere in North America. We also note that the increased use of digital imagery and electronic mail in herbarium work greatly facilitates inter- national collaboration and the identification of an unknown, extracontinental and potentially invasive species; “virtual annotation” can provide rapid results without the expense and risk of loaning uniquely valuable specimens. Hybanthus parviflorus (Mutis ex Lf.) Baill., Bot. Med. 2:841. 1884. Herbs up to 30 cm high, branched, erect or suberect, internodes (1.7-)4-20 mm long; leaves alternate or opposite, frequently opposite only at the base of the branches, petiole ca. 0.5-4 mm long, blade (3.5-)6-30 mm long x 2-10(-15) mm wide, elliptic, lanceolate or oblong, less frequently oblanceolate, widely ellip- tic, ovate or obovate, acute to obtuse at apex, margin serrate, base acute, attenu- ate, rounded or obtuse, glabrous to pubescent or puberulent on the midrib; stipules narrowly lanceolate; flowers white, frequently with purplish lines on 1214 BRIT.ORG/SIDA 21(2) the anterior petal, solitary, axillary or arranged in poorly defined racemose in- florescences; floral pedicel 3-11(-24) mm long, bracteoles absent; sepals equal or subequal, ca. 1-1.7 mm long x ca. 0.5 mm wide, narrowly lanceolate or rarely ovate to suborbicular, subfalcate, acuminate or rarely round or obtuse at apex, margin entire; posterior petals ca. 1 mm wide x ca. 0.5 mm long, oblong, lateral petals ca. 1.5-1.8 mm long x ca. 0.7 mm wide, oblong to lanceolate, falcate, ante- rior petal (1.5-)3-3.7 mm long x 1.2-1.4 mm wide, clawed; stamens subsessile, anthers ca. 0.5mm long, connective appendages orange-brown, ca. 0.3 mm long, frequently asymmetric, anterior stamens provided with noduliform nectariferous appendages; style 0.6-0.8 mm long, straight or subsigmoid, ovary ca. 0.6 mm long, glabrous, capsule 3-4 mm long x 3.5 mm wide, ovoid to glo- bose, seeds ca. 1.5 mm long x ca. 1-1.2 mm wide. Voucher specimens: GEORGIA. Chatham Co he picnic area, south central Cockspur Island, 7 Apr 1998, Gove 946 (GAD. NEW JERSEY. Camden Co.: on ballast, Kaigns Pt., N.J., Oct 1885, Isaac Burk s.n. (PH!). Hudson Co.: ballast, near Communipaw Ferry, ae Sep ed as Brown s.n.(GHI, NY!, US); Communipaw, NJ. (ballast), 20 Jul 1880, Jos. Schrenk s.n.(NYD. ACKNOWLEDGMENTS We thank the curators of GA, GH, NY, PH, and US for access to collections. We thank Harvey Ballard for pointing out the existence of the 1885 PH collection, and Harvey Ballard and Julie Ballenger for helpful reviews of the manuscript. REFERENCES Battarb, H.E., Jr. [submitted] Violaceae. In: Kubitzki, K., ed. Families and genera of vascular plants. Institut fur Allgemeine Botanik, Hamburg, Germany. Britton, N.L. 1889. Catalogue of plants found in New Jersey. Final Report of the State Ge- ologist, vol. II, Trenton, N.J. Dowett, P.1906. North American species of Calceolaria. Bull. Torrey Bot. Club 33:547-556. Fernald, M.L. 1950. Gray's manual of botany, eighth (centennial) edition. D. Van Nostrand Co., New York, N.Y. GLEASON, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United States and adjacent Canada. New York Botanical Garden, Bronx, NY. Gieason, H.A. and A. Cronauist. 1991, Manual of vascular plants of northeastern United States and adjacent Canada, second edition. New York Botanical Garden, Bronx. Govus, T.E. 1998. Fort Pulaski National Monument inventory final report, Part A: Vascular plants. National Park Service, Southeast Region, Atlanta, GA. Kartesz, J.T. 1999.A synonymized checklist and atlas with biological attributes for the vas- cular flora of the United States, Canada, and Greenland. First Edition.In:Kartesz, J.T.,and C.A.Meacham. Synthesis of the North American flora, Version 1.0. North Carolina Bo- tanical Garden, Chapel Hill. Maseertey, D.J. 1997. The plant-book: a portable dictionary of the vascular plants. Second edition. Cambridge Univ. Press, Cambridge, England. ASTERAGEAE FROM WOOL MIBESITES IN SOUTH CAROLINA, INCLUDING NEW RECORDS FOR NORTH AMERICA Guy L.Nesom Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Documentation is given for 73 taxa of Asteraceae collected in 1957-1960 from wool mill sites in Ber- Sl County an Florence County, South Carolina. Twenty-eight (28) of these taxa are first reports r the flora of South Carolina; seventeen (17) others represent the first documentation for earlier canines state. Six (6) are first reports for North America: Chevreulia sarmentosa, Gamochaeta argentina, and Panphalea heterophylla (native to South America), and Peripleura arida, Stuartina hamata, and Vittadinia sulcata (native to Australia). Calotis cuneifolia (native to Australia) is re- ported for North America for the second time. Fifty-nine (59) of the 73 taxa are not native to South Carolina; of these, 29 are native to continents other than North America, while 30 are native to North America but from regions outside of the state. RESUMEN ] | ae see BG tt ne Se ofrece documentacién de 73 taxa de 1960 en los condados de Berkeley y Florence County, Carolina del Sur. Veintiocho (28) de estos taxa son primeras citas para la flora de Carolina del Sur; otros diez y siete (17) representan los primeros testigos para Citas previas en el estado. Seis (6) son ae citas para Norte América: Chevreulia sarmentosa, Gamochaeta argentina, and akaeases heterophylla (nativa de Sur América), y Pe ee es Stuartina hamata, y Vittadinia sulc de Australia) a ae ativa de A ) para Norte América por segunda vez. Cincuenta y nueve (59) de los 73 taxa no son nativos de Carolina del Sur; de estos, 29 son nativos de otros continentes diferentes - Norte América, mientras que 30 son nativos de Norte América pero de reg iones fuera del e In the years 1957 through 1960, as part of the Flora of the Carolinas project and working from the University of North Carolina-Chapel Hill herbarium, Harry Ahles and companion John Haesloop made repeated collecting trips to two ‘wool combing mills’ in South Carolina. One of these was in Jamestown in Berkeley Co., the other north of Johnsonville in Florence Co.; both sites are located on the coastal plain, 40-50 kilometers from the coast and about 60 kilometers from each other. Some of the collections were identified and incorporated into the general collections at NCU several decades ago; others were mounted but were not identified or inserted into the general collections until 2001. Only a few of these were included in the account of the Flora of the Carolinas (Radford et al. 1968). Some have been the basis of recent reports (without documentation) for distribution records (Kartesz 1999 and others). SIDA 21(2): 1215-1223. 2004 1216 BRIT.ORG/SIDA 21(2) —— The wool combing mills received raw fleece and processed it toward pro- duction of clean wool for spinning. Early stages in the process are mechanical shaking (removes dirt and some plant matter), “scouring” (removes chemical substances and sand), and “combing” (removes finer plant material as well as shorter, weaker fibers). Waste from these processes at the South Carolina mills was the source of propagules for species collected by Ahles and Haesloop. De- livery of the raw wool probably was by railroad, because both of the sites are along the CSX Railroad, which runs roughly parallel to the coast. Both mills apparently are still in operation. Seventy-three taxa (73) of Asteraceae in 54 genera are represented in the col- lections identified here. Twenty-eight (28) of these are first reports for the flora of South Carolina; seventeen (17) others represent the first documentation for ear- lier reports for the state. Six (6) are first reports for North America: Chevreulia sarmentosa, Gamochaeta argentina, and Panphalea heterophylla are native to South America; Peripleura arida, Stuartina hamata, and Vittadinia sulcata are native to Australia. Calotis cuneifolia (native to Australia) is reported for North America for the second time. Fifty-nine (59) of the species are not native to South Carolina; of these, 29 are native to continents other than North America, while 30 are native to North America but from regions outside of the state. It would be useful to reinvestigate the status of these taxa in South Caro- lina, in order to determine which have persisted or increased their distribution. A number of the taxa reported here may have the potential to spread and be- come damaging weeds, or they may simply be waifs (sensu Nesom 2000). Collections by Ahles and Haesloop were made in 1957 (April, May, July, Sep- tember, October), 1958 (April, May, June, July, September), 1959 (May, July), and 1960 (May, July). From each of the two wool mill sites, collections were provided with essentially the same label data, except for the collection number and date. SOUTH CAROLINA. Berkeley Co.: waste ground around Santee Wool Combing Mill, Jamestown on S.C. Rte. 45, [Apr-Oct 1957-1960], H.E. Ahles[col- lection number] with J.G. Haesloop. SOUTH CAROLINA. Florence Co.: waste ground around the Wellman Wool Combing Mill, north of Johnsonville on S.C. Rte. 41, [Apr-Oct 1957-1960], H.E. Ahles[collection number] with J.G. Haesloop. PLANT LIST The genera and species are arranged alphabetically. If more than one collection was made for a single species, the citations are arranged chronologically. Vouch- ers are located at NCU, unless otherwise indicated. Achillea millefolium L. U.S.A.; previously known from South Caro- Berkeley Co.: 13 Jun 1958, Ahles 42939 lina (Radford et al. 1968). Berkeley Co.:27 May 1959, Ahles 52717 Amblyolepis setigera DC. INaUlve tO western Asia, widely naturalized in the Berkeley Co:8 Apr 1957, Ahles 22614 NESOM MILL Native to Texas and Mexico; first report for South Carolina. Ambrosia artemisiifolia L Berkeley Co.: oe He 1959, Ables 52805 Niloti tr NI vide ly |: +1 INGAUVE QlouIwWu previously known from Suit Caroline (Rad- ford et al. yi Amphiachyris dracunculoides (DC.) Nutt. Berkeley Co.: 28 Sep 1957, Ahles 35585 Berkeley Co.: 30 Oct 1957, Ahles 38195 Berkeley Co.: 22 Sep 1958, Ahles 49216 Native to the south-central U.S.A. as far east as Alabama and Tennessee; reported for South Carolina by Kartesz (1999), documented here Anthemis cotula L. Berkeley Co.: 20 May 1957, Ahles 25827 Florence Co.: 28 May 1957, Ahles 26690 Berkeley Co.: 27 May 1959, Ahles 52797 Florence Co.: 11 Jul 1960, Ahles 53756 NI |e Fal Native t Asia turalized in widely Ila U.S.A.; rerio known from South an (Radford et al. 1968). Arctium minus Bernh. Berkeley Co. 11 Jul 1960, Ahles 53824 Native to Europe, widely naturalized in the U.S.A, eviously known from South Carolina (Rad- et al. 1968). Artemisia annua L. Berkeley Co: 11 Jul 1960, Ahles 53848 Native to eastern Europe and Asia, natural- ized in various states of the USA; first report for South Carolina. Artemisia biennis Willd. var. biennis Berkeley Co.: 20 May 1957, Ahles 25897 Berkeley Co.: 27 May 1959, Ahles 52762 Berkeley Co.: 11 Jul 1960, Ahles 53814 Native to western and northern U.S.A. and Canada; first report for South Carolina. Artemisia vulgaris L. var. vulgaris Berkeley Co.: 25 May 1960, Ahles 53402 Berkeley co 225 Mey. 1960, Ahles 53403 Native t d Asia, widely naturalized | in Canada and the eastern U.S.A. previously known from South Carolina (Radford et al, 1968). 1217 Bidens bipinnata L. Berkeley Co.: 20 May 1957, Ahles 25836 (NCU, SMU) Berkeley Co.: 14 May 1958, Ahles 40488 Native to the southern and eastern U.S.A. and Mexico; previously known from South Caro- lina (Radford et al. 1968). Bidens pilosa L Florence Co.: 14 May 1958 Ahles 40446 Florence Co.:30 Oct 1957, Ahles 38153 Florence Co.:28 May 1957, Ahles 26699 Florence Co.:22 Sep 1958, Ahles 49139 Native to Mexico and Central America, natural- ized in the southwestern and southeastern U.S.A. and a few other states; previously known from South Carolina (Radford et al. 1968) Bidens polylepis Blake Berkeley Co.:5 Jul 1957, Ahles 30809 Native to the eastern U.S.A.; previously known from South Carolina (Radford et al. 1968). Bidens frondosa L. Berkeley eo: tas Sep oe aS cok Native and erica; pr ously Hk from South Carolina (Radford etal. 1 Calotis cuneifolia R Berkeley Co.:8 Apr 1 ie ee ee Co.: 14 May 1958, Ahles 40564 Berkeley Co.: 13 Jun 1958, Ales ae 42973 Native to Australia } the U.S.A only (Sorrie 1992); first er for Massacl for South Carolina. Carduus pycnocephalus Berkeley ue . ae ee aes paced Native to! rm Asia, natu- alized in sestenaie states oer western and Gulf coast U.S.A.; reported for South Caro- lina I Kartesz (1999), documented here. Carthamus baeticus (Boiss. & Reut.) Lara (syn = Carthamus lanatus | & Reut aN man Berkeley Co.: 13 Jun 1958, Ahles 42899 Berkeley Co.: 27 May 1959, Ahles 52774 (det. DJ. Keil 1997) Berkeley Co.: 27 May 1959, Ahles 52854 Berkeley Co.: 25 May 1960, Ahles 53372 Native to southern Europe and northern Africa, subsp. baeticus (Boiss 1218 naturalized in scattered states, mostly along the Pacific coast; reported for South Caro- lina by Kartesz (1999), documented here. Centaurea americana N Berkeley Co. 27 May fa : les 52776 Berkeley Co.:27 May 1959, Ahles 52843 Berkeley Co.:25 May 1960, Ahles 53492 Native primarily to the central U.S.A.; reported for South Carolina by Kartesz (1999), docu- mented her ~— Centaurea melitensis L. Berkeley Co.:8 Apr 1957, Ahles 22624 age Co. 14 May 1958, Ahles 40542 (det. L. Shinners) a - 13 Jun 1958, Ahles 42919 Native to southern Europe and northern Africa, naturalized in the western U.S.A.and various other states; reported for South Carolina by Cronquist (1980), documented here. ate solstitialis Berkeley 25 May ea Ahles 53487 ey . 11 Jul 1960, Ahles 53846 (det. L. Shinners) Native to western Asia and the Mediterranean region of Europe, naturalized and weedy in most of the U.S.A; reported for South Caro- lina by Hill and Horn (1997). Chaetopappa asteroides (Nutt.) DC. var. asteroides Berkeley Co.:8 Apr 1957, Ahles 22551 Native to northern Mexico and the south-cen- tral U.S.A. first report for South Carolina. Chevreulia sarmentosa (Pers.) Blake Berkeley Co.: 14 May 1958, Ahles 40509 Native to South America (Brazil, Paraguay, Uru- guay, Bolivia, Argentina); first report for the U.S.A. and South Carolina. eee Conyza bonariensis (L.) Crona. Berkeley Co.: 14 May 1958, Ahles 40568 Berkeley Co.:21 Jul 1958, Ahles 46996 Native to South America, naturalized in most outhern states of the U.S.A.; previously known from South Carolina (Radford et al. 1968) Coreopsis tinctoria Nutt. var. tinctoria Berkeley Co.:27 May 1959, Ahles 52825 Berkeley Co.: 11 Jul 1960, Ahles 53822 BRIT.ORG/SIDA 21(2) Native and wide North Americ revi- ously nn aie South Carolina (Radford etal. 1 Coreopsis verticillata L. Berkeley Co.: 13 Jun 1958, Ahles 42900 Alar + th th INGUIVE LO UIE SOUTTTIETT from South Carolina (Radford et al. ] “previously ky IOVVE] 1968). Cotula australis (Sieber) Hook. f. Berkeley Co.:8 Apr 1957, Ahles 22592 Native to Australia and New Zealand; ee ized in various western U.S.A. states and Florida; first report for South i Dittrichia graveolens (L.) W.Greuter Berkeley Co. 28 aca 957,Ahles 35595 Native tothe M naturalized in various regions, including Australia, New nd, and California (Preston 1997), and reported from various other localities in the U.S.A. (Gleason & Cronquist 1991; Mitchell & Tucker 1997;Preston 1997;Tucker 1995): first report for South Carolina.|n view of the other species reported here from Australia, it seems likely that the wool mill plants of Dittrichia graveolens also originated from there rather than from the native region of the spec Dracopis amplexicaulis (Vahl) Cass. Berkeley Co.: 20 May 1957, Ahles 25840 Berkeley Co.: 25 May 1960, Ahles 53369 Native primarily to the south-central U.S.A.; re ported for South Carolina by Kartesz (1999), documented here Dyssodia papposa (Vent.) A.S. Hitchc. Berkeley Co.:8 Apr 1957, Ahles 22620 Berkeley Co.: 20 May 1957, Ahles 25892 Berkeley Co.: 14 May 1958, Ahles 40483 Berkeley Co.: 14 May 1958, Ahles 40553 Native to the southwestern and central U.S.A; first report for South Carolina. Eupatorium dubium Willd Berkeley Co.: 15 Jul 1959, ie 52950 Native to Atlantic coast states of the U.S.A, from South Carolina northward; previously known from South Carolina (Radford et al. 1968). Evax multicaulis DC. Berkeley Co.:8 Apr 1957, Ahles oy es Co. 14 May 1958, Ahles 4051 NESOM MILL Native to Mexico and various states of the south- ern U.S.A. reported for South Carolina by Cronquist (1980), documented h Evax prolifera Nutt. ex DC. Berkeley Co.: 14 May 1958, Ahles 40512, 40513 Berkeley Co.:27 May 1959, Ahles 52737 Native to Mexico and various states of the Great Plains and southern U.S.A, first report for South Carolina. Facelis retusa (Lam.) Schultz-Bip. Berkeley Co.: 27 May 1959, Ahles 52731 Berkeley Co. 27 May 1959, Ahles 52858 Native to South America and naturalized in states of the Gulf coast and southern Atlan- tic coast; previously known from South Caro- lina (Radford et al. 19 Flaveria trinervia (Spreng.) Mohr Berkeley Co.: 28 Sep 1957, Ahles 35584 Berkeley Co: 21 Jul 1958, Ahles 47035 Berkeley Co. 22 Sep 1958, Ahles 49172 Berkeley Co.: 22 Sep 1958, Ahles 49215 Native to northwestern Mexico and the south- western United States, naturalized in a few states of eastern U.S.A.;first report for South Carolina Gaillardia pulchella Foug. var. drpummondii (Hook.) B.L. Turner Berkeley Co.:8 Apr 1957, Ahles 22528 Berkeley Co.: 20 May 1957, Ahles 25852 14 May 1958, Ahles 40460 Berkeley Co.: 13 Jun 1958, Ahles 42902 Berkeley Co.: 13 Jun 1958, Ahles 42950 Berkeley Co.: 13 Jun 1958, Ahles 42969 Berkeley Co.: 25 May 1960, Ahles 53367 Native primarily to the south-central U.S.A. but naturalized in various other states; reported for South Carolina by Kartesz (1999), docu- mented here. The collections were deter- mined by annotation as var.drummondii by B.L. Turner & TJ.Watson in 1995. Gamochaeta antillana (Urb.) Anderb. Berkeley Co.:8 Apr 1957, Ahles 22529 Florence Co.:8 Apr 1957, Ahles 22628 Probably native to the Caribbean region and perhaps South America, perhaps southeast- ern USA; first recognized by this name and circumscription in the U.S.A. by Nesom (2004) 1219 makes: argentina Cabr. erkeley Co. 8 Apr 1957, Ahles 22594 . to South America (Argentina and Uru- guay); first report for the U.S.A. and South Carolina Gamochaeta calviceps (Fern.) Cabr. Berkeley Co.: 27 May 1959, Ahles 52842 Perhaps native to the southeastern USA, appar- ently naturalized worldwide; recognized by this name and circumscription in the U.S.A. by Nesom (2004). Gamochaeta pensylvanica oS ) Cabr. Bewncley Co.: ae r 1957, Ahles h Ame fateson ai fe aaraie in the southern U.S. nd throughout the world; pr A con from South Carolina (Radford et al. 1968). P robably TGV cr L - VALIA ex Spreng.) Cabr. Berkeley Co.:5 Jul 1957, Ahles 30825 Berkeley Co.: 21 Jul 1958, Ahles 47038 Berkeley Co.: 15 Jul 1959, Ahles 52959 Berkeley Co.: 11 Jul 1960, Ahles 53783 Native to South America, naturalized in a few states of Atlantic gulf coast U.S.A.; previously known from South Carolina, first reported by Nesom (1999). Grindelia lanceolata Nutt. var. lanceolata Berkeley C 1958, Ahles 42920 Native to the south-central U.S.A,;first report for South Carolina. The plant is sterile, but the identification is probably correct. The plant has relatively broad leaves like plants from east Texas. Gutierrezia sarothrae (Pursh) Britt.& Rusby Berkeley Co.: 28 Sep 1957, Ahles 35269 Native to Mexico and the western U.S.A, first report for South Carolina. Gutierrezia texana (DC.) Torr. & A. Gray var. texana Berkeley Co.: 11 Jul 1960, Ahles 53821 Berkeley Co.: 22 Sep 1958, Ahles 49193 Native to Mexico and Texas, naturalized or pos- sibly native in a few other states of central U.S.A.;first report for South Carolina. Helenium amarum (Raf.) H. Rock var. badium (A. Gray ex S.Wats.) Waterfall 1220 Berkeley Co.:20 May 1957, Ahles 25829 Berkeley Co.: 20 May 1957, Ahles 25834 Berkeley Co.: 14 May 1958, Ahles 40478 Berkeley Co.: 13 Jun 1958, Ahles 42936 Berkeley Co.: 27 May 1959, Ahles 52727 Native to Texas and Oklahoma; reported for South Carolina by Kartesz (1999), docu- mented here Helenium elegans DC. var. elegans Berkeley Co.: 25 May 1960, Ahles 53357 Native to Texas, Oklahoma, and Louisiana; re ported for South Carolina by Kartesz (1999), documented here Helenium microcephalum DC. var. microcephalum Berkeley Co.: 20 May 1957, Ahles 25844 (NCU, SMU Berkeley Co.:5 Jul 1957, Ahles 30803 Berkeley Co.: 21 Jul 1958, Ahles 47002 Berkeley Co.:27 May 1959, Ahles 52719 Berkeley Co.:25 May 1960, Ahles 53405 Florence Co.: 26 May 1960, Ahles 53560 Florence Co.;11 Jul 1960, Ahles 53750 (NCU,SMU) Berkeley Co.: 11 Jul 1960, Ahles 53816 Native to Texas, Oklahoma, New Mexico, and olorado; reported for South Carolina by Kartesz (1999) documented here. Helianthus annuus L. sae Co. 20 May 1957, Ahles 25862 Berkeley Co.: 13 Jun 1958, Ahles 42987 Florence Co.: 22 Sep 1958, Ahles 49145 Berkeley Co.: 27 May 1959, Ahles 52777 Berkeley Co.: 25 May 1960, Ahles 53456 Berkeley Co.: 11 Jul 1960, Ahles 53770 Berkeley Co.: 11 Jul 1960, Ahles 53820 Native or naturalized across the entire U.S.A; previously known from South Carolina (Rad- ford et al. 1968). sane Heliomeris multiflora (Nutt.) Blake var. multi- flora (syn = Viguiera multiflora (Nutt.) Blake) Berkeley Co.: 14 May 1958, Ahles 40541 Berkeley Co.:27 May 1959, Ahles 52778 Native to the western USA and Mexico; first re- port for South Carolina. Hymenoxys odorata DC. Berkeley Co.:8 Apr 1957, Ahles 22512 Berkeley Co.:8 Apr 1957, Ahles 22603 BRIT.ORG/SIDA 21(2) Native to the southwestern U.S.A, first report for South Carolina. Hypochaeris brasiliensis (Less.) Griseb. var. dei (Hook. & Arn.) Baker Berkeley Co.:8 Apr 1957, Ahles 22623 Berkeley Co: 28 Sep 1957, Ahles 35593 Berkeley Co: 27 May 1959, Ahles 52730 Native to South America, naturalized in states along the Gulf coast and southern Atlantic; Previously known from oe ne es 10K by Shinners (1 collection. IOUT ted Hypochaeris glabra |. Berkeley Co. 14 May 1958, Ahles 40469 Berkeley Co.: 13 Jun 1958, Ahles 42954 Berkeley Co. 27 May 1959, Ahles 52738 Native to South America, naturalized in states along the Gulf coast and southern Atlantic and various others; previously known from South Carolina (Radford et al. 1968). Iva annua L. Berkeley Co.: 28 Sep 1957, Ahles 35606 malice | Co. 30 ee 957, ay es 38194 nd eastern U.S.A. P robably lal Bieviously known from South Carolina Anonymous 1997), documented here. —_ Ilva axillaris Pursh Berkeley Co. 13 Jun 1958, Ahles 42961 Nat rn North America; first report for Soot Esteli: Ilva xanthifolia Nutt. Berkeley Co.: 14 May 1958, Ahles 40531 Berkeley Co.:27 May 1959, Ahles 52803 Native to most states of the U.S.A. except the Southeast; first report for South Carolina. Krigia virginica (L.) Willd. Berkeley Co. 8 Apr 1957, Ahles 22627 Florence Co.:8 Apr 1957, Ahles 22546 Florence Co.:9 Apr 1958, Ahles 38274 Native to the eastern U.S.A, previously known from South Carolina (Radford et al. 1968). Lactuca graminifolia Michx. Berkeley Co.: 13 Jun 1958, Ahles 42968 Native to coastal states of the southeastern U. eviously known from South Caro faa (Radford et al. 1968). all NESOM MILI John Pruski) Berkeley Co.:8 Apr 1957, Ahles 22585 Native to South America (Brazil, Uruguay, Ar- gentina);first report for the U.S.A.and South Carolina (Pruski & Nesom 2004). a Parthenium hysterophor Berkeley Co.: 20 May 1957, ae 25850 Berkeley Co.:5 Jul 1957, Ahles 30810 Berkeley Co.: 22 Sep 1958, Ahles 49161 Berkeley Co: 15 Jul 1959, Ahles 52960 ee native to the West Indies and adja- cen rth America and South America, ae in various states of south-cen- tral U.S.A. and elsewhere; first report for South Carolina. Peripleura arida (Burbidge) Nesom (syn = Vittadinia arida Burbidge) Berkeley Co. 14 May 1958, Ahles 4054/7 Native to Australia; first report for the U.S.A. and South Carolina. Pseudognaphalium stramineum (Kunth) n E Berkeley Co.: 11 Jul 1960, Ahles 53825 Native to Mexico and the western U.S.A, natu- ralized in Virginia, North Carolina, and South Carolina; previously known from South Caro- lina (Radford et al. 1968) Ratibida columnifera (Nutt.) Woot. & Stand. Berkeley Co.: 20 May 1957, Ahles 25830 Berkeley Co.: 14 May 1958, Ahles 40468 Berkeley Co.: 27 May 1959, Ahles 52712 Berkeley Co.: 27 May 1959, Ahles 52801 Berkeley Co.: 27 May 1959, Ahles 52860 Berkeley Co.: 25 May 1960, Ahles 53370 Berkeley Co.: 25 May 1960, Ahles 53371 Berkeley Co.: 25 May 1960, Ahles 53491 Florence Co. 11 Jul 1960, Ahles 53753 Native to Mexico and the central U.S.A, widely naturalized; reported for South Carolina by Kartesz (1999), documented here Rudbeckia hirta L. var. angustifolia (TV. Moore) Perdue Berkeley Co.: 20 May 1957,Ahles 25863 Berkeley Co.: 13 Jun 1958, Ahles 42916 Berkeley Co.: 27 May 1959, Ahles 52830 Native to the coastal states of southeastern 1221 U.S.A.; previously known from South Caro- lina (Anonymous 1997), documented here. Soliva sessilis Ruiz & Pavon (syn = Soliva pterosperma (Juss.) Less.) Berkeley Co.: 27 May 1959, Ahles 5271 Native to South America, naturalized in the southeastern U.S.A.;previously known from South Carolina (Radford et al. 1968), — Sonchus oleraceus L. Berkeley Co.: 8 Apr ca Ahles 22615 Native to Europe, northern and western Asia,and northern Africa, widely naturalized in North America; previously known from South Caro- lina (Radford et al. 1968). Stuartina hamata Philipson Berkeley Co.:8 Apr 1957, Ahles 22593 Native to Australia; first report for the U.S.A.and South Carolina. Symphyotrichum divaricatum (Nutt.) Nesom (syn = Aster subulatus Michx. var. ligulatus Shinners) Berkeley Co.: 30 Oct 1957, Ahles 38169 Native to northern Mexico and the south-cen- tral U.S.A. first report for South Carolina. Tagetes minuta Berkeley Co.: 20 May 1957, Ahles 25845 Berkeley Co.: 28 Sep 1957, Ahles 35609 Berkeley Co.: 22 Sep 1958, Ahles 49188 Native to South America, in the U.S.A. natural- ized mainly in Atlantic coastal states; previ- ously known from South Carolina (Radford et al. 1968 Tetraneuris linearifolia (Hook.) Greene var. linearifolia (syn = Hymenoxys linearifolia Hook.) Berkeley Co.:8 Apr 1957, Ahles 22513 Native to Texas, New Mexico, Oklahoma, and Kansas; first report for South Carolina. Thymophylla tenuiloba (DC.) Small var. tenuiloba) (syn = Dyssodia tenuiloba (DC) obins. var. tenuiloba) Berkeley Co.:8 Apr 1957, Ahles 22580 Native to Mexico and Texas, naturalized in a few Gulf coast states (e.g. Florida and Mississippi, Cronquist 1980; Alabama,Lelong 1988); first report for South Carolina 1222 Uropappus lindleyi (DC.) Nutt. Florence Co.: 26 May 1960, Ahles 53543 Native to the southwestern U.S.A.;first report for South Carolina. bahar been (Cav.) var. encelioides Berkeley O May 1957, Ahles 255 Florence ae May 1957, Ahles 26719 Berkeley Co.:5 Jul 1957, Ahles 30812 Berkeley Co.: 14 May 1958, Ahles 40486, 40554 Florence Co.:21 Jul 1958, Ahles 46950 Berkeley Co.: 22 Sep 1958, Ahles 49180 Berkeley Co.:27 May 1959, Ahles 52810 Berkeley Co.: 15 Jul 1959, Ahles 52945 Florence Co.: 26 May 1960, Ahles 53546 Florence Co.:11 Jul 1960, Ahles 53762 Native to Mexico and the western U.S.A., natu- ralized in several states of the southeastern U.S.A. previously known from South Caro- lina, reported by Coleman (1974), docu- mented here BRIT.ORG/SIDA 21(2) Vittadinia sulcata N. Burbidge Berkeley Co.:8 Apr 1957, Ahles 22538 es to Australia; first report for the U.S.A.and South Carolina. Xanthium spinosum |. Berkeley Co.:8 Apr 1957, Ahles 22608 Berkeley Co.:20 May 1957, Ahles 25872 Florence Co.: 28 May 1957, Ahles 26689 Berkeley Co.: 30 Oct 1957, Ahles 38199 Berkeley Co.: 13 Jun 1958, Ahles 42924 Native to South America, widely naturalized in North America; reported for South Carolina by Kartesz (1999), documented here. Xanthium strumarium L. var. canadense (P. Miller) Torr. & A. Gray pale ea ns 1957, bile pee weed; cenit owe eon south on (Radford et al. 1968). : it d ACKNOWLEDGMENTS lam grateful for help from the staff of herbarium NCU and for comments on the manuscript by Linda Lee (USCH). John Pruski (MO) provided the identifi- cation of Panphalea heterophylla. REFERENCES ANONYMOUS. 1 June 2004. 997. South Carolina plant atlas. . Accessed Coteman, J.R. 1974. Verbesina section Ximenesia (Compositae): Biosystematics and adap- tive radiation. Amer. J. Bot. 617:25-35, Cronauist, A. 1980. Flora of the southeastern United States. Vol. 1 Carolina Press, Chapel Hill. Asteraceae. Univ.of North Gteason, H.A.and A.Cronauist. 1991. Manual of vascular plants of the northeastern United States and adjacent Canada. New York Botanical Garden, Bronx. Hitt, S.R. and C.N. Horn. 1997. Additions to the flora of South Carolina. Castanea 62: 194-208. Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. First Edition. In: Kartesz, J.T, and C.A.Meacham. Synthesis of the North American flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill, NC. Leona, M.G. 1988. Noteworthy dicots of Mobile and Baldwin counties, Alabama. Sida 13:223-240. MircHett, RS. and G.C. Tucker. 1997. Revised checklist of New York state plants. New York State Museum Bull. 490. New York State Museum, Albany. NESOM MILI 1223 Nesom, G.L. 1999. Gamochaeta simplicicaulis (Asteraceae: Gnaphalieae) in four southeast- ern states and new for North America. Sida 18:1259-1264. Nesom, G.L. 2000. Which non-native plants are included in floristic accounts? Sida 19: 189-193. Nesom, G.L. 2004. New distribution records for Gamochaeta (Asteraceae: Gnaphalieae) in the United States. Sida 21:1175-1185. Preston, R.E. 1997. Dittrichia graveolens (Asteraceae), new to the California weed flora. Mad- rono 44:200—203. Pruski, J. and G.L. Nesom. 2004. Panphalea heterophylla (Compositae: Mutisioideae: Nassauvieae), a genus and species new for the flora of North America. Sida 21: 1225-1227, Raprorb, A.E., H.E. AHLes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill. SHINNERS, L.H. 1966. Hypochoeris microcephala var. albiflora (Compositae) in southeastern Texas: New to North America. Sida 2:393-394. Sorrie, B.A. and P. Somers. 1999. The vascular plants of Massachusetts: a county checklist. Massachusetts Division of Fisheries and Wildlife, Natural Heritage & Endangered Spe- cies Program. Westborough, MA. Tucker, G.C. 1995. The vascular flora of southeastern Connecticut: an annotated checklist. Mem. Connecticut Bot. Soc. 3:1-205. PANPHALEA HETEROPHYLLA (COMPOSITAE: MUTISIOIDEAE: NASSAUVIEAE), A GENUS AND SPECIES NEW FOR THE FLORA OF NORTH AMERICA John F. Pruski Missouri Botanical Garden PO. Box 299 St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT Panphalea peeroE te nee ee is a new genus and species for the United States. This also is the first I e genus from outside of its native range in southern South America. RESUMEN Panphalea heterophylla (Cot i sas y eee nueva para los Estados Unidos. Este es también el primer reporte ade género fuera d urrencia nativa en Sur América. Determinations of Compositae in conjunction with the study of Nesom (2004) identify Ahles & Haesloop 22585 (NCU) as Panphalea heterophylla Less. (Compo- sitae: Mutisioideae: Nassauvieae),a genus and species new for the flora of North America. This is the first known occurrence of Panphalea Lag. from outside of its native range in Argentina, Brazil, Paraguay, and Uruguay and increases to three the number of genera of Nassauvieae known in North America. In this paper I amplify comments on the taxonomic significance of the record, past the summary provided in Nesom (2004), with a description and typification of the species, U.S.A. voucher, anda key to the genera of Mutisioideae that occur in North America north of Mexico. Nassauvieae Cass. were treated as Mutisieae subtribe Nassauviinae (Cass.) Benth. & Hook.f. by Bentham and Hooker (1873) and placed in Compositae subfamily Mutisioideae (Cass,) Lindl. [syn. Nassauvioideae (Cass.) Lindl] by Pruski and Sancho (2004), Nassauvieae were resurrected from synonymy of Mutisieae and reinstated at the tribal level by Pruski (2004) Nassauvieae differ from Mutisieae Cass. by longer style branches with tr ly covered bya tuft of collecting hairs, by anthers with a well-defined appendage and shorter broader thecae, but they have bilabiate corollas and caudate anthers typical of most Mutisioideae [tribe Stifftieae D. Don is characterized by discoid capitula with short, abaxially peop style branches and mutisioid anthers (Pruski 2004)].N tered in South America, but two genera ly known tooccurin North A ica (Si & Anderson 1978): Trixis P. Browne i is largely South American, but extends northward into the West In- dies, Mesoamerica, Mexico, and the southwestern United States, Acourtia D. Don oc- curs from Mesoamerica northwards into the southwestern United States. SIDA 21(2): 1225-1227. 2004 1226 BRIT.ORG/SIDA 21(2) Panphalea heterophylla differs by epappose cypselae from the species of the five genera of Mutisioideae treated by Simpson and Anderson (1978) in North America north of Mexico [Acourtia D. Don (5 spp.), Chaptalia Vent. (4 spp.), Gochnatia H.B.K. (1 sp.), Hecastocleis A. Gray (1 sp.),and Trixis P Browne (2 spp]. Additionally, the combination of the following features serves to further dis- tinguish Panphalea heterophylla from these 13 species of Mutisioideae: annual herbaceous habit; glabrous herbage; leaves basal and cauline, lyrate to sharply lobed; corymbiform capitulescences; few-flowered, bilabiate capitula; thinly chartaceous, apiculate phyllaries; homogamous florets; and bilabiate, isomor- phic corollas with an apically shortly bidentate inner lip. KEY TO MUTISIOIDEAE IN NORTH AMERICA NORTH OF MEXICO 1. Style branches short, apically rounded, glabrous or sparsely papillose;cypselae pa- ppose (Mutisieae). 2. Plants rosulate herbs; capitula bilabiate- ee 2. Plants subshrubs to shrubs; oo a discoid. 3. Leaves abaxially tomentose, not subtended by spiny aT bracts; pollen tricolporate Chaptalia Vent. entire; c Se many-flowered, not glomerate, _ ss Gocchnatia hypoleuca (DC) A. Gray 3. Leaves not abaxially tomentose, at least some leaves with spiny nc pitula uniflorous, glomerate, subtended by spiny subinvolucral bracts; po Hecastocleis shockiey) A. Gray tricolpate Style branches elongate, apically truncate, crowned by a tuft of collecting hairs; Fare pappose or epee uNeenavicae) Glabrous annual t herbs: innel | | lly cypselae Pan phaleas heterophylla Less. 4. Perennial pubescent herbs to shrubs; inner corolla lip bifid, divided to bas cypselae pa 5. Rosulate pereni herbs, caudex commonly tomentose; corollas violet, pink: Acourtia D.Don Trixis P Browne 2 ia) ish, or whit 5 cae to shrubs; corollas yellow Panphalea heterophylla Less., Linnaea 5:8. 1830. Typr: BRAZIL: In humidis ees meridionalis, 1823-1828 [sic], Sello 1328 (HOLOTYPE: B, destroyed, photographs: MOI, NY!, US all photographs from C.B.G.N. negative 16096). Glabrous annual herbs; stems slender, few to several, branched at base, spread- ing to ascending, 5-17 cm long. Leaves basal and cauline, alternate, chartaceous; basal leaves petiolate, oblong to oblong-lyrate, petiole 7-12 mm long, blade or- bicular or more commonly lyrate with about 3 pairs of lobes, 4-15 mm long, 4- 7mm wide, margins entire; cauline leaves commonly sessile and clasping, ovate- lyrate, palmatifid, pedate, or lanceolate, 4-12 mm long, 4-6 mm wide. Capitulescence corymbiform, few-headed; peduncles 10-30 mm long. Capitula 8-12-flowered, bilabiate, homogamous; involucre 1-2-seriate, ca. 2.2 x 2 mm: phyllaries subequal, ovate-lanceolate, thinly chartaceous, apically truncate- apiculate, apiculum to ca. 0.5 mm long, margins broad, hyaline; receptacle PRUSKI, PANPHALEA IN NORTH AMERICA 1227 epaleate. Florets bisexual; corollas bilabiate, isomorphic, white, ca. 5 mm long, weakly puberulent with scattered uniseriate, 5-7-celled trichomes, apical cell larger but neither bulbous nor inflated, tube ca. 2mm long, outer lip ovate, ca. 3 x L5mm, minutely tridentate apically, adaxial epidermal cells longitudinally elongated (mutisioid epidermal pattern), outer wall acute-ridged in transverse section, inner lipca. 1.2 mm long, coiled, undivided proximally, shortly bidentate apically; style branches ca. 0.7 mm long, apically truncate, papillate with many collecting hairs; anthers ca. 2mm long, caudate, apical appendage well-defined, obtuse, ca. 0.5 mm long, tails broadly lanceolate, ca. 0.4 mm long, smooth, tips rounded, not tapered. Cypselae obconical, ca. 1.5 mm long, densely pubescent, apically truncate; pappus absent. Pollen spheroidal, tricolporate, tectum mi- nutely granular (subpsilate). Specimen examined: UNITED STATES. South Carolina. Berkeley Co. aes ae on route 45, waste p 22585 (NCU). Al] ground around the Santee Wool Combing Mill, 8 Apr 1957, Ahles & I Panphalea contains nine species (Cabrera 1953; Cabrera & Klein 1973) found in northeastern Argentina, adjacent Brazil, Paraguay, and Uruguay. Three species are annual herbs, and among these only the smallest, Panphalea heterophylla, is glabrous. Cypselae of this species were presumably imported into the United States in fleece brought from South America for processing. Species of tribe Nassauvieae are not commonly weedy, and it is unlikely that this subtropical species will become a pest, if it has persisted at all in South Carolina. ACKNOWLEDGMENT I would like to thank Dra. Gisela Sancho (LP) for helpful discussion. REFERENCES BentHam, G. and J.D. Hooker. 1873. Compositae. In: Genera Plantarum, vol. 2, part |. Reeve and Co., London. Pp. 163-533. Caprera, A.L. 1953. Las especies del género Pamphalea [sic] (Compositae). Notas Mus. La Plata 16:225-237. Caprera, A.L.and R.M.Ktein. 1973. Compostas, 1 tribo:Mutisieae.In:PR. Reitz, ed. Flora Illustrada Catarinense, | Parte: As Plantas, Fasc. COMP:1—124 Nesom, G.L.2004. Asteraceae from wool mill sites in South Carolina, including new records from North America. Sida 21:1215-1223. Pruski, J.F. 2004. Missouri Botanical Garden, Research: Asteraceae (Compositae). Missouri Botanical Garden, St. Louis. Pruski, JF. and G. SANcHo. 2004. Asteraceae or Compositae (Aster or Sunflower family). In: Smith, N.P, S.V. Heald, A. Henderson, $.A.Mori,and D.W. Stevenson, eds. Flowering plants of the Neotropics. Princeton University Press, Princeton., NJ. Pp. 33-39 + pl. 5. Simpson, B. and C. Anperson. 1978. Compositae tribe Mutisieae. North Amer. Flora, ser. 2, LO 13: CUSCUTA INDECORA (CONVOLVULACEAE) NEW FOR CANADA Mihai Costco! Francois J. Tardif Department of Plant Agricultur Department of Plant Agriculture University pp CuE lah University of Guelp Guelph, Ontario, NIG 2W1, CANADA Guelph, Ontario, NIG 2W1, CANADA coste_amihai@hotmail.com ftardif@uoguelph.ca Vernon L. Harms WP. Fraser Herbarium Department of Plant Sciences University of Saskatchewan, 51 Campus Dr, Saskatoon Saskatchewan, S7N 5A8, CANADA vharms@shaw.ca Cuscuta indecora Choisy is frequent in southern US.A. (Yuncker 1932, 1965) and extends northward to Michigan, Minnesota, North Dakota, Montana and Idaho (USDA NRCS 2004). The species al urs in Mexico, Central and South America (Yuncker 1932, 1965). We are reporting here the first record of this spe- cies for southern Saskatchewan, Canada. Its presence in the Canadian Provinces adjacent to its northern range was to be expected, and further floristic research may reveal its occurrence in southern Alberta, Manitoba and Ontario. Although C. indecora has definite weedy tendencies (Parker & Riches 1993), taking into account the natural habitats it was found in, the species may be considered na- tive to southern Canada as well. The species was cultivated experimentally in Ontario and Alberta (vouchers at DAO and ALTA, respectively), but it has never spread (see also Crins & Ford 1988). The host range of C. indecora is wide, in- cluding numerous herbaceous and woody plants (Yuncker 1965). Cuscuta indecora and C.coryli Engelm from subsection Indecorae Yuncker are probably sister species (Costea et al. unpublished). Cuscuta coryli in Canada, is rare in Saskatchewan, Manitoba, Ontario and Québec (Argus & Pryer 1990). Both species have in common the fleshy, papillate flowers with corolla lobes acute and inflexed. Cuscuta indecora differs from C. coryli mainly through the 5-merous flowers, 2—5 mm long, having usually large and abundantly fringed intrastaminal scales. Cuscuta coryli has flowers commonly 4-merous, ca. 2mm long, and infrastaminal scales are reduced to winged or toothed ridges. Descrip- tions, identification keys and iconography can be found in Yuncker (1932, 1965). — se specimens: CANADA. Say etchenats oe of Rosetown- cae iS anton Leu ca see mi of Demaine, ca. 50° 40' N, 107° 20°'W la d, 30 ‘Corres rrespondi ng author ee SIDA 21(2): 1229-1230. 2004 1230 BRIT.ORG/SIDA 21(2) Jul 1956, B. Boivin et al. 11648 (SASK- 2 collections; ALTA, DAO, MT, NY); Crooked Lake Provincial Park, 50° 35'N, 102° 40'W, in bog on sow thistle and sunflower, 12 Aug 1976, VJ. Huffen s.n. (SASK). ACKNOWLEDGMENTS Appreciation is extended to Luc Brouillet and an anonymous reviewer for their helpful suggestions on an earlier version of the manuscript. We thank direc- tors/curators from ACAD, ALTA, DAO, HAM, MT, MTMG, NEFLD, NSPM, NY, OAC, QFA, QUE, RBG, SASK, SFS, TUPR UBC, UNB, USAS, UWO, UWPG, WAT, WIN, and WIS for the herbarium loans. REFERENCES Arcus, G.W. and K.M. Prayer. 1990. Rare vascular plants in Canada: our national heritage. Canadian Museum of Nature, Ottawa. Crins, WJ. and B.A. Foro. 1988. The parasitic dodders (Cuscuta; Cuscutaceae) in Ontario. Canad. Field-Nat. 102:209-215. Parker, C.and CR. Riches 1993. Parasitic weeds of the world. Biology and control. CAB Inter- national, Wallingford, UK. USDA, NRCS. 2004. The PLANTS Database, Version 3.5 (plants.usda.gov). National Plant Data Center, Baton Rouge, LA. Yuncker, T.G.1932.The genus Cuscuta. Mem. Torrey Bot. Club 18:113-331. Yuncxer, 1.6. 1965. Cuscuta. North American flora, Ser. 2,4:1—51. CONTRIBUTIONS TO THE MOSS FLORA OF THE AMAZONIAN LOWLANDS OF MADRE DE DIOS, PERU Piers Majestyk John P. Janovec Institute of Systematic Botany Botanical Research Institute of Texas (BRIT) The New York Botanical Garden (NY) 09 Pecan St. 200th St. and Kazimiroff Blvd Fort Worth, Texas 76102-4060, U.S.A. Bronx, New York 10458, U.S.A. jjanovec@brit.org pmajestyk@nybg.org (corresponding author) ABSTRACT A recent collection of mosses from the Department of Madre a Pio in the low lance.< of alae Peru contains 58 different species. Fifty-four taxa are new new to Peru. Previous collections from the district are listed which brings the total species ina n from the area to 74. This is the first major study of mosses from Madre de Dios, Peru, and it provides a foundation for continued work and production of a field guide for the region. RESUMEN Una coleccion reciente de m ee del Departamento de Madre de Dios en la zona baja del Sureste de Pertt contiene 58 ntes. Cincuenta y cuatro taxa son nuevos para el distrito y se citan cuatro especies nuevas para Pert. Se listan las col pe ias pace listrito que elevan el total de especies conocidas del area a 74. Este es el primer di gos de Madre de Dios, Peru cree bo See aa ] J | ed A ‘i ees tes 5 ces bs la region INTRODUCTION The southwestern Amazon of the trinational corner of SW Brazil, NW Bolivia, and SE Peru contains what is probably the largest and least disturbed area re- maining of Upper Amazonian and Lower Andean ecosystems (Foster et al. 1994). Threats to the forest occur in the form of hunting, gold mining, timber extrac- tion, impending road construction, and slash and burn agriculture. The Depart- ment of Madre de Dios (12°-14.5°S), Peru, dominated by the Madre de Dios River basin and tributaries, is an important geopolitical region in the pristine SW Amazon, which is divided equally between lowland, premontane, and mon- tane forest (Davis et al. 1997). Madre de Dios is located at the southwestern edge of the Amazon basin near the Andean foothills and is covered primarily by lowland tropical/subtropical moist forest (Gentry & Le6n 1997). The region has a distinct six month dry season and receives 2000-2500 mm of rain annually (Gentry & Leén 1997). Madre de Dios has been recognized as an epicenter of botanical diversity and holds world record numbers of other organisms, such as birds, tabanid flies, tiger beetles, damselflies and dragonflies, and butterflies (Stewart 1988). Botanical research over the past 25 years in Madre de Dios, Peru, has been SIDA 21(2): 1231-1238. 2004 1232 BRIT.ORG/SIDA 21(2) important for identifying the extent and significance of the region’s plant diver- sity, producing various publications and datasets (e.g., Gentry 1982; Phillips @ Gentry 1993a-b; Foster et al. 1994; Terborgh & Andersen 1998; Pitman et al. 1999, 2001). There have been two geographic areas of botanical investigations in Madre de Dios, with most of the resources focused on quantitative inventory of trees (trunk diameter > 10 cm) in scattered one-hectare forest plots and 0.1-hectare transects. The Cocha Cashu Biological Station in Manu National Park has been the focus of significant field research during the last three decades (Terborgh 1994), producing one of the most comprehensive plant checklists for the region. The Tambopata-Candamo Reserved Zone (Foster et al. 1994) in lower Madre de Dios near the Peru-Bolivian border has been the focus of long-term monitoring of one- hectare forest plots and complementary O.l-hectare transect work (Phillips et al. 2003), plus associated programs of general botanical collection. Despite the research that has been focused on these selected areas, there has been no major, long-term systematic inventory of plants and habitats, and no flora has been published for the region. Between July ll and August 19, 2002, as part of a long-term program of botanical inventory and monitoring, the au- thors conducted expeditions in the Amazonian lowlands of the Department of Madre de Dios. Led by the first author, cryptogamic collections were made from several localities in the region (Table 1, Fig. 1). Fifty-eight species of mosses were identified from approximately 226 collections. Of the 58 taxa identified, 54 are new records for the district and four of these are new records for Peru (Table 2). The only previously published records of moss collections for Madre de Dios, as far as we know, were made by Vargas (1974) and recorded in a checklist of mosses for Peru by Menzel (1992). Vouchers of the Vargas material were searched for but not located in either of the two larger Peruvian herbaria—the Herbarium of the Museo de Historia Natural, Universidad Nacional Mayor de San Marcos (USM) and the Herbario Vargas, Facultad de Ciencia Bioldgicas, Universidad Nacional San Antonio Abad del Cuzco (CUZ). We conclude that future work should be focused on comparative studies of the diversity of mosses and other bryophytes in time and space between differ- ent sites in the lowlands of Madre de Dios, and along the altitudinal gradient from the lowlands up the slopes of the Andes into the Department of Cusco. This work is produced as a component of long-term botanical exploration and collaboration in the Andes-Amazon region of southeastern Peruand it will serve as a foundation for the production of field guides to serve the multidisciplinary science, education, and conservation programs that are ongoing in the region. ANNOTATED LIST OF SPECIES Species are listed alphabetically by family. New records for the country are marked with an asterisk (*). The nomenclature for the species follows Crosby et al. (2000). Voucher collections are deposited at NY. Duplicates of some taxa are MAJESTYK AND JANOVEC, MOSS FLORA OF THE AMAZONIAN LOWLANDS IN PERU 1233 Taste 1. Summary of nine collection sites in Madre de Dios, Peru, and the corresponding moss col- lections made between July 11 and August 19, 2002. Locality Elevation Geographic Coordinates Collection Numbers (P. Majestyk) 1.Manu Province, Puerto Maldonado 300m 12°36' S,69°11'W 3720-3727 2.Manu Province, Los Amigos field 280m 12° 34'S, 70° 06'W 3728-3902; 3951.5-41 78; station, ca. 3 km N of the jct of the 4301-4364 Madre de Dios and Los Amigos rivers. 3. Manu Province, Community of 220m 12° 36'S, 70° 05'W 3903-3951 Boca Amigo, ca 1.6 km N of the jct of the Los Amigos and Madre de Dios rivers on the W side of the river 4. Tambopata Province, E side of the 300m 12° 28'50"S, 4179-4209 Los Amigos River 70° 11°01" W 5.Tambopata Province, E side of the 300m 12° 27' 48'S, 4210-4215 Los Ami igos 5 River, small waterfall 70° 13' 09" W emptying into r 6. Tambopata acne Eside ofthe 300m 12°25) 37'S, 4718-4231 Los Amigos River 70° 15' 35" W 7.Tambopata Province, E side of the = 230-250m_ 12° 31'52"- 4240-4265 Los Amigos River, 12° 32' 05"S from banks of river to inland 70° 05' 24"- 70° 05' 29"W 8 Manu Province, Inambari River ca. 220m 12°43'87" S, 4266-4267 2 km from jct of Madre de Dios River 69°45'51" W 9. Manu Province, ca. 13 km W of 300 m 12°40' S,69°20' W 4768-4300 Puerto Maldonado in small park dominated by Mauritia flexuosa L (Arecaceae). also deposited at BRIT, CUZ, MO, SMF and the first author's personal herbarium. Numbers following the species name are the first author's collection numbers. Bartramiaceae *Bryum renauldii ROll ex Renauld & Cardot; 4212 Philonotis uncinata (Schwagrichen) Bridel; 3743, 3820, 3876, Calymperaceae 3882, 3885, 3997, 4145, 4165, 4180, Calymperes afzelii Swartz; 3948, 4205, 4303, 4310, 4213,4215, 4231, 4267 4314 Brachytheciaceae Calymperes palisotii Schwagrichen; 4336 Rhynchostegium conchophyllum (Taylor) A. Jae- Aue eee Dozy & Molkenboer; ger; 4220 Sthopoton eae Schwagrichen; 3838, Bruchiaceae TF raatoue nilonaicolia Nicneoae4s 2, 3985, 4017, 4018, 4108, 4110, 4141, er 4264, 4301 Bryaceae Syrrhopodon parasiticus (Swartz ex Bridel) Bryum apiculatum Schwagrichen,; 3875, 3888, Bescherelle: 4038, 4309 3995, 4134,4151 1234 BRIT.ORG/SIDA 21(2) 04.5.9 18 27 30. ee Kilometers Dr a7) Uy 6 “ ““Ilos om @5 mY cal va (*) +, © . - rf } : b) 4 2497 Q q 4 (alk, : SO! o OAL o> ! & —— 3° Pe = VV ~, 9 5 a § ) w : r C9 iy of o~ .*) a : on" Fegpbe Seg aan "Mad : ae ae he cS > Pres $ f 6 1 : , oa Fic. 1 AA I L . g . Tl 4° . L L ly rs I RA j inf; Peru Table 1 provides descrip- tions of +h I tee L y L i° g L of eact 5 e Dicranaceae acta 1 tenerum (Swartz) Mitten; 375 1,3775, Dicranella hilariana prety Mitten; 3722, 3780, 3828, 3852, 3973, 4004, 4277, 4286, 430? 3728, 3730, 3781, 4102, 4130, 4197 stenornamn um reptans (Hedwig) Cardot; Ditrichaceae 4338,4 fheeonienes trinitensis (C. Muller) E. Britton & Dixon; 4169, 4252 Ditrichum rufescens (Hampe) Hampe; 4178, 4202 Fissidentaceae Sea gulanensis Montagne; 3939, 4023, bee 041.5,4061, 4062, 4096, 4100, 4101,4109 41 4125,4195, 4262 Fissidens flaccidus Mitten; 4211, 4226 Fissidens eee vatus (Hampe) Mitten; 3750 Fissidens pellucidus Hornschuch; 3756, 388 1, 4246 Fissidens prionodes Montagne; 3777, 3783, 3966, Vesi cularia sp.;4344,.We were unable to identity collection from South American mate- el in NY. This species has branch leaves measuring ca. 2.5mm in length. It is perhaps a new species but we cannot without fur- ther study of the genus make that decision now. Vesicularia vesicularis (Scwagrichen) Brotherus Mess . 7 var. vesicularis; 3784, 3877, 4072, 4104, 4183 Fissidens submarginatus Bruch; 3873, 4138,4259 —— Vesicularia vesicularis var. rutilans (Bridel) WAR. Fissidens zollingeri Montagne; 3871, 3928, 4132 Buck: 4064 Leucobryaceae Hypnaceae Leucobryum martianum ee Hampe Chryso-hypnum diminutivum (Hampe) W.R. Buck; ne Muller; 3847, 3861,4149, 41534159 4167, 4188 an ; bo tum (Hampe) Brot! : (Brotherus) B.H. F : *Le anne subobtusifolium 4067,4116,4192 Allen: 4092 MAJESTYK AND JANOVEC, MOSS FLORA OF THE AMAZONIAN LOWLANDS IN PERU 1235 Taste 2. A summary of moss collections made by family, species, and localities in Madre de Dios, Peru. Family Number of Species Number of Collections — Collection Localities (see map, Fig. 1) Bartramiaceae ] 13 1,2,4,5,6,8 Brachytheciaceae 1 1 6 Bruchiaceae 1 1 7 ace 2 6 2 Calymperaceae 5 23 24,7 Dicranaceae 1 7 1,2,4 Ditrichaceae 1 2 ; Fissidentaceae 7 3] 2,3,4,5,6,7 Hypnaceae 8 25 Leucobryace 5 28 2,4,7 Meteoriaceae 2 9 2,3,4,7 Neckeraceae 2 7 2 Othotrichaceae 2 2 2 Pilotrichacea 4 18 24 Pottiacea 2 2 6 Pterobryaceae 2 5 2,3 Sematophyllaceae v/ 23 2,3,4,7,9 Splachnobryaceae ] 2 2 Stereophyllaceae 1 9 2 Thuidiace 5 12 2 tal 55 212 Octoblepharum albidum Hedwig; 3738, 3755, 3778, 3859, 3965, 3969, 4013, 4081, 4121, 4155, 201, 4299 4291, 4253 Octoblepharum cylindricum W.P. Schimper ex Montagne; 4075 Octoblepharum pulvinatum (Dozy & Molkenboer) Mitten; 3874, 3961, 4005,4107, 4161 Meteoriaceae | (Hed ig)M |. 3724, 3917, 4099 4115,4117, 4200, 4204, 4254 Zelometeorium recurvifolium (Hornschuch in Martius) Manuel; 395 Neckeraceae ae disticha (Hedwig) Kindberg; 3935, Mocken DSi ta (Hedwig) Reichardt; 3762, 3909, 4078, 4085, 4249 Orthotrichaceae Groutiella tomentosa (Hornschuch) Wijk & Margadant; 38 Schlotheimia rugifolia (WJ. Hooker) Schwa- grichen; 4307 Pilotrichacea Callicostella pal ee (Hornschuch in Martius) Angstrém; 3757, 3759, 3869, 3872, 3968, 4026, 4097, 4103, 4136, 4184, 4186, 4203, 4315, 4322 oa ium beet te - Muller; 4787 uiller oaae surinamense e Nace 3752 Pottiaceae Barbula indica (W.J. Hooker) Sprengel in Steudel; 23 Dolotortula mniifolia (Sullivant) R.H. Zander; 4225 Pterobryaceae Henicodium geniculatum (Mitten) W.R. Buck; 3930, 3988, 4127, 4137 Pireella pohlii (Schwagrichen) Cardot; 4152 Sematophyllaceae *Potamium lonchophyllum (Montagne) Mitten; 425] 1236 *Sematophyllum adnatum 89 SOS NO Michx.) E. Britton; BRIT.ORG/SIDA 21(2) eophyllacea a chl ch Jum (Hornschuch) C. Muller Sematophyllum subpinnatum (Bridel) E. Britton; in Brotherus; 3956, 4003, 4012, 4034, 4037, 3/ 4087, 4256, 4305, 4312 um subsimplex toe a 2, 3960, 3967, 4008, 4025, 4 SS Thuidiaceae Cyrto-hypnum involvens (Hedwig) W.R. Buck & H. een planum (Bridel) Mitten; 833, 3934, Crum: 3938, 4124,4147,4198 Ses Hla Eade Anse Abaca heal Cyrto-hypnum leptocladum (Taylor) W.R. Buck & Trichosteleum cf. pusillum (Hornschuch) A. Jae er aS Cyrto-hypnum scabrosulum (Mitten) W.R. Buck & Trichosteleum subdemissum (Schimper ex H. Crum; 3761, 3857, 3951, 4010, 4084, 4128, Bescherelle) A. Jaeger; 4123, 4129, 4279 Splachnobryaceae Splachnobryum obtusum (Bridel) C. Muller; 3996, 3998 Taxa recorded in Vargas (1974): Anomobryum julaceum (Schrader ex PG. Gartner, B. Meyer & Scherbius) WP. Schimper, Bryum apiculatum Schwagrichen, Dicranella hilariana (Montagne) Mitten, Entodon jamesonii (Taylor) Mitten, Leucobryum martianum (Hornschuch) Hampe ex C. Miller, Philonotis longiseta (Michx.) Britton, P unci- nata (Schwagrichen) Bridel, Pilopogon guadalupensis (Bridel) Frahm , Thamnobryum fasciculatum (Hedwig) lL. Sastre, Sematopyllum subpinnatum (Bridel) E. Britton, Sphagnum subsecundum Nees in Sturm, Squamidium nigricans (WJ. Hooker in Kunth) Brotherus. Taxa listed as occurring in Madre de Dios in the Tropicos database with collec- tor and collection number (mobot.mobot.org/W3T/Search/most.html): Fissidens allionti Brotherus (Matthews B-86580, MO); Fissidens lagenarius Mit- ten (Matthews B-86577, COLO, PAC); Fissidens perfalcatus Brotherus (Matthews B-85860, MO), Phyllogonium viscosum (P. Beauv.) Mitten (Chavez 857, MO). ACKNOWLEDGMENTS We would like to thank the Gordon and Betty Moore Foundation and the Bene- ficia Foundation for their support of botanical studies in Madre de Dios, Peru. The first author thanks William Buck and the New York Botanical Garden for financial assistance which made this trip possible. The second author thanks Scott Mori for support during postdoctoral studies at the New York Botanical Garden. For aiding in the identification of several collections we thank Will- iam Buck, Robert Ireland, and Ron Pursell. We thank Asha McElfish for her as- sistance in preparation of tables and for helpful comments to improve the origi- nal manuscript. And we thank Barney Lipscomb for his excellent editorship and the two reviewers (Robert E. Magill and William R. Buck) who helped us improve the manuscript. Fieldwork would not have been possible without the MAJESTYK AND JANOVEC, MOSS FLORA OF THE AMAZONIAN LOWLANDS IN PERU 1237 logistical support provided by the Amazon Conservation Association (ACA), the Asociacion para la Conservacion de la Cuenca Amazonica (ACCA), and the staff of the Los Amigos Biological Station, including Jorge Herrera, Juan Carlos Flores, Giovana Espino, and Sonia Guerra. We also thank Asuncioén Cano of USM, Carlos Reynel of MOL, and staff of the Peruvian Ministry of Natural Resources (INRENA) for their support of our ongoing botanical research in Peru. Fernando Cornejo, Piher Maceda, Mathias Tobler, and Euridice Honorio pro- vided essential assistance and collaboration during the field expeditions that made moss collections possible. REFERENCES Crossy, MLR., R.E. Maaitt, B. ALLEN, and S. He.2000.A checklist of the mosses. Missouri Botani- cal Garden, St. Louis. Dawis, S.D., V.H. Hevywoon, O. HerreRA-MacBryoe, J. VitA-Losos, and A.C. Hamitton. 1997.The Ameri- cas. Volume 3.V.H. Heywood and S.D. Davis, eds.Centres of plant diversity, a guide and strategy for their conservation. World Wildlife Fund for Nature (WWF) and IUCN-The World Conservation Union. Foster,R., LA. Parker Ill, A.H. Gentry, L.H. Emmons, A. CHICCHON, T. SCHULENBERG, L. RoDRIGUEZ, G. LAMAS, H. Orteca, J. OcocHea, W. Wust, M. Romo, J. A. Castitto, O. PHILLIPS, C. REYNeL, A. KRATTER, P. K. Donaxue, and L. J. Barktey. 1994. The Tambopata-Candamo Reserved Zone of southeast- ern Peru: A biological assessment. RAP Working Papers, Vol 6. Conservation Interna- tional. Gentry, A.H. 1982. Patterns of neotropical plant species diversity. Evol. Biol. 15:1-84. Gentry, A.H. and B. Leon. 1997. Tambopata region, Peru. In: S.D. Davis, V.H. Heywood, O. Her- rera-MacBryde, J. Villa-Lobos, and A.C. Hamilton, eds. Centres of plant diversity:a guide and strategy for their conservation, Volume 3—The Americas. WWF and IUCN, Oxford, England. Pp. 355-259. Menzet, M. 1992. Preliminary checklist of the mosses of Peru (Studies on Peruvian bryo- phytes IV). J. Hattori Bot. Lab. 71:1 75-254. Puities, O. and A.H. Gentry. 1993a. The useful plants of Tambopata, Peru: |. Statistical hy- potheses tests with a new quantitative technique. Econ. Bot.47:15-32. Puities, O.and A.H. Gentry. 1993b. The useful plants of Tambopata, Peru: II. Additional hy- potheses testing in quantitative ethnobotany. Econ. Bot.47:15-32. Puicties, O.L., R. Vasquez Martinez, P Nunez Varcas, A. Lorenzo Monteacupo, M.-E. CHuspe ZANS, W. GALIANO SANCHEZ, A. Penta Cruz, M. Timand, M.Yu-HaLta, and S. Rose. 2003. Efficient plot-based tropical forest floristic assessment. J. Trop. Ecol. 19:629-645. Pitman, N.C.A., J.W. Tersorch, M.R. Sian, and P. Nunez V. 1999. Tree species distributions in an upper Amazonian forest. Ecology 80:2651—2661. Pitman, N.C.A., J.W. TersorcH, M.R. Sitman, P. Nunez V., D.A. Nei, C.E. Ceron, WA. PALacios, and M. Autestia. 2001. Dominance and distribution of tree species in upper Amazonian terra firme forests. Ecology 82:2101-2117. 1238 BRIT.ORG/SIDA 21(2) Stewart, P.D. 1988. Tambopata Reserved Zone, southeast Peru. Oryx 22:95-99, Tersorcu, J. 1994. An overview of research at the Cocha Cashu Biological Station. In: A.H. Gentry, ed. Four Neotropical Rainforests. Yale University Press, New Haven and London. Pp. 48-59, TerborGH, J.and E. Anberson. 1998. The composition of Amazonian forests: patterns of local and regional scales. J. Trop. Ecol. 14:645-664. Varaas, C. 1974. La flora del departamento de Madre de Dios (Pert). Lima, Peru. KATHLEEN M. PETERSON 1948-2003 James L. Reveal! University of Maryland College Park, Maryland 20742, U.S.A. The New York Botanical Garden Bronx, New York 10458-5126, U.S.A. = = ABSTRACT Kathleen M. Peterson (1948- 2003) 1 is remembered as a student, gifted teacher, skilled botanist, law- | aria (Lamiaceae) from Mexico named yer, and friend allasani of anew species of Scutel 1 her honor KEY WORDS: Scutellaria, Lamiaceae, Mexico, obituary RESUMEN Kathleen M. Peterson (1948-2003 lad studiante, prof talento, botanica de pericia, eee y sida ag como una ere de una nueva especie de Scutellaria (Lamiacede) de Méxi 1 su honor. The notice in the Harrisburg, Pennsylvania, paper, the Patriot-Ne ws, published on November 26, 2003 was short: “Kathleen M. Peterson, 55, of Camp Hill, died Monday, November 24, 2003 at Hospice of Central Pennsylvania, Carolyn Croxton Slane Residence. She was an attorney for the Pennsylvania Treasury Department. Services will be held at the convenience of the family. Arrange- ments are by the Cremation Society of Pennsylvania, Harrisburg.” Nothing was mentioned of the person or her life as a botanist; we hope our words and actions in proposing Scutellaria petersoniae (Turner & Reveal 2004) in some way, will rectify the oversight for it is with sadness that we note to the botanical commu- nity the passing of Kathy Margaret Peterson. Kathy Peterson wasa graduate student at the University of Maryland from 1973 until 1978, where she received her doctoral degree, working on a portion of the mint genus Salvia. Her master’s thesis was done with Dr. Willard Payne (The genus Hymenoclea (Compositae: Ambrosieae), Brittonia 25:243-256. 1973) at the University of Illinois (1969-1971) where she took her undergraduate de- gree in 1969. The two of them also published “Observations of the hypoder- mises of ferns” in the American Fern Journal (63:34-42. 1973). Based on her work at Maryland, she published “Salvia reptans. Labiatae” with Raymond M. Harley of the Royal Botanical Garden, Kew, England, in Curtis’s Botanical Magazine Mailing address: 18625 Spring Canyon Road, Montrose, Colorado 81401-7906, U.S.A. SIDA 21(2): 1239-1243. 2004 1240 BRIT.ORG/SIDA 21(2) (182:13-16. 1978). As in all papers published in that journal, their comments were accompanied by a beautiful, full-page colored painting of the species. Her research on Salvia was supported by a grant from the National Geo- graphic Society and a Doctoral Dissertation Improvement Grant from the Na- tional Science Foundation. Together, she and Reveal presented a report entitled “Biosystematic and evolutionary studies of Salvia subgenus Calosphace” that was published in the Society’s Research Reports (15:557-564. 1983). Of a more personal nature was an article of their travels in Mexico “Exploring the green frontier” by Broome, Reveal and Peterson published in the University of Maryland’s Graduate School Chronicle 10(4):3-6. 1977). C. Rose Broome and Kathy spent a month collecting prior to being joined by Reveal and Harley at the silver mining town of Toluca. Together they explored a portion of Guerrero that had not been visited botanically for several decades. The new finds were numerous. Among them were Bouvardia hintoniorum, Galium iltisii, Hansteinia harleyi, Montanoa revealii, Salvia turneri, Sinclairia broomeae, Tibouchina patens, and a new genus Revealia. The most beautiful addition made during their joint trip across the Sierra Madre del Sur was a wonderful new species named Utricularia petersoniae by Peter Taylor (Kew Bull. 41:8.1986). It was a fitting tribute to Kathy, tall and slen- der yet graceful with an elegant beauty. Kathy was born in Oak Park, Illinois, on March 10, 1948. Here she spent her youth, attending local grade and high schools before entering the University of Illinois in 1965. There she majored in botany with minors in chemistry and ge- ology. Asa youngster she was tall, taller than most boys her age, and this would continue to be the case throughout her life. She had elegance about her, borne by her height so that in the classroom she was a presence. At Maryland she tu- tored struggling athletes, and was most effective as she could intimidate the largest football player and look many a basketball player in the eye. Her wit was appreciated but her strength was asa teacher. In 1978 she received the Car- roll E. Cox award as the outstanding graduate student, during the 1978-1979 academic year she was appointed an instructor at College Park. She departed soon thereafter to accept a position as an assistant professor at the University of Montana. Her skill in the classroom continued, for Kathy taught a variety of classes and soon had a small group of students working on graduate degrees, being named “KP” to distinguish her from the other Kathys in the group. Aug- mented by grants from the National Science Foundation and National Geo- graphic Society she continued her work on Salvia. Other funding allowed her and her students to work on various rare and endangered species in Montana. Among her students were Jeffrey Strachan now with the Plant Variety Protec- tion Office, U.S. Department of Agriculture, Lisa Campbell of The New York Botanical Garden, Jennifer Ramstetter, a professor of biology at Marlboro Col- lege,and Kathy Ahlenslager of the U.S. Forest Service who wrote a recent obitu- — REVEAL, KATHLEEN M. PETERSON 1241 ary (Ahlenslager 2004). Perhaps her greatest contribution to the University of Montana came in the form of her effort to upgrade the herbarium by hiring a collections manager, finding volunteers to mount and file thousands of plant specimens, and to distribute a large backlog accumulated over the decades. Professionally Kathy moved upward. Besides her teaching duties she di- rected the operations of the University’s natural history museum, wrote sev- eral research papers, served on the editorial board of Systematic Botany, and was president of the University of Montana Sigma Xi chapter. In 1984 she was promoted to the rank of associate professor, and in 1986 she received tenue. Along with tenure the University awarded her a “meritorious teaching and re- search salary increase.” Most of her publications at this time dealt with rare plant species in Montana. They include “Vascular plants of limited distribu- tion in Montana” (Lesica, P, G. Moore, K.M. Peterson, and J.H. Rumley, Montana Acad. Sci. Monogr. 2:1-61.1984) and a summary report on rare plants in the state (Peterson, K.M., P Lesica &J.S. Shelly, Proceedings of the 1986 Natural Areas Con- ference, pp. 97-113). Then, in 1987, she suddenly resigned her position, left Montana, and botany as well, all for what proved to be an unfortunate, short-lived marriage which ended formally in divorce in 1994. Finding herself in Pennsylvania, Kathy began a new career. She was certi- fied as a paralegal by Pennsylvania State University in March of 1989 and then attended The Dickinson School of Law where she was awarded her J.D. in 1992. During this time she was a member of the Dickinson Law Review and the Ap- pellate Moot Court Board; she also served on the editorial board of the Dickin- son Journal of Environmental Law and Policy. In the Review she published an article entitled “Federal regulation of artificial insemination donor screening practices: An opportunity for law to co-evolve with medicine,” a title only a bi- ologist would think of. Her interest initially was in environmental law, work- ing as an intern for the Department of Environmental Resources (1991-1992) and as a law clerk for the state’s Treasury Department. In August of 1992, two months before being admitted to the Pennsylvania Bar, she became the Assis- tant General Counsel for Treasury. Asa lawyer, Kathy drafted banking service contracts between the Treasury and banks, prepared white papers on State Workers’ Insurance Fund and the Fed- eral Reserve System, did general legal research on banking and ethics issues, and provided litigation support for the General Counsel. In September of 1995 she was promoted to the Department’s Associate General Counsel. In this position, she represented the State Treasurer on various committees, including the Board of Directors of Pennsylvania's chapter of The Nature Conservancy. In 1999, Kathy discovered that she had ovarian cancer. Initially the combi- nation of chemotherapy and radiation was successful, and during most of this time she continued to serve the citizens of Pennsylvania. In 2001 her doctors BRIT.ORG/SIDA 21(2) 1242 48-2003). \ Kathleen M Peterson (19 REVEAL, KATHLEEN M. PETERSON 1243 found a lesion on the brain. It was removed surgically, and she returned to work determined to beat the newly found cancer. Two weeks after surgery she fell and a MRI found more lesions. A second round of surgery was performed, but her chances of survival were greatly diminished. Up to the end, Kathy was a strong, stalwart woman, loosing a battle to cancer that two and half years early claimed her brother, and months before her own death, took her mother. It is in Kathy’s memory that we dedicate the following new species. ACKNOWLEDGMENTS We are grateful to Carl Peterson, Kathy Ahlenslager and Jeff Strachan for pro- viding us with insights into Kathy’s life. REFERENCES AHLENSLAGER, K.2004. Kathy Peterson (1948-2003). Friends Univ.Montana Herb. Spring 2004:1, Turner, B.L. and J.L. Reveat. 2004. Scutellaria petersoniae (Lamiaceae), a new species from Guerrero, Mexico. Sida 21:679-681., BOOK REVIEWS 1245 BOOK REVIEWS ANpREW ParKER. 2003. In the Blink of an Eye. (ISBN 0-465-05438-2 pbk.). Basic Books, 387 Park Avenue S., New York, N.Y. 10016, U.S.A. (Orders: 800-255- 1514 or special. marl p books.com). $15.00, 316 pp., b/w figures, index, 5'* 3’: he new book In The Blink Of An Eye, author Andrew Parker puts a new spin on why animals first developed body armor during the Cambrian era. Parker contends that the cee and sudden development of hard body coverings about 543 million y g take advantage of the simultaneous development of higher- heey vision in some animals. It was usef a in attracting mates, and fighting off enemies, and being ive predator. This book takes the reader on an ne: expedition, through numerous fossil beds ane to examine creatures living today throughout the world to discover the role of color and light in the evolution of Cambrian-era ae pa presents a poweunl: ane ev nies packed allen armament suggesting the cause of th and distinct shapes sea 543 mallicnt years ago, as the onset of organisms otaducug true vision and the ability to pursue prey. opening chapters present a well- eae mini “history of life” book and discussion of n appearance of and the living environ- how fossils are investigated to give ment of those fossil organisms. These introductory ee are helpful for the reader to get a good understanding of the argument about to be discussed. Following the background chapters, a good deal of information and numerous interesting examples are presented on the importance of visual appearance and behavior of various creatures as a response to light. For instance, exam ples included multilayer reflectors of Messel fossil beetles and Hercules beetles that alter the color of their appear- ance as humidity conditions change within layers of their body. Adaptations discussed included pig- ments, structural colors (such as iridescent film on wings), natural liquid crystals, diffraction, re- flection layers and broltiminescenes all of which cause specific visual appearances. The uses of these clude camouflage, warning coloration, defense, and mating attractants. Infor- mation ang examples aes included both fossil and extant organi idence. Many of the ad- can only en if light is present. Therefore, many of the color, s es and be- havior oradaptaton ae give ae as to the environment an organism could be f th he Cambrian explosion i is supported by evidence of the ae evolu- tion of the eye in a short period of time (geologically speaking), and the ability of organisms to see their environment and more oe (their) prey. It was determined possible for true eyes, which not only bee perceive light and darkness, but form a true visual image in which the eens can ae its surroundings, could evolve over less than half a million years. In addition, - t nce for a ae theory is the occurrence of soft-bodied trilobites with no eyes in the Pre re-Cambrian, then the abundance of trilobites with eyes and body armor in the Cambrian. The development of the first true eyes was followed by a rapid evolution of body color, body structure and behavior; all of these increased the capacity to be a predator and the avoidance of becoming prey through the formation of body armor and deterrents such as spines. The author does leave the reader with a parting question: “What prompted the development of the eye in the first place?” Andrew Parker has presented a well-organized argument for vision as a trigger for the Cam- brian explosion. He managed to explain the theory in a logical pattern while using both an educa- tional and entertaining writing style. However, because each chapter entails so many details, | rec- ommend you read each chapter beginning to end nonstop; doing so helps keep the ideas together. visual < SIDA 21(2): 1245. 2004 1246 BRIT.ORG/SIDA 21(2) O 5 lle plai tistl il reference to image plates that are not in the book. These plates would have enhanced the inderseandiie of the information presented. This book is recommended lor those interested in the Cambrian age, evolution, the development of vision, and body structural colors in insects and aquatic life —Lee Luckeydoo, Botanical Research Institute of Texas, 509 Pecan Street, kort Worth, TX 76102-4060, U.S.A. Grape Man of Texas SHERRIE S. McLeRoy and Roy E. ReNrro, Jr. 2004. Grape Man of Texas: The Life of T.V. Munson. (ISBN |-57168-819-6, hbk.) Eakin Press, PO. Drawer 90159, Austin, Texas 78709-0159, U.S.A. (Orders: 512-288-1771; 800-880-8642: 512-288-1813 fax; Service and Sales: sales@eakinpress.com). $39.95, 296 pp., color and b/w figures, 7" = 10" Grape Man of Texas: The Life of T.V. Munson is the first biography written about this eminent Texas horticulturist. He became one of the leading experts in native American grape specie ies, and his stud- 1 } ] ies WETC instr umental wine indus uly from disasterin tne tate nin c teenth century. Munson d vel 7 >d ov 300 n new grapes but is perhaps best known for his efforts in fig Merite Agricole and inducted into the French Legion of Honor. He was also a leader in the vinicul- iting the phyll oxera epidemic of the late 19th century, for which he received the Chevalier du ture movement of the period for his cutting-edge work in breeding new grape and plant varieties. His articles, experiments, correspondence, speeches, exhibits, grape classifications, and numerous in- ventions reveal his scientific thirst for knowledge and his sleek interests. Over 100 years ago Cognac, France and ears Texas shared two ol it’s citizens with the world tosolve a major destructive force—tl t ,pn phy yllo oxera. It was 1880 and the phyllox- era plague was rampant in France, arene in the Chi irante ‘Region where Cognac is located. The phy the economy of France. France assigned the preeminent French scientist, Pierre Viala the task of loxera root louse was destroying the prestigious French winegrapes and therefore, destroying finding a cure for the phylloxera plague. This investigation brought him to the United States and to the home of Thomas Volney Munson in Denison, Texas. The two scientists collaborated on the prob- lem for several days in Denison and various other locations in Texas to view the native grapes of i; Texas in their carnal habitat. The spe the C ales in France, and Denison are very similar anc therefore should support gra] S ble of growing in either location. T.V. Munson suggested that the only way to save the F euch abeyatdls was to graft the Mess vin ae ra varieties to resistant rootstocks. Munson knew that the Texas rootstocks were resistant to ] t his suggestion, Pierre Viala agreed that it was a good plan of action. Thousands ol bundles of Texas rootstocks were shipped to France where Pierre Viala and other French scientists started educating the French wine- growers on the enormous ee: tasks that lay ahead. The grafting literally continues to this day. For hi France, T.V. Munson was awarded the highest award that could be given a foreign civil = ued du Merite eeelcoe and was inducted into the | pepe of Honor in 1888. In 1898 he ig g member of the Societe Nationale d’Agriculture de France and as an honorary member in ite Societe des Viticulteurs de France. Sev- eral statues honoring Munson have been erected in France. arly 300 pages long, the Grape Man of lexasincludes more than 100 illustrations, many never belore published; the first listing compiled since Munson’s death of his 300+ grape h ybrids and their parentages, a list of his wild grape discoveries, several of which remain important in modern viticulture and research; and a list of all of Munson’s known speeches and publications. SIDA 21(2): 1246. 2004 BOOK REVIEWS 1247 Released in June 2004, Grape Man of Texas soon received its first international accolade when it was awarded a prestigious Uanens stars from the Gourmand World Cook book pyres which recog- nize the finest in i d wine books. T1 his summer to rank the 4,000-plus books entered each yeae in its “Best in 1 the World” eompeniion: Gourmand defines these 3-star books as “masterpieces—buy and read immediately.” Grape Man, the Gourmand press release states, is “a very important book for wine acl [tis one of only four Ameri- can books and 19 worldwide to be honored on the inaugural lis The book was co-written by award-winning Texas ae and historian Sherrie S. McLeRoy and by Dr. Roy E. Renfro, Jr, director of the T.V. Munson Viticulture and Enology Center in Denison, Texas.—Gary L. Jennings, Librarian, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, ct Preservation of the Big Thicket James J. CoZINE, JR. 2004. Saving the Big Thicket: From Exploration to Preserva- tion, 1685-2003. (ISBN 1-57441-175-6, hbk.). University of North Texas Press, PO. Box 311336, Denton, TX 76203-1336, U.S.A. (Orders: 1-800-826-8911, 940- 565-4590 fax, rchrisman@unt.edu, www.unt.edu/untpress). $34.95, 272pp, 5 maps, 25 illustrations, 6" x 9". Saving the Big Thicket: From Exploration to Preservation, 1685-2003 is the fourth in the Temple Big Thicket Series pune ms U —e of North Texas Press. Originally written by Cozine in 1976 as part of a dissertation li d Assault on a Wilderness, the work has been updated to the present time and now includes a foreword and afterword by Pete A.Y. Gunter, author of another book in this series, The Big Thicket: An Ecological Reevaluation. Saving the Big Thicket is an account of the regional history and “play-by- play” or the political fight lor the ee Thicket Nenone: Preserve (BT NP) with the afterword by Gunter ’s original 1976 manuscript was published. The foreword by Gunter describes the work of the author and his involvement and persever- ance on this work, presents a general construction of the book and provides some insight into the process of constructing an overview of a region with such a long and complicated history. Introduc- tory material gives a brief description of the units that make up the preserve, general ecosystems and plant diversity found in each, along with information about the creation of the units, trails and structures. Two maps from Cozine’s ont work accompany this first section. Cozine divides the history of the region into six time periods of human involvement: early Native American use, French and Spanish arrival and attempted settlement, the coming of early American backwoods settlers, initiation of sae railroads (for large-scale a . aa oil exploration, the beginning of an early conservation m ent and finally the formation of the second Big Thicket Association and creation of BTNP. Baus section prov ides en cise and historical information on the people, orga- nizations and industries that with the land, how they used it, settled it (or attempted to), helped destroy it and eventually helped save it. The first few ae involve pre-industry hu- man effects on the Big Thicket, outlining the Native American tribes that lived in the area leaving a minimal impact, the arrival of the Spanish and French explorers that interacted with the Native American tribes, and finally the ae impact of backwoodsmen, escaped slaves, criminals and hunters seeking refuge in the Big Thicket. The following chapters address the arrival of timber rail- re ncredible fe damage to the region that came with them, a aa ways, oil exploration and t SIDA 21(2): 1247. 2004 1248 BRIT.ORG/SIDA 21(2) the rise and fall of the original East Texas Big Thicket Association who first raised the environmen- tal flag in the Big Thicket and the eventual creation of the Big Thicket National Preserve with the aid of the second (and successful) Big Thicket Association in 1974. Each of these chapters is riddled with political and social struggles among the groups involved and with the tale of destruction and hu- man impact on the region’s incredible biodiversity. Throughout the book there are a total of 5 maps from Cozine’s 1976 publication. The quality of these maps is that of the original publication era and they are somewhat difficult to interpret. There is also a pcan: section ContalNnS 19 black and white photographs from Cozine’s original work. These ly 1900's of bear hunts and early scenes in the big thicket, but the section camels contains portraits and casual photographs of prominent politicians, activists, industry representatives and other people involved in the history of the area. These photographs are of poor quality and add little useful information or ene to the book, besides presenting a _— record of the prominent figures in the Big Thicket’s hist e afterword by Gunter is a well-written account of the pda environmental and park management struggles since the creation of the park. Itis a detailed account of complications a have arisen since the initial creation of the BTNP. Gunter divides this afterword into five sections addressing the bureaucratic beginnings of the park, the acquisition of new units and river property, further development of the park and facilities, and what Gunter calls a “New Big Thicket Crisis” about a struggle to add controversial new tracts of land, and finally what he calls a “postscript to a postscript” introducing, but not expounding on, peripheral issues affecting the Big Thicket. Each section discussed above ends with a set of notes and references and a thorough index makes for easy location of subjects, organizations and persons mentioned in this book. Saving the Big Thicket provides a well-written historical account of the Big Thicket and is a good addition to the UNT Press series on the region. This work could be useful to historians, scientific researchers, resto- ration project managers sets -nts, and the casual reader alike interested in the region’s complex and “entang oe as Hayley Gillespie, University of Texas at Austin, Integrative Biology, Austin, 1X — 78712, | leye ie@mail.utexas.edu SIDA 21(2): 1248. 2004 his illustrated monograph is the most comprehensive generic treatment of legumes for any geographical area of Mexico. The book covers the genera of legumes known to be present in northern Mexico from Tamaulipas to Baja California Sur. The introductory material covers topography. climate, and vegetation types of northern Mexico. Included are 121 genera treated in three families: and Caesalpiniaceae. The Fabaceae is the most abundant family with 87 genera followed by Mimosaceae (19) and Caesalpiniaceae (15). Keys to the families, tribes (only in Fabaceae), genera, and detailed line drawings of distin- guishing characteristics are included for most of the genera to facilitate identification. Each genus is provided with a description, followed by distributional information, pecies number, chromosome number, and economic, ecological, and potential food importance. Available from Botanical Research Institute of Texas Press 9 Pecan Street Fort Worth, TX 76102-4060, USA E-mail: sida@brit.org * Fax 1-817-332-4112 www.brit.org/Sida/Sida25.htm Sida, Bot. Misc. No. 25, 2004 issn 0833-1475, isbn 1-889878-13-8=7" « 10",x + 134 pp, over 500 individual drawings. $25 + p&h = (USA $3; International $7) Texas residents add $2.06 sales tax Fabaceae, Mimosaceae, Los Géneros de Leguminosas del Norte de México Los Géneros de Leguminosas del Norte de México (The Genera of Legumes of Northern Mexico) By A. Eduardo Estrada C. and Alfonso Martinez M. ¢ BRIT PRESS he Atlas of Texas covers about 6000 taxa. This is the result of 54 years of herbarium and fieldwork by B.L. Turner, beginning in 1948 at Sul Ross State University, Alpine, Texas. In short, the senior author has examined personally, touched, or “pored over” an estimated several hundred thousand sheets in the preparation of the forth- as volumes. Contents include an introduction, coming Atl atlas of Texas plants arranged alphabetically by by genus, by species, and an index. family, Available from Botanical Research Institute of Texas Press 509 Pecan Street Fort Worth, TX 76102-4060, USA E-mail: sida@brit.org * Fax 1-817-332-4112 www.brit.org/Sida/Sida/24.htm Sida, Bot. Misc. No. 24, 2003, issn 0833-1475, isbn (vol. 1) 1-889878-08-1 = isbn (vol. 2) 1-8 *USA:$10 (vol. 1); $9.50 (vol. 2); $12 (set) *International: $12.50 (vol. 1) «$11.50 (vol. 2); $25 (set) Texas residents add sales tax:$4.13 9878-09-X = 7 1/2" * 10" Atlas of the Vascular Plants of Texas By B.L. Turner, Holly Nichols, Geoffrey Denny, ¢ (vol. 1) = $3.30 (vol. 2) « $7.43 (set) BRIT PRESS a RKANSAS FLOR DA (OWA MIs sts ExICO CAL IF OF YUISIANA TEXAS. GE ORGIA oo tA NEW. MEX NDLAND -MANATOBA ou | ERICA a CHILE | ee ONDU URAS, GUYANA | DIA “As BURMA : BAN a4 ‘UROPE. Ve STR. RAL I Annas A GREECE PAKISTAN: SYR A Bey salar PAPU a NE SU ENEA. NEPAL L SYR IA A 4 ut EENSLAND. VICTORIA WES . JSTRAL AN |ORTHE RN TERRN ORY VG COE _ AUSTRALIA A NEW ZEA i SUMATRA 8 ru JAYANA ISRAEL MOROCCO SOUTH 4 A EGYPT | NIGERIA i. OP A s¢ A LIBYA JORL DON KENYA CONC » St OZAMBIQUE ZAMBIA SAUDI Al iw a j NICO fe i ONTEREY : MICHOACAN: OA u { IDALGO GUADALAJARA Ni _COAHL ILA YUCATAN SIN Lu. ° U c =) ° “ Barney L. Lipscoms, EDITOR Botanical Research Institute of Texas 909 Pecan Street Fort Worth, Texas 76102-4060, USA 817 332-4441 / 817 332-4112 FAX Electronic mail: sida@brit.org Home page at the URL: http://www.brit.org/sida/ SIDA CONTRIBUTIONS TO BOTANY WAS FOUNDED BY JOHN W. THIERET, Associate EDITOR Ltoyo H. SHINNERS (LEFT) Dept. of Biological Sciences Northern Kentucky University IN 1962. INHERITED BY Highland Heights, KY 41099 USA WILLIAM F. MAHLER (RIGHT), Director EMERITUS FELIX LLAMAS, CONTRIBUTING SPANISH EDITOR oF BRIT in 1971, AND Dpto. de Botanica, Facultad de Biologia Universidad de Leon SINCE 1993, IT HAS BEEN E-247] Leon, SPAIN PUBLISHED BY Brit Press. oes . [he views expressed in this journal do not necessarily reflect those of the editors or of the Botanical Research Institute of Texas Guidelines for contributors are available upon request and on our SIDA home page as well as the last pages of each volume. Cc. ) Subscriptions for year 206 $37. Individual $65. USA Institutions $75.Outside USA numbers issued twice a yeal OSIDA, CONTRIBUTIONS TO BOTANY VOLUME 21, NUMBER 3, PAGES 1249-1954, 5 Au 2005 Copy tT 2005 T BOTANICAL RESEARCH INSTITUTE OF TEXA PRESS PRINTED IN THE UNITED STATES OF AMERICA ISSN 0036-1488 TABLE OF CONTENTS SYSTEMATICS A new species of Sabatia (Gentianaceae) from Saline County, Arkansas JAMES S. PRINGLE AND THEO WITSELL Aulonemia nitida (Poaceae: Bambusoideae: Bambuseae), a new species from Guyana EMMET J. JUDZIEWICZ Four new species of Ericaceae (Vaccinieae) from Ecuador JAMES L. LUTEYN Nomenclature of Ipomoea arborescens (Convolvulaceae) in Sonora, Mexico DANIEL F AUSTIN, RICHARD FELGER, AND THOMAS R. VAN DEVENDER Ipomoea seaania, a new species of Convolvulaceae from Sonora, Mexico RICHARD FELGER AND DANIEL F. AUSTIN Infrageneric classification of Liatris (Asteraceae: Eupatorieae) Guy L. Nesom Broadened concept of Liatris helleri (Asteraceae: Eupatorieae) Guy L. Nesom A revision of the Psidium grandifolium complex (Myrtaceae) LesLiE R. LANDRUM Ptilagrostis luquensis (Poaceae: Pooideae: Stipeae: Stipinae), a new species from China PauL M. Peterson, Rosert J. SORENG, AND ZHEN-LAN Wu Eragrostis (Poaceae: Chloridoideae: Eragrostideae: Eragrostidinae) from northeastern México Paut M. PETERSON AND JESUS V ALDES-REYNA A revision of Trisetum and Graphephorum (Poaceae: Pooideae: Aveninae) in North America north of México Victor L. Finot, PAuL M. PETERSON, ROBERT J. SORENG, AND FERNANDO O. ZULOAGA New combinations in Festuca californica (Poaceae) STEPHEN J. DARBYSHIRE Eugenia inversa (Myrtaceae), a new species from Espirito Santo, Brazil Marcos SOBRAL A multivariate morphometric study of the aster genus Sericocarpus (Asteraceae: Astereae) MICHELLE R. LEONARD, RACHEL E. COOK, AND JOHN C. SEMPLE Neotypification of Ceropegia palustris and Lyonia maritima (Apocynaceae: Asclepiadoideae) ALEXANDER KRINGS 1249 1263 1269 1283 1293 1305 1323 1335 1355 1363 1419 1455 1465 1471 1507 A new combination in Matelea (Apocynaceae: Asclepiadoideae) for an endemic Jamaican vine ALEXANDER KRINGS A new species of Matelea (Apocynaceae: Asclepiadoideae) from Hispaniola ALEXANDER KRINGS hem laa} snare ast Noteson Asclepiadoideae: Gonolobinae) i in the Greater Antilles and Bahamas ALEXANDER KRINGS igrina COTT 1plex (Apocy TlaACCAC, Estudios en las Apocynaceae neotropicales X: definicion de la verdadera identidad de Mandevilla leptophylla, con la descripcion de dos nuevas especies y una nueva combinacién de Mandevilla (Apocynoideae: Mesechiteae) para Sur América J. FRANCISCO MORALES Estudios en las Apocynaceae neotropicales XI: una nueva especie de Mandevilla (Apocynoideae: | Mesechiteae) para Sur América, con un nuevo reporte para las Apocynaceae de Paraguay J. FRANCISCO MORALES New species and new combinations in Ericameria (Asteraceae: Astereae) ROLAND P. Roperts, LOWELL E. UrBAtsCcH, AND JOHN ANDERSON Taxonomic revision of the neotropical genus: Erithalis (Rubiaceae: Chiococceae VIVIAN NEGRON-ORTIZ Una nueva especie de Aiphanes (Arecaceae) de la Cordillera de Mérida, Venezuela SANTOS MIGUEL NINo, LJ. Dorr, AND FRED W. STAUFFER Three previously undescribed species of Vaccinium (Ericaceae) from Costa Rica and Panama Ropert L. WILBUR AND JAMES L. LUTEYN Cuniculotinus and Lorandersonia, two new genera of Asteraceae: Astereae and new combinations in Chrysothamnus Lowe E. Ursatscu, ROLAND P. Roperts, AND Kurt M. NEuBIG Symphyotrichum pygmaeum: transier of Eurybia pygmaea from the eurybioid grade to the subtribe Symphyotrichinae (Asteraceae: Astereae) Luc BROUILLET AND SUGIRTHINI SELLIAH New combinations in Artemisia (Asteraceae: Anthemideae) LeitA M. SHULTz New combination in Conoclinium (Asteraceae: Eupatorieae) THOMAS FE. PATTERSON 1515 1519 1525 1535 1549 1557 1565 1599 1607 1615 1633 1637 1641 Cc : \ A new combination in the genus Packera (Asteraceae Desra K. TROCK Correct author citation for Cirsium eatonii var. eriocephalum (Asteraceae: Cardueae): erratum for Sida 21:212. 2004 Davin J. KEIL Nestotus and Toiyabea, two new genera of Asteraceae: Astereae from the western United States and Canada RoLAND P. Roperts, LOWELL E. URBATSCH, AND Kurt M. NEuBIG CHROMOSOME NUMBERS Chromosome numbers of Glandularia (Verbenaceae) from central and Trans-Pecos, Texas B.L. TURNER AND A. MICHAEL POWELL Documented chromosome numbers 2005: 1. Miscellaneous counts from western Texas, mostly Trans-Pecos A. MICHAEL POWELL AND B.L. TURNER Documented chromosome numbers 2005: 2. Counts from western Texas, mostly Trans-Pecos cacti A. MicHAEL POWELL AND JAMES F WEEDIN Documented cl numbers 2005: 3.Chromosome numbers in North Ameri pecies of Silene and Stellaria (Car yophyllaceae) J.K. Morton DEVELOPMENT AND STRUCTURE Morfologia de tricomas foliares en especies de Solanum secci6n Acanthophora (Solanaceae), presentes en Venezuela WILLIAM A. GRANADA-CHACON Y CARMEN E. BENITEZ DE ROJAS FLORISTICS, ECOLOGY, AND CONSERVATION Introduced species in Kansas: floristic changes and patterns of collection based on an historical herbarium TeresA M. Woops, SUZANNE C. STRAKOSH, MADHAV P. NEPAL, SEEMANTI CHAKRABARTI, NICHOLAS B. SIMPSON, MARK H. MAYFIELD, AND CAROLYN J. FERGUSON The vascular plants of a forest fragment in southern Bahia, Brazil Anpre M. Amorim, PEDRO FIASCHI, JOMAR G. JARDIM, Wm. Wayt THOMAS, BRENDA C. CLIFTON, AND ANDRE Mauricio V. DE CARVALHOT The vascular flora of Rattlesnake Falls: a potential state natural area on the Western Highland Rim Escarpment in Tennessee Dwayne ESTES AND JEFFREY L. WALCK Endemic vascular plants of the Interior Highlands, U.S.A. DoucLas ZOLLNER, MicHAEL H. MacRoserts, BARBARA R. MACROBERTS, AND DouGLas LADD 1643 1645 1647 1657 1663 1665 1669 1675 1695 1727 1753 1781 Reference conditions of the Red River floodplain and upland, Caddo Parish, Louisiana MICHAEL H. MACROBERTS AND BARBARA R,. MACROBERTS Annotated checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin counties, Texas LARRY E. Brown, BARBARA R. MACRoserts, MICHAEL H. MACRopeRTS, PAUL A. HARCOMBE, WARREN W Pruess, I. SANDRA ELSIK, AND DAN JOHNSON Checklist of the vascular plants of Greene County, Pennsylvania Rosert Coxe, CYNTHIA M. Morton, Mary Joy Haywoop, Bonnie L. ISAAC, AND JOSEPH A. ISAAC Vegetation and flora of American Beech Woods Nature Preserve, Clark County, Illinois Bos EpdGIN, GORDON C. TUCKER, AND JOHN E. EBINGER Soil and ecological features of Hexalectris (Orchidaceae) sites ANNIE B. CoLLins, JAMES E. VARNUM, AND MARGARET BROWN-MARSDEN The ecology of Trillium texanum (Trilliaceae) on the Angelina National Forest, Texas Michare H. MACRoserTS AND BARBARA R. MACRoBERTS Dichanthium (Poaceae) new to Arizona: open door for a potentially invasive species RICHARD FELGER, TONy L. BURGESS, SAMUEL Dorsi, JOHN R. REEDER, AND THomaAS R. VAN DEVENDER State records and other noteworthy collections for Kentucky Ross C. CLARK, RONALD L. JONEs, TIMOTHY J. WECKMAN, RALPH L. THOMPSON, JOHN W. THIERET, KENTUCKY STATE NATURE PRESERVES COMMISSION, AND KIM FEEMAN Rediscovery of Ponthieva brittoniae (Orchidaceae) in Everglades National Park Jimi L. SADLE, STEVEN W. WOODMANSEE, GEORGE D. GANN, AND THOMAS V. ARMENTANO Quercus montana (Fagaceae), new to Missouri MARIAN SMITH AND NANCY PARKER Sporobolus heterolepis (Poaceae), new to Tennessee DwayNe Estrs AND JOHN BECK A second population of Agalinis navasotensis (Scrophulariaceae) confirmed from Tyler County, Texas MoniQue DuBRULE REED, JOHN F Hays, JupitH M. CANNE-HILLIKER, DANA PRICE, JASON SINGHURST, AND Eric KEITH Gibasis pellucida (Commelinaceae), a new and potentially weedy genus and species for Texas Davib J. ROSEN AND Ropert B. FADEN 1793 1807 1829 1861 1879 1893 1905 1909 1917 1921 1923 1927 1931 Stemodia coahuilensis (Scrophulariaceae), a new record for the United States B.L. TURNER 1935 Emilia fosbergii (Asteraceae: Senecionae) in Texas revisited WP. SPENCER AND J.K. WILLIAMS 1937 A new record of the devil’s cigar, Chorioactis geaster (Pezizales: Ascomycota), from Collin County, Texas JOHN E. UBELAKER AND JANE K. STARKS 1939 COMMEMORATIVE Wilbur Howard Duncan, 1910-2005 Wenpy B. ZOMLEFER AND Davib E. GIANNASI 1941 Book reviews AND NoTices 1268, 1304, 1322, 1334, 1454, 1464, 1470, 1506, 1514, 1518, 1524, 1534, 1548, 1556, 1636, 1640, 1644, 1646, 1656, 1662, 1726, 1792, 1828, 1860, 1892, 1904, 1930, 1936, 1951 Announcements 1321, 1333 Index to new names and new combinations in Sida 21(3), 2005 Aiphanes stergiosii M. Nino, Dorr & FW Stauffer, sp. nov—1600 Artemisia arbuscula subsp. longiloba (Osterhout) Shultz, comb. nov—1637 Artemisia globularia subsp. lutea (Hultén) Shultz, comb. nov—1638 Aulonemia nitida Judz., sp. nov—1263 Ceratostema oyacachiensis Luteyn, sp. n0ov—1271 Ceratostema pendens Luteyn, sp. nov—1272 Ceratostema pubescens Luteyn, sp. nov—1275 Chrysothamnus scopulorum (M.F. Jones) Urbatsch, R.P. Roberts & Neubig, comb. nov.—1626 Chrysothamnus scopulorum var. canonis (S.L. Welch) Urbatsch, RP. Roberts & Neubig, comb. nov—1627 Chrysothamnus stylosus (Eastwood) Urbatsch, R.P. Roberts & Neubig, comb. nov.—1627 Cirsium eatonii var. eriocephalum (A. Gray) DJ. Keil, comb. nov—1645 Conoclinium betonicifolium var. integrifolium (A. Gray) T.F. Patterson, comb. nov.—1641 Cuniculotinus Urbatsch, R.P. Roberts & Neubig, gen. nov—1618 Cuniculotinus gramineus (H.M. Hall) Urbatsch, R.P. Roberts & Neubig, comb. nov.—1619 Disterigma bracteatum Luteyn, sp. nov—1279 Ericameria arizonica RP. Roberts, Urbatsch & J. Anderson sp. nov—1558 Ericameria linearis (Rydberg) R.P. Roberts & Urbatsch, comb. nov—1560 Ericameria winwardii (R.D. Dorn & CH. Delmatier) R.P Roberts @ Urbatsch, stat. NOV—1562 Eugenia inversa Sobral, sp. nov—1465 Festuca californica subsp. hitchcockiana (E.B. Alexeev) S.J. Darbyshire, comb. et stat. nov.—1461 Festuca californica subsp. parishii (Piper) S.J. Darbyshire, comb. nov—1461 Ipomoea seaania Felger & Austin, sp. nov—1296 Liatris series Elegantes (Alexander) Gaiser ex Nesom, comb. et stat. nov—1312 Liatris series Garberae Nesom, ser. nov—1316 Liatris series Graminifoliae Gaiser ex Nesom, ser. nov—1317 Liatris section Graminifolium Nesom, sect. nov—1315 Liatris series Pauciflorae (Alexander) Gaiser ex Nesom, comb. et stat. nov—1315 Liatris section Pilifilis Nesom, sect. nov—1314 Liatris series Virgatae Nesom, ser. nov—1316 Liatris section Vorago Nesom, sect. nov—1312 Lorandersonia Urbatsch, R.P. Roberts @ Neubig, gen. nov—1619 Lorandersonia baileyi (Wooton & Standley) Urbatsch, R-P. Roberts & Neubig, comb. nov—1621 Lorandersonia linifolia (Greene) Urbatsch, RP. Roberts @ Neubig, comb. nov—1622 Lorandersonia microcephala (Cronquist) Urbatsch, R.P. Roberts & Neubig, comb. nov.— 1622 Lorandersonia peirsonii (D.D. Keck) Urbatsch, R.P. Roberts @ Neubig, comb. nov.i—1623 Lorandersonia pulchella (A. Gray) Urbatsch, R.P Roberts & Neubig, comb. nov—1624 Lorandersonia salicina (S.E Blake) Urbatsch, R.P Roberts & Neubig, comb. nov—1624 Lorandersonia spathulata (L.C. Anderson) Urbatsch, R.P. Roberts & Neubig, comb. nov—1625 Mandevilla amazonica J.) Morales, sp. nov—1536 Mandevilla colombiana J. Morales, sp. nov.—1538 Mandevilla gracilis (Kunth) J.-F Morales, comb. nov—1541 Mandevilla matogrossana J.F. Morales, sp. nov—1551 Matelea pentactina Krings, sp. nov—1519 Matelea rhamnifolia (Griseb.) Krings, comb. nov—1515 Nestotus R.P. Roberts, Urbatsch & Neubig, gen. nov—1650 Nestotus macleanii (Brandegee) R.P. Roberts, Urbatsch @ Neubig, comb. nov—1651 Nestotus stenophyllus (A. Gray in Torrey) R.P. Roberts, Urbatsch & Neubig, comb. nov.—1652 Packera musiniensis (S.L. Welsh) Trock, comb. nov—1643 Psidium australe var. argenteum (O. Berg) Landrum, comb. nov.—1342 Psidium australe var. suffruticosum (O. Berg) Landrum, comb. nov—1344 Ptilagrostis luquensis P.M. Peterson, Soreng & Z.L. Wu, sp. nov—1356 Sabatia arkansana J.S. Pringle @ CT. Witsell, sp. nov—1250 Symphyotrichum pygmaeum (Lind|.) Brouillet & S. Selliah, comb. nov—1635 Toiyabea R.P. Roberts, Urbatsch & Neubig, gen. nov—1652 Toiyabea alpina (L.C. Anderson & S. Goodrich) R.P. Roberts, Urbatsch & Neubig, comb. nov.—1653 Vaccinium almedae Wilbur & Luteyn, sp. nov—1607 Vaccinium furfuraceum Wilbur & Luteyn, sp. nov—1609 Vaccinium luteynii Wilbur, sp. nov—1611 A NEW SPECIES OF SABATIA (GENTIANACEAE) FROM SALINE COUNTY, ARKANSAS James S. Pringle Theo Witsell Royal Botanical Gardens Arkansas Natural Heritage Commission P.O, Box 399 1500 Tower Building Hamilton, Ontario enter St. CANADA L8N 3H8 Little Rock, Arkansas 72201, U.S.A. jpringle@rbg.ca theo@arkansasheritage.org ABSTRACT Sabatia arkansana, a new species from shale and igneous glades in central Arkansas, is described. It differs from S. campestris in its narrower leaves and narrower, more deeply colored corolla lobes rounded at the apex. RESUMEN Se describe Sabatia arkansana, especie nueva de claros de esquistos y de rocas igneas en Arkansas central. Se diferencia de S. campestris por las hojas mas angostas y por los lébulos de la corola mas angostos, coloreados mas intensamente, y redondeados en el apice. HISTORY OF THE DISCOVERY In June 2001, while collecting plants for the Flora of Arkansas Project and for his Master’s thesis, the junior author was shown a small shale glade (Womble Formation) northwest of Owensville, Saline County, Arkansas, by nature pho- tographer and amateur botanist John Pelton. Pelton, a long-time student of the flora of Arkansas and of Saline County in particular, was especially interested in showing Witsell plants of a Sabatia that he was unable to identify to species. Witsell collected specimens of this plant, as well as specimens of Sabatia campestris Nutt., which occurred in the same area. Studies using all relevant botanical references available at the Arkansas Natural Heritage Commission and the University of Arkansas at Fayetteville Herbarium (UARK) supported the idea that this species was significantly dif- ferent from any described in the literature. In any of the regionally appropriate keys, specimens of this plant keyed out to S. campestris. However, seeing S. campestris and this new species occurring at the same site made the differences between the two obvious (Fig. 3). Conversations with the landowner at this site revealed the existence of another, larger glade opening 0.2 mile (0.3 km) northwest of the known glade. This glade was impressive in its botanical diversity and in that two intermit- tent spring-fed streams flow into the glade and join before flowing out of the glade downslope. More plants of this new Sabatia were found in this opening, SIDA 21(3): 1249-1262. 2005 1250 BRIT.ORG/SIDA 21(3) which should be considered part of the same complex rather than a completely separate site. In June 2002, while conducting surveys for the rare small-headed pipewort (Eriocaulon kornickianum) on igneous glades near Bauxite, Saline County, Ar- kansas, Witsell and Pelton located several more populations of this Sabatia. Witsell compared the Saline County specimens with Sabatia specimens from a broader region at the herbarium of the Missouri Botanical Garden (MO) in July 2002. When no matches were found at MO, the Saline County speci- mens were sent to the senior author for confirmation that this was indeed a species new to science. DESCRIPTION OF THE SPECIES Sabatia arkansana JS. Pringle & CT. Witsell, sp. nov. (Figs. 1-5). Type: ARKANSAS: Saline Co.: Alcoa Lake Glades Natural Area, 3.5 mi (5.6 km) SE of Bauxite, NE 1/4 of SW 1/4 of Section 26, T2S R14 W, northernmost glade opening on W shore of lake, 18 Jun 2002, Witsell 02-0832 (HOLOTYPE: UARK; ISOTYPE: MO). CHhagtaca + cf: Vfrlise] } ] ] Tes 1; a (pe Ps spathulatis apices versus rotundatis differens. ° Annuals; roots fibrous. Plants 7-25 cm tall, single-stemmed at base. Stems + terete but narrowly 4-winged; proximal diameter 0.4-1.3 mm, wings ca. 0.1-0.2 mm high; branching generally alternate (rarely opposite proximally), mostly at 30-50°, lowest branching ca. mid-height. Proximal internodes 0.8-2 x as long as subtending leaves, distal internodes 0.6-1 x as long. Leaves all cauline, bright green, membranous, spreading, linear to lanceolate; most leaves 7-30 X 1-4.5(- 6) mm; bases narrowed, not clasping, apices obtuse (proximal leaves) to acute or acuminate (mid-stem and distal leaves): leaves smaller near base of stem, lowermost leaves minute. Inflorescence a + corymboid, monochasial cyme; flowers 1-12(-18) per plant. Pedicels (2-)10-40 mm. Perianth and androecium pentamerous. Calyx (0.7-)1.0-L4 as long as corolla; tube turbinate, green along commissural veins, otherwise whitish and nearly hyaline, 2.8-5.5 mm, 0.7-0.9x as long as corolla tube; ridges present along commissural veins of tube, trian- gular in cross-section, rounded or distally acute along apex but not winged, widening from 0.2-0.5 mmat base to 0.5-0.8 mm at summit of tube, where there is sometimes a minute protuberance, dividing into lower ridges along submar- ginal veins of lobes; lobes green, spreading at 60-90°, nearly filiform to linear, 9-13 x 0.6-1.2 mm, apices acute. Corolla deep but bright magenta-pink, with an oblong-triangular, sometimes apically notched yellow zone extending from tube into each lobe along midrib, alternating with shorter, yellowish-white zones flanking sinuses; tube 3-7 mm; lobes narrowly spatulate-obovate, 8-18 x 3-6 mm, apices rounded. Filaments (3-)4-5.5 mm, ca. 1.5 as long as uncoiled anthers; anthers yellow, 2.3-3.5 mm long before coiling. Pollen grains subprolate, 33-38 um long, 27-34 um in diameter. Uncleft portion of style 2-3.5 mm, style PRINGLE AND WITSELL, A NEW SPECIES OF SABATIA FROM ARKANSAS 1251 a ‘ ; & D) << @ a NX Fig. 1. Sabatia arkansana, holotype. branches plus stigmas when uncoiled (3-)4.5-6 mm, L.7-2.5xX as long as uncleft portion. Paratypes: ARKANSAS: Saline Co.: Bauxite, sandy outwash on rock outcrop, 6 Jun 1950, Moore 50-0161 (UARK); ee and date uncertain (see note below), Moore 50-0168 (UARK);, Womble Shale glade on W side of Burk Rd. NW of Owensville, 21 Jun 2001, Witsell 01-0474; roe capi Lake Glade, 2.5 mi (4.0 = ee of Bauxite, N 1/2 of NW 1/4 of NW 1/4 of Section 23, T25 R14W, 18 Jun 2002, Witsell 02-0829, Womble Shale glade E of Burk Road and W of the Middle Fork of a Saline River, NW of Owensville, NE 1/4 of SE 1/4 of Section 17, TIS RI7W, 18 Jun 2002, Witsell 02-0825: International w BRIT.ORG/SIDA 21(3) 1252 Fic. 2. Sabatia arkansana. A. Plant (Witsell 02-0829, UARK). B. Flower, w with petals removed, anthers newly dehisced and the stigma not yet receptive wel 02- Obs uaa: C rower aia view with stamens and sepals removed (Witsel! 02-0832, UARK).D. Capsule, largely i Nitsell 02-1125, UARK). Paper Pipewort Glades Preserve [pow Dunnahoo Preserve, The Nature Conservancy], 3.5 mi (5.6 km) SE of Bauxite, NE 1/2 of NE 1/4 of Section 34, T25 R14W, 19 Jun 2002, Witsell 02-0838; Dry Lost Creek Glades, 1.75 mi (2.8 km) SE of Bauxite, E 1/2 of SE 1/4 of Section 21, T2S RI4W, 19 Jun 2002, Witsell 02-0840; oo. Shale g] rr of Burk Road and W of the Middle Fork of the Saline River, NW ol Owensville, N 4 of SE 1/4 of Section 17, TIS RI7W, 10 Aug 2002, Witsell 02-1125. Unless Arkansas N ] Heritage Commission — otherwise indicz ited, specimens are in the herbarium ol the PRINGLE AND WITSELL, A NEW SPECIES OF SABATIA FROM ARKANSAS 1253 The two specimens collected by D.M. Moore in 1950 were originally identified as Sabatia campestris. Moore 50-0161 was labeled as being collected at Bauxite 6 June. Moore 50-0168, although bearing a higher number, was labeled as being collected 15 miles south of Little Rock, near Ferguson Lake, the previous day. Ferguson Lake is in extreme southeastern Saline County, on unconsolidated Tertiary and Quaternary sediments (Haley et al. 1976). No rock outcrops or glades—that is, no suitable habitats for S.arkansana—are known from the area. It is unlikely that S.arkansana actually occurred at this site, and it may be that specimens from the two sites were mixed up. The genus Sabatia Adans. and its sections and subsections have been de- scribed by Wilbur (1955). Sabatia arkansana is unequivocally a species of sect. Campestria J.D. Perry. This section, which is well defined morphologically and isolated genetically, is most readily recognizable by its calyx morphology (Wilbur 1955; Perry 1971). In all species in the section, prominent ridges extend along the commissural (fused lateral) veins from the base of the calyx to the sinuses between the lobes. At the sinuses they divide and extend along the sub- marginal veins of the adjacent lobes. The commissural and submarginal veins, therefore, are more prominent than the midveins, which are not ridged. In the other sections the calyx tubes either lack ridges along the veins or have low- ridged commissural veins that are no more prominent than the midveins. Plants — insect. Ca mpest ria are annuals, and the branching is entirely or predominantly alternate. The flowers are pedicellate and are pentamerous except lor the carpels. Exceptin S. arenicola, the eye of the corolla is of the shape and pattern described above for S. arkansana, as contrasted with the more widely triangular yellow zones of most species with pink corollas in the other sections. The stamens are inserted immediately below the sinuses of the corolla. The combinations of style branches plus stigmas are linear. The morphology of S.arkansana is consistent with that of the section in all of these respects. Sabatia arkansana exhibits the syndrome of floral morphology associated with predominantly allogamous pollination in Sabatia (Hill 1891; Perry 1971). The corollas are showy and brightly colored, with sharply contrasting eyes. The flowers are protandrous. Initially the styles and stigmas are bent nearly hori- zontally to one side, and the style branches are helically coiled around each other, so that the stigmatic surfaces are not exposed. At this stage the stamens are nearly erect. The anthers, when mature, coil circinately at the tip, with the rest of the anther still being straight and nearly erect when the pollen sacs de- hisce. Subsequently the stamens diverge and the whole anther curves intoa bass- clef shape. The stigmas then become receptive. The uncleft portion of the style becomes erect, and the branches diverge and uncoil more or less completely. Etymology and common name.—We name this new species Sabatia arkansana for the state to which it is apparently endemic. We propose the common name 1254 BRIT.ORG/SIDA 21(3) “Pelton’s rose-gentian” in honor of John Pelton, who first noticed that this spe- cies was distinct from S. campestris, but was too humble to allow a scientific name to be given in his honor. He has done much to further our knowledge and appreciation of the flora of Arkansas through his insight, his photography, and his encouragement and tutelage of younger students of the flora. COMPARISON WITH RELATED SPECIES Only three species, or in some treatments only two, have generally been recog- nized in sect. Campestria (Wilbur 1955; Perry 1971; Bell & Lester 1980). The leaves of S.arkansana are narrower than those of any other species in the section. The mid-stem leaves of the largest plants of S. arkansana are linear to narrowly lan- ceolate, 4-8 times as long as wide, with three primary veins. Except for one pair of leaves on one plant that were 6 mm wide, the maximum width of any leaf seen was 4.5 mm. All leaves of the smaller plants are linear, 1-2 mm wide, with one primary vein. Also uniquely in the section, the corolla lobes of S. drkansana are narrowly spatulate-obovate, generally more than L8x and of- ten more than 2.2x as long as wide. They are widest at ca. 0.8X their length. The apex is rounded. Sabatia campestris Nutt. the most widespread species in sect. Campest ria, isthe most similar to S. arkansana. Both S. arkansana and S. campestris (Witsell 01-0473) were found at the locality where Witsell 01-0474 was collected. At this PRINGLE AND WITSELL, A NEW SPECIES OF SABATIA FROM ARKANSAS 1255 1256 BRIT.ORG/SIDA 21(3) site as elsewhere S. arkansand appears distinctly different in the field, and does not intergrade with S. campestris. Plants of S. campestris are larger than those of S.drkansana in nearby sites. Its leaves are ovate to lance-elliptic, 8-40 mm long < 5-20 mm wide, with clasping, rounded to subcordate bases. The ridges on its calyx tube have a distinct wing or keel along the apex, ca. 1.0 mm high, which projects slightly above the sinus. Its corollas are a paler, less purplish shade of pink. Its corolla lobes are obovate, less than 1.8% as long as wide, and are widest at ca. 0.6% their length. The apex is usually abruptly acute, occasionally obtuse. The pollen grains of Sabatia are tricolporate, with a finely reticulate exine. Perry (971) concluded that pollen size was not useful as an indicator of ploidy or as a species characteristic, and that neither shape nor exine sculpturing dif- fered significantly among the species. The pollen of S. arkansana, however, is subprolate, and grains in equatorial view are as numerousas those in polar view or more so on a microscope slide. That of S. campestris, as seen in the present study, is more nearly spherical, 26-30 tm in diameter, ie. slightly smaller than that of S. arkansana, and, as noted by Perry (1971, nearly always appears in polar view when placed on a slide. At the Womble Formation locality near Owensville, S. arkansana is con- fined to open, flat, seasonally wet, narrow floodplains and seepage areas, whereas S. campestris grows in better-drained, steeper, drier microhabitats. Each of the two species occupies a distinct zone within the glades. Occasional plants of S. campestris occur within the microhabitat occupied by S. arkansana (Fig. 3) and, as noted above, no intergradation has been observed. Sabatia campestris is apparently absent from all of the glades on the nepheline syenite batholith near Bauxite. The two species also differ distinctly in phenology. Based on ob- servations from 2001 through 2004, S. arkansana typically begins flowering 7 to 10 days before S. campestris, and S. campestris continues flowering at least 7 days alter S. arkansana. Sabatia formosa Buckley is known from Louisiana, Oklahoma, and Texas. It is recognized here following Bell and Lester (1978, 1980), but has often been included in S. campestris (Wilbur 1955). Sabatia arkansana is more similar to S. formosa than to S.campestris in its corolla color and markings but not in other respects. Sabatia formosa has closely spaced basal and near-basal leaves, which are larger than the mid-stem leaves and are generally present at flowering time, whereas in S. arkansana there is no evidence from crowded leaf-scars that a basal rosette is present at any stage. The proximal and mid-stem leaves of S. formosa are lanceolate to ovate, 8-25 x 3-13 mm. Its calyces are usually shorter than the corollas. Its corolla lobes are elliptic-rhombic, less than L8X as long as wide, and are widest near the middle, tapering to an obtuse to acute apex. Sabatia arenicola Greenm. (including S. carnosa Small) is a seabeach spe- cies, native along the Gulf Coast from Louisiana to Tamaulipas. Unlike S. PRINGLE AND WITSELL, A NEW SPECIES OF SABATIA FROM ARKANSAS 1257 arkansana it frequently branches from near the base, and its relatively copious and dense branching is proximally more often opposite than that of the other species in section Campestria. Its leaves are succulent when fresh and blacken upon drying. They are elliptic to ovate or obovate, 6-27 x 2-13 mm. Its calyx lobes are oblong-lanceolate to narrowly ovate-triangular. It differs further in exhibiting the floral morphology associated with autogamy in Sabatia. It has relatively small corollas, which are white or light pink with the eyes whitish and less sharply defined than those of the allogamous species. The styles are nearly erect when the flowers open, and the stigmas are receptive concurrently with rather than after the dehiscence of the anthers. Only two other validly published names have been associated with sect. Campestria, and it is evident that neither was based on specimens of S. arkansana. Sabatia nervosa Raf. was included in S. campestris by Wilbur (1955). Its leaves were described as ovate-lanceolate and its corolla lobes as broadly obovate. The identity of S. concinna Alph.Wood, which presumably was based on specimens from Indiana, is uncertain. It was described as having opposite branching, which would be inconsistent with its inclusion in sect. Campestria, and calyces only half as long as the corollas. Small plants of S. campanulata (L.) Torr., in sect. Sabatia subsect. Campanulatae S.FBlake, mewhat similar to S. arkansana in aspect. Sabatia campanulata is an extremely rare (Sl) species in Arkansas. It has narrow leaves and calyx lobes, but the plants are perennial, usually with clustered stems, and the pedicels are mostly 40-70 mm. The calyx tube of S.campanulata is obconic, only 1-3 mm long and less than 0.5 as long as the corolla tube. Ridges along the commissural veins are absent or low and no more prominent than those along the midveins. Sabatia arkansana cannot plausibly be interpreted as a recurrently pro- duced interspecific hybrid. In most populations it is the only Sabatia species present. Other than S. campestris, the only Sabatia that has been found in the vicinity of S. arkansana is S. angularis (L.) Pursh, in sect. Sabatia subsect. Angulares S.FBlake. Sabatia angularis does not intergrade with S. arkansana, and its morphology, which includes opposite branching, stem wings 0.2-0.3 mi high, and lanceolate to ovate leaves 5-30(-40) mm wide, does not suggest that it might bea parent of S.arkansana. The only other Sabatia species known from Arkansas are S. brachiata Ell, in sect. Sabatia subsect. Angulares, and S. gentianoides Ell. in sect. Pseudochironia Griseb. The first is uncommon in Ar- kansas, and the latter is extremely rare (SL) in the state. No intersectional hy- brids in Sabatia are known in nature (Wilbur 1955; Perry 1971). All of Perry's (1967) attempts to make intersectional crosses involving species in sect. Campestria, including S. campestris x both S. angularis and S. campanulata, were unsuccessful. 1258 BRIT.ORG/SIDA 21(3) KEY TO THE SPECIES OF SABATIA SECT. CAMPESTRIA . Leaves succulent when fresh, blackening in drying, elliptic to ovate or obovate; corolla lobes 4-10(- 3) mm _Sabatia arenicola |. Leaves not succulent nor blackening, linear to ovate; corolla lobes 8-25 mm. 2. Basal leaves usually present at flowering time; calyces usually shorter than corol- las; corolla lobes elliptic-rhombic, widest near mid-length Sabatia formosa Basal leaves absent at flowering time; calyces usually as long as or longer than corollas; corolla lobes spatulate to obovate, widest distally. 3. Leaves all lanceolate to linear, generally less than 5 (rarely to 6) mm wide; corolla lobes deep magenta-pink, more than 1.8X as long as wide, widest at ca.0.8% or more of their length, rounded at apex Sabatia arkansana 3. Leaves except in distal portions of inflorescence lance-elliptic to ovate, more than 5 mm wide;corolla lobes light pink or occasionally white, less than 1.8X as long as wide, widest at ca.0.6X their length, + acute at apex ___ Sabatia N campestris HABITAT AND ASSOCIATED SPECIES Sabatia arkansana isa plant of flat, seasonally wet microhabitats in shale and igneous glades in the eastern Ouachita Mountains and igneous batholiths of the Upper West Gulf Coastal Plain of central Arkansas. It appears to be restricted tosmall flat areas along the narrow floodplains of intermittent streams or along seepage areas within these glades. These habitats are typically wet in the win- ter and spring but usually become dry by July and remain so until the late fall. The substrate consists of thin soil over bedrock and of loose, weathered frag- ments of the same type as the local bedrock. No standing water was observed around the plants at the time of flowering. Sabatia arkansana is known from glades with two different geologic sub- strates: shale from the Womble Formation and the igneous rock nepheline sy- enite (Fig. 6). The Womble Formation was deposited during the Middle Ordovi- cian and consists of mostly black shale with thin layers of limestone, silty sandstone, and some chert (McFarland 2004). It is confined to the Central Oua- chita Mountains physiographic province, a subsection of the Ouachita Moun- tains (Foti & Bukenhofer 1998). Large igneous intrusions of Late Cretaceous age outcrop in the Upper West Gulf Coastal Palin (Saline and Pulaski counties) and consist largely of nepheline syenite. These intrusions consist of light gray or bluish feldspathic and feldspathoidal igneous rocks in a batholith containing pendants of altered rocks of Paleozoic age, and are weathered locally to kaolin or bauxite (Haley et al. 1976, Gordon et al. 1958). These are the largest out- croppings of igneous rocks in Arkansas and are located within the Upper West Gulf Coastal Plain physiographic province, an area consisting of mostly un- consolidated surface geology of Cretaceous age and younger (McFarland 2004). Associated species include Talinum ca lycinum, Croton wildenowii, Croton capitatus, Croton monanthogynus, Euphorbia cyathophora, Bulbostylis capillaris, Fimbristylisautumnalis, Valerianella nuttallii, Calamintha arkansana, Allium nd PRINGLE AND WITSELL, A NEW SPECIES OF SABATIA FROM ARKANSAS 1259 * eens of Sas pS Location of fea h Je ph é Detailed Map r PULASKI we OU chita iGulf Coastal . Plain ee F Bo Urban A —— County Pe nderes SALINE * GARLAND ] ® KE Bauxite 5 & oy s = a s s GRANT oy < HOT SPRING es N A L . hol +; ra Iact, Cot en Pere} 1 Cid ae ia Nol 4 ¢L halt Fic.6 ite batholith southeast of Bauxite. canadense var. mobilense, Hedyotis nigricans, Asclepias longifolia var. hirtella, Sedum pulchellum, Polygonum tenue, Aristida dichotoma var. curtissii, Digitaria cognata, Sporobolus ozarkanus, Panicum flexile, Chamdesyce missurica, Nemastylis nuttallii, Silphium laciniatum, Ptilimnium nuttallii, Amsonia hubrichtii, Aster oblongifolius, Eriocaulon kornickianum, Isoetes butleri, Grin- delia lanceolata, Mimosa quadrivalis var. nuttallii, Astragalus distortus var. engelmannii, Selenia aurea, Ranunculus pusillus, Astranthium integrifolium, Minvartia patula, Eryngium yuccifolium, Spiranthes vernalis, and Nostoc sp. (Nomenclature follows Kartesz 1999). Of these associates, the following (given with their conservation status ranks) are tracked as elements of special concern by the Arkansas Natural Heri- tage Commission: Valerianella nuttallii(G1G2SD, Nemastylis nuttallii (G482), Amsonia hubrichtii (G3S3), Eriocaulon kornickianum (G2S2), Chamaesyce missurica (G5S2), and Bulbostylis capillaris (G5S3) (Arkansas Natural Heritage Commission 2002). Valerianella nuttalliiand Amsonia hubrichtii are endemic to the Ouachita Mountains and Arkansas Valley of Arkansas and Oklahoma. This new species of Sabatia is one of more than 15 endemic plant taxa from the Ouachita Mountains (including upland portions of the Arkansas River Valley and 1260 BRIT.ORG/SIDA 21(3) the igneous batholith that outcrops in the Upper West Gulf Coastal Plain near the edge of the Ouachita Mountains in Saline and Pulaski counties, Arkansas). In addi- tion to V. nuttalliiand A. hubrichtii, these endemic taxa include Amorpha ouachi- tensis, Carex latebracteata, Galium arkansanum vat. pubiflorum, Liatris compacta, Hydrophyllum brownei, Polymnia cossatotensis, Monarda stipatatoglandulosa, Houstonia ouachitana, and Quercus acerifolia (Zollner et al. 2005). eu CONSERVATION STATUS Though S. arkansana has been collected from a number of glade openings, all of these are components of only two larger glade complexes, the nepheline sy- enite complex near Bauxite and the Womble Formation (shale) complex near Owensville. Therefore it can accurately be stated that this species is known from just two sites in the world, both in Saline County, Arkansas. Glades with appro- priate microhabitat in adjacent counties (Pulaski, Garland, and Montgomery) were searched in 2003 and 2004 and no new sites for S. arkansana were found. Because S. arkansana is an annual, population size fluctuates from year to year. Two shale glade openings at the Womble Formation site support populations of S. arkansana, which ranged from approximately 200 to more than 1000 in- dividuals per opening from 2001 to 2004. Five glade openings in the nepheline syenite complex support populations on S. arkansana, which ranged from hun- dreds to thousands of plants per opening from 2001 to 2004. These glades have historically been the sites of mines (in the case of the economically important nepheline syenite) and borrow pits for road fill mate- rial Cin the case of the shale glades). The absence of significant past mining at the Womble Formation locality makes it an especially rare site among shale glades in the Ouachita Mountains. The open character of these glades is main- tained in part edaphically by the thin soil and in part, at least historically, by fire. The absence of fire in recent years has allowed glades throughout the re- gion to be invaded by woody plants, particularly eastern redcedar Juniperus virginiana). This encroachment has led to the decline of many plant species dependent upon open glade habitat. These factors, along with significant pressure from encroaching residen- tial development, make these glades a high conservation priority. Several glades immediately west of the Womble Formation locality (and part of the same glade complex) have recently been destroyed for an addition toa large gated residen- tial golf course community. Fortunately, the Arkansas Natural Heritage Com- mission was recently able to acquire and protect 136 acres (55 ha) that include the S. arkansana locality, several other glades, and associated woodlands and forest. Meanwhile, all of the known S. arkansana sites on the nepheline syenite batholith are being protected by joint efforts of the Arkansas Field Office of The Nature Conservancy and Alcoa Corporation. PRINGLE AND WITSELL, A NEW SPECIES OF SABATIA FROM ARKANSAS 1261 ACKNOWLEDGMENTS Theo Witsell extends special thanks to John Pelton for sharing his enthusiasm and knowledge of the flora of Arkansas, and for showing him that beautiful pink flower in the glade back in the summer of 2001. Thanks also to Linda Ellis for the illustration and to Tanya Miller-Witsell for spending a day of our honey- moon in the herbarium at the Missouri Botanical Garden. Jim Peck (LRU), Johnnie Gentry (UARK), and George Yatskievych (MO) all helped facilitate this research. Seth Young, Chris Tracey, and Bill Shepherd reviewed and improved earlier versions of this manuscript. Thanks also to Meryl Hattenbach, Scott Simon, and Doug Zollner of the Arkansas Field Office of The Nature Conservancy, Jarvis Harper of Alcoa Corporation, and the staff of the Arkansas Natural Heritage Commission. Special recognition should go to Hugh and Steve Davis for grant- ing permission to collect on their land at the Womble Formation site and for agreeing to sella portion of that land for dedication as the Middle Fork Barrens Natural Area. James S. Pringle thanks the curators and staff of the herbaria of the Arkansas Natural Heritage Commission and the University of Arkansas at Fayetteville for the loan of specimens, and those at Harvard University and the Missouri Botanical Garden for the opportunity to study and compare specimens at those institutions. REFERENCES ARKANSAS Natural HeritaGe Commission. 2002. State species of special concern—plants. Arkansas Natural Heritage Commission, Little Rock. Bett, N.B. and L.J. Lester. 1978.Genetic and morphological detection of introgression in a clinal population of Sabatia section Campestria (Gentianaceae). Syst. Bot. 3:87-104. Beit, N.B. and L.J. Lester. 1980. Morphological and allozyme evidence for Sabatia formosa (Gentianaceae) in the section Campestria. Amer. J. Bot.67:327-336. Fon, T.L. and G.A. BukenHorer. 1998. A description of the sections and subsections of the Interior Highlands of Arkansas and Oklahoma. J. Arkansas Acad. Sci. 52:53-62. Goroon, M., J.|. TRAceY, and M.W. Eitis. 1958. Geology of the Arkansas bauxite region. Geological Survey Professional Paper 299.U.S.Government Printing Office, Washing- ton, D.C Hacey, B.R., E.E. Guick, W.V. Bush, B.F. CLarby, C.G. Stone, M.B. Wooowarp, and D.L. ZacHary. 1976. Geologic map of Arkansas. Arkansas Geological Commission and U.S. Geological Survey, Little Rock. Hitt, EJ. [(“E.G."]. 1891. The fertilization of three native plants. Bull. Torrey Bot. Club 18: 111-118. Kartesz, J.7.1999. A synonymized checklist and atlas with biological attributes for the vas- cular flora of the United States, Canada, and Greenand. First edition. In: Kartesz, J.T.,and C.A.Meacham. 1999. Synthesis of the North American flora, Version 1.0.North Carolina Botanical Garden, Chapel Hill. Compact disc. 1262 BRIT.ORG/SIDA 21(3) McFarvano, J.D. 2004. Stratigraphic summary of Arkansas. Arkansas Geological Commis- sion Information Circular 36. Arkansas Geological Commission, Little Rock. Perry, J.D. 1967. Biosystematic studies in the North American genus Sabatia (Gentianaceae). Ph.D. dissertation, Duke University, Durham, North Carolina. Perry, J.D.1971.Biosystematic studies in the North American genus Sabatia (Gentianaceae). Rhodora 73: 309-369. Wieur, R.L.1955.A revision of the North American genus Sabatia (Gentianaceae).Rhodora 57:1—33, 43-71, 78-104. Zoutner, D., M. MacRoserts, B. MacRoserts, and D. Lapp. 2005.Endemic vascular plants of the Interior Highlands, U.S.A. Sida 21:1781-1791. AULONEMIA NITIDA (POACEAE: BAMBUSOIDEAE: BAMBUSEAE), A NEW SPECIES FROM GUYANA Emmet J. Judziewicz Department of Biology University of Wisconsin-Stevens Poin Stevens Point, Wisconsin 54481, U : emmet.judziewicz@uwsp.edu ABSTRACT A new species, Aul ia nitida Jud is described from the Pakaraima Mountains of Guyana, South America. It is a large-leaved species that differs from the Mesoamerican species A. laxa and A. pa- triae in its smooth, shining, non-striate, non-maculate foliage leaf sheaths. An illustration of the new species and a key to the species of Aulonemia from the Guayana Highlands are provide RESUMEN Se describe una especie nueva, Aulonemia nitida Judz., de las montanas de Pakaraima, Guyana, América del Sur. Es una especie con laminas foliares grandes y anchas parecida a las especies Mesoamericanas A. laxa y A. patriae, pero se distingue por sus vainas foliares lisas y lustrosas, sin estriaciones o manchitas. Se incluyen una ilustracion de la nueva especie y una clave de las especies de Aulonemia de las montanfias de Guayana. Since the publication of the grass treatment for the “Flora of the Guianas” project Judziewicz 1991), several more specim ene of a new bamboo (Poaceae: Bambusoideae: Bambuseae: Arthrostylidi from the Guayana High- lands have come to light. The name Aulonemia nitida has long been in print, appearing as a nomen nudum in Boggan et al. (1997: 168) checklist of Guianas plants, and, based on the Boyan specimen, was recognized asa likely new species as long ago as 1988. Even though all five known collections are sterile, the taxon is so distinctive that I have chosen to propose it here as a new species, following the example of Clark (1989, 1992) who has named several distinctive species in the bamboo genus Chusquea Kunth based on material lacking inflorescences Specimens were examined from the following herbaria: F (2002), FDG (1988), K (1989), MO (2004), NY (1989), US (2004), UWSP (2004), and WIS (2004). Aulonemia nitida Judz., sp. nov. (Fig. 1). Type: GUYANA. cep a REGION: upper slopes of Mt. Wokomung, 5°05'N-59°50"W, 1530 m, Hedyosmum-bamboo dominated moist forest, bamboo with culms at base to 2 cm diam., overall height 4 m, common and at times dense on upper oe sterile, 14 Jul 1989, B.M. Boom & GJ. Sa ae 9224 (HOLOTYPE: FDG! ISOTYPES: MO-3 sheets!, NY! Graminum cespitosum. Culmi usque ad 5 m longi, 2.5 cm lati. Vaginae foliorum glabrae, nitidae, fimbriatae ad apicem tantum; fimbriae 7-10 cm longae; laminae foliorum 25-37 cm longae, 7-10.5 cm latae, lanceolatae-ovatae. Inflorescentia non vidi. SIDA 21(3): 1263-1267. 2005 1264 BRIT.ORG/SIDA 21(3) cm Fic. 1. A.B: f plant st i Jial rhi ( 1 17 J 4 " «it one . toh] *oay\ B. Culm and branch. C. Branch with foliage | D. Detail of ligul Scale bar = 1 cm. Illustration by Emily Lain. A and B based on Boom & Samuels 9224 (MO), Cand D based on Henkel, Williams, Fratello & Williams 4493 (WIS). JUDZIEWICZ, A NEW SPECIES OF AULONEMIA FROM GUYANA 1265 Cespitose perennial, thicket-forming woody bamboo from short, sympodial rhizomes; glabrous throughout. Culms up to 5 m long and at least 2.5 cm in diameter, shiny, hollow, the walls only ca. 1 mm thick with a 1-15 mm thick annular ring present at the lower nodes; buds and branches one per node. Culm leaves not seen, perhaps not differentiated from foliage leaves. Foliage leaves at least 5 per complement; sheaths glabrous, smooth, shiny, and stramineous throughout most of their length, strongly keeled 3-5 cm below the apex, striate only in the area 0.5-1 cm below the apex; fimbriae ca. 12-20, each 7-10 mm long, pale, flexuous and spreading, confluent basally and forming an indurate rim at the summit of the leaf sheath; outer ligules ca. 0.2 mm long, indurate, rim-like; inner ligules 1-15 mm long, membranous; pseudopetioles 7-10 mm long, pale; blades 25-37 X 7-10.5 cm, narrowly ovate to broadly lanceolate, rounded to very slightly oblique or subcordate at the base, acuminate at the apex, not evidently tessellate, the abaxial surface slightly whitened or bluish green. Inflorescence not seen. Additional collecti ined ( yPES): GUYANA. Cuyuni-Mazaruni Region: Ayanganna [a sand- stone mountain with a summit a ation ol over 2100 m| slope, among rocks in low forest, sterile, 2 Mar 1960, R. Boyan 120 [= FD7944] (FDG, NY): ie araima Mts., 2 km transect gene summit ridge et Mt. Ayanganna, 5°23'N- comet 1800-2000 m stone, 3 Nov 1992, B. Hoffman & T. Henkel 3209(US, WIS). atk: ae Region: : Mt. oe E- most 2 of massif, 5°05'34"N-59°50'13"W, 1524 m, coarse herb 4 m, sterile, 13 Ju Clarke, R. Williams, C. Perry, E. Tripp, D. Gittens & S. Stern 10808 (US, WIS); Mt. Wokomung, summit ridge - Ka-mie-wah pao NE to § pinnacle, “Little Ayanganna,” 5°04'N-59°52'W, 1550-1650 m, od cl ana scrub [orest on pinnacle escarpments; on sandstone, mixed hardwoc grading stout bambusoid grass from 1-5 m tall, forming thickets on rocky ridges, sterile, 17 Nov 1993, T.W. Henkel, R. Williams, S. Fratello, L. Williams 4493 (FDG, MO, US, WIS). } DISCUSSION Aulonemia nitida is endemic to elevations of 1500-2100 m in cloud forests on sandstone tepuis in the Pakaraima Mountains (Mt. Ayanganna and Mt. Wokomung; all collections come from an area of about 30 km in length) of Guyana near the Brazilian frontier. The species epithet derives from the nitid or shiny foliage leaf sheaths. A vernacular name is “reroballi” (Boyan 120). There are 35 described species of Aulonemia throughout tropical America (Clark et al. 1997; Judziewicz et al. 1999, 2000) and lam currently working ona revision of the genus. Aulonemia nitida appears to be most closely related to the Mesoamerican species A. laxa (F Maek.) McClure and A. patriae R. Pohl (Pohl & Davidse 1994). All three taxa have broad, fimbriate foliage leaves with dis- tinctive keeled sheaths, but A. laxa and A. patriae have sheaths that are striate and maculate their entire lengths. Aulonemia nitida differs in its smooth, shiny, non-maculate sheaths that are striate, if at all, only in the final 1 cm or so below the ligular area. The two Mesoamerican species also differ in their generally 1266 BRIT.ORG/SIDA 21(3) longer 10-32 mm) fimbriae; those of A. laxa are straight and appressed to the culm, not spreading as in A. nitida and A. patriae. There are eight species of the Aulonemia in the Guayana Highlands Qudziewicz 2004). In the “Flora of the Venezuelan Guayana” Judziewicz et al 1991; Judziewicz 2004) Aulonemia nitida would key to either A. deflexa (N.E. Brown) McClure, A. chimantaensis Judz. & Davidse, or A. jauaensis Judz. & Davidse. However, these three taxa have smaller foliage leaf blades (a maximum of 22 x 4.5cm versus at least 25 X 7 cm in A. nitida), longer fimbriae (10-20 mm long versus 7-10 mm in A. nitida), and completely striate foliage leaf sheaths. Aulonemiad deflexa, the only other species known from Guyana, is present at higher elevations on Mt. Roraima (ca. 2800 m), as well as on several Venezuelan tepuis. Aulonemia patula (Pilg.) McClure from Andean Colombia and Ecuador has similarly large foliage leaves, but the sheaths are non-keeled, non-striate, and the fimbriae are conspicuous all along the sheaths margins as well as at the sheath summit. A key differentiating the species of Aulonemia found in the Guayana Highlands follows. KEY TO THE SPECIES OF AULONEMIA FROM THE GUAYANA HIGHLANDS 1. Foliage leaf sheaths with oe fimbriae 2. Foliage leaf blades wit h midrib excentric, “piaced 5-7 mm from one margin of a a 22 ~25 mm wide; 600-700 m, Cerro Huachamacari, Amazonas, Venezuela A.sp.A (Judziewicz 2004 2. Foliage leaf blades with midrib placed centrally on a blade 30-60 mm wi 2200 m, Bolivar and Amazonas, Venezuela 1. Foliage leaf sheaths lacking marginal fimbriae. 3. Foliage leaf sheath summits prominently auriculate; 1000-2000 m, Cerro Duida, Amazonas, Venezuela 3. Foliage leaf sheath summits lacking au 4, Dwarf plants ca.0.5 m tall; foliage at blade ca.4.cm long, 0.7 cm wide; 2600 m, Cerro Marahuaka, Amazonas, Venezuela ena 2004) 4, Plants 1 ~3 or More m tall; foliage leat blades 11—37 cm ret ne A. aff. subpectinata (Kuntze) McClure Buavianie: 2004 a 5. Piss leaf sheaths smooth and shiny, pe blades 25-37 cm long, 7-10.5 cm wide; fimbriae 7-10 mm long; 1500-2100 igo na A. nitida Judz. 5. Foliage leaf sheaths striate and dull, the ae ~20 cm long, 1.8-4.5 cm a fimbriae 10-20 mm long. Spikelets 12-20 mm long, 4—-5-flowered; 2100-2800 m, Bolivar, Venezuela and adjacent Guyana A. deflexa (N.E. Brown) McClure 6. Spikelets 22—70 mm long, 9-23-flowered. 7, Spikelets 22-40 mm ai Sen 1;lemmas 8-10 mm long, ob- tuse, slightly tridentate at apex, glabrous to sparsely puberulent on ck, the margins ee ae 2200 m, Macizo de Chimanta, Bolivar, Venezuela A. chimantaensis Judz.& Davidse 7. Spikelets (20-)40-70 mm long, 11-23-flowered; lemmas 10-13 mm long, acute, densely puberulent throughout with prickle-like hairs; 1900-2100 m, Cerro Jaua, Bolivar, Venezuela A. jauaensis Judz. & Davidse JUDZIEWICZ, A NEW SPECIES OF AULONEMIA FROM GUYANA 1267 ACKNOWLEDGMENTS I thank Lynn G. Clark for encouraging me to publish this paper; the curators of MO, US, and WIS for loans; Emily Lain for the illustration; and Paul M. Peterson for a thorough and helpful review. REFERENCES Bocean, J., V. FUNK, C. Kettorr, M. Horr, G. Cremers, and C. Feuitter. 1997. Checklist of the plants of the Guianas. 2nd ed. Biological Diversity of the Guianas Program, Smithsonian Institu- tion, Washington, DC. Cark, L.G. 1989. Systematics of Chusquea section Swallenochloa sect. Swallenochloa, sec- tion Verticillatae, section Serpentes, and section Longifoliae (Poaceae: Bambusoideae). Syst. Bot. Monogr. 27:1- Ciark, L.G. 1992. Chusquea sect. Swallenochloa (Poaceae: Bambusoideae) and allies in Brazil. Brittonia 44:387-422. Ctark, L.G., X. Lonpono, and M. Kosayasuii. 1997. Aulonemia bogotensis (Poaceae: Bambusoideae), a new species from the Cordillera Oriental of Colombia. Brittonia 49:503-507. Jupziewicz, E.J. 1991. Family 187. Poaceae. 1991 [as 1990]. ln: A.R.A. Gorts-Van Rijn, ed., Flora of the Guianas, Series A: Phanerogams. Koeltz Scientific Publications, KOnigstein, Germany. Juoziewicz, EJ. 2004. Aulonemia. |n: Steyermark, J.A., PE. Berry, K. Yatskievych, and B.K. Holst, eds. Flora of the Venezuelan Guayana, Vol. 8: Poaceae-Rubiaceae. Missouri Botanical Garden Press, St. Louis. Pp.40-45. Jubziewicz, E.J., L.G. CLARK, X. LONDONO, and MJ. Stern. 1999. American Bamboos. Smithsonian Institution Press, Washington, DC. Juoziewicz, E.J.,G. Daviose, and L.G. CLark. 1991. Six new bamboos (Poaceae: Bambusoideae: Bambuseae) from the Venezuelan Guayana. Novon 1:76-87. Juoziewicz, E.J., RJ. suas P.M. Peterson, T.S. Fitcueiras, and F.O. Zutoaca. 2000. Cata- logue of New World gr (Poaceae):|.Subfamilies Anomochlooideae, Bambusoideae, Ehrhartoideae, and prarelieae Contr. U.S. Natl. Herb. 39:1-128. Pout, R.W.and G. Daviose. 1994. Aulonemia. In: Davidse, G.,M. Sousa-S.,and A.O. Chater, eds. Flora Mesoamerica. Vol. 6, Alismataceae a Cyperaceae. Universidad Aut6énoma de México, Mexico, D.F. Pp. 198-199. 1268 BRIT.ORG/SIDA 21(3) BOOK REVIEWS RoLanp H. Wauer. 2001. Naturally...South Texas: Nature Notes from the Coastal Bend. (ISBN 0-292-79144-5, hbk.). The University of Texas Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A. (Orders: 800-252-3206, fax 800-687-6046, www.utexas.edu/utpress). $22.95, 240 pp., 6" x 9" This isa collection of brief essays arranged chronologically asa “natural history calendar” and sorted into 12 chapters, January through December. All were originally published in the Victoria Advocate, the regional newspaper. After the author retired from the National Park Service, he moved to Victoria (about halfway between Houston and Corpus Christi) in south Texas, which “undoubtedly is the best birding area anywhere in the United States.” This region “encompasses four rather distinct eco- systems, all within a mile circle of Victoria: the northeastern edge of the South Texas Plains, the southern edges of the Post Oak Savannah and Blackland Prairie, and the heart of the Gulf Prairie and Marshes.” Wauer’s attention is strongly turned to birds, but various plants, insects, and other animals are ated The essays are built around his own observations and interpretations and are sup- ported by “technical” insertions from various sources—all easily readable i enn Botanical Research Institute of Texas, Fort Worth, TX, 76102-4000, U.S.A. JASON F SHOGREN (Ed.). 2005. Species at Risk: Using Economic Incentives to Shelter Endangered Species on Private Lands. (ISBN 0-292-70597-2, pbk.). The University of Texas Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A. (Orders: 800-252-3206, fax 800-687-6046, www.utexas.edu/utpress). $21.95, Zi Dps0' xo, Regarding the rapid decline and loss of species: “The stakes are high, and they go to the heart of our collective responsibility to leave this land a better place than we found it. This book is an attempt to b y develop and refine a workable, practical, and equitable set of incentives lor preserving species and the habitat they need for survival” (from the Foreword). About ha f of all endangered species rely on private land for their habitat, but private landowners have often opposed the regulations of the En- dangered Species Act, which, they argue, uniairly limits their right to Be it from their property. “ this pace lawyers. economists, political sciet s, historians, and zoologists come together to assess th c P 2 | for protecting spe- cies at neko on ae proper 1. Introduction Part I. Current and proposed incentive options for species protection on private lands. 2. The Endangered Species Act and its current set of incentive tools for species protection. 3. An economic review of incentive mechanisms to protect species on private lands. Part Il. Challenges to using economic incentives for species protection. 4. Endangered species protection and ways of life: Beyond economy and ecology. 5. A critical examination of economic incentives to promote conservation. 6. Appraising the conservation value of private lands. 7. Markets 8. The role of private information in designing conservation incentives for property owners. or conserving biodiversity habitat: Principles and practice. Part II]. Economic incentives for ESA reauthorization 9. Evaluating the incentive tools. SIDA 21(3): 1268. 2005 FOUR NEW SPECIES OF ERICACEAE (VACCINIEAE) FROM ECUADOR James L.Luteyn Institute of Systematic Botany The New York Botanical Garden Bronx, New York 10458-5126, U.S.A. jluteyn@nybg.org ABSTRACT Four new species of endemic cua from montane Ecuador, Ceratostema oyacachiensis, Ceratostema pendens, C I Disterigma bracteatum, are described. illustr ated, and their relationships discussed. RESUMEN Se describen y se ee t ies nuevas endemicas de mortinos de la sierra del Ecuador, Ceratost hi Coriesene cy Ceratostema pubescens, y Disterigma bracteatum, con sus ilustraciones y racine. INTRODUCTION In Ecuador, the Ericaceae, with 21 genera and about 222 species, are one of the largest and most conspicuous montane, flowering plant families (Luteyn 1996, 1998, 2002). Despite the recent treatment of the family in the Flora of Ecuador series, the number of new species continues to increase due to additional col- lecting efforts in previously res or FUG erexD One regions. This paper documents some of these new sf ,once again, that our basic knowledge of the numbers of species in this family, even in a country as well collected and studied as Ecuador, is still uncertain. CERATOSTEMA Jussieu Ceratostema is a montane genus of about 35 species of blueberries that is char- acterized by stamens usually as long as the corolla and of equal lengths, pedicels usually articulate with the calyx, anther thecae that are coarsely papillate, an- ther tubules that are elongate and about half the diameter of the thecae, and large corollas with lobes that are proportionately elongate. It ranges from Ven- ezuela and Guyana south through the Andes to northern Peru. Luteyn (1984, 1986) considered the genus morphologically related to Semiramisia Klotzsch, although recent molecular studies (Powell & Kron 2003) place it ina clade with Macleania Hook. and Psammisia Klotzsch. The genus is currently being inves- tigated by the author. SIDA 21(3): 1269-1282. 2005 1270 BRIT.ORG/SIDA 21(3) - Ceratostema- l: G yi Cyace chrensis Fic. 1. Ceratostema oyacachiensis. A, habit. B, detailed habit showing close-up of axillary bud. C, detail of leaf base (undersurface). D, flower showing pedicel, calyx, corolla, and detail of calyx lobe margins. E, longitudinal section of feides lati tae Let FE] tudinal cacti f calvy G howina lateral. ventral a J d baa ry ee | £ * at dah? ld £ hal type, Stahl et al. 2512) LUTEYN, NEW SPECIES OF ERICACEAE FROM ECUADOR 1271 Ceratostema oyacachiensis Luteyn, sp. nov. (Fig. 1). Type: ECUADOR. Napo: Rio Chalpi at confluence with Rio Oyacachi, 00°15'S, 77°58'W, 2500-2550 m, 21 May 1996 ({1), B. Stahl, P Asimbaya & H. Navarrete 2512 (HOLOTYPE: NY; ISOTYPES: AAU, K n.v, MO n.v, QCA n.v., QCNE nv). I ibus differt foliis quoad s, base late cuneatis vel obtusis, lyci costato, rlmbocalyein incnepcun rotato, HOB TeX ie r 5 [ calycibus ad pedicellos articulatis, tub ] ee ol a1 dh ] Jon 13 3 ] sal ] a Fea | &lat 1dUIOSIS, o atque corollis magnis crasso-carnosis lobis earum brevibus latisque. Epiphytic shrubs; mature stems terete or subterete and bluntly angled, glabrous, the bark grayish, cracking longitudinally and exfoliating in thin strips; twigs terete to subterete, striate, glabrous, reddish-brown; axillary buds arising up to 3.5mm above leaf-nodes, the outer pair of scales 2, valvate, relatively obscure, narrowly triangular, acuminate, up to 2.5 mm long. Leaves alternate, flat, the blades thick-coriaceous, lanceolate, 5.2-l1 X 1.2-2.4 cm, basally broadly cuneate to obtuse, apparently decurrent onto petiole, apically long-acuminate, glabrous, the venation weakly 3-5-plinerved from near base, the midrib thickened and raised in proximal ca. 5 mm then plane to weakly impressed distally adaxially, raised and conspicuous abaxially, the lateral nerves plane to very weakly im- pressed adaxially and raised abaxially, the reticulate veinlets obscure adaxially and weakly raised abaxially; petioles subterete, slightly flattened adaxially, slightly winged to blade, ca. 4-11 mm long, glabrous. Inflorescences axillary, sometimes located along tips of branches where leaves have fallen, racemose, 2-8-flowered, somewhat short-pedunculate with flowers congested distally; rachis subterete, striate, 1.5-2.5 cm long, glabrous; floral bract caducous, not seen; pedicel terete, striate, 15-18 mm long, glabrous, articulate with calyx; bracteoles 2, located near base, caducous, ovate, ca. 2.2 mm long, apically long- acuminate, marginally glandular-fimbriate. Flowers 5-merous, pendent; calyx 6-9 mm long, glabrous, the tube obconic, truncate, terete to bluntly 10-ribbed, 2.5-3 mm long, the limb open, spreading to rotate, 5-6 mm long, the lobes broadly ovate, short-acuminate, 3-4 X 5mm, with margins thin and seemingly lacerate-glandular, the sinuses acute; corolla thick-carnose, bistratose, cylin- drical but slightly broadening distally, terete in cross-section, ca. 37-46 mm long, 7-9 mm basal diam. and 9-14 mm diam. at throat, orange, glabrous exter- nally, the lobes broadly deltate, bluntly acute, 4-7 x 6-7 mm, green, densely floccose internally with flat, translucent trichomes to 2 mm long; stamens 10, + equaling corolla in overall length, equal with each other, ca. 36-43 mm long, the filaments distinct, glabrous, ca. 4-6 mm long, the anthers ca. 33-39 mm long, the thecae ca. 9.5-11 mm long, basally conspicuously granular-papillate, the tubules ca. 24-28 mm long, seemingly connate in proximal 2/3, dehiscing by terminal pores ca. 0.2 mm diam, style exserted, to 56 mm long, glabrous. Fruit not seen. 1272 BRIT.ORG/SIDA 21(3) Distribution —Endemic to northeastern Ecuador, where it occurs in both primary and disturbed forest, at ca. 1500-2550 mm. Ceratostema oyacachiensis is characterized by having leaf blades that are basally broadly cuneate to obtuse with plinerved venation, calyces that are ar- ticulate with the pedicels, calyx tubes that are terete to 10-ribbed, calyx limbs that are inconspicuous and rotate, calyx lobes that are short and lack basal, circular glands but do possess instead lacerate-glandular margins, corollas that are large and thick-carnose, broadening slighty distally and having short and broad lobes. In Luteyn’s (1996) key to the Ecuadorean species of Ceratostema, this new species would be found in the vicinity of C. pedunculatum Luteyn, C. prietoi AC. Sm,,C. nubigenum(A.C. Sm.) A.C. Sim.,and C. ventricosum Luteyn. It is distinct from all those species, however, based on its combination of charac- ters mentioned above. If it were not for the articulate calyx/pedicel, this new species might be placed in the genus Semiramisia due to its rotate calyx limb and corolla that broadens slightly distally with relatively short, broad lobes. Additional collections of this species are needed to determine its morphologi- cal range of variation and relationships. — Additional collections examined: ECUADOR. Sucumbios: Sinangoe Station, Shishicho Ridge, Alto Aguarico drainage, above (south of) Rio Cofanes, W of Puerto Libre, NW of Lumbaqui, 00°12'N, 77°32'W, 1500-1570 m, 13 Aug 2001 U1), Aguinda, Pitman & Foster 1673 (F QCA nv, QCNE nw). Ceratostema pendens Luteyn, sp. nov. (Fig. 2). Tyee: ECUADOR. MoRONA-SANTIAGO: Limon-La Union road, trail beyond end of road (beginning at 13.6 km from Limén) towards La Union, ca. 2°59'S, 78°25'W, 1340-1370 m, 18 Nov 1998 (EL, fr), J.L. Luteyn & H. Mogollon 15376 (HOLOTYPE: NY; ISOTYPES: AAU, CAS, MO, QCA, QCNE, US) Ab C. auriculato Luteyn foliis breve pilosis (non glabris), calice breviore 8-9 mm longo (non 12-14 mm), tubo calycis tereti vel quinquangulo (non 5- aaa rae calycis onus mGORS PICO: lone ) calycinis brevioribus 4.8-5 mm longis (non 9-1 g Epiphytic shrubs, arising from lignotubers; mature stems long- pendent, terete, striate, glabrous, the bark thin, reddish, cracking longitudinally; twigs subterete to terete, striate, brownish, densely spreading short-pilose ea simple (uniseriate, unicellular) trichomes ca. 1-13 mm long, glabrate; axillary buds not seen. Leaves alternate, petiolate, flat, amplexicaul, involute at base so as to conceal flowers and fruits, the blades broadly ovate, 7-10 x 4-7 cm, basally deeply cordate and slightly auriculate with the lobes imbricate when fresh, apically acuminate to short-acuminate, densely soft, white, short-pilose on both surfaces with simple trichomes ca. 1 mm long, the venation pinnate with 2-4 lateral veins or weakly 5(-7)-plinerved with inner pair of lateral nerves arising in the proximal | cm, the midrib thickened and raised in proximal 1 cm then plane to weakly impressed distally adaxially, weakly raised abaxially, the lat- eral nerves anastomosing distally and along with reticulate veinlets weakly raised but obscure on both surfaces; petioles terete, rugose, 2-3 mm long, densely short-white-pilose with simple trichomes. Inflorescences axillary, racemose, 1273 LUTEYN, NEW SPECIES OF ERICACEAE FROM ECUADOR } tral. lateral, and A U a uu 4 + + J a) + we % ai [oe <= 2 Y o < =~ fs x = < b=) = < = = 3 same a + ~ 7 a > “ ws 4 Ms x = * + Me _ uo 4 uo “uO =x gd - => ow Lo oc uw A 1274 BRIT.ORG/SIDA 21(3) |-4-flowered but evidently only 1-2 flowers develop per rachis; rachis subterete, ca. 8-10 mm long, short-pilose with white, simple trichomes; floral bract 1, tri- angular, acuminate, ca. 2mm long, densely short-pilose with simple trichomes; pedicel subterete, 5-6 mm long, articulate with calyx, short-pilose with simple trichomes; bracteloles 2, located near base, ovate, acute, ca. 1.3 mm long, short- pilose with simple trichomes. Flowers 5-merous; calyx ca. 8-9 mm long, short- pilose with white, simple trichomes, the tube terete to slightly pentagonal in cross-section, obconic, ca. 2.7-3.5mm long, densely matted short-pilose, the limb spreading, ca. 5.3-5.5 mm long, moderately short-pilose, the lobes triangular- ovate, short-acuminate, ca. 4.8-5 mm long, striate, eglandular, moderately short- pilose externally and weakly so internally, the sinuses acute; corolla carnose, bistratose, cylindric to broadly and bluntly pentagonal in cross-section, slightly widening distally, ca. 45-48 mm long and 1] mm diam. at throat, dark maroon- red to pinkish-red, densely short-pilose with white, simple trichomes ca. 1 mm long, the lobes narrowly triangular, long-acuminate, ca. 12-13 X 3mm; stamens 10, + equaling corolla in overall length, alternately slightly unequal with each other, ca. 45mm and 46.5 mm, the filaments equal, connate intoa tube ca. 8-8.5 mm long, glabrous, the anthers 39.5 mm and 41 mm long, the thecae equal, ca. 7 mm long, conspicuously papillate, the tubules 2, alternately slightly unequal, distinct to base, ca. 34 mm and 35.5 mm long, dehiscing by introrse, oblique, oval pores ca. 0.6-0.8 mm long. Fruit a spherical, translucent cream-colored to waxy white, juicy berry, 15-22 mm diam., weakly short-pilose; seeds numer- ous, surrounded by translucent, mucilaginous sheath. Distribution —Endemic to Ecuador, where it occurs in primary forest on sandstone substrates, at 1OOO-1600 m. Ceratostema pendens is characterized by its long-pendent, epiphytic habit with generally short-pilose vegetative and floral organs, amplexicaul leaves with cordate blades that are basally involute thus concealing the flowers and fruits when living, short floral bracts, terete to slightly pentagonal calyx tube, relatively inconspicuous calyx limb and lobes, and translucent whitish berry. In Luteyn’s (1996) key to the Ecuadorean species of Ceratostema, this new spe- cies would be found in the final couplet containing C. silvicola and C. amplexicaule. It may be easily distinguished from those species by its involute leaves, fewer-flowered inflorescences, connate staminal filaments, and white berry (although berry color is unknown for C. silvicola). It is morphologically most similar to C.auriculatum Luteyn, having in commona long-pendent, epi- phytic habit, amplexicaul leaves with blades that are cordate and pinnately- nerved, and few-flowered inflorescences that are hidden by the leaves. Ceratostema pendensditters from C.auriculatum, however, by having leaves that are short-pilose (vs. glabrous), shorter calyces (8-9 mm ys. 12-14 mm long), ter- ete to pentagonal calyx tubes (vs. conspicuously 5-winged), and calyx limbs that are relatively inconspicuous possessing shorter lobes (4.8-5 mm vs. very LUTEYN, NEW SPECIES OF ERICACEAE FROM ECUADOR 1275 conspicuous and 9-10 mm long) that lack glandular fimbriae. There are very few collections of these species, however, and so interspecific relationships are uncertain at this time. In Flora of Ecuador (Luteyn 1996), the sterile collection van der Werff & Palacios 10428 (MO, NY) was determined as Ceratostema macbrydiorum Luteyn, but the pubescence of its young leaves now characterizes it as an ex- ample of this new species. This points out further that sterile material of C. pen- dens, C.auriculatum, and maybe C. cutucuense Luteyn may be confused, due primarily to their having in common amplexicaul leaves with blades that are rounded to broadly ovate and deeply cordate basally, and short-acuminate apically. Table | compares and contrasts these species. Additional collections examined: ECUADOR. Morona-Santiago: Limon Indanza, Cordillera de Huaracayo, E of Cordillera del Condor and Blo Coangos, E of Shuar village of Tinkimints, 3°15'S, 78°11'W, 1600 m, 24 Mar 2001 ({1, fr), Neill & es 13192 (MO, NY); along unfinished road E of Limon, 1000 m, 5 Feb 1989 (ster), van der Werff & Palacios 10428 (MO, NY). — Ceratostema pubescens Luteyn, sp. nov. (Fig. 3). Type: ECUADOR. EL ORO: Manu-Chilla road, Km 36, 10 km W of Guanasan, 3°28'S, 79°33'W, 2600 m, 4 Oct 1996 (£1), G.P. Lewis, P. Lozano, N. Aguirre & I. Aldaz 2640 (HOLOTYPE: NY, ISOTYPES: AAU n.v, E nv, K nv, LOJA nv QCNE n.v.) AbC fasciculato Luteyn foliis ad | is | i dese Pees eet inflorescentia e fasciculis 4-6-floris (non e racemis usque 30 Fc ris) composita, bracteis floralibus ongioribus 20-26 mm longis (non 17-20 mm), corolla breviori 38- 43 1 mm longa (non 45-57 mm), staminibus brevioribus 36-40 mm longis (non 45-51 mm) differt. — Coarse, terrestrial shrubs, sometimes semi-scandent, 2-3 m tall with stems to 10-12 cm diam., arising from lignotubers; mature stems somewhat contorted, erect or pendulous, subterete, coarsely and bluntly angled, densely short-pilose with whitish, simple trichomes, the bark grayish; twigs subterete, bluntly angled, striate, reddish-brown, densely pilose with whitish, simple trichomes to ca. 2 mm long; axillary buds with outer scales 2, valvate, pseudostipular, narrowly lanceolate, long-acuminate, 6-11.5 x 1.5-2 mm, carinate, short-pilose with whitish, simple trichomes. Leaves alternate, congested, petiolate, the blades coriaceous, flat to slightly revolute, sometimes slightly bullate, ovate, 4-10.5 x 2.5-6.5 cm, basally rounded and often subcordate, apically short-acuminate, glabrous to weakly short-pilose with whitish, simple trichomes adaxially and there also bearing multicellular-multiseriate, reddish, glandular-fimbriate trichomes, densely white pilose abaxially and there also reddish, glandular- fimbriate, when fresh dark to yellowish-green adaxially and whit- ish-green abaxially, when dry olive-green adaxially and reddish-brown abaxially), the venation 3-5(-7)-plinerved with inner lateral nerves arising 1-2 cm above the base, the midrib thickened and raised in proximal 1 cm then plane to slightly impressed distally adaxially, raised and conspicuous abaxially, the lateral nerves plane to impressed adaxially and raised abaxially, the reticulate — 1276 BRIT.ORG/SIDA 21(3) Taste 1. Salient features that characterize and distinguish four closely related species of Ceratostema ~ C.auriculatum, C. cutucuense, C. macbrydiorum, and C. pendens. Ceratostema Ceratostema Ceratostema Ceratostema pendens auriculatum cutucuense macbrydiorum Twig pubescence = Glabrous Glabrous Densely hirsute — Short-pilose Leaf Posture Amplexicaul, flat Amplexicaul, flat Amplexicaul, flat to ag to somewhat volute incurved thus hiding flowers Apex Acuminate Cuspidate to Short-acuminate Acuminate acute Pubescence Glabrous Glabrous Pilose (glabrate — Pilose both surfaces adaxially) Venation Pinnate 5-plinerved 5-7(-9)- Pinnate to weakly plinerved plinerved alyx Overall length 12-14 8.5-10 ca. 28 8-9 mm) Tube cross- 5-winged 5-winged 5-winged Terete to 5-angled section Tube length 3-45 6.5-7 6 27-35 (mm) Lobe length 9-10 <0.5 ca.21 mm 4.8-5 (mm Lobe glands Glandular- Eglandular Eglandular Eglandular fimbriate Pedice mee (mm) 5-8 12-13 9 - Pubescence Pilose Glabrous Pilose Pilose Corolla Length (mm) 45-47 ca. 50 nv. 45-48 Cross-section Terete to bluntly 5-winged over — nv. Terete to bluntly 5-angled entire 5-angled length Pubescence Glabrous to Short-pilose nv Glabrous sparsely pilose along angles Stamens Length (mm) ca. 43 ca. 50 nv. 45-46.5 Filaments Connate Connate nv. Connate veinlets inconspicuous to obscure, weakly impressed adaxially and weak raised abaxially; petioles subterete, rugose 4-10 x 2-4 mm, densely long-pilose with simple trichomes. Inflorescences axillary, racemose, pendent, to (10-)ca. 30-flowered; rachis persistent, subterete, bluntly angled, ca. 4-8 cm long, the SS LUTEYN, NEW SPECIES OF ERICACEAE FROM ECUADOR 1277 Fic. 3.0 b nn habit By i t howing leaf und f 1i f put C,portion f stem showing leaf peti ipular bud scales. D, ‘flower bud with calyx and bracteoles. E, corolla with longitudinal section showing relative Sicition of stamens. F, longitudinal section of calyx and inset of lobe margin showing ample halts by aandhlat fimbriae. G, stamens showing lateral, dorsal, ventral views with inset of dehis- Lewis et al. 2640) t 7F 1278 BRIT.ORG/SIDA 21 3) proximal several (to 4) nodes bearing sterile bracts; floral bract 1, lanceolate, long- acuminate, 11-16 x 3-5 mm, the venation conspicuous, moderately short-pilose with simple trichomes, marginally glandular-fimbriate with multicellular, multiseriate trichomes; pedicel subterete, striate, 10-14 mm long, densely short- pilose with simple trichomes and also short-glandular-fimbriate with multi- cellular, multiseriate trichomes, articulate with calyx; bracteoles 2, alternate, located basally to distally along pedicel, similar to floral bract but 6-16 x 2.5- 3 mm. Flowers 5-merous, pendulous; calyx 17-20 mm long, short-pilose with simple trichomes and also sometimes short-glandular-fimbriate with multi- cellular, multiseriate trichomes, the tube cylindric to obconic, terete in cross- section, 4-6 mm long, densely short-pilose with white to yellowish trichomes, the limb slightly spreading, 13-17 mm long, moderately short-pilose, the lobes membranous, concave, ovate, acuminate, 11-13 x 4-5 mm with venation con- spicuous, the sinuses acute; corolla membranous (fleshy when fresh), weakly bistratose, broadly and bluntly pentagonal in cross-section, cylindric and only slightly expanded basally, 45-57 x 6-9 mm, red to scarlet when fresh, short- pilose with whitish to reddish simple trichomes, also short-glandular-fimbri- ate with multicellular, multiseriate trichomes, the lobes wide-spreading and slightly reflexed exposing the stamens, lanceolate, bluntly acute, 7.5-13.5 x 2- 5mm, green when fresh; stamens 10, nearly equaling corolla in overall length, alternately slightly unequal with each other, 45-51 mm and 46.5-53.5 mm long, the filaments distinct, alternately 9-11 mm and 9.5-13 mm long, glabrous, the anthers alternately 38-42 mm and 40-44 mm long, the thecae alternately 12- 13 mm and 13-14 mm long, the tubules 2, alternately 25-29 mm and 28-31 mm long, distinct in distal 1/2-1/3, dehiscing by introrse, oblique, short clefts ca. 15-2 mm long; style shortly exserted, 48-59 mm long, glabrous, red to pink with green apex when fresh. Fruita spherical, short-pilose berry at least 13 mm diam., apparently translucent pale greenish when mature. Distribution —Endemic to Ecuador, where it occurs in rocky outcrops of “Southern Ecuadorean Scrub” vegetation along a very narrow and local cloud belt zone, at ca. 2600-3100 m. Common associates include Puya and Pitcairnia (Bromeliaceae), Macleania (Ericaceae), and lichen-covered boulders. Some co- rollas have holes at their bases made by nectar robbing birds. The fruit is said to be edible and a local common name is “salapa blanca grande.” Ceratostema pubescensis characterized by having a coarsely shrubby habit, densely pubescent leaf blades, long and narrow bud scales that appear pseudo- stipular, multi-flowered and racemose inflorescences, elongate bracteoles, large calyces and corollas, terete calyx tubes, conspicuously veined calyx lobes, and bluntly 5-angled corollas with proportionately short lobes. In Luteyn’s (1996) key to the Ecuadorean species of Ceratostema, this new species would be found near- est C. fasciculatum Luteyn, which differs morphologically by its basally cuneate and short-attenuate leaf blades (vs. rounded to subcordate), fasciculate and LUTEYN, NEW SPECIES OF ERICACEAE FROM ECUADOR 1279 4-6-flowered inflorescences (vs. racemose and to 30-flowered), longer floral bracts (20-26 mm vs. 11-16 mm), longer bracteoles (5-24 mm vs. 6-16 mm), overall longer calyx (20-27 mm vs. 17-20 mm), shorter corolla (38-43 mm vs. 45-57 mm), shorter stamens (36-40 mm vs. 45-51 mm), and eastern slope geo- graphical distribution (ie, Zamora-Chinchipe vs. western slope El Oro). The exact phylogenetic relationship of the new species awaits further study. Additional collections examined: ECUADOR. El Oro: same as ie 6 Nov 1997 (£1), Lewis et al. 3687 (AAU, Env, GBn.v, K n.v, LOJA n.v, MO nv, NY, Pn.v, QCA nv, QCNE n.v, US n.v.), 1 May 1997 (ster), Luteynet al. 15066 (NY, QCA): Chilla, Km 7, track to e antennas and paramo, 3°27'S, 79°306'W, 3100 m,7 Nov 1997 (fl, Lewis et al. 3700 (AAU, Env, GB nv, Aen ., LOJA n.v., MO nv, NY, Saae ce n.v., US n.v.); Chilla-Pueblo Viejo road, trail above Pueblo Viejo, 3° 55S 79°36'W, ca. 2800 May 1997 (im fr), Luteyn et al. 15071 (NY, QCA), Chilla-Pueblo one 3°28'S, 79°43'W, 2780 m, 28 ee 1996 (fr), Van den Eynden & Cueva 630 (LOJA n.v, NY). DISTERIGMA (Klotzsch) Niedenzu Disterigma is a montane genus of about 35 species that is characterized by its usually small leaves, sessile to subsessile flowers, and pedicellary bracteoles that are apical and surround (sometimes tightly clasp) the calyx and some- times the proximal parts of the corolla. It ranges from Guatemala south to Bo- livia and east to Guyana. The genus has been considered related to Vaccinium on the basis of morphology, although recent molecular studies (Powell & Kron 2003) place it in a clade with Sphyrospermum. The genus is currently being monographed by graduate student Paola Pedraza at The New York Botanical Garden saree bracteatum Luteyn, sp. nov. (Fig. 4). Type: ECUADOR. Azuay: Jesus Maria- uro-Cuenca road, 22.3-25.2 km E of Coastal Highway at Jesus Maria, ca. 2°37'S, 79°14'W, ae ee m, 23 Nov 1998 (£1), J. L. Luteyn & H. Mogollén 15401 (HOLOTYPE: NY, ISOTYPES: AAU, 5G. h, MO2,0GA, OCNE. 5, U5). ae ee a sete ne are ; 1 ae] We ina DO latis | li il 11mm longis, L lobis calycinis anguste lanceolatis usque 5-6 mm longis atque Sau 5 geniculatis. Terrestrial to epilithic, spreading shrubs with branches somewhat pendent to 3 m long; mature stems terete, striate, glabrous, brownish, the bark cracking longitudinally into parallel strips; twigs subterete, bluntly and broadly angled, glabrous to weakly puberulent, grayish-brown; axillary buds with outer pair of scales 2, valvate, ovate, acuminate, glabrous, ca. 2.5 mm long, the inner series of scales numerous, lanceolate to ovate-lanceolate, acuminate, striate, glabrous, brown, to 20 x 7mm, persistent at base of stems for at least three seasons. Leaves alternate, congested, the blades succulent and thick-coriaceous when fresh, wrinkled when dry, slightly revolute, elliptic to ovate-elliptic, 18-35 x 12-2.5cm, basally rounded to broadly obtuse, apically rounded to broadly acute, glabrous on both surfaces, the venation obscurely 3-5-pliner ved from the base with only the midrib scarcely visible adaxially; petioles terete, rugose, 25-5 mm long, weakly 1280 BRIT.ORG/SIDA 21(3) Disterigma- racteaktum- Fic. 4. Dist section. C, infl 7 oer J A, habit. B ti f st howina | 1 inf] wth dapail n€] 7 7 In ¢£I L . g le, calyx and corolla. E, | y J h immature inflorescences. H, details of Mogollon 15401 & 15404). ufey Jt |, dorsal, and lateral views. G, portion of stem showing fruits and L > + L rT 1 IAL ref a J f, n 2, \ Pi LUTEYN, NEW SPECIES OF ERICACEAE FROM ECUADOR 1281 Te fl f. aa puberulent m inshape, of solitary flowers, seemingly aris- ing from the axils of each leaf of a current season’s growth, circumscribed by a series of numerous (ca. 23), ovate to lanceolate, acuminate to long-acuminate, weakly striate, scarious or brownish, glabrous but deciduously fimbriate-mar- gined bracts up to ca. 11 x 4 mm that cover the calyx and lower ca. half of the corolla and persist at least until the fruits mature, the third innermost bract (i.e., the floral bract) morphologically indistinguishable from the other inflo- rescence bracts ca. ll x 4 mm, the two innermost bracts (i.e., the bracteoles) also morphologically indistinguishable from the other inflorescence bracts ca. 7-9 x 3-4 mm; pedicel none, replaced by a series of overlapping nodes covering <0.5mm length. Flowers 5-merous; calyx ca. 8-11.5 mm long, glabrous or some- times weakly short-pilose (especially the lobe tips), the tube barrel-shaped, ca. 2-4.5 mm long, the limb cylindric, ca. 6-7 mm long, the lobes narrowly lan- ceolate, long-acuminate, ca. 5-6 mm long, marginally fimbriate, the sinuses acute; corolla cylindric, narrowing at base and to throat, somewhat pentagonal to 5-angled in cross-section, ca. 7.7-15 X 6.5 mm, bright red, glabrous, the lobes deltate, ca. 1.2-2 mm long, bluntly acute; stamens 5, shorter than corolla in over- all length, equal with each other, ca. 10-11 mm long, the filaments geniculate, distinct, ca. 4-4.5 mm long, glabrous, the anthers ca 7-7.4 mm long, the thecae ca. 3.6-3.9 mm long, the tubules 2, distinct to base, ca. 3.4-3.5 mm long, dehise- ing by introrse, elongate clefts ca. 25-3 mm long; style included, + equaling corolla. Fruit a spherical, dark purple berry, 7-8 mm diam., crowned by persis- tent calyx lobes. Distribution —Endemic to Ecuador, where it occurs on rocks and rock out- crops asa low, spreading shrub within montane cloud forest habitats, at ca. 975- 2600 m Disterigma bracteatum is characterized by having succulent leaves, numer- ous (ca. 23), lanceolate, brownish, persistent bracts to 11 mm long that surround the vegetative branches, inflorescences and fruits, calyx lobes that are narrowly lanceolate to 5-6 mm long, and five geniculate stamens. In Luteyn’s (1996) key to the Ecuadorean species of Disterigma, this new species would be found near D. pentandrum SF Blake and D. rimbachii (A.C. Sm.) Luteyn, all three species characterized by possessing succulent leaves, solitary flowers surrounded by a series of brownish bracts, and five stamens. Disterigma bracteatum differs from D. rimbachii, which has few, caducous bracts to 2 mm long, calyx lobes deltate ca. | mm long, and bracteoles 1-2 mm long, and from D. pentandrum, which has about six persistent bracts to 7 mm long, calyx lobes ca. 3.5 mm long, and bracteoles ca. 7 mm long. — Additional collecti 1: ECUADOR Azuay: Jesus Maria-Molleturo-Cuenca road, 0.9 km to- wards Molleturo from turn-off from highway that is 56 km E of Jesus Maria, ca. 2°42'S, 79°13'W, ca. 2438 m, 23 Nov 1998 (fr), Luteyn & Mogollén 15404 (AAU, COL, GB, NY, QCA, QCNE, W), 2600 m, 27 1282 BRIT.ORG/SIDA 21(3) Dee 2003 ED, Pedraza & Pedraza 1016 (COL, NY, QCA, QCNE), 10 kms ane Molleturo, ca. 2°46'N, 79° 24'W, 2600 m, 27 Dec 2003 (f1), I t -edrazad & Pedraza lOI7 (NY, QC A, QCNE ACKNOWLEDGMENTS | wish to thank Gwilym P. Lewis and Veerle Van den Eynden for making special efforts to collect Ceratostema pubescens for me and guiding me to its location; Bertil Stahl for providing pickled floral material; Robin B. Foster for providing beautiful photos of C. oyacachiensis; Bobbi Angell for the wonderful illustra- tions; and Patricia Eckel for providing the Latin diagnoses. The National Sci- ence Foundation (grants BSR-9024221 and DEB 9628841) provided funds for field work in Ecuador, and the Missouri Botanical Garden provided facilities to write portions of this paper. Thanks to Walter Judd and Larry Dorr who made helpful review comments. REFERENCES Luteyn, J.L. 1984. Revision of Semiramisia (Ericaceae: Vaccinieae). Syst. Bot. 9:359-367. Luteyn, J.L. 1986. New species of Ceratostema (Ericaceae: Vaccinieae) from the northern Andes. J. Arnold Arbor. 67:485-492. Luteyn, J.L. 1996. Ericaceae. In:G. Harling and L. Andersson, eds. Flora of Ecuador 54:1-404, color plates I-VIII. Luteyn, J.L. 1998 [ver. 2002]. Neotropical blueberries: The plant family Ericaceae. nw.nybg.org/bsci/res/lut2, Luteyn, J.L. 2002. Diversity, adaptation, and endemism in neotropical Ericaceae: biogeo- graphical patterns in the Vaccinieae. Bot. Rev.68:55-87 Powett, EA. and KA. Kron. 2003. Molecular systematics of the northern Andean blue- berries (Vaccinieae, Vaccinioideae, Ericaceae). Int. J. Plant Sci. 164:987-995. — NOMENCLATURE OF IPOMOEA ARBORESCENS (CONVOLVULACEAE) IN SONORA, MEXICO Daniel F. Austin Richard Felger Arizona-Sonora Desert Museum Drylands Institute 2021 N. Kinney Road PMB 405, 2509 N. Campbell Avenue Tucson, Arizona 85743, U.S.A. Tucson, Arizona 85719, U.S.A. Thomas R.Van Devender Arizona-Sonora Desert Museum 2021 N. Kinney Road Tucson, Arizona 85743,U.S.A. ABSTRACT Ipomoea arborescens (tree morning glory) has two varieties in the state of Sonora, Mexico. Author citation of I. arborescens var. glabrata (a nomenclatural synonym of I. arborescens var. arborescens) is corrected. The second taxon, |. arborescens var. pachylutea, was named by Howard Scott Gentry from the Rio Mayo region. These plants differ considerably from I. arborescens var. arborescens in having different bark morphology and corolla tube colors. RESUMEN En el estado de Sonora, México, Ipomoea arl pal ) ti bres para dos variedades. ral Ceri, Perea nar teentoa | do] 1 eel hasan ees : 1 o £ oO L var. arborescens. Howard Scott Gentry nombro I var. pachylutea de la region del Rio Mayo, la cual se a principalmente de la var. arborescens por su corteza amarillenta y corolas con tubos morados During studies of the Convolvulaceae for both the Flora Mesoamericana (Aus- tin et al. in prep.), and the Trees of Sonora, Mexico (Felger et al. 2001), it became apparent that there is nomenclatural confusion with plants called Ipomoea arborescens. In addition, there is biological uncertainty about the taxonomic delimitations of these Mexican trees. This paper will address the nomencla- tural problems. When Gentry (1942) was ee the plants of the Rio Mayo region of southern Sonora, he encountered tw varieties of this tree. One of these he called I. arborescens var. fe and the other he named I. arborescens var. pachylutea. Although the most recent revision of the group by McPherson (1981) does not mention either variety, subsequent field studies by various botanists (e.g., Martin et al. 1998; Van Devender et al. 2000; Felger et al. 2001) make it clear that in Sonora there are indeed two distinct morphotypes subsumed by the binomial I. arborescens. There is as well a third morphotype farther south for which we have insufficient data to completely compare with Sonoran plants. SIDA 21(3): 1283-1292. 2005 1284 BRIT.ORG/SIDA 21(3) —— The species is widespread and may be characterized by the following: Ipomoea arborescens (Hlumb. & Bonpl. ex Willd.) G. Don, Gen. Syst. 4:267. 1838. Convolvulus arborescens Willd., Enum. Pl. 204. 1809. Type: MEXICO. GUERRERO: between Acaguisotla and Chilpancingo, Humboldt & Bonpland 3927 (HOLOTYPE: P; microfiche seen). Trees 5-15 m tall; the trunk thick, often 50-70 cm diameter, the bark gray, whit- ish or pale yellowish, stems with abundant latex, tomentose when young with trichomes 0.1-2.5 mm long, glabrescent. Leaves 9-27 cm long, 6-9+ cm wide, ovate to lanceolate, glabrescent, the apex acuminate, the base cordate, with tri- chomes longer that those on the branches, velvety at least below, the midrib base Gust above the petiole) with a pair of blisterlike glands 1-3 mm in diam., turgid on young, enlarging leaves (these glands are the same color as the midrib and may be difficult to see, especially on older or dried specimens). Inflores- cences terminal or axillary, monochasial, racemose. Flowers |(-2); sepals 6-14 mm long, ovate to rarely orbicular, more or less equal, tomentose, the apex ob- tuse to obtuse-mucronate; corollas 4-4.3 cm long, often 6.5-9.5 cm wide, fun- nelform, tomentose at least on the lobes, white, tube green without. Fruits 17- 25 mm long, capsular, 4-valvate, brown, glabrous; seeds 1-4, 9-16 mm long, brown, pilose on the margins with trichomes 10-15 mm long. Illustrations.—Martinez (1969: 237), Felger et al. (200L 139-141). Common names.—Tree morning glory; cazdhuate (from Distrito Federal to Oaxaca), juituguo (Mayo, southern Sonora), osi (Tarahumara, Chihuahua), palo blanco (Sonora), palo santo (Sonora), patancdan blanco and rosi (Durango, Jalisco, Michoacan, Guerrero), tochiyé (Guarijio, SE Sonora). Flowering November-April; near sea level-1800 m; Sonoran desertscrub, thornscrub, tropical deciduous forest, and oak woodland or rarely at the lower edge of “tropical” pine-oak forest (Fig. LD. The type locality is between Acapulco and Cd. México, in a mid-elevation y dry tropical zone. The original vegetation at the type locality was — seasonal probably tropical deciduous forest. That region is the southern part of the “typi- cal” habitat of the modern known range of 1. arborescens, although the species ranges into the highlands of Edo. México, Michoacan, and Morelos. Gentry (1942) called the “smaller, less pubescent-leaved form characteris- tic of the species throughout the foothill regions of southern and central Sonora” var. glabrata. The trees that extend into the Sonoran Desert North of Hermosillo are probably the same taxon as Gentry’s variety glabrata. Trees of the lowland and northern Sonora populations have conspicuously lighter-colored (whiter) bark, and flowers with the purple coloration much reduced or lacking when compared with var. pachyluted. AUSTIN ET AL., NOMENCLATURE OF IPOMOEA ARBORESCENS 1285 ee ee ne: \ oe) e | e oceee fo [on ff ° . a eS , oR ‘J (* 7" R e vA y | | J ° q ; e " e e | , @ R R e e ® e | e e eo e f ® : RI Mexico | e | e | ; O setions! captal | ed 2 e J | Fic.1.Distributi ££] I AOE pres Nat harl Turner et al. 1995; R=reportby Torres: -R, (2004). Based on Le eae Matuda (1963, 1966), McPI (1981), Cowan (1983), Torres-R. (2004) + MEX KEY TO SYNOPTIC TRAITS Habitat Sonoran desert-scrub to tropical deciduous forest and lower oak woodla white. Flowers white with yellow or diffuse pale purple dots and short bands within the tube (Figs. 2, 3). Flowers in Sonora visited by bees, hawkmoths, and hum- mingbirds, south of Sonora by bats Ipomoea arborescens var. arborescens Habitat humid tropical deciduous forest and oak woodland. Bark yellowish. Flowers solid or almost solid dark-purplish within the tube. Flower visitors unknown Ipo moea arborescens var. pachylutea NOMENCLATURE Ipomoea arborescens (Humb & Bonpl. ex Willd.) G. Don var. arborescens. Ipo- moea arborescens (Humb & Bonpl.ex ie Don var. glabrata Gentry, Car Poe West Publ. 527:212. 1942. TPE: MEXICO. SONORA: Arroyo Cuchijaqui, G 870 entry 870 (SYNTYPE: D Se EVE: cited by McPherson); San Bernardo, Gentry 1158 (LECTOTYPE, here chosen: ARIZ! ISOLECTOTYPE: ; i murucoides var. glabrata Rose, Contr. US. — Herb. L107. 1891, non A. Gray (1887). Type: XICO. Sonora: Alamos, Palmer 316 GioLotyee: US 1286 BRIT.ORG/SIDA 21(3 > + ~ | =a D ral v _ + - 2. A Fic. 2. Corollas of /pomoea arborescens var y purple in the corolla throats are distinctive. corolla throats are distinctive. AUSTIN ET AL., NOMENCLATURE OF IPOMOEA ARBORESCENS 1287 When Gentry made this combination, he cited the correct protologue by Rose, but incorrectly listed Gray as the author. Since the varietal name by Rose was a later homonym of var. glabrata A. Gray, we are interpreting Gentry’s action as creation of a new name (Article 58). Tentatively, we consider this a nomencla- tural synonym of L arborescens. McPherson (1981) al nsidered the glabrous forms of the plants asa synonym, because he cited the type of L murucoides var. glabrata Rose insynonymy with L. arborescens. As pointed out by Gentry (1942), there are differences between these northern plants and those farther south, but we do not have enough data on the southern populations to determine if the two should be considered nomenclaturally distinct. These are the plants that Soderholm and Gaskins (963) called I. wolcottiana. The USDA collection forming the basis of their report grows in Miami at both the Plant Introduction Station and the Fairchild Tropical Garden and its iden- tity has been verified as I. arborescens var. glabrata. Presumably that report is also the basis of the incorrect report of 1. wolcottiana from Sonora. That species has not been documented in Sonora. — Ipomoea arborescens (Humb & Bonpl. ex Willd.) G. Don var. pachylutea Gen- try, Carnegie Inst. Wash. Publ. 527:213. 1942. TYPE: MEXICO. SONORA: Sierra de Alamos, Gentry 3000 (LECTOTYPE: ARIZ! IsoTypEs: MO!, UC, n.v., US!). PARATYPE: MEXICO. SONORA: Algodones, Sierra Charuco, Gentry 2299 (ARIZ), selected 3000 ft {6lO-915 ml]. Vernacular: palo sunto amarillo. Large trees with massive trunks & yellowish park browning with age; 8-15 m high. Petioles and twigs with milky juice, Gentry 4888 (ARIZ); Sinaloa Las Mesas, Sierra Surotato, 25 Aug 1941. Oak-Ipomoea savanna; volcanic ash, elev. ca. 3000 ft [915 m]. Vernacular: palo blanco. Tree with yellow bark. Co-dominant with oak. Gentry 6144 (ARIZ). Addi- tional specimens were cited by Martin et al.(1998), McPherson (1981), and Turner et al. (1995) mapped additional specimens seen: Sonora, Sierra de Alamos, rocky slopes and canyon bottoms, 2000- the species. Common name.—Palo santo amarillo. sentry (1942) named the higher elevation or montane populations in southeastern Sonora and adjacent southwestern Chihuahua var. pachylutea,and distinguished these trees from I. arborescens var. glabrata in having “...yellowish bark, larger and more pubescent leaves, longer and stouter pedicels, larger and more numerous flowers, larger sepals, and generally heavier inflorescences.” Other differences include wood that is apparently not as soft, corollas with a prominently maroon-purple throat, and pale lavender filaments. Ipomoea arborescens var. pachylutea occurs at ca. 5300-1220 m in tropical deciduous for- est and the lower oak zones. To date, variety pachylutea is known only from Sonora and Chihuahua, but no comparison has been made with living plants farther south. The southern limits of var. pachylutea remain unknown. Both varieties occur in southeastern Sonora, but do not occur intermixed with minor excep- tions. Near Alamos, Sonora, a single tree of var. pachylutea was found in an area 1288 BRIT.ORG/SIDA 21(3) of var. arborescens trees. Such trees, however, are in disturbed. partially cleared areas at elevations near the usual lower elevational limits of var. pachylutea. —— Trees of intermediate character are not known, and the varieties abruptly re- place each other. We suspect that the two “varieties” are actually cryptic species. Grown side-by-side at the Arizona-Sonora Desert Museum in Tucson from seed collected in Sonora, plants of var. pachylutea bear leaves nearly twice as large as those of var. arborescens (var. glabrata sensu Gentry 1942). In the field, Sonoran populations of these varieties exhibit overlap in leaf sizes. In Arizona plantings, variety pachylutea tends to branch near the base of the plant with large horizontal branches, w — hile living specimens of var. arborescens generally do not share these features and developa thick trunk at an earlier age. However, — in dense forests near Alamos, Ipomoea arborescens var. pachy tree. uted isan upright One of the difficulties in applying these names to herbarium specimens is that there typically are few obvious traits retained that can be used in distin- guishing the two taxa, even though living trees are quite distinctive. When flow- ers are present, dissection of the corollas will reveal the needed ‘comparative’ differences in purple within the base of the tube. Even when both forms are not available, the two may be separated relatively easily. The inside of the tube of var. pachylutea is solid or nearly solid purple throughout; tubes of var. arborescens are variable but usually lighter in color—diffusely colored with bands and dots of light purple. Beyond that trait, we have found no consistent herbarium differences in the sizes of leaves or flowers that Gentry noted, as there uL is considerable variation throughout the range of the species. Although no field phenology data are available for Sonora, trees grown in Tucson flower at different times. Data taken over 10 seasons on var. arborescens indicate that its beginning flowering dates vary from the third week in Octo- ber to the third week in January. However, the trees cease flowering before the third week in February. By contrast, two season's data show that var. pachylutea does not begin flowering until the second week in March. Thus, a full two weeks separate the flowering periods. Based on herbarium specimens at ARIZ, these differences do not hold for wild plants in Sonora where flowering seasons of both varieties overlap. — The floral biology is variable and is largely temperature-dependent (Al- berto Burquez M., pers. comm. 1998; Francisco Molina F, pers. comm. 2001). The flowers commonly open in the late afternoon. During warmer weather, the co- rollas fall the next morning with warming daytime temperatures, but on cooler days they often do not fall until the second night. This is consistent with a num- ber of matinal and/or nocturnal species in the family (Austin, unpubl. data). The trees are self-incompatible (Alberto Burquez M., pers. comm. 1998): buds attract both red and black large ants (Felger et al. 2001), perhaps Camponotus. In Sonora, Hyles lineata (white-lined sphinx moth) and perhaps other oa AUSTIN ET AL., NOMENCLATURE OF IPOMOEA ARBORESCENS 1289 hawkmoths and hummingbirds are the primary pollinators of var. arborescens (Felger et al. 2001). The flowers are visited by hummingbirds in tropical de- ciduous forest and foothills thornscrub from the Alamos area north to Santa Ana de Yécora and Tepoca, and northwest to the Hermosillo area in the Plains of Sonora subdivision of the Sonoran Desert (Van Devender et al. 2004). Cynanthus latirostris (broad-billed hummingbird) and Calypte costae (Costa’s hummingbird) are the most common visitors to I. arborescens while Calpyte anna (Anna’s hummingbird) are sporadically seen. Other avian visitors such as montane species that come down from the pine-oak forest and oak wood- land of the Sierra Madre Occidental into the tropical deciduous forest for the winter, include Amazilia beryllina (berylline hummingbird), Heliomaster constantii (plain-capped starthroat), and Hylocharis leucotis white-eared hum- mingbird). Bees also visit the plants near Hermosillo (Francisco Molina, pers. commn., 2004). While agaves (Agave spp.), ceibas (Ceiba grandiflora), yuccas (Yucca spp.), saguaros (Carnegied gigantea), organ pipe cactus (Stenocereus thurberi), cardon (Pachycereus pringlei), and other cacti are famous for being pollinated by bats (Arizaga et al. 2000; Casas et al. 1999; Fleming et al. 1998; Nassar et al. 1997; Quesada et al. 2003; Stoner et al. 2003; Valiente-B. et al. 1997), these flying mam- mals do not visit only these plants exclusively. Indeed, flowers on tree Ipomoea may supply nectar during the season when few other bat-pollinated plants are in bloom (Hevly 1979; Turner et al. 1995). Pollinators of Ipomoea arborescens var. pachylutea are not known, although the white limb and lavender throat may suggest bat-pollination, but bats have not been seen visiting the flowers of I. arborescens in Sonora. Farther south in Guerrero, México (Baker et al. 1977; Butanda C. et al. 1978), L arborescens and some other species of tree morning glory are visited and presumably pollinated by Choeronycteris mexicana (Mexican long-tongued bat), Glossophaga soricinia (long-tongued bat; Villa R.1966), and Leptonycteris curasoae (lesser long-nosed bat). Although L. curasoae is reported feeding from ‘I. arborescens’ in Hidalgo (Baker et al. 1977), that species does not occur there (Carranza-G. et al. 1998); instead the plants probably are I. rzedowshii, although L. murucoides also grows there. Leptonycteris curasoae is known to consume Ipomoea pollen in Sonora (Hevly 1979) and L.arborescensis one of the few species of Ipomoea sturdy enough for bats in the state. Because the mammals migrate north from central and southern Mexico to southern Arizona and back south during different seasons (Moreno-Valdez et al. 2000; Newton et al. 2003; Wilkinson et al. 1996), a variety of food sources are critical for their survival and reproduction. It is noteworthy that the flowering seasons of the two varieties of IL. arborescens are different in the Arizona-grown plants because they correspond with the migration dates of the bats. Leptonycteris arrives in Organ Pipe Cactus National Monument in April and 1290 BRIT.ORG/SIDA 21(3) May (Hoffmeister 1986; K. Krebbs & T. Tibbitts pers. comm., Sep 2004) just af- ter var. pachylutea has flowered in Sonora. Adult female Leptonycteris have left Arizona by late September, and the juveniles leave later (K. Krebbs pers. comm., Sep 2004). Ipomoea arborescens var. arborescens in Sonora is in flower from October through the end of the year. Therefore, when the bats leave Arizona and fly through northern Sonora, the tree [pomoed are not in flower. However, in about the southern half of Sonora, there are at least a few ani- mals present during the winter. Francisco Molina (pers. comm., 2004) informed us that “we have captured Leptonycteris in Hermosillo (Centro Ecologico) in mid-February visiting Agaves. We have also captured Leptonycteris in Febru- ary at Rancho San Francisco (between San Jose de Pimas and Tecoripa) visiting etchos [Pachycereus pecten-aboriginum].” These winter resident Leptonycteris apparently are not present much north of Hermosillo (K. Krebbs, pers. comm. 2004); hence they are absent from most of the October to January flowering period of I. arborescens var. arborescens. On the other hand, Choeronycteris is present throughout the year in northern Sonora. Much less is known about Choeronycteris because it is not a colonial spe- cies gathering in maternity colonies like Leptonycteris (Hoffmeister 1986; K. Krebbs, pers.comm., Sep 2004). Some individuals remain in Arizona and north- ern Sonora throughout the winter (K. Krebbs, pers. comm., Nov 2004), although it appears that the majority migrate southward. Their young are born in late June (Hoffmeister 1986), and presumably the migrants arrive from Mexico near the same time as Leptonycteris. Thus, bats may utilize lL arborescens on their migration north through Sonora, but do not appear to do so when going south. — ~~ ACKNOWLEDGMENTS We thank Alberto Burquez M., Karen Krebbs, Ana Lilia Reina-G., Curtis McCasland, Rodrigo A. Medellin, Francisco Molina F, Guillermina Murguia S., Yar Petryszyn, Tim Tibbitts, and Michael F Wilson for information and aid in examining these plants, herbarium material, and photographs. RSF thanks the Wallace Research Foundation for support. Ana Lilia Reina G. kindly provided the Spanish abstract. Andrew McDonald and an anonymous reviewer provided helpful comments on an earlier draft of the manuscript. REFERENCES Arizaca, S.,£. Ezcurra, E. Peters, FR. of ARELLANO, and E.Veca.2000. Pollination ecology of Agave macroacantha (Agavaceae) in a Mexican tropical desert.|I. The role of pollinators. Amer. J.Bot.87:1011-1017. Austin, D.F., J.A. McDonato, and G, Murcuia-S. (in prep.). Convolvulaceae In: Flora Mesoamericana Baker, R.J., J. KNox Jones, Jr, and D.C. Carter (eds.). 1977. Biology of bats of the New World family Phyllostomatidae, Part Il. Texas Tech Press, Lubbock. AUSTIN ET AL., NOMENCLATURE OF IPOMOEA ARBORESCENS 1291 Butanpa-C., A., C. VAsquez-Y., and L. Treo. 1978. La polinizacién quiropterdfila: una revision bibliogrdfica. Biotica 3:29-35. Carranza-G., E., S. ZAMUDIO-R., and G. Murauia-S. 1998. Una especie nueva de /|pomoea (Convolvulaceae), de los Estados de Guanajuato, Hidalgo y Querétaro, México. Acta Bot. Mex. 45:31-42. Casas, A., A. VALIENTE-BANUET, A. RosAs-Martinez, and P. Davita. 1999. Reproductive biology and the process of domestication of the columnar cactus Stenocereus stellatus in central Mexico. Amer. J. Bot. 86:534-542. Cowan, C.P. 1983. Listados floristicos de México. |. Flora de Tabasco. Instituto de Biologia, Universidad Nacional Aut6noma de México, D-F. Feicer, R.S., M.B. JoHNsoN, and M.F. Witson. 2001. Trees of Sonora, Mexico. Oxford University Press, New York, NY. FLemina, T.H., S. Maurice, and J.L. Hamrick. 1998. Geographic variation in the breeding system and the evolutionary stability of trioecy in Pachycereus pringlei (Cactaceae). Evol. Ecol. 12:279-289. Gentry, H.S. 1942. Rio Mayo plants. A study of the flora and vegetation of the Valley of the Rio Mayo, Sonora. Carnegie Inst. Wash. Publ. 527,Washington, DC. Heviy, R.H. 1979. Dietary habits of two nectar and pollen feeding bats in southern Arizona and northern Mexico. J. Arizona-Nevada Acad. Sci. 14:13-18. Horrmeister, D.F. 1986. Mammals of Arizona. Univeristy of Arizona Press and Arizona Game and Fish Department, Tucson. Mastin, P.S.,D. YETMAN, M. FISHBEIN, P JENKINS, T.R. VAN Devenber, and R.K. WiLson (eds.).1998.Gentry’s Rio Mayo Plants. The tropical deciduous forest & environs of northwest México. Uni- versity of Arizona Press, Tucson. Martinez, M. 1969. Las plantas medicinales de México. Ediciones Botas, México. Matupa, E. 1963. El genero Ipomoea en Mexico (|). Anal. Inst. Biol. Univ. Nac. Méx. 34(1—2): 85-145. Matupa, E. 1966.Las Convolvulaceas del Estado de Mexico.Govierno del Estado de Mexico, Direccion de Agricultura y Ganaderia, Toluca, Mexico. McPuerson, G. 1981. Studies in Ipomoea (Convolvulaceae) |. The Arborescens group. Ann. Missouri Bot. Gard. 68:527-545. Moreno-Vatpez, A.,W.E. Grant, and R.L.Honeycurt. 2000. A simulation model of Mexican long- nosed bat (Leptonycteris nivalis) migration. Ecol. Modelling 134:117-127. Nassar, J.M.,N. Ramirez, and O. Linares. 1997. Comparative pollination biology of Venezuelan columnar cacti and the role of nectar-feeding bats in their sexual reproduction. Amer. J. Bot. 84:91 8-927. Newron, L.R., JM. Nassar, and T.H. Fieminc. 2003. Genetic population structure and mobility of two nectar-feeding bats from Venezuelan deserts: inferences from mitochondrial DNA. Molec. Ecol. 12:3191-3198. Parra-Tasia, V. 2002. /pomoea wolcottiana Rose (Convolvulaceae). Ozote. In: F.A. Noguera, J.H. Vega Rivera, A.N. Garcia Aldrete, and Q. Avendano. Historia Natural de chamela. Universidad Nacional Auténoma de México, D.F., México. Pp. 159-161. 1292 BRIT.ORG/SIDA 21(3) QuesabA, M., K.E. Stoner, V. Rosas-Guerrero, C. PAtacios-Guevara, and J.A. Logo. 2003. Effects of habitat disruption on the activity of nectarivorous bats (Chiroptera: Phyllostomidae) in a dry tropical forest: implications for the reproductive success of the neotropical tree Ceiba grandiflora. Oecologia 135:400-406. Rose, J.N. 1894.Some notes on the tree lpomoeas of Mexico. Gard. and Forest 7:364-367. SODERHOLM, P.K. and M.H. Gaskins. 1963. lpomoea wolcottiana, a tree morningglory. Amer. Hort. Mag. 42:111-113. STONER, K.E., K.A.O. SALAZAR, R.C. FeRNANDEZ, and M.Quesava. 2003. Population dynamics, repro- duction, and diet of the lesser long-nosed bat (Leptonycteris curasoae) in Jalisco, Mexico: implications for conservation. Biodivers. & Conservation 12:357-373. Torres Rojo, J.M. 2004. Especies con usos no maderables en bosques tropicales y subtropicales en los estados de Durango, Chihuahua, Jalisco, Michoacan, Guerrero y Oaxaca. URL: http://148.233.168.204/pfnm2/fichas/ipomoea_arborescens.htm. Turner, R.M., J.E. Bowers, and T.L. Buraess. 1995. Sonoran Desert plants. An ecological atlas. University of Arizona Press, Tucson. VaviENTE-BANUET, A., A. Rosas-Martinez, M. bet Coro Arizmenpi, and P. Davita. 1997. Pollination biology of two columnar cacti (Neobuxbaumia mezcalaensis and Neobuxbaumia macrocephala) in the Tehuacan Valley, central Mexico. Amer. J. Bot. 84:452-455. Van Devenper, T.R., A.C. Sanpers, R.K. Witson, and S.A. Meyer. 2000. Vegetation, flora and sea- sons of the Rio Cuchujaqui, a tropical deciduous forest near Alamos, Sonora. In: R.H. Robichaux and D.Yetman, eds. The tropical deciduous forest of Alamos. biodiversity of a threatened ecosystem in México. University of Arizona Press, Tucson, Pp. 36-101. VAN DEeveENDER, T.R., W.A. Cacper, K. Krepes, A.L. Reina-G., S.M. Russett, and R.O. Russett. 2004. Hum- mingbird plants and potential nectar corridors for the rufous hummingbird in Sonora, Mexico. In: G.P. Nabhan, R.C. Brusca, and L. Holter, eds. Conserving migratory pollinators and nectar corridors in western North America. Arizona-Sonora Desert Museum and University of Arizona Press, Tucson. Vitta-R.,B. 1966.Los Murciélagos de México. Universidad Nacional Aut6énoma de México. Instituto de Biologia, México, D.F. WILKINSON, G.S., T.H. Femina, and J. Witkinson. 1996. Migration and evolution of lesser long- nosed bats Leptonycteris curasoae, inferred from mitochondrial DNA. Molec. Ecol. 5: 329-339 IPOMOEA SEAANIA, A NEW SPECIES OF CONVOLVULACEAE FROM SONORA, MEXICO Richard Felger Daniel F. Austin Drylands Institute Arizona-Sonora Desert Museum PMB 405, 5, 2509 N. Campbell Ave. 2021 N. Kinney Road Tucson, Arizona 85719, U.S.A Tucson, Arizona 85743, U.S.A. ABSTRACT Ipomoea seaania is described as a new species from the vicinity of Guaymas, Sonora. These plants are confined to the Sierra El Aguaje, although they perhaps grow in other nearby unexplored moun- tains. The relationships of this species to the other members of Ipomoea series Arborescentes are dis- cussed, and all taxa in the group are listed and ranges given. Ipomoea seaania brings the total num- ber of taxa in the series to 13 species, with I. arborescens having two varieties and L. pauciflora and L wolcottiana each having two disjunct subspecies. RESUMEN describe Ipomoea seaania dee as cercanias de Guaymas, Sonora, como una especie nueva. Estas la Sierra El Aguas jue q | 1 i I de Ipomoea serie cercanas sin ie ae Se discuten mismo modo se listan ged: los taxa del grupo y se proporcionan sus rangos de distribucion. Con el reconocimiento de ee da seddnid como nueva especie, se reconocen un total de 13 especies en el gru elas cuales | variedadese I. duct lorae £ r I. wolcottiana dos ee When Old World botanists began discovering morning glories in the New World, most species they found had life forms like the twiners Calystegia, Con- volvulus, and Cuscuta they knew at home. Their concept of the family was some- what broadened when they found erect and sprawling herbs in the Americas, and they were amazed when they found morning glory trees (Austin 2004). In 1809, Humboldt, Bonpland and Willdenow called the first known tree species Convolvulus arborescens, the distinctions between Convolvulus and Ipomoea being unclear at the time. These trees still are considered “odd” or “unusual” in the family, and the only other genus in the family that achieves tree stature is the Malagasian Humbertia (cf. Pichon 1947; Deroin 1992). Anatomically, these American trees are distinct from other shrubby and woody members of the family (Austin 1971; Carlquist @ Hanson 1991; McDonald 1992; Deroin 2001). The arborescent species of Ipomoea in the New World have long been of interest to the people who lived with them. Indigenous people use several spe- cies (Hersch-M. 1995; Yetman & Felger 2002; Yetman & Van Devender 2001), and the chemistry of the group is somewhat distinctive (Pérez-A. et al. 1982, 1983, 1992a, b). Three of these alkaloids (3a-(4-hydroxybenzoyloxy )tropane, 3a- SIDA 21(3): 1293-1303. 2005 1294 BRIT.ORG/SIDA 21(3) (4-meth ybe 1ZC yl xy )nortr opane, phyllal bine) are considered rare constitu- ents, spe ily in ie genus Ipomoea (E. Eich, pers. comm., 26 Jul. 2004). Of additional interest was the discovery that nectar-feeding bats are at least sea- sonal flower-visitors and pollinators in some species (Butanda-C. et al. 1978; Carranza-G. etal. 1998; Casas et al. 1999; Fleming et al. 1998; Hevly 1979; Moreno- V.et al. 2000; Nassar et al. 1997; Newton et al. 2003; Quesada et al. 2003; Stoner et al. 2003; Turner et al. 1995: Valiente-B. et al. 1997; Wilkinson et al. 1996). These bats are Choeronycteris mexicana, Glossophaga soricina, and Leptonycteris curasode (= L. sanborni, L. yerbabuenae). The Leptonycteris has been listed as endangered in the United States (Reid 1997; US. Fish and Wildlife Service 1997) since 30 September 1988. (See accompanying paper by Austin, Felger & Van Devender p. 1283-1292 for discussion.) As summarized in Table |, there are 13 species in the American series Arborescentes (Austin & Huaman 1996; McDonald 1991; McPherson 1981: Murguia-S. et al. 1995; Carranza-G. &@ McDonald 2004). Most of the species are confined to Mexico and nearby Mesoamerica (Austin 2001; Austin @ Huaman 1996, Austin et al.in preparation), but there are two with disjunct subspecies in western South America (Austin 1982; McPherson 1981). The first author found L seadania in Sonora during 1980 and again in 1985 and located additional herbarium specimens. Although we talked about the plants in 1989, neither of us had the opportunity to pursue them further. Fi- nally, we have been able to compare the known taxa with these plants. Regarding morphology and range, this isa markedly distinct species (Table Ll). Using the key in McPherson (1981) these plants come out at I. chilopsidis. Leaves on the two are the most obvious distinction on herbarium specimens. Both have narrow leaves, often less than one cm wide, but they are 10-20 cm long in Ll chilopsidis and only 4-8 cm in I. sedania. Flowers are large (8-9.5 cm long) in I. chilopsidis, but only 4-6 cm in I. seaania. Furthermore, the overall architecture of the plants is profoundly different. Ranges and altitudinal differences also are pronounced. Ipomoea chilopsidis is a plant of “high and arid craigs” (Gentry 2391, ARIZ) of the Sierra Madre Oc- cidental, ranging from the eastern border of southeastern Sonora through about half of the southern end of Chihuahua. Near the border between Chihuahua and Sonora L. chilopsidis grows at 1100-1800 m in oak and pine-oak forest (Gen- try 1942, Martin et al. 1998). On the other hand, I. seaania is known only from the vicinity of Guaymas in west-central Sonora where it grows near the southern margin of the desert on rocky slopes near the Gulf of California. The Guaymas region uplands are more than 1,000 m lower than those of Chihuahua. All records for I. sedania are from essentially the same locality, below ca. 20 m elevation and near a road, except one collection (Felger 80-36 et al.) which is from a nearby canyon probably one kilometer eastward. The rugged slopes immediately above this area have yet to be explored. The canyons where the — . 5 16570 Scirpus polyphyllus Vahl, Lohr 160 DIOSCOREACEAE Dioscorea villosa L., (1996) Isaac, B.L.; Isaac, JA. COXE ET AL DIPSACACEAE *Dipsacus fullonum L., (2003) Isaac, J.A. 16629; Exotic ELAEAGNACEAE *“Elaeagnus umbellata Thunb., (2003) Isaac, J.A.; Isaac, B.L. 15820; Asia ERICACEAE Epigaea repens L.,(1951) Buker, W.E. Gaultheria procumbens L., (1950) Henry, L.K.; Buker, W.E / inh aylussa accata (Wangenh.) K. Koch, (2004) Isaac, J.A. 17375 Kalmia latifolia L., (1951) Henry, L.K.; Beer, FH. ydendrum arboreum (L.) DC., (2004) Isaac, J.A. 17638; G5:$354 Rhododendron maximum L., Bell 580 Rhododendron periclymenoides (Michx.) shinners, (2004) Isaac, J.A. 17377 \ corymbosum L., (1907) Jennings, O.E. Vacant a lidum Aiton, (2002) Isaac, J.A.; Takacs, M. 14402 Vaccinium stamineum L., (2002) Isaac, J.A.; Takacs, EUPHORBIAC a he aca Raf.,(1992) Haywood,M.H. Acalypha virginica L. _(1993) Haywood, M.J.211 Chamaesyce maculata (L.) Small, (1955) Buker, W.E (1993 nee apace nutans (Lag.) Small, »MJ.17 sea commutata Engelm., We ae Folman, J.; Gardne neha ae : (2003) ae JA rae FABACEAE Amphicarpaea bracteata (L.) Fern., (1984 son, S.A.; Nishida, J.H.; Bier, C.W. 1945 Apios americana Medik., (1993) Haywood, M.J. 159 SS Cercis canadensis L., (2003) Isaac, J.A.; Isaac, B.L. 15821 Chamaecrista nictitans (L.) Moench, (1953) Buker, W.E *Coronilla varia L., (1995) Isaac, B.L.; Isaac, J.A. 7505; Europe p Desmodium canadense (L.) DC.,(1993) Haywood, M.J. 194 1849 Desmodium glutinosum (Muhl. Ex Willd.) Wood, (2003) Isaac, J.A. 16349 Desmodium marilandicum (L.) DC., (1966) Buker, W.E. Desmodium nudiflorum (L.) DC, (1951) Henry, LK; eer, FH. Desmodium paniculatum (L.) DC., (2003) Isaac, J.A. 7156 Desmodium perplexum Schub., (1993) Haywood, MJ. 157 Desmodium rotundifolium DC.,, (1954) Henry, L.K.; au ker WE. editsia triacanthos L., (2004) Isaac, A.A. 17508 Ginnod ie dioicus (L.) K.Koch, Donley 160 *Lathyrus sylvestris L., (1995) Isaac, B.L.; Isaac, J.A. vi otic Lespedeza hirta (L.) Hornem., (1974) Buker, W.E. Lespedeza procumbens Michx., (1921) Jennings, O.E.; Jennings, G.K,;et al. Lespedeza repens (L.) W. Bart., (1954) Henry, L.K.; Buk E er, W Lespede a violacea (L (L .) Pers., ‘a 969) Buker, W.E. L pede a virg inica (L.) B Lohr 174 wea, ritt, aa corniculatus L., (2003) Isaac, J.A. 16176; pe ee lupulina L.,(1995) Isaac, B.L;lsaac,J.A. 7495; Eurasia “Medicago sativa L., (1958) Buker, W.E,; Eurasia *Melilotus officinalis (L.) Lam., (2003) Isaac, J.A. 16113; Eurasia Robinia pseudoacacia L., (2003) Isaac, J.A. 16039 enna hebecarpa (Fern.) Irwin & Barneby, (2003) Isaac, J.17153 Senna marilandica (L.) Link, (1951) Henry, L.K.; ,F.H;G5:S1 ee aureum Pollich, (2003) Isaac, J.A. 16324; urasia al : ium ee Schreb., (2003) Isaac, J.A. urope =e ium Fyoraine LS on Thompson, S.A.; Nishida, J.H. 2351; Eu *Trifolium pratense L., (1 ae i B.L.; Isaac, J.A. 7 : ope *Trifolium repens L., (2003) Isaac, J.A.16117;Europe Vicia caroliniana Walt., (1 te He Ai M.J.446 *Vicia cracca L., (1896) Hoge, M.K.; Euras *Vicia villosa Roth, (2003) ee JA, fort pune FAGACEAE Castanea dentata (Marsh.) Borkh., (2003) Isaac, A. 16586 1850 Fagus grandifolia Ehrh., (2004) Isaac, J.A. 17633 Quercus alba L., (1985) Thompson, S.A. 2435 Quercus bicolor Willd. (1921) Dickey, S.S. Quercus coccinea Muenchh., (1954) Henry, L.K; Buker, W.E. pasieauean aan a (2003) Isaac, J.A. 16159 Engelm., (1984) Thomp- son, S.A.; Nishida, J.H.; Bier, C.W. 2002 Quercus prinus L., (1991) Isaac, J.A. 3647 Quercus rubra L., (1985) Thompson, S.A. 2432 Quercus stellata Wangenh., (1948) Bryner,C.L.501 Quercus velutina Lam., (1985) Thompson, S.A. Quercus x leana Nutt. (pro sp.), (1948) Bryner, C.L. 502 FUMARIACEAE Corydalis aurea Willd. (1996) Haywood, M.J.634 Corydalis Havula (Raf) DC., (2003) Isaac, B.L.;lsaac, 827 Dicentra canadensis (Goldie) Walp., (2003) Isaac, B.L.; Isaac, J.A. 15844 Dicentra cucullaria (L.) Bernh., (2003 Isaac, J.A. 15858 GENTIANACEAE Gentiana andrewsii Griseb. var.andrewsii, (2003) R.; Block, R.; Bradburn, M Gentiana clausa Raf. (1954) Henry, L.K; Buker, W.E. Obolaria virginica L., (1952) Henry, L.K.; Buker, W.E. Sabatia angularis (L.) Pursh, (1998) Grund, S.P. wo Isaac, B.L.; — GERANIACEAE “Geranium columbinum L., (1993) Isaac, J.A.4505; Europe Geranium maculatum L., (2003) Isaac, J.A.; lsaac, B.L. *Geranium sibiricum L., (1984) on S.A, Nishida, J.H., Bier, C. 1954: Eur “Geranium thunberel Sieb. & ae ex Lindl. & eae . is Thompson, S.A,; Nishida, J.H. GROSSULARIACEAE Ribes cynosbati L., (2003) Isaac, J.A. 16334 *Ribes rubrum L., Rhoads, A.; Eurasia HALORAGRACEAE Myriophyllum heterophyllum Michx.,(2003) Coxe, R;G5:S1 HAMAMELIDACEAE Hamamelis virginiana L., (2004) Isaac, J.A. 17635 BRIT.ORG/SIDA 21(3) HIPPOCASTANACEAE Aesculus flava Aiton, (2003) Isaac, J.A.;|saac, B.L.; Morton, C.M. 15811 Aesculus glabra Willd., (2003) Isaac, J.A. 15883 HYDRANGEACEAE Hydrangea arborescens L.,(2003) Isaac, J.A. 16366 *Philadelphus coronarius L., (1896) Hoge, M.K.; Eurasia HYDROPHYLLACEAE Hydrophyllum appendiculatum Michx., (1995) Isaac, BL; Isaac, J.A. 7183 Hydrophyllum canadense L., (1995) Isaac, B.L.; Isaac, J.A. 7164 phyllum Nutt., (2003) Isaac, JA. 15882; G5:S1 Hydrophyllum virginianum L., (1996) Isaac, B.L.; Isaac, J.A.8846 Phacelia purshii Buckl., (2003) Isaac, J.A. 16040 IRIDACEAE Iris cristata Aiton, (2004) Isaac, J.A. 17380; G5:S1 Iris pseudacorus L., (2003) Isaac, J.A. 16193 Sisyrinchium angustifolium P. Mill., (1993) Isaac, J.A.4377 JUGLANDACEAE Carya alba (L.) Nutt. ex Ell, (1921) Jennings, O.E. Carya cordiformis (Wangenh.) K. Koch, (1993) Haywood, M.J. 197 ets dea (P.Mill.) =swers (1905) Jennings, O.E. valis (Wang ) Sarg. (1954) Henry, L.K,; ‘Buren W Carya ovata . Mill.) K.Koch, (1921) Jennings, O.E. uglans cinerea L.,(2003) Isaac, J.A. 1639 Juglans nigra L., (1951) Henry, L.K.; Beer, FH. JUNCACEAE Juncus acuminatus Michx., (2003) Coxe, R.; radburn, M. Juncus effusus L.var.pylaei (1954) Henry, L.K.; Buker, Juncus effusus L.var. solutus an & Wieg,, (2003) Isaac, J.A. 1619 Juncus marginatus Rostk.,(1951) Henry, L.K,; Beer, F.H Laharpe) Fern.& Wieg., Juncus tenuis Willd. (2003) Isaac, JA. 16307 Juncus torreyi Coville, (2002) Paluh, A.T. 1 Luzula acuminata Raf., (1996) Isaac, B.L.; Isaac, J.A. — Luzula echinata (Small) FJ. Herm., Bell 422 COXE ET AL Luzula multiflora (Ehrh.) Isaac, J.A. 8887 MIACEAE Agastache nepetoides (L.) Kuntze, (1975 a W.E. Blephilia ae on Benth., (1995) Isaac, B.L.; Isaac, .A. 752 ‘npn aoe L., (2003) Isaac, J.A. 16370; rope a, insonia ae Lee oe S.A.; Nishida, J.H.; Bier, C.W. 1 Cunila ae a (L.) ane eas Henry, L.K,; Beer, ee one L., (1996) Isaac, J.A,; Isaac, B.L. 8840; Eur Hedeoma pu ae (L.) Pers., (1994) Thompson, S.A. Rawlins, J.E.11731 Isanthus brachiatus (L.) B.S.P., 1921) Jennings, O.E. *Lamium amplexicaule L., (1996) Haywood, M.J 631; Eurasia *Lamium maculatum L,,(1 947) Henry, LK; Eurasia *lamium purpureum L., (2003) Isaac, J.A.; Isaac, n, C.M. 15802; Eurasia *Leonurus cardiaca L.,(1995) Isaac, B.L.; Isaac, J.A. ;Asia Lycopus americanus Muhl. ex W. Bart. (1984) Th- mpson, S.A.; Nishida, J.H. 2026 Lycopus uniflorus Michx. var. uniflorus, (1993) Haywood, MJ. 154 Lycopus virginicus L., (2003) Isaac, J.A. 16572 Meehania cordata (Nutt.) Britt. Bell 619;G5:S1 Mentha arvensis L., (1950) Henry, L.K.; Buker, W.E. *Mentha x piperita L.(pro sp.), (1951) Henry, LK; Beer, F.H.; Eurasia *Mentha spicata L., (1951) Henry, L.K.; Beer, FH; Europe Monarda clinopodia L., (2003) Isaac, J.A. 16388 Monarda fistulosa L., (1995) Isaac, B.L.; Isaac, J.A. 7538 Monarda media Willd., (1907) Jennings, O.E. *Nepeta cataria L., (1954) Henry, L.K,; Buker, W.E,; Europe Prunella vulgaris L.ssp. lanceolata (W. Bart.) Hulten, (1995) Isaac, B.L.; Isaac, JA. 7529 Pycnanthemum incanum (L.) Michx., (195 Henry, L.K; Buker, W.E. Pycnanthemum tenuifolium Schrad., (1951) Henry, L.K; Buker, W.E. Salvia lyrata L., (2003) Isaac, J.A. 16048 Scutellaria incana Biehler, (2003) ie 16579 paar, — Lej., (1996) Isaac, B.L.; Buker, 1851 — Scutellaria lateriflora L., (1984) Thompson, S.A.; ishida, J.H. 2077 Scutellaria nervosa Pursh, (2003) Isaac, J.A.16332 Scutellaria saxatilis Riddell, (2003) Isaac, J.; Haywood, M.J.;Coxe, R.;Brandburn, M. 17230; G3:S1 cee is Hii Schuttlw.ex. oun (2003) Isaac, ‘Bradburn, M Stachys teuole Willd., (2003) 7 eneme R.; Bradburn Teuauin cane ae Haywood,M.J.127 mL.,(1921) Dickey, S.S. LAURACEAE Lindera benzoin (L.) Blume, (2003) Isaac, J.A.;lsaac, orton, C.M. 15799 Sasso albidum ea Nees, (1995) Isaac, B.L.; ac, J.A. 7095 LEMNACEAE Lemna minor L., (2003) Isaac, J.A. 16402 LILIACEAE Allium canadense L.,(2003) Isaac, J.A. 16132 Allium cernuum Roth, (2003) Isaac, J.A. 16583 Allium tricoccum Aiton, (1996) Haywood,M.J.41] “Allium vineale L.,(2003) Isaac, J.A.16139; Europe Clintonia umbellulata (Michx.) Morong, Bell 377 *Convallaria majalis L., (1896) Hoge, M.K.; Europe Erythronium albidum Nutt., (1993) Haywood, MJ. $3 Erythronium americanum Ker-Gawl.,(2003) Isaac, A, Isaac, B.L. 15832 ee is fulva (L.) L., (2003) Isaac, J. 16317; er yeseaaint (1941) Henry, L.K.; Exotic Hypoxis hirsuta (L ea Sebben Lilium superbum L., (2003) Coxe, R.; Bradburn, M Maianthemum racemosa (L,) Link, (2003) J.A.lsaac 16023 Medeola virginiana L., (1993) Isaac, J.A.4367 eae wane s L., (1896) Hoge, M.K.; Europe alum umbellatum L., (1995) Isaac, B.L.; ee JA. 7182: Europe Polygonatum biflorum (Walt.) Ell. var. com- mutatum (J.A.& J.H. Schultes) Morong {(1975) Buker, W.E Pane! pubescens (Willd.) Pursh, (1996) ac, B.L c,J.A.8899 anes: pies, (Michx.) D.Don, (1970) Bu .E.; [Disporum lanuginosa (Michx. ae ae 1852 Trillium erectum L., 15854 Trillium ‘dats Gen A. |sa ‘Mort zy J. sas ee Aceh ea sa : P; Gray, A,; Folman, J.; Gardner, M.1818;G4:53 Trillium sessile L., (2003) Isaac, J.A.;[saac, B.L.; Mor- .M. 15808 Wuedaria grandiflora Sm., (2003) Isaac, J.A.; |saac, B.L. 15841 Uvularia perfoliata L., (1993) Haywood, M.J. 79 LIMNANTHAC EAE Floerkea proserpinacoides Willd., (2003) Isaac, J.A.; Isaac, B.L. 15830 LINACEAE Linum virginianum L.,(1951) Henry, L.K.; Beer, FH. LYTHRACEAE -uphec 1 VISCOSISSIMA Jacq,, Q 1991) Isaac, J.A. *Lythrum salicaria L., (1993) Haywood, MJ. Europe MAGNOLIACEAE ma reees tulipifera L., (1996) Isaac, B.L.; Isaac, 8868 ears iaacuminata (L) Lb MALVACEAE “Abutilon theophrastii Medik., (1951) Henry, L.K.; Beer, F.H.; Asia *Hibiscus trionum L., (1984) Thompson, S.A. Nishida, J.H. 2044; Europe *Malva moschata L., Lohr; Europe *Malva neglecta Wallr., Lohr 260; Eurasia/ N. Africa “Sida spinosa L., (1951)Henry, L.K.; Beer, F.H.; Tropics MELASTOMACEAE Rhexia virginica L., (2003) Isaac, J.A. 17150 MENISPERMACEAE Menispermum canadense L., (2003) Isaac, J.A. 39 . (1904) Jennings, O.E. MOLLUGINACEAE *Mollugo verticillata L.,(1951 ropic America MONOTROPACEAE MOnIETOPE hypopithys L., (2004) Isaac, J.A.17653 tropa uniflora L., (2003) Isaac, J.A. 16414 Henry, L.K.;Beer, FH.; (2003) Isaac, B.L.; Isaac, JA. nee ot Isaac, BRIT.ORG/SIDA 21(3) MORACEAE Maclura pomifera (Raf.) Schneid., (2003 oe Isaac, JA. *Morus alba L.,(1951) Henry, L.K.; Beer, F.H.; Asia Morus rubra L., (1996) Isaac, B.L.; Isaac, J.A. 8864 OLEACEAE Chionantheus virginicus L., (1896) Hoge, M.K. Fraxinus americana L., (2003) Isaac, J.A. 16403 Fraxinus nigra Marsh., Bell *“Syringa vulgaris L., (1896) Hoge, M.K.; Europe ONAGRACEA Circaea lutet pes ssp. canadensis (L alae Magnus, (1996) Haywood, M.J.6 pilobium co ae see (1984) ee S.A,; Nishida, J.H. 2 pate BIER L. (l i ees MJ. . (2003) Isaac, a 49 .) Ell, (2003) aie R.; — ae pie 7 Bradburn, M enothera biennis L., pee eines: 16605 Oenothera futicosa - (1896) Ho Oenothera flora L., Donley, — Oenothera perennis ? (1994) Haywood, M.J.468 ORCHIDACEAE Aplectrum hyemale (Muhl. ex Willd.) Torr, (1997) Shriver, J.S.; Shrive Hace C. 868; G5:S3 n, (1922) Dickey, S.S. as pubescens aie R. Br. ex Aiton f,, 97) Shriver, J.S.; Shriver, A; Smith, C.867 ee medeoloides (Pursh) Raf., (1 ae Bright, J. Isotria verticillata Raf., (1922) Dicke Liparis liliifolia (L.) L.C. Rich. ex Ker- La (1995) 9 C\ Nrinediijma qule A SZ Isaac, B.L.; Isaac, 15 Platanthera lacera ict ) G. Don, (2003) Isaac, xe, R. 1629 Platanthera oe (Pursh) Lindl. (1953) Spiranthes rT | .) LC. Rich., Lohr, WN Spiranthes lacera (Raf.) Raf. var. gracilis {Bigelow Luer, (2004) Isaac, J.A.; Smith, C. 18 Spiranthes ochroleuca (Rydb.) Rydt oe ales — W.E. Spiranthes ovalis Lindl., (2003) Isaac, J.; Coxe, R.; Bradburn, M. 17225 OROBANCHACEAE Conopholis americana (L.) Wallr. f., (2003) Isaac, 6165 Epitagus virginiana (L.) W.Bart., (1969) Buker, W.E. COXE ET Al Orobanche uniflora L., (1996) |saac, B.L.;Isaac, J.A. 8 871 = ALIDACEAE xalis dillenii Jacq., (2003) Isaac, J.A. 16376 xalis grandis Small, (2003) Isaac, J.A. 16326 ser stricta L., (1993) Haywood, M.J. 1 Oxalis violacea L., (1996) Haywood, M.J.629 PAPAVERACEAE *Chelidonium majus L.,(1994) Haywood, M.J.426; urope Sanguinaria canadensis L., (2003) Isaac, J.A.;|saac, B.L. 15855 PASSIFLORACEAE Passiflora lutea L., (2003) Isaac, J.; Coxe, R.; radburn, M.17221;G5:S1 PHY TOLACCACEAE Phytolacca americana L., (1993) Haywood, M.J. PLANTAGINACEAE Plantago aristata Michx., (1954) Henry,L.K,; Buker, W.E. *Plantago lanceolata L., (1996) Isaac, B.L.; Isaac, J.A.8900; Europe *Plantago major L., (1984) Thompson, S.A.; Nishida, J.H. 2050; Europe flee bain Dene., (2003) Isaac, J.A. 16399 1tago virginica L., (1975) Buker, W.E. PLATANACEAE Platanus occidentalis L.,(1995) Isaac, B.L; Isaac, J.A. 7491 POACEAE “Agrostis gigantea Roth, (2003) Isaac, J.A.; Coxe, Agrostis fete a (Walt.) Tuckerman, (1994 n, S.A, ieee 11969 Boe nee Willd., Pohl 592 Andropogon virginicus L. es Henry, L.K.; Beer, eae FH. ees odoratum L., (2003) Isaac, J.A. 989 asia Aristida fa eee Michx. var. dichotoma, Bell 334 *Arrhenatherum elatius (L.) Beauv.ex J.& C. Presl, (1995) Isaac, B.L.; Isaac, J.A. 7516; Europe *Avena fatua L., (1971) Buker, W.E.; Europe Bromus ciliatus L., (2003) Coxe, R. 1853 “Bromus commutatus Schrad., (2003) Isaac, J.A. 16175; Europe “Bromus inermis Leyss.,(1995) Isaac, B.L.;lsaac,J.A. 7489; Europe “Bromus japonicus Thunb. ex Murr. (1995) Isaac, B.L.; Isaac, J.A. 7488; Eurasia Bromus latiglumis (Shear) A.S. Hitchc., Fogg 18323 Bromus pubescens Muhl. ex Willd. (2003) Isaac, JA. 16416 “Bromus racemosus L., (1 a Thompson, S.A.; Nishida, J.H. 2419; Euro “Bromus sterilis L., (1996) = B.L.; Isaac, J.A.8844; Europe a um latifolium ae Yates, (2003) Coxe radburn, M Cinna onda are 7 aenieen S.A. Nishida, J.H.; Bier, C.W. 1987 *Dactylis glomerata L., (2003) Isaac, J.A.;|saac, B.L.; Isaac, H.R. 16096; Europe Danthonia compressa Austin ex Peck, (2003) Isaac, J.A.16151 Danthonia spicata (L.) Beauv. ex Roemer & J.A. Schultes, (2003) Isaac, J.A. 16352 Diarrhena americana Beauv., (2002) Isaac, B.L.; Isaac, J.A. 15787; G4?:S1 Dichanthelium acuminatum i Gould & CA. Clark, (2004) Isaac, J.A.17 Dichanthelium boscii (Poir.) tae & CA. Clark, (1995) Isaac, B.L.; Isaac, J.A. 7162 Dichanthelium clandestinum (L.) Gould, (1995) Isaac, B.L.; Isaac, JA. 7535 Dichanthelium commutatum (J.A. Schultes) Gould, (2003) Isaac, J.A. 16409 Dichanthelium dichotomum (L.) Gould, (1904) Jennin Di hanthen ian Paonamie (L.) Gould & CA. Clark, (2003) Isaac, J.A. 16346 Dichanthelium linearifolium (Scribn. ex. Nash) Gould, (2004) Isaac, J.A. 176 Dichanthelium sabulorum (Lam.) Gould & C.A. Clark var. thinium (A.S. Hitchc.& Chase) Gould &C.A.Clark, (2002) Isaac, J.A.; Takacs, M. 14416 *Digitaria ischaemum (Schreb.) Schreb.ex Muh, (1984) Thompson, S.A,,Nishida,J.H.2075;Eurasia “Echinochloa crus-galli (L.) Beauv., (1993) Haywood, M.J. 165; Eurasia *Eleusine indica (L.) Gaertn., (2003) Isaac, J.A. 16616; Old World Tropics ae — 1854 Elymus canadensis L., (1951) Henry, L.K.; Buker, W.E Elymus hystrix L., (2003) Isaac, J.A. 16299 “Elymus repens (L.) Gould, (2003) Isaac, J.A. 16119; Exotic Elymus riparius Wieg., (2003) Isaac, J.A. 16172 Elymus villosus Muhl. ex Willd. (2003) Isaac, J.A. 16135 Elymus virginicus L., (1951) Henry, L.K.; Buker,W.E. Eragrostis spectabilis (Pursh) Steud.,(1941) Davis, A. etal. Festuca subverticillata (Pers.) Alexeev, (2003) Isaac, J.A,;lsaac, B.L.; Isaac, H.R. 16103 Glyceria ed AS. Hitche., Bell 271 Glyceria striata (Lam.) A.S.Hitchc., (2003) Isaac, J.A. ae *Holcus lanatus L.,(2003) Isaac, J.A.16172;Europe Leersia oryzoides (L.) Sw.,(1951) Henry, L.K.; Beer, FH Leersia virginica Willd., (1984) Thompson, S.A.; Nishida, J.H.; Bier, C.\W. 1999 “Lolium perenne L., (1995) Isaac, B.L.; Isaac, JA. 498; Europe — *Lolium wound (Huds.) S.J. Darbyshire, (2003) c,J.A.; Isaac, B.L.; Isaac, H.R. 16097; Europe Muhlenbergia ie (Poir.) Fern, Bell 3645 ay lea Ree at F, So (1984) Thomp- Nishida, bn is a oon fou ics (1984) Thomp- A.; Nishida, J.H ee philadetphicum rae ex Trin., (1985) h on, S.A.; Nishida, J.H. 2516 ries iene (L.) R.Br. (1993) Haywood, 4. 164; ous Phalaris ar a L., (2003) Isaac, J.A. 16309 *Phleum pratense L.,(1954) Henry, L.K.; Buker, W.E.; Poa alsodes A. Gray, (1951) Henry, L.K.; Beer, FH. *Poa annua L., (1996) Thompson, S.A.; Rawlins, rasia “Poa compressa L.,(1993) Isaac, J.A.4375; Europe Poa cuspidata Nutt., (2003) Isaac, J.A.; Isaac, B.L.; Morton, C.M. 1 “Poa pratensis L.,(1 984) Nishida, J.H.;Bier, C.W. 709; Europe sylvestris Gray, (1951) Henry, L.K; Beer, FH. Poa trivialis L., (2003) Isaac, J.A.16198; Europe Schizachyrium scoparium (Michx.) Nash, (1904) Jennings, O.E. — — BRIT.ORG/SIDA 21(3) Schizachyrium scoparium ues, Nash var. scoparium, (1904) Jennings, O *Setaria faberi Herrm., (1951) Dalene F.H.; Asia *Setaria viridis (L 7 : Beauv. var. viridis, (2003) Isaac, Shenoenol: meme (Rydb.) Rydb., (1996) T son, S.A, Rawlins, J.E. 12459 Sphenopholis witida eae Scribn,, (2003) Isaac, JA. 16160 Sporobolus vaginiflorus (Torr. ex Gray) Wood, (2003) Isaac, J.; Coxe, R.; Bradburn, M. 17227 Tridens flavus (L.) A.S. Hitchc., (2004) Isaac, J.a. 18557 POLEMONIACEAE Phlox divaricata L., (1999) Haywood, M.J.607 Phlox maculata L. ssp. maculata, Wherry Phlox Sa ‘a L., (2003) Isaac, J.A. 16607 Phlox stolonifera Sims, (1952) Henry, L.K.; Buker, W.E. Isaac, J.A. 16170 peel Polemonium reptans L., (2003 POLYGALACEAE Polygala sanguinea L., (2003) Isaac, J.A. 16639 Polygala verticillata L. var. verticillata,(1954) Henry, POLYGONACEAE *Polygont viculare L., (1994) Thompson, S.A.; Rawlins, J.E. 11730; ate *Polygonum caespitosum Blume, (2003) Isaac, .A. 16189; Asia ba a convolvulus L., (1954) Henry, L.K; Pune W.E.; Europe tum L., (2003) Isaac, J.A. 16571 eee dae ct (1984) Thompson,5.A.; Nishida, J.H.; Bier, CW. 1986; Europe Polygonum Ree ee Michx., (2003) Isaac, JA. 16589 Polygonum pensylvanicum L., (1984) Thompson, S.A.; Nishida, J.H. 2054 Polygonum persicaria L.,(1984) Thompson, S.A.; Nishida, J.H. 2055; Europe Polygonum punctatum EIl. var. confertiflorum (Meisn.) Fassett, (2003) Isaac, J.A. 16590 Polygonum sagittatum L., (1984) Thompson,S.A.; Nishida, J.H.; Bier, C.W. 1975 Polygonum. scandens L. var. scandens, (2004) Isaac, 8515 Polygonum virginianum L.,(1998) Grund, S.P.2048 COXE ET Al NALCWE LUVIN I, PENNSYLVANIA *Rumex acetosella L., (1996) Isaac, B.L.; Isaac, J.A. 8874; Europe *Rumex crispus L.,(2003) Isaac, J.A.16110;Europe *Rumex obtusifolius L., (2003) Isaac, J.A. 16302; Europe PORTULACACEAE — caroliniana Michx., (2003) Isaac, J.A.; Isaa 1585 eae virginica L., (2003) Isaac, J.A.; !saac, B.L.; Morton, C.M. 15797 Portulaca oleracea L., (2003) Isaac, J.A. 16624 PRIMULACEAE Lysimachia ciliata L., = (1995) Isaac, B.L,; Isaac, J.A. Lysimachia lanceolata Walt.,(1993) Haywood, M.J. 69 *Lysimachia nummularia L., (1995) Isaac, B.L.; Isaac, J.A. 7508; Europe Lysimachia quadrifolia L., (1995) Isaac, B.L; Isaac, JA. 793 | PYROLACEAE Chimaphila maculata (L.) Pursh, (2004) Isaac, J.A. 17654 Pyrola americana Sweet, (1968) Buker, W.E. ees AE nitum uncinatum L., (1932) Bright, J. 7641; oe Actaea pachypoda EIL, (1969) Buker, W.E. Actaea racemosa L. var. racemosa, (2003) Isaac, J.A. 16575 Anemone quinquefol ia L., Lohr Anemone virginiana L., (2003) Isaac, J.A.16121] 2003) Isaac, J.A. 15900 Caltha, palustris L. var. palustris, Bell 423 Clematis occidentalis (Hornem.) DC.,(1896) Hoge, M.K Aq Peyalial anadensis L., ( Clematis virginiana L., (2003) Isaac, J.17138 Delphinium tricorne Michx., (1996) Isaac, B.L.; Isaac, J.A. 8850 *Helleborus viridis L.,(1940) Baker; Exotic Hepatica nobilis Schreber var. acuta (Pursh Steyermark, (2003) Isaac, B.L,;lsaac, JA; Mor- ton C.M. 15814 Hepatica nobilis Schreber var. obtusa (Pursh Steyermark, (1952) Henry, L.K.; Buker, W.E. . (2003) Isaac, J.A. 16335 eanwacul us abortivus L., (2003) Isaac, J.A,; Isaac, B.L.; lsaac, H.R. 16098 = lydra ft] qAnadensic 1855 “Ranunculus acris L., (1995) Haywood, MJ. 561; Europe Ranunculus all lensis Britt., (1996) Isaac, B.L.; Isaac, J.A. 8909 *Ranunculus ficaria L. ene Isaac, J.A.\saac,B.L.; orton, C.M. 15812; Ranunculus hispidus ne var. hispida, (2003) saac, J.A,; Isaac, B.L. 15835 Ranunculus hispidus Michx. var. nitidus (Chap- man) T. Duncan, (1994) Haywood, M.J. 25 Ranunculus micranthus Nutt., (2003) Isaac, J.A.; Isaac, 8 perene nti recurvatus Poir., (1996) Isaac, J.A.; D (Panlineulus repens L.,(1995) Isaac, J.A;lsaac, B.L. 7178; Europe Thalictrum dioicum L., (2003) Isaac, J.A.;|saac, B.L. 1584 Thalictrum pubescens Pursh, (2003) Isaac, J.A. 16356 Thalictrum thalictroides (L.) Eames & Boivin, (2003) Isaac, J.A.; Isaac, B.L.; Morton, C.M. 15815 RHAMNACEAE Ceanothus americanus L.,(2003) Isaac, J.A. 16378 ROSACEAE Agrimonia gryposepala Wallr., (1993) Haywood, MJ.95 Agrimonia parviflora Aiton, (1951) Henry, L.K;; er, F. PA. Agrimonia pubescens Wallr., (1998) Grund, SP. 205 > tellata Wallr., (2003) Coxe, R.; Isaac, * haying MJ. a (Mi eae (2003) Isaac, Amelanchier stolonifera Wieg,., : 949) Henry, LK. Aruncus dioicus (Walt.) Fern., (2003) Isaac, J.A. 16178 Crataegus crus-gallii L., (2003) Coxe, R.;Isaac, J.A.; Ernst Crataegus punctata Jacq.,(2003) Isaac,J.A. 16020 Fragaria virginiana Duchesne, (1994) Haywood, Geum aleppicum Jacq., (1993) Haywood, MJ. 68 Geum canadense Jacq. var. canadense, (1995) Isaac, J.A.; Isaac, B.L.751 Geum vernum (Raf.) Torr. & Gray, (1996) Isaac, J.A.; Isaac, B.L.8856 1856 Geum virginianum L., (1950) Henry, L.K.; Buker, W.E Malus coronari (L.) P.Mill., (2003) Isaac, J.A. 16417 *Malus pumila P. Mill, (2003) Isaac, J.A,; Isaac, B.L. 15822; Eurasia Physocarpus opulifolius (L.) Maxim., Haywood, M.J.477 Ih densis L.,(1999) Haywood,M.J.05 eae norvegica L. ssp. monspeliensis (L.) Aschers.& Graebn., (1995) Isaac,B.L.;lsaac, J.A. 7501 *Potentilla recta L., (1995) Isaac, B.L.; Isaac, J.A. (1994) 533; Europe Potentilla simplex Michx., (1996) Isaac, B.L.; Isaac, A. 8896 Prunus americana Marsh.,(1951) Henry, L.K.; Beer. FH. “Prunus avium (L.) L., (2003) Coxe, R.; Bradburn, M.; Eurasia Prunus serotina Ehrh., (2003) Isaac, J.A. 16027 Prunus virginiana L., (1975) Buker Lee scandens (Thunb.) Meaning: (2003) c,J.A. 15977; Japan a carolina L. var. carolina, (2003) Isaac, J.A.: xe, R. 16281 *Rosa eglanteria L., (1992) Haywood, M.J. 377; Europe *Rosa multiflora Thunb.ex Murr. (1995) Isaac, B.L.; Isa a 7539; a Sia h.,(1951) Henry, L.K; Beer, FH. rice al so ensis Porter, (1997) Grund, S.P. 18 Rubus alumnus Bailey, (1950) Davis, H.A.; Davis, T. 9145 ose i Viale Bailey, (1950) Davis, H.A.; Davis, ae an is Willd, (2003) Isaac, J.A. 16046 Rubus laudatus Berger, (1951) Davis, H.A.; Davis, 7.9495 Rubus multifer Bailey, (1951) Davis, H.A.; Davis, T. 9479 Rubus esi L., (2003) Isaac, J.A. 16030 Rubus odoratus L., (1958) Buker, W. ian coer Poir, (1951) Davis, H.A.; Davis, T.9496 Rubus tes Blanch.,(1951) Davis, H.A.; Davis, T.948 — ror roa (Bailey) Rydb., (1949) Davis, H.A,; 8882 BRIT.ORG/SIDA 21(3) Spiraea alba Du Roi var. latifolia (Aiton) Dippel, (1993) Haywood, MJ. 102 *Spiraea japonica L. f., (1998) Grund, S.P. 2056; Ja pan Waldsteinia fragarioides (Michx.) Tratt., (1895) Guttenberg, G. RUBIACEAE Diodia teres Walt., (1951) Henry, L.K.; Beer, FH. Galium aparine L., (1996) Isaac,B.L.; Isaac, J.A.8872 Galium aes lum Michx.,(1984) Thompson, S.A.; Nishida, J.H. 2031 Galium circaezans Michx. var. circaezans, (2003) Isaac, J.A.; Isaac, B.L.; Isaac, H.R. 16102 Galium circaezans Michx. var. hypomalacum rn., (1950) Henry, L.K.; Buker, WE. os jum concinnum Torr.& Gray, (1 966) Buker, W.E. eolatum Torr. (1921) Jennings, O.E. *Galium mollugo L., (2003) Isaac, J.A.; Coxe, R. 16284; Eurasia Galium pilosum Aiton, (1951) Henry, L.K.; Beer, FH. Galium tinctorium (L.) Scop., Donley 151 Galium triflorum ee (2003) Isaac, J.A. 16012 *Galium verum L., (1921) Dickey, S.S.; Eurasia Houstonia caerulea L., (1996) Isaac, B.L.;!saac, J.A. alitsnm lan Houstonia ene Willd. ex Roemer & J.A. Schultes, (1921) Dickey, S.S. Mitchella ae L., (1970) Buker, W.E,; Henry, LK. SALICACEAE Populus grandidentata Michx., (1974) Buker, W.E. Salix caroliniana Michx., (1951) Henry, L.K.; Beer, FH; G5:$1 Salix discolo i , (1985) Thompson, S.A,; Nishida, J Salix eset cane (2003) Isaac, J.A.; Isaac, Bils /C.M.1 Salix exigua Nutt., (1995) Isaac, B.L.;lsaac, J.A.7122 le nigra Marsh., (1952) Henry, L.K.; Buker, W.E. alix sericea Marsh., (1955) Henry, L.K.; Buker, WE. SAXIFRAGACEAE Chrysosplenium (2003) Isaac, J.A. 15903 Heuchera americana L.,(1995) Isaac, B.L.jlsaac, JA. Schwein.ex Hook.,, 7118 Mitella diphylla L., (2003) Isaac, J.A.; Isaac, B.L. 15859 Saxifraga virginiensis Michx., i Isaac, J.A.; Isaac, B.L.; Morton, C.M. 15816 COXE ET AL., VASCULAR FLORA OF GREENE COUNTY, PENNSYLVANIA Tiarella cordifolia L., (1996) Isaac, B.L.; Isaac, JA. 8895 SCROPHULARIACEAE Agalinis tenuifolia (Vahl) Raf., (1969) Buker, W.E. “Antirrhinum majus L.,(1896) Hoge, M.K.; Europe Aureolaria flava (L.) Farw. var. flava, (1975) Buker, Aureolaria laevigata (Raf.) Raf., (1993) Haywood, 97 Abiesiae ia virginica (L.) Pennell, (1969) Buker,W.E. *Chaenorhinum minus (L.) Lange, (2003) Isaac, J.A. 16138; Europe Co poe verna Nutt., (2003) Isaac, J.A.; |saac, B.L. 5850 are neglecta Torr., (2003) Isaac, J.A. 16397 *Linaria vulgaris P.Mill.,(1974) Buker,W.E.,; Eurasia Lindernia dubia (L.) Pennell var. dubia, (1974) Mimulus alatus Aiton, (2003) Isaac, J.A. 16587 Mimulus ringens L., (1 951 ) Henry, L.K; Beer, FH. Pediculari d L., (1995) Isaac, B.L.; Isaac, J.A.7166 Penstemon digitalis Nutt. ex Sims, (2003) Isaac, Penstemon laevigatus Aiton, (2002) Isaac, J.A.; Takacs, M. 14399; G5:$3 ae marilandica L., (1984) Thompson, hida, J.H.; Bier, C.W. 1973 Pec blattaria L., (1995) Isaac, B.L.; Isaac, A. 7504; Eurasia *Verbascum thapsus L.,(1995) Isaac, B.L.;lsaac,J.A. 7540; Eurasia Veronica americana Schwein. ex Benth., (2002 c, J.A.; Takacs, M. 14403 Veronica anagallis-aquatica L., (1996 M.J.627 = wa Haywood, *Veronica arvensis L., (2003) Isaac, J.A. 15867; Eurasia *Veronica hederifolia L., (2003) Isaac, J.A. 15880; Eurasi a *Veronica officinalis L., (1996) Isaac, B.L.; Isaac, J.A. 8881; Europe *Veronica persic Poir., (1920) Dickey, S.S.; Eurasia ce ica serpyllifolia L., (1995) Isaac, B.L,; Isaac, 7177;Europe SMILACACEAE Smilax glauca Walt., (1951) Henry, L.K.; Beer, F.H. Smilax herbacea L., (1993) Haywood, M.J. 327 1857 Smilax rotundifolia L., (1995) Isaac, B.L.; Isaac, JA. 7105 Smilax tamnoides L., (1970) Buker, W.E. SOLANACEAE “Datura stramonium L., (1994) Isaac, J.A.6015 "Nicanata A ae .) Gaertn., | Jen- ennings, G.K,; et al.;P ea ae Nutt., (1951) i ae Beer, FH Ww Physalis pubescens L. var. pubescens, (1921) Jen- nings qanrim carnlinen L , (1993) Haywood, ee 60 *Solanum nigrum L., (2004) Isaac, J.A. 56; Europe STAPHYLEACEAE Staphylea trifolia L., (2003) Isaac, J.A,; Isaac, B.L. 15834 THYMELAEACEAE Dirca palustris L., (2002) Isaac, J.A.; Haibach, M. 14412 TILIACEAE Tilia americana L.var. americana, (2003) Isaac, J.A. elle Tilia AM ANA phylla (Vent.) Loud., (1951) aaa oe Beer, F.H. TYPHACEAE ee ee L., (2003) Isaac, J.A.; Coxe, R. ae = ae 7 Tenis 97 ULMACEAE Celtis occidentalis L., (1921) Jennings, O.E. Ulmus americana L., (1985) Thompson, S.A,; Nishida, J.H. 2293 Ulmus rubra Muhl., (2003) Isaac, J.A. 16026 URTICACEAE Boehmeria cylindrica (L.) Sw., (2003) Isaac, J.A. 16362 Laportea canadensis (L.) Weddell, (2003) Isaac, J.A. 16319 Parietaria pensylvanica Muhl. ex Willd., (2003) Isaac, J.A. 16157 Pilea pumila (L.) Gray, (1993) Haywood, M.J. 142 Urtica dioica L.ssp. gracilis (Aiton) Seland., (2003) Isaac, J.A. 16320 VALERIANACEAE Valeriana pauciflora Michx., (1992) Isaac,J.A.3910 1858 BRIT.ORG/SIDA 21(3) Valerianella chenopodiifolia (Pursh) DC., (1996) Ex Torr. & Gray, (2003) Isaac, J.A.; Isaac, B.L. Isaac, B.L.; Isaac, J.A.8855 15846 Valerianella radiata (L.) Dufr,(1999)Haywood,MJ. — Viola sagittata Aiton var. sagittata, (1993) 606 Haywood 540 VERBENACEAE Viola sororia Willd. (2003) Isaac, J.A.; Isaac, B.L.; Phryma leptostachya L., (2003) Isaac, J.A. 16408 Verbena hastata L., (1985) Thompson, S.A.; wlins, J.E. 12882 Verbena urticifolia L. var. urticifolia Perry & Fern., (1993) Haywood, MJ. 169 VIOLACEAE Hybanthus concolor (7.F. Forst.) Spreng., (2003) Isaac, B.L.; Isaac, J.A. 15839 Viola bicolor Pursh, (1950) Henry, L.K.; Buker, W.E Viola blanda Willd. (1951) Henry, L.K.; Beer, FH. aes ; ; ; Vitis cinerea (Engelm.) Millard var. baileyana Viola canadensis L. var. canadensis, ( Haywood 422 Viola striata Aiton, (1996) Isaac, B.L.;lsaac,J.A.8901 Viola x eclipes H.E. Ballard, (1976) Buker, W.E. Viola x palmata L.(pro.sp.), (1994) Haywood, MJ. 4] E Parthenocissus quinquefolia (L.) Planch., (1951) Henry, L.K,; Beer, F.H. Vitis aestivalis Michx., (1995) Isaac, B.L.; Isaac, J.A. 7104 1994) (Munson) Comeaux, (1995) Isaac, J.A. 6017; Viola cucullata Aiton, (1996) Haywood, MJ.432 4G5?:SH Viola hirsutula Brainerd, (2003) Isaac.J.A;Isaac,B.L, “it’s fabrusca L.,(1994) Isaac, J.A. 6024 15824 Vitis riparia Michx., (1994) Isaac, J.A.6026 Vitis vulpina L., (2003) Isaac, J.; Coxe, R.; Bradburn M. 17222 — Viola labradorica Schrank, (1976) Buker, W.E. Viola pubescens Aiton var. scabriuscula Schwein. ACKNOWLEDGMENTS Our thanks go to the Wild Resource Conservation Fund of the Pennsylvania Department of Conservation and Natural Resources for partial funding of the field work. We would also like to thank Dr. Ann Rhoads, an anonymous reviewer and the editor for their helpful comments. We are also thankful to Melanie Van Olffen and Matthew Oborski for data entry. REFERENCES Brummitt, R.K.and C.E. Powett. 1992. Authors of plant names. Royal Botanic Gardens, Kew. Fike, J. 1999. Terrestrial & palustrine plant communities of Pennsylvania. Bureau of For- estry, PA.and Department of Conservation and Natural Resources, Harrisburgh, PA. Kartesz, J.T. 1999. A synonomized checklist and atlas with biological attributes for the vascular flora of the United States, Canada and Greenland. 1st. ed. In: Kartesz, J.T. and C.A. Meacham. Synthesis of the North American flora, Version 1.0. North Carolina Bo- tanical Garden, Chapel Hill. RHoabs, A.F.and W.M. Ktein, Jk. 1993. The vascular flora of Pennsylvania: annotated checklist and atlas. American Philosophical Society, Philadelphia, Pennsylvania. RHoaos, A.F. and T.A. Bock 2000. The Plants of Pennsylvania. University of Pennsylvania Press, Philadelphia, Pennsylvannia. SMITH, G.W. 1996. History of Greene County, Pennsylvania. Cornerstone Genealogical Soci- ety, Waynesburg, PA. COXE ET Al GREENE COUNTY, PENNSYLVANIA 1859 THe Nature Conservancy (1996 version). Element ranking list—Pennsylvania Natural Heri- tage Program htt WWW dc nrstate Da US forestry/pl \di/rat if Atm USDA. 1983. Soil survey of Greene and Washington counties Pennsylvania. USDA Soil Conservation Service.US Government Printing Office, Washington, D.C ~ 1860 BRIT.ORG/SIDA 21(3) BOOK NOTICE Doucias J. Furuyma, H. BRADLEY SHAFFER, and DANIEL SiMBERLOFF (eds). 2004. Annual Review of Ecology, Evolution, and Systematics: Volume 35, 2004. (ISBN 0-8243- 1435-2, hbk; ISSN 1543-592X). Annual Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto,CA 94303-0139, US.A. (Orders: www.AnnualReviewsorg, 800-523-8635, 650-493-4400, 650-424-0910 fax). $168.00 (USA), $173.00 (nel), (22.0, Oe o. Volume 35, 2004 of Annual Review of Ecology, Evolution, and Systematics, has 24 articles covering an array of topics. Vertebrate Dispersal of Seed Plants Through Time. B.H. TiFFNEY Are Diseases increasing in the Ocean? K.D. Larrerty, J.W. Porter, and S.E. rorp Bird Song: the interlace of Evolution and Mechanism. J. Popos, $.K. Huser, and B. Tart Application of Ecological indicators. GJ. Niemi and M.E. McDonald E pecan Impacts of Deer Overabundance. $.D. Corr, TP. Rooney, J.-P. Trempiay, C. Dussautt, and D.M. LER Fogel ‘fects of Transgenic Crops and the Escape of Transgenes into Wild Populations. D. PiLson nd H.R. Prenpev eerie iesantie Community Structure: the Enemy of my Enemy Nb Friend. M.E. Hay, J.D. >ARKER, D.E. BuRKEPILE, C.C. CAupiit, A-F. WiLson, Z.P. HALLINAN, and A.D. CHEQUER Operational Criteria for Saimiane Species. J.W. Sites, Jk. and J.C. Marsiatt ALANYCH The New View of Animal Phylogeny. K.M. | Landscapes and Riverscapes: the influence of Land Use on Stream Ecosystems. J.D. ALLAN Long-Term Stasis in Ecological Assemblages: evidence from the Fossil Record. W.A. DiMicuete, A.K. BEHRENSMEYER, TD. OLSZEWskI, C.C. LABANDEIRA, J.M. PANDOLFI, S.L. WING, and R. Bose Avian Extinctions from Tropical and Subtropical forests. N.S. Sonu, L.H. Liow, and FA. Bazzaz Evolutionary Biology of Animal Cognition. R. Dukas Pollination Syndromes and Floral Specialization. C.B. Fenster, WS. ARMBRUSTER, P. WiLson, M.R. Dupasi, and J.D. Tiomson On the Ecological Roles of Salamanders. R.D. Davic and H.H. Weisu, JR. Ecological and Evolutionary Consequences of Multispecies Plant-Animal interactions. S.-Y, SrRAuss and R.E. IRwIN Spatial Synchrony in Population Dynamics. A. Lirsitoip, W.D. Kornic, AND O.N. BjoRNSTAD Ecological Responses to Habitat Edges: mechanisms, Models, and Variability Explained. L. L. Ries, RJ. FLETCHER, JR. J. Battin, and TD. Sisk Evolutionary Hea and i aes mpacts of Marine Eukaryotic Phytoplankton. ME. Katz, Z.V. Finke, D. Grzesyk, A-H. KNoiL, and P.G. FALKOWsKI Regime Shilts, rere and Sie in eye oo C. Fouke, S. CARPENTER, B. WALKER, M. SCHEFFER, T. ELMavist, L. GUNDERSON, and C.S, HOLLIN Ecology of Woodland Herbs in Temperate Deciduous eee DF WHIG! _ Southwest Australian Floristic Region: evolution and Conservation 7 a Global Hot Spot of 3iodiversity. $.D. Hopper and P. Gioia Benn induced Phenotypic Plasticity in Organisms with Complex Life Histories. M.F. Benarp The Evolutionary Ecology of Novel Plant-Pathogen interactions. LM. Parker and G.S. GILBERT SIDA 21(3): 1860. 2005 VEGETATION AND FLORA OF AMERICAN BEECH WOODS NATURE PRESERVE, CLARK COUNTY, ILLINOIS Bob Edgin Gordon C.Tucker! and John E. Ebinger Illinois Nature Preserves Commission Department of Biological Sciences Springfield, Illinois 62701, U.S.A. Eastern Illinois University Charleston, Illinois 61920, U.S.A ABSTRACT American Beech Woods N Preserve is located on Cea till in ae Wabash Border Division of eastern Illinois. The plant life of this 8 ha si 1999-2001 growing seasons. We documented a total of 207 vascular oa species in 148 gene a n families (LO pteridoy J] gymnosperm, +9 monocots, and 147 dicots). We also nat vegetation using a ee es -ran- dom line-strip method. Tree density iene 249 trees/ha with a basal area of 22.9] m?/ha. nee grandifolia Ehrh. (American beech) was the dominant tree species with 63 trees/ha, a basal a 7.34 m?/ha,and an importance value a | (possible 100). Acer sacc ae um Marsh. (sugar maple) r ae second in es uate IV =15.7) with most individuals in the 10-19.9 cm diameter class. Quercus velutinda Lam. | d Carya glabra (Mill) Sweet (pignut ee were the only other over- stony trees witha an iouegtanes yelue aes as an 10. In comparison with an earlier study in 1973, (399 trees/ha vs. 249 trees/ha) and basal area (33.35 ye vs. 22.9] m?/ha). RESUMEN eeoeua Napimaw A : ta ubicad li la none del Rio WabaSe en Illinois oriental. La flora del ae ide 8 hecta I inad crecimiento en los anos 1999-2001. Hemos identificado 207 especies de plantas vasc ones que incluyen . géneros de 71 familias ene 10 pteridofitas, ae aera y 147 dicotiledoneas. Muestreamos - — usance el método de linea-franja ale: a. La densidad promedia era 249 arboles por | basal de 22.91 m2/ha. Be come id Ehrh. (haya) era la especie de arbol dominante. Habia 63 arboles ors hectarea con una area basal de 7.32 m?*/ha y un valor de importancia de 26.1 (posibilidad de 100). Acer saccharum (arce de azticar) — tenia una importancia secundaria con un valor de im postanca de 167 7.La mayoria de los individuos pertenecian a la clase de tamanoentre 10-19.9cm. Quercus velutina Lam. (roble negro) y Carya glabra Mill. (nuez de cerdo) eran los tnicos arboles que tenian un alse de importancia de mas de 10. En comparacion con el estudio de 1973 el t BOsaHe ae ufrido una perdida de densidad (399 arboles por ha v. 249 arboles por ha) y area basal ( 33.35 m’/ ha v. 22.91 m’/ha.) INTRODUCTION At the beginning of extensive European settlement (ca. 1800), about 61% of Illi- nois was prairie and savanna. The remainder, mostly the more rugged terrain, was woodland and forest (Ktchler 1964, Anderson 1970; Iverson et al. 1991. Ebinger 1997). In such areas of rugged terrain, tree species composition varied ‘Corresponding author; cfgct@eiu.edu. SIDA 21(3): 1861-1878. 2005 1862 BRIT.ORG/SIDA 21(3) locally with oaks (Quercus spp.) and hickories (Carya spp.) being the common forest species on drier mostly upland sites. Mesophytic species such as elm (Ulmus spp.), ash (Fraxinus spp.), and sugar maple (Acer saccharum Marsh.) were associated with the dissected ravines and narrow river floodplains (Braun 1950; Anderson 1983; Cowell & Jackson 2002). At the eastern edge of Illinois, particularly in the Wabash Border Natural Division, many of these forests con- tained American beech (Fagus grandifolia Ehrh.), tulip tree (Liriodendron tulipifera L.), and other tree species typically found in forests to the east of IIli- nois (Schwegman 1973). American beech has a wide range comparable to that of other major Eastern Deciduous Forest trees. Beech-maple forests usually included some species of oaks and hickories and reached the western limit of their range in east-central and southern IIli- nois. In Illinois, the few remaining examples of this community type are asso- ciated with steep, deeply dissected ravine systems, narrow valleys, and narrow to broad ridges. The beech-maple component has a rich herbaceous layer on the mesic slopes and an oak-hickory component on the ridges and more level uplands. These remnants have been variously disturbed by logging, grazing, and exotic species invasion. Three examples of this forest community located in the Wabash River Valley have been dedicated as Illinois Nature Preserves (McFall & Karnes 1995). Oc- currences of American beech in this region have enhanced significance as these populations represent the western edge of the range of a wide ranging eastern North American species. The American Beech Woods Nature Preserve contains one of these protected beech-maple forests. The objectives of our study were to document the vascular flora; to determine the composition and struc- ture of the woody and herbaceous vegetation; and to analyze changes in the forest composition that occurred since the forest was last studied in 1973. —_— DESCRIPTION OF THE STUDY AREA The American Beech Woods Nature Preserve, dedicated as a nature preserve in 1985, is located in Lincoln Trail State Park (Fig. 1), about 5 km south of Marshall, Clark County, Illinois (SE/4, NW/4, S2, TION, R12 W, 39°20'30'N, 87°42'45"W). Located in the Southern Upland Section of the Wabash Border Natural Divi- sion about 15 km from the Indiana state line, the preserve is situated on IIlinoian glacial till about 20 km south of the terminal moraine of Wisconsin glaciation (Schwegman 1973). The preserve, about 8 ha in size, has rugged topography, ranging in elevation from 167 m at the edge of Lincoln Trail Lake to 190 m at the highest point. Topographic features include steep-sided ravines, valley walls of various slope aspects, and narrow ridges. The western boundary of the pre- serve follows the shoreline of Lincoln Trail Lake. Presently most of the preserve is high quality, old second growth, mesic and dry-mesic upland forest (White & Madany 1978). Based on the original Government Land Office survey records ~— EDGIN ET AL., FLORA OF AMERICAN BEECH WOODS NATURE PRESERVE 1863 \ ets KK P ivincston ff \ atl {7 | + a % _ bss incout : TRAIL, ® TATE Fic. 1. The location of Clark County in eastern Uline, ane me incauon GY uae Pee Woods pane Preserve in Lincoln Trail State Park, S of Marshall, | an at the Illinois State Archives in Springfield (Hutchison 1988), the woods still retains many of the species present prior to settlement by Europeans. The overstory of the preserve was sampled in 1973 as part of an extensive study that examined the structure and composition of beech-maple forests in Illinois, Indiana, Michigan, and Wisconsin (Dunn 1978). Two prescribed burns (Nov 1993 and Dec 1998) have been conducted in the preserve, while seedlings and saplings of sugar maples have been removed recently from the flat uplands and ridgetops. The soils of the ridgetops are me silt loam, a somewhat poorly drained soil that formed in loess underlain by Ilinoian glacial till (Awalt 1979). Soils of the wooded slopes and drainages are Hickory loam, a well-drained soil that developed in Hlinoian glacial till. These soils overlay bedrock composed of Penn- sylvanian shale and sandstone (Dunn 1978). e climate is continental, characterized by hot, humid summers and cold winters. Weather station records for Marshall, Illinois, about 6 km north of the preserve, indicate that the area receives an average annual precipitation of 104 cm which falls mostly as rain during the period of April through September (Weather.com 2002). January is the coldest month with an average high tem- perature of 1° C and an average low temperature -8° C. The record high for the month was 27° C on January 9, 1932 and the record low was -31° C on January 18, 1930. July is the hottest month with an average high temperature of 31° C and an average low of 17° C. The record high for the month was 43° C on July 14,1936 and the record low was 7° C on July 1, 1937 (weather.com 2002). 1864 BRIT.ORG/SIDA 21(3) MATERIALS AND METHODS The area was visited numerous times during the 1999, 2000, and 2001 growing seasons. During each trip, all new flowering or fruiting species encountered were collected, the specimens identified, and deposited in the Stover-Ebinger Herbarium (EIU) of Eastern Illinois University, Charleston. Native status and nomenclature follows Mohlenbrock (2002). All vascular plant taxa observed are enumerated in the Annotated Species List (Appendix D. Vegetation sampling to determine quantitative abundance of woody and herbaceous species was conducted on August 29 and 30, 2000. We employed the stratified-random line-strip method of Lindsay (1955) as modified by Donselman (1973), Levenson (1973), and Dunn (1978). Using this method, over- story trees, saplings, shrubs, and ground layer strata were sampled simulta- neously in rectangular plots positioned along transect lines. Sample plots for the overstory trees (=10.0 cm dbh) were delimited using a 100 m tape divided into 25 m sections. Overstory trees were sampled in 10 m x 25 m (0.025 ha) with four located along each transect. All trees whose centers were located within the plots were included in the sample. Aspect of the plot, species, and diameter at breast height (dbh) were recorded for each individual located within the boundaries of each plot. Large saplings (5.0 cm dbh - 9.9cm dbh), intermediate saplings (=2.5 cm dbh; = 4.9 cm dbh), small saplings (= 50.0cm tall; < 2.4cm dbh), shrubs, and the groundlayer (woody seedlings <50.0 cm tall and all herbaceous taxa) were sampled in rectangular plots located at the zero, 25m, 50 m, and 75 m mark of the tape. Aspect, species, and the num- ber of individuals were recorded for all vascular plants in each category that fell within one meter from the tape along a section 2.5m long (0.00025 ha plot). When all plots along the 100 m transect line were sampled, a section of 1/ 2" steel conduit marked “Edgin 2000” was driven at each end of the tape to fa- cilitate the relocation of the transect line for future studies. A new 100 m transect line, located a minimum of 25 m distant from the first line and perpendicular to the ravine was then established and the sampling procedures repeated. This process was replicated along ten 100 m transect lines providing a total of 40 plots in each category. Density (trees/ha), basal area (m*/ha), frequency (%), relative density, rela- tive dominance, relative frequency, importance value (relative density + rela- tive dominance + relative frequency /3) and average basal area were determined for each species in the overstory tree stratum. Density (stems/ha), frequency (%), relative density, relative frequency, and importance value (relative density + relative frequency /2) were determined for each species in the small, interme- diate, and large sapling, shrub, and ground layer strata. The Floristic Quality Index (FQD of the site was determined using the Co- efficient of Conservatism (CC) assigned to each species by Taft et al. (1997). The EDGIN ET AL., FLORA OF AMERICAN BEECH WOODS NATURE PRESERVE 1865 CC for each species in the Illinois flora was determined by assigning an integer from O to LO for each species based on its tolerance to disturbance and its fidel- ity to habitat integrity. The FQI is a weighted index of species richness (N = number of species present), and is the arithmetic product of the average Coeffi- cient of Conservatism (C-Value = the average of all species CC’s) multiplied by the square root of the species richness (\/N): FQI = C-Value (\/N). Therefore the FQI indicates the level of habitat degradation and provides an assessment of the quality of each tract based on the taxa present. It is particularly useful when combined with quadrat-based sampling methods and provides a way of mak- ing quantitative comparisons among sites. The Sorenson Coefficient of Com- munity (Serenson 1948) was used to determine the similarity of the ground layer vegetation on the different slope aspects in the study area. The index is calculated as 2c/(a+ b +2c), where a is the number of species unique to sample a, bis the number of species unique to sample b, and c is the number of species shared by both samples (Small & McCarthy 2001). RESULTS AND DISCUSSION Overstory and woody understory composition and structure.— During the sam- pling of the overstory, a total of 17 tree species was encountered, including two understory species having little chance of reaching the canopy; seven additional tree species were found elsewhere on the preserve, ie., outside the sampling transects, accounting for a total of 24 species. Overall tree density was 249.0 trees/ha and total basal area was 22.91 m?/ha. American beech ranked first in basal area, relative density, relative dominance, and importance value (Table LD). It was the most frequently encountered species, occurred in 70% of the plots, and was evenly distributed throughout most diameter classes. Sugar maple ranked second in importance value and relative density and third in basal area. It occurred in 57.5 % of the plots and was most abundant in the smaller diam- eter classes with 63% of the individuals encountered being in the 10-19.9 cm diameter class. Quercus velutina Lam. (black oak) (IV = 11.7) and Carya glabra (Mill) Sweet (pignut hickory) (IV = 10.0) were the only other taxa encountered with importance values greater than 10. Black oak was most abundant in the medium and large diameter classes while pignut hickory was most abundant in the smaller diameter classes. Of the remaining overstory trees, tulip tree was the only species to be rep- resented in most diameter classes, being present in low numbers in all but the largest diameter class (Table 1). Carya ovata (Mill.) K. Koch (shagbark hickory) was present only in the smaller diameter classes with no individuals over 39.9 cm dbh being encountered. Carya tomentosa (Poir. ex Lam.) Nutt. (mockernut hickory) was present in low numbers in the small and medium diameter classes while Quercus alba L. (white oak) was present only in the medium diameter TA Ble 1. Density (#/ha) by diameter classes : ey mportance value, and average dbh are given for tree taxa encountered during sampling of American Beech Woods nti ne Clark County, Illinois ( cm), total density (#/ha), basal area (m*/ha), frequency (%), relative dé it lativ Ae included is the importance value and average dbh per tree taxa from the 1973 study (Dunn 1978). ice, relative frequen Avg. 1973 Total Basal Basal Basal 10.0 200 300 400 500 600 70.0 Density Area ‘Freq. Rel. Rel. Rel. Area/) 1973 Area/ 19.9 -29.9 -39.9 -49.9 -59.9 -69.9 -79.9 80.0+ (#/ha) (m*/ha) (%) Den. Dom. Freq. IV tree IV ree Fagus cela ia 160 120 12.0 10.0 8.0 40 - 1.0 63.0 7.34 700 253 320 21.1 26.1 0.12 25.9 0.09 cer sa m 29.0 6.0 40 5.0 : 2.0 -- - 46.0 259 575 185 11.3 17.3 157 0.06 15.5 0.06 cen eee - 40 9.0 6.0 2.0 3.0 1.0 25.0 3.72 30.0 100 16.2 90 11.7 0.15 7.3 0.13 Ca aya gla 12.0 7.0 8.0 2.0 1.0 -- 24 30.0 1.87 325 12.1 8.2 98 10.0 0.06 43 0.08 Liriodendron tulipifera 3.0 3.0 40 2.0 1.0 1.0 1.0 - 15.0 1.80 25.0 6.0 79 75 71 0.12 54 0.18 Carya ovata 11.0 9.0 4.0 - - - 24.0 0.95 25.0 9.6 41 75 7.1 0.04 08 0.02 Carya tomentosa 40 2.0 2.0 3.0 1.0 1.0 - -- 13.0 1.02 20.0 5.2 45 6.0 5.2 0.08 41 0.06 Quercus alba -- 4.0 1.0 1.0 1.0 - 7.0 1.12 17.5 2.9 49 5.3 44 0.16 11.0 0.11 Quercus rubra - : - -- 1.0 1.0 - 1.0 3.0 1.06 10.0 1.2 4.6 3.0 2.9 0.36 8.2 0.14 Ulmus americana 6.0 1.0 - -- ; 7.0 0.13 10.0 2.8 0.6 3.0 2.1 0.02 0.4 0.03 Fraxinus pennsylvanica 1.0 -- 2.0 7 - “ 3.0 044 75 1.2 19 2.2 1.8 0.15 0.7 0.11 Nyssa sylvatica - -- 3.0 - -- - 3.0 0.29 75 1.2 1.3 2.2 1.6 0.10 6.2 0.04 Ulmus rubra 1.0 1.0 -- 1.0 -- -- 3.0 O22 £5 1.2 1.0 2.2 1.5 0.08 -- Sassatra | 3.0 = - == = 3.0 0.04 5.0 1.2 0.2 1.5 1.0 0.01 0.7. 0.09 Acer rubrum - - 1.0 1.0 - 2.0 0.23 2:5 0.4 1.0 0.8 0.9 0.23 - Juglans nigra - - 1.0 - -- 1.0 0.08 2.5 0.4 0.3 0.8 0.5 0.09 Se Cornus florida 1.0 -- - -- - - - 1.0 0.01 2.5 0.4 0.0 0.8 0.4 0.0 0.8 0.01 Others (7 taxa) - - -- -- 8.7 Totals 86.0 46.0 10 320 160 14.0 20 20 2490 22.91 100.0 100.0 100.0 100.0 100.0 998L (€)LZ VdIS/9¥O'LINA EDGIN ET AL., FLORA OF AMERICAN BEECH WOODS NATURE PRESERVE 1867 classes. Of the remaining trees, most were present as widely scattered individu- als with two, Sassafras albidum (Nutt.) Nees (sassafras) and Cornus florida L. (flowering dogwood) being understory trees. The overall tree density declined from 399.0 trees/ha in 1973 to 249.0 trees/ ha in this study (Dunn 1978) (Table 1). Total basal area also d d from 33.35 m2/ha in 1973 to 22.91 m2/ha in 2000. However, the importance values for American beech, sugar maple, and most other species were very similar to those reported in the previous study. The importance value of shagbark hickory was considerably higher in the present study while those of white oak and red oak were considerably lower. Seven species having a combined importance of 8.7 in the 1973 study were not encountered during the sampling in this study. Of those species, Carya cordiformis Wang.) Koch (bitternut hickory) and Fraxinus americana L. (white ash) had IV’s totaling L5 in the 1973 study. Amelanchier arborea (Michx. f.) Fernald (shadbush), Carpinus caroliniana Walt. (musclewood), and Ostrya virginiana (Mill.) K. Koch Gironwood) are understory trees that had a combined importance value of 4.8 in the 1973 study. Since no permanent transects were established in the previous study, these changes may be more reflective of sam- pling error rather than changes in the composition of the forest stand. Tilia americana L. (basswood) (IV = 2.4 in 1973) was encountered neither during the sampling, nor the site visits and appears to have been extirpated from the preserve. In the understory, sugar maple and American beech ranked first and sec- ond, respectively in all three sapling categories. Of the remaining understory trees, only flowering dogwood and ironwood were present in all of the sapling categories (Table 2). Hickories were not common in the understory and no oaks were encountered. A total of 26 dead-standing saplings were encountered in 11 plots. These stems were all in the medium and small sapling categories and occurred in plots located on ridgetops or slopes with an east, west, or southwest aspect. Of 26 stems encountered, 19 appeared to have been top-killed by fire 14 sugar maple, two ironwood, two American beech, and one hickory). Seven dead-standing dog- wood saplings were encountered, but it was not clear as to whether these indi- viduals were dead asa result of fire or anthracnose. Hydrangea arborescens L. (wild hydrangea) and Lonicera maachii (Rupr) Maxim. (bush honeysuckle) were the only shrub taxa encountered. Wild hy- drangea was present in one plot located in a creek bottom while one bush hon- Estee ely was encountered in a plot with a north-facing aspect. ition and structure.—A total of 70 taxa was encountered in the groundlayer (Table 3). The Carex spp. (sedges) as a group ranked first in importance value (IV =88) and occurred in 50% of the plots. Pilea pumila (L.) A. Gray (clearweed) ranked second in pores vale being most abundant in plots that occurred in creek bott t-facing slopes. Sanicula Taste 2. Density (stems/ha) arranged by aspect, total density (#/ha), frequency (% of plots in which each taxon was observed), relative density, relative fre- quency, and importance value for large saplings (=5.0 cm dbh-9.9 cm dbh), intermediate saplings (= 2.5 cm dbh-< 5.0 cm dbh), and small saplings (>50 cm tall-2.5 cm dbh) encountered during sampling of American Beech Woods Nature Preserve, Clark County, Illinois. Aspect All plots Large Saplings (5—10cm dbh) Total Crk. Ridge Density —_ Freq. Rel. Rel. Bot SW N W E S NE SE #/ha (%) Den Freq IV Acer saccharum - 114 66 198 -- 400 400 a2 = 130 17.5 65.0 46.7 55.9 Fagus grandifolia 57 -- -- 66 80 -- -- -- =+ 30 7.5 15.0 20.0 17.5 Carpi lini 114 ie = -- ae a -- -- = 20 5.0 10.0 13.3 11.7 Cornus florida te 57 -- -- -- -- -- -- -- 10 5.0 5.0 13.3 9.1 Carya ovata -+ -- -- == -- -- -- -- -- 10 25 5.0 6.7 5.8 Totals 171 171 66 264 80 400 400 -- == 200 100.0 100.0 ~=100.0 Intermediate Saplings (2.5-4.9 cm dbh) Acer saccharum 228 22 198 132 a 400 133 800 400 200 30.0 52.6 46.3 49.4 Fagus grandifolia 57 160 66 198 160 _ 266 400 120 22.5 31.7 34.6 33.2 Cornus florida -- i 132 -- 80 -- -- -- = 30 5.0 79 77 78 arpir arolini ta = -_ -- 80 -- -- -- -- 10 2.5 2.6 3.8 3.2 Ulmus rubra -- -- a -- 80 ee -- am 10 25 2.6 3.8 3.2 Carya ovata = 80 = = se as = -- ie 10 2.5 2.6 3.8 3.2 Totals 285 240 396 330 320 480 399 800 800 380 100.0 100.0 ~=100.0 Small Saplings (> 50 cm tall—2.4 cm dbh) Acer saccharum 513 720 462 924 -- 80 266 3200 800 600 47.5 43.4 31.1 37.3 Fagus grandifolia 456 320 66 330 320 =a 133 200 ee 240 375 17.4 245 20.9 Ostrya virginiana 25 -- -- 528 80 -- -- 400 400 120 12.5 8.7 8.2 8.5 Ulmus rubra 114 80 -- 66 aes -- <5 800 -- 90 15.0 6.5 9.9 8.2 Cornus florida 57 80 198 66 80 a= -- =< -- 70 15.0 5.1 9.9 75 Asimina triloba ais =i -- -- a 1280 == a = 160 25 11.6 1.6 6.6 898L (€)L? VOIS/9YO LINE Taste 2. continued Aspect All plots Small Saplings (>50 cm tall—2.4 cm dbh) Total Crk. Ridge Density Freq. Rel. Rel. Bot. SW top W E S NE SE (#/ha) —(%) Den. Freq. IV Carpinus caroliniana -- - -- 80 -- -- -- -- 20 5.0 1.5 3.4 25 ae eed vanica 57 oe = 66 i as as ae =i 20 5.0 des 3.4 25 Prunus serotin a == eS a ee =< 400 ae 20 255 15 1.6 1.6 Liriodendron tu eee ee o- - - == a as 200 -- 10 2.5 0.7 1.6 1.1 oe — - - == ae Ass 200 a 10 2.5 0.7 1.6 1.1 q -- -- - 66 -- : -- -- -- 10 25 0.7 16 11 Fraxinus americana ao : ae 66 oe : ai = 10 2.5 0.7 1.6 1B Totals 1254 200 726 2112 560 360 399 5400 1200 1380 100.0 100.0 100.0 Taste 3. Density (#/ha) cine by aspect, total density (#/ha iF impor tar We value for Clark County, Illinois. groundlay er taxa in cluding woody species (< <50¢ frequency vias of pees in nen a) taxon was observed), relative density, relative frequency, g sampling of American Beech Woads Nature Preserve, Aspect All plots Crk. Ridge Density Freq. Rel. Rel. Bot. W E S NE SE (#/ha) (%) Den Freq. IV Carex spp. 9324 2880 25974 13320 1332 : 6660 2000 : 14100 50.0 10.4 Fe? 8.8 Pilea pumila 46620 —-- 1998 1332 Be Bios! 70000 - - 14600 = 22.5 10.8 3.3 7.0 Sanicula spp. 1332 2664 666 22644 800 4662 28000 —-- 10500 =30.0 Le 43 6.0 Asa anaden -- -- a 87246 = 800 4662 =e eke 14600 LS 10.8 fl 5.9 Viola sororia 800 10656 666 4995 4000 1998 1400 - 7800 35:0 5:7 al 5.4 Lirfodendron t tul ifera 888 7200 1998 3996 999 800 3 ie 4500 35.0 3.3 5.1 4.2 Solidago c ae 800 7992 9324 333 =e 333 18000 —-- 4800 32.5 3.5 47 4.1 Impatiens a 11544 - 6660 2664 : 333 2000 = 4600 20.0 3.4 2.9 3.1 JAYISIYd JYNLYN SCOOM HI3938 NVDINIWY 40 VHOTd “TY 13 NIDG3 6981 Taste 3. continued Aspect All plots Crk. Ridge Density Freq. Rel. Rel. Bot. SW top N WwW E $ NE SE (#/ha) (%) Den. Freq. IV Acer saccharum 332 1600 7326 1998 333 3200 333 s 500 30.0 1.8 43 3.0 Polystichum 332 1600 666 1332 - 333 8000 6000 ~=1700 27.5 1.3 40 2.7 soli Sas 8800 4662 - 333 600 333 - : 2200 22.5 1.6 3.3 2.5 Parthenocissus 1776 3200 66 6660 : 600 : + -- 2200 20.0 1.6 2.9 2.3 quinquefolius Arisaema triphyllum 444 4800 1998 =e -- 2000 333 - - 2600 17.5 1.9 25 2.2 Agerati [tissi 2664 7326 666 = 0) = 2000 = 2000 20.0 1.5 2.9 2.2 Leersia virginica 20424 -- 2 = - 4600 75 3.3 1.0 2.1 Antenoron virginianum 11544 - _ 999 == oe 2000 4000 10.0 2.9 14 ZA oe ma ine a 776 800 666 a= 1332 -= <5 32000 - 2600 15.0 1.9 2.2 2.) m 444 : 1332 333 333 40000 - - 2500 15.0 1.8 2.2 2.0 Ulmus ae. 444 800 2664 = -- 3200 666 -- : 1200 20.0 0.9 29 1.9 888 800 2664 ae 333. 2000 900 20.0 0.7 2.9 1.8 Osmorhiza C el 1332 : 666 330 ao 5600 - = 1600 15.0 12 2.2 1.7 ti 8436 2400 : 7 : 1600 2400 75 18 (lee 1.4 Pinas serotina - 1998 3330 : 332 7 - 900 12.5 0.7 1.8 1.3 Aster spp. 888 666 = = 800 2000 - 600 15.0 0.4 2.2 1.3 Carya spp. ~~ 3200 3996 = -- -- =< - 1000 12.5 0.7 1.8 1.3 Equisetum a 12432 -- 1332 == aS ae -- 2800 2.5 2.0 0.4 1.2 Vitis aestivalis ae - 666 _ ae 8325 Ss = 2500 2.5 1.8 0.4 1.1 ae SPP. 9768 : - =f ae -- 2200 5.0 1.6 0.7 1.1 == : 4662 =< 999 - a 2000 - 1100 Lo 0.8 11 1.0 Hepatica acuti ieba => - -- - - : -- 8000 1800 5.0 1.3 0.7 1.0 Muhlenbergia spp - 7326 7 2331 -- aa - 1800 5.0 13 0.7 1.0 Cornus florida 66 - - -- -- - 100 25 0.1 0.4 0.3 Others (39 taxa) 12876 6000 7992 7326 3996 12800 5661 3200 800 12800 140.0 9.5 20.3 14.9 Totals 170496 63680 109224 146850 43623 49600 44289 214600 24800 136100 100.0 100.0 = 100.0 OZ8L (€)LZ VOIS/OYOLINS Taste 4. Ranking of tree species by importance value for plots that occurred on 9 slope aspects at American Beech Woods Nature Preserve, Clark County, illinois. The species listed are those with the 10 highest overall importance values throughout the preserve and are arranged by descending importance value. Ranking by importance value for each aspect is in parentheses. Aspect All plots Creek Bottom Southwest Ridge top North Southeast West East Northeast Species (7 plots) (7 plots) (5 plots) (5 plots) (5 plots) (4 plots) (4 plots) (3 plots) Fagus grandifolia 28.3 (1) 33.9 (1) 10.0 (5) 30.2 (1) 20.4 (2) 26.7 (1) 33.3 (1) 53.7 (1) Acer saccharum 17.7 (3) 19.2 (2) 5.4 (7) 16.9 (2) 4.5 (10) 27.2 (2) 15.3 (3) 20.2 (2) Quercus velutina — 12.2 (3) 26.5 (1) 6.8 (5) 13.8 (3) 12.1 (3) oo 12.7 (4) Carya glabra ~ 6.8 (6) 259?) 15.0 (3) 4.8 (9) 9.2 (4) — — Liriodendron tulipifera 28.1 (2) 6.8 (5) — 5.6 (6) 10.2 (4) 6.3 (5) — —— Carya ovata — 40 (7) 8.4 (6) 4.9 (8) 22.0 (1) 9.2 (6) 12.7 (4) — Carya tomentosa 5.9 (6) 2.3 (9) 10.3 (4) — 6.2 (6) — 47 (8) 13.4 (3) Quercus alba — 10,7 (3) 47 (9) 5.9 (7) — 16.4 (2) — Quercus rubra — 7.2 (A) — 10.7 (4) = — Sassafras albidum 2.8 (8) — — — -- 45 (7) — Taste 5. Sorenson Index for groundlayer taxa encountered during sampling of American Beech Woods Nature Preserve, Clark County, Illinois. Creek Bottom Southwest Ridge top North West East South Northeast (30 taxa) (20 taxa) (27 taxa) (23 taxa) (20 taxa) (18 taxa) (25 taxa) (18 taxa) Southwest 440 Ridge top 56.1 63.8 North 49.1 32.6 52.0 West 36.0 40.0 55.3 41.9 East 45.8 42.1 48.9 39.0 26.3 South 43.6 48.9 53.8 50.0 44.4 46.5 Northeast 50.0 31.6 57.8 39.0 474 22.2 41.9 Southeast (3 species) 6.1 8.7 6.7 15.4 8.7 0 7.1 9.5 JAWISINd JUNLVN SCOOM H3994 NVIINAWY 40 VYOTd “TW 13 NIDGI LZ8L 1872 BRIT.ORG/SIDA 21(3) wu spp. (snakeroot), Asarum canadense L. (wild ginger) and Viola sororia Willd. (woolly blue violet) were the only other herbaceous taxa with IV's greater than five. Snakeroot was most abundant in plots that had a west and northeast aspect while wild ginger was most common on the north-facing slopes. Woolly blue vio- let was present in low to moderate numbers in most plots. Tulip tree, sugar maple, and sassafras were the most commonly encountered nce seedlings. Oak seed- lings were rare with only one seedling being enc ered on a north-facing slope. Slope aspects.—Among the overstory trees, American beech ranked first in importance value in plots located on most slope aspects (Table 4). It ranked second in plots with a south or southeast aspect and fifth in plots that occurred on ridge tops. Sugar maple ranked second in importance value in plots having a southwest, north, west, or northeast aspect. Black oak ranked first in plots that occurred on the ridge tops, but no higher than third on the remaining as- ects. White oak ranked second and third in plots with an easterly aspect and on the ridge tops, respectively, but was only a minor component or was absent from plots occurring on the remaining aspects. Red oak ranked fourth in plots with a southwest or north aspect and was not encountered in the remaining plots. Among the hickories, shagbark hickory ranked first in plots with a south or southeast aspect while pignut hickory ranked second in plots that occurred on the ridge tops. The plots located on the ridgetops and the southwest-facing slopes had greatest similarity (63.8% Sorenson Coefficient of Community, Sorenson 1948) (Table 5). Plots located on the southeast-facing slopes were considerably dis- similar to plots located on other slope aspects having Sorenson Coefficient of Community percentages that ranged from 0 to 15.4. These low values may be attributed to the low number of plots with southeast aspects (1) and the low species richness of the plot (3). Most other plots had similarity indices that ranged from 36.1% and 57.8%. During the study, we observed 207 vascular plant taxa in the study area: 11 ferns, fern allies, and gymnosperms, 49 monocots, and 147 dicots. Of that num- ber, 23 (11.1%) had a Coefficient of Conservatism (CC) of seven or greater and 10 (4.8%) were non-native taxa. The average CC, when calculated for all taxa, was 3.88 and the FQI was 55.8. When calculated for native taxa only, the average CC and FQI were 4.04 and 57.0, respectively. Sites that have an FQI greater than 35 are considered regionally noteworthy, while sites with an FQI greater than 45 are defined as statewide-significant natural areas (Taft et al. 1997). The composition of American Beech Woods is similar to other beech-maple forests in Illinois and Indiana, having American beech and sugar maple as co- dominants on the mesic slopes with oaks and hickories predominating on the drier slopes or more level uplands (Ebinger 1997, Cowell & Jackson 2002). The decline in overall tree density and total basal area in this preserve is typical of many similar forest stands in the region (Petty & Lindsey 1961; Lindsey & EDGIN ET AL., FLORA OF AMERICAN BEECH WOODS NATURE PRESERVE 1873 Schmelz 1964; Barton & Schmelz 1987). Oak density declines as mature indi- viduals die while shade-intolerant and successional species such as tulip tree, ash, and sassafras, persist as minor components because of gap-phase distur- bances (Cowell & Jackson 2002). APPENDIX I Vascular flora of American Beech Woods Nature Preserve, Lincoln Trail State Park, Clark County, Illinois arranged alphabetically by taxonomic group. No- menclature follows Mohlenbrock (2002). Collection numbers with the T prefix are those of Tucker; while the E prefix indicates specimens collected by Ebinger. All specimens were deposited in the Stover-Ebinger Herbarium at Eastern IIli- nois University, Charleston, Illinois, with some duplicates at ILLS. Taxa preceded by an asterisk (*) are non-native. FERNS AND FERN ALLIES ANNONACEAE Asimina triloba (L.) Dunal £30430 ADIANTAC Adiantum ae sea Fa rd) APIACEAE Cicuta maculata Late 14 739 ASPLENIACEAE ‘ densis (L.) DC.T11738 Cry Asplenium platyneuron (L.) Oakes £29720 Osmorhiza caeny i (Michx.) Clarke 111752 ee Blase) Beatneloyer|) 750 miclancis Woe cama onSeTTIAk hx) Schott £29605 Sai ical F ata (Raf) Pryer & Phillippe E29622 ARALIACEAE Aralia racemosa L.E30 OPHIOGLOSSACEAE Panax quinquefolius L. a Botrychium dissectum Spreng. var. dissectum ARISTOLOCHIACEAE EQUISETACEAE Equiset L.111736 sae hia serpentaria L.E29618 Botrychium dissectum Spreng. var. obliquum 4.0 dense |. £29497 (Muhl.) Clute £30427 Botrychium virginianum (L.) Sw. £29501 ASTERACEAE Ophioglossum vulgatum L. £29889 Ageratina altissima (L.) R.M. King & H. Robins. 30158 THELYPTERIDACEAE es ; Antennaria plantaginifolia (L.) Richards. E29500 Phegopteris hexagonoptera (Michx.) Fée E29604 Arnoglossum atriplicifolium (L.) H.Robins.£30233 Aster lateriflorus (L.) Britt. E 7 CYMNOOEERM= Aster sagittifolius Wedem. ex Willd. £30438 CUPRESSACEAE Aster shortii Lindl. £30436 Juniperus virginiana L.E29890 Erechtites hieracifolia (L.) Raf. £30439 Erig L.) Pers.111742 DICOTYLEDONS Erigeron philadelphicus L.112047 ACERACEAE gad sessilifolium L.var. brittonianum Por- er E30159 Acer saccharum Marsh. E2989 Acer rubrum L. (Observed) ANACARDIACEAE Hieracium gronovil L.E30235 Toxicodendron radicans (L.) Kuntze E30232 Krigia biflora (Walt.) Blake 112041 Lact densis L.E30236 Eutham eLannelg (L.) Nutt.ex Cass. £30293 L.E30160 1874 Prenanthes Hticci ma L.E30440 Senecio glabellus Poir. E29608 Senecio obovatus ae 112044 ae os caesia L. 1g ae £30294 : Salidad i s Aiton £30443 Solidago ulmifolia Muhl. ex Willd. E30237 BAP a MINACEAE is Meerb. £30161 BERBERIDACEAE Podophyllum peltatum L.E29487 CORYLACEAE Carpinus caroliniana Walt. var. virginiana (Marsh.) Fernald 111775 Corylus americana Walt. E29895 Ostrya virginiana (Mill.) K. Koch E29897 BORAGINACEAE Cynoglossum virginianum L. £29620 Hackelia vit giniana (L.) LM. Johnst. E29894 BRASSICACEAE Dentaria laciniata Muhl. £29490 CAESALPINIACEAE Cercis canadensis L.111765 CAMPANULACEAE Campanulastrum americanum (L.) Small £30162 manele mee L.TI1782 lia siphilitica L.E30295 CAPRIFOLIACEAE *Lonicera maackii (Rupr.) si E29610 Sambucus canadensis L.T Symphoricarpos orbi ae a £30164 iburnum prunifolium L.E29898 CARYOPHYLLACEAE Silene stellata (L.) Aiton E30165 CORNACEAE Cornus florida L.E29726 EBENACEAE Diospyros virginiana L. E2 9900 aa ieianinaiies stifolia L.111773 EUPHORBIACEAE Acalypha rhomboidea Raf, £30238 aes BRIT.ORG/SIDA 21(3) FABACEAE Amphicarpaea bracteata (L.) Rickett & Stafleu nodium nudiflorum (L.) DC.111741 Desmodium paniculatum (L.) DC. £30298 “Robinia pseudoacacia L.E29901 FAGACEAE Fagus grandifolia Enrh.112051 uercus alba 2 Quercus imbricaria Michx. E30296 Quercus palustris Muenchh £30433 Quercus rubra 0434 Quercus velutina wil m. 712305 eee is Walt.E29904 GERANIACEAE Geranium maculatum L.E29492 HAMAMELIDACEAE Liquidambar styraciflua L.E30239 HYDRANGEACEAE Hydrangea arborescens L.E29903 HYDROPHYLLACEAE | ly UP hyllum virginianum L. E301 68 oe ctatum Lam.1T11776 JUGLANDACEAE Carya cordiformis (Wangenh.) K. Koch E3043 1 Carya glabra (Mill.) Sweet E30301 Carya ovata (Mill.) K. Koch E30432 Carya tomentosa (Poir.) Nutt. E30300 Juglans nigra L. E3024] LAMIACEAE is L.E30242 haan viDINIUS L. £3030 eck £29617 Prineiia rent ie 1 ee rentiol ium Schrad. £29905 loriaq incanga Ri hl.E29906 olen Teucrium canadense L.T11735 ee ac alhid utt.) Nees 111759 Sincere benzoin (L.) Blume 111764 MAGNOLIACEAE Liriodendron tulipifera L.E29728 EDGIN ET AL., FLORA OF AMERICAN BEECH WOODS NATURE PRESERVE MENISPERMACEAE Menispermum canadense L.E29615 MORACEAE Morus rubra L.E30169 NYSSACEAE Nyssa sylvatica Marsh, E29908 OLEACEAE Fraxinus americana L.E29729 Fraxinus pennsylvanica Marsh. £30435 *Ligustrum vulgare L. £29730 ONAGRACEAE Circaea lutetiana L. ssp. canadensis (L.) Aschers. & Magnus E29910 OROBANCHACEAE Conopholis americana (L.) Wallr.E29619 OXALIDACEAE Oxalis stricta L.E29624 Oxalis violacea L.E29625 PAPAVERACEAE Sanguinaria canadensis L.£29498 PHRYMACEAE Phryma leptostachya L.111760 PHY TOLACCACEAE Phytolacca americana L. E2991] POLEMONIACEA Phlox divaricata i spp. laphamii (Wood 712045 Polemonium reptans L. £29731 POLYGONACEAE Persicaria punctata (Elliott) Small £30303 Antenoron virginianum (L.) Roberty & Vautier a) Wherry £30170 *Rumex crispus L.111767 PORTULACACEAE Claytonia virginica iz E29488 nner ea pachypoda Pe — 6 ee acutilo Ranunculus eae ‘ See Ra ulus recurvatus Poir. 112046 Ranunculus cantentniondll G in £29494 Thalictrum dioicum L.T1203 = ROSACEAE Agrimonia g i] AJ>\] Wallr.E30173 1 ee 1875 Agrimonia parviflora Soland. ex Aiton E30243 Agrimonia pubescens Wallr.E30172 Amelanchier arborea (Michx. fil.) Fernald 712050 um canadense Jacg.T11761 Geum vernum (Raf.) Torrey & A. Gray E29629 Potentilla simplex sur = 630 Prunus serotina Ehrh. E *Rosa multiflora ee ex Sane E29628 Rubus allegheniensis fiers L.H. Bailey E29634 Rubus flagellaris Willd. £29632 Rubus occidentalis L.11 ; 777 Rubus pensilvanicus Poir.ex Lam.T11732 UBIACEAE Cephalanthus occidentalis L.111734 ral paeaiins L.E29489 aezans ee TI17 orrey . ay £29733 Galium triflorum ee T11 Houstonia purpurea L. Se SAXIFRAG Penthorum ae ae L.E30244 SCROPHULARIACEAE i Ns alatus Aiton 111774 adensis L.E29735 arilandica L.E30174 alitim byl iq CIOPATIGIGlicl Ii ULMACEAE Ulmus americana L.111737 Ulmus rubra Muhl.111768 URTICACEAE Boehmeria cylindrica (L. Laportea canadensis (L.) Wedd. £30246 Parietaria pensylvanica Muhl. ex Willd. £29915 Pilea pumila (L.) A. Gray E30304 VERBENACEAE Verbena urticifolia L.E29917 VIOLACEAE Viola palmata L.E29503 Viola pratincola Greene E29502 Viola sororia Willd. £29623 VITACEAE Parthenocissus quinquefolia Vitis aestivalis Michx. E30305 (L.) Planch. E30248 MONOCOTYLEDONS ALISMACEAE Alisma triviale Pursh 111733 1876 eae (L.) Schott E29626 Arisaema triphyll um (L.) Schott 112043 MELINACEAE Tradescantia subaspera Ker 112033 Tradescar Ita virgit val Le de £29505 CYPERACEAE Carex albicans Willd. ieee Carex blanda Dew Carex eae ae ex ie E29637 Carex glaucodea Tuckerm. £30176 Carex gracilescens Steudel T12039 Carex hirsutella Mack. E29927 Carex hirtifolia Mack.T12048 Carex lurida Wahl. Tl] 762 Carex pensylvanica Lam. E29509 Carex rosea Schk. ex Willd. 112037 Scirpus georgianus Harper T1763 cui wiped jaternata (Walt.) J.F. Gmel. £29925 JUNCACEAE Juncus tenuts Willd.T11772 Luzula multiflora (Retz.) Lej.T11753 LILIACEAE Allium tricoccum Aiton E29508 ae acina racemosa (L.) Desf.E2961 1 Beck E29507 Uoulana ee ee sm. E2961 2 CHIDACEAE =< OR Corallorhiza odontorhiza (Willd.) Nutt. (Ob- ved) BRIT.ORG/SIDA 21(3) Galearis spectabilis (L.) Raf.E29926 Liparis liliifolia (L.) Rich. E30307 POACEAE oe gigantea Roth 111766 grostis hyemalis (Walt.) BSP. E30230 Brachyelytr rectum eas Beauv. E29918 pioak Siig: ens Muhl.T Cinna arundinacea L. E301 . Danthonia spicata (L.) Roem. & Schult. £29721 ise aicaie 1 boscli st ed acer osole 748 thol) Dichanthelium dichotomum (L.) Gould E29723 Dichantheliurm lindheimeri (Nash) Gould 711746 ae jum microcarpon (Muhl.) Mohlenbr. 711747 5 e) es a) e) c on N = Elymus hystrix L.E29922 Elymus villosus Mubl. oo pea mg us L.E30178 1 Schreb.T11780 verticillata (Pers.) Alekseev 112036 sera striata (Lam.) Hitche. 111754 Leersia virginica Willd. E29923 Muhlenbergia schreberi J.F.amel. £30429 Muhlenbergia sobolifera ene £30179 “Poa compressa L.E30306 Poa sylvestris A. Gray 112049 SMILACACEAE Smilax tamnoides L. var. hispida (Muhl.) Fernald 30180 ACKNOWLEDGMENTS — We wish to thank the Illinois Department of Natural Resources (IDNR) and the Illinois Nature Preserves Commission for granting permission to conduct the study and collect specimens. Appreciation is also extended to Matt Bowyer, Robert Gillespie, and Amy Young (IDNR) for their assistance with data collec- tion and manuscript preparation. Scott Meiners provided helpful comments on data analysis and manuscript revision. Identification of Carex specimens was confirmed by Tony Reznicek (MICH). The abstract was translated into Spanish by Daniel Johnson (U.S. State Dept.). REFERENCES ANberSON, R.C. 1970. Distribution of forest and prairie in Illinois about 1820. Illinois Nature Preserves Commission, Rockford (Map.) EDGIN ET AL., FLORA OF AMERICAN BEECH WOODS NATURE PRESERVE 1877 Anperson, R.C. 1983. The eastern prairie-forest transition - an overview. In: R. Brewer, ed. Proceedings of the 8th North American Prairie Conference, 1982, Western Michigan University, Kalamazoo. Pp. 86-92. Awatt, F.L. 1979. Soil Survey of Clark County, Illinois. U.S. Dept. Agric., Washington, D.C Barton, J.D. and D.M. ScHmeLz. 1987. Thirty years of growth records in Donaldson's Woods. Trans. Indiana Acad. Sci. 96:209-214. Braun, E.L. 1950. Deciduous forest of Eastern North America. Blakiston Books, Philadelphia. Cowett, C.M.,and M.T. Jackson. 2002. Vegetation change ina forest remnant of the eastern presettlement prairie margin, USA. Nat. Areas Jour. 22:53-60. Donsetman, H.M. 1973. An ecological analysis of the shrub stratum of the beech-maple forest type. M.S. Thesis #1085, Indiana State University, Terre Haute, Indiana. Dunn, C.P.1978.An ecological analysis of the tree stratum of the beech-maple forest type. M.S. Thesis #1299, Indiana State University, Terre Haute, Indiana. Esincer, JE. 1997. Forest communities of the Midwestern United States. In: M.W. Schwartz, ed. Conservation in Highly Fragmented Landscapes, Chapman and Hall, New York. Pp. 3-23. Hutcuison, M. 1988. A guide to understanding, interpreting, and using the public land survey field notes in Illinois. Natural Areas Jour. 8: 245-255. Iverson, L.R.,G.L.Rotre, TJ. Jacos, A.S. Honains, and M.R. Jerroros. 1991.Forests of Illinois. Illinois Council on Forest Development, Urbana, and Illinois Natural History Survey, Cham- paign, Illinois. Kucuier, A.W. 1964. Potential natural vegetation of the coterminous United States. Am. Geograph. Soc. Spec. Publ. 36. Levenson, J.B. 1973.The herbaceous stratum of the beech-maple forest:A community struc- ture analysis. M.S. Thesis # 1088, Indiana State University, Terre Haute, Indiana. Linosey, A.A. 1955. Testing the line-strip method against full tallies in diverse forest types. Ecology 36:485-495. Linosey, A.A. and DV. ScHmetz. 1964. Comparison of Donaldson’s Woods in 1964 with its 1954 forest map of 20 acres. Trans. Indiana Acad. Sci. 74:169-177. McFaut, D. and J. Karnes. (eds.). 1995. A directory of Illinois nature preserves. Volume 2. Northwestern, Central, and Southern Illinois. Illinois Department of Natural Resources, Division of Natural Heritage, Springfield, Illinois Moxtensrock, RH. 2002.Vascular flora of Illinois. Southern Illinois University Press, Carbon- dale and Edwardsville, Illinois Petty, R.O. and A.A. Linpsey. 1961. Hoot’s Woods, a remnant of virgin timber, Owen County, Indiana. Trans. Indiana Acad. Sci.71:320-326. ScHwecman, J.E. 1973. Comprehensive plan for the Illinois nature preserves system. Part 2. The natural divisions of Illinois. Illinois Nature Preserves Commission, Springfield, Illinois 32 pp.+ map. Smatt,C.J.and B.C.McCartHy.2001.Vascular flora of the Waterloo Wildlife Research Station, Athens County, Ohio. Castanea 66:363-382. 1878 BRIT.ORG/SIDA 21(3) SaRENSON, T.1948.A method of establishing groups of equal amplitude in plant sociology based on similarity of species content. Det. Kong. Danske Vidensk. Selsk. Biol. Skr. (Copenhagen) 5:1-34. Tart, J.B.,G.S.WitHeLm, D.M. Lapp, and L.A. Masters. 1997. Floristic quality assessment for veg- etation in Illinois,a method for assessing vegetation integrity. Erigenia 15:1-95. WearHer.com. 2002. Monthly ges and records for Marshall, Illinois. The Weather Channel. Waite, J.and M.H. Mapany. 1978. Classification of natural Communities in Illinois. In:White, J. Illinois Natural Areas Inventory Technical Report. Natural Areas Inventory, Urbana, IL. Pp. 311-357 SOIL AND ECOLOGICAL FEATURES OF HEXALECTRIS (ORCHIDACEAE) SITES Annie B. Collins James E.Varnum Biology Department 14524 Tamerisk lane University of Dallas Farmers Branch, Texas 75234, U.S.A. 1845 E. Northgate Drive Irving, Texas 75062, U.S.A. Margaret Brown-Marsden' Biology Department University of Dallas 1845 E. Northgate Drive Irving, Texas 75062, U.S.A ABSTRACT Soil and ecological features of the orchid genus I lectri ined to obtai t description of the factors in aS its distribution aad to direct future conservation efforts. Data on canopy and ground cover, tree species diversity, and soil series were obtained for Hexalectris in Dallas County, and over a on historic data on Hexalectris in Texas. We determined that Hexalectris does associate with oak alee juniper, but the amount of cover at Hexalectris sites did not exceed 60%. Ground cover. ity, and percent oak/juniper did not differ between sites with and without Hexalectris. The soil series associated with Hexalectris in this region were Eddy- Brackett entisols of | 8-20% Soil ee was an accurate a of areas in which Hexalectris could be found. Hexalect ob on soil series, a factor which can aid in predicting areas in Bra Hes xd ate ctrisis likely < ae a fi has not yet been located, as wel conservation of this less well-studied genus. asin RESUMEN Se examinaron las caracteristicas ecologicas y del suelo de la especie de orquidea Hexalectris para obtener una descripcion mas precisa de los Boos que eyes su ibe StIECIon pela ucuros S esfuerzos de conservacion I arboreas, 5 le Hexalectri ] dado de Dallas, y sobrepuestos con datos orice: sobre Hexalectris en Ieee Detsriningies aus gee tris si se asocia con roble y Sree: po la cantidad de los sitios de Hexalec y cubierta, | aba 1 iA aise 1 | Hexalectris. especies arboreas, y el } ] es) Las series Hexalectris en esta region eran suelo Eddy-Brackett con 8 al 20 por ciento de eee El ope de suelo fue un NGOSIO preciso para pose aces el area en que se encontraria Hexalectris. n factor que puede ayudar a predecir areas en las quel pueda encontiatse Hexalectris ae todavia no ee ser descritas, al igual que la conser g I TAuthor to whom correspondence should be addressed. SIDA 21(3): 1879-1891. 2005 1880 BRIT.ORG/SIDA 21(3) INTRODUCTION Most orchids begin life by forming a mycorrhizal relationship, as seed germi- nation is dependent on a mycorrhizal association to supply the seedling with carbon during its early stages; a relationship known as myco-heterotrophy (Dressler 1981; Leake 1994; Smith @ Read 1997). Ultimately, approximately 80% of orchids switch from the myco-heterotrophic lifestyle to one in which car- bon exchange occurs in the opposite direction, from orchid to fungus (Atwood 1986). Only 20% of orchid species maintain this symbiosis Uyeusnou: their lifetime, which can evolve toa high degree of mycorrhizal specialization (Ras- mussen 1995; Taylor et al. 2002). Within the recognized orchid subfamilies, the appearance of myco-heterotrophic species is nearly ubiquitous, and these kinds of orchids can be found within all tribes of the Orchidaceae (Dressler & Dodson 1960; Chase et al. 2003). Although recent work has sought to understand the nature of the mycorrhizal associations for orchids and how they relate to or- chid taxonomy (Zelmer et al. 1996), less is known about how myco-heterotro- phy is related to geographic distribution. It is thought that a high degree of speci- licity between orchid and fungus may have broader conservation implications, as protection of endangered myco-heterotrophic forms requires both the main- tenance of the orchid itself as well as its associated fungus (Taylor et al. 2003). Because these orchids have a relatively low ability to withstand transplanta- tion from the wild (Liggio 1999), determining the specific features found in the habitat of myco-heterotrophic orchids can provide a key to understanding their geographic distribution, and ultimately aid their conservation worldwide. Corallorrhiza Gagnebin and Hexalectris Rafinesque are the only two gen- era of myco-heterotrophic orchid that occur in Texas. Members of both genera are commonly called “coral root” orchids, due to the presence of anthocyanin in the rhizome, stalk, and flowers (Liggio 1999), although the ce differ in their broader appearance, habitat, and distribution. Corallorrhiza includes ten species, of which nine are native to North and Central America (Freudenstein 1997). The genus Corallorrhiza is found within all the lower 48 states and Alaska. Hexalectris is found in a much narrower range, with a center of diver- sity in northern Mexico (Luer 1975). As a result, only five of the Hexalectris species occur in the United States, and of these species four are limited to parts of Texas (H. warnockii Ames & Correll, H. revoluta Correll, H. nitida L.O. Will- iams, and H. grandiflora (A. Richard & Galeotti) L.O. Williams), Arizona (H. warnockii), and New Mexico (H. nitida) (Fig. D. Only Hexalectris spicata (Walter) Barnhart ranges widely, occurring along the eastern seaboard as far north as Maryland and West Virginia. The range of two Hexalectris species (H. grandifloraand H. revoluta)is restricted to only two counties in west Texas (Jeff Davis County for H. grandiflora, Jeff Davis and Culberson counties for H. revoluta; Liggio 1999; Hatch et al. 1990). COLLINS ET AL., SOIL AND ECOLOGICAL FEATURES OF HEXALECTRIS SITES 1881 Number of Hexalectris species per county My yay: Wyilyfy 1, Wily ly, Wie Wipe SS \ Y »s Fic. 1. Distribution map of Hexalectris in Texas (based ties, divided | ies, using inf ion from Hatch et al. 1990) RA ‘anc t¢L 1 », a spicata and H. nitida. 1 ¥ >I ither H. spicata orH. nitida.M 7 Fr In Texas, most of the counties with Hexalectris populations are within only three of the state’s 11 ecological regions: the Trans-Pecos, the Edwards Plateau, and the Blackland Prairies (Fig. 1; Hatch et al. 1990; Turner et al. 2003). Only H. spicata extends beyond these three regions, occurring in all but the High Plains and Rolling Plains of the Texas panhandle. Dallas County (in the Blackland Prairies ecological region) is only one of four counties (Gillespie in the Edwards Plateau, Jeff Davis and Brewster in the Trans-Pecos are the other three) that have three or more species of Hexalectris. Dallas County shows a high recorded diver- sity for Hexalectris, but this is in part related to recently reported range exten- sions for H. warnochii and H. nitida (Engel 1987; Mahler 1988), and the new combination H. spicata var. arizonica, which was described in part based on speci- mens from the Dallas Nature Center (now Cedar Ridge Preserve) in southwest 1882 BRIT.ORG/SIDA 21(3) Dallas County. However, as most information on Hexalectris has appeared only within the last fifty years (Liggio 1999), and relatively few herbarium collec- tions have been made for this genus, Hexalectris may perhaps be present over a very large geographic area, and thus be more common than previously thought (Goldman et al. 2002). In this study we wished to expand the information known about Hexalectris abundance and distribution by conducting a detailed census of Cedar Ridge Preserve (CRP) in southwest Dallas County. This is an ideal site for a broad study of Hexalectris due to its large area (approximately 256 hectares) and its protected status as both a Dallas County Open Space Preserve and a park within the Dallas Parks and Recreation Department. CRP is also the loca- tion of extensive historic study by several orchid hobbyists (V. Engel, D. Will- iams), long-term plant research and inventory by the Dallas Nature Center (G. Stanford, J. Varnum) and the University of Dallas (M. Brown, A. Collins), as well as the range expansions for H warnockii and H. nitida, and the discovery of H. spicata var. arizonica (Catling and Engel 1993). The goals of this study were to |) compile historic data for Hexalectris at Cedar Ridge Preserve, 2) assess the number of Hexalectrisat the preserve in 2004, 3) determine the ecological char- acteristics of Hexalectris sites, to help provide a more complete description of its habitat, and 4) provide a map of orchid locations at the preserve, to deter- mine whether there are any predictors that may be used to help identify other potential Hexalectris sites in Texas. MATERIALS AND METHODS All data were collected at Cedar Ridge Preserve, in southwest Dallas County, Texas. CRP is located in one of the few remaining undeveloped areas of the Aus- tin Chalk Escarpment, a geological region of lower Cretaceous limestone that extends northeast from Dallas to the Oklahoma border, and southwest past Waco and Austin into the Edwards Plateau (Dallas Department of Urban Plan- ning 1977)). In Dallas County the escarpment formsa series of steep slopes, with erosion of the bedrock creating a variety of diverse habitats (Kennemer 1987). CRP has been a subject of longtime plant study and monitoring by virtue of its historic role as an environmental center (Greenhills Environmental Center, Dallas Nature Center) and research site (University of Dallas, M. Brown). Asa result, we were able to use historic data as well as newly-collected data to create amore complete picture of Hexalectris occurrence at the preserve. Historical data on specific Hexalectris locations were obtained by conduct- ing a walk-through of the site (outside of the Hexalectris blooming season in November 2003) with Dale Williams, who had significant background knowl- edge of past orchid records at CRP (Williams 1986). At each site identified by Williams, GPS coordinates were recorded using a Garmin eTrex Legend. Infor- mation on dates of specific range extensions for particular species were identi- COLLINS ET AL., SOIL AND ECOLOGICAL FEATURES OF HEXALECTRIS SITES 1883 fied from published accounts by V. Engel, who had conducted surveys similar to Williams (Engel 1987) and had co-described H. spicata var. arizonica 1] years earlier (Catling & Engel 1993). Recent data were obtained through both casual sightings as well as detailed censusing. Casual sightings of Hexalectris were recorded during ongoing bo- tanical inventory of the preserve (Brown et al., in prep). During that botanical inventory of approximately 75 hectares of the preserve in 2003 and 2004 we recorded GPS coordinates for any Hexalectris observed on study transects. Each Hexalectris found during surveys was identified by species, and photographed whenever possible. On July 23 and 24, 2004 we conducted more extensive surveys to specifi- cally count and map all Hexalectris found blooming at the preserve. Survey dates corresponded to dates when Hexalectris were found on the preserve in 2003 (S. McCabe, pers. obs.). Survey areas were of two different types: 1) areas where historic data on Hexalectris available or 2) areas that were ecologi- cally similar to places where Hexalectris were found in the past. GPS coordi- nates for these sites were logged and mapped. Censusing was conducted with the help of volunteers from the Master Naturalist Program and other volun- teers with significant background knowledge of plants. For most census loca- tions we obtained data from small transects on both the right and left sides of the trail whenever possible, and each transect counted as a sampling point. Transects were 20 m long and approximately 3 m wide. In each transect we counted lL) number of Hexalectris colonies, 2) total number of Hexalectris stems, and 3) number of Hexalectris of each species. Data on plant height were re- corded for some Hexalectris if they appeared to fall outside the typical height values for the species. In each transect, general ecological data on tree species, canopy, and ground cover were obtained for all sites regardless of whether Hexalectris were counted or not. Canopy cover was measured as a percentage value, and ground cover values were estimated as percentage of deciduous leaves, juniper scales/leaves, and bare ground. Statistical analyses were conducted using Microsoft Excel 2000. During analysis, the actual value of the canopy was used, as well as canopy class (< 20%, 21-40%, 41-60%, 61-80%, >80%). Ground cover was divided into five dif- ferent groups as a ratio of juniper leaves to deciduous leaves (0/100, 25/75, 50/ 50, 75/25, and 100/0). Diversity of tree species was calculated for each transect using the Shannon Index, to account for both the diversity and evenness of tree species within the transect. Diversity was compared between sites where or- chids were present and sites where orchids were absent. We also compared the percent of trees belonging to the genus Juniperus at each sampling point and orchid presence/absence, as well as the percent of trees belonging to genus Quercus and orchid presence/absence. GPS data for all Hexalectris sites (both current and historic) were converted — 1884 BRIT.ORG/SIDA 21(3) to ArcView shape files using DNR Garmin Version 4.0.28 (Minnesota Depart- ment of Natural Resources 2001), and imported into ArcView 8.3. Arc inter- change files for soil data were obtained from the Soil Survey Geographic (SSURGO) Database, available from the Soil Survey Laboratory, National Soil Survey Center (Soil Survey Staff 2004). Details on soil series found in Dallas County were obtained from the Soil Survey of Dallas County (Coffee et al. 1980). RESULTS Historic and current Hexalectris distribution at Cedar Ridge Preserve The oldest records of Hexalectris at Cedar Ridge Preserve are those described in the paper by Engel (1987). In that paper he describes several orchids origi- nally thought to be Corallorrhiza in 1981, although their identification was not confirmed until 1986, when they reappeared and were identified as H. nitida. A second species, H. warnockii, was identified in that same year by Williams (1986). These records remained the last published account of Hexalectris at Cedar Ridge Preserve, until the description of H. spicata var. arizonica in 1991, partly based on specimens collected at the preserve (Catling @ Engel 1993). We were able to identify eight sites at Cedar Ridge Preserve where Hexalectris were historically found (Fig. 2, based on D. Williams, pers. comm.), all of which fell on two trails in the southeastern part of the preserve, within an area dominated by mixed hardwoods and the two coniferous species Juniperus virginiana (Eastern red cedar) and Juniperus ashei (ashe juniper). This matched the common description of Hexalectris habitat, variously described as conifer woods on calcareous soils (Diggs et al. 1999), oak litter and decaying juniper scales/leaves (Engel 1987), leaf mold in the shade of cedars or oaks (Luer 1975), and often upon a slight slope (Coleman et al. 2002). In 2003, the preserve’s Hexalectris were rediscovered during botanical inventories (S. McCabe, pers. obs.),and ultimately we counted a total of 39 H. warnockii that year Uk. Gempel, J. Varnum, M. Brown, pers. obs.). However, in 2003 H. spicata and H. nitida were not found/counted anywhere on the preserve. In 2004 we conducted transect sampling of 89 different locations which were either L) areas for which historic data on Hexalectris were available (N=12) or 2) areas that were ecologically similar to places where Hexalectris were found in the past (N=77) (Fig. 2). In 39 (43.8%) of the 89 sites Hexalectris was present. In seven out of eight of the sites identified by Williams Hexalectris was present, indicating a reasonable degree of accuracy in the historical data obtained out- side the Hexalectris blooming season. We counted a total of 308 stems in 141 colonies, or an average of 2.2 stems/colony. The breakdown according to spe- cies was as follows: 176 stems of H. nitida at 25 different locations (57% of all stems, 64% of all locations), 113 stems of H. warnockii at 12 different locations (37% of all stems, 31% of all locations), and 15 stems of H. spicata at two loca- tions (5% of all stems, 5% of all locations). In addition, we found four stems of en — COLLINS ET AL., SOIL AND ECOLOGICAL FEATURES OF HEXALECTRIS SITES 1885 / c aoe \ f OP STS \ \ aN x \ / \ \ RA = : / \ = \ \ \ / e. \ \ \26 \ | aie oe “: Faas Ok NS 27 om \ \ \ = eo SS 202~«O ae ae ee oo < ca y Se A2 | fs — 0 005 0.1 0.2 0.3 0.4 Legend EE es Miles + Orchids found in eres census /. Additional H nd in 2004 a Soil series ID nu * Hexalectris sites ere by D. Williams Fic 9) Pye eed oa Lu L wae: yi ae ae! J pd: dD ¢ dan f£all 16 (Brackett loam, 3—5% slopes), 26 (Eddy clay loam, 1 slopes), 27 (Eddy clay loam, 3—8% slopes), 28 (Eddy-Brackett complex, 8-20% slopes), 34 (Ferris-Heiden complex, 5—12% slopes), 44 (Houston Black day, t= -3% slope 7 (Vertel ales Pasiye: The category “Additional oe sites found in 2004” f th formal survey on July 23/24 200 Hexalectris in one colony that were atypical, in that they appeared to completely lack any anthocyanin pigment, and thus were pale yellow to light green. Will- iams had also noted these atypical individuals during his exploration of the preserve. A sample of this type was later tentatively identified as H. nitida by researchers at the Botanical Research Institute of Texas (B. Lipscomb, pers. comm.), although further examination of the sample is pending. Due to the large numbers of Hexalectris counted in 2004 and limited time, we were unable to 1886 BRIT.ORG/SIDA 21(3) identify plants of the variety H. spicata arizonica, but more detailed censusing with trained orchid observers is recommended for inclusion of this type in fu- ture censuses. Of the transects studied, only three out of the 39 (8%) had more than one species (2 sites with H. warnockii and H. nitida, | site with H. nitida and H. spicata). In the course of sampling we identified several individuals that were taller than the plant heights recorded in the literature. H. warnockii is described as ranging up to 30 cm tall (Luer 1975; Diggs et al. 1999), although more recent published accounts have them within a range of 15-40 cm tall (Coleman et al. 2002). In our study we routinely found H. warnockii within 30 to 40 cm, with the tallest of this species being 64 cm. For H. nitida, published ee heights range from 10-32 cm (Coleman et al. 2002), 15 - 30 cm (Diggs et al. 1999), and up to 30 cm (Luer 1975). Our H. nitida were frequently found to be greater than 30 cm tall, with the tallest at 44 cm. Ecological characteristics of Hexalectris sites Data on canopy cover, ground cover, and tree diversity were obtained from 89 different locations in 2004 Hexalectris censusing. To determine whether the presence/absence of orchids is affected by level of canopy cover, canopy was divided into five categories (<20%, 21-40%, 41-60%, 61-80%, and >80%), which were compared. We found that there was a significant association between canopy cover and orchid presence/absence (x? of association = 13.36, P < 0.01, df=4, Fig. 3). Fifty-four percentof the sites with orchids had canopy of between 40 and 60%, and 71% of the sites without orchids had over 60% cover. We found no significant association between the type of ground cover present and orchid presence/absence (2 of association = 5.38, df=3), although all areas on which orchids were found had a ground cover of < 50% juniper leaves. Diversity of tree species was not significantly different between sites with and without Hexalectris Independent two-tailed to.95,g5= 1.054, n-s.). Sites with- out Hexalectris had a Shannon diversity index of 1.09, compared to 1.17 for sites with Hexalectris. Overall, 59% of the trees counted in transects were Juniperus spp. (either J. virginiana or J. ashei), followed by oaks (28% of all trees counted). Sites with and without Hexalectris did not significantly differ from one another in the percent of Juniperus spp. present (y- of association = 5.43, df=3, n-s.), or in the percent of Quercus spp. present (2 of association = 11.42, df=5, ns.) Only 14% of trees were something other than oak or juniper, and included (in order from highest to lowest number of individuals counted): Cornus drummondi (rough-leaf dogwood), Ilex decidua (possumhaw), Fraxinus texensis (Texas ash), Viburnum rufidulum (rusty blackhaw viburnum), and Rhus (sumac) spp. — Soil characteristics of Hexalectris sites When soil survey maps were overlaid with maps of Hexalectris sites, we were able to show some association between soil type and Hexalectris presence (Fig. COLLINS ET AL., SOIL AND ECOLOGICAL FEATURES OF HEXALECTRIS SITES 1887 M@ Present CO Absent 30 No on 1 ho [ao] I _ on ! Number of sites = oO it <20% 21 - 40% 41-60% 61 - 80% >80% Canopy level (% cover) Fic. 3. Rel 2). Nearly all (93.7% total) of the orchids found were on the Dallas County soil series Eddy-Brackett complex (8-20% slopes). This soil series is classified as a loamy-skeletal, carbonatic, thermic, shallow typic ustorthent (within the entisols), and is often found on strong to moderately sloping hillsides, with a soil depth to approximately ll inches and a surface layer of grayish brown clay loam 4 inches thick (Coffee et al. 1980). Soils within this complex have rapid runoff, with severe erosion hazard. Unfortunately, although the data appear to point to an association between soil series and Hexalectris presence, interpretation of these data is hampered by the fact that only 16 of the 89 sites examined were something other than the Eddy-Brackett complex (8-20% slopes). To further examine the relationship between soil type and Hexalectris, following our initial two-day survey we spe- cifically searched two other areas of the preserve with this soil series, and also mapped datapoints for Hexalectris detected during ongoing botanical surveys for other projects at the preserve. Overall, nine additional H. nitida were found outside of the 89 areas that we originally surveyed (bringing the total Hexalectris count for 2004 to 317). Of these sites, seven (77.7%) were within the Eddy-Brackett complex (8-20% slopes), and the remainder were found on simi- lar soils with less slope. Hexalectris found outside of Eddy-Brackett complex (8-20% slopes) were on Eddy clay loam (1-8% slopes) (Coffee et al. 1980). 1888 BRIT.ORG/SIDA 21(3) DISCUSSION In this study we have been able to provide what is perhaps the largest known count of multiple species of Hexalectris orchids at a single research site in the United States. With the initial census from this study, we will be able to follow up our data with future censusing at the preserve and perhaps expansion of t study area into other sites with ecological and soil characteristics similar to those found in this year’s census. It is possible that the large number of Hexalectris seen this year may be a result of the late spring rains that occurred in Dallas County. In June 2004 Dallas experienced record-breaking rainfall, reaching over 10 inches of rain for the month, or over 250% above the normal June precipita- tion (Office of the Texas State Climatologist 2004). It is thought that generous rainfall in late spring is necessary for flowering of Hexalectris (Engel 1987), although currently there are no published data showing the relationship be- tween rainfall and Hexalectris abundance. However, with ongoing censusing of these orchids, we should be able to better elucidate the climatological factors — 1e that influence flowering. Based on the general ecological data collected in this study, we cannot nec- essarily provide any new information on the plant community with which Hexalectris is associated. Oak and juniper are clearly the primary genera that make up both the canopy and ground cover, providing both shelter anda source of decaying organic matter for the fungal symbiont. Yet oak and juniper alone do not necessarily make for good Hexalectris habitat. Having a relatively open canopy may also be important, as our study has shown these orchids to be al- most completely absent in oak-juniper sites with 60% canopy or greater. As this is not believed to be due toa need for sunlight for photosynthesis (these species do not have chlorophyll and are nonphotosynthetic), cover may influence other factors such as soil or air temperature. Temperature records of microclimate at key sites where Hexalectris have been found may be a useful future direction for studies of these species. The most important result derived from this study is that we were able to predict the occurrence of Hexalectris orchids based on soil maps After our ini- tial census efforts, we were able to identify areas on the soil map where a par- ticular soil complex, and consequently the orchids, should occur, and confirm their occurrence through targeted searches. Predictions based on soil type were also corroborated by information from other areas in other parts of Dallas County. Hexalectris have frequently been found in Dogwood Canyon, an area approximately 2 kilometers to the southwest of our study area (D. Hurt, pers. comm.). Soil maps reveal that most of the canyon is composed of Eddy-Brackett complex (8-20% slope), with the exception of the lowest levels of the canyon along the creekbed. In addition, long-term observations of areas less than half a kilometer west of the preserve and north of Dogwood Canyon that fall within Cedar Hill State Park show the only recorded orchids to be Spiranthes L.C. Rich. COLLINS ET AL., SOIL AND ECOLOGICAL FEATURES OF HEXALECTRIS SITES 1889 and Corallorrhiza, rather than Hexalectris (Paul Baldon, CHSP, pers. comm.). Cedar Hill State Park lies mostly on chromustert soils of the Heiden or Vertel complex, which are vertisols, rather than an entisol such as Eddy-Brackett. Fi- nally, we were advised of two small colonies of Hexalectris in east Dallas, adja- cent to Lower White Rock Creek known as the Scyene overlook (J. Flood, pers. com.) These colonies were located in city parks within an area geologically simi- lar to Austin Chalk Escarpment, and were confirmed to be on Eddy-Brackett complex (8-20% slope). This confirmation helps to solidify the connection be- tween soils and Hexalectris incidence in Dallas County. Based on the information from this study, we have planned to extend this research to other areas with similar soil types. In Dallas County, approximately 1.3% of the land area (3127 hectares) falls within the Eddy-Brackett (8-20%) soil series, and nearly all of these soils are found at 36 locations in the county. With a broader search area, we have the potential to expand the known range of Hexalectris within Dallas County, and to protect these areas from expand- ing development within the county. If the soil-orchid relationship is confirmed in Dallas County, it can potentially be applied to all of Texas. The map of Hexalectris distribution in Texas indicates that Dallas County has a high Hexalectris diversity compared to most other counties, yet this may be an arti- fact of a lack of censusing in other areas, or perhaps a limited knowledge of the precise soil and ecological characteristics that this genus requires. With the in- formation from this study, we are confident that the missing pieces of the Hexalectris distribution map can be filled, and our knowledge about this ge- nus can be expanded. ACKNOWLEDGMENTS Many thanks to Dale Williams for providing his background knowledge of the orchids at Cedar Ridge Preserve, and for his historical information about the orchid search at Cedar Ridge Preserve. The initial observations of orchids in 2003 that set in motion this study would not have been possible without the efforts of Sussann McCabe. Karen Gempel assisted in our informal census of Hexalectris in 2003. Stephanie Varnum assisted greatly in survey efforts, data compilation and manuscript editing. Survey efforts in 2004 were aided by mem- bers of North Texas Chapter, Master Naturalist programs (Marguerite Kaufman, Annie Smirmaul, Holly Toland, Dana Wilson), other plant enthusiasts Joann Cross, Shirley Lusk, and Kathy Saucier) and the University of Dallas (Heather McWilliams, John Rueda). Ken Garrison provided photography expertise and photos of these species. Richard Marsden, Winwaed Software Technology, con- structed the range map for Hexalectris used in this paper. We acknowledge the support of the Botanical Research Institute of Texas, particularly Gary Jennings, Barney L. Lipscomb, and George M. Diggs, Jr. and Austin College), Jim Flood (North Texas Chapter, Master Naturalist programs), Paul Baldon (Texas Parks 1890 BRIT.ORG/SIDA 21(3) and Wildlife, Cedar Hill State Park) and David Hurt (Dogwood Canyon) pro- vided historic data on Hexalectris in other Dallas locations. REFERENCES Atwoob, J.T. 1986. The size of the Orchidaceae and the systematic distribution of epi- phytic orchids. Selbyana 7:171-186. Carine, PM. and V.S. Encet. 1993. Systematics and distribution of Hexalectris spicata var. arizonica (Orchidaceae). Lindleyana 8:119-125. Cuase, M.W., K.M. Cameron, R.L. Barrett, and J.V.FREubeNsTEIN. 2003. DNA data and Orchidaceae systematics: a new phylogenetic classification. In: Orchid conservation. K.M. Dixon, S.P. Kell, R.L. Barrett, and PJ. Cribb, eds. Natural History Publication, Kota Kinabalu, Sabah Malaysian. Pp. 69-89 Corree, D.R.,R.H.Hitt,and D.D.Ressett. 1980. Soil survey of Dallas County, Texas. United States Department of Agriculture, Washington, DC. Dattas DeparTMeNT oF URBAN PLANNING. 1977. The escarpment report. Department of Urban Planning of the City of Dallas, Texas. Diccs, Jr, G.M., B.L. Liescome and RJ. O’Kennon 1999 Shinners & Mahler's illustrated flora of north central Texas Botanical Research Institute of Texas, Fort Worth. Dresster, R.L.1981.The orchids: Natural history and classification. Harvard University Press, Cambridge, MA. Pp. 76-79. Dresster, R.L.and C.H. Dopson. 1960. Classification and Phylogeny in the Orchidacea. Ann. Missouri Bot. Gard. 47:25-68 Encet, V. 1987. Saprophytic orchids of Dallas. Amer. Orchid Soc. Bull. 56:831—835. FREUDENSTEIN, J.V. 1997.A monograph of Corallorhiza (Orchidaceae). Harvard Pap. Bot. 1:5-51. GOLDMAN, D.H., R.A. COLEMAN, L.K.MacratH, and PM. Catune. 2002. Hexalectris. In Flora of North America, Vol. 26. Oxford University Press, New York and Oxford., Pp. 603-607. ] Hartcn, S.L., K.N.Ganoui, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Stat. Misc. Publ. 1655: 1-158. Kennemer, G.W.1987.A quantitative analysis of the vegetation on the Dallas County white rock escarpment. Sida, Bot. Misc. 1:1-10. Leake, JR. 1994, Tansley review no. 69. The biology of myco-heterotrophic (‘saprophytic’) plants. New Phytologist 127:171-216. Licaio, J. 1999. Wild orchids of Texas. University of Texas Press, Austin. Luer,C.A.1975.The native orchids of the United States and Canada,excluding Florida. The New York Botanical Garden, Bronx. Manter, WF. 1988. Shinners’ manual of the North Central Texas flora. Sida, Bot. Misc. 3: -313. Orrice oF tHe Stare Cumarovoaist. 2004. Texas Climatic Bulletin. Vol. 17. Department of Atmo- spheric Sciences, College of Geosciences, Texas A & M University, College Station. Rasmussen, H.N. 1995. Terrestrial orchids: from seed to mycotrophic plant. Cambridge Uni- versity Press, New York, NY. COLLINS ET AL., SOIL AND ECOLOGICAL FEATURES OF HEXALECTRIS SITES 1891 Smit, S.E.and DJ. Reao. 1997. Mycorrhizal symbiosis. 2nd ed. Academic Press, San Diego, California. Soi Survey Starr. 2004. National Soil Survey Characterization Data, Soil Survey Laboratory. National Soil Survey Center. USDA-NRCS, Lincoln, NE. (Thursday July 29, 2004) Taytor, DL, T.D. Bruns, J.R. Leake, and D.J. Reap. 2002. Mycorrhizal specificity and function in myco-heterotrophyic plants. In: The ecology of mycorrhizas. Vol. 157. M.G.A. van der Heijden and I.R. Sanders, eds. Springer-Verlag, Berlin, Germany. Pp. 375-414. Taytor, D.L., T. D. Bruns, T.M. Szaro, and S.A. Hopces. 2003. Divergence in mycorrhizal special- ization within Hexalectris spicata (Orchidaceae), a nonphotosynthetic desert orchid. Amer. J. Bot. 90:1168-1179. Turner, B.L, H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas Volume 2. Sida, Bot. Misc, 24:769-770. Witiams, D.1986. Native orchid species. Friends of Greenhills (a publication of the Greenhills Environmental Center, Dallas, TX, USA 5:1-2. Zeimer, C.D., L. CUTHBERTSON, and R.S. Curran. 1996. Fungi associated with terrestrial orchid mycorrhizae, seeds, and protocorms. Mycoscience 37:439-448. 1892 BRIT.ORG/SIDA 21(3) BOOK NOTICES Blackwell Publishing Joun A. MirANowskiand Coin G. SCANEs (Eds). 2005. Perspectives in World Food and Agriculture, Volume 2. (ISBN 0-8138-2021-9, hbk.). Blackwell Publishing Professional, 2121 State Avenue, Ames, [A, 50014, U.S.A. (Orders: 800-862- 6057; 1-515-292-3348; www.blackwellprofessional.com). $124.99, 334 pp., b/w figures, graphs, tables, index, 6" x 9" From the back cover—“The second cone in a series designed to keep agricultural leaders abreast of the most up-to- date int t global agriculture, Perspectives in World Food and Ag- ture, Volume 2 brings together cutting-edge agricultural research and the latest views on agri- cultural policy.” ricu ALEXANDER Krincs and RicHArb R. Branam. 2005. Guide to Tendrillate Climbers of Costa Rican Mountains. (ISBN 0-632-05249-X, hbk.). Blackwell Publishing Professional, 2121 State Avenue, Ames, IA, 50014, U.S.A. (Orders: 800-862- 6657; 1-515-292-3348; www.blackwellprofessional.com). $99.99, 182 pp., b/w line drawings, tables, index, 7" x 10" aay Krings and Braham include all native and naturalized tendrillate climbers, lianas and vines, known to occur at elevations over 700 m in the mountains of Costa Rica. This taxonomic guide includes ili 5, genera, and species. “Most family and ¢ information on distin- ishing the family or genus from other similar- ee taxa.” Synonymy, Description, chee and Distribution are provided for each taxon as well as detailed b/w line drawings. ALAIN Peeters with contributions from Crci_é VANBELLINGHEN and JOHN FRAME, 2004. Wild and Sown Grasses: Profiles of a Temperate Species Selection: Ecolo ncaa and Use. (ISBN Blackwell 0-4051-0529-1; FAO ISBN 92-5- 105150- 3, hbk.). Food and Agriculture Organization of the United Nations and Blcewell Publishing Professional, 2121 State Avenue, Ames, [A, 50014, U.S.A. (Orders: 800-862-6657; 1-515-292-3348; www.blackwellprofessional.com). $119.99, 311 pp., color photos, b/w line drawings, tables, index, 7" x 10" From the back cover—..this book contains profiles of 43 temperate species, many of which are re- gaining meee tance in the context of agricultural extensification. Amply illustrated, each profile iption of the plant, its ecological peau its agronomic onan acteristics and its uses, einenade innovative and unusual.” In addit Etymology, Common Names (English, French, German), and Distri bution. SIDA 21(3): 1892. 2005 THE ECOLOGY OF TRILLIUM TEXANUM (TRILLIACEAE) ON THE ANGELINA NATIONAL FOREST, TEXAS Michael H.MacRoberts and Barbara R. MacRoberts Bog Research, 740 Columbia Shreveport, Louisiana 71104, U.S.A. an Herbarium, Museum of Life Sciences Louisiana State University in Shreveport, Shreveport, Louisiana 71115, U.S.A. ABSTRACT Trillium texanum Buckley,a member of the Trillium pusillum Michx. complex, is rare over its range, occurring in scattered populations in baygalls, streamsides, and wooded seeps. We collected infor- mation on the habitat, phenology, population, and soils of Trillium texanum on the Angelina Na- tional Forest in southeast Texas. KEY WORDS: Trillium pusillum, Trillium texanum, baygall, Angelina National Forest, Texas. RESUMEN Trillium texanum Buckley, miembro del complejo Trillium pusillum Michx.,, es raro en todo su areal, apareciendo en poblaciones dispersas en arroyos ¢ infiltraciones boscosas. Hemos colectado informacion sobre su habitat, fenologia, poblacion, y suelos de Trillium texanum en el Angelina Na- tional Forest en el Sureste de Texas. INTRODUCTION Except for taxonomy, morphological variation, and distribution, little is pub- lished about the Trillium pusillum Michx. complex, of which Trillium texanum Buckley (or Trillium pusillum Michx. var. texanum (Buckley) J. Reveal & Broome), isa member (Kral 1983; Freeman 1994; Cabe 1995; Cabe & Werth 1995; Case & Case 1997; Farmer & Schilling 2002; Singhurst et al. 2002; Timmerman- Erskine et al. 2002a, 2002b). Only a decade ago, Freeman (1994:49) pointed out that “ecological parameters have not been measured in any population of T. pusillum.” More recently, Singhurst (1996), in hissummary of T-texanum, stated that not only is there virtually nothing known about T. texanum ecologically, there are currently no research programs that include it. Since we now know a great deal about the distribution, morphology,and taxonomy of the T pusillum complex, what would be of interest would be ecological/autecological descrip- tions of T. pusillum over its range. What is known is that all members of this complex are shade plants of moist hardwood bottoms, creek sides, or swamps. They occur on a variety of soils from sandy to cherty-flinty, fertile to infertile, alkaline to basic. They occur in the Appalachians and Interior Highlands and on the Coastal Plain (Kral 1983). SIDA 21(3): 1893-1903. 2005 1894 BRIT.ORG/SIDA 21(3) Trillium texanum is rare, occurring in small, disjunct populations (Singhurst et al. 2002). It has been found at several locations in Caddo Parish in northwest Louisiana (MacRoberts 1977; Teague & Wendt 1994) and nine east Texas counties (Nixon et al. 1977; MacRoberts & MacRoberts 1998; Singhurst et al. 2002; Turner et al. 2003). It is rated state critically imperiled in Louisiana and imperiled in Texas and globally rare/imperiled (Louisiana Natural Heri- tage Program 1999; Poole et al. 2004). Its habitat preference has been variously reported as baygalls and wooded seeps (Ajilsvgi 1979; Teague @ Wendt 1994, Singhurst et al. 2002). From 1995 to 1997, we collected information on population, associated flora, and soils for T. texanum on the Angelina National Forest in Angelina and Jas- per counties in southeast Texas. General descriptions of the area and edaphic conditions are given in Orzell (1990) and Bridges and Orzell (1989). METHODS 1. In 1995 and 1996, while conducting overall surveys for rare plants on the Na- tional Forests and Grasslands in Texas (MacRoberts & MacRoberts 1998), we surveyed for I. texanum by walking baygalls, streamsides, and branchbanks in various mesic to xeric sandy longleaf pine uplands in southern Sabine National Forest, Sabine County, and in southern Angelina National Forest in Angelina and Jasper counties. This was slow work because the plants are rare, scattered, and inconspicuous. On the Angelina National Forest, I. texanum puts up only a few early spring (March-May) flowers, and although it puts up many incon- spicuous single leaves, these are soon overtopped and hidden by later develop- ing herbaceous species, particularly ferns such as Osmunda and Woodwardia. Since our time was limited and there are many kilometers of potentially suit- able habitat along streams in the area, this survey is not exhaustive. 2. In May 1995, we established seven permanent one meter sq. plots in three I. texanum populations. Because of the limited number of known populations the sites were chosen as typical of what we had encountered and were acces- sible to repeated visits. Number of plants and their developmental stage (e.g., single leaf, three leaves but no flower, three leaves with flower, Fig. 1) were re- corded for each plot. 3. To define the plant community in which T. texanum occurs, we estab- lished two permanent plots centered on two of our one meter sq. plots (called plot 2 and plot 4 hereafter). These were chosen because they were typical and accessible. Each measured 18m x 13.5 m and was divided into three 6m X 13.5 m sections running parallel to the topographic/moisture/light gradient (Fig. 2). The upper section (highest elevation) was farthest from the stream course; the lower section was closest to the stream course. In the center of the middle section was the one meter sq. plot with I. texanum. We surveyed the flora in each plot every two to three weeks between February and November 1996. ECOLOGICAL REQUIREMENTS OF TRILLIUM TEXANUM 1895 Fic. 1, Above-g 4. We collected soil samples to a depth of 15cm in each of the center plots near the Trillium and had them processed at A & L Laboratories, Memphis, Tennessee. RESULTS 1. We found eight populations of T. texanum in Angelina and Jasper counties in the Angelina National Forest. These populations were scattered over an area about 6 km east-west along the Angelina-Jasper county line and were near the headwaters of Trout Creek, Buck Branch, Clear Creek, and Shearwood Creek. These are south-flowing, intermittent streams. The eight populations ranged from a single group of plants consisting of only a few single leaves and scapes occurring within an area less than one meter square to fairly large populations with thousands of leaves and dozens of flowering scapes scattered over a hect- are. We did not find T. texanum in the Sabine National Forest, although there appeared to be suitable baygall habitat. The dominant habitat of the whole area where T. texanum occurs on the Angelina National Forest is arenic longleaf pine uplands grading into grossarenic uplands (see Bridges & Orzell 1989; Orzell 1990; Harcombe et al. 1993; Turner et al. 1999, for habitat classifications). This area is locally know as Longleaf Ridge.Common upland vegetation of this area consists of Andropogon ternarius Michx., Croton argyranthemus Michx., Cnidoscolus texanus (Muell.- Arg.) Small, Berlandiera pumila (Michx.) Nutt., Dichanthelium aciculare (Desy. 1896 BRIT.ORG/SIDA 21(3) ™~S si ( 6m 6m~ x 6m \ ( ( = : : ) [| om \ LOWER CENTER UPPER SECTION SECTION SECTION STREAM UPsSLoOE;- ——p> Fic. 2. Study plot layout for plant communities of Trillium texanum. me ‘beimanent plots were established, ache cen- tered onaone eee pet Each Se mx 13.5 ti I I pograp Vie £. 1 £, Pa + tha | J J rr 1 ttrnth + Th + +1 taj th t 1 plot with 7. texanum ECOLOGICAL REQUIREMENTS OF TRILLIUM TEXANUM 1897 ex Poir) Gould & CA. Clark, Froelichia floridana (Nutt.) Mogq,, Ilex vomitoria Ait., Pinus palustris P. Mill. Pinus echinata P. Mill, Pityopsis graminifolia (Michx.) Nutt., Pteridium aquilinum (L.) Kuhn, Quercus incana Bartr., Q. stellata Wang., O. marilandica Muenchh., Schizachyrium scoparium (Michx.) Nash., Tephrosia virginiana (L.) Pers., Toxicodendron radicans (L.) Kuntze, Tradescantia reverchonii Bush, Tragia urens L.,and Vernonia texana (A. Gray) Small. 2. Table 1 gives the data on the developmental stage of plants in the seven one meter square plots. The vast majority of above-ground growth was single leaves, not scapes with leaves or flowers. In the seven plots, there were only 15 flowering stems in three years, and a total of only 31 scapes. This pattern was typical for all T. texanum we observed on the Angelina National Forest. In April and May, the ground cover for the seven plots was always below 30%. These figures for flowering stems are exceptionally low when compared with recent surveys by Singhurst (1996), who reported populations of T. texanum in Cass and Nacogdoches counties of 1000 to 2000 flowering scapes. 3. Table 2 lists the taxa in the different sections of Plots 2 and 4. Counting Sphagnum, plots 2 and 4 had 59 species, 50 genera, and 33 families. Plot 4 had 52 species; plot 2 had 37 species. The index of similarity (Sorenson’s) between plots 2 and 4 was 67 and between the middle sections of 2 and 4 (where T. texanum was) was 65 indicating that both plots were the same community. Sev- enty percent of these species were found in six baygalls in central Louisiana (MacRoberts et al. 2004) suggesting that these T. texanum sites may be part of the general baygall community type that is widespread over much of the West Gulf Coastal Plain (Brooks et al. 1993; Nesom et al. 1997). However, the species has not been found in central Louisiana although there are many baygalls and we have searched for it. Table 3 examines some of these data further. The upper sections of the plots were the richest, both in number of species and in number of herbaceous spe- cies. The lowest section was the least diverse, and the middle section, where T. texanum occurred, was intermediate. The lower section was dominated by woody vegetation; whereas the upper section was dominated by herbaceous species. The upper edges of baygalls in the Angelina National Forest are often essentially narrow bogs, with such characteristic bog genera as Sphagnum, Pogonia, Eriocaulon, and Xyris.Just upslope of the plots (and of most baygalls in this region) was arenic/grossarenic longleaf pine upland. The difference in vegetation among the three sections of the plots is un- doubtedly due to differences in soil moisture and sunlight. The upper sections received the most sunlight (thinnest canopy and located on the baygall edge next to arenic/grossarenic, relatively open longleaf pine uplands) and had the least saturated soil. The middle section was intermediate, and the lower sec- tion received the least sunlight (dense canopy; no open edge) and was often mucky wet. 1898 BRIT.ORG/SIDA 21(3) Tasce 1. Trillium texanum developmental stage in seven one meter sq. plots. Year single leaf three leaves flowering plant 1995 169 3 5 1996 345 3 3 1997 308 10 7 Taste 2. Vascular plants occurring in three sections of two permanent plots. Plants in plot 2 are designated "2" and plants in plot 4 are designated “4."The sections within each plot are: upper, middle, lower depending on their elevation and proximity to the stream. Nomenclature follows Kartesz and Meacham (1999). Family/species Upper Middle Lower Aceraceae Acer rubrum L. 24 24 24 Anacardiaceae Toxicodendron Vernix 24 24 24 Centella erecta (L.f.) Fern. 24 Eryngium integrifolium Walt. 4 Oxypolis rigidior (L.) Raf. 4 4 Prilit Yalitioa! ipilla yur 4 (Michx.) Raf. Aquifoliaceae llex corlacea (Pursh) Chapman 4 24 24 llex opaca Ait. 24 2 Araceae Arisaema triphyllum (L.) Schott 4 Asteraceae mogl 4 4 4 (W Valt) F H. E Robins Doellingeria sericocarpoides Small 4 4 abcd oul Hist osum peal tt 24 24 Eupatorium | dif L. 24 2 Holianth tito] 4 Liatris pycnostachya Mic Ay 4 Solidago rugos ill. 4 4 Symphyot in aun 4 4 (L.) Nesom Symphyotrichum lateriflorum 4 (L.) A.& D. Love Blechnaceae lA Ayardin or lat 1 (L.) 24 24 T,Moore Woodwardia virginiana (L.) Sm. 4 4 ECOLOGICAL REQUIREMENTS OF TRILLIUM TEXANUM 1899 Taste 2. continued. Family/species Upper Middle Lower Burmanniacea ga aphylla ne Barnh. mall 24 24 24 ec ycuekee Lobelia puberula Michx. 4 Cornaceae Nyssa biflora Walt. 2 Clusiaceae Hypericum crux-andreae 2 (L ee Hyperict alioides Lam. 4 4 Cyperaceae Carex glaucesc 4 Rhynchospora pee A. Gray 24 - 4 Scleria ee Michx. 4 4 4 Dennstaedtiace Pteridium Se (L.) Kuhn 2 Ericaceae Lyonia ligustrina (L.) DC. 4 24 Rhododendron canescens aun le chx whe W. 24 24 4 \ iu rymbosum L 2 24 2 Eriocaulaceae ae eeane? pp aecn gi are Ell. 4 24 (Walt.) Mrong. melidaceae Liquidambar styraciflua L. 2 Lauraceae Persea palustris (Raf) Sarg. 24 24 24 Lentibulariaceae Pinguicula pumila Michx. 24 Liliaceae Aletris aurea Walt. 24 Melanthium virginicum le 4 4 Loganiacea elsemium sempervirens 4 24 (L.) Aition.f. Magnoliaceae Mag! 0lia virgit var ue L. 4 4 24 Melastom Rhexia petiolata Walt. 24 Orchidaceae 24 24 Pogonia ophioglossoides (L.) Ker.Ga 1900 BRIT.ORG/SIDA 21(3) Taste 2. continued, Family/species Upper Middle Lower Osmundaceae Osmunda cinnamon L 4 24 Osmunda regalis | 4 24 2 Pinaceae Pinus palustris P. Mill. 2 2 2 Pinus taeda L. 4 4 Poaceae Chasmanthium laxum (L.) Yates 24 Di hantheli) 4 i -} t ) 24 24 (L.) Gould Polygalaceae Polygala nana (Michx.) DC. 2 Rubiaceae Mitchella repens L. 24 4 24 Smilacaceae Smilax laurifolia L. 24 4 4 Trilliaceae Trillium texanum Buckley 24 enaceae Callicarpa americana L. 4 Violaceae Viola primulifolia L. 24 4 Xyridaceae Xyris ambigua Bey. ex Kunth 4 Xyris caroliniana Walt. 2 Xyris scabrifolia Harper 4 Sphagnum 24 4 Taste 3. Floristic breakdown of plots by section. Plot 2 Upper Middle Lower Total species in sections: 27 19 13 Total % of all species: 73 51 35 Total woody species: 8 9 7 Total % woody species: 30 4/7 54 Plot 4 Total species in sections: 4] 35 Total % of all species: 79 65 35 Total woody species: 9 1] 8 Total % woody species: 22 2 44 ECOLOGICAL REQUIREMENTS OF TRILLIUM TEXANUM 1901 Taste 4. Soil characteristics of two baygalls. Exchangeable lons (ppm) Organic Sample pH P K Ca Mg Matter % Plot 2 5.0 7 21 90 20 4] Plot 4 5:2 15 22 110 25 3.3 4. Trillium texanum occurred in the Tehran-Letney-Melhomes soil series. These are sandy soils that are deep, gently sloping, poorly drained but rapidly permeable (Dolezel et al. 1988; Neitsch et al. 1982) (Table 4). In the baygalls, these soils are wet most, if not all, of the year. Like bog and baygall soils throughout the West Gulf Coastal Plain, they are acidic and nutrient poor (Nesom et al. 1997; MacRoberts & MacRoberts 2001; MacRoberts et al. 2004). DISCUSSION While our observations are local and limited, some ecological information has been gained. In the area of this study, Trillium texanum is associated with stream courses, and the typical flora of baygalls that occur below arenic/grossarenic longleaf pine uplands. These sandy uplands hold and slowly discharge water and are often associated with hillside bogs, baygalls, and seepage areas. Tril- lium texanum prefers wet but not inundated soils that are acidic and nutrient poor. It occurs under a deciduous canopy, putting up leaves and scapes in March and April before the canopy blocks light and before other herbaceous plants overtop it. In our study, most of the above-ground vegetation was single leaves with a few scapes. Why these populations should be weighed so heavily in ju- venile, non-reproductive individuals, is not known. Habitat preferences and population characteristics in other parts of its range definitely need study before this species complex will be both taxonomically and ecologically understood. ACKNOWLEDGMENTS The work was undertaken as part of a Cost-Share Agreement with the National Forests and Grasslands in Texas. Larry Brown, Rob Evans, Stanley Jones, John Logan, Tom Wendt, Joseph Wipff, Jason Singhurst, and Edwin Smith aided in various ways. Jason Singhurst, Guy Nesom, and two anonymous reviewers made many useful comments on the paper. REFERENCES Auivscl, G. 1979. Wild flowers of the Big Thicket. Texas A & M Univ. Press, College Station. Brioces, E.L. and S.L. Orzeut. 1989. Longleaf pine communities of the West Gulf Coastal Plain. Natural Areas J. 9:246-263. 1902 BRIT.ORG/SIDA 21(3) Brooks, A.R..E.S. Nixon, and J.A. Neat. 1993. Woody vegetation of wet creek bottom com- munities in eastern Texas. Castanea 58:185-263. Case, PR. 1995. The Trillium pusillum Michaux (Liliaceae) complex in Virginia. |. morpho- logical investigations. Castanea 60:1—14. Case, PR.and C.WertH .1995.The Trillium pusillum Michaux (Liliaceae) complex in Virginia. Il: isozome evidence. Castanea 60:15-29, Case, F.W.and R.B. Case. 1997. Trilliums. Timber Press, Portland, Oregon. Dotezet,R.,C. Fucus, L.Gray.T. Hout, and L. Stertoe. 1988. Soil survey of Angelina County, Texas. U.S.D.A. Soil Conservation Service, Washington, D.C FARMER, 9.B. and E.E. ScHiLLinG. 2002. Phylogenetic analysis of Trilliaceae based on morpho- logical and molecular data. Syst. Bot. 27:674-692. Freeman, J.D.1994. Status survey of the Trillium pusillum complex. Unpublished report, Ala- bama Natural Heritage, Montgomery, Alabama. Harcomer, P.A., J.S. GUTZENSTEIN, R.G. KNox, S.L. OrzeLL, and E.L. Brinces. 1993. Vegetation of the longleaf pine region of the West Gulf Coastal Plain. Proc. Tall Timber Fire Ecol. Conf. 18:83-103 Kartesz, J.T. and CA. Meacham. 1999. Synthesis of North American flora. Version 1.0. North Carolina Botanical Garden, Chapel Hill. Kral, R. 1983. A report on some rare, threatened, or endangered forest-related vascular plants of the South. Vol. 1.U.S.D.A. Forest Service, Tech. Publ. R8-TP2: 1-718. Louisiana Natura Heritact Procram. 1999. Rare plant species of Louisiana. Unpublished re- port. Baton Rouge. MacRoserts, D.T. 1977. Additions to the Louisiana flora. Sida 7:220-222. MacRoeerts, M.H. and B.R. MacRoseets. 1998. Noteworthy vascular plant collections on the Angelina and Sabine National Forests, Texas. Phytologia 84:1-27. MacRoeerts, M.H. and B.R. MacRoserts. 2001. Bog communities of the West Gulf Coastal Plain:a profile. Bog Res. Pap. Bot. Ecol. 1:1-151. MacRoeerrs, B.R., M.H.MacRoserts, and L.S. Jackson. 2004. Floristics of baygalls in central Loui- siana. Phytologia 86:1-15. Neitscu, C.L., K.L. GrieritH, N.L.McCaces, LF Maruca, and D.E. McKay. 1982. Soil survey of Jasper and Newton counties, Texas. U.S.D.A. Soil Conservation Service, Washington, D.C Nesom, G.L., B.R. MacRoserts, and M.H. MacRoserts. 1997.A new community type in south- east Texas related to baygalls. Phytologia 83:371-383. Nixon, E.S., N. Lewis, and J.D. Freeman. 1977. Trilliums in trouble. Texas Parks & Wildlife 15(1): 12-14. Orzett,S.L.1990.Texas Natural Heritage Program inventory of National Forests and National Grasslands in Texas. Unpublished report. Texas Parks and Wildlife, Austin. Poote, J.M., J.R. SincHurst, D.M. Price, and W.R. Carr. 2004. A list of the rare plants of Texas. Texas Parks and Wildlife Department, Austin. SincHursT, J.R. 1996.The status of nine endangered plants of east Texas: historical, ecologi- cal, and phytogeographical notes. M.S. Thesis, Stephen F. Austin State University, Nacogdoches. ECOLOGICAL REQUIREMENTS OF TRILLIUM TEXANUM 1903 SincHurst, J.R., E.S. Nixon, W.F, Cacbwett, and W.C. Hoimes. 2002. The genus Trillium (Liliaceae) in Texas. Castanea 67:316-323. Teacue, J.and T. Wenor. 1994. Caddo and Bossier parishes, Louisiana: Natural areas survey. Unpublished report, The Nature Conservancy, Baton Rouge. TIMMERMAN-Erskine, M., R.L. Dute, and R.S. Bop. 2002a. Morphometric analysis of the Trillium pusillum Michaux complex (Trilliaceae) of the southeast United States. Castanea 67: 109-119 TiMMERMAN-ErskINE, M., R.R. Dute, and R.S. Boyo. 2002b.The Trillium pusillum Michaux complex (Trilliaceae): Analysis of pollen and leaf epiderman micromorphology.J. Torrey Bot.Soc. 129:175-186. Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida, Bot. Misc. 24: 1-888. Turner, R.L., J.E. VAN Kiey, L.S. Smit, and R.E. Evans. 1999. Ecological classification system for the national forests and adjacent areas of the West Gulf Coastal Plain. The Nature Con- servancy, Nacogdoches. 1904 BRIT.ORG/SIDA 21(3) BOOK NOTICES Blackwell Publishing Jim L. Bowyer, RuBin SHMULsKY, and JOHN G. HaYGREEN. Drawings by KAREN LILLEY. 2003. Forest Products and Wood Science: An Introduction. Fourth Edition. (ISBN 0-8138-2054-3, hbk.). lowa State University Press, A Blackwell Pub- lishing Company, 2121 State Avenue, Ames, IA, oe U.S.A. (Orders: 800- 862-6057, 1-515-292-3348; wwwohblacl prot lcom). $69.99, 554 pp., b/w figures, graphs, tables, index, 7" x 10" as From the back cover—*Forest Products and Wood Science provides a comprehensive overview of the anatomical and physical nature of wood and the relationship of these characteristics to its use as an industrial raw material.” Missouri Botanical Garden Press Monographs in Systematic Botany JAMes S. MILLER, Mary Sur Taytor, and Erin Rempata. 2005. Ivan M. Johnston’s Studies in the Boraginaceae. (ISBN 1-930723-44-X; ISSN 0161-1542, pbk.). Mono- graphs in Systematic Botany from the Missouri Botanical Garden, Vol. 101. Missouri Botanical Garden Press, P.O. Box 299, St. Louis, MO 63166-0299, U.S.A. (Orders: Missouri Botanical Garden Press, Dept. 46, PO Box 299, St. Louis, MO 63166-0299, US.A.; 877-271-1930; http://www.mbgpress.org/). $29.95, 132 pp., 7" x 10". Ivan Johnston was the chief student of the borage family until his sees in 1960. He published 200+ new taxa in Boraginaceae along with numerous new combinations; 13 of the 16 new genera described by him were borages. Among his 107 publications was a series of : journ al articles entitled “Studies in the Boraginaceae.” He also studied various other families, though with less intensity, and con- ducted floristic studies in South America, Central America, and Mexico—including long periods in the field. Johnston began his botanical career as a high school student in southern ia ae anc con- tinued there for an A.B. (1920 and M.A, (1922), then spent the next 38 years cen t Harvard, asa student (through 1925) and Research Associate and faculty member (1931- on rom 1925 to 1931, re traveled, collected, and worked as an assistant in the Gray Herbarium. The volume by Miller, Taylor, a Rempala presents various indexes and lists intended as a “tool for further ene studies on the Boraginaceae.” There is an interesting and nicely writ- ten botanico-bio ogr aphical I sketch and a ‘A catalog of the correspondence of | * An index to the botanical names aie in IMJ’s “Studies in the Boraginaceae” * Publications of IMJ * Types of IMJ by binomial * Types of IMJ by collector —Guy Nesom, Botanical Research Institute of texas, Fort Worth, 1X, 76102-4060, U.S.A. SIDA 21(3): 1904, 2005 DICHANTHIUM (POACEAE) NEW TO ARIZONA: OPEN DOOR FOR A POTENTIALLY INVASIVE SPECIES Richard Felger Tony L. Burgess Drylands Institute Institute of Envi ronmental 5 tudies PMB 405, 2509 N. oe Tucson, Arizona 85719 Texas Christian University Ft. Worth, Texas 76129, U.S.A. Samuel Dorsi ea of Earth and Environmental Science a Uni ersity, 2336 a ea ae New York ey U.S.A. John R. Reeder Thomas R.Van Devender Her Oath ium, a Hall Arizona-Sonora Desert Museum Ol 2071 North Kinney Road Teen re izona 085721 U SA. Tucson, Arizona 85743, U.S.A. ABSTRACT Dichanthi innulatum has become locally established in southern Arizona and is a new generic record for the state. This tenacious perennial grass is a potentially invasive species of concern and we offer specimen-vouchered documentation for its entrada into Arizona. There are three, simulta- neous, and independent routes for its spread. RESUMEN Dichanthi lat | blecido local 1 Sur de Arizona y es una cita de un nuevo género para el estado. Esta graminea perenne y tenaz, es una especie potencialmente invasora y ofrecemos documentacion con especimenes testigo de su entrada en Arizona. Hay tres rutas simultaneas e independientes para su dispersion. Three species of Dichanthium, a genus native to the Old World, have been in- troduced into North America as forage grasses and are established from Texas to Florida and in northern Mexico (Barkworth 2003), including Sonora: D. annulatum (Forsskal) Stapf, D. aristatum (Poir.) CE. Hubb. (Reeder & Reeder 1998), and D.sericeum(R. Br.) A.Camus (Beetle & Johnston 1991). Ringed dichan- thium, D.annulatum, isa highly variable species with a long history as a valued forage and fodder grass widely introduced in tropical and subtropical regions (Barkworth 2003; Bor 1960; Duke 1983; Reeder & Reeder 1998). It was “intro- duced for forage” and has become well established in thornscrub and tropical deciduous forest in southern Sonora (Reeder & Reeder 1998:504; they report it from 10 localities). SIDA 21(3): 1905-1908. 2005 1906 BRIT.ORG/SIDA 21(3) Ringed dichanthium was introduced into the Savanna Biome section in- side the very large greenhouse/habitat of the Biosphere 2 near Oracle, AZ, shortly before the system was sealed in 1991 for a 2-year manned mission and then a 5-month and another 6-month mission. After the first two years people went in and out frequently. This wiry perennial is a tropical grass that is “wide- spread in India and Burma, and tropical and North Africa” (Bor 1960:133). It was selected for the Savanna Biome largely because it was not expected to sur- vive the hard freezes in desert grassland at nearly 1220 meters elevation. It was grown from seeds obtained from CSIRO in Australia. Sometime after 1994, it escaped from the habitat and became established out-of-doors (Dorsi Bur- gess 2003). After 1994 the Biosphere 2 campus was managed by Columbia Uni- versity until they abandoned the project in summer 2004. Large numbers of people went in and out of the greenhouse prior to closure in 199] and after it was reopened. In 1995, Biosphere 2 was opened to tourists who passed through the Sa- vanna area on a narrow, well-trodden trail. Tony Burgess, then a faculty mem- ber at the Biosphere 2 campus, first noted ringed dichanthium outside of the closed system in 2001. By 2003, a population of this grass had become locally established and was ee oe (Dorsi & Burgess 2003). We speculate that the readily disarticul tsor spikelet clusters with their long awns might have been carried outside inadvertently on shoes or clothing. During the first few years of partial opening people exiting the greenhouse had their shoe soles disinfected with Lysol for control of a nematode in the rainforest areas of the greenhouse but no control was done for pant legs, shoe tops nor socks. The southern Sonora populations of ringed dichanthium, the only previ- ously known ones from west of the continental divide in southwestern North America, occur in an essentially frost-free region about 620 km south of the Biosphere 2 locality. In southern Sonora and elsewhere this C-4 grass is repro- ductive with hot-season rains of summer and fall, and when it was introduced into the Biosphere 2 greenhouse it was presumed to be winter-spring dormant. The population sampled in June 2004 at Biosphere 2 had recently-disarticu- ated as well as fully-ripened spikelets, demonstrating that this species can be reproductive in late spring even at higher elevation. We also found that the plants in this population are tenaciously rooted in very hard, rocky soil and are very difficult to dig up. In March 2005, Tom Van Devender and Ana Lilia Reina collected ringed dichanthium south of San Nicolas, Sonora, in tropical deciduous forest about 150 km north of the previous collections in the Alamos area (Reeder & Reeder 1998). It apparently was a recent arrival in 2005. They also found the grass in 2002 near Querobabi, about 380 km north of the previously known Sonora records in the non-desert southern part of the state. The northern Sonora popu- lation occurs in the Sonoran Desert in an area of presumed minimal winter- — FELGER ET Al RDITAKIK ae freezing. In 2004, Tom and Ana Lilia collected an unusual grass near the Pima County Fairgrounds at the southeastern edge of greater Tucson (about 65 km south of Biosphere 2) that John Reeder identified as Dichanthium annulatum. In April 2005, it was discovered in Nogales just north of the Sonora border. Both the fairground and Nogales experience moderate freezes. Ringed dichanthium has been shown to be potentially invasive in a rather wide range of environments (e.g. Duke 1983). We predict ringed dichanthium will spread widely from the three presently known Arizona sites and at the time of this writing it is probably too late to control it effectively in Arizona except by immediate and concerted effort. The northern Sonora population is also likely to spread, even to southern Arizona only 150 km to the north. Thus this newly ar- rived non-native perennial grass has three potential and simultaneous routes of expansion in southern Arizona and could become a seriously invasive species ies specimens: Dichanthium annulatum. U.S.A. ARIZONA. Pima Co.: W side of Tucson Kart peed way, just W of Houghton Road (south of I-10), S part of greater nee rs os0! COuShi@eseLScnUP: ae 02°N, 110°47'12"W, 938 m elevation: ally abundant f 21 Sep 2004, TR. Van Devender 2004-1093 & AL. Reina G. (ARIZ, ASDM, ASU, NMC, TEX). Pinal Co.: Columbia University Biosphere 2 Gap pus, E side of access road W of Biosphere 2 Savanna Biome, 32°34.692'N, 110°50.963'W, NAD 27, grass 80 cm tall, growing near roadway in elongated patch, 25 Sep 2003, S. Dorsi 1 & T.L. Burgess (ARIZ, MO, NMCR, SD, TEX, UC, US); N of Biosphere 2 Rainforest Biome, 32°34.762'N, 110°50.966'W, NAD 27, grass 60 cm tall, growing in Dichanthium dominated patch along railing in landscaped area with a tendency to collect runoff, 25 Sep 2003, S. Dorsi 2 & ARIZ); W side of access road W of Biosphere 2 Savanna Biome, 32°34.692'N, 110°50.963' W, NAD 27, grass 50 cm tall, growing along roadway as single isolated plants, S. Dorsi 5& LL. Bur- gess, 25 Sep 2003 (BRIT); handicap ramp west of Biosphere 2 Savanna Biome near Savanna airlock, 32°34.70I'N, 110°51.034'W, NAD 27, grass 60 cm tall, Le along walkway in dispersed patches and coexisting with other grasses, 25 Sep 2003, S. Dorsi 4 & T.L. Burgess (USON); Biosphere 2 Cam- s, W of Oracle, above Canada del Oro, 70 m E of a biome of Biosphere 2 structure (also an- ne small colony about 200 m to the E), ca. 3950 ft, roadside and also adjacent semi-landscaped area, Sepa wee: rocky grassland with some uae ere | with hard, knotty bases, 6 Jun 4, RS. Felger 04-2, TL. Burgess & S. Schneider (ARIZ, ASU, RSA). Santa Cruz Co.: Fiesta Market on aes Road, ie 31°21'25"N 110°57'27"W, ee m ene common perennial on edge of pavement, 27 Apr 2005, TR. Van Devender 2005-731, A.L. Reina G. (ARIZ, ASC, CAS, MEXU, NMC, TEX, US, USON). MEXICO. Sonora: ee SeNICiile de Opodepe: 2.6 km W of peucrebee! Plains of Sonora desertscrub; 30°03'14"N LL0°03'31"W, 65 ide; 21 Aug 2001, TR. Van Devender 2001-734 & ALL. Reina G. (ARIZ Z, USON). Municip de Yécora: Road to La Quema on SON 117, 1.8 km S$ of San Nicolas junction with MEX 16; tropical deciduous forest; 28°22'11"N 109°15'42"W;, 558 m elevation; locally abundant perennial on ae 18 Mar 2005, T.R. Van Devender 2005-389 & A.L. Reina G. (ARIZ, ASU, MEXU) T.L. Bu ress REFERENCES Barkworth, M.E. 2003. Dichanthium Willemet.In: Flora of North America Editorial Commit- tee. Flora of North America North of Mexico, vol.25.Oxford University Press, New York. Pp. 637-638. Beette, A.A. and D. JoHNson. 1991. Gramineas de Sonora. Gobierno del Estado de Sonora, Secretaria de Fomento Ganadero. Hermosillo, Sonora 1908 BRIT.ORG/SIDA 21(3) Bor, N.L. 1960. The grasses of Burma, Ceylon, India and Pakistan (excluding Bambuseae). Pergamon Press, New York. oe Wert, JM.J. and J.R. Hartan. 1968. Taxonomy of Dichanthium section Dichanthium (Gramineae). Bol. Soc. Argentina Bot. 12:206-227. be Wert, JMJ.and J.R. Har.an. 1970. Apomixis, polyploidy and speciation in Dichanthium. Evolution 24:270-277. Dorsi, S. and T. Burcess. 2003. Spread of the exotic grass Dichanthium annulatum (Forssk.) Stapf at the Biosphere 2 Center, Oracle, Arizona. Unpublished manuscript, on deposit at ARIZ. Duke, J.A. 1983. Handbook of energy crops, unpublished. www.hort.purdue.edu/newcrop/ duke_energy/Dichanthium_annulatum.html (accessed Ee coay Te 2005). Reever, J.R. and C.G. Reeper. 1998. Poaceae. In: PS. Martin, D. Yetman, M. Fishbein, P. Jenkins, T.R.Van Devender, and R.K.Van Devender, eds. Gentry’s Rio Mayo Plants. University of Arizona Press, Tucson. Pp. 498-520. ot SIAPE RECORDS AND OTTER NOTE WORTTY COEEEGTIONS FOR KENTUCKY Ross C. Clark, Ronald L. Jones Raloh L. Thompson and Timothy J.Weckman Department of Biology Department of Biological Sciences Berea College a oe University Berea, Kentucky 40404, U.S.A Rich icky 40475, U.S.A rOss. a edu John W.Thieret Kentucky State Nature Department of Biological Sciences Preserves Commission Nome CGY ot Vey 801 Suen Bae Highland Hei USA Frankfort. k 601, U.S.A. Kim Feeman Department of Biological and Environmental Sciences Morehead State University Morehead, Kentucky 40351, U.S.A. Lf ABSTRACT Twenty-four species or varieties of vascular plants, all native to or naturalized in the southeastern eported a 7 or noteworthy for Kentucky. Fourteen of these are native and ten are non- native. Fifteen of the taxa are reported as state records, and the accounts for the other nine taxa pro- vide clarifications or updates on their status in the state. Taxa reported as state records are: Acanthopanax sieboldianus, Callicarpa dichotoma, Castanea sativa, Cladium mariscoides, Hyd ro- cotyle ranunculoides, Ilex conti: Ipomoea quamoclit seeaneas turbinata, pails x dicaiaie: eles Lonicera xylosteum, M C aie oleifolia, and Symphyotrichum steer Nees RESUMEN Se citan ticuat variedades de plantas vascular | lizad | Sureste de U.S.A., como nuevas o notables para Kentucky. Catorce de ellas son nativas y diez son introducidas. Quince de los taxa son citad u sy a el estado, y de otros nueve se cha aclaraciones o puestas al dia de su estatus en el eds Los taxa citados stado son: Acanthopanax sieboldianus, Callicarpa dichotoma, Se sativa, ae marisc se Hydrocotyle ranunculoides, Ilex cornuta, nea oclit, Ipomoea turbinata, Lonicera X minuti- flora, Lonicera xylosteum, Magnolia grandiflora, Quercus texana, Ribes americanum, Salix cinerea subsp. oleifolia, y Symphyotrichum eee INTRODUCTION Recent field and herbarium work has resulted in the discovery of 24 species of native and naturalized flowering plants new or noteworthy for Kentucky. These SIDA 21(3): 1909-1916. 2005 1910 BRIT.ORG/SIDA 21(3) findings update occurrences and distributions as reported by Beal and Thieret (1986), Browne and Athey (1992), Medley (1993), and Jones (2005). Kentucky rarity status—Special Concern, Threatened, Endangered, or Historical—is based on lists published by Kentucky State Nature Preserves Commission (KSNPC 2000, 2001). Nomenclature, as well as abbreviations for physiographic regions (AP—Appalachian Plateaus, IP—Interior Low Plateaus, ME—Mississippi Embayment), are based on Jones (2005), and herbarium abbreviations follow Holmgren et al. (1990). State distributions in the United States are based on USDA, NRCS (2004). Many of these records have resulted from an on- gone statewide survey of woody plants by R.C. Clark, and also from analyses by R. Clark and R.L. Jones of recent additions to EKY of major sets of collections ae E.T. Browne, H.R. Athey, and M.E. Wharton. Additional records have resulted from recent field work by staff of the KSNPC and other authors listed above. TAXA NEW OR NOTEWORTHY FOR KENTUCKY Acanthopanax sieboldianus Makino [Eleutherococcus pentaphyllus (Siebold & Zucc.) Nakai] (Araliaceae). Previous reports of fiveleaf aralia were based on speci- mens persistent after cultivation (Medley 1993). This species has been docu- mented as an escape in Utah, Ohio, West Virginia, Pennsylvania, and in a few states in New England. It was included in Jones (2005) only asa brief note un- der the family account. The following records from the n. IP of Kentucky are from plants escaped from cultivation. — Voucher specimens: Grant Co.: roadside thicket near Zion Station, 7 Jun 1994, Thieret & Buddell 57455 (KNK). Jefferson Co.: woodland edge, Jefferson Hill Rd, ca. 2 mi SW of jet Key’s Ferry Road, | Jun 1994, Medley 20021-94 (KNK). Acer floridanum (Chapm.) Pax (Aceraceae). There has been disagreement on the existence of the Florida maple (or southern sugar maple) in Kentucky. It was accepted by Browne and Athey (1992), rejected by Guetig and Jones (1991) and Medley (1993), but included as A. barbatum Michx. by Jones (2005). Ward (2004) concluded that the Michaux name, A. barbatum, was originally linked tospecimens of A. saccharum, and therefore cannot be used for the Florida maple. The taxon is known from all contiguous states except Indiana, Ohio, and West Virginia. Most of the following records are from the ME of Kentucky. Voucher specimens: Butler Co.: woods along KY 105, 3.8 mi W of Grayson Co. line, 8 Jul 1963, Browne & Browne 7556 (kY). Calloway Co.: Hancock Biological Station, Murray State University, mesic up- land oak-hickory forest, 14 Jun 1998, Thompson 98-162 (BEREA). Carlisle Co.: | age road off KY 62 to Beech Grove Church Rd, 13 May 1971, Browne & Browne 71EI3.1(EKY, MDKY lower slopes of loess bluffs, 8 Oct 1986, Campbell s.n (KY). Graves Co.: KY ~ 1.5 mi W a jet ] S.45, deciduous woods, 27 Jun 1973, Browne et al. 73D1.1(EKY). Hickman Co.: oak-hickory woods, base of loess blutts, 21 Jul 1995, Weckman & Rozeman 1758 (EKY ). Marshall Co.: woods along KY 80, 1.8 mi F Graves Co. line, 12 May 1963, Browne & Browne 7046 (KY). Owen Co.: upland woods near Kentucky River off KY 355, 2 Jun 1996, Weckman etal. 2815 (EKY). CLARK ET AL., 1911 Callicarpa dichotoma (Lout.) K. Koch (Verbenaceae). The purple beautybush, a native of China and Japan, was previously known as an escape only in North and South Carolina, Tennessee, and Virginia. This report from the n. IP is the first for Kentucky. Voucher specimen: Madison Co.: adventive at woodland edge in Hilltop Acres Subdivision, ca. 6 mi W of Richmond, 4 Oct 2004, Clark 25570 (EKY). Castanea sativa Mill. (Fagaceae). Spanish chestnut, native in western Asia, has previously been documented for Alabama, Pennsylvania, and several New England states. It has not been reported in previous literature as naturalized in Kentucky. The rene specimens from the AP and IP were determined by R.C. Clark. Voucher Fl Co.: swamp forest near Plummer’s Landing, 15 Sep 1974, Meijers.n.(KY). McCreary ioe near Whitley City, Summer, 1989, Campbell s.n. (KY). Cladium mariscoides (Muhl.) Torr. (Cyperaceae). Smooth sawgrass is known from all surrounding states except West Virginia and Missouri. It has been re- ported from Kentucky (see Beal & Thieret 1986) but no previous vouchers are known, and it was not included in Jones (2005). This recent collection from the ne. IP now confirms the presence of this sedge in Kentucky. Voucher specimen: Bath Co.: shallow drainage ditch of wooded wetlands, Hog Hollow drainage of Licking River just SW of Cave Run Dam, elevation 820 ft, 23 Nov 2004, Feeman s.n. (MDKY). Cornus sericea L.[C. stolonifera Michx.](Cornaceae). Red-osier dogwood is known from all contiguous states (except Missouri) to the east, north, and west of Ken- tucky. The species was accepted for Kentucky by Browne and Athey (1992), but rejected by Medley (1993), and considered a species of questionable documen- tation by Jones (2005). The following specimens determined by R.C. Clark docu- ment the presence of red-osier dogwood in the AP and IP of Kentucky. Voucher specimens: Henry Co.: no locality, 23 Aug 1910, Garman s.n. (KY). Wolfe Co.: oak-pine woods, along trail to Sky Bridge; single clone of 4 stems, 4 Oct 2004, Clark 25569 (ERY). Drosera intermedia Hayne (Droseraceae). Narrow-leaved sundew has been docu- mented from most of the eastern United States, but is known in Kentucky only from sf collected in the 1800s (see Beal & Thieret 1986 and Medley 1993), and considered Historical in the most recent listings by the KSNPC (2000, 2001). It was recently rediscovered in the s. IP of the state by the KSNPC, in wet, heavily disturbed woodland openings. Voucher: a digital image deposited at EKY: Russell Co.: several hundred plants observed in ruts of wet fields in bush-hogged (previously bulldozed) ings of flatwoods, plants still persistent the following summer, ca. 3 miles N of Russell Sere on a m along Berry Road, 23 Jul 2003, Hines & Drozda s.n.(EKY). Hydrocotyle ranunculoides Lf. (Apiaceae). Buttercup penny wort has been docu- mented from all contiguous states except Indiana and Missouri. It was not listed by Beal & Thieret 1986), Browne and Athey (1992), or Medley (1993), but was 1912 BRIT.ORG/SIDA 21(3) included in Jones (2005), based on the following voucher collected in the far western pane of the ME. Vouche Graves Co.: Terrapin Creek Nature Preserve, edge of large marsh, forming float- ing mats, 3 Jul 2003, White sm (ERY). Ilex cornuta Lindl. & Paxton (Aquifoliaceae). Chinese holly has previously been reported as an escape only in Alabama and North Carolina, and has not been listed in earlier publications on the Kentucky flora. This collection is from the n. IP. Voucher specimen: Madison Co.: several plants oo campus of Eastern Kentucky University, behind Keith Hall, 4 Oct 2004, Clark 25571(EKY Ipomoea quamoclit L. (Convol V ulaceae 0) Cypressvine is native to tropical America, and is known to escape in all states surrounding Kentucky except Indiana, Ohio, and West Virginia. Medley (1993) rejected this taxon from the Kentucky flora, and it was treated as a “to be expected” species in Jones (2005). This collection is from the n. IP. — Voucher | Co.: twining on Heliant! sina landfill off South Dogwood Drive and KY 21, Berea, where the species has persisted for two years, 30 Aug 2003, Thompson & FitzGerald 04-1264 (BEREA). Ipomoea turbinata Lag. (Convolvulaceae). Purple moontlower, a native of In- dia, has been reported as an escape from most southern states (North Carolina to Texas to Arkansas, except for Alabama and Tennessee). This collection from the far western portion of the ME is a considerable range extension, and is the northernmost record. It was not observed at the site in a follow-up visit in sum- mer, 2003 (MJ. McWhirter, pers. comm.). Voucher specimen: Hickman Co.: Wolf Island, in open field, a cottonwood plantation in bottomland of Mississippi River, Jul 2002, McWhirter s.n. (EKY). Determination by R.L. Jones, verified by D.F Austin, Arizona-Sonora Desert Museum Lonicera < minutiflora Zabel [Lonicera muendeniensis Rehder](Caprifoliaceae ) This taxon has a complex hybrid origin, and has previously been documented only in Illinois, Indiana, Michigan, and Wisconsin. This collection determined by R.C. Clark is from the n. IP of Kentucky. Voucher specimen: Woodford Co.: along railroad tracks, near U.S. 60 bypass at 2nd railroad crossing W of Lexington- Versailles Pike, 1] May 1962, Browne & Browne 5176 (EKY) Lonicera xylosteum L.(Caprifoliaceae). European fly honeysuckle is known from all contiguous states except West Virginia and Tennessee. It was not listed for Kentucky by Browne and Athey (1992), rejected by Medley (1993), but was in- cluded in Jones (2005) on the basis of inaccurate specimen determinations Voucher sae Laurel Co.: uplands of Rock Creek Gorge, ruderal community near white pine stand, 4 Jul 1989, Thompson & Skeese 89-1425 (EKY Magnolia grandiflora L. ieccdinece Southern magnolia occurs across the Coastal Plain from North Carolina to east Texas, north to Arkansas, Tennessee, CLARK ET AL., NEW OR NOTEWORTHY VASCULAR PLANTS FOR KENTUCKY 1913 and Virginia. It is not native to Kentucky, and both Browne & Athey (1992) and Medley (1993) rejected it as occurring outside of cultivation in the state. Speci- mens listed below document the first spontaneous establishment of Magnolia grandiflora in Kentucky. It was included in Jones (2005) as a rare adventive on the basis of these vouchers. The collections are from the ME and the n. IP A similar distribution pattern has been documented in Tennessee (Chester et al. 1997). Both sites in Kentucky were similar, remnant woodlands with poorly drained, acidic soils, adjacent to urban areas. | Vi Cracken Co age below eee ball field, Paducah Communiey College 5 Nov 1994, Weckman & Weckman L190 (EKY). Madison Co.: low remnant pin oak-red maple woodlot, between KY 595 and Rash Rd, just N of Berea, 23 Jan 1999, Weckman et al. 4527 (EKY). Nyssa biflora Walter (Nyssaceae). Swamp tupelo occurs in all contiguous states except Indiana, Ohio, and West Virginia. It was accepted by Browne and Athey (1992), rejected by Medley (1993), and treated as a questionable taxon by Jones (2005). It has now been verified by R.C. Clark for several counties in the ME and IP of Kentucky. Voucher specimens: Calloway Co.: flatwoods on Tobacco Road near Murray, L1 Jul 1996, Campbell s.n. (KY). Cumberland Co.: low ground near Cloyd’s Landing, 2 Oct 1999, Clark 25351 (EK Y). Fleming Co.: low ground along Big Run Creek, 0.3 mi N of jet KY 1013, 19 Jul 1999, Clark & Bauer 25226 (EKY). aera Co.: 1.2 mi E of Ledbetter, 2.2 mi NE of Tennessee River bridge, 24 Mar 1972, Wilson s.n. (MUR). Marshall Co.: Sledd Creek emb t, Sof KY 641, ca. 2 mi W of Kentucky Dam, shoreline, 30 May 1989, Gr Abb bs 1437 (MUR) and Rockcastle Co.: bank of Copper Creek, 15 Jun 1938, Wharton 2091 (KY). Populus balsamifera L. (Salicaceae). Balsam poplar is native to mesic woods of northeastern North America, and has been documented in all contiguous states to Kentucky except Tennessee and Missouri. There long has been confusion about whether this species occurs naturally in Kentucky. Medley (1993) was of the opinion that previous reports of the plants in Kentucky should be referred to P X jackii Sargent, a hybrid between P. balsamifera L. and P. deltoides W. Bartram ex Marshall. Some specimens have been referred to Populus x gileadensis Rouleau (Balm-of-Gilead poplar), but this taxon is known to be a pistillate clone (Hardin et al. 2001), and several of the cited specimens have staminate catkins. These collections, all verified by R.C. Clark, document the species in both the AP and IP of Kentucky. Voucher specimens: Carter Co.: KY 182, 5.9 mi W of jct U.S. 60, flood plain, 30 Jul 1965, Browne & Browne 10978(EKY ). Casey Co.: Turkey Creek Road, 6 ae 1962, Murphy & Browne 413 (KY). Lee Co.: Beartrack, 8 Sep 1974, Black 10 (KY). Letcher Co.: old strip mine area, ae mi from Virginia border, near Eolia, 4 Oct 1978, Hannan & Phillippe KEP-O1- ae 00637 (EKY). Menifee Co.: bench above Wolf pen Creek, ca. | mi N of KY 715, 6 Sep 1989, Clark & Taylor oe (EKY). Powell Co.: Anders Branch, 1 May 1993, Campbell s.n. (KY). Potamogeton amplifolius Tuck. (Potamogetonaceae). Largeleaf pondweed is known from most eastern states. Medley (1993) noted that the species was col- 1914 BRIT.ORG/SIDA 21(3) lected in Kentucky in the mid-1800s by C.W. Short, but that it was likely later extirpated. It was included in Jones (2005), based on the following voucher col- lected by the KSNPC from the s. IP. Voucher specimen: Pulaski Co.: submersed in stream over a 2 x 3 meter area, in Buck Creek near Reynold Hollow, 10 Aug 1999, White & Fields sn. KNK). Verified by J.W. Thieret. Prenanthes racemosa Michx. (Asteraceae). Purple rattlesnakeroot is known from most northeastern and northcentral states. The only record of this species in Kentucky was a historical collection from Pendleton County, and the species was considered to be extirpated in the state by Medley (1993). It was listed as “to be expected” in Jones (2005), and the following recent collection by the KSNPC from the n. AP reaffirms its presence in Kentucky. —_— Voucher specimen: Lewis Co.: siltstone/calcareous shale glade, in Crooked Creek Barrens State Na- ture Preserve, 27 Sep 2004, Hines & Evans s.n. (EKY). Quercus nigra |. (Fagaceae). There have been persistent reports of water oak in Kentucky. Browne and Athey (1992) accepted the species, based on reports from the 19' century. Medley (1993) questioned many of the records, suggesting that some were based on misidentifications or were from trees in cultivation. The species has now been firmly documented from the s. IP by the KSNPC, and other records confirm the ability of the species to naturalize in the ME of Kentucky. The native range of the species in south-central U.S. now includes Kentucky and all contiguous states except Indiana, Ohio, and West Virginia. Voucher specimens: Wayne Co.: large tree on edge of bottomland in Meadow Creek Swamp, S of KY 90, 14 Jul 2000, Hardin et al. s.n. (EK Y). Quercus texana Buckley (Fagaceae). Nuttall’s oak was not accepted as a mem- ber of Kentucky's flora by Browne and Athey (1992) or by Medley (1993). It was included in Jones (2005), based on the following voucher from the ME. It is also known from adjacent Tennessee, Missouri, and Illinois. Voucher specimen: Calloway Co.: common along trail, end of gravel road off KY 444, seeps into Blood ul 1995, Weckman & Rozeman 1778 (EKY). — River, N of New Concord, 22 Ribes americanum Mill. (Grossulariaceae). Wild black currant grows from Mon- tana to New England to Missouri, Illinois, Indiana, Ohio, West Virginia, and Virginia. It was not accepted by Browne and Athey (1992) or by Medley (1993). It was included in Jones (2005), based on the following collections from the AP and n. IP. Voucher specimens: Lee Co.: mixed woods, S-facing ravine of Walker Creek, 27 Apr 1996, Kirk & Clark 41 (EKY). Madison Co.: bottomland forest, Bluegrass Army Depot, 14 Jul 1993, Libby & Mears OB-526 (EKY). Rosa virginiana Mill. (Rosaceae). Virginia rose has been documented from all states contiguous to Kentucky except Indiana, Ohio, and West Virginia. It was accepted by Browne and Athey (1992), rejected by Medley (1993), and treated as CLARK ET AL., 1915 a taxon of unknown status by Jones (2005). The following collections from the AP and IP have now been verified for Kentucky by R.C. Clark. Voucher specimens: Harrison Co.: upland pasture and woodlots, W of Dividing Ridge Road and N of KY 32,7 Jun 1999, Clark & Bauer 24680 (EKY ). Menifee Co.: KY 77, roadside, 0.25 mi from iron bridge, 9 Jul 1969, Higgins 1618 (KY). Mercer Co.: woodland edges between Shakertown and High Bridge, 23 Jul 1955, Wharton 9217 (KY). Rowan Co.: KY 174, 0.6 mi W of Haldeman PO,, fallow land, 28 Jun 1965, Browne & Browne 10492 (EKY). Salix cinerea L. subsp. oleifolia (Sm.) Macreight (Salicaceae). Large gray willow, a native of the Mediterranean region, is known to escape in the eastern United States, and has previously been documented from North Carolina, Pennsylva- nia, New York, Massachusetts, and Maine. Earlier reports of this species in Ken- tucky were based on specimens from cultivated plants (Argus 1986), and it was not included in Jones (2005). This collection from the ME is therefore the first documentation of the species in Kentucky. Voucher specimen: Hickman Co.: N on Old Milburn Road, E on new logging road to Obion Cree slough, 23 Aug 1988, Grubbs 1226 (MUR). Determined by R.C. Clark. K, Symphyotrichum divaricatum (Nutt.) G.L. Nesom [Aster subulatus Michx. var. ligulatus Shinners] (Asteraceae). Southern annual saltmarsh aster was previ- ously known from most southern and midwestern states, including adjacent Tennessee and Missouri, but has not been reported in previous literature on the Kentucky flora. It was known from adjacent counties (Lake and Obion) in the Reelfoot Lake region of Tennessee (Chester et al. 1997). This species was discov- ered in Kentucky during an Eastern Kentucky University class trip to western Tennessee and Kentucky. The plants were first observed by two students, Amy V. McIntosh and James Storm, who noted the plants growing along a roadside just north of the upper reaches of Reelfoot Lake. This discovery from the ME came too late for inclusion in Jones (2005), where it was treated as “to be expected.” Voucher specimen: Fulton Co.: open wet fields, along KY 1282, at jct gravel road, 2.7 mi W of jet KY 311, 2 Oct 2004, Bio 525/725 class collection # 45 (EKY). ACKNOWLEDGMENTS The authors wish to thank Robert Paratley (KY) and an anonymous reviewer whose helpful comments improved the manuscript. REFERENCES Arcus, G. 1986. The genus Salix (Salicaceae) in the southeastern United States. Syst. Bot. Monogr. 9:1-170. Beat. E.O.and J.W. THierer, 1986. Aquatic and wetland plants of Kentucky. Kentucky Nature Preserves Commission, Scientific and Technical Series, Number 5, Frankfort. Browne, E.T., Jr. and R. AtHey. 1992. Vascular plants of Kentucky: an annotated checklist. University Press of Kentucky, Lexington. Cuester, E.W., B.E. Worroro, and R. Krat. 1997. Atlas of Tennessee vascular plants. Vol. 2. Misc. 1916 BRIT.ORG/SIDA 21(3) Publ. No. 13. The Center for Field Biology, Austin Peay State University, Clarksville, Tennessee. Haron, J.W., DJ. Leopoio,and FM. Wire. Harlow & Harrar’s textbook of dendrology.McGraw- Hill, New York, NY. Houmoren, P.K., NH. Houmaren, and L.C. Barnett (eds.). 1990. Index herbariorum. Part |: The herbaria of the world, 8th ed. New York Botanical Garden, Bronx, NY. Jones, R.L. 2005, Plant life of Kentucky, an illustrated guide to the vascular flora. University Press of Kentucky, Lexington. [KSNPC] Kentucky State Nature Preserves COMMISSION. 2000. Rare and extirpated biota of Ken- tucky. J. Kentucky Acad. Sci.61:115-132. [KSNPC] Kentucky Stare Nature Preserves COMMISSION. 2001. Rare and extirpated biota of Ken- tucky: 2001 update. J. Kentucky Acad. Sci. 62:145-146. Meotey, M.E. 1993. An annotated catalog of the known or reported vascular flora of Ken- tucky. Ph.D. dissertation, University of Louisville, Louisville, Kentucky. REDISCOVERY OF PONTHIEVA BRITTONIAE (ORCHIDACEAE) IN EVERGLADES NATIONAL PARK! Jimi L.Sadle, Steven W.Woodmansee, orge D.Gann Thomas V. Armentano The Institute for Regional Conservation South Florida Natural Resources Center 2601 SW 152 Ave. South Florida Ecosystem Office Miami Florida 33170, U.S.A. 950 N. Krome Ave. sadle@regionalconservation.org Homestead, Florida 33030, U.S.A. ABSTRACT The rediscovery of Ponthieva brittoniae is reported. T1 ions f E les National Park, Miami-Dade County, Florida, represent the only aens cumently ae the United States. RESUMEN ee cita _ ned ese uP de note sais oniae. Las dos poblaciones encontradas en el Parque Dade, estado de Florida, representan actualmente las Gnicas plantas Reeee de los Estados Unidos. Reports of Ponthieva brittoniae Ames in the United States have been few since its initial discovery. This species was first collected in southern Florida by bota- nists J.K. Small and JJ. Carter. Plants were collected in 1909 near Perrine, Florida, and in the Long Pine Key area of what is now Everglades National Park. Fifty- two years later, FC. Craighead Sr. made a collection in the eastern portion of Long Pine Key near Osteen Hammock. The next report of P. brittoniae was made by R.L. Hammer, who found a population of plants growing along a firebreak road in central Long Pine Key in 1979 (McCartney 1997; Gann et al. 2002). Plants persisted along the road until 1986, when the re-grading of the firebreak is thought to have destroyed them. The last recorded sighting of P. brittoniae in the United States occurred in 1987 when a single plant was observed in a solu- tion hole by Chuck McCartney northeast of the Hammer station (McCartney 1997). Since then, multiple searches of the historical locations in Everglades National Park have been carried out by Hammer, McCartney, staff of The Insti- tute for Regional Conservation (IRC) and others. The failure of these efforts to locate plants resulted in the listing of this species as “Historical” in South Florida by IRC (Gann et al. 2002). Currently, P brittoniae is listed as endangered by the state of Florida and critically imperiled by Florida Natural Areas Inventory (Chafin et al. 2000) 'Disclaimer:"The views and lusi tained in this document are those of the authors and should not be interpreted as representing the opinions or policies of the U.S. Government. Mention of trade names or commercial products does not constitute their endorsement by the U.S. Government.” SIDA 21(3): 1917-1920. 2005 1918 BRIT.ORG/SIDA 21(3) Although Ponthieva brittoniae has been reported for two other Florida counties, Brown (2002) identified these populations as the closely related P racemosa (Walter) Mohr. These two species closely resemble one another but can be distinguished on the basis of sepal and labellum shape (Sauleda & Ad- ams 1980). In addition, P. racemosa has petals with green striping while P brittoniae does not. The leaves of P racemosa are present at anthesis while the leaves of P. brittoniae are usually absent at anthesis (Luer 1972). Outside of Florida, Ponthieva brittoniae is known from the Bahamas (Ackerman 2002), the Turks and Caicos Islands ().L. Sadle, unpublished), Cuba (Correll & Correll 1982; Nir 2000) and Hispaniola J.D. Ackerman, pers. comm.). The current status of this species in the Bahamas is unknown, but it has been reported from New Providence and Andros Island. Pine rockland is still present and protected in some areas on both islands. However, the decline of other pine rockland species on New Providence has been attributed to land clearing (Ba- hamas Environment Technology and Science Commission 1999). In the Turks and Caicos, plants are known from a single, small, protected population on North Caicos Island. The status of P. brittoniae in Cuba and Hispaniola is un- certain. This species is considered to be rare and threatened throughout its range J.D. Ackerman, pers. comm.). While conducting rare plant surveys as part of the Critical Ecosystems Study Initiative (CESD in the Long Pine Key region of Everglades National Park, two populations of Ponthieva brittoniae were encountered. The first, initially seen in December 2003, was located in the vicinity of where the last plant was seen in 1987. The site was revisited several times between December and Feb- ruary and 102 plants were observed. These plants undoubtedly represent the population last seen by McCartney. The second population, discovered in Janu- ary 2004, was located approximately 2 km southwest of the original site. In this population, 139 plants were observed. Of the 241 plants, 72 were flowering. Populations were found in open pine rockland characterized by exposed limestone substrate with extensive solution holes and minimal leaf litter. The majority of the plants were growing on the vertical walls of solution holes within 0.5m of the pineland’s exposed limestone surface. A few plants were found in soil filled cracks on limestone surfaces of the limestone and a single plant was observed growing between a downed log and limestone. Prescribed fires had been carried out in both locations between 2003 and 2004. In an effort to characterize the habitat of this species, three 5m radius plots were centered on solution holes in which Ponthieva brittoniae was growing. Plots were located in both populations. A total of 73 species of flowering plants were recorded in the plots. Associated species found within the solution holes of all plots were Anemia adiantifolia (L.) Sw., Bletia purpurea (Lam.) DC., Eu- patorium leptophyllum DC, Metopium toxiferum (L.) Krug & Urb., Mikania om SADLE ET AL., ALVIOLYVVEAT VE PARK 1919 scandens (L.) Willd., Mitreola sessilifolia J.F Gmel.) G. Don, Phyllanthus pentaphyllus C. Wright ex Griseb. var. floridanus G.L.Webster, Polygala grandi- flora Walt., Pteris bahamensis(J.G. Agardh) Fée, Samolus ebracteatus Kunthand Sideroxylon salicifolium (L.) Lam. Species growing on the upland portions of all three plots were Anemia adiantifolia, Echites umbellata Jacq., Guapira dis- color (Spreng.) EL. Little Jr, Ilex cassine L., Jacquemontia curtisii Peter ex Small, Mikania scandens, Myrica cerifera L., Passiflora suberosa L., Phyllanthus pentaphyllus var. floridanus, Physalis walteri Nutt., Pteris bahamensis, Rapanea punctata (Lam.) Lundell, Sabal palmetto (Walt.) Lodd. ex J.A. & J.H. Schultes, Samolus ebracteatus, Schizachyrium gracile Spreng.) Nash and Tetrazygia bi- color (P. Mill.) Cogn. Voucher Specimen: FLORIDA. Miami-Dade Co.: Everglades National Park, Long Pine Key,S of Winkley Hammock; in recently burned, ep fire suppressed pine rockland, in solution hole 30cm from top on vertical face; collected with Everglades National Park permit #EVER-2003-SCI-0084 and Florida Department of Agriculture Consumer Services Regulated Plant Index Harvesting Per- mit #56110 Feb 2004, Sadle 396 (EV — (on ACKNOWLEDGMENTS The authors thank Chuck McCartney, J.D. Ackerman and Roger L. Hammer for providing information, comments and inspiration. Bryan Naqqi Manco, the Turks and Caicos National Trust and the United Kingdom Overseas Territories Conservation Forum provided financial and logistical support for work con- ducted in the Turks and Caicos Islands. Barbara S. Carlsward, David L. Martin and an anonymous reviewer provided useful comments on this manuscript. Fairchild Tropical Botanic Garden (FTG) and New York Botanical Garden (NY) provided information on herbarium specimens and label data. Funding for the project that led to this rediscovery was provided by the Critical Ecosystems Study Initiative of the U.S. Department of the Interior through Cooperative Agreement #H5284-03-0044, Rare Plant Monitoring and Restoration on Long Pine Key, Everglades National Park, G.D. Gann and T.V. Armentano, principal investigators. REFERENCES Ackerman, J.D.2002.Ponthieva. In:Flora of North America Editorial Committee,eds.Fl.North Amer. 26:547-549. Oxford Univ. Press, New York and Oxford. Bavamas ENVIRONMENT TECHNOLOGY AND SCigNCE Commission. 1999. The Commonwealth of the Bahamas National Biodiversity Strategy and Action Plan. Published online by the ONE OD on plogie! Diversity, United Nations Environment Programme /world/bs/bs-nbsap-01-en.pdf Brown, P.M. 2002. Wild orchids of Florida. University Press of Florida. Gainesville. CHarIN, L.G., J.C.P. Hancock, and G. Netson. 2000. Field guide to the rare plants of Florida. Florida Natural Areas Inventory. Tallahassee, FL. 1920 BRIT.ORG/SIDA 21(3) Corrett, D.S. and H.B. Corre. 1982.Flora of the Bahama Archipelago (including the Turks and Caicos Islands). A.R.G. Gantner Verlag, Vaduz. Gann, G.D., K.A. Braotey, and S.W. Wooomansee. 2002. Rare plants of South Florida: their his- tory, conservation, and restoration. Institute for Regional Conservation. Miami, FL. Luer, C.A. 1972. The native orchids of Florida. New York Botanical Garden, Bronx. McCartney, C.L. 1997. Mrs. Britton’s shadow witch, Ponthieva brittoniae. Palmetto 17:10-11. Nir, M.A. 2000. Orchidaceae Antillanae. DAG Media Publishing, Inc., NY. Saucepa, R.P and R.M. Apams. 1980. The taxonomic status of Ponthieva brittonae Ames (Orchidaceae). Bull. Torrey Bot. Club 107:544—-545. QUERCUS MONTANA (FAGACEAE), NEW TO MISSOURI Marian Smith Nancy Parker Southern Illinois University Southern Illinois University Edwardsville Illinois 62026, U.S.A. Edwardsville Illinois 62026, U.S.A. msmith@siue.edu njrparker@charter.net ABSTRACT Quercus montand (Fagaceae) is a new species for Missouri, extending the western edge of the range for the species in North America. RESUMEN Quercus montana (Fagaceae) es una nueva especie para Missouri, L de su area en Norte América. Quercus montana Willd. rock chestnut oak, iscommon in the NE US, and known tooccur in AL, CT, DE, GA, IL, IN, KY, ME, MD, MA, MI, MS, NH, NJ, NY, NC, OH, PA, RI, SC, TN, VT, VA, WV (Nixon & Muller 1997). The species also has been called Q. prinus L. in some of the North American botanical literature, but be- cause of persistent problems with the typification and application of that epi- thet, we are following Nixon and Muller (1997) in using the name Q. montana. There has been a recent proposal to reject the name Q. prinus (Whittemore & Nixon 2005). We report it from four sites in Wayne Co in southern Missouri, on land owned and managed by the US Army Corps of Engineers (USACE) sur- rounding Lake Wappapello. The lake was created in 1941 by the USACE to con- trol flooding of farmland on the St. Francois River (USACE 2002). The Wappapello Project consists of 44,000 acres of land and water; the lake varies in size from 5,200 to 23,200 acres in surface area, depending upon the season. The original vegetation of the area consisted of woodlands that were part of the eastern temperate deciduous forest (Yatskievych 1999). This is the first re- port of Q. montana in Missouri, and a range extension of ca. 50 km for the spe- cies on its western boundary. Because of the proximity of the Q. montana popu- lations in southern Illinois (Nixon & Muller 1997; see distribution map in FNA, Vol. 3, pg. 476) and the number of populations discovered in Wayne Co, MO, the current report suggests that forested areas in counties lying between the Missouri and Illinois populations should be surveyed for the species. The populations reported below were from three glades and an oak-hickory forest. In all cases, the specimens were collected from saplings (1-1.5 m) grow- ing in well-drained rocky soil. All specimens were verified by Alan Whittemore (US National Arboretum, Washington, DC). Although saplings appeared to be abundant in the areas, the parent trees were not identified. At glade 1, associ- SIDA 21(3): 1921-1922. 2005 1922 BRIT.ORG/SIDA 21(3) ated species included Acer saccharum Michx., Carya ovata (Miller) K. Koch, Platanus occidentalis L., Quercus alba L., Q. imbricaria Michx., Q. marilandica Moench., Q. rubra L., Q. Xtridentata Engelm. and Ulmus americana L; at glade 2, A. negundo L., Cercis canadensis L., Q. alba, Q. imbricaria, Q. muehlenbergii Engelim., Q. rubra, and Q.Xtridentata;, and A. saccharum, Q. marilandica, Q. muehlenbergii, Q. rubra and U. alata Michx. at glade 3. Associated species at the oak-hickory site included A. saccharum, Celtis occidentalis L., C. canadensis, Q. imbricaria, Q. marilandica, Q. stellata Wang var stellata, Q. xtridentata, Q. rubra and U. alata. a Voucher specimen: MISSOURI. Wayne Co.: go S on US67 to Hwy 34; g0 . USA Institutions S85. Outside USA numbers issued twice a yeal v a BOTANICAL RESEARCH INSTITUTE OF 1 PRINTED IN THE UNITED STATES OF AMERICA SSN 0036-1: TABLE OF CONTENTS SYSTEMATICS Two new species of Calyptranthes (Myrtaceae) from Ecuador MariA LUCIA KAWASAKI AND Bruce K. Hoist Revision of Siolmatra (Cucurbitaceae: Zanonieae) GEORGE L. ROBINSON AND RICHARD P. WUNDERLIN Revision of Fevillea (Cucurbitaceae: Zanonieae) GEORGE L. ROBINSON AND RICHARD P WUNDERLIN Bromus hallii (Poaceae), a new combination for California, U.S.A., and taxonomic notes on Bromus orcuttianus and Bromus grandis JEFFERY M. SAARELA AND Pau M. PETERSON Taxonomic review of Astranthium integrifolium (Asteraceae: Astereae) Guy L. NEsomM Studies of Neotropical Compositae—I. Novelties in Calea, Clibadium, Conyza, Llerasia, and Pluchea JOHN F Prusk1 Una nueva especie de Guarea (Meliaceae) para Costa Rica ALEXANDER RODRIGUEZ — Diospyros torresii (Ebenaceae): a new black zapote from tropical Mexico MITCHELL C. PROVANCE AND ANDREW C. SANDERS Staurochilus leytensis, a Philippine segregate of Staurochilus fasciatus (Orchidaceae: Aeridinae) Eric A. CHRISTENSON Estudios en las Apocynaceae Neotropicales X VII: una revision del género Galactophora (Apocynaceae: Apocynoideae) J. FRANCISCO MORALES Lectotypification and a new combination in Matelea (Apocynaceae: Asclepiadoideae) for an endemic Hispaniolan vine ALEXANDER KRINGS Pal hella huangii gen.and sp. nov, an Early Cretaceous flower (Angiospet mae) in Burmese amber GEORGE POINAR JR. AND KENTON L. CHAMBERS A new combination in Stenotus (Asteraceae) Cates A. Morse Pharus primuncinatus (Poacae: Pharoideae: Phareae) from Dominican am ber GEORGE POINAR JR. AND Emm Et J. JUDZIEWICZ Notes on Libertia (Iridaceae: Sisyrinchieae) in South America PETER GOLDBLATT AND MARCELA CELIS 1955 1961 1971 1997 2015 2023 2039 2045 2051 2053 2081 2087 2093 2095 2105 Aristidae eludendae: Aristida hitchcockiana (Poaceae)—a valid species? KELLY W. ALLRED A lectotype for Stachys floridana (Lamiaceae) JOHN B. NELSON AND DANIEL B. WARD Taxonomy of the Symphyotrichum (Aster) subulatum group and Symphyotrichum (Aster) tenuifolium (Asteraceae: Astereae) Guy L. Nesom Pappus variation in North American asters. |. Double, triple and quadruple pappus in Symphyotrichum and related aster genera (Asteraceae: Astereae) JOHN C. SEMPLE AND JENNIFER L.A. Hoop Dahlia sublignosa (Asteraceae): a species in its own right DayLe E. SAAR AND PAUL D. SORENSEN Correction to the type citation of Sarracenia alabamensis and vali- dation of the name Sarracenia alabamensis subsp. wherryi : ; : (Sarraceniaceae) FW. Case, JR. A case olf disputed orthography: is it Echinochloa colona; or is it Echinochloa colonum (Gramineae)? DANIEL B. WarD ICBN clarification needed: use of ranks Guy L. NESOM AND BARNEY L. LipscoMB Herbarium acronym correction for Ipomoea seaania (Convolvu- laceae) holotype: Sida 21(3). 2005 RICHARD FELGER AND DANIEL F AUSTIN CHROMOSOME NUMBERS 1 1] ~ IVIISCeCLLANCOUS C North America ROBERT J. SORENG ETHNOBOTANY hromosome number reports for Poa (Poaceae) in Relationships between plant folklore and antitumor activity: an historical review RICHARD W. Spjut Ethnobotanical report from mangroves of Pichavaram, Tamil Nadu State, India K. VENKATESAN, V. BALAKRISHNAN, K.C. RAVINDRAN, AND V. DEVANATHAN Pinguicula vulgaris (Lentibulariaceae) and its uses in Norway TORBJORN ALM 2113 2119 2125 2141 2161 2169 2171 2185 2193 2195 2205 2243 2249 BOTANICAL HISTORY Notes on Louisiana botany and botanists, 1718-1975 JosePpH A. EWAN FLORISTICS, ECOLOGY, AND CONSERVATION Additions and emendations to The Wild Orchids of North America, North of Mexico PAUL MARTIN BROWN New records of pter idophy tes for the flora of Peru ALAN R. SmitH, BLANCA LEON, HANNA TUOMISTO, HENK VAN DER WERFF, Rossin C. Moran, MARCUS LEHNERT, AND MICHAEL KESSLER The vascular flora of Giles County, Tennessee Dwayne Estes The vascular flora of Montgomery County, Arkansas Travis D. MARSICO Mikania buchtienii (Asteraceae: Eupatorieae) new to Argentina Marid M. CeRANA AND Luis ARIZA-ESPINAR Ranunculus ficaria (Ranunculaceae), new to North Carolina and an updated key to Carolina congeners ALEXANDER KRINGS, ALAN S. WEAKLEY, JOSEPH C. NEAL, AND EDWARD C. SwaB The occurrence of Cerastium pumilum (Caryophyllaceae) in Oklahoma Bruce W. HoaGLanb, AMY BUTHOD, AND GLORIA CADDELL First record of Nymphoides indica (Menyanthaceae) in Texas KEN SAUNDERS Muscari comosum (Liliaceae) new to Texas Lee M. LuckEYDOO Hydrocotyle sibthorpioides (Apiaceae) new for Texas and notes on introduced species Barney L. Lipscoms AND GEORGE M. Diacs, JR. 2275 2297 2321 2343 2389 2425 2429 2439 2441 2445 2449 Book reviews and notices 1970, 2014, 2022, 2038, 2080, 2086, 2094, 2104, 2160, 2168, 2184, 2192, 2194, 2204, 2242 2320, 2424, 2438, 2444, 2448, 2458 Announcements 2448, 2457 Sida, Contributions to Botany update—2467 Reviewers for Volume 21, 2004-2005—2469 Index to volume 21, 2004-2005 Titles of Articles With Authors—2471 Authors—2483 Botanical Names and Subject—2488 New Names and New Combinations—2501 Sida, Contributions to Botany Guidelines for Contributors—2509 Index to new names and new combinations in Sida 21(4), 2005 Astranthium ciliatum (Raf.) Nesom, comb. nov.—2016 Bromus hallii (Hitchc.) Saarela @ P.M. Peterson, comb. nov—2004 Calea mediterranea (Vell.) Pruski, comb. nov—2024 Calea triantha ( Vell.) Pruski, comb. nov—2027 Calyptranthes glandulosa M.L. Kawasaki & B. Holst, sp. nov—1955 Calyptranthes ishoaquinicca M.L. Kawasaki & B. Holst, sp. nov—1957 Clibadium arriagadae Pruski, sp. nov—2029 Conyza popayanensis (Hieron.) Pruski, comb. nov-—2032 Dahlia sublignosa (P.D. Sorensen) D.E. Saar & P.D. Sorensen, comb. & stat. nov— 2165 Diospyros torresii M.C. Provance W A.C. Sanders, sp. nov—2046 Fevillea subgenus A (Silva Manso) G. Robinson & Wunderlin, comb. et stat. nov—1993 Fevillea bahiensis G. Robinson & Wunderlin, sp. nov—1977 Galactophora angustifolia J.-F. Morales, sp. nov—2060 Guarea subsessilifolia Al. Rodr., sp. nov—2040 Llerasia macrocephala (Rusby) Pruski, comb. nov-—2033 Matelea domingensis (Alain) Krings, comb. nov—2081 Palaeoanthella Poinar G Chambers, gen. nov—2088 Palaeoanthella huangii Poinar @ Chambers, sp. nov—2088 Pharus primuncinatus Judz. & Poinar, sp. nov—2096 Pluchea baccharis (Mill.) Pruski, comb. nov.—2035 Sarracenia alabamensis FW. Case & R.B. Case, sp. nov—2169 Sarracenia alabamensis subsp. wherryi FW. Case & R.B. Case, subsp. nov—2169 Staurochilus leytensis (Ames) F.A. Christenson, comb. nov—2051 r Stenotus lanuginosus var. andersonii (Rydb.) C.A. Morse, comb, nov.—2093 TWO NEW SPECIES OF CALYPTRANTHES (MYRTACEAE) FROM ECUADOR Maria Lucia Kawasaki Bruce K. Holst Botany De Ururricilt, The Field Museum A Mar le Se Iby Botanical Gardens 1400 South Lake Shore Drive 811 South Palm Avenue hicago, Illinois 60605-2496, U.S.A. Sarasota, Florida 34236, U.S.A. av eat erence bholst@selby.org ABSTRACT Calyptranthes glandulosa and Calyptranthes ishoaquinicca, two new species from Ecuador, are de- scribed and illustrated. RESUMEN Se describen y se il d peci le M del Ecuador, Calyptranthes glandulosa y Calyptranthes ishoaquinicca. INTRODUCTION Calyptranthesisa genus of more than 100 species ranging from Mexico to north- ern Argentina (Landrum & Kawasaki 1997). It is characterized by the usually paired panicles, the closed calyx opening as a calyptra, and, in many species, dichotomous branching. About 30 species are found in Ecuador, many of them new to science (Holst 1999). Two of these, Calyptranthes glandulosa and Calyptranthes ishoaquinicca, are described and illustrated in this paper. Calyptranthes glandulosa M.L. Kawasaki & B. Holst, sp. nov. (Fig. 1). Type: ECUA- IR. ORELLANA (“NAPO” on label): Reserva Biologica Jatun Sacha, Rio Napo, 8 km al E de Misahualli, 01°04'S, 77°36'W, 450 m, 21-25 May 1987 (£D, C. Cerén 1431 (HOLOTYPE: QCNE; ISOTYPES: F, MO, NY, SEL). Arbuscula. BONA RECO oblonga, siccata supra oli btus | trinque ¢ pellucid punctata; ner | Icato. Panicul if] lal h pilosa. Bacca globosa, Small shrubs or trees 2-6 m tall, the trichomes where present yellowish, bifur- cate. Leaf blades elliptic to oblong or less commonly obovate, 16.6-28(-34) x 5.1-10.5(-20) cm, coriaceous, with conspicuous, dark, convex glands on both surfaces, discolorous when dry, the upper surface olive-green, glabrous, the lower surface brownish-green to light-brown, nearly glabrous, with a few scattered trichomes; apex abruptly acuminate, the acumen 1-2 cm long; base obtuse to cuneate; midvein sulcate above, convex below; lateral veins 20-25 pairs, con- vex on both surfaces; marginal veins 2, the innermost 3-5 mm from blade mar- gin; petiole 1.1-2 cm long, canaliculate, glabrous, drying blackish. Inflorescences paired reduced panicles with ca. 5-20 flowers per panicle, 1.5-5.5 cm long, the SIDA 21(4): 1955-1960. 2005 1956 Hl a a 2 Digitai Scans of Type Collection - Scale 10 cm Marie Selby Botanical Gardens Herbarium (SEL) Fic. 1 Lf, lypt th BRIT.ORG/SIDA 21(4) MYRTACEAE Isotype ECUADOR PO: Reserva Biologica Jatun Sac eee Napos 8& km al_F de Nisahualli. Bosque muy hiimedo Flores axilares. Frutos verdngos como un 8 de tiernos. Bosaue di rbado. N e = mayo 1987 Carlos B. ron M. 143 MISSOURI pe al peels HERBARIUM (MO) M.L. Kawasaki & B. Holst (Cerdn 7437: isotype, SEL). OF CALYT 1957 axes pubescent; flower buds ovoid, somewhat constricted in the middle, ca. 4 mm long, distinctly gland-dotted, sessile; bracteoles deciduous, not seen; calyx calyptrate, glabrescent, deciduous; petals absent; stamens ca. 90, the filaments ca. 6 mm long, the anthers ca. 0.5 mm long; style ca. 6 mm long; hypanthium prolonged 1-2 mm beyond the ovary, tomentose without; ovary 2-locular, wit 2 ovules per locule. Fruits globose, ca. 1.5-2 cm diam., crowned by a circular hypanthium scar, gland-dotted, glabrous; seeds 1-2, ca. 9 x 7 mm, the seed coat membranous; embryo myrcioid, the cotyledons leafy and folded, the radicle well developed, equaling cotyledons in length. Distribution.—Known only from Amazonian Ecuador, in lowland humid forests, at 200-450 m elevation. Among the large-leaved species of Calyptranthes in Ecuador, Calyptranthes glandulosa is readily recognized by the leaf-blades with conspicuous, dark glands on both surfaces. The flowers appear distorted in bud as though dis- eased. No specimens are known at anthesis, though pollination is evidently occuring as several specimens bear fruit. It is possible that this species has cleis- togamous flowers or perhaps a pathogen is present that is interrupting the nor- mal flowering and fruiting sequence. Additional collections examined: ECUADOR. Orellana: Payamino, Reserva Floristica “El Chuncho,” bosque primario, Estacion Experimental INIAP-Napo, 5 km al NW de Coca, 00°30'S, 77°OL'W, 250 m, 13 Dec 1987 (£1), C. Cerén & W. Palacios 3002 (MO, SEL). Yasuni Forest Reserve, 1-2 km E of Pontificia Universidad Catolica del coe Sci. Station, 00°40.853'S, i 23.697'W, 225 m, 23 Jun ie ‘ ‘r), P. T Acevedo-Rodriguez & J.A. Cedeno 7502 (SEL); Orellana, P: ional Yasuni, Car de Maxus en construccion . 32, al S del Rio Tiputini, 00° s 7 76° 29'W, 250 m, 8- 10 Feb 1004 aD, M. Aulestia 1720 (MO, SEL); Estacion Cientifica Yasuni, Tiputini River, NW of confluence with Tivacuno River; 6 km E of Km 44 on main Maxus Road, on spur road toTivacuno ee 00°38'S, 76°30'W, 200-300 m, 23 Oct 1996 (st), R. Foster, K. Romoleroux, M. Bass & G. Villa 15700 (F QCA); Estacion Cientifica Yasuni, Rio Tiputini, al NO de la confluencia con el Rio Tivacuno, E de la carretera Maxus, Km 44, desvio hacia el pozo Tivacuno, parcela de 50 ha, 00°38'S, 76°30'W, 200-300 m, 12 Jun 1995 (fr), K. Romoleroux & R. Foster 1697 (F QCA); Estacion Cientifica Yasuni, Rio Tiputini, al NO de la confluencia con el Rio Tivacuno, 6 km E de la carretera Maxus, Km 44, desvio hacia el pozo Tivacuno, pare ela de 50 ha, 00°38'S, 76°30'W, 200-300 m, 21 Nov a fl). K.R | & R. Foster 2031 (E QCA), Estacion Cientifica Yasuni, Rio Tiputini, al NO de nfluencia con el Rio Tivacuno, E de lacarretera Maxus, Km 44, desvio hacia el pozo Tivacuno, parcela de 50 ha, arbol #102694, an 77°45'W, 200-300 m, 30 Oct 1997 (fl), K. Romoleroux, G. Villa & P Asimbaya 3191 (FE QCA), Estacion Cientifica Yasuni, Rio Tiputini, al NO de la confluencia con el Rio Tivacuno, E de lacarretera Repsol-YPF, Km 7 desvio hacia el pozo Tivacuno, Laguna Herradura, 00°38'S, a 200-300 m, 15 Oct 1999 (f1), G. Villa & C. Flores 192 (F QCA). Pastaza: Pastaza Canton, Pozo petrolero “Ramirez,” km al S de la poblacion de Curaray, 01°32'S, 76°51'W, 300 m, 21-28 Feb 1990 (il. a V. Zak & S. Espinoza 4906 (MO, SEL). aun eee ishoaquinicca M.L. Kawasaki & B. Holst, sp. nov. (Fig. 2). Type: EC- SUCUMBIOS: Fundacion Sobrevivencia Cofan, Sinangoe Station, Rio Sieguyo, near ae with Rio Alto Aguarico, across from Puerto Libre, NW of Lumbaqui, foothills of the Andes, 40m tall lowland hill-forest on gentle ridgeslopes with clay soils, 00°L0'45'N, 1958 BRIT.ORG/SIDA 21(4 ) o € oc NW? 225361 «J FIELD MUSEUM OF NATURAL HISTORY Th Id Museum (F) Isotype 0 Calyptranthes ishoaquinicea ML Kawasaki & B Holst, sp d asakt & Holst, 204 2 ECUADOR 4 Sucumbios Roberto Aguinda IMs 16 Agosto 20 Nigel Pitman, Robin f FUNDACION SOUREVIVENCIA COPAN Fic. 2. Cal’ hes j A.L. Kawasaki & B. H Ist (Ag ind tal. 1345: isotype, F). KAWASAKI ,NDW SFEMILDO VE LALTE 1959 77°29'50"W, 600-800m, 16 Aug 2001 (buds, fr), R. Aguinda, N. Pitman & R. Foster 1345 (HOLO- TYPE: QCNE: ISOTYPES: F, SEL) Frutex. Folia lanceolata, supra olivacea, subtus flavo- vel pallide-viridia; nervo medio supra sulcato; apice longiuscule acuminata. Inflorescentia 3-flora; alabastra ochraceo-pilosa, apiculata. Bacca globosa, glabra Shrubs 1-2 m tall, the trichomes where present yellowish-brown, bifurcate; young stems narrowly 4-winged to 4-angled, quadrangular in cross section. Leaf blades lanceolate, 8-14.3 x 1.8-3.6 cm, chartaceous, discolorous when dry, the upper surface olive-green, glabrous, the lower surface yellowish- to brownish- green, nearly glabrous, with a few scattered trichomes especially on the midvein; apex narrowly acuminate to caudate-acuminate with a slender acumen to 3.5 cm long; base obtuse to cuneate; midvein sulcate above, convex below; lateral veins ca. 14-17 pairs, impressed to strongly impressed above, raised below; mar- ginal veins 2, the innermost 1-3 mm from blade margin; glandular dots indis- tinct on the upper surface, numerous and convex on the lower surface; petiole 1-2 mm long, canaliculate, puberulous, drying blackish. Inflorescences paired, (1-)3(-4)-flowered spikes, 2-6 cm long, the axes glabrous, nodding; bracts lan- ceolate, ca. 5 X 1mm, puberulous, deciduous; flower buds obovoid, 2-3 mm long, pubescent, apiculate, sessile; bracteoles deciduous, not seen; calyx calyptrate, glabrescent, deciduous in mature fruit; petals absent; stamens ca. 50, the fila- ments ca. 6 mm long, the anthers ca. 0.5 mm long; style ca. 10 mm long; hy- panthium prolonged ca. 1mm beyond the ovary, tomentose to strigose without, ovary 2-locular, with 2 ovules per locule. Fruits globose, 7-9 mm diam., crowned by circular hypanthium scar, orange-red to dark-purple, glabrous; seeds 1-2, ca. 6 X 5mm, the seed coat membranous; embryo myrcioid, the cotyledons leafy and folded, the radicle well developed, equaling cotyledons in lengt Distribution —Known only from Ecuador (Sucumbios and Pastaza), in low- land to montane forests, at 430-800 m elevation. Calyptranthes ishoaquinicca, well known as “ishoa quinicco” in the Cofan villages (Pitman et al. 2002), has traditionally been used in a Cofan coming-of- age ceremony for young men (12-15 years old), as a purgative to impart strength for their adult lives; they drink a concoction prepared from this plant for ca.10 days, accompanied by much vomiting. It is distinguished from all other species of the genus in Ecuador by having short, 3-flowered inflorescences and chartaceous, narrowly acuminate leaves, with impressed lateral veins. Additional collections examined: ECUADOR. Pastaza: Villano, Pandanuque, encima de colina al S del pozo petrolero Villano 2 de ARCO, 01'28'S, 77°27'W, 550 m, 30 Ago 1987 (fr), A. Alvarez, H. Vargas & E. Freire 2410 (F MO, QCNE, SEL); Canton Arajuno, Parroquia Villano, linea propuesta por ARCO para el oleoducto, Campamentos 4 a 5, Km 10 de Villano, 430 m, 0128'S, 77°31'W, 3-7 Jul 1998 (£D, E. Freire & M. Innunda 3182 (MO, SEL). Sucumbios: Alto Rio Aguarico, Rio sae upriver from Sinangue along Northern border of Reserva Cayambe-Coca, small tributary across Aguarico SE of Puerto Libre, steep forested ridgeslopes, a x 82"N, 77°2983'W, 580-700m, 13 Jul 2000 (fr), R. Aguinda, R. Foster, M. Metz & T. Theim 955(F QCN 1960 BRIT.ORG/SIDA 21(4) ACKNOWLEDGMENTS We thank Robin Foster and the herbaria cited in the manuscript for the collec- tions. Thanks to Anna Balla for preparing the digital images. Helpful sugges- tions from Fred Barrie, Robin Foster, and Les Landrum are also acknowledged. REFERENCES Hots, B. 1999. Myrtaceae. In: P.M. Jorgensen and S.Le6n-Yanez, eds. Catalogue of the vas- cular plants of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard. 75:618-622. Lanorum, L.R.and M.L. Kawasaki. 1997. The genera of Myrtaceae in Brazil: an illustrated syn- optic treatment and identification keys. Brittonia 49:508—5 36. Pimman, N., R. Foster, and R. AGuinba. 2002. Endemic plants. In Pitman, N., D.K. Moskovits, WS. Alverson, and R. Borman A., eds. Ecuador: Serranias Cofan - Bermejo, Sinangoe. Rapid Biological Inventories Report 3. Chicago, Illinois: The Field Museum. REVISION OF SIOLMATRA (CUCURBITACEAE: ZANONIEAE) George L. Robinson Richard P Wunderlin USDA APHIS, Plant Protection and Quarantine Department of Biology, SCA 110 t Everglades University of South Florida He Drive, Suite 414 Tampa, Florida 33620-5200, U.S.A. Fort nb fai Florida 33316, U.S.A. ABSTRACT A treatment of the neotropical genus Siolmatra is presented. Two species are recognized with de- scriptions, illustrations, distribution data, and a key to the species provided. RESUMEN Se presenta un tratamiento del as neon! Siolmatra. Se reconocen dos especies con des- dat cripciones, ilustraciones y se aporta una clave de identificacion de especies. The genus Siolmatra (Cucurbitaceae) was erected by Baillon (1885) to accom- modate Siolmatra brasiliensis (Cogn.) Baill. which had resided in the heterog- enous genus Alsomitra (Blume) M. Roem., now restricted to the Old World. The name Siolmatra isan anagram of Alsomitra. Subsequently, Baillon (1886) aban- doned Siolmatra, but Cogniaux (1893) resurrected it and described a second species (S. paraguayensis). Cogniaux (1916) later added one new species (S. amazonica) and two transferred from Alsomitra (S. pedatifolia (Cogn.) Cogn. and S. peruviana (Huber) Cogn.). Harms (1926) described a sixth species (S. pentaphylla), later a seventh (Harms 1933)(S. simplicifolia), and Standley (1937) an eighth (S. mexiae). Jeffrey (1962) noted the incongruent mixture of taxa in Siolmatra which had been distinguished from Fevillea on the basis of leaf type; viz., simple in Fevillea versus 3- to 5-foliolate in Siolmatra. He redefined the two genera and transferred all taxa to Fevillea except S. brasiliensis, S. pentaphylla, and S. paraguayensis. These were maintained by Jeffrey (1978) in his enumeration of the New World Cucurbitaceae. Two species of Siolmatra are recognized here. The main distinctions between Fevillea and Siolmatra are presented in the fol- lowing key. . Leaves with petiolar or laminar glands; calyx with 5 free lobes, these with glandular squamellae (except F. passiflora); petals with a medial adaxial flap-like appendage or ridge; staminate flowers with bilocular anthers; fruit globose, usually indehis- cent; seeds not winged Fevillea . Leaves lacking glands; calyx with two pairs of calyx lobes connate, the fifth one free (calyx appearing 3-lobed), glandular calycine squamellae lacking; petals lacking a medial adaxial flap-like appendage or ridge; staminate flowers with unilocular an- thers; fruit opening apically by 3 triangular valves; seeds with marginal wings Siolmatra 7 SIDA 21(4): 1961-1969, 2005 1962 BRIT.ORG/SIDA 21(4) SYSTEMATIC TREATMENT Siolmatra Baill, Bull. Mens. Soc. Linn. Paris 1:458. 1885. Tyre: Siolmatra brasiliensis (Cogn.) Baill. BASIONYM: Alsomitra brasiliensis Cogn. Dioecious tendriled vine or liana; stems slender, sulcate. Leaves pedately 3- to 5-foliolate, petiolate, the leaflets ovate to elliptic, petiolulate or rarely subsessile. Tendrils slender, slightly sulcate, bifurcate distally, coiling both above and be- low the bifurcation. Staminate inflorescences many-flowered, in axillary panicles on the upper part of the stem, the leaves often reduced upwards; flower buds globose, pedicellate; bracts minute; hypanthium pedicelloid, slender; ca- lyx 5-merous, with 2 pairs of calyx lobes connate, the fifth free (the calyx ap- pearing 3-lobed); petals 5, white, greenish white, or greenish yellow, obdeltoid or obcordate, short-clawed, the inner surface papillate; stamens 5, the filaments slender, the anthers I-loculate, the locules horizontal or vertical, the filament extension with a dorsal, glandular, hornlike projection. Pistillate inflorescences many-t lowered, in axillary racemes or panicles, the leaves often reduced on the flowering branches; flower buds conical; bracts minute; hypanthium conical; calyx 5-merous, with 2 pairs of calyx lobes, these connate, the fifth free (the calyx appearing 3-lobed); petals 5, white, greenish white, or greenish yellow, obovate, the apex emarginate, papillose on the ventral surface; ovary 3-locu- late, the styles separate, conical, the stigmas 2-lobed, the lobes strongly diver- gent. Fruit conical, 3-loculate, obscurely ribbed, shallowly and obscurely pit- ted, coriaceous, opening apically by 3 triangular valves, the perianth scars evident at the distal end below the area of dehiscence: seeds com pressed, ob- long or elliptic, with broad marginal wings, woody with chartaceous tips or wholly chartaceous. |. Leaves 3-foliolate (rarely 4- to 5-foliolate), if the lower leaflets further divided then usually havinc \gacommon oe . not articulate from the petiole; petals of the staminate flowers obcordate; fruit 6-8 cm long S. brasiliensis iF ae 5-foliolate; petiolule articulate from i es petals of tr the staminate flov ers obdeltoid; fruit 3.5-4.5(-7) cm long s. pentaphylla Siolmatra brasiliensis (Cogn.) Baill., Bull. Mens. Soc. Linn. Paris 1:458. 1885. (Fig. 1). Alsomitra brasiliensis Cogn. in Martius, FL. Bras. 6(4):115. 1878. TYPE: BRAZIL. sé Hilaire sn. (HOLOTYPE: P, n.v., photo ex P: EUS). Saint- Alsomitra haaola var. pubescens Griseb., Symb. Fl. Argent. 136. 1879. Siolmatra brasiliensis var. pubescens (Griseb.) Cogn., in Engler, Pllanzenr. 4(Heft 66):29. 1916. Type: ARGENTINA Juwuy: ao Lorenzo, 4 Nov 1873, Lorentz & Hieronymus 228 (LECTOTYPE: BR, lectotype here designated; ISOLECTOTYPES: B (destroyed), NY; photo ex B: F MO, NY, US.). Siolmatra paraguayensis Cogn., Bull. Herb. Boissier L611. 1893. Tyre: PARAGUAY. ALTO PARANA: ear Guarapi, 1880, Balansa 3184 (HOLOTYPE: G?, nv; ISOTYPES: B (destroyed), BM, BR, F-frag- ment, K; photo ex B: F MO, NY, US). Vine or liana; stem glabrous, to 5 cm in diameter, the bark scaly, light brown. Leaves 3-foliolate, the lower leaflets occasionally further divided into 2 seg- ments, but usually with a common petiolule, the leaves rarely 4- to 5-foliolate, ROBINSON AND WUNDERLIN, REVISION OF SIOLMATRA 1963 flower (Venturi 5582, Siol brasiliensis. A. Habit, staminate infl Fic. US). C. Pistillate flower (Hassler 6793, BM). D. Fruit (Vent the blade chartaceous to subcoriaceous, the margin entire or irregularly undu- late to lobed (lateral leaflets), ovate to elliptic, the base of the lateral leaflets broadly cuneate to cordate, oblique, unequal, the base of the central leaflets broadly cuneate to subcordate, occasionally oblique and unequal, the apex acuminate, 6-10(-20) cm long, 4-8(-10) cm wide, the upper surface moderately rugose-veined, minutely pustulate, but smooth or nearly so to the touch, tomentellous to glabrate on the veins, occasionally with a few scattered tri- chomes on the blade, the lower surface with scattered short glandular-capitate trichomes and straight or curved non-glandular trichomes; petiolule 0.5-2 cm 1964 BRIT.ORG/SIDA 21(4 long, the median one slightly longer than the laterals, tomentellous; petiole 4-8 (-12) cm long, canaliculate, tomentellous to glabrate; tendrils glabrous or occa- sionally sparsely strigose near the base. Staminate inflorescences many-llow- ered, in axillary panicles on the upper part of the stem, 8-30(-40) cm long, the branches with glandular-capitate and straight to curved non-glandular tri- chomes; pedicel to | mm long; bracts up to 0.5 mm long, lanceolate, strigose- tomentellous; flower buds globose, 2-3 mm in diameter; hypanthium pedi- celloid, slender, 1-2 mm long; calyx glabrous, the lobes ovate-lanceolate, ca. 2 mm long; petals white, greenish white, or greenish yellow, obcordate, ca. 3 mm long, 2-2.5 mm wide, broadly clawed in the lower 1/4, the apex retuse, the in- ner surface papillate distally; stamens ca. 2 mim long, glabrous, the filaments slender, the theca oblong, horizontal, ca. 0.4 mm long, the filament extension with a dorsal glandular projection. Pistillate inflorescences many-flowered, in axillary racemes or panicles, 8-15 cm long, the leaves often reduced on the flow- ering branches and the entire structure to 40 cm long, the indumentum as in icel and bracts as in the staminate; flower buds conical, 8-15 cm long; hypanthium conical 5-8 mm long, glabrous; calyx lobes triangular, y Cc Qo, the staminate; pec 3-4 mm long, glabrous; petals white, greenish white, or greenish yellow, obo- vate, 4-6 mm long, the margins erose, the apex emarginate, minutely papillose on ventral surface; styles conical, ca. 2mm long, the stigma branches diverging at right angles, papillate. Fruit narrowly conical, yellow or yellow-brown, 6-8 cm long, 2-3 cm wide, obscurely ribbed, the surface shallowly and obscurely pitted, the perianth scar at the distal end evident; seeds compressed, with broad marginal wings, narrowly elliptic-oblong, 4.2-5.5 cm long, 0.8-1.5 cm wide (in- cluding the wings), the central portion elliptic, 5-7 mm long, 5-6 mm wide, both ends acuminate, the surface papillate, the wings submembranaceous, smooth, with the median rib extending from the central portion to the funicular end. Distribution and ecology.—The species occurs from eastern Peru south to northwestern Argentina, east to Paraguay and eastern Brazil. It is widespread but generally uncommon, although locally abundant in western chaco forests in Depto. Tarija, Bolivia (Michael Nee, pers. comm.). It occurs in wet forests at low elevations. Siolmatra brasiliensis is usually readily distinguished from S. pentaphylla by its 3-foliolate leaves. However, three collections from Loreto, Peru (Vdsquez et al. 2827, Vasquez et al. 2829; Vasquez & Jaramillo 5532) have 3-, 4-, or 5- foliolate leaves. However, in S. pentaphylla the petiolules are articulate from the petiole, while in they are not in S. brasiliensis. — Siolmatra paraguayensis was distinguished by Cogniaux (1916) from S. brasiliensis on the basis of its more membranaceous leaves and the ovate rather than triangular sepals of the staminate flowers. Siolmatra brasiliensis var. pubescens was distinguished by the pubescent lower leaf surface and peduncles. ROBINSON AND WUNDERLIN, REVISION OF SIOLMATRA 1965 However, these distinctions are trivial and inconsistent, so the segregates are here reduced to synonymy. Additional specimens examined. PERU. Loreto: Reserva Nacional Pacaya-Samiria, 04°51'-05°12'S, 73°50'-74°40'W, 90 m, 1993, Carpio 2104 (MO), Florida, Rio Putumayo, mouth of Rio Zubineta, ca. 200 m, Mar-Apr 1931, Klug 2033 (F K, MO, NY, US); Rio Maranon basin, near mouth of the Rio San- tiago at Pongo Manserichi, ca. 77°30'W, 1924, Tessmann 4527 (G, NY); Estacion Bioldgica Callicebus Rio Nanay-Mishana, | Jan 1982, Vasquez et al. 2827 (MO), Vasquez et al. 2829 (MO); Puerto Almendras (Rio Nanay), 122 m, aa 1984, ane & Jaramillo 5532 (MO); lquitos, 120 m, 4 Apr 1930, Williams 8112 (F). Madre d o Amazonico, 15 km ENE of Puerto Maldonado: 12°35'S, 69°O5'W, 200 m 12 Dec 1989, Gentry et al. 68606 (MO); Cuzco Amazonico, across Rio Madre de Dios on road to Lago Sandoval, 12°35'S, 69°05'W, 200 m, 19 Dec 1989, Gentry et al. 68963 (MO), Las Piedras, Cusco Amazonico, 12°29'S, 69°03'W, 200 m, 9 Oct 1991, Timand & Jaramillo 2502 (MO), Las Piedras, Cusco Amazonico, near the river and Quebrada Cicha, 12°29'S, 69°03'W, 200 m, 15 Oct 1991, Timand & Jaramillo 2605 (MO); Las Piedras, Cusco Amazénico, 12°29'S, 69°03'W, 200 m, 2 Nov 1991, Timand & Jaramillo 2958 cae BRAZIL. Acre: basin of Rio Purus, right bank of Rio Iaco, Novo Olinda, between eae Santo Antonio and Igarapé Boa Esperang¢a, 10°07'S, 69°13'W, 21 Oct 1993, Daly et al. 7831(M 0 Oct 1993, Daly et al. 7981 Hae Sao Francisco, Aug 1911, Ule 9378 (G, K, US). Bahia: Beiuad ae peas km 33, 520 m, 16 Jun 1983, Coradin et al. 5727 (MO, NY); Ferreira, Nov 1912, Zehntner 4097 (M); Faixao, Nov 1912, Zehntner 5005 (M). Maranhao: 27 km S of Entroncamento, in- tersection of Hwy 6 & Hwy 222, along Hwy 6, 4°23'S, 46°14'W, 20 Mar 1983, Schatz et al. 943 (NY). Rio de Janeiro: near Rio de Janeiro, s.d., Burchell 1685 (K); Cabo Frio, s.d., Glaziou 10071 (x). BOLIVIA. El Beni: Espiritu, floodplain of Rio Yacuma, 200 m, 5 Jul 1984, Beck 5648 (NY); Cachuela Esperanza, Rio Beni, Oct 1922, Meyer 235 (NY). El Beni/Pando: junction of Rio Beni and Rio Madre de Dios, Aug 1887, Rusby 547 (NY). La Paz: Parque Nacional Madidi, near Arroyo Aguapolo and Rio Tuichi, 270 m, 16 Mar 2002, Macia et al. 6855 (NY). Santa Cruz: Santa Cruz Botanical Garden, 12 km E of Santa Cruz, 17°46'S, 63°04'W, 375 m, 9 May 1991, Gentry et al. a uy Campamento El Rufugio, 14°45'20"S, 61°01'32"W, 180 m, 29 Jun 1994, Guillén 1987 (MO); Par ional Noel Kempff Mercado, 24 km W of San José de Campamento, on way to Piso Firme, ae 61°14'34"W, 300 m, 28 Apr 1996, Guillén et al. 4240 (NY): Las Trancas, 16°32'40'S, 61°59'28"W, 500 m, 11 Nov 1994, Killeen et al. 7116 (MO); study area of “BOLFOR?” project, Las Trancas-95, 16°31'13"S, 61°50'47"W, 450 m, 12 Dec 1994, Mamani & Jardim 390 (MO, NY); Cerro San Miguel, Mar 1989, Mereles & Ramella 2784 (FCQ, G); Parque Na- cional eee steep slopes above and 1 km S of Rio Saguayo, 17°41'S, 63°44'W, 750 m, 20 Jan 1988 Nee 36027 (MO, NY); Estancia — Rafael de Ambor6, 17°36'S, 63°36'W, 420, 11 Jun 1998, Nee 49747 (NY): ee ree de Santa z, 12 km E of center of Santa Cruz, on road to Cotaco, 17°47'S, 63°04'W, 375 m, 5 Jun 1998, Nee a Bohs 49613 (MO, NY); Cerro San Miguel, 7 Mar 1989, Ramella & Mereles ae ) PARAGUAY.Canindeyu: Estacion Biologica Mbaracayu, ca. LO km E of Villa Ygatimi, trail from main road through reserve to Mirador de los Chanchitos de Monte (Mirador Bojerkue), 24°07.41'S, 55°30.57'W, ca. 200 m, 24 Nov 2003, Bohs & Nee 3184 (MO, NY); Mbaracayt Reserve, around eee 23°59'30"S, 55°28'44"W, 27 May 1999, Zardini & Chaparro 50833 (NY). Chaco: eae Biosphere Reserve “Gran Chaco Americano”, Agua Dulce, 19°59'04'S, 59°45'28"W, 170 m, 8 Feb 2 ee & Apestegui 58247 (NY). Concepcion: Estancia Primavera-Vallemi, 22°24'07"S, 57°37’ ee at 150 m, 3 Nov 2001, Zardini & Guerrero 57291 (NY). Guaira: Cordillera de Ybytyruza, W of Cerro Peré, 2 km E of Destacamento Tororo, 12 Nov 1988, Zardini 8044 (MO); Rio Yhaca, 10 km N of Tebicuary, 16 Nov ra raileas, ae ee ee NY, USF). La Cordillera: Cordillera de Al- tos, Dec 1902, Fiebrig rade Altos, Dec 1898, Hassler 3635 (G, NY); near Lago Ypacarai, Nov 1913, Hassler 12370 (BM, G, MO, oo Dec 1913, oe 12370a (BM, G, K, MO, NY). San Pedro: Primavera, 15 Nov 1959, Woolston 1149 (K, NY, US). Paraguari: Rio Yaca Gallen near Hes Dec 1900, Hassler 6793 (BM, BR, G, NY). ARGENTINA. Jujuy: s.d., Lorentz s.n. (BR, K); Campame aimancito de YPE 7 Dec 1986, Zuloaga et al. 2525 (MO). Salta: Rio Seco, 340 m, 2 Apr 1945, ne pe (NY); 30 1966 BRIT.ORG/SIDA 21(4) km from Colonia San Andrés on road to Oran, Morrone et al. 3998 (MO), Senda Hachada, 9 Dec 1979, Schinini 19560 (kk); Finca San Andrés, La Marona, bank of Rio San Andrés, ca. 23°04'23"S, 64°45':07'W, 800 m, 30 Oct 1997, Schinini et al. 33083 (NY); Abra Grande, 12 Nov 1927, Venturi 5582 (US). Siolmatra pentaphylla Harms, Notizbl. Bot. Gart. Berlin-Dahlem 9:989. 1926. (Fig 2). Type: PERU. LORETO: upper Rio Maranon, mouth of Rio Santiago at Pongo Manserichi, ca. 77°30'W, 160 m, 18 Nov 1924, Tessmann 4575 (LECTOTYPE: G, here designated to replaced de- stroyed B holotype; photo ex B: F MO, NY, US; ISOLECTOTYPES: NY, Bassler Herb. [Peru], n.v. photograph of isotype in Bassler Herb. US). Vine or liana; stem glabrous or rarely sparsely pilose when young, soon gla- brescent. Leaves 5-foliolate, the petiolules articulate from the petiole, the blade chartaceous to subcoriaceous, the margin entire, narrowly ovate to elliptic, the base cuneate to rounded, often unequal, the apex acuminate, (5-)7-11 cm long, 3-5.5 cm wide, the upper surface moderately rugose-veined, minutely pustu- late, but smooth to the touch, occasionally with a few short trichomes on the midvein, otherwise glabrous, the lower surface glabrate or occasionally pubes- cent and sparsely short glandular-capitate; petiolules 0.5-1.5 cm long, rarely subsessile, the median one slightly longer than the lateral, pilose and sparsely short glandular-capitate or glabrous; petiole 5-10 cm long, caniculate, glabrate; tendrils glabrous. Staminate inflorescences many-flowered, in axillary panicles on the upper part of the stem, 20-50 cm long, the leaves often reduced upward and the entire floriferous part of the stem to 2 m long, the branches sparsely pilose and short stipitate-glandular or glabrous; pedicel up to 1 mm long; bracts up to 2.5mm long, linear, with scattered stipitate-glandular and non-glandular trichomes; flower buds globose, 2-3 mm in diameter; hypanthium pedicelloid, slender, 1-3 mm long; calyx glabrous, the lobes ovate-lanceolate, 2-3 mm long; petals white, greenish white, or greenish yellow, 2-3 mm long, ca. 5 mm wide, narrowly obdeltoid, clawed on the lower 1/4, the apex retuse, the inner surface papillate distally and at the base; stamens ca. 1.5 mm long, free or variously connate, the filament ca. 1mm long, the anther vertical, the filament extension with a dorsal triangular, glandular projection. Pistillate inflorescences many- flowered, axillary racemes or panicles, 10-15 cm long, the leaves often reduced on the {lowering branches and the entire structure to 40 cm long, the indumen- tum as in the staminate; pedicels and bracts as in the staminate; flower buds conical, 7-10 mm long; hypanthium conical, 5-7 mm long, glabrous; calyx lobes triangular, ca. 3 mm long, glabrous; petals white, greenish white, or greenish yellow, obovate, +-6 mm long, the margins erose, the apex emarginate, papillose on the ventral surface; styles conical, ca. 2 mm long, the stigma branches di- verging at right angles, papillate, the staminodes 5, ca. 0.5 mm long. Fruit short- conical, yellowish brown (rarely reddish brown), 3.5-4.5(-7) cm long, 2-3 cm wide, obscurely ribbed, the surface shallowly and obscurely pitted, the perianth scars at the distal end evident; seeds compressed, with broad marginal wings, oblong, 3-3.5 cm long, I-13 cm wide (including wings), the central portion — ROBINSON AND WUNDERLIN, REVISION OF SIOLMATRA 1967 Fic. Sif f | hyll A Hahit ctaminate inf] (P tal 8039, NY) B.S i fl (Gentry Revilla 20798, USF). C. Pistillate fl (Willi 2453,F).D. Fruit (Mori et al. 9134, NY).E. Seed (Mori et al. 9734, NY) ° a - ¢ Pe 13-17 mm long, 5-6 mm wide, both smooth, woody, the distal 3-6 mm submembranaceous and yellow-colored. Distribution and ecology.—The species occurs in the Amazonian basin from southeastern Colombia and northern Peru, south to northern Bolivia, and east to Para in Brazil and Guyana. Although apparently widespread in the Amazonian the surface smooth, the wings 1968 BRIT.ORG/SIDA 21(4) basin, the species is not commonly collected. It occurs in wet forests along riverbanks and flooded lake shores at 1OO-300 m. The staminal condition is surprisingly highly variable, more so than that of the other species. The stamens may be entirely free, the filaments connate at base into two groups (2 + 3), two pairs of filaments connate for part or nearly their entire length (2 + 2 + 1), or apparently all short-connate at base Although the leaves are typically glabrate on the lower surface, two collec- tions have leaves distinctly pubescent (Pires 3857; Prance et al. 8039). The fruits are usually 3.5-4.5 cm long and light in color. However, Pires 3857 has fruits 7 cm long and dark in color. These two collections may represent a distinct taxon, but with so little material available, we are reluctant to describe it. Additional specimens examined. COLOMBIA. Amazonas: Rio Loreto-yacu, ca. 100 m, Oct 1945, Schultes 6732 (F). GUYANA. Oronoque, New River and Amazon divide, ca. 280 m, Nov 1937, Beddington 29 (K). ECUADOR. Napo: Aguarico, Reserva Etnica Huaorani, km 60-61 along road and oil pipeline Maxus, Sof Rio Tivacuno, 00°51'S, 76°26'W, 250 m, 21-25 Oct 1993, Aulestia & Andi 890(MO). Pastaza: Via Auca, 115 km S of Coca, 10 km S$ of the Napo-Pastaza border, near the Rio Tigtino, Petro-Canada road, O1°15'S, 76°55'W, 320 m, 26-31 Jan 1989, Hurtado & Neill 1550 (MO). PERU. Amazonas: Yamayakat Bosque, 04°55'S, 78°L9'W, 320 m, 22 Jan 1996, Jaramillo et al. 951 (MO); Quebrada Kusu, 05°03'20"S, 78°20'23'W, 380 m, 6 Nov 1996, Vasquez et al. 21531 (MO). Lereto: Rio ee between Emilia and Brazilian village of Paumari (above Atalaia del Norte), 22 Nov 1977, Gentry & Revilla 20798 (MO, USF); Airico (native community of Shimaco-Santa Rosa), 150 m, 11 Dec 1984, Vasquez 6069 (MO, NY); Explor Camp at Rio Sucusari, 03°20'S, 72°55'W, 120 m, 19 Mar 1996, van der Werff & Vasquez 13921 (MO); Caballococha on the Rio Amazonas, 13 Aug 1929, Williams 2453 (PF), Pastaza: Via Auca, 115 km S of Coca, 10 km $ of Napo-Pastaza border, near Rio Tigtiino, along Petro-Canada highway under construction, O1°15'S, 76°55'W, 320 m, 26-31 Jan 1989, Hurtado & Neill 1550 (MO). BRAZIL. Amazonas: Rio Japura, 01°50'S, 65°40'W, 3 Nov 1982, Cid & Lima 3492 (NY); Rio Solimoes, Igarapeé Jandiatuba, 6 Jan 1949, Froes 23837 (NY); basin of Rio Solimoes, basin of creek Belém, 26 Oct-I1 Dec 1936, Krukoff 9046 (BM, EF MO, NY); mouth of Rio Iga, on bank of Rio Solimoes, 24 Feb 1977, Mori et al. 9073 (NY); Rio Jandiatuba, 10 km upstream from mouth, 26 Feb 1977, Mori et al. 9134 (kK, NY); Lago Preto on Rio Purus, 3 km N of Labrea, 29 Oct 1968, Prance et al. 8039 (K, NY). Para: Igarape Ipixuna, tributary of Rio Xingu, 04°49'S, 52°31'W, 5 km S of settlement, Araweté Indian Reserve, 23 Mar 1986, Balée 2024(NY); left bank of the Rio Sao Manuel [Telespires], Igarape Ferné anne de aig downstream from Cachoeira do Cladeirao, 7 Jan 1952, Pires 3857 (US). BO Campamento 18, 18 km N of airstrip, 10°39'S, 66°46'W, 160 m, 2 Jul 1992, Gentry et al. 77663 (MO). EXCLUDED SPECIES The following five taxa previously placed in Siolmatra by various workers are here referred to Fevillea. Siolmatra amazonica Cogn in Engler, Pflanzenr 4(Heft 66):30. 1916. [-Fevillea pedatifolia (Cogn.) C. Jeltrey Siolmatra } me — Standl., Aa F Macbride, Publ. Field Mus. Nat. Hist., Bot. Ser. 1937. [=Fevillea cordifolia L| Siolmatra pec fatifol id (Cogn.) Cogn., in Engler, seen us lett o 30. 1916. BASIONYM: Alsomitra pedatifolia Cogn., in Martius, FI. Bras. 6(4):115. 187 (Cogn.) C. Jeffrey]. Sota peruviana (Huber) Cogn., in Engler, Pflanzenr. 4(Heft 66):30. 1916. ROBINSON AND WUNDERLIN, REVISION OF SIOLMATRA 1969 BASIONYM: Alsomitra peruviana Huber, Bol. Mus. Paraense Hist. Nat. 4: 616. 1908. |=Fevillea pedatifolia (Cogn.) C. Jeffrey]. Siolmatra simplicifolia Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11:769. 1913. [- Fevillea pedatifolia (Cogn.) C. Jeffrey]. ACKNOWLEDGMENTS We gratefully acknowledge the curators of BM, BR, FE G, K, MO, NY, and US for making specimens available for our study. We thank Lorenzo Ramella (G) for providing information from his collections in Bolivia, Bruce F Hansen (USF) for his many helpful suggestions on the manuscript, and Kathleen Hotchkiss (USF) for assistance with graphics. We also thank Michael Nee (NY) and an anonymous reviewer for their helpful comments. REFERENCES Baiton, H. 1885. La fleur femelle de l’Alsomitra brasiliensis. Bull. Mens. Soc. Linn. Paris 1: 457-4 BAILLON, H. 1886. Histoire des plantes. 8:417-418. L. Hatchette et Cie, Paris. Cocniaux, A. 1893. Genre Siofmatra H. Baill. et la tribe des Zanoniées. Bull. Herb. Boissier 1:609-613. Cocniaux, A. 1916. Cucurbitaceae: Fevilleae et Melothrieae.In:A.Engler,ed.Das Pflanzenreich 66(4) 275(1):1-277.W. Engelmann, Berlin Harms, H.1926.Cucurbitaceae.In:J.Mildbraed, Plantae tessmannianae peruvianae lll. Notizbl. Bot. Gart. Berlin-Dahlem 9:989-996. Harms, H. 1933. Cucurbitaceae americanae novae. Notizbl. Bot. Gart. Berlin-Dahlem 11: 769-776. Jerrrey, C. 1962. Notes on some species of Fevillea L., Siolmatra Baill., and Pseudosicydium Harms (Cucurbitaceae) in the Amazon Basin. Kew Bull. 16:199-202. Jerrrey, C. 1978. Further notes on Cucurbitaceae: lV, some New-World taxa. Kew Bull. 33: 347-380. STaNDLEY, PC. 1937. Cucurbitaceae. In: J.F. Macbride, ed. Flora of Peru. Publ. Field Mus. Nat. Hist., Bot. Ser. 13(6):329. 1970 BRIT.ORG/SIDA 21(4) BOOK REVIEW Rosert H. MoHLENBROCK. 2005. Aquatic and Standing Water Plants of the Cen- tral Midwest: Cyperaceae-Sedges. (ISBN 0-8093-2628-0, hbk.). Southern Illinois University Press. PRO. Box 3697, Carbondale, IL 62902-3697. (Orders: 618- 459-6633, 618-453-1221 fax, wwwsiuedu/‘siupress). $65.00, 272 pp, 183 line drawings, 6" x 9" Aquatic and Standing Water Plants of the Central Midwest: Cyperaceae-Sedges by Mohenbrock is a flora, which consists of the Cyperaceae portion of the larger Aquatic and Standing Water Plants of iana, Kentucky, Ih the Central Midwest series. The book’s central Midwest range includes: Ohio, Inc nois, lowa, Missouri, Nebraska, and Kansas. The flora includes information on 183 Cyperaceae species, including members of the genera Carex, Schoenoplectus, Eriophorum, Fimbristylis, Fuirena, Trichophorum, Dulichium, Cyperus, Eleocharis, Psilocarya, Scirpus, Hemicarpha, Lipocarpha, and Rhynchospora. There is an overall key to determine genus, as well as within genus keys to determine species. Each species is presented in the book by black and white drawings, which includes i ig of the whole plant and the spikelet, achene, perigynium, scales and/or sheath, and a detaile ipti The description for each species contains a ial deal ol infor mation, eee oe current accepted specific ana ey nonymas, as well | each specific epithet. Each species’ eee includes plant h abit information, descriptions and mea- surements of culms, root types, bracts, scales, spikelets, notable colors and, w here a Sees details on perigynium. The descr eon includes habitat information, witha ee of the spe licabl (ie. FACW) within the U.S. Fish cies are typically found in, and any and Wildlife Wetlands Inve — The author has induced a handy quick field traits section wi ‘thin the Bai description to hel ine that species from similar looking others. Aquatic and Siandine Wate rP lants oft} 1€ C G ntral Midw est: Cyperac ede Sedgestlora Ww oul d be a nice ae to the library of any person who works with aquatic plants. Although focused on the Midwest, a pas o oS species included in this flora are found in various regions of North America. The black and gs P. isy to read images that show major identification traits for the species. The key is straightforward, - the flora does contain a glossary for any unfa- miliar terms, and is fairly user friendly. The description for each species is thorough, which can help you decide that you have determined the correct Species or not; and includes those very helpful “in the field” identification hints. If you need to identify sedges, be sure to look into this helpful book. —Lee Luckeydoo, Herbarium, epicnieal Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. a SIDA 21(4): 1970. 2005 REVISION OF FEVILLEA (CUCURBITACEAE: ZANONIEAE) George L. Robinson Richard P Wunderlin USDA APHIS, Be eee and Quarantine Department of Biology, SCA 110 verglades University of South Florida ee Suite 414 Tampa, Florida 33620-5200, U.S.A. Fort eae Florida 33316, U.S.A. ABSTRACT A revision of the neotropical genus Fevillea with keys, descriptions, and distribution is provided. Two subgenera are recognized: subgenus Fevillea with six species and subgenus Anisosperma with one species. Fevillea bahiensis G. Robinson & Wunderlin from Brazil is described as new and subge- nus Anisosperma (Silva Manso) G. Robinson @ Wunderlin is proposed. RESUMEN Se aporta una revision del género neotropical Fevillea con claves, descripciones, y distribucion. Se reconocen dos SU PECRETOS: ene Fevilled con seis especies y subgénero Anisosperma con una especie. Se d lea bahicnsis G. Robinson & Wunderlin de Brasil y se propone vil el subgénero Anisosperma ee ees G. Robinson & Wunderlin Fevillea (Cucurbitaceae: Zanonieae), a neotropical genus of seven species, is characterized by leaves with glands, calyx with glandular squamellae (except F passiflora), petals with a medial adaxial flap-like appendage or ridge, stami- nate flowers with five bilocular anthers, and a globose, usually indehiscent, large-seeded fruit. TAXONOMIC HISTORY Fevillea was established by Linnaeus (1753) in honor of Louis Econches Feuillée, (1660-1732), a French clergyman, explorer, astronomer, and botanist. Linnaeus recognized two species, F trilobata and F cordifolia. Adanson (1763) published the pre-Linnaean name Nhandiroba, of Marcgrave (Piso & Marcgrave 1648), but placed Linnaeus’s name Fevillea in synonymy, thereby making Nhandiroba illegitimate. No species were listed. The name Nhandiroba remained unused until resurrected by Kuntze (1891-1898). The first comprehensive treatment of Fevillea since Linnaeus (1753) was that of Seringe (1828) who recognized four species. In addition to F. cordifolia and EF trilobata, Seringe recognized FE punctata (L.) Poir.[=Trichosanthes sp.]and F. javilla Kunth [=F. cordifolia]. Silva Manso (1836) established the monotypic Hypanthera with H. guapeva [=Fevillea trilobata] and Anisosperma with A. passiflora (Vell) Silva Manso [=Fevillea passiflora Vell.]. Both monotypic genera were accepted by most sub- sequent workers until recently. SIDA 21(4): 1971- 1996. 2005 1972 BRIT.ORG/SIDA 21(4) Roemer (1846) recognized Hypanthera and Fevillea. He placed ten species in Fevillea, incorporating some taxa now placed in Trichosanthes L., Pteropepon Cogn.,and Sicydium Schltdl. Fevillea passiflora was also recognized, but with- out reference to Silva Manso’s placement of the species in Anisosperma. Roemer erected two sections: section Fevillea with nine species and section Javilla with only F. javilla Kunth =F. cordifolia L. Cogniaux (1878) recognized three species of Fevillea for Brazil: F. trilobata, F. albiflora Cogn., and F. deltoidea Cogn. (the latter now in Pterope pon). In addi- tion, he recognized the monotypic genus Anisosperma and expanded the ge- neric limits of the previously Old World genus Alsomitra (Blume) M. Roem. to include two new neotropical species: A. brasiliensis [=Siolmatra brasiliensis (Cogn.) Baill.] and A. pedatifolia [=Fevillea pedatifolia (Cogn.) C. Jeffrey]. Ina more comprehensive treatment, Cogniaux (1881) expanded Fevillea to six spe- CK cies, maintaining Anisosperma and Alsomitra. The genus Siolmatra, a segregate of Alsomitra created by Baillon (1885) to acommodate Alsomitra brasiliensis Cogn. [=Siolmatra brasiliensis (Cogn.) Baill], was accepted by Cogniaux (1893) who described a second species (S. paraguayensis). Three additional species were later added by Cogniaux (1916), one new (S. amazonica) and two transferred from Alsomitra (S. pedatifolia (Cogn.) Cogn. and S. peruviana (Huber) Cogn.). Harms (1926) added a sixth spe- cies (S. pentaphylla), later (Harms 1933)a seventh (S. simplicifolia),and Standley (1937) an eighth (S. mexiae). Jeffrey 1962b), noting the incongruent mixture of taxa in Siolmatra, recircumscribed the genus and transferred four species to Fevillea. In reviewing the New World taxa of the Cucurbitaceae, Jeffrey (1978) listed nine species in Fevillea, including F. passiflora, considering Anisosperma congeneric with Fevillea. FLORAL AND FRUIT MORPHOLOGY The staminate inflorescence consists of numerous, small, pentamerous flow- ers that are paniculate in subgenus Fevillea or fasciculate to subumbelliform in subgenus Anisosperma. In subgenus Fevillea, the midrib of the adaxial surface of the sepals is fused with the lower margins of the petals. At or above this point of fusion extends a small glandular protuberance of uncertain ontogenic origin which is here re- ferred toasa “glandular calycine squamella.” The exudate from the squamellae is clear and remains visible on most herbarium specimens. Each petal has a median, adaxial, uncinate appendage or slightly raised glandular midrib which is adnate with the base of the stamen filament. In subgenus Anisosperma, the sepals and petals are united at their base and lack squamellae. Instead, the pet- als have a median, adaxial, glandular ridge. An articulation occurs between the filiform hypanthium and the pedicel. The indumentum on the staminate flowers is quite variable in most species and ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1973 is similar (when present) on both the hypanthium and pedicel. However, F. trilobata has a stipitate-glandular pubescence on the hypanthium above the articulation in rather sharp contrast to the non-glandular trichomes on the pedicel below. The presence of five free, bilocular anthers in Fevilleais unique for the fam- ily and is considered plesiomorphic. The general trend within the Cucurbitaceae is the reduction of stamen number from five to three or to two and the reduc- tion in locule number from two to one. The pistillate flowers are rarely collected and are thus imperfectly known or unknown for some species. We have seen them in only four of the seven spe- cies. Those of F. pergamentacea and F. passiflora are known to us only through the literature while those of F. moorei are unknown. The petals of the pistillate flowers have a slightly raised median append- age extending from the base to the center, reminiscent of those of the staminate flowers. Two small glands occur at the base of the petal on either side of this median appendage. A large, subrotund, flattened, glandular protuberance ex- tends from the base of the calyx lobe between each petal. These protuberances may represent staminodes. Cogniaux (1878, 1881, 1916) reported these structures, counting 20 small “glands” at the base of the petals. This probably included the sum total of glands, ridges, and protuberances. The fruits of subgenus Fevillea are large, subglobose, mottled green or brown, and gourd-like. The size ranges from 8 to 16 cm in length and from 7 to 13cm in diameter. An individual plant may produce as many as 50 to 100 fruits at a time (Gentry & Wettach 1986). The fleshy rind of the fruit is zonate above the middle with the hypanthium lip scar (ovary partly inferior). The fruits are typically indehiscent, but reportedly sometimes dehiscent along the hy- panthium lip scar in F. pedatifolia (A. Gentry, pers.comm.). In contrast, the fruits of subgenus Anisosperma are ovoid or oblong, subtrigonous, short-apiculate at the apex, and not zonate above the middle (ovary fully inferior). The seeds (up to 15 per fruit) vary from 3 to 6 cm in diameter and weigh 3 to 9 g when dry. They are among the largest in the Cucurbitaceae and are comparable in size only to those of the paleotropical genera Telfairia (Cucurbitoideae: Joliffieae) and Hodgsonia (Cucurbitoideae: Trichosantheae). The seed coat consists of three layers. The innermost layer surrounding the cotyledon is spongy and aerifer- ous. This layer is enclosed in a thin, hard, woody layer. The outermost layer is thin, smooth, and of a corky texture which tends to obscure the margin of the woody layer below but usually does not persist. DISTRIBUTION AND HABITAT Fevillea cordifolia has the widest distribution, ranging from southern Mexico, east into the Caribbean to Puerto Rico, south through Central America and into South America to northern Argentina. Dieterle (1976) notes that it is cultivated 1974 BRIT.ORG/SIDA 21(4) in some or all Central American countries and is found in most Central Ameri- can markets, especially in Guatemala. Three species (F. trilobata, F. bahiensis,and F. passiflora) are endemic to eastern Brazil and F. pedatifolia and F. pergamentacea occur in Ecuador, Peru, and Bolivia, with F. pedatifolia extending into adjacent Amazonian Brazil. Fevillea moorei is known only from the type material culti- vated in England and is probably from Guyana or Amazonian Brazil. Fevillea typically occurs along river banks, along the edge of tropical pri- mary or secondary forests, and along the edge of seasonally inundated riverine forests, occasionally climbing to heights of 35 m in forest canopy openings. It also is found in forest clearings and along roadsides. It occurs at elevations from near sea level to about 500 m, less commonly up to 1,700 meters. Fruits and seeds of Fevillea are quite buoyant and thus apparently are well suited to dispersal in fresh water. Gentry and Wettach (1986) report that at least one species (F. cordifolia or F. pedatifolia) of Amazonian Peru occurs in season- ally inundated forests, a habitat in which water dispersal is prevalent. Seed drift materials of Fevillea cordifolia have been found within the Caribbean basin well outside the species natural range. Gunn and Dennis (1976), Morton (1981), and the senior author have identified seeds of F. cordifolia collected from beaches of southern Florida. Guppy (1917) reported materials found along beaches of the Turks Islands, Tobago, and Grenada. Guppy (1917) and Gunn and Dennis (1976) found that seeds of F. cordifolia germinated in fresh water while afloat alter the disintegration of the fruit wall, but were generally rendered non-vi- able in salt water. Although F. cordifolia is sometimes listed as an estuarine plant, the seeds are probably not capable of over-sea transport for any distance. How- ever, dispersal by seed drift via salt water can not be disregarded. Guppy (1917) estimated that 5% of the Fevillea cordifolia drift seeds reaching the Turks Is- lands were viable while Gunn and Dennis (1976) found 20% of the undamaged drift seeds on Florida beaches were viable. The high salinity of the beach is probably lethal, thus preventing colonization. — ECONOMIC IMPORTANCE The high seed oil content of Fevillea trilobata was recognized centuries ago by indigenous Brazilians whose use of it was first documented by Marcgrave (Piso & Marcgrave 1648). Fevillea cordifolia is similarly well known in the ethnobo- tanical literature (cf. Gentry & Wettach 1986). It has been used as a purgative, reputed antidote for many kinds of poisoning, and asa treatment for numerous diseases. In Jamaica it is called “antidote caccoon” or “antidote vine” (Adams 1972; Gunn & Dennis 1976; Morton 1981). Lindley and Moore (1870) first re- ported the use of Fevillea seeds by Peruvians as candles. Gentry and Wettach (1986) report that “abiria” (Fe villea pedatifolia) is used as candles by the Campa Indians of the Pichis Valley of Peru. Fevillea seeds may have potential as an edible or fuel oil source. Calculated ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1975 ona weight per fruit basis, the seed oil content of Fevillea is higher than that of any other dicotyledon (Gentry & Wettach 1986). Preliminary analysis of oils extracted from the seeds of two Peruvian species (F. cordifolia and F. pedatifolia) by Gentry and Wettach show the oil to be simple triglycerides, slightly heavier than those of refined cottonseed oil. All species were rich in the saturated low- weight fatty acids, palmitic (21-60%) and stearic (10-42%), and the unsatur- ated oleic (17-17%) and linoleic (6-7%). When compared with previous results from the Brazilian F. trilobata (Tulloch & Bergter 1979), Gentry and Wettach suggest that the Peruvian species with 60-70% low-weight, saturated fatty ac- ids would be a good sources of fuel oil while the Brazilian species with 57% unsaturated fatty acids would be a good source of polyunsaturated edible oils. However, the relatively high concentration of high molecular weight fatty ac- ids in F. trilobata, probably correlated with its purgative properties, may reduce its value as an edible oil. The high percent of stearic acid in F. cordifolia might also suggest its use in the candle industry where this chemical is used to harden waxes and in the rubber industry where used as an extender. Preliminary labo- ratory analysis of F. cordifolia at the University of South Florida gave highly variable results thought to be related to the different ages of the seeds tested, further complicating the use of Fevillea seeds as a potential commercial oil source (unpublished data). Although Fevillea as an oil source is documented in the literature, to date it remains a genus of little or no economic importance INFRAGENERIC RELATIONSHIPS Jeffrey (1962a) recognized two genera within the subtribe Fevilleinae, Fevillea and Anisosperma. In our treatment, Anisosperma is reduced to a subgenus of Fevillea. Subgenus Anisosperma differs from subgenus Fevillea by the shape of the corolla lobes, the character of the median adaxial glandular ridge of the staminate petals, the congested staminate inflorescence, the lack of glandular calycine squamellae on the staminate flowers, and the fruit shape. However, with the presence of the foliar glands, the median adaxial glandular ridge on the petals, and overall similarity in fruit and seed morphology, the single spe- cies of subgenus Anisosperma is easily accommodated in Fevillea. Within subgenus Fevillea, three species groups can be distinguished on the basis of foliar gland characters. The first group consists of F. pergamentacea and F. pedatifolia which have conspicuous petiolar glands and inconspicuous laminar glands. The others are characterized by having laminar glands only. Of these, F. cordifolia and F. trilobata have glands terminating the veins on the lamina and lack basal laminar glands while F. moorei and F. bahiensis both have basal laminar glands only. SYSTEMATIC TREATMENT Fevillea L., Sp. Pl. 1013. 1753. Nhandiroba Adan., Fam. PL. 2:139. 1763, nom. illegit. 1976 BRIT.ORG/SIDA 21 4) Type: Fevillea trilobata L. Lectotype designated by M.L. Green (in eal et al., Nom. Prop. Brit. Bot. 190. 1929). This lectotypification replaces the lectotype of Fevillea cordifolia L. of Britton and P. Wilson. (Sci. Surv. Porto Rico 6:270. 1925) under Art. 10.5b of the St. Louis Code. Anisosperma Silva Manso, naa Subst. Braz. 38. 1830. Typr: Anisosperma passiflora (Vell) Silva Manso |=Fevillea passiflora ell]. Hypanthera Silva Manso, am Subst. Braz. 37. 1836. Type: Hypanthera guapeva Silva Manso [=Fevillea trilobata L.). — Dioecious vines or lianas; stems sulcate; tendrils axillary, sulcate, distally 2-fid, coiling both above and below the bifurcation. Leaves alternate, petiolate, the blade unlobed, or palmately 3- to 7-lobed, or 3- to 5-foliolate, with glands on the leaf margins terminating the primary lateral veins and/or 2 glands at the blade base or on the petiole, the petiole canaliculate, sometimes bearing 2 glands at or above the middle. Staminate inflorescences paniculate or subumbelliform, many-t lowered, bracteate; flowers short-pedicellate; hypanthium pedicelloid; calyx lobes 5, fused to the petals above, not completely enclosing the petals in bud, witha glandular calycine squamella on each calyx lobe at or near the point of fusion with the petals (except in F. passiflora), petals 5, fused to the sepals below, the lobes each with a median, adaxial, uncinate, flap-like appendage or a slightly raised ridge or (in F. passiflora) witha thick, glandular ridge; stamens 5, equal, free, inserted near the center of the flower, the anthers bilocular, extrorse, dehiscing longitudinally, the connective with an adaxial glandular protuber- ance or projection; pollen prolate, 18-22 win length, tricolporate, coarsely striate. Pistillate flowers solitary or in pairs; hypanthium deeply cupular; sepals and petals as in the staminate flowers or sometimes the petals differing in shape; ovary partly inferior, 3-locular, the styles 3, free, outwardly curved, the stigmas reniform, capitate, the ovules pendulous, usually 4 in each locule. Fruit globose, gourd-like with a thick, fleshy rind, zonate above the middle with the hy- panthium lip scar or non-zonate, indehiscent or rarely circumscissile dehiscent along the hypanthium lip scar; seeds large, orbicular, somewhat compressed, the seed coat consisting of a thick, spongy, aeriferous inner layer surrounded by a thin woody layer, and an outer, usually non-persistent layer, the lateral surface smooth or striate-verrucose, the outer edges smooth or tuberculate, the inner kernel disk-like, oily. = Two subgenera are distinguished as follows: 1. Corolla lobes of the staminate flowers suborbicular, the base cuneate, with a me- dian adaxial uncinate appendage or sharply defined ridge; glandular calycine squamellae present between the petals and the calyx lobes; staminate flowers in spreading panicles; fruit subglobose, zonate above the middle, the apex rounded subg. Fevillea 1, Corolla lobes of staminate oblong-hastate, the base with a median adaxial glandular ridge broadening downward; glandular calycine squamellae absent: staminate flowers in congested oo or subumbelliform; fruit ovoid to oblong, not zonate, the apex short-apiculate subg. Anisosperma f| ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1977 Fevillea subgenus Fevillea Javilla M. Roem., Fam. Nat. Syn. Monogr. 2:116. 1946. TyPE: Fevillea javilla Kunth [ Fevillea cordif li LL. Staminate inflorescences paniculate; glandular calycine squamellae present; corolla lobes suborbicular with a median adaxial uncinate appendage or slightly raised ridge. Fruit subglobose, zonate above the middle, the apex rounded. 1. Leaves pedately 3- to 5-foliolate F. pedatifolia 1. Leaves lobed or unlobed, but not pedately foliolate. 2. Leaf blade without glands at the base or on the petiole, with marginal glands terminating the veins. 3. Leaf blade with angled or rarely with rounded lobes, the marginal glands inconspicuous; staminate flowers with the hypanthium densely stipitate- glandular pubescent, the pedicel with non-glandular trichomes F.trilobata 3. Leaf blade unlobed or occasionally with rounded lobes, the marginal glands conspicuous; staminate flowers with the hypanthium and pedicel variously ubescent but not as above F. cordifolia 2. Leaf blade with glands at the base or on the petiole, with or without marginal glands iernlner ioe the Fat veins. Leaves wit ly at the blade base, without marginal glands terminat- ing the leat veins of the blade. 5. Staminate flowers 3-5 mm wide; sae drying reddish-brown, the blade with a conspicuous uncinate-ciliate margin F. bahiensis 5. Staminate flowers 15-20 mm wide; ae drying green, the blade with a smooth margin F. moorei 4. Leaves with glands either at the blade base or on the petiole, also with mar- ginal glands terminating the veins of the blade 6. Leaves with glands at the blade base F. pergamentacea 6, Leaves with glands on the petiole F. pedatifolia Fevillea bahiensis G. Robinson & Wunderlin, sp. nov. (Fig. 1). Tyre: BRAZIL. BAHIA: 8 km to the N of Ubaitaba on BR 101, 16 Jun 1972, dos Santos 2307 (HOLOTYPE: CEPEC; ISOTYPE: K). Species haec a Fevillea moorei Hook. f. differt floril i foliis in siccitate badiis usque atrobrunneis marginibus uncinato-ciliatibus. Vine or liana; stem glabrous to lightly appressed golden brown-pubescent; ten- drils glabrous to lightly pubescent. Leaves unlobed, the blade ovate, (2.5-)6.5- 10.5(12) cm long, (3.5-)5.5-9 cm wide, membranaceous, drying dark brown to reddish brown, 5-nerved, the apex acuminate, the base cordate to truncate, the margin entire, with two irregularly shaped glands at the base near the petiole, the upper and lower surfaces glabrous or with scattered, appressed, golden brown trichomes, these usually denser along the leaf veins, the margin unci- nate-ciliate, the petiole (1.5-)3.5-5 cm long, glabrous to lightly pubescent. Stami- nate flowers in a paniculate inflorescence on reduced subterminal, lateral branches, the branches subtended by a reduced leaf; pedicel 1.5-2 mm long, glabrous or sparsely pubescent; bracts linear, ca. 1 mm long; hypanthium 1.5-2 mm long, lightly golden brown-pubescent; calyx shallowly cupular, the lobes 1978 BRIT.ORG/SIDA 21(4) Fic. 1. Fevillea bahiensis. A. Habit, st te infl (dos S$ 2307). B. Staminate fl (dos Santos 2307). up to 15 mm long, | mm wide, glabrous to sparsely pubescent abaxially, the margin entire, the apex rounded, with scattered stipitate glands, with a glan- dular calycine squamella at or near the point of fusion with the petals; petals suborbicular, 2.5-3 mm long, ca. 1.5 mm wide, cream-colored, the margin en- tire, the median adaxial ridge slightly raised; stamens 1-15 cm long, the an- thers ca.0.25mm long, slightly longer than wide, the filaments ca. 0.5mm long. Pistillate flowers solitary; calyx deeply cupular, the lobes ca. 1.5 mm long, ca. | mm wide, dark brown, fleshy; petals strap-shaped, ca. 2 mm long, ca. 1.5 mm wide, cream-colored. Immature fruit subglobose, 2-4 cm long and wide, the surface smooth; mature fruit not seen. Distribution and ecology.—Endemic to Brazil in southern Bahia. — ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1979 Additional specimens examined. BRAZIL. Bahia: Nova Esperanca, Sao Lourenco, 32 km W of Wenceslau Guimaraes, 26 Jul 2001, Mattos-Silva et al. 4479 (NY); Ramal da Torre da Embratel, en- trance 15 km [rom the Ubaitabe/Itacaré Road (BR 654), 5.8 km from the entrance, 6 Jun 1978, Mori& bs Santos 10135 (CEPEC, kK, NY); km 3, Urucuca-Taboquinha highway, 19 Jun 1972, dos Santos 2316 EPEC); access road to Torre da Embratel, entrance to the right of road to Ubaitabe/Itacaré (BR 3 24 Sep 1977, ae Santos 3130 (CEPEC, K); Almadina Mata da Serra Pancadinho, 10 Mar 1971, Pinheiro 1076 (CEPEC, kK). Fevillea vee is most similar and probably most closely related to Fe villea moorei of Guyana Amazonian Brazil. Both species have glands only at the base of the leaf blade. Fevillea bahiensis differs by having smaller staminate flow- ers (petals 2.5-3 mm long vs. ca. 1 cm long in F. moorei) and the leaves drying a dark reddish brown and with conspicuous uncinate-ciliate margins. Fevillea cordifolia L., Sp. Pl. 1013. 1753. (Fig. 2). Fevillea scandens L. Sp. PL. ed. 2 1763, nom. ee Deas sibs ns Descourt., Fl. Méd. Antilles, see = 198} ee nom. illegit. ) Kuntze, Revis. Gen. Pl. oa 1891. ea cordifolia L. var. typica Stehlé, M. Stehlé SiOuentin, Fl. Guadeloupe 2(3):133. 1949, nom. inadmiss TYPE: “Habitat in America Calidiore” (LECTOTYPE: “Nhandiroba,” Plumier, Pl. Amer. 20, t. 27. 1703). Lectotypified by C.Jeffrey, (in CE. Jarvis et al, eds. Regnum Veg. 127:47. 1993). Fevillea hederacea Poir, in Lamarck, Encycl. 4:418.1798. ele cc een ala ine Cogn., in _de Candolle & C. de Candolle, Monogr. Phan. 3:943. 1881. TyPE: “On la cultive au jardin des plantes. Elle est ea del ene (vs.)” CHOLOTYPE: P?, aa Fevillea javilla Kunth, in Humboldt, Bonpland, & Kunth, Nov. Gen. Sp. 2:124. 1817. TyPE: CO- LOMBIA. Bolivar: Turbaco, Hum at & Bonpland 1403 (HOLOTYPE: P, n.v. [microfiche IDC 6209 39:11 7). Fevillea karstenii Cogn., in Alph. de Candolle & C. de Candolle, Monogr. Phan. 3:943. 1881. ndiroba karstenii (Cogn.) Kuntze, Revis. Gen. PL. 1:257. 1891. TyPE VENEZUELA. DISTRITO FEDERAL: Capaya, near Caracas, Karsten s.n. (HOLOTYPE: W, n.v; photo ex W: F, MO). The type sheet contains a mixed collection of F. cordifolia and probably Selysia prunifera (Poepp. & Endl.) C he material of the latter in the lower left portion of the sheet is excluded. Fevillea t ene See Gain, Fl. Mexic. ed. 2. 231.1894. TYPE: MEXICO. n mapa te mexiae Stand, in J.F Macbride, Publ. Field Mus. Nat. Hist., Bot. - 13(6):329. 1937. ee Loreto: left bank of Rio Maranon, above Rancho Indiana, 110 m, 22 Jan 1932, i i HOLOTYPE: F; photo ex F: F; 1so Fevillea fine ipetals Kuhlm., Arch. Jard. Bot. Rio ane 4:365. 1925. TYPE: BRAZIL. PARA: Rio Branco de Obidos, Castanhal Grande, 4 Nov 1919, Ducke s.n. (RB 15924) (HOLOTYPE: RB, n.v; ISOTYPE: B, destroyed; photograph ex B: F MO, NY, US). Vine or liana; stem glabrous or lightly to densely pubescent or tomentose; ten- drils glabrous to lightly glandular-pubescent. Leaves with the blade unlobed or occasionally 3- to 5-lobed, suborbicular to cordate, (4-)7.5-13(-18) cm long, (2.5-)5-12(-19) cm wide, membranaceous or coriaceous, 5-nerved, the apex acute, the base cordate to truncate or rarely rounded, the main lateral veins terminating in small irregularly shaped glands, the upper and lower surfaces glabrous to densely pubescent, the petiole (2-)3-7 cm long, glabrous or pubes- cent. Staminate flowers in a paniculate inflorescence on reduced subterminal, lateral branches, the branches subtended by a glandular bract 0.5-4 mm long; 1980 BRIT.ORG/SIDA 21(4) Fic. 2. Fevillea cordifolia.A. Habit. staminate inf] (Lent 3288 ).B. Staminate fl (Klug 3090).C Fruit (Huashikat 1078).D. Seed (Huashikat 1078). pedicel 2-4 mm long, glabrous to densely pubescent, bracts linear, glandular, ca. 0.5mm long; hypanthium 3-4 mm long, glabrous to densely pubescent; ca- lyx shallowly cupular, the lobes 1-2.5 mm long, 1-15 mm wide, obtuse or rounded, densely to sparsely glandular-pubescent, the margin entire, the apex rounded, with a small glandular calycine squamella protruding from each se- pal at or near the point of fusion with the petal; petals suborbicular, 3.5-5 mm long, 2.5-4 mm wide, white, whitish green, cream-colored, light brown, pink or pinkish orange, dark red, or reddish purple, the margin undulate, the median, adaxial ridge with an uncinate appendage; stamens ca. | mm long, the anthers ca. 0.5mm wide, slightly longer than wide, the filaments ca. 0.5 mm long. Pis- tillate flowers solitary or in pairs; calyx deeply cupular, the lobes suborbicular, ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1981 ca. 3mm long and wide, pustulate, fleshy; petals oblong, ca. 5 mm long, ca. 3.5 mm wide, cream-colored, the median adaxial appendage slightly raised, with 2 small suborbicular glands on each side of the ridge at the base. Fruit subglobose, gourd-like, 10-12 cm in diameter, 10-16 cm long, the surface pustulate, mottled green, zonate above the middle with the hypanthium lip scar and marked at the apex by a raised triradiate line; seeds orbicular, compressed, 1-2.5 cm thick, 4-6(-7.5) cm wide, the lateral surface of the woody layer striate-verrucose or pustulate, with the outer edges smooth or occasionally tuberculate, the tubercles 2-3 mm long. Distribution and ecology.—Southern Mexico east to Puerto Rico, south through Central America, and into South America to Bolivia. A canopy plant of wet forests from sea level to 1,700 meters. — mate Sas examined. MEXICO. Guerrero: Acapulco, 1894-1895, Palmer 335 (MO). GUATE- : near Antigua, 1500-1600 m (cultivated, found in market), Nov 1938-Feb 1939, . a ine yo ie er on NICARAGUA. Rio San Juan: | km E of the village of Sabalo, 11°02'N, 84°29'W, 50 m, 6 Sep 1985, Moreno 26249 (MO), between Pueblo de San Juan del Norte Nuevo and La Casa de Ramon Castillo Viajando by San Juanillo, 10°55'N, 83°49'W, 0-100 m, 7 Jul 1994, Rueda et al. 1839 (MO), Rio Pigibaye, 18 Feb 1995, Rueda et al. 3199 (MO); Reserva Indio-Maiz, along Rio Indio, 11°06'N, 83°58'W, 5-20 m, 19 Sep 1998, Rueda et al. 8822 (MO). Rivas: Isla Ometepe, Volcan Maderas, Hacienda “La Argentina,” 11°27'-28'N, 85°31'W, 700-900 m, 15 Jun 1984, Robleto 864 (MO); “Las Cuchillas,” Isla Ometepe-Volcan Maderas, 11°27'N, 85°28 W, 400-800 m, 2 Jun 1985, Robleto 1972 (MO). Zelaya: Cano Monte Cristo, “La Grupera,” 11°33'N, 87°48'W, ca : m, 4 Feb 1982, Moreno & Sandino 14743 (MO, NY). COSTA RICA. Heredia: Finca La Selva, OTS field station on the Rio Puerto Viejo just E of its junction with the Rio Sarapiqui, 24 Mar 1980, ae 8251 (MO). Limon: Tortuguero-Sierpe basin, near Rio Sierpe and Rio Penetencia, 10°32'40'N, 83°32'50"W, 20 m, 21 Jan 1997, Hammel & Grayum 20720 (MO); Rio Jiménez, 18 Mar 1973, Lent 3288 (F MO); Cordillera de Talamanca, Reserva Biologica Hitoy Cerere, road between Estacion de la Reserva and Cerere, 9°40'20"N, 83°01'35'W, 100 m, 23 Feb 1989, Herrera & Chacén 2434 (MO), Puntarenas: Peninsula de Osa, Estacion de Oro, along the Aquaduct, 08°42'00"N, 83°29'10"W, 150 m, 10 Feb 1996, Angulo 517 (MO, NY); Peninsula de Osa, La Tae Guadalupe, Finca de Efrain Gonzalez, 08°3830"N, 83°28'00"W, 50 m, 17 Aug 1993, Aquilar 2119 (MO); Peninsula de Osa, Rancho Quemado, road to Draque, 08°42'00"N, 83°33'00W, LOO m, 30 Jan ior Nielsen 895 (MO), Valle de Coto Colorado, 08°46'00'N, 83°15'00"W, 100 m, 25 Jun 1993, Quesada & Segura 705 (MO); Playa San Josecito, Peninsula de Osa, 08°37'00"N, 83°44'00"W, 10-100 m, 10 Dec 1993, Quesada et al. 848 (MO); Valle de Coto Colorado, shore of Rio Esquinas, mouth of Rio Esquinas, 08°44'00'N, 83°20'00"W, 30 m, 17 Dec 1993, Segura eta 5(NY); Forest de Santo Domingo de Golfo Dulce, Mar 1896, Tonduz 10078 (BR). San José: Cordillera de Talamanca, Las Nubes, Estacion Santa Elena, 09°23'30"N, 83°36'30"W, 1150 m, 14 Feb 1996, Alfaro 477 (MO). PANAMA. Chiriqui: Burica Peninsula, Rabo de Puerco, 8 km along road W from Puerto Armuelles, 150 m, 19 Feb 1973, Busey 440 (F, MO, NY, USF). Colon: Barro Colorado Island, 100 m S$ of Zetek Trail, 600 m, 26 Aug 1970, Croat 11918 (F, MO, NY, USF). Darién: Rio Sabana, above Sante Fe, 14 Sep 1967, Duke 14107 (MO). Los Santos: 17.8 miSof Macaracas, ca. 300 m, 25 May 1967, Burch 1605 (MO). Panama: 12.4 km E of Canita, 10 Oct 1975, Witherspoon 8704 (MO). CUBA. Oriente: Bayate at Rio Jagua, 4 May 1919, Ekman 9613 (G, K, NY, US). JAMAICA. Portland: gorge of the Swift River at Eden, 0.5 mi N of Paradise, ca. 30 m, 19 Mar 1956, Proctor 11871 (MO). St. Andrew: Hope Gardens, 29 May 1902, Harris 8381 (BM, NY). St. Ann: cave near St. Ann’s Bay, Dec 1873, Purdie s.n. (K). St. Mary: Wapping Stairs, N side of Guys Hill, 20M ar 1960, Proctor 20713 (NY). St. Thomas: near Dove Hall, 10 Feb 1850, Alexander s.n. (kK, NY). HAITI. ae du Nord, Le Borgne, edge of RiviPre du Borgne, 12 Sep 1925, Ekman 4851 (US). DOMINICAN REPUBLIC. Kho 1982 BRIT.ORG/SIDA 21(4) El Seibo: Cordillera Oriental, ca. 6-8 km S of Miches-Las Lgunas de Nisibon Highway, on road to Batey Arroyo Santiago, basin of Rio Yeguada (S of Miches), 18°55'N, 69°04'W, 80-100 m, 28 Jun 1990, Zanoni & Jiménez 44609 (MO). La Vega: Jarabocoa, Monabao, Los Calabazos, Arroyo Frio entrance, 19°4'23"N, 70°43'34.6"W, ca. 774 m, 19 Mar 2001, Ososki & Saborio 299 (NY). PUERTO RICO, Along road between Utuado and Adjuntas, km 40, 2 Feb 1997, Acevedo & Angell 9419 (NY); Bayamon, 31 Mar 1885, Sintents 986 (BM, BR, G, K, M, NY, US). COLOMBIA, Amazonas: Loreto-Yactt River, ca. 100 m, Sep 1946, Schultes & Black 83351 (K). Antioquia: near Rio Leon ca. 20-30 km upstream and S$ of the river mouth ca. 15 km W of Chigorodo, ca. 7°45'N, 76°50'W, ca. LOO m, 14 Mar 1962, Feddema 1907 (NY); Murri la Blanquita, Rio Murri, 06°35'N, 76°50'W, 960 m, 28 Feb 1992, Gentry et al. 75799 (MO): km 28.8, Nutibara-La Blanquita road, 06°40'N, 76°27'W, 1020 m, 5 Nov 1988, Zarucchi et al. 7172 (MO). Atlantico: Barranquilla an hee Jan 1934, Elias 1173 (F US). Bolivar: aes of Turbaco, Nov 1920, Heriberto 469 (F, US). Chocé: Rio San Juan, Quebrada del Taparal, 5-20 m, 30 May 1946, Cuatrecasas 21504 (FP): Rip Chiniatlo 1-2 !/2 hrs. above La Nueva, 6 Feb 1967, ae sere right bank of Rio Baudo, ca. 18.5 km upstream of estuary, between estuary of Quebrada P. sie ee the sawmill Porquera, ca. 5m, 6 Feb 1967, Fuchs & Zanella 21791 (NY). Cundinamarca: Sierra de Subia, 6.6 km N of Cumaca along road to Viota, 1700 m, 22 Jun 1972, Barclay et al. 3521 (US). alee Santa Marta, 5 miS of Cienaga, near sea level, 12 Sep 1898-1899, Smith 1607 (BM, BR, FG, MO, NY, US). Meta: Serrania de la Macarena, Plaza Bonita, bank of eae 400 m, 14 Noy 1949, Philipson etal. 1423 (F BM, US). Putamayo: Frontera Colombia-Ecuatoriana, along Rio San Miguel, at mouth of the Rio Conejo, 300 m, 9 Dec 1940, Cuatrecasas 10915 (US); San Antonio, ‘Alto es path, Finca La mariposa, 1350-1420 m, OL°12'N, 76°38'W, 10 Apr-l May 1994, Ferndndez et al. 10745 (NY). Vaupés: Rio Guayabero, 240 m, 8 Nov 1939, Cuatrecasas 7499 (US). VENEZUELA. Apure: Reserva Forestal San Camilo, vicinity of Chirocoa, 9-10 km E of Caserio San Camilo (El Nula), 200 m, | Apr 1968, Steyermark et al. 101665 (MO, NY). Aragua: Parque Nacional Henri Pittier, Estacion Biologica de Rancho Grande 6 Jun 1987, Rojas & Rojas 3617 (MO, NY); Parque Nacional Henry Pittier, between trail up Periquito and Fila de Periquito, along upper slopes of tributary to Quebrada Palo Vaco on side tow oe Ss Valencia, opposite Rancho Grande Biological Station, 1300-1400 m, 25 Oct 1961, Steyerm CE NY, US). Carabobo: Rio Moron, E of Moron, L0°17-18'N, 68°1L0-16'W, 0-50 m, 20- ore 199], Digs & Jiménez 487 (MO). Delta Amacuro: between La Margarita and Puerta Miranda, Rio Cure, 80-100 m, 26 Nov 1960, Ste yermark 87780 (NY, US); Rio Acure, 29 Jan 1980, Trujillo & Sulbaran 16234 (MO). Distrito Federal: Hacienda Chichiriviche, ca. 300 m, Jul 1958, Aristeguieta 3225 (NY, US). Mérida: 0.5-2 km ae dam site on Rio Guaimaral, 7°45'N, 71°29'W, 15 Mar 1981, Liesner & Gonzdlez 10637 (MO). Miranda: Carretera Santa Teresa~Guatopo, ca. 300 m, 4 Jun 1959, Trujillo 4149 (US). Yaracu Sierra de ae Jul 1953, Aristeguieta & Pannier 1854 me Zulia: ca. 5 km SSE of Desieatien Guasare No. | (La Yolanda), 10°52'10"N, 72°29'30"W, 250-350 m, 16 Nov 1982, Bunting et al. 12420 (NY), 8 km from San José de Los Altos on road to Cano Colorado, 390 m, 27 Nov 1977, Jeffrey & Trujillo 2396 (EK, MO); 6 km W of main road and 2 km $ of Rio Catatumbo, 09°6'N, 72°42'W, ca. 20-100 m, 29 Mar 1982, Liesner & Gonzdlez 13347(MO, NY). ECUADOR. Morona Santiago: El Centro Shuar Pampants, Rio Kankaim (Cangaime), 02°47'S, 77°36'W, 300 m, 10 Sep 1985, Warush RBAES85 (NY). Napo: 5 km SE of Las Sachas, 300 m, 13 Apr 1985, Baker et al. 5995 (NY); Parroquia Dureno, indig- enous Cofan-Dureno community, 00°02'S, 76°42'W, 350 m, 29-31 Dec 1987, Ceron & Cerén 3107 (MO, NY); Estacion Biologica Jatun Sacha, Rio Napo, 8 km E of Misahualli, 01°04'S, 77°36'W, 450 m, 22 Oct 1988, Ceron & Iguago 5510 (MO), Hollin-Loreto-Coca highway, between Avila and Rio Pucuno, 00°39S, 77°22'W, 800 m, 10 Dec 1987, Ceron et al. 2871 (MO, NY); Parque Nacional Yasuni, along Maxus road and pipeline construction project, km 54-54, 13-16 Sep 1993, Dik 436 (MO, USF); km 2, new Cotundo- Coca highway, 1130 m, 5 Aug 1984, Bedson: etal. 15057 (MO); Coca-Auca oilfields road, km 53, 00°50'S, 76°52'W, 400 m, 20 Aug 1979, Jaramillo & Coello 19710 (NY, US); Maxus road, km 1.8, 00°27'S, 76°38'W, 21 Sep 1997, Klitgaard et al. 617 (NY); Parque Nacional Yasuni, Anangu, along Rio ee near junc- tion with Rio Napo, 0°31'S, 76°23'W, ca. 270 m, 16 Jun 1982, L uteyn etal. 8498 (NY); near NW corner bray 69951 (MO); along Rio Indillama, ne of Rio Napo, of Lake Liméncocha, Sep 1969, Mow ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1983 Comuna Pompeya, 00°30'S, 76°40'W, 220 m, 5 Dec 1992, Neill et al. 10192 (MO, NY); right bank of Rio Napo, 8 km from Puerto Misahualli, 01°04'S, 77°37'W, 450 m, 7-16 Sep 1988, Palacios 3006 (MO, NY); Codo Sinclair, 00°08'S, 77°27'W, 650 m, 16-20 Sep 1990, Palacios 5719 (MO); Rio Huataraco, towards Ishpano, 00°44'S, 77°23'W, 700 m, 30 Nov 1992, Palacios 10510 (MO, USF); 6 km N of Shushufindi, towards Dureno, 0°10'S, 76°40'W, 450 m, ens Palacios et al. 829 (MO); Dureno on Rio Aguarico 2 Jul 1966, Pinkley 103 (MO), Yusuni National Park, along Maxus road os ee construction project, km 21, 00°33'S, 76°31'W, 250 m, 24 Jul 1994, Pitman 646 (MO, USF); 20 km N of Coca, Palmoriente property, 00°20'S, 77°05'W, 250 m, 3-21 Nov 1989, Rubio 332 (MO, i Pastaza: ARCO oil well Villano 2, 01°25'S, 77°20'W, 400 m, 1-18 Dec 1991, Hurtado 2908 (MO); Petro-Canada highway under eee Via Auca, 115 km S$ of Coca, 5 km S of Rio Tigtiino, 01°15'S, 76°55'W, 320 m, 1-6 Mar 1989, Zak 4 MO). Pichincha: Santo Domingo de los Colorados, 800 m, 10 Aug 1945, Solis see (F). Carchi: se ndigena Awa, Parroquia Tobar Donoso, sector El Baboso, 78°20'W, 00°53'N, | m, 3 Oct 1991, Rubio & Talcuz 295 (MO, USF). PERU. Amazonas: S of Huampami across Rio ae 700-900 m, 27 Dec 1972, Berlin 716 (MO), Rio Santiago, Canton de la Quebrada Caterpiza, 1 km from community of Caterpiza, 200 m, 11 Sep 1979, Huashicat 552 (MO, USF); Yamayakat Brosque, 04°55'S, 78°19'W, 320 m, 16 Jan 1996, Jaramillo et al. 881 (MO). Huanuco: vicinity of Tingo ee Insupte, 670 m, 16 Aug 1961, Schunke 5645 (F US). Loreto: Flor de Yarina-Rio Samiria, 20 Oct 1982, Ayala et al. 3922 (NY); Explorer's Inn, Rio Amazonas near Indiana, 03°30'S, 73°03'W, 130 m, 20 Feb 1988, Gentry et al. 61736 (MO); Santa Rosa, lower Rio Huallaga below Yurimaguas, ca. 135 m, 1-5 Sep 1929, Killip& Smith 28720 (NY), Balsapuerto, ca. 220 m, May 1933, Klug 3090 (BM, FG, MO, NY, US); Quebrada de Tamishaco above Tamishaco, 7 Nov 1978, Rimachi 4046 (MO); Florida, 8 Feb 1980, Rimachi 4856 (NY): Rio Itaya, highway to San Antonio, near Venezia, 90 m, 21 Sep 1994, Rimachi 11120 (NY); Bosque Na- cional de Iparia, along the Rio Ucayali near Iparia (80 km at the confluence with Rio Pachitea), 250- 300 m, 23 Aug 1968, Schunke 2670 (NY); San Antonio, Rio Itaya, 04°10'S, 73°20'W, 150 m, 13 Dec 1982, Vdsquez & Jaramillo3597 (MO); Cocha Pastor, Isla Padre, 03°45'S, 76°10'W, 116 m, 21 Dec 1982, ee et. al. 3683 (NY); Indiana, Explorama Inn, 03°30'S, 73°05'W, 108 m, 12 Apr 1992, Vasquez et al. 182 (MO). Madre de Dios: Parque NES iat Manu, Cocha Cashu Biological Station, 21 Aug 1976, a — & Augspurger 3394 (K, MO, NY, ycha Cashu Cam p, Parque Nacio nal de Manu, along Rio Manu, 380 m, 22 Oct 1979, Gentry etal. ae nae )Cu Amazonico Lodge, 15 km NE of Puerto Maldonado, 12°35'S, 69°03'W, 200 m, 18 Jun 1990, Nunez 12192 (MO): Las Piedras, Cusco Am Seen Rio Madre de Dios, 12°29'S, 69°03'W, 200 m, 13 Aug 1991, Timand 1997 (MO). San Martin: Valley of San Martin, E of Tarapoto, Funde de San Isidro near Codo Creek, 1000 m, 15 Aug 1937, Belshaw : 30 (NY); ee a Cainarachi, Rio Cainarachi, ErDubany of Rio Huallaga, ca. 230 m, Sep- 1932, Klug 2749 (BM MO, NY. ioja-Pomacochas road, below Venceremos, ca. 20 km NW of Rioja, 05°45'S, 77° oe Ww. 1600 m, 8 Feb 1984, Gentry & Smith 45128 (MO): san Chazuta, W o ree Chazuta, 06° 34'S, 76°12'W, 200-300 m, 28 Aug 1986, Knapp 8177 (NY); W of Nueva Aspusana (2 hrs down Rio Huallaga from La Roca), 8 Aug 1962, Mathias & Taylor 6116 ne Fundo La Campina, 2 km below Tocache Nuevo, right bank of Rio Huallaga, ca. 400 m, 23 Aug 1969, Schunke 3377 (NY); Tananta (left bank of Rio Huallaga), 6 Oct 1970, Schunke 4479 (NY); Nueva Union below Puerto Huicte (right bank of Rio Huallaga), 450-500 m, 1 Aug 1974, Schunke 7965 (NY). Ucayali: Bosque Nacional de Iparia, along the Rio Ucayali near the village of Iparia (ca. 80 km above the confluence with Rio Pachitea), 200-300 m, = 1968, paar k Se S Gh Rio Novi ia, right bank at native community San José, 10°12'S, 70°57'W, 189 m, 26 Feb 2002, Sc & Graham S14909(NY). BRAZIL. Acre: margin of Rio Azul, ca. O7° pe 73°39 o 13 Oct te Campbell et al. 8995 (NY); Sena Madureira, 28 Sep 1980, Cid & Nelson 2596 Pe basin of Rio Jurua. right bank of Rio Tarauaca, 8°32'51'S, 71°28'39"W, 17 Nov 1995, Daly et al.8562 (MO, NY); near mouth of Rio ieesae eens of Rio laco), 9°20'S, 69° W, 23 Aug 1933, Kru ned 5610 (BM, a G, M, MO, US). Amazonas: near mouth of Rio Embira (tributary of Rio Tarauaca), 7°30'S, 70°15'W, 6 Jul 1933, Krukoff 5209 (BM, EG, o ve NY, ae Para: Belém, 20 Dec 1950, Black 50- oe (NY); Rio Pacaja 2°50'S, 50°50'W, 15 Oct 1965, Prance et al. 1636 (NY); Rio Mocooes, 45 min. below Frances, 00°45'S, 49°41'W, 10 Nov 1987, Prance et al. 30399 (MO, NY): Altamira, left bank of lower 1984 BRIT.ORG/SIDA 21(4) Rio Xingu, 19 Oct 1986, Souza et al. 385 (NY); Travessao do CNEC, between D13 and edge of Rio Xingu, 2 Dec 1986, Souza etal. 635 (NY); ha de Marajo, Cuanta, above Anajas, Rio Anajas, 00°57'S, 49-48° W, 2 Nov 1987, Tavares 334 (NY). Rondonia: FE bank of Rio Madeira at Misericordia between Cachoeiras Misericordia and Madeira, 30 Jul 1968, Prance etal. 6620 (NY). BOLIVIA. Cochabamba: Proyecto Valle del Sacta, km 240 on Santa Cruz-Villa Tunari highway, 17°00'S, 64°46 W, 290 m, 12-14 Jul 1989, Smith etal. 13711 (MO). El Beni: Rio Beni, above confluence with Rio Quiquibey, 3.5 hrs. upstream from Rurrenabaque, 14°44'S, 67°25'W. 320 m, 23 May 1990, Daly et al. 6590 (MO, NY). La Paz: Parque Na- cional Madidi, 10.2 km NW of turnoff in pera 200-500 km from summit, 14°09'57"S, 67°55'02"W, 830 m, 9 Aug 2000, Croat et al. 84416 (MO); Santa Fe, NE of community13°40'S, 68°12'W, 250 m, 10 Aug 1995, De Walt et al. 823 (MO, NY); basin of Rio Bopi, San Bartolomé near Calisaya, 750-900 m, |- 22 Jul 1939, Krukoff 10528 (FG, K, MO, NY, US); Parque Nacional Madidi, near Arroyo Aguapolo and Rio Tuichi, 270 m, 16 Mar 2002, Macia et al. 6850 (NY). Santa Cruz: Parque Nacional Amboro, along Rio Isama [*Rio Pitasama’], 17°42-43'S, 63°37-38'W, 475 m, 1] Oct 1990, Nee 39252 (NY); Estancia San Rafael de Amboré, 15 km (by air) SSE of Buena Vista, 17°35'S, 63°37'W, 375 m, 28 Jul 1987, Nee et al 35391 (NY): Rio Palometilla, 400 m, 16 Jun 1927, Steinbach 7904 (FG, MO, NY, BM) Fevillea cordifolia is a polymorphic species exhibiting considerable variation throughout its range. The species can be readily distinguished from other mem- bers of the genus by its conspicuous marginal laminar glands. It is most similar to F. trilobata of eastern Brazil and is distinguished from that species by the typically rounded leaf lobes Gwhen lobes are present), rather than the angular lobes characteristic of F. trilobata. Fevillea trilobata also has a distinctive stipi- tate-glandular pubescence on the hypanthium of the staminate flowers, which is sharply differentiated from the uncinate pubescence of pedicel. This pattern of pubescence is lacking in F. cordifolia. Fevillea moorei Hook. f., Bot. Mag. t. 6356. 1878. Nhandiroba moorei (Hook. f.) Kuntze, Revis. Gen. PL. 1:257.1891. Type: GUYANA or BRAZIL: Cultivated at Liverpool Botanic Garden, 1871, Tyreman s.n. HOLOTYPE: K; ISOTYPE: BR-Lragment). Vine or liana; stem glabrous; tendrils glabrous. Leaves with the blade unlobed, broadly ovate, 6-12 cm long, 3.5-7 cm wide, membranaceous, drying light green, 3-nerved, the apex acuminate, the base rounded, with 2 small, irregularly shaped glands near the petiole, the upper and lower surfaces glabrous, the petiole 1-2.5 cm long, glabrous. Staminate flowers in a racemose inflorescence ca. 6 cm long, (10-12 cm fide Hooker f.); pedicel 1-4 mm long, glabrous; hypanthium 5-6 mm long, glabrous; bracts linear, ca. 0.5 mm long; calyx shallowly cupular, the lobes oblong, ca. 6 mm long, 4 mm wide, the apex obtuse, the margin entire, with a glandular calycine squamella protruding at or near the point of fusion with the petals; petals suborbicular, somewhat broader at the apex, ca. | cm long, | cm wide, pale brick-red, the margin undulate; stamens ca. 4 mm long, the an- thers ca. 1 mm long, slightly longer than wide, the filament ca. 3 mm long. Pis- tillate [lowers and fruit not seen. an — Distribution and ecology.—Known only from the type material received by J.D. Hooker from David Moore, curator of the Glasnevin Botanic Garden, Dub- lin, Ireland, who received it from Mr. Tyreman of the Liverpool Botanic Garden. ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1985 The material was labeled as “Strychnos curari” and said to have come from Gambia. As suggested by Hooker in the protologue, it is probable that the plant is from South America, possibly Guyana or the Amazon River basin of Brazil where Strychnos toxifera, the source of the drug curare is native and was known to occur at that time. Hooker named the plant in honor of his friend, Dr. Moore. Fevillea moorei is most similar to F. bahiensis of southern Bahia, Brazil. It is easily distinguished by its larger staminate flowers, its leaf margins lacking uncinate trichomes, and its 3-nerved leaves which dry toa light green color. Fevillea pedatifolia (Cogn.) C. Jeffrey, Kew Bull. 16:199 1962. (Fig. 3). Alsomitra pedatifolia Cogn., in Martius, Fl. Bras. 6(4):116. 1878. Siolmatra pedatifolia (Cogn.) Cogn., in Engler, Pflanzenr. 4(Heft 66):30. 1916. Type: BRAZIL. AMAZONAS: Rio Solimées, 1819-1820, Martius s.n. (HOLOTYPE: M; photo ex M: F MO, NY, US; IsotyPes: BR-fragment). Alsomitra peruviana Huber, Bol. Mus. Paraense Hist. Nat. 4:616. 1908. Siolmatra ae ees (Hur ber) Cogn., in Engler, Pflanzenr. 4(Heft 66):30. 1916. Fevillea peruviana (H Kew Bull. 16:200. 1962. Type: PERU. UCAYALE Rio Ucayali, near Concha 28 Oct 1898, Huber 1390 (HOLOTYPE: MG, t n.v.; photo ex MG: F; ISOTYPE: F-fragment). Siolmatra amazonica Cogn., in Engler, Pflanzenr. 4(Heft 66):30. 1916. le ama zonica (Cogn.) C. Jeffrey, Kew Bull. 16:199. 1962. Type: BRAZIL. AMAZONAS: Rio Jurua, Jurua Miry, Oct 1901, Ute: 5819 (LECTOTYPE: here sae K; ISOLECTOTYPES: F-fragment, MG, n.v; photo ex MG: F; 5 eens ae ex B: F MO, NY. Notiz i Bot Gart. Berlin-Dahlem 11:769. 1933. Fevillea simplicifolia (Harms Cc Jeff frey, Kew Bull. 16:200. 1962. Type: PERU. LORETO: Lower Rio Huallaga, Yurimaguas, Puerto Arturo, 15 Nov 1929, Williams 5072 eee B, destroyed; LECTOTYPE: here designated, F; ISOLECTOTYPE: US). = z Vine or liana; stem glabrous to densely glandular-pubescent,; tendrils sparsely pubescent or glabrous. Leaves with the blade unlobed, or 3- to 5-lobed, or 3- to 5-foliolate, membranaceous to coriaceous, the unlobed or the lobed leaves ovate to ovate-oblong, (6-)8-15(-18) cm long, 9-14(-17.5) cm wide, 5- to 7-nerved, the divisions of the lobed leaves or the leaflets of the foliolate leaves (6-)8-15(-18) cm long, +-8 cm wide, l- to 2-nerved, with a petiolule up to 2.cm long, the apex of the blade or leaf divisions acuminate, the base of the blade or leaf divisions oblique or rounded, the margin entire or coarsely crenate-toothed, the primary lateral veins terminating in a small irregularly shaped marginal gland, the up- per and lower surfaces glabrous or lightly pubescent, especially along the leaf veins, the petiole 3.5-5(-7) cm long, glabrous to sparsely pubescent, with 2 op- posite, prominent, irregularly ae median to subapical glands. Staminate flowers in a paniculate inflorescence on reduced lateral branches, each branch subtended by a thin scale-like glandular-pubescent bract 0.5-2 mm long; pedicel 0.5-1 mm long, sparsely glandular-pubescent to glabrate; hypanthium 0.5-1.5 mm long, sparsely glandular-pubescent to glabrate; calyx shallowly cupular, the lobes light green or greenish brown, suborbicular, 0.5-1 mm long and wide, the apex rounded or obtuse, glandular-pubescent or glabrous, the margin slightly erose and glandular-ciliate, with a glandular calycine squamella 1986 BRIT.ORG/SIDA 21(4) Fic. 3. Fevillea pedatifolia. A. Habit, staminate inflorescence (Klug 4787). B. Staminate flower (Klug 2678). C. Fruit (Barbour 5533). D. Seed (Barbour 5533). protruding at or near the point of fusion with the petals; petals suborbicular, |- 2mm long, l-1.5 mm wide, white or greenish white, greenish yellow, or yellow- ish, the margin slightly erose, the median adaxial ridge slightly raised; stamens 0.25- 0.5 mm long, the anthers ca. 0.25 mm long, slightly longer than wide, the filament ca. 0.5 mm long. Pistillate flowers in pairs; pedicel 1.2-2 mm long; ca- lyx deeply cupular, the lobes ca. 3mm long, ca. 2mm wide, dark greenish brown, the surface pustulate; petals white. Fruit subglobose, ca. 14 cm long, 13cm wide, gourd-like, indehiscent or rarely circumscissile dehiscent along the hy- panthium lip sear (A. Gentry, pers, comm.), the surface smooth or pustulate, usually a mottled green color; seeds orbicular, compressed, 4-5 cm long and om ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1987 wide, 1.5-2.5 cm thick, silver-white upon drying, the woody layer smooth to slightly pustulate, the winged margin ca. 0.5 cm wide. Distribution and ecology.—Amazonian Ecuador and Peru, south to Bolivia, and east to Acre and Amazonas, Brazil. A plant of moist to wet forests, occur- paca at 130-800 meters in elevation. Sel ined. ECUADOR. Napo: Estacion Bioldgica Jatun Sacha, 8 km E of Misahualli, 01°04'S, 77° 36'W, 400 m 23-31 Jan 1989, Cerén 6076 (MO); km 2, new road from Cotundo to Coca 1130 m, 5 Aug 1984, Dodson et al. 15072 (NY). PERU. Amazonas: | km an La Poza, W of Rio Santiago 180 m, 21 Aug 1979, Huashikat 111(MO), va i of Rio Santiago, Quebrada Caterpiza, 2-3 km behind the community of Caterpiza, 3°50'S, 77°40'W, 200 m, 8 Feb 1980, nna Oe (MO). ic heapuaael be tween Santa Rosa and Hacienda Luisiana, 640 m, 9 Sep 1976, Wasshaus Y) Huanuco: Codo de Puzuzo, floodplain of Rio Puzuzo, S of settlement to main river, 9°40'S, 75°25 W 450 m, 21 Oct 1982, Foster 9370 (USF). Junin: Rio Negro, 800 m, 14 Aug 1960, Woytkowski 5795 (G, MO, US). Loreto: Yanamono, Explorama Tourist Camp on Rio Amazonas between Indiana and mouth of Rio Napo, 03°28'S, 72°48'W, 120 m, 26 Jul 1980, Gentry et al. 29043 (MO); Yanamono, Explorama Tourist Camp, Rio Amazonas half way Hoe een Indiana and aes of Rio Napo, 03°28'S, 72°50'W, m, 13 Jul 1983, Gentry et al. 42937A (MO); Balsapuerto, 22 Feb 1933, Klug 2896 (BM, EG, MO, Y, US); Indiana, Reserva cope aap a 03°30'S, ee W, 90 m, 28 Sep 1990, Pipoly et al es (MO), Yanamono tourist camp, 50 mi. ae ae He 72°50',ca. 106 m, 19 Oct 1980, Vasquez & Jaramillo 586 (NY); 1 ae Iquique, 03°30! °58'W, 115 m, 16 Dec 1987, Vasquez & Jaramillo 10164 (MO), Indiana, Explorama Reserve, 03° i : a 50'W, 7 m, 9 Nov 1989, Vasquez & Jaramillo 13140 (MO). Madre de Dios: Tambopata, 12°50'S, 69°17'W, 260 m, 8 Nov 1988, Alexiades et al. 81 (MO, NY): Rio Piedras, near confluence with Rio Pariamanu, 12°40'S, 69°17'W, 260 m, 1] Jan 1991, Alexiades et al. 1071 (NY); Lago Tres Chimbadas, ca. 65-70 river km SSW of Puerto Maldonado, ca. 10-15 air km NW effluence of Rio La Torre (Rio D’Orbigny) Rio Tambopata, 12°49'S, 69°17'W, ca. 260 m, 7 Jun nee eee 5533 (MO); Rio Manu, Cocha Cashu Biologial Station, 350 m, 1 Oct 1980, Foster 5439 (N a Cashu, vicinity of ox-bow lake of Rio Manu, between Panaqua and Tayakome, 17-24 ae ie ph et al. 3492 (K, MO, US). Las Piedras, Cusco Amazonico, 12°29'S, 69°03'W, 200 m, 24 Jun 1991, Timand 1844 (MO). Pasco: Rio Pichis, | hr. below Puerto Bermudez, between Puerto Bermedez and Paujil, 10°10'S, 74°50'W, 200 m, 17 Jun 1983, Gentry et al. 42148 (MO), Iscozacin, near confluence of Rio Palcazu and Rio Iscozacin, 10°12’, 75°13'W, 430 m, 16 Jun 1982, Smith 1911 ree Palcazu Valley, Rio Chuchurras drainage, 10°09'S, 75°20'W, 400 m, 13 May 1983, Smith 4009 (MO, US). San Martin: Pongo de Cainarachi, Rio Cainarachi, tributary of Rio Huallaga, 230 m, Sep-Oct = Klug 2755 (BM, FG, K, MO, N Eee Quebrada Cachiyacu de Huaquisha, ca. 5300-650 m, 7 Dec 1980, Schunke 12451 (NY). Ucayali: Bosque Nacional von Humboldt, Pucallpa-Tingo Maria Road, 8°40'S, 75°O'W, 250 m, rn Feb 1981, sae et al. 31307 (MO, USF). BRAZIL. Acre: N bank of Rio Jurua opposite Cruzeiro do 27 Oct 1966, Prance et al. 2936 (K, MO, NY, US). Amazonas: near Palmares, 11 Sep 26-Oct 1936, Kru 2 8475 (BM, BR, EG, K, MO, NY, US). BOLIVIA. La Paz: basin of Rio Bopi, Asunta (near Evenay), 690- 750 m, 27-31 Jul 1939, Krukoff 10666 (FG, K, MO, NY) Plants with 3- to 7-foliolate leaves are easily distinguished from other species of Fevillea. They previously had been considered a species of Siolmatra because of this feature, but the presence of the foliar glands, the bilocular anthers, and the large globose fruit with large unwinged seeds clearly separates it. Siolmatra, in contrast, lacks foliar glands, has unilocular anthers, and has a cylindric fruit with winged seeds. Fevillea peruviana and F. amazonica were separated from F. pedatifolia on the basis of leaflet number (3 rather than 5). Fevillea amazonica was further 1988 BRIT.ORG/SIDA 21(4) separated on the basis of the petiolar gland position (median rather than sub- apical). These characters are not constant and thus F. peruviana and F. amazonica are here reduced to synonymy as suggested by Jeffrey (1962b). Plants with unlobed leaves have been called F. simplicifolia. Since speci- mens with leaves intermediate between simple and 3-foliolate are occasionally found (although rare) and there are otherwise no other floral or vegetative dif- ferences, F. simplicifolia is here reduced tosynonomy. Plants with unlobed leaves are sometimes confused with F. pergamentacea. However that species is readily distinguished by the presence of a pair of glands at the base of the blade, while Me the glands in F. pedatifolia are on the petiole. Fevillea pergamentacea (Kuntze) Cogn., in Engler, Pflanzenr. 4(Heft 66):8. 1916. (Fi g. 4). Nhandiroba pergamentacea Kuntze, Revis. Gen. Pl. 3(2):104. 1898. Type: BOLIVIA. ae CRUZ: Rio Yapacani, 400 m, Jun 1892, Kuntze s.n. (LECTOTYPE: here designated, NY: ISOLECTOTYPES: NY, B, destroyed: photo ex B: F MO, NY, US). Nhandiroba harmsii Kuntze, Revis. Gen. Pl. 3(2):103. 1898. Fevillea harmsii (Kuntze) kK. Schum., Just’s Bot. Jahresber. 20:383. 1898. TypF: BOLIVIA. SANTA CRUZ: Velasco, 200 m, Jul 1892, Kuntze sn. (LECTOTYPE: here designated, NY, ISOLECTOTYPEs: US, B, destroyed; photo ex B; F MO, NY, US). Vine or liana; stem glabrous to lightly pubescent; tendrils glabrous. Leaves with the blade unlobed or occasionally 2- to 3- lobed, ovate, 1O-11(-15) cm long, (4-) 6-8(-15) cm wide, subcoriaceous, drying very light brown or green, the apex or the lobe tips acuminate or acute to slightly rounded, the base cordate to trun- cate, the upper surface glabrous, the lower surface lightly pubescent along the veins, the margin entire, occasionally with scattered trichomes, the lateral veins occasionally terminating with a small gland, with 2 small, opposite, irregu- larly auriculate glands at the base of the blade, the petiole 2.5-4 cm long, gla- brous to sparsely pubescent. Staminate flowers in a paniculate inflorescence on reduced lateral branches; pedicel 1-2 mm long, densely to sparsely pubes- cent; hypanthium 0.5-1 mm long, glabrous; calyx crateriform, the lobes lan- ceolate, 0.5-1 mm long, ca. 0.5 mm wide, obtuse, slightly erose-margined, sparsely to densely pubescent, witha prominent glandular calycine squamella protruding from each ae at or near the point of fusion with the petals; petals narrowly obovate, 2-2.5 mm long, 15-2 mm wide, cream to yellowish brown, the margin entire, the median appendage slightly raised; stamens ca. 0.5 mm long, the anthers ca. 0.25 mm wide, slightly longer than wide. Pistillate flowers not seen. Fruit globose, 6-7 cm long and wide, green, the surface minutely pus- tulate; seeds orbicular, compressed, 1.5 cm thick, 3.5-4.5 cm long and wide, the lateral surface of the woody layer smooth to minutely pustulate, the marginal nerve ca. 0.5 cm wide, completely enclosing the seed. Distribution and ecology.—Amazonian Colombia, Ecuador, eastern Peru, and central Bolivia. A plant of wet forests, occurring between 180 and 500 meters. ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1989 Fic. 4. Fevillea | t A. Habit i infl (Klug 2198). B. Staminate flower (Klug 2798). Additional specimens examined. COLOMBIA. Guaviare: Vereda Miraflores, 02°19'N, 72°26'W, 300 m, 27 Jan 1990, Marulanda & Marquez 1766 (MO). Putamayo: Vereda “La Kofania,” 1°1'N, 77°17'W, 500- 700 m, 2 Sept 1993, Cogollo et al. 6839 (MO). ECUADOR. Napo: Yasuni Forest Reserve, along road from PUCE Scientific Station to end of ae towards Waoroni Territory, 00°40.901'S, 76°24.348'W, 240- 310 m, 29 Jun 1995, Acevedo & Cedeno 7571 (NY); Rio Eno NE of coe ca. 00° 10'S, 76°40'W, 300 m, I] Apr 1982, Balslev 2321 (NY); Estacion Biologica Jatun Sacha, 8 km E of Misahualli, 01°04'S, 77°30'W, 400 m, 23-31 Jan 1989, Cerén 6076 (MO, NY); Estacion se Sacha, Rio Napo, 8 km E of Misahualli, 01°04'S, 77°36'W, 400 m, 10 Aug 1989, Cerén 7379 (MO, NY); Hollin-Loreto high- way, between Avila and Loreto, Huiruno (Quichua community), 00°43'S, 77°L9'W, 450 m, 29 Nov 1982, Cerén 7853 (MO); km 2, new Cotundo-Coca highway, 1130 m, 5 Aug 1984, Dodson et al. 15072 (MO ae 7 Pompeya, S of Rio Napo, Rio Jivino, Maxus highway, km 1-5, 00°25'S, 76°37'W, 220 m, 23-29 2, Grijalva et al. 231B (MO), E of the mission in Ahuano on Rio Napo, 550 m, 15 Feb 1973, ee 6214 (F, MO); Yasuni National Park, Rio Indillama, small southern tributary of Rio Napo, Comuna Pompeya, 00°30'S, 76°40'W, 220 m, 26 Aug 1992, Neill & Gudino 10120(MO, NY); S of Volcan Sumaco, Hollin-Loreto highway, km 31,Comuna Challua Yacu, 00°43'S, 77°40'W, 1200 m, 20-25 Mar 1989, Palacios 4102 (MO); Rio Aguarico, Shushufindi, 244 m, 14 Feb 1975, Vickers 116 (F). PERU. Loreto: 1990 BRIT.ORG/SIDA 21(4) Florida, Rio Putamayo, at mouth of Rio Yubineto, 180 m, May-Jul 1931, Klug 2198 (BM, E.G, K, MO, NY, US). BOLIVIA. Cochabamba: Cochabamba, Bang 1264 (BM, kK, MO). El Beni: Trinidad, 164 km towards Santa Cruz, 28 Aug 1985, Beck 12212 (NY). Santa Cruz: kim 11, Rio res Suet adas road, 14 Aug 1991, Acevedo et al. 4598 (NY); border of Rio Pirai, along Montero-Por , Mostacedo 2288 (NY); 12. km SE of Comunidad Don Lorenzo, 0.5 km E of Estancia Caracore, W side of Rio Grande, I7°51'S, 62°47'W, 300 m, 17 Nov 1990, Nee 39986 (MO, NY); Parque eee Ambor6, 0-2 km SW ol El Carmen, along trail to Quebrada Yapoje and Rio Saguayo, 17°32'S, 63°42'W, 360 m, 15 Jun 1991, Nee 41079 (NY); Parque Nacional Ambor6, l-2 km NE of El Carmen on all to crossing of Rio Surutu, I7°31'S, 63°4'W, 350 m, 21 Jul 1991, Nee 41796 (MO, NY); along Quebrada Salada and oil pipeline, first ae foothills of the Andes, 4 km W of highway bridge over Rio Pirai, 18°O6'S, 63°30'W, 750 m, 1] Dec L991, Nee 42017 (MO, NY); 2 km (by air) NW of center of Bermejo, around Laguna Volcan, 18°07'S, 63°39'W, 1125-1175 m, 24 Dec 1994, Nee 46129 (MO, NY); Parque Nacional Ambor6, along Rio Verde, Lkm E of Campamento Mataract, 17°33'S, 63°52'W, 375m, 31 May 1998, Nee & Bohs 49540 (NY); 4 km (by air) NE of Bermejo, valley of “Rufugios Los Volcanes,” 18°06.3'S, 63°36'W, 1070-1150 m, 29 Jul 2003, Nee 52385 (NY); Estancia San Rafael de Amboro, 15 km SSE of Buena Vista, 17°35'S, 63°37'W, 375 m, 30 Jul 1987, Nee & Saldias 35454 (NY, USF); Parque Nacional Ambor6, 0.5 km upstream from the last Andean foothills, 17°40'S, 63°43'30"W, 450 m, 20 Dec 1988, Nee & Saldias 37265 (NY), Rio Piray, 450 m, 19 Jul 1924, Steinbach 6259 (G, K); Parque Nacional Amboro, before Rio Saguayo, 5 km SW of El Carmen, ca. 17°33'S, 63°44'W, 400 m, 15-22 Nov 1991, Vargas et al. 1225 (MO, NY); Reserva de Vida Silvestre Rios Blanco y Negro, Rio Negro de Caimanes, ca. 4 km SE of Rio San Pablo, 14°45'07'S, 63°56'W, 250 m, 18 Jun 1993, Vargas 2605 (MO, NY); Reserva de Vida Silvestre Rios Blanco y Negro, 5 km from meeting of Rios San Pablo and Negro de Caimanes, 14°48'05"S, 63°58'W, 200 m, 18-20 Jun 1993, Vargas 2623 (NY) Fevillea pergamentacea is most similar to simple-leaved forms of F. pedatifolia, but differs in the position and form of its foliar glands and in the shape of the staminate flowers. Fevillea pergamentacea is characterized by flat, eae elliptic glands at the base of the leaf blade rather than petiolar glands well be- low the base of the leaf blade as in F. pedatifolia. Fevillea pergamentacea also has a distinctive crateriform staminate flower as opposed to the shallow cup- shaped flowers of F. pedatifolia. The distinction between F. pergamentacea and F. harmsii on the basis of the leaf shape cannot be maintained and F. harmsii is here reduced to synonymy as suggested by Jeffrey (1962b). Fevillea trilobata |... Sp. Pl. 1014. 1753. Fig. 5). Fevillea dia ats Hist. Nat. Drog. Simpl. ed. 4. 3:244. 1850, nom. illegit. Nhandiroba trilobata ( e, Revis, Gen. Pl. 1:257. 1891, Type: BRAZIL: (LECTOTYPE: here designated, Marcgrave, in | nero een Hist, Nat. 16048). iy Bras. 2:46, Seana pl, Fevillea eis id Vell, Fl. Flumin., Icon. 10:t. 102. 1831 1827”); non L., 1753. Fevillea triangu- ., Fam. Nat. Syn. Monogr. 2:114. 1846. TYPE: BRAZIL. RIO DE JANEFIRO: (LECTOTYPE: ere ears Vellozo, FI. I Fiend, Icon. 10:t. 102. 1831 61827")). Hypanthera Suapeva Silva Manso, Enum. Subst. Braz. 38. 1836. Type: BRAZIL. SAO PAULO: Sao Ignacio (iy. = Fevillea tomentosa Gardner, London J. Bot. 2:355. 1843. Fevillea saree var. tomentosa (Gard- ner) Cogn.,in Martius, FL. Bras. 6(4):118. 1878. Type: BRAZIL. RIO DI : Serra dos Orgaos, s.d., Gardner 425 (HOLOTYPE: BM; ISOTYPES: G, K; photo ex G: F MC ), US ). Fevillea albiflora Cogn., in Martius, FL. Bras. 6(4):118. 1878. Nhandiroba albiflora (Cogn.) Kuntze, Revis. Gen. PI. 1:257. 1891. Type: BRAZIL. BAHIA: without precise local yy s.d., Blanchet 2380 (LECTOTYPE: here designated, G; ISOLECTOTYPFS: BR, G, K, NY). SYNTYPE MATERIAL: BRAZIL. ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1991 Fic.5. Fevillea trilobata. A. Habit i infl (Saint-Hilaire s.n. [Glaziou 87201}).B.$ i fl (Sai Hilaire s.n. [Glaziou 8720)). 1992 BRIT.ORG/SIDA 21(4) MINAS GERAIS: near Contendas, 1818, Martius 1581 (M): near Contendas, s.d., Saint-Hilaire s.n. (BR, F). Fevillea albiflora var. glaziovii Cogn., Bull. Acad Roy. Sci. Belgique, ser. 3. 14:364. 1887. TyPF: BRA- ZIL. MINAS GERAIS: Serra do Inficionado, near Caracas, 6 Sep 1882, Glaziou 13906 (LECTOTYPE: here designated, BR; ISOLECTOTYPES: B, destroyed, K, NY; photo ex B; F MO, NY, US). Fevillea trilobata var. longipedicellata Cogn., in Martius, FI. Bras. 6(4):118. 1878. Type: BRAZIL. RIO DE JANEIRO: near Copacabana, s.d., Riedel {et Langsdorff, fide Cogn. (1881)] 706 (LECTOTYPE: here designated, BR; ISOLECTOTYPES: K, M). SYNTYPE MATERIAL: BRAZIL. RIO DE JANEIRO: Copacabana, Dec 1840, Gardner 5469 (BM), Copacabana, Lund 213 (nv): s.d., Sello s.n. (BM). Fevillea trilobata var. subintegrifolia Cogn., in Martius, Fl. Bras. 6(4):118. 1878. Type: BRAZIL, RIO DE JANEIRO: Copacabana, s.d., Luschnath 552 (LECTOTYPE: here designated: BR). SYNTYPE MATE- RIAL: BRAZIL. RIO DE JANEIRO: Copacabana, Luschnath 950 (n.v.); Blanchet 955 (nw). lea trilobata var. subuniflora Cogn., in Martius, Fl. Bras. 6(4):118. TYPE: BRAZIL. RIO DE JAN- Fevi EIRO: Gavea, Glaziou 3986 (n.w.). Vine or liana; stem densely pubescent, tomentose to lightly pubescent, or subglabrous; tendrils pubescent or glabrous. Leaves with the blade 3-lobed, or occasionally 5-lobed, 6-10 cm long, (2-)5-10(-15) cm wide, membranaceous, the apex acute or acuminate, the main lateral veins occasionally terminating inasmall apical gland, the upper and lower surfaces densely to lightly pubes- cent. Staminate flowers in a paniculate inflorescence on reduced lateral branches; pedicel with uncinate trichomes, 1-3(5) mm long; hypanthium (2-) 3-5 mm long, densely to sparsely stipitate-glandular pubescent; bracts glan- dular, linear, 0.5-1 mm long, densely to sparsely pubescent, the ones subtending the panicle branches, larger, less glandular, more leaf-like, often witha distinct stipe; calyx shallowly cupular, the lobes slender, ca. 2.5mm long, ca. 1 mm wide, obtuse, densely to sparsely glandular-pubescent with a glandular calycine quamella protruding from each sepal at or near the point of fusion with the etals; petals suborbicular, 3-6 mm long, 1-4 mm wide, pale yellow, cream- olored or pink, each with a median adaxial uncinate appendage; stamens ca. mm long, the anthers ca. 0.5 mm long, slightly longer than wide. Pistillate lowers (fide Cogniaux 1878, 1881, 1916) 1-3; pedicel ca. 6 mm long, densely to sparsely pubescent; hypanthium ca. 2 mm long, densely to sparsely pubescent: calyx campanulate, the lobes with a glandular calycine squamella at the point of fusion between the sepals and the petals; petals strap-shaped, broadened at the base, cream-colored; styles 3; ovary 5-7 mm long. Fruit (fide Cogniaux 1878, 1881, 1916) subglobose, reddish brown, 7-9 cm wide, pubescent or lightly glandular-pubescent, the locules 4-seeded; seeds orbicular, compressed, ca. 4 cm long and wide, ca. 1 cm thick, the lateral surface of the woody layer striate- verrucose, the outer edge tuberculate. Distribution and ecology.—Brazil from Ceara southwest to Minas Gerais and — some >ro Sao Paulo. Selected specimens examined. BRAZIL. Bahia: [|heus, 1836, Blanchet 2380 (G, F K, MO, NY, US); km 22 on Ilheus/Itabuna road (BR 415), near Palmoreto, 50 m, 1 Sep 1997, Jardim et al. LIO] (NY). Ceara: Serra do Araripe, Sep 1828, Gardner 1629 (BM, K). Espirito Santo: Reserva Florestal de Sooretama, 9 — ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1993 Aug 1965, Belém 1499 (CEPEC, NY); Reserva Florestal da C.V.R.D., Linhares, Estrada Farinha Seca, on 221, km 1100, 17 Jul 1986, Folli 130 (NY); Reserva Florestal da C.V.R.D., Linhares, Estrada Farinha Seca, ant. 221, km 1100, 7 Aug 1990, Folli 373 (NY); Reserva Florestal da C.V.R.D., Linhares, 28 Oct 1992, Folli 1715 (NY). Goias: Formosa, 1814-1817, Bowie & Cunningham 913 (BM). Minas Gerais: near Vicosa, 8 km from Fazenda de A. Cocerro toward Sao Miguel, 900 m, 26 Jun 1930, Mexia 4798 (BM, F, G, MO, NY, US). Paraiba: Cidade Universitaria, 6 km SE of Centro de Joao Pessoa, 7°57'S, 34°53'W, 30- 45 m, 10 Aug 1990, Agra 1165 (MO). Parana: Barra do Turvo, 10 Oct 1973, Hatschbach 32631 (K, MO, NY); Barra do Lagead de, 31 Aug 1978, Hatschbach 41560 (NY), along road to landing of Porto Byington on bank of Rio Parana, 200 m, 26 Jun 1966, Lindeman & de Haas 1778 (NY); Parque Estadual Vila Rica do Espirito Santo, 18 Feb 1998, Silva s.n. (NY). Pernambuco: Tapera, border of Rio Tapacura, Oct 1933, Pickel 524 (US). Rio de Janeiro: Gavea, s.d., Glaziou 1344 (BR, US); Serra dos Orgaos, Alto Macabé, s.d., Glaziou 17010 (NY). Sao Paulo: Campinas, 27 Aug 1938, Krug 1593 (US) Fevillea trilobata is most similar to F cordifolia from which it can be distin- guished by its usually angular (vs. rounded) leaf lobes and staminate flowers with stipitate-glandular trichomes on the hypanthium which are sharply dif- ferentiated from the uncinate ones on the pedicel. Fevillea subgenus Ani (Silva Manso) G. Robinson & Wunderlin, comb. et stat. nov. BASIONY m: Anisosperma Silva Manso, Enum. Subst. Braz. 38. 1836. TYPE: Anisosperma passiflora (Vell.) Silva Manso [=Fevillea passiflora Vell. Staminate inflorescences of congested panicles or subumbelliform; glandular calycine squamellae absent; corolla lobes oblong-hastate with a thick glandular adaxial ridge. Fruit ovate or oblong, subtrigonous, not zonate the apex apiculate. Fevillea passiflora Vell., Fl. Flumin., Icon. 10:t. 104. 1831 1827”). (Fig. 6). Anisosperma passiflora Vell.) Silva Manso, Enum. Subst. Braz. 38.1836. TYPE: BRAZIL. RIO DE JANEIRO: (LECTOTYPE: here designated, Vellozo, Fl. Flumin., Icon. 1O:t. 104. 1831 1827"). Vine or liana; stem glabrous or sparsely glandular-pubescent; tendrils glabrous. Leaves with the blade unlobed, ovate to broadly lanceolate, 5-13 cm long, 4.5- 75cm wide, membranaceous, with a distinct marginal nerve, the apex acute to acuminate, the base rounded, with 2 small irregu petiole, the upper and lower surfaces minutely pustulate, sometimes sparsely glandular-pubescent, the petiole 1-3 cm long. Staminate flowers in a congested panicle or subumbelliform inflorescence; pedicel (2-)4-5 mm long, sparsely glandular-pubescent; hypanthium 4-5(-13) mm long, sparsely glandular-pu- bescent; calyx shallowly cupular, glandular-pubescent, the lobes ca. 2.5 mm long, lanceolate, acute; petals oblong-hastate, 5-7 mm long, green or cream-col- ored, the median adaxial ridge glandular, slightly raised, broadest at the base, gradually tapering to about the center; stamens 1.5-2 mm long, the anthers ca. 1 mm long, slightly longer than wide. Pistillate flowers 2-4 on a short, thick pe- duncle; styles with the lower part somewhat thickened, the upper part dilated and broadly obcordate, curved inward longitudinally; ovary 6-9 mm long and 4-5 mm wide, glabrous. Fruit ovoid to oblong, 8-15 cm long, 5-11 cm wide, subtrigonous, smooth or irregularly verrucose, fleshy with thickened outer —_— LAT, auric ulate glands near the 1994 BRIT.ORG/SIDA 21(4) infl (Hatschbach 19205).B. Staminate flower (Hatschbach 9106 => ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1995 cortex, the apex short-a ple ulate See dssuborbiculat , compressed 3.5-3.5¢m long, 3-4 cm wide, ca. 1.5cm thick, pale brown, witha thin erose-margined cover. Distribution and ecology—Coastal Brazil in southeastern Bahia, Rio de Ja- neiro, eastern Sao Paulo, and eastern Parana. In gallery forests near sea level. Additional specimens examined. BRAZIL. Bahia: ca. 5 km W of Itamaraju, 20 Sep 1878, Mori et a L. 10742 (CEPEC). P 4: Guaratuba, Rio Cubatao, 27 Dec 1911, Dusén 13640(F NY); highway between Rio Branco do Sul and Cerro Azul along Rio P. Grossa, 26 Apr 1962, Hatschbach 9106 (US); Ribeirinha, 7 May 1968, Hatschbach 19205 (kK); Morro do Inglés, 18 Feb 1976, Hatschbach 38090 (K). Sao Paulo: Campinas, 1873, Corréa de Mello s.n. (BR); Cabital, 26 Dec 1941, Pickel 5566 (US), 1816-1821, Saint- Hilaire D600 (F). Rio de Janeiro: 1894, Glaziou 20335 (BR); Nov 1987, Glaziou 10870 (K); Cantagalo, 1859, Peckolt 8 (BR); Cantagalo, 1861, Peckolt s.n. (BR); TeresOpolis, Serra dos Orgaos, 27 Feb 1887, Schenck 2898 (BR). Without precise locality: 1859, Peckolt s.n. (BR): s.d., Martius s.n. (BR). EXCLUDED NAMES Fevillea deltoidea Cogn., in Martius, Fl. Bras. 6(4):119. 1878. [=Pteropepon deltoideus Cogn.) C .) Cogn]. Fevillea monosperma Vell. Fl. Flumin., Icon. 10:t. 103. 1831 (“1827”). [-Pteropepon monospermus (Vell.) Cogn]. Fevillea pedata Smith ex Sims, Bot. Mag. t. 2681. 1826. [-Telfairia pedata (Smith ex Sims) Hooker. Fevillea puneiate (L.) Poir, in Lamarck, Encycl. 4:418. 1796. BasionyM: Bryonia punctata L., Demonstr. Pl. 26.1753. TyPF: Herb. Linn. No. 1180.1 (two sheets), apparently a mixed collection, possibly a Trichosanthes and a Momordice Fevillea tamnifolia Kunth, in Humboldt, Bonpland é¢ Kunth, Nov. Gen. Sp. 7:175. 1825. [=Sicydium tamnifolium (Kunth) Cogn. Fevillea trilobata Reichard, Syst. Pl. 4:253. 1780, nom. illegit.; non L., 1753. (BASIONYM: Bryonia punctata L., 1753.) [See Fevillea punctata above|] ACKNOWLEDGMENTS We gratefully acknowledge the curators of BM, BR, CEPEC, F G, K, M, MO, NY, US, and USF for making specimens available to us for study. We thank Bruce Hansen (USF) for his many helpful suggestions on the manuscript and Kath- leen Hotchkiss (USF) for assistance with graphics. We also thank Michael Nee (NY) and an anonymous reviewer for their helpful comments. REFERENCES Apams, C.D. 1972. Flowering plants of Jamaica. The University Press, Glasgow. Baitton, H. 1885. La fleur femelle de |'Alsomitra brasiliensis. Bull. Mens. Soc. Linn. Paris 1: 457-458 Cocniaux, A. 1878. Cucurbitaceae: Zanonieae-Fevilleae. In: CFP. von Martius, ed. Flora Brasiliensis 6(4):114-122.MUnchen. Cocniaux, A. 1881. Cucurbitaceae: Zanonieae-Fevilleae. In: Alph. de Candolle & C. de Candolle, eds. Monographiae phanerogamarum. 3:925—946. G. Mason, Paris. Cocniaux, A. 1893. Genre Siolmatra H. Baill. et la tribe des Zanoniées. Bull. Herb. Boissier ].609-613. = 1996 BRIT.ORG/SIDA 21(4) Cooniaux, A. 1916. Cucurbitaceae: Zanonieae—Fevillieae. In: A. Engler, ed.Das Pflanzenreich 66, IV, fam. 275, |:3-41.Wilhelm Engelmann, Berlin. Dierere, J.V.A. 1976. Cucurbitaceae. Flora of Guatemala. Fieldiana, Bot. 24(11):306-395. Gentry, AH. and R.H. Wettacn. 1986. Fevillea—a new oil seed from Amazonian Peru. Econ. Bot. 40:177-185. Gunn, C.R.and J.V. Dennis. 1976.World guide to tropical drift seeds and fruits. Quadrangle/ The New York Times Book Co., New York. Guppy, H.B. 1917. Plants, seeds, and currents in the West Indies and Azores. Williams & Norgate, London. Haras, H. 1926.Cucurbitaceae.In J. Mildbraed, Plantae Tessmannianae Peruvianae Ill. Notizbl. Bot. Gart. Berlin-Dahlem 9:989-996. Harms, H. 1933. Cucurbitaceae americanae novae. Notizbl. Bot. Gart. Berlin-Dahlem 11: 769-776. JerFrey, C. 1962a, Notes on Cucurbitaceae, including a proposed new classification of the family. Kew Bull. 15:337-373. JerFrey, C. 1962b. Notes on some species of Fevillea L., Siolmatra Baill., and Pseudosicydium Harms (Cucurbitaceae) in the Amazon Basin. Kew Bull. 16:199-202, Jerrrey, C. 1978. Further notes on Cucurbitaceae: IV. Some New World taxa. Kew Bull. 33: 347-380. Kuntze, C. 1891-1898. Revisio generum plantarum. G.E. Schechert, Leipzig. Linotey, J.and T. Moore. 1870. The treasury of botany, ed.2. Longmans, Green & Co.,London. Linnaeus, C. 1753. Species plantarum. Impensis Laurentii Salvii, Stockholm. Morton, J.F.1981.Atlas of medicinal plants of Middle America—-Bahamas to Yucatan. Charles Thomas, Springfield, IL. Piso,G.and G. Marcarave. 1648. Historia naturalis Brasiliae. Leiden/Amsterdam, Roemer, MJ. 1846. Nnandirobaceae ["Nhandirobée"], in Familiarum naturalium regni vegetabilis synopses monographicae 2:111-118.Landes-Industrie-Comptior, Weimar. SeriNGe, N.C. 1828. Cucurbitaceae. In: A.P. de Candolle, ed. Prodromus systematis naturalis regni vegetabilis. 3:297-320.Treuttel & Wurtz, Paris. Siva Manso, A.L.P.ba.1836.Enumeracgao das substancias Brazileiras. lypographia Nacional, Rio de Janeiro. STANDLEY, PC. 1937. Cucurbitaceae. In: J.F. Macbride, ed. Flora of Peru. Publ. Field Mus. Nat. Hist., Bot. Ser. 13(6):321-383 Tuttoch, A.P. and L. Bercrer. 1979. Analysis of the conjugated trienoic acid containing oil from Fevillea trilobata by '’C nuclear magnetic resonance spectroscopy. Lipids 14: 996-1002. BROMUS HALLII (POACEAE), A NEW COMBINATION FOR CALIFORNIA, U.S.A.. AND TAXONOMIC NOTES ON BROMUS ORCUTTIANUS AND BROMUS GRANDIS Jeffery M. Saarela Paul M. Peterson i ee and Department of Botany UBC Botanical Garden and Centre for Plant Research National Museum of Natural History ea of British clan id Smithsonian Institution Vancouver, B.C., CANADA a DC 20013-7012, U.S.A. Pe eee eterson@si.edu ale nian National P. lt h—-Biodiversit Agia ture and Agri-Food Canada m. Saunders Building (#49) eee Experimental Farm Ottawa, Ontario, CANADA, K1A 0C6 cayouettej/@agr.gc.ca ABSTRACT {B g lis. B.orcuttianus, and B.orcuttianus var. hallii from western North Herbarium study of America has indicated that these taxa are saga sa and see ie distinct, and best Peterson is made. Bromus treated as species. A new combination B hallii differs from B. orcuttianus by having densely eat Pade (verses glabrous in B. orcuttianus), lower sheaths with dense short stiff hairs 1 mm long or less (vs. long soft hairs 2-4 mm long), and 1-2(-3) culm nodes (vs. 2-4). Bromus hallii differs from B. grandis by having blades that are 7.5-16.5 cm long [vs. (13-)18-38 cm long in B. grandis], 1-2(-3) culm nodes (vs. 3-7), and lower glumes that are l(-3)-nerved [vs. 3(1)-nerved]. We provide the first report of differences in the type of pubescence on the lower sheaths between B. orcuttianus and B. hallii. We confirm the diploid chro- mosome number of 2n = 14 for B. grandis. We include descriptions, synonymies, and representative specimens examined for B. grandis, B. hallii, and B. orcuttianus, a key to all sections of Bromus in California, and a key to Bromus sect. Bromopsis in California RESUMEN EA estlicio de Bromus grandis, B. genie: y B. Crean var, zat de Norteamérica occidental stan tratados mejor como especies disinies Se hace una nueva Poneeeaes Bromus hallii (Hitche) Saarela y P.M. Peterson. Bromus hallii se diferencia de B. orcuttianus por tener laminas densamente pupesceace oe a glabras en B. orcuttianus), vainas inferiores con los pelos rigidos cortos de 1 mm larg tra los pelos suaves largos de 2-4 mm), y 1 6 2(-3) nudos en el culmen (contra 2-4). Bromus hallii se diferencia de Bromus grandis por tener las laminas de 7.5-16.5 cm de largo (contra 18-38 cm en B. ee 1 6 2 nudos en el culmen (contra 3-6), y glumas inferiores I(-3)-nervadas [contra 3(1)- das]. Proporcionamos el primer informe de diferencias en el ee ee pupesccnce en las vainas nery eae B.orcuttianus y B. hallii. Confirmamos el nu I le 2n= para B. grandis. Incluimos descripciones, sinonimias, y especi I los d SIDA 21(4): 1997-2013. 2005 1998 BRIT.ORG/SIDA 21(4) B. grandis, B. hallii y B. orcuttianus, una clave de todas las secciones de Bromus en California, y una clave de Bromus sect. Bromopsis en California. Bromus L. is a large genus of approximately 160 species that are distributed worldwide in temperate regions. The genus is distinguished from other grass genera by the combination of leaf sheaths that are connate for most of their length, awns that are inserted subapically, and hairy appendages on the apices of the ovary (Clayton & Renvoize 1986). In the New World, 79 native and intro- duced species are currently recognized (Pavlick et al. 2003), although new spe- cies are being described (e.g, Saarela et al. in review) and the taxonomic bound- aries of several difficult species complexes have been reevaluated (e.g., Peterson etal. 2002; Oja et al. 2003; Massa et al. 2004). Molecular phylogenetic investiga- tions have clarified several aspects of the evolutionary history of Bromus, and have identified major lineages in the genus, some of which correspond to com- monly recognized intrageneric taxa (Pillay & Hilu 1990, 1995; Ainouche & Bayer 1997; Saarela et al. in press). A new classification of the genus has not yet been proposed, pending further study of morphological and molecular variation. Bromus sect. Bromopsis Dumort, the largest of the traditionally recognized sections in the genus, is a non-monophyletic taxon comprised of several lin- eages that occur in North America, Mexico, South America, and Eurasia (Saarela et al. in press). The section is characterized morphologically by 1(3)- and 3(5)- nerved first and second glumes, lemmas that are dorsally flattened, and a pe- rennial habit (Smith 1970), although some of these characteristics may be symplesiomor phies. In North America north of Mexico, there are 16 native spe- cies in section Bromopsis that have diversified in a variety of habitats (Pavlick 1995; Pavlick & Anderton, in press). Some species are wide-ranging (e.¢., B. ciliatus L.), whereas others are highly restricted in distribution le.g., B. texensis (Shear) Hitchce.]. Several taxa (B. laevipes Shear, B. orcuttianus Shear, B. orcuttianus var. hallii Hitche., B. pseudolaevipes Wagnon, and B. suksdorfii Vasey) are restricted to mountainous regions in the southwestern part of the continent, largely in California (Pavlick 1995). Others|[e.g., B. porteri J.M. Coult.) Nash, B. richardsonii Link, B. vulgaris Hook.) Shear] have broader distributions in western North America. While studying B s specimens from California to support preparation of the taxonomic treatment of Bromus for the Second Edition of the Jepson Manual (Saarela & Peterson, in prep.), it became clear that B.orcuttianus var. hallii, despite its varietal rank, is a readily identifiable taxon that can be consistently and reliably distinguished from both B. orcuttianus var. orcuttianus and B. grandis. These preliminary observations stimulated a — fou — more detailed investigation of its mor phological variation and taxonomic status. Vasey (1885) named and described B. orcuttianus based on collections by C.R. Orcutt from the mountains near San Diego, and collections by W.N. Suksdorf from Mt. Adams in Washington. In his revision of the North American SAARELA ET Al ERICA 1999 species of Bromus, Shear (1900) recognized B. orcuttianus, and described larger plants from La Maite, San Diego, with pubescent sheaths, leaves, culms, and spikelets asa new variety, B.orcuttianus var. grandis Shear. Hitchcock (in Jepson 1912) described plants from the mountains of southern California with densely pubescent blades and cataphylls, and pubescent glumes and lemmas as a new variety, B.orcuttianus var. hallii Hitche. Hitchcock (in Jepson 1912) also elevated B.orcuttianus var. grandis toa species, B. grandis (Shear) Hitchc., indicating that it is similar to B. orcuttianus var. hallii, but differs in its drooping panicle and distinctly 3-nerved lower glumes. Subsequent authors have recognized B. grandis, B. orcuttianus, and B. orcuttianus var. hallii in their treatments (e.g., Hitchcock & Chase 1951; Wagnon 1952; Munz & Keck 1959; Hitchcock et al. 1969; Munz 1974; Wilken & Painter 1993; Pavlick 1995; Pavlick @& Anderton, in press) although Clayton and Williamson (2002 onwards) synonymized B. orcuttianus var. hallii with B. orcuttianus. In his revision of Bromus sect. Bromopsis in North America, Wagnon (1952) hypothesized that B. orcuttianus var. hallii is equally related to B. grandis and B. orcuttianus, and indicated that it requires further study. Among species of Bromus sect. Bromopsis in North America, B. orcuttianus var. hallii is the only taxon recognized currently at the varietal level, even though the characteristics separating it from B. orcuttianus var. orcuttianus and B. grandis are similar in degree to those separating other morphologically similar species of Bromus. Here we discuss the morphologi- cal, cytological, geographical, and ecological characteristics of these three taxa, and propose a new combination, B. hallii (Hitchc.) Saarela & P.M. Peterson. We include synonymies, species descriptions, distributions, and specimens exam- ined for B. grandis, B. hallii,and B.orcuttianus, and keys to the sections of Bromus in California, and Bromus section Bromopsis in California. — MATERIAL AND METHODS This study is based on examination of over 425 herbarium specimens from CAS, DAO, K, OSU, UBC, UC/JEPS, US, W,and WTU, including most type specimens. Specimens were measured to generate morphological descriptions. Data on geo- graphic distribution, elevation, and habitat were obtained from herbarium specimen labels. Cytological observations of B. grandis were carried out by the third author on living material germinated from seeds collected in 1994 (Cayouette & Darbyshire C7947). Detailed procedures for the chromosome count are provided in Peterson et al. (2002). The taxonomic key was generated through study of the literature and representative herbarium specimens at US. RESULTS AND DISCUSSION — Bromus orcuttianus, B. orcuttianus var. hallii, and B. grandis are morphologi- cally similar taxa that have been recognized as distinct for almost a century. Examination of morphological variability among individuals from across the 2000 BRIT.ORG/SIDA 21(4) range of these three taxa indicates that they are each easily distinguished by a combination of quantitative and qualitative characteristics, and each has a dis- tinctive geographic distribution. The degree of difference among the three taxa is approximately the same, although one taxon is recognized currently at the varietal level while two are recognized as species. It would be equally logical (although not valid nomenclaturally) to classify individuals currently included in B. orcuttianus var. hallii as an intraspecific taxon of B. grandis, if emphasis were placed on vegetative rather than reproductive characteristics when mak- ing taxonomic decisions. To minimize such ambiguity, either one wide-rang- ing polymorphic species could be recognized, or each morphologically and geo- graphically distinct taxon could be treated as a species. We prefer the latter approach, and propose a new combination, B. hallii (Hitchc.) Saarela & P.M. — Peterson to accommodate this. This mor phological-geographical approach to circumscribing species follows existing treatments of Bromus sect. Bromopsis in North America, in which taxa with consistent morphological characteris- tics throughout their ranges are treated as species (e.g., Pavlick 1995; Pavlick & Anderton, in press; Saarela & Peterson, in prep). Bromus hallii and B. orcuttianus are distinguished morphologically by sev- eral qualitative characteristics, including the presence or absence of pubescence on the blades, the type of pubescence, and the number of culm nodes (Table 1). Bromus hallii is characterized by blades that are densely pubescent abaxially and adaxially, lower sheaths that are densely pubescent with short, stiff hairs up to | mm long, and culms with 1-2(-3) nodes. In contrast, B. orcuttianus has blades that are glabrous abaxially and adaxially (sometimes with pilose mar- gins towards the base), lower sheaths that are sparsely to densely pilose (occa- sionally glabrous) with long, soft hairs up to + mm long that are easily visible with the naked eye, and culms with 2-4 nodes. Previous authors (e.g., Hitch- cock 1951; Pavlick 1995) noted that these two taxa differ in the distribution of the pubescence on the leaves, but they were apparently not aware of the differ- ences noted here in the type of pubescence on the lower sheaths. Additional morphological characteristics distinguish B. hallii and B.orcuttianus, but there is some overlap in these character states. Bromus hallii consistently has pubes- cent glumes, lemmas, and blades that range in length from 7.5-16.5 cm long, whereas B.orcuttianus has glumes that are usually glabrous but sometimes sca- brous or pubescent, lemmas that range from glabrous to pubescent, and leaf blades that range in length from 7-24 cm long. The morphology of the inflo- rescence is the same in both taxa, with stiff branches that are erect and ap- pressed to spreading not more than 90° from the rachises. Good illustrations of the general habit and inflorescence morphology of B. orcuttianus and B. hallii are found in Pavlick (1995), but these do not adequately emphasize the diagnos- tic differences of the type of pubescence found on the lower leaf sheaths or the differences in culm node number. SAARELA ET Al 2001 Taste 1. Diagnostic characters separating Bromus grandis, B. hallii, and B. orcuttianus. Characters B. grandis B. halli B. orcuttianus Leaf blades pubescent pubescent glabrous, occasionally pilose near base Lower leaf densely pubescent, densely pubescent, sparsely to moderately sheaths hairs up to 1(-3) hairs up to 1 mm long pilose, hairs 2-4 mm m long long, occasionally glabrous Culm nodes 3-7 1=2(-3) 2-4 Leaf blade length (13)18-38 cm 7.5-16.5 cm 7-24 cM Glume surface pubescent pubescent glabrous, scabrous, or pubescent Lemma surface pubescent pubescent glabrous or pubescent Lower glumes, 3(1) 1(3) 1(3) number of veins Awn length 3-6 mM 3.5-7 mm (4-)5.5-8 mm Inflorescence flexuous, usually stiff, erect, ascending and _ stiff, erect, ascending branches spreading > 90° from appressed, to spreading and appressed, t culm axis not > 90° from culm axis spreading not > 90° rom culm axis Stebbins and Love (1941) reported a diploid chromosome count of 2n = 14 for B.orcuttianus, and this was confirmed by counts by Wagnon (1952). Stebbins and Love (1941) synonymized B. orcuttianus var. hallii with B.orcuttianus, thus it is unclear if they obtained counts for material that we are treating here as B. halli. Wagnon (1952) was unable to obtain living material of B. hallii, and at- tempts to germinate seeds of B. hallii for chromosome counts in this study were unsuccessful. The chromosome number of B. hallii remains unknown. Bromus halliiand B.orcuttianus differ in their geographic distributions and mean (1) elevations. Bromus halliiisa narrowly distributed species that occurs in southern California in the mountains of Kern, Fresno, Los Angeles, Monterey, Santa Barbara, San Bernardino, and Tulare counties, at high elevations (ut = 6658 ft; n= 29). Bromus orcuttianus has a much wider vanes occurring in the moun- tains throughout California, Oregon, southern Washington, eastern Nevada, and southeastern Arizona (Piper 1906: Wagnon 1952; Kearney et al. 1960; Pavlick 1995), generally at lower elevations (u = 5306 ft; n = 40). Bromus hallii and B. grandis are distinguished morphologically by a com- bination of vegetative and reproductive characteristics (Table 1). The distribu- tion and type of pubescence on the blades, sheaths, and spikelets is similar in B. hallii and B. grandis, although two specimens of B. grandis seen (Wolf 6888 and 2002 BRIT.ORG/SIDA 21(4) Silveus 2829) have longer hairs up to 3mm long on the blades and sheaths. The taxa differ in their blade lengths, number of culm nodes, number of nerves on the lower glume, and inflorescence morphology. Bromus hallii has blades that are 7.5-16.5 cm long [vs. (13-)18-38 cm long in B. grandis], 1-2(-3) culm nodes (vs. 3-7), and lower glumes that are 1(3)-nerved [vs. 30.)-nerved]. Bromus hallii has panicles with stiff branches that are erect or spreading not more than 90° from the culm axis, whereas B. grandis has panicles with flexuous branches that are spreading often more than 90° from the culm axis. In immature speci- mens of B. grandis (those collected in late May and early June), the panicles ap- pear similar in morphology to those found in B. orcuttianus and B. hallii, thus this characteristic should only be used on fully mature specimens. A good il- lustration of the inflorescence morphology of B. grandis is found in Hitchcock (1951). Wagnon (1952) indicated in his taxonomic keys that B. halli differs from B. grandis in having cucullate (boat-shaped) blade tips, but thorough examina- tion of this character indicates that several individuals of B. grandis share this character state, making this an unreliable character. Stebbins and Love (1941) reported a diploid chromosome number of 2n = |4 for B. grandis. Wagnon (1952) did not have fresh material of B. grandis for chromosome counts. Our cytologi- cal observations of B. grandis confirm the diploid number 2n = 14. Like B. hallii, B. grandis is a narrowly distributed species endemic to southern California, known from the mountains in the southern counties and in the coastal ranges from Santa Cruz to San Diego counties. Bromus grandis generally occurs at much lower elevations (ut = 3267 ft; n = 25) than B. hallii (ut = 6658 ft; n = 29). Recognition of B. hallii as a distinct species increases the number of spe- cies in Bromus section Bromopsis in California to 11, 10 of which are native, and one that is introduced (B. inermis Leyss.). Keys distinguishing the sections of Bromus and the species of sect. Bromopsis in California are presented below. A revised key distinguishing all 34 native and introduced species of Bromus in California will be published in the Second Edition of the Jepson Manual (Saarela and Peterson, in prep). TAXONOMIC TREATMENT Bromus grandis (Shear) Hitche., Fl. Calif. 1:175. 1912. Br cut r grandis Shear, Bull. Div. Agrostol., U.S.D.A. 23:43. 1900. nen a ean rai a Geobot Phytotax. 8(2):167. 1973. Type: U.S.A. CALIFORNIA. San Diego Co. La Maite, 25 Jun 1883, C.R. Orcutt 472 (HOLOTYPE: US-81613)) Bromus porteri var. assimilis Burtt Davy, Univ. Calif. Publ. Bot 1:55, 1902. TYPE: U.S.A. CALIFORNIA: south side, San Jacinto Mts, 1901, H.M. Hall 2228 (HOLOTYPE: UC-37692!: ISOTYPE: US-865453 fragm. ex UC!) Loosely caespitose perennial. Culms 70-180 cm tall, erect, glabrous to pubes- cent; nodes 3-7, dark brown, retrorsely pilose below, often covered by sheaths. Leaf sheaths 3.5-12 cm long, closed for most of their length; sheaths densely SAARELA ET AL 2003 pubescent, hairs up to | mm long; th, ionally hyaline at apex; collars densely pilose, with hairs up to 2 mm long, auricles sometimes present: cataphylls and basal sheaths pubescent, sometimes shredding; ligules 1-3 mm long, membranous, densely pubescent to pilose adaxially, glabrous abaxially, apex obtuse, erose; blades (13-)18-38 cm long, 3-12 mm wide, flat, membra- nous, apically cucullate, sparsely to densely pubescent adaxially and abaxially, hairs up to I(-3) mm long; margins glabrous, sparsely papillate or scabrous. Panicles 15-26 cm long, 6-17.5 cm wide, open; lower branches 5-26 cm long with l-6 spikelets, flexuous, nodding and usually spreading more than 90° from the rachises, pubescent; lower inflorescence nodes with 1-4 branches. Spike- lets 2.5-3.5(-4.5) cm long, 4-9-flowered, terete to conspicuously distichous at maturity with rachis visible; glumes subequal, shorter than lemmas, pubes- cent, margins sometimes hyaline; lower glumes 5-8.5 mm long, 3(.)-nerved, apex acute; upper glumes 7-10(-12) mm long, 3(5)-nerved, apex acute; lemmas L1-14 mm long, margins pubescent, backs glabrous to densely pubescent, marginal hairs sometimes longer; backs flattened dorsally; apex entire or minutely bifid, the teeth not greater than 0.2 mm long; awned just below apex, awns 3-6 mm long, straight; paleas 8-11 mm long, shorter than lemma, backs glabrous or slightly pubescent, margins usually pubescent; apex acute; anthers 3-5 mm long, yellowish-orange. Caryopses about 9 mm long, linear, dark brown. Chro- mosome number 2n = 14 Distribution.—Endemic to southern California; San Gabriel Mts., Santa Lucia Mts., San Bernardino Mts., San Antonio Mts., San Jacinto Mts., San Rafael Mts., and Topatopa Mts. Habitat.—Dry areas in open pine woods, hillsides, and rocky slopes; eleva- tion 1200-8000 ft. Specimens examined. U.S.A. California: Fresno Co.: junction N of South Fork of Kings River, | May 1923, Duncan s.n. (CAS); Kings Canyon road 4.3 mi W of Horseshoe Bend, 3600 [t, 6 Jun 1960, Howell 35330 (CAS), Kaiser, 27 Jun 1935, Bullard 95 (UC/JEPS). Los Angeles Co.: San Dimas Canyon, San Gabriel Mts, 1750 ft, 4 Jul 1933, Wheeler (CAS, US); San Gabriel Mts, 3500 ft, 4 Jul 1933, Ewan 7857 (US); San Gabriel Mts, East Fork, B a Anita ees 2250 ft, 30 May 1931, Ewan 4249(US): Little Santa Anita Canyon, 2 July 1902, vie 2632 (CAS, US); San Gabriel Mts, Bell Canyon, 2300 ft, 8 Jun 1936, Wheeler 4134 (CAS); San Gabriel Mts, a Canyon Camp, 2 mi off Soledad Canyon, 3000 ft, 27 May 1936, Wolf7845 (CAS, W); San Gabriel Peak, 9 Jul 1900, Dudley s.n. (CAS); San cl Mts., 990 m, | Sep 1994, Cayouette & Darbyshire C7947 (DAO); Summit of Mt. Wilson, 29 Jun 19 Abrams 2600 (CAS, US); San Gabriel Mts, Big Tujunga Creek between Coldwater and Wickiup Can yons, LO Jul 1992, Ross 6671 (UC/JEPS); Horse Flats, San Gabriel Mts, 16 Aug 1991, Ross 5873 (UC/JEP Roundtop Mtn, San Gabriel Mts, Chilao Flat, 14 Aug 1991, Ross 5780 (UC/JEPS). Madera Co.: 0.5 mi S - Southfork, 17 Jun 1933, Hormay 26 (UC/JEPS). Monterey Co.: Pico Blanco, 11 May 1901, Davy 7339 (US); Santa Lucia Mts, Jun 1901, Davy 7691 (US); Tassajara Hot Springs, Jun 1901, Elmer 3398 (CAS, US); Little Sur, 14 May 1901, Davy 7385 (US); coast range W of King City, 8 Jun 1927, Swallen 596 (US); Gavilan Peak, 3000 ft, 1 Jul 1861, Brewer 740 (US); Alder Creek, Santa Lucia Mts, 3000 ft, 23 Jul 1958, Hardham 3789(CAS), Arroyo Seco River Canyon near Hanging Valley, 2 Jun 1957, Howell 32566 (CAS): Arroyo Seco River, Santa Lucia Mts, 5 Jun 1959, Hardham 4796 (CAS); Cruikshank Trail, Santa Lucia 0 2004 BRIT.ORG/SIDA 21(4) Mts, 26 May 1903, Howitt 1589 (CAS), Frances Simes cae Natural History Reservation, Santa Lucia Mts, 22 Jul 1944, Linsdale s.n. (CAS); Frances Sime s Natural History Reservation, up- per Carmel Valley, about 2 mi NE of Jamesburg, 22 Aug 1942, Oumar sn. (CAS); Nacimiento Sum- mit, Santa Lucia Mts ts 18 Jun 1955, Munz 20886 (CAS): near Carmel River, 26 Jun 1905, Dudley sn. (CAS); near Naci it — Santa Lucia Mts, 26 Jun 1957, Raven 10987 (CAS), Santa Lucia Mts, bank by road from Arroyo Seco Cam idido Camp, 24 Jun 1956, Howitt 980 (CAS), = Santa Lucia Mts, between Middle and South F aie af Dev il’s Creek near Canogas Falls, 2000 ft, 23 May 1983, Carpenter 218 (CAS); Santa Lucia Mts, Landels-Hill Big Creek Reserve, Gamboa Point Sec- tion, 27 May 1982, Genetti & Engles 393 (CAS); Santa Lucia Mts, San Miguel Creek, below confluence with Anthony Creek, ca. 5 air km WNW of San Miguelito Ranch Ruins, 390 m, 5 Jun 1999, Wilken 15707 (CAS), Santa Lucia Mts, South Fork Devil’s Canyon, along Gamboa Trail near saddle of Twin Peak se Peak 3850, 4 Jun 1983, Carpenter 257 (CAS); Santa Lucia Mts, Jun 1901, Davy 7653, 7718, 7720 (UC/JEPS), Orange Co.: Santiago peak, 20 Jun 1927, Swallen tee ), Silverado Canyon., 640 m, 3 Sep a Cayouette & Darbyshire C7953 (DAO), Santa Lucia Mts, Mi i Creek, 25 May 1941, Hoovet 5260 (K). Placer Co.: N branch of Middle Fork of American River, at road crossing E a i orest Hill, 20 Jul 1952, Stebbins et al. 5132 (CAS). Riverside Co.: San Jacinto Mts, Idyllwild Area, N of Strawberry Valley, between Bear Trap Cyn and Lily Creek, 5200-5300 ft, | Aug 1996, White 4442 (CAS); Colo rado Desert, 1889, Palmer s.n. (US); Fuller Creek, 29 Jun 1936, Yates 6383 (UC/JEPS). San Benito Co.: Pinnacles National Monument, 3 Jun 1939, Stebbins 2773 (UC/JEPS). San Bernardino Co.: San Ber- nardino Mts, 29 May & 28 Jun 1888, Parish 2053 (US); San Bernardino Mts., 3300 ft, 13 Jun, Reed 1011 (US); San Bernardino Mts, 29 May 1890, Parish s.n. (US); San Bernardino Mts, road below Highlanc and Running Springs, 26 Jun 142, Beetle 3645 (UBC); San Antonio Mts, 8000 ft, 28 Jul L917, Johnston 1407 (UC/JEPS, US): N of Snow Canon, 5500 ft, 20 Jun 1901, Parish 5038 (CAS); San Bernardino Mts, 2500 ft, 5 Jun 1917, Parish 11304 (CAS, UC/JEPS); San Bernardino Mts, road between Highland anc Running Springs, 26 Jun 1942, Beetle 3645 (CAS); San Bernardino Mts, 29 May 1888, Parish s.n. (CAS): San aoe Mts, 3000 ft, 29 Jun 1888, Parish 2053 (CAS); San Bernardino Mts, Dobbs trail, Mill Creek, 2 Jul, Crawford s.n. (CAS); San Bernardino Ntl Forest, road 38, near Angelus, 1654 m, 2 Sep 1994, Cayouette & Darbyshire C7949 (DAO), 0.5 mi Sof Santa Ana River, 26 Jun 1937, Yates 6688 (UC/ Pee San Diego Co.: Cleveland National Forest, 30 Jun 1915, Hitchcock 13160 (US). San Luis Obispo « between Rocky Butte and Pine Mt. Santa Lucia Mts, 21 Jun 1950, Hoover 7998 (CAS, UC/JEPS); near Rocky Butte Fire Lookout, 12 June 1964, Hoover 9076 (CAS). Santa Barbara Co.: San Rafael Mts, Potrero on SE side of Cachuma Peak, 4000 ft, 12 Jun 1961, Blakley 4491 (CAS, US), N side of La Cumbre Peak, Santa Ynez Mts, 3600 ft, 5 Dec 1958, Pollard s.n. (CAS); Romero Canyon, Santa Ynez Mts, 7 Jul 1951, Pollard s.n.(CAS); W fork of Cold Spring Canyon, Santa Barbara, 16 Jun 1965, Pollard s.n. (CAS), Santa Ynes Mountains, 5 mi E of Santa Barbara, 12 Sep 1994, Cayouette & Darbyshire C7971 (DAO), bank of Camino Cielo, E of and near San Marcos Pass, 2000 ft, LO Jun 1958, Pollard s.n. CW); Zaca Lake Forest Reserve, 19-30 Jun 1906, Eastwood 719 (UC/JEPS). Santa Cruz Co.: near Eagle Rock, 2500 ft, Hesse 2731 (CAS); Santa Cruz, 27 Jun 1938, Silveus 2829 (CAS). Tuolumne Co.: Tuolumne River at Early Intake dam, 10 mi W of Mather, | Jul 1951, Stebbins 5000 (UC/JEPS). Ventura Co.: Red Reet Canyon, Topatopa Mts, 2800-3500 ft, 8 Jun 1908, Abrams & McGregor lol (US); above Murietta-Santa Ynez divide, Santa Ynez Mts, Ojai District, 4300 ft, 27 Jun 1963, Pollard s.n. (CAS); Matilija Canyon, 3 Jun 1945, Pollard s.n. (CAS); Murrieta Canyon, 15 Jun 1946, Pollard s.n. (CAS); Red Reef Canyon, Topatopa Mts, 2800-3500 ft, 8 Jun 1908, Abrams & McGregor 161 (CAS); Santa Ynez Mts, Camino Cielo, 3500 ft, 29 Jul 1967, Pollard s.n(CAS); Upper North Fork, Matilija Canyon, Ojai District, 27 Jun 1962, Pollard s.n. (CAS): Ventura River Basin, Camino Cielo, 4000 ft, 25 May 1946, Pollard s.n. (CAS): Santa Ynez Mts, Ojai to Cuyama Valley Road, N Fork of Ventura River, 2.2 mi below Wheelers Hot Springs, 1200 ft, 21 May 1935, Wolf 6888 (CAS); 0.5 mi N of Whiteacre Pk, 18 Jun 1935, Simontacchi 120 (CUC/JEPS). Ww es Qa Bromus hallii (Hitchc.) Saarela @ PM. Peterson, comb. nov. Bromus orcuttianus var. hallii Hitche., Fl. Calif 1175. 1912. Type: U.S.A. CALIFORNIA: west side, San Jacinto Mts, 27 Jun L9OL, H.M. Hall 2301 (HOLOTYPE: US-4128941). SAARELA ET Al 2005 Loosely caespitose perennial. Culms 90-150 cm tall, erect, puberulent or occa- sionally glabrous; nodes 1-2(-3), dark brown, retrorsely pilose to densely pu- bescent below. Leaf sheaths 6-12.5cm long, 1/4-3/4 as long as internodes, closed for most of their length, densely pubescent, hairs 0.3-1 mm long; margins smooth, occasionally hyaline at apex; collars densely pilose, with hairs up to 2 mm long, auricles absent; cataphylls and basal sheaths densely pilose, some- times shredding; ligules 0.5-2.5 mm long, membranous, sparsely to densely pubescent adaxially, glabrous abaxially, apex obtuse, erose; blades 7.5-16.5 cm long, 3-12 mm wide, flat, membranous, apically cucullate, densely pubescent abaxially and adaxially, hairs 0.2-0.5 mm long, occasionally longer hairs up to 1.2 mm on margins near base; margins scabrous or smooth. Panicles 5-16 cm long, 2-11 cm wide, open to densely branched; branches erect, ascending and appressed to spreading not more than 90’ from the rachises, pubescent; lower inflorescence nodes with 1-2 branches, lower branches 3.5-11 cm long with l- 2(-3) spikelets, upper spikelets occasionally sessile and spike-like Spikelets 2.5- 3.5(-4.5) cm long, 3-7-flowered, terete to conspicuously distichous at maturity with rachis visible, lowest rhachilla 2.5-4 mm long; glumes subequal, shorter than lemmas; sparsely to densely pubescent, margins sometimes hyaline; lower glumes 5-8(-9) mm long, 1(3)-nerved, apex acute; upper glumes (7-)8-9 mm long, 3-nerved, apex acute, acuminate, or mucronate, the mucro less than 1 mm long; lemmas 10-14 mm long, 5- to 7-nerved, sparsely to densely pubescent across back, marginal hairs sometimes longer, up to 0.5 mm long, backs flat- tened dorsally to slightly keeled; apex entire or minutely bifid, the teeth not greater than 0.2 mm; awned just below the apex, awns 3.5-7 mm long, straight; paleas 9-13 mm long, shorter than the lemmas, backs glabrous or slightly pu- bescent, margins usually pubescent; apex acute; anthers 3-6 mm long, yellow- ish-orange. Caryopses 9-11 mm long, linear, amber to dark brown. Chromosome number unknown. Distribution.—Endemic to southern California; known only from Fresno, Kern, Los Angeles, Monterey, Santa Barbara, San Bernardino, and Tulare counties. Habitat.—Dry, open or shady areas on hillsides, rocky slopes, and pine woods in the mountains; elevation 5200-8800 ft. Specimens examined. U.S.A. California: Fresno Co.: between Vidette Meadow and Bullfrog L 9 Aug 1940, Howell 16079 (CAS); Bubbs Creek Canyon in vicinity of Vidette Meadows, 9,500-10,000 ft, 23 Jul 1948, Howell 24942 (CAS); Simpson Meadow, Middle Fork of the Kings River, 6000 n ae 1958, Howell 33926 (CAS): Wood's Creek, 19 Jul 1910, Clemens s.n. (CAS). Kern Co.: 1.5 mi W of Green- horn Summit, 5200 ft, 22 Jun 1970, Howell & True 46599 (CAS); Breckenridge Mountain Road, 0.4 mi E of Barrel Spring, 6500 ft, 19 Aug 1958, Twisselmann 4725 (CAS); Greenhorn Mountains, 1.5 mi SE of Summit, 5000 ft, 8 Jul 1962, Howell 38312 (CAS); Greenhorn Pass Road, about 0.5 mi E of Neat Greenhorn Range, 5750 ft, 13 Aug 1958, Twisselmann 4666 (CAS); Greenhorn Range, NW of Calf Creek 17 Sep 1963, Twisselmann 9010 (CAS), Greenhorn Range, Greenhorn Pass Road just $ of a Flat, 6600 ft, 13 Aug 1958, Twisselmann 4653 (CAS): Kern Plateau, road to Bartolas Country E of Little Cannell Meadow, 7200 ft, 12 Jul 1966, Howell & True 41851 (CAS): Kern Plateau, trail to Little Cannell 2006 BRIT.ORG/SIDA 21(4) Meadow above Pine Flat, 7450 ft, 24 Jul 1964, Twisselmann 9880 (CAS); Old Kernville Road, 4.4 mi E of Greenhorn Summit, Greenhorn Range, 5200 ft, 21 Jun 1957, Twisselmann 3716 (CAS). Los Angeles Co.: Buckhorn, San Gabriel Mts, 6500 ft, 20 Jul 1933, Duran 3521(CAS, OSU, US); San Antonio Mts, $ Fork of Lytle Creek, 6000 ft, 15 Jul 1917, Johnston 1454 (CAS, US), San Antonio Canyon, 27 Jun 1927, Swallen 679 (US): San Bernardino Mts, 2 mi N of Big Pine Camp, 25 Jun 1942, Beetle 3640 (CAS); San Gabriel Mts, Blue Ridge, 8000-8500 ft, 18 Jul 1947, Howell 23389 (CAS); San Antonio Mountains, Prairie Fork of San Gabriel River, 6500 ft, 6 a 1918, Johnston 2070 (CAS); San Gabriel Mts., 1826 m, | Sep 1994, Cayouette & Darbyshire C7948 (DAO). Monterey Co.: Tassajara Hot Springs, Jun 1901, Elmer 3314 (CAS, K, OSU, US); Santa Lucia — 1901, Davy 7709 (US), 7710 (OSU); NW of Tassajara Road suminit, Chews Ridge, Los Padres National Forest, 15 Jun 1973, Griffin 3625 (UC/JEPS). San Bernar- dino Co.: San Bernardino Mts, Little Bear Valley, Aug 1907, Wilbur 1071 (US); San Bernardino Mts, 22 Jul 1902, Abrams 2799 (CAS, K); San Ber nardino Mts, Bear Valley, ot 19 Jul 1900, Jones s.n. (CAS); San Jacinto Mts, W fork of Snow Creek, 5000 ft, Jul 1901, Hall 2538 (CAS, K); San Gorgonio Mts., S of Barton Flats, 2251 m, 2 Sep 1994, Cayouette & Darbyshire C7 950 (DAO), San Bernardino Mts, Deep Creek, 6500 ft, 19 Jul 1899, Hall 1348 Ck); above Arrowhead Lake, San Bernardino Mts, | Jul 1927, Swallen 700 (OSU, US); 2 mi W of Barton Flat, 26 Jun 1937, Yates 6694 (UC/JEPS). Santa Barbara Co.: Mission Pine, San Rafael Mts, 6200 ft, 25 Apr 1930, Hoffman 90 (US); San Bernardino Mts., Santa Ana Canyon above Clark’s Ranch, 7400 ft, Jul 1926, Quibell 59 (US). Tulare Co.: trail from Bakeoven to Templeton Meadows, 8800 ft, 17 Jul 1950, Howell 27005 (US, CAS); Kaweah River Valley, 29 Jul 1891, Coville & Funston 1346 (US); Kings Canyon National Park, below Mist Falls, S. Fork Kings River, 13 Jul 1927, Swallen 770 (US); Copper Creek trail to Granite Basin, 1] Jul 1927, Swallen 753 (US); Aug 1897, Dudleys.n. (CAS); Kern Plateau, about 2 mi NW of Beach Meadow, 8200 ft, 7 Aug 1967, Howell & True 43419 (CAS); Kern Plateau, N of Cain Meadow, ca. 7200 [t, 24 Jun 1966, Twisselmann 12421 (CAS): Kern Plateau, North Manter Creek, Big Meadow, 7700-8000 [t, 24 Jun 1970, Howell & True 46696 (CAS): Monarch Lakes Trail, ca. 8000 ft, 19 Jul 1951, Howell 27943 (CAS); Portugese Pass, 7400 ft, 5 Aug 1957, Iwisselman 3946 (CAS); Quaking Aspen Road, 1.3 mi N of Ice Creek, 6600 ft, 8 Jul 1959, Twisselmann 5497 (CAS); Sequoia National Forest, Lloyd Meadows Basin, 1.25 mi W of the Pyles Boys Camp in the Freeman Redwood Grove, 5800 ft, 19 Jun 1973, Shevock 2770 (CAS); Sequoia National Forest, Lloyd Meadows Basin, along USFS road 22582, FE base of the Needles, about 3 mi from the Pyles Boys Camp, 5800 ft, 4 Jun 1974, Shevock 3580 (CAS), Sequoia National Forest, Lloyd Meadows Basin, approxi- mately 2 mi NW of the Pyles Boys Camp, 5700 [t, 10 Jul 1974, Shevock 3716 (CAS); Sequoia National Forest, near Belknap Creek, 26-28 Jul 1941, Bacigalupi, Wiggins, & Ferris 2655 (CAS); Sequoia Nt Forest, N limit of Sequoia Ntl Park road, 2010 m, 5 Sep 1994, Cayouette & Darbyshire C7955 (DAO): Long Meadow, eg 9000 ft, 14 Jun 1888, Palmer 233 Cx, US); Mt. Silliman, Clover Creek, 29 Jul 1896, Dudley l481(C Bromus orcuttianus Vasey, Bot. Gaz. 10:223. 1885. Bromopsis orcuttiana (Vasey) Holub, Folia Geobot. Phytotax 8(2):168, 1973. Typr: U.S.A. CALIFORNIA: in the mountains near San Diego, 1884, CR. Orcutt s.n. C(HOLOTYPE: USD. Bromus brachyphyllus Merr., Rhodora 4:146. 1902. Type: U.S.A. OREGON: Crook Co. Black Butte, open dry pine forests, 19 Jul L901, Cusick 2677 (HOLOTYPE: unknown; ISOTYPE: USI). Loosely caespitose perennial. Culms 90-150 cm tall, erect, glabrous to pubes- cent; nodes 2-4, dark brown, retrorsely pilose to densely pubescent below. Leaf sheaths 3.5-11 cm long, usually 1/3-2/3 as long as internodes, closed for most of their length; lower sheaths sparingly to densely pilose, hairs 2-4 mm long, occasionally glabrous; upper sheaths pubescent, hairs up to 1 mm long; mar- gins smooth, occasionally hyaline at apex; collars pilose with hairs up to 4mm long or glabrous, auricles absent; cataphylls and basal sheaths glabrous, some- SAARELA ET Al 2007 times shredding; ligules 1-3 mm long, membranous, glabrous, occasionally pi- lose adaxially, apex obtuse, erose; blades 7-24 cm long, 3-12 mm wide, flat, membranous, apically cucullate, glabrous, edges sometimes pilose with hairs up to 2mm or pubescent on lower 15-25%; margins scabrous or smooth. Panicles 7-13.5 cm long, 2-10 cm wide, open to densely branched; branches erect, as- cending and appressed to spreading not more than 90° from the rachises, pu- bescent; lower inflorescence nodes with 1-2(-3), lower branches 3-7.5 cm long with 1-5 spikelets, upper spikelets occasionally sessile and spike-like. Spike- lets 2-3.7 cm long, 3-9-flowered, terete to conspicuously distichous at matu- rity with rachis visible; glumes subequal, shorter than lemmas, glabrous, occa- sionally scabrous on veins or pubescent, margins sometimes hyaline; lower glumes 5-9 mm long, 1(3)-nerved, apex acute; upper glumes 7-11 mm long, 3- nerved, apex acute or mucronate, the mucro up to lmm long; lemmas 9-15mm long, glabrous, scabrous, or sparsely to densely pubescent, marginal hairs some- times longer, the hairs up to 0.5 mm long; backs flattened dorsally to slightly keeled; apex entire or minutely bifid, the teeth not greater than 0.2 mm, awned just below the apex, awns (4-)5.5-8 mm long, straight; paleas 8-11 mm long, shorter than lemmas, backs glabrous or slightly pubescent, margins usually pubescent; apex acute; anthers 3-5 mm long, yellowish-orange. Caryopses 6-9 mm long, linear, brown to purple-black. Chromosome number 2n = 14. Distribution.— Known from California, Nevada, Oregon, Washington, Ari- zona, and Utah. Habitat—Dry areas in open pine woods, on hillsides and rocky slopes, and in meadows in the mountains; elevation 1850-11500 ft. —_ Specimens examined. U.S.A. ARIZONA: Cochise Co.: Huachuca Mts, Sept 1883, Lemmon s.n. (US). CALIFORNIA: Alpine Co.: N side of Pigeon Flat, 22 Jul 1940, Hoover +423 (K). Amador Co.: Pioneer, 3500 ft, 13 Jul 1896, Hansen 1835 (K, US): Jackson, 7 Sep 1937, Johannsen 1247 (UC/JEPS). Butte Co.: Butte Meadows, 4600 ft, 26 Jul 1917, Heller 12819 (CAS, OSU, US); Butte Creek, 5000 ft, 21 Jul 1930, oa 1348 (CAS); Jackson Ranch Ridge, along Road 21N25Y ca 0.6 mi E of the W end of the ridge, 3750 ft, 27 Jul 1995, Janeway 4877 (CAS). W side of Hwy. 32 ca. 1/4 mi N of Forest Ranch, 2357 ft, 18 Jul 1978, Taylor 1674 (CAS); along Big Bar Mountain Road, about 3 mi SW of Coyote SE of Pulga, 8 Jul 1987, Ahart 5796 (CAS). Calaveras Co.: near zi eae 1978, Howell, Menzies,&S hockey) 53063 (CAS); Dorrington, 1976, Menzies s.n. (CAS). Colusa Co.: Trout Creek, SE Snow Me 30 Jul 1981, Heckard & Hickman 5763 (UC/JEPS). El Dorado Co.: along US ie 3 mi E of Camino, 3400 ft, 15 Jul 1945, Robbins 2062 (US); Chute Champ Road, 3000 ft, 15 Jun 1937, White 1078 (US); Fallen Leaf Lake, 27 Jul 1928, Abrams 12630 (CAS): Fallen Leal Lake, 9 Jul 1928, Abrams 12627 (CAS): Glen Al- pine region, Camp Agassiz, 21 Jul-15 Aug 1906, East wood 998 (CAS); Echo Summit, | Sep 1946, Howell 22986 (CAS); 0.5 mi W of Omo Ranch Post Office, 22 Jun 1956, Crampton 3596 (CAS). Fresno Co.: Pine Ridge, 5300 ft, 15-25 Jun 1900, Hall & Chandler 316 (UC/JEPS, US); Huntington Lake, 22 ae 1927, Swallen 810 (US); Bearskin Meadow, 2 Jul 1899, Eastwood s.n. (US); Granite Basin, 10 July 192 Swallen 743 (US); above Deer Creek, N shore of Huntington Lake, 7100 ft, 8 Aug 1951, Quibel ase (CAS); Dinkey Creek, Jul 1901, Dudley s.n. (CAS): ae Lake and vicinity, near ranger station, 7000 ft, 3 Aug 1951, Pollard s.n. (CAS); John Muir Trail N of Bear Creek, 9400 ft, 5 Aug 1954, Raven 7789 (CAS); Mono Creek, 8200 ft, 9 Aug 1953, Raven 6142 aes Tel a Valley, Middle Fork - the Kings River, 4650 ft, 26 Jul 1958, Howell 33958 (CAS); drainage of Deer Creek, above Lakeshore Post 2008 BRIT.ORG/SIDA 21(4) Office, Huntington Lake, 23 Aug 1958, Bacigalupi & Quibell 6717 (UC/JEPS); E slope of Converse Mt, drainage of Indian Creek, along Kings Canyon Road, 23 Jul 1958, Bacigalupi & Alava 6498 (UC/JEPS): Huckleberry Creek, Huntington Lake, 28 Jul 1928, Jepson 13330 (UC/JEPS). Glenn Co.: Plaskett Meadows, 6000 ft, 4 Aug 1943, Howell 19020 (CAS, US); road 168, a few miles E of Shaver Lake, 2100 m, 6 Sep 1994, Cayouette & Darbyshire C7958 (DAO); Mendocino National Forest, 5500 ft, 21 Jul 1956, Burcham 368 (k); near Fish Pond Plaskett Meadows, 5 Aug 1943, Baker 10583 (CAS). Humboldt Co.: Orleans Mt, 2 Sep 1946, Pollard s.n. (CAS); Trinity Summit, on ridge 2 mi E of Grove’s Prairie, 15 Aug 1948, Tracy 18125 (UC/JEPS); Grouse Mt, 27 Jun 1934, Clarks & Tracy 11079 UC/JEPS). Lake Co.: N of Hoberg’s, 14 Jul 1944, Hoffman 1967 (UC/JEPS, US); foothills S of Mt. Sanhedrin, midway between Potter Valley and Hullville, 25 Jul 1902, Heller 5987 (US); S base of Mt. Sanhedrin, 25 Jul 1902, Heller & Brown 5987 (US): Bz ile Mtn, 23 Jun 1948, Wagnon 1509 (ik): 15 mi N of Lakeview, 6 Jul 1927, Peck 5517 (CAS), summit of Bartlett Mt, 1] Jul 1939, Stebbins 2971 (UC/JEPS); NW of Timber Lake, Snow Mt, 16 Jun 1979, Heckard & Hickman 5086a (UC/JEPS). Lassen Co.: 5 mi S of Susanville, 27 Jul 1927, Swallen 872 (US); Black’s Mt, LO Jul 1934, Howell 12554 (CAS); Elysian Valley, W of Janesville, 28 Jun 1973, Howell, True & Williams 49472 (CAS), Gold Run Creek cad Sof Susanville, Diamond Range, | Aug 1973, Howell & True 50180 (CAS); Gold Run Creek SW of Susanville, 19 Jul 1976, Howell 51973 (CAS); on road from Janesville to Thompson Peak, Diamond Range, 31 Jul 1973, Howell & True 50029 (CAS). Madera Co.: 9 mi above Bass Lake on Beasore Meadow Road, 18 Jul 1933, Springer 505 (OSU); Bass Lake Fish Hatchery, 1 Sep 1951, Holt s.n. (CAS). Mariposa Co.: || Aug 1895, — s.n. (US); Signal Peak, Chowchilla Mt., 25-31 Jul 1938, Quick 2003 (CAS ); road 41, Sierra Ntl F miles of Yosemite Ntl Park, 1914 m, 8 Sep 1994, Cayouette & Darbyshire C7964 (DAO); ee to Wawona, 28Jun LOL, Jepson 4295 (UC/JEPS). Mendocino Co.: along ro. ier nroute tosummit of he Sanhedrin from Towhead Flat, 7 Jul 1981, Knight & Se 4295 (CAS). Modoc Co.: Forestdale, 4500 ft, 25 Aug 1894, Baker & Nutt ings.n. (US). 13 Jul y, Baker & b Nutting eee Nevada Co.: Truckee, ene 1913, Hitchcock 10516 (x, US, W); Banner Mt, 4 mi E of Nevada City, 15 Sep 1961, True 261(CAS): Hwy 20, ca. 2.5mi —E of Washington ee 5000 ft, 10 Jul 1968, True 4341 (CAS); Omega Rd, between Omega Dig- i and Diamond Creek 14 mi NE of Skillman Flat on Hwy 20, 15 Jul 1965, True & Howell 2359 ); Mayflower spi 5 ie 1966, Mott s.n. (CAS); Banner Mt, ca. 4 mi E of Nevada City, 15 Jul 1965, a ue & Howell 2319A (CAS); on road to Pierce Meadows, short distance from the S Fork of the Yuba River, Tahoe ee ra 30 Jun 1931, Smith 2575 (UC/JEPS). Placer Co.: Tahoe, 5 Aug 1908, 6225- 7000 ft, Hitchcock 3091 (US); railroad crossing at Blue Canyon, 17 Jul 1956, Crampton 3723 (CAS); about 2. mi SW of French Meadows on the road to Big Meadow, 2 Aug 1981, Best s.n. (CAS); | mi from Blue Canyon on road to Emigrant Gap, 4 Aug 1956, Raven 9999 (CAS): Truckee, 8 Aug 1936, Yates 5981 (UC/JEPS):; Eldorado Canyon, above Bullion Mine, Tahoe National Forest, 4 Jun 1926, Smith 1883 (CAS, UC/JEPS); Antone Meadow, 7 Sep 1967, Hoover 10858 (UC/JEPS). Plumas Co.: between Blairsden and Gold Lake, 5500 ft, 5 Jul 1938, Wood 7 (US); Truckee River, July 1888, Sonne 21 (US, CAS); Butterfly Botanical Area, ol luence of See ee reek and Butterfly Creek, 2900 ft, Ll Jun 1968, Howell, Knight, Knight, & True 2352 (CAS), Fern Glen, 26 Jul 1966, Knight, Knight & Howell 1576 (CAS); Drakesbad, 5500-6000 ft, 17 Jul 1960, aa 35494 (CAS); Drakesbad, 5500-6000 ft, 20 Aug 1960, Howell 36274 (CAS); Gold Lake Road above Blairsden, 5500 ft, 25 Jun 1934, Ewan 8227 (CAS): Johnsville, 5200 ft, 28 Jun 1951, Howell 27656 (CAS); Lassen Volcanic National Park, trail to Little Willow Lake, 21 Jul 1960, Howell 35860 (CAS); near Prattville, Lake Almanor, 14-26 Aug 1944, Kearney 7 (CAS); near the sum- mit of Soapstone ridge 12 mi W of Bucks, 5500 ft, 7 Jul 1915, Heller 12053 (CAS, OSU); Quincy, 31 Jul 1942, Quick 42-46 (CAS); Willow Lake Meadow near the east inlet, 5450 ft, 17 Jul 1957, Gillett 805 (CAS, UC/JEPS);, 3 mi W of Keddir, Butterfly aie 3600 ft, 10 Jul 1967, Rose 67152(W). near Prattville, 20 Jul 1926, Howell 2088 (CAS); Round Valley, 9 Jul 1973, Howell 49722 (CAS); 2 mi NW ol Spring Garden, 22 Jul 1975, Howell 51363 (CAS); Butterfly Valley, 3 mi. W > Keddie, 27 Jul 1966, Rose 66060 (CAS); about 0.5 mi N of Humboldt Summit, 23 Jul 2001, Ahart & Oswald 8998 (UC/JEPS). San Diego Co.: Cleveland National Forest, E of San Diego, near Cuyamaca hake 1700 r m, 29, 30 Jul 1915, Hitch- cock 13168 (kK, UC/JEPS, US, W); Cuyamaca Mts, 30 Jun 1903, Abrams 3945 (OSU). Shasta Co.: Goose SAARELA ET Al 2009 Valley, 29 Jun-11 Jul 1912, East wood 930, 985 (CAS, US), 4 Jul 1914, Smith 735 (CAS); 0.25 mi above here Creek on Chaos Crags trail, 11 Jul 1957, Gillett 705 (UC/JEPS, CAS); Goose Valley, 29 11 Jul 1912, Eastwood 930 (CAS); Redding, 5 Jun 1934, Kraebel 25 (UC/JEPS); Logan Mt, 13 mi N of Lassen Peak, Lassen National Forest, 12 Jul 1934, Whitney 2163 (UC/JEPS); Little Hatchet Creek oo 1940, Jepson 20131 (UC/JEPS). Sierra Co.: near summit of Yuba Pass, 6350 ft, 17 Aug 1944, Beetle 3041 (US); Tahoe Forest, 10 Sep 1925, Smith 1727 (CAS); Tahoe Forest, 10 Jun 1926, Smith 1906 (CAS); Tahoe forest, 19 Jul 1926, Smith 1994 (CAS): Tahoe Forest, 4 Jul 1926, Smith 1883 (CAS); Independence Lake watershed, W of Independence Lake, 7200 ft, 26 Jul 1977, True 8380 (CAS), 2 mi E of Bassett station, North Fork of Yuba River, 4 Aug 1934, Jepson 16850 (UC/JEPS). Siskiyou Co.: base of Mt. Eddy, 18 Jul 1915, Heller 12124 (CAS, OSU, US); S Fork of Shasta River, Mount Eddy, Shasta Forest, 1850-2000 m, 11, 12 Aug 1915, Eggleston 11635 (US); Sisson, 30 Jul 1894, Howe 122 (US); near Shasta Springs, 13 Jun 1905, Heller 8026 (CAS, US); between upper reaches of China and Blind Horse Creeks, S side of South Fork of Salmon River, 24 Jul 1955, Wiggins 13497 (CAS, UC/JEPS), toward Black Fox, 15 Aug 1899, sie n.(CAS); Trinity Mts, about 6 mi SE of Cecilville, W side of Rush Creek, 21 Jul 1954, Thomas Thomas 4432 (CAS); on Everitt Memorial Hwy, 4.7 mi N of McCloud River railroad crossing, 13 Jul on Frenkel 229 (CAS); English Lake, Salmon Mts, 16 Aug 1969, Oettinger 1392 (UC/JEPS), below Pine Lake Basin, Salmon Mts, 6 Aug 1969, Oettinger & Thorne 1246 (UC/JEPS); Humbug Creek, 20 Jul 1908, Butler 469 (UC/JEPS). Tehama Co.: Deer Creek Canon, 17 Jul 1911, Eggleston 7284 (US), 5.1 mi from Whitlock Camp, 12 Jul 1953, Baker & Wagnon 12861 (UC/JEPS); E side of Willow Creek, about 1.25 mi N of Jonesville, 24 Jul 1994, Ahart 7513 (UC/JEPS). Trinity Co.: Grasshopper Public Camp, 2.5 mi NW of Stuart Gap, North Yolla Bolly Mts, 17 Jul 1951, Munz 16575 (CAS); North Fork of Trinity River, Hobo Gulch Camp vicinity, 18 mi NW of Weaverville, along backbone Creek Trail near Key- stone, 15 Jun 1972, Carter 435(CAS). Tulare Co.: S Fork of Kaweah River, 20 Jul 1904, Culbertson 4512 (US); above Mineral King, 30 Jul 1927, Swallen 883 (US); Sequoia National Park, 18 Jul 1927, Swallen 882 (US); Grant Park, 4000-8000 ft, 1] Aug 1895, Dudley 1205 (CAS), Hollow Log Camp, Jul 1900, Dudley s.n.(CAS); middle Kaweah River, 2 Aug 1900, Dudley 3031 (CAS): Mineral King road, 6000 ft, 15 ie 1951, Howell 27780 (CAS); Sequoia National Park, 12 Aug 1896, Dudley 1625, Sequoia Forest, man Creek, 2 Aug 1916, 7500 ft, Cunningham 3 (CAS); Sequoia National Forest, 11500 ft, 23 Jul os Hopping 22 (CAS), Kaweah River, 20 Jul 1904, without collector (US); Sequoia National Park, road to Crystal Cave, near start of Black Oak trail, 21 Jul 1948, Bailey & Bailey 2446 (UC/JEPS); L loyd Meadow, near trail to Quaking Aspen on the Springville Road, 24 Jul 1964, Smith 1316 (UC/JEPS). Tuolumne Co.: Yosemite National Park, Echo Creek Canyon, 17-25 Aug 1908, Hitchcock 3352, 3355 (US); Yosemite National Park, Tenaya Trail, 6500 ft, 21-22 Jul 1915, Hitchcock 13136 (US); Yosemite to Wawona, 28 Jun 1911, Jepson 4295 (US); Yosemite Valley, 4060 ft, 5Jul 1909, Jepson 3125 (US), Yosemite National Park, 1 Jul 1938, Silveus 2873 (CAS); Yosemite National Park, Glacier Point, I] Aug 1915, Abrams 5428 (CAS); Yosemite Valley, Sierra Nevada Mts, 5000-8000 ft, 4-12 Jul 1901, Parish 4360 (CAS); Yosemite, Jul 1902, Bacon s.n. (CAS); along Hwy 108, between Twain Harte and Confidence, 10 Jul 1972, 4000 ft, Wiggins 21785 (CAS); along South Fork road, 2.6 mi E of Twain Harte, 4000 ft, ll Jul L971, Wiggins 21679 (CAS), Dodge Ridge SE of Pinecrest, 21 Jul 1953, Quick 53-42 (CAS), Herring Creek 4 mi from Strawberry, 17 Jul 1936, Wiggins 8543 (CAS); Mather, in the Sierra Nevada in the lower borders of the Transition Zone, 1400 m, 6 Jun 1931, Keck 1252 (CAS), near Cow Creek Research Sta- tion, 5750 ft, 11 Jul 1941, Quick 41-69 (CAS); Twain Horte, 3700 ft, 6 Jun 1954, Howell 29944 (CAS), Long Barn, 16 Jul 1941, Hoover 5470 (K). Yuba Co.: about 2 mi NE of Dobbins on Oregon Hill road, 22 Jun 1981, Howell, Fuller & Barbe 54081 (CAS); about 200 yards S of the 4-H Camp, on W side of Oregon Hill Road, ae 0.5 mi N of Marysville Road, about 2 mi NE of Dobbins, 2 Jul 2003, Ahart 10340 (UC/JEPS). NEVADA: Washoe Co.: Incline, | Aug 1928, Smith s.n. (CAS). OREGON: Gayhart Buttes, 1850 m, 8 Aug 1896, Letberg 2887 (OSU); East Eagle Creek, 5500 ft, 10 Aug 1909, Cusick 3369, 3370 (OSU). Crook Co.: base of Black Butte, 19 Jul 1901, Cusick 2677 (UC/JEPS, US). Douglas Co.: along Golden Stairs Trail, E fork of Abbott Creek, ca. 20 mi W of Crater Lake, near Abbott Butte, 19 Jul 1972, Mitchell 241 (OSU). Grant Co.: Strawberry Lake, Strawberry Mts, 17 Jul 1910, Cusick 3525 (OSU, US). 2010 BRIT.ORG/SIDA 21(4) Jackson Co.: Klamath Forest, 4800 ft, 19 Jul 1934, W Pdi 1(CAS, UC/JEPS, US); Ashland Butte, Siskiyou Mountains, 20 Jul 1887, Howell 253, 369 (OSU, UC/JEPS, US, W, WTU): summit of Cascades, 10 Jul 1931, Peck 16747 (OSU); Carberry Creek, 3 mi ae mouth of Applegate River, 25 Jun 1931, Peck 16386 (OSU). Jefferson Co.: Suttle Lake, 18 Jul 1925, Peck 14421 (CAS, K, OSU); Abbot Butte Spring Road, 15 September 1959, Swed berg 100 (OSU), NW of Sisters, 23 July 1960, Johnson 543 (OSU); NW ol Sisters, 27 Aug 1960, Johnson 611 (CAS, OSU); near Hwy. 20, W of Black Butte, 20 Jul 1962, West s.n. (OSU). Siskiyou, 21 Jul 1908, Hitchcock 2879, 2909, 2916 (US). Klamath Co.: Klamath Valley at Keno, 27 Jun 1902, Cusick 2838 (CAS, OSU, UC/JEPS, US, W); Pelican Bay, W side of Upper Klamath, 16 Jul 1920, Peck 9516 (CAS, WTU): Fort Klamath, 29 Jul 1908, Hitchcock 3012 (US), Lake Co.: 15 mi N ol Lakeview, 6 Jul 1927, Peck 15517 (OSU). Wallowa Co.: Imnoha River, Wallowa Mountains, 10 Aug 1909, Cusick 3310 (OSU, ae Maen Co.: Dixie ices 15 Jul 1910, Cusick 3518 (US). UTAH: 1876, McLeans.n.(US). WASHING ackima Co.: Klickitat River, near Mt. Paddo, 12 Jun 1885, Suksdor{ sn. (CAS, K, Lb - i iddo, a e 1905, Suksdorf 5265 (US); Mt. Paddo, 27 Aug 1884, Suksdorf 120 (US). Klickitat Co.: Falcon Valley, 13 Jul 1924, Suksdorf 11754 (CAS, K, OSU, UBC, UC/JEPS, US, WTU): 6 Jun 1891, nea 826 (UC/JEPS). KEY TO THE SECTIONS OF BROMUS IN CALIFORNIA 1. Spikelets laterally compressed; lemmas compressed and keeled sect. Ceratochloa (P. Beauv.) Griseb. 1. Spikelets not laterally compressed: lemmas not keeled. 2. Lemma apex bidentate, teeth 3-7 mm, awn-like to acuminate. wn of the lemma geniculate and/or twisted _sect.Neob (Shear) Hitche. 3. Awn of the lemma straight, not twisted sect. Genea Dumort. 2. Lemma apex entire or bidentate, teeth 0-3 mm, not awn-like or acuminate. 4. Plants perennial, with or without rhizomes, the bases fibrous; lower glume 1- or 3-nerved; upper glume 3- or 5-nerved sect. Bromopsis 4. Plants annual; lower glume 3- or 5-nerved; upper glume 5- or 7-nerved sect. Bromus |. KEY TO BROMUS SECTION BROMOPSIS IN CALIFORNIA |. Plants with rhizomes; awns absent or up to 3 mm long B.inermis 1, Plants without rhizomes; awns present, 1.5-11 mm long. 2. Most lower glumes on a plant 3-veined. . Upper glumes 5-veined. 4. Ligule 0.4—1 mm long; glumes scabrous or pubescent; upper glume 6-9 mm long; blades and sheaths pubescent or glabrous . pseudolaevipes 4. Ligule (1.5-)2—4 mm long; glumes glabrous; upper glumes 7-11mm long; blades and ro glabrous B. laevipes 3. Upper glumes 3-veine 5. Lemma awns 1.5-3(-4) mm long; blades 2-5 mm wide; anthers 1.5—3.5 (—4) long B. porteri 5. Lemma awns 3-9 mm long;blades 3-12 mm wide; anthers 3.5—5 mm long. 6. Blades glabrous; lower sheaths pilose, hairs 2-4 mm long; nodes 2-4; inflorescence branches mostly ascending to spreading < 90° from ae rachises B. orcuttianus 6. Blades Spubescet lower sheaths pubescent, hairs up to 1(-3) mm long; no -7; inflorescence branches mostly spreading >90° from the I se B. grandis a 2. Most lower glumes on a plant 1-veined. 7, Glumes pubescent. SAARELA ET Al 2011 8. Panicles narrow at anthesis, = 2 cm wide; branches erect or tightly ascending; blades and sheaths glabrous B. suksdorfii 8. Panicles broad at anthesis, > 2 cm wide; branches erect,ascending, or nod- ding, usually spreading or divaricate; blades and sheaths pubescent or glabrous. 9. Ligules (2—-)3-6(—7) | | (4-)6-11 mm long; branches of inflorescence glabrous or scabrous B. vulgaris 9. Ligules 0.5-3 mm long; lemma awns 3-7(-9) mm long; branches of in- florescence pubescent. 10. Lower sheaths pilose with hairs 2-4 mm long, or glabrous; blades glabrou B. orcuttianus 10: ee oes igs pubescent with hairs up to 1 mm long;blades caries pubescen . Longest aie 7.5-16.5 cm long; 1—2(-3) nodes per culm B. hallii _ Longest blades (13-)18-38 cm long; 3-6(—7) nodes per culm B. grandis 7. Glumes glabrous. 12. Panicles narrow at anthesis, < 2 cm wide; branches erect or tightly as- ndin B. suksdorfii ce g 12. Panicles broader at anthesis, > 2 cm wide; branches erect, ascending, or nodding, usually spreading or divaricate. 13. Ligules (2-)3-6(-7) mm long; awns (4-)6-11 mm long B. vulgaris 13. Ligules 0.5-3 mm long; awns 4-7(-9) mm long 14. Lemmas not densely pubescent along margins; lemma awns (4-) 5-7(-9) mm long; inflorescence branches mostly ascending to spreading = 90° from the rachises B. orcuttianus 14. Lemmas d ly pubescent along margins; lemma awns (2-) 3- 5(-6) mm long; ‘inflorescence branches spreading to drooping >90° from the rac 15. Lemmas setae on ee back between pubescent mar- ae (0.9-)1-1.4(-1.6) mm;second glumes (6.2-) 7.1 8.5(-9.5) mm long; baa sheaths glabrous or with long hairs; top culm blad S n upper surfac Eton culm nodes sheath ety pubescent B. ciliatus . Lemmas with scattered hairs on lower back between pubes- cent margins; ets (1.2-)1.6-2.7(-3.4) mm long; second glumes (7.8-)8.9-11.3(-13.2) mm long; basal sheaths with dense, short, or oe ea hairs; top culm blades glabrous on the upper surface; top culm nodes usually glabrous; caryopses (6.9-)7.7-9.7(-10.5) mm long; top culm sheath glabrous B. richardsonii Sal ACKNOWLEDGMENTS We thank Dieter Wilken, an anonymous reviewer, and Barney Lipscomb for constructive comments on the manuscript; the curators of the cited herbaria for the loan of specimens or assistance during visits; Cindy Sayre (UBC) for as- sistance in the herbarium; and Nicole Fillion (Ottawa) for assistance with the 2012 BRIT.ORG/SIDA 21(4) chromosome count. J.M.S. was supported by a Canada Graduate Scholarship from the Natural Sciences and Engineering Research Council of Canada, the University of British Columbia, anda travel grant from the Lawrence R. Heckard Fund of the Jepson Herbarium to visit the U.S. National Herbarium. REFERENCES Ainouche, M.L.and R.J.Baver. 1997.On the origins of the tetraploid Bromus species (section Bromus, Poaceae): insights from the internal transcribed spacer sequences of nuclear ribosomal DNA. Genome 40:730-743. Crayton, W.D. and S.A. Renvoize. 1986. Genera graminum. Grasses of the world. Kew Bull. Add. Ser. 13:1-389. Crayton, W.D. and H. WituiAmson. 2002 onwards. World grass flora: Nomenclature. Royal Botanic Gardens, Kew. HitcHcock, A.S.1951.Manual of grasses of the United States (ed. 2, revised by A.Chase).U.S. Department of Agriculture, Washington, D.C HircHcock, C.L., A. CRonauist, M. Owneey, and J.W. THomeson. 1969. Vascular cryptogams, gym- nosperms, and monocotyledons. Vascular plants of the pacific northwest. University of Washington Press, Seattle. 1:1-194. Jepson, W.L. 1912.A flora of California. Volume 1.University of California, Berkeley Kearney, [.H.,R.H. Peestes, and coutasorators. 1960. Arizona flora. University of California Press, Berkeley. Massa, A.N., K. B. Jensen, S.R. Larson, and D.J. Hote. 2004. Morphological variation in Bromus sect. Ceratochloa germplasm of Patagonia. Canad. J. Bot. 82:136-144. Munz, PA. 1974. A flora of southern California. University of California press, Berkely. Munz, PA. and D.D. Keck. 1959. A California flora. University of California Press, Berkely. 1-168]. Oia, T.,V. Jaaska, and V. Vistar, 2003. Breeding system, evolution and taxonomy of Bromus arvensis, B. japonicus and B. squarrosus (Poaceae). Pl. Syst. Evol. 242:101-117. Pavuick, L.E. 1995. Bromus L.of North America. Royal British Columbia Museum, Victoria. Pavuick, L.E.and L.K. ANperton. In Press. Bromus. In: Barkworth, M.E., K.M. Capels, S.Long, and M.B. Piep, eds. Magnoliophyta: Commelinidae (in part): Poaceae, part 1. Flora of North America North of Mexico, volume 24. Oxford University Press, New York. Pavuck, L.E., A.M. PLANCHUELO, PM. Peterson, and R.J. Soren. 2003. Bromus.Pp.154-191.In:Soreng, R.J.,P.M. Peterson, G. Davidse, EJ. Judziewicz, F.O. Zuloaga, T.S. Filgueiras, and O.Morrone, Catalogue of New World grasses (Poaceae): IV. Subfamily Pooideae.Contr.U.S. Natl. Herb. 48:1—730. Peterson, P.M., J. Cavouette, ¥.S.N. FeRDINANDEZ, B. COULMAN, and R.E. CHAPMAN. (2001) 2002. Rec- ognition of Bromus richardsonii and B. ciliatus: evidence from morphology, cytology, and DNA fingerprinting (Poaceae: Bromeae). Aliso 20:21-36. Pittay, M.and K.W. Hitu. 1990. Chloroplast DNA variation in diploid and polyploid species of Bromus (Poaceae) subgenera Festucaria and Ceratochloa. Theor. Appl. Genet. 80: 326-332 SAARELA ET AL 2013 Pittay, M.and K.W.Hitu. 1995.Chloroplast-DNA restriction site analysis in the genus Bromus (Poaceae). Amer. J. Bot. 82:239-249. Piper, C.V. 1906. Flora of the State of Washington. Contr. U.S. Natl. Herb 11:1-635. SAARELA, J.M., PM. Peterson, R.M. Keane, J. Cavouette, and S.W.GraHam. 2005. Molecular system- atics of Bromus (Poaceae: Pocideae) based on chloroplast and nuclear DNA sequence data. Pp. xxx-xxx .In: Columbus, J.T. E.A. Friar, J.M. Porter, L.M. Prince and M.G. Simpson, eds. Monocots: comparative biology and evolution, 2 vols. Rancho Santa Ana Botanic Garden, Claremont, CA. In Press. Saarela,J.M., PM. Peterson, and N.F. ReFutio-Ropricuez.In review.Bromus ayacuchensis (Poaceae: Pooideae: Bromeae), a new species from Peru. Syst. Bot. SHear, C.L. 1900. A revision of the North American species of Bromus occurring north of Mexico. Bull. U.S.D.A. (1895-1901) 23: 1-66. Smity, P. 1970. Taxonomy and nomenclature of the brome-grasses (Bromus L. s.|.). Notes Roy. Bot. Gard. Edinburgh 30:361-375. Steesins, G.L., Jk., and R.M. Love. 1941.A cytological study of California forage grasses. Amer. J. Bot. 28:371-382. Vasey, G. 1885. Some new grasses. Bot. Gaz. 10:223-224. Waanon, H.K. 1952. A revision of the genus Bromus, section Bromopsis, of North America. Brittonia 7:415-480. Witken, D.H. and ELL. Painter. 1993. Bromus.|n:J.C. Hickman, ed., The Jepson manual: higher plants of California. University of California Press, Berkeley. Pp. 1239-1243. 2014 BRIT.ORG/SIDA 21(4) Book REVIEW VirGINIA D. NAZAREA. 2005, Heirloom Seeds and Their Keepers: Marginality and Memory in the Conservation of Biological Diversity. (ISBN 0-8165-2435-1, hbk.). The University of Arizona Press, 355 S. Euclid, Ste. 103, Tucson, AZ 85719, U.S.A. (Orders: 520-621-1441, fax 520-621-8899, ww W.UAPTess. arizona.edu). $35.00 pbk., $29.95 hbk., 190 pp., 31 b/w, author notes, glos- sary, index, 6" X 9” The term “seed savers” refers to those persons who grow traditional, heirloom, or handed-down vari- eties of fruits, grains or vegetables in personal, family or community gardens. In her book, Heirloom Seeds and Their Keepers, author Virginia Nazarea presents an analytical approach concerning of the role of seed savers in the preservation and conservation ol plant genetic diversity and cultural practices. Such analyses of this mechanism of biodiversity conservation are very timely given in- creasing concern over the loss of both crop diversity and tradition agricultural practices. Nazarea uses both metaphors and personal histories collected during research to orelate the ipore ince of the seed savers, which in continuing to grow heirloom varieties at 2 cultural heritage, family / locality memories, as well as different plant varieties for Rene ate use. Alternative species of food crops, such as heirloom varieties, help protect farmers and gardeners from complete loss of spe cific crops asa result of damage or disease, and are important resources for the future of ang The author includes short personal stories of persons met through a variety of different research interviews. These stories put a real face on those persons who participate in seed saving. Often, these seed savers are not intentionally saving seeds as a method of retaining biodiversity, but may be subsis- tence farmers, collect interesting varieties from the wild, retaining seeds as part of their family history, planting seeds that have been handed down over generations, or preserving plants given to them by friends and neighbors. Often these “seed savers” cultivate plants in a way familiar to their family or region; practices that are dying out as a result of increasing development of commercial farming or highly managed agricultural sites. Other — savers, may have ~~ to new locations, bringing ult t r with them plant species from at y with uni n practi You may think this review Pavan surface of this ee ee youare correct. a found the topic of Heirloom Seeds and Their Keepers to be very interesting, however it was a very challenging book for me to read. Perhaps the ses saan in this book is common for books written on topics in e familiar with the descriptive phrases used anthropology and/or sociolo m«¢ to present research in this bad, or mays the niior ‘ee her thesaurus, but the need to decipher many phrases left me feeling as though | may not have gleaned all the information the author was providing. A suggestion | offer other readers is to be sure to take your time in reading this book, and keep a dictionary handy for careful consideration of terms and phrases such as “.decolonization of texture and vu human consciousness and imagination..,” “hermeneutic,” aoe press serendipity of encounters.” If life stories of seed y, &/orananalysis Se ecnite Seedsand Their Keep- en of seed Savers 1n onesies biodiversity int terests you, reac ers for yoursell.—Lee pee ie Herbarium, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 21(4): 2014. 2005 TAXONOMIC REVIEW OF ASTRANTHIUM INTEGRIFOLIUM (ASTERACEAE: ASTEREAE) Guy L.Nesom Botanical Research Institute of Texas ecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Astranthium integrifolium (Michx.) Nutt. has been treated as comprising two taxa (var./subsp. integrifolium and var. triflorum/subsp. ciliatum) or simply as a single species without formal in- fraspecific designation. The two taxa are completely allopatric and separated by features of vegeta- tive, floral, and cypselar morphology. Consistent with taxonomic ranking of other Astranthium spe- cies py ies two aes are Sees here at cuss rank: Astra a cee sensu stricto and af.) Nesom, comb. n I ellis ciliata Var. tri, lora Raf. (= A. ciliatum). RESUMEN Astranthium ee eMECEA) Nate se ha tatane comprenalende dos t taxa as paunep: INCE? ifolium ar. tri lor formal pie etnies Los dos taxa son completamente alopelees ie se separan por ‘ morfologia vegetativa, floral, y de la cipsela. De acuer stranthium, estos dos taxa se tratan aqui con rango enc Astranthium ae sensu stricto y Astranthium ciliatum (Raf.) Nesom, comb. nov. Se selecciona un neotipo para Bellis ciliata var. triflora Raf. (= A. ciliatum). The genus Astranthium Nutt. includes 12 species (De Jong 1965), all but three of them restricted to Mexico. Astranthium integrifolium (Michx.) Nutt. occurs in northeastern Mexico, but its range is mostly in the central U.S.A. Larsen (1933) and De Jong (1965) treated eastern and western population systems of A. integrifolium (Fig. 1) as var. integrifolium and var. ciliatum (Raf.) Larsen, re- spectively, or as subsp. integrifolium and subsp. triflorum (Raf.) De Jong. Shinners (1950) added a third taxon at varietal rank, A. integrifolium var. robustum Shinners, which subsequently was raised to specific rank by De Jong (1965). Since De Jong’s treatment, Texas botanists have maintained A. robustum (Shinners) De Jong as a Texas endemic (Fig. 1), but the taxonomy of the more broadly distributed A. Eee eiuoLae sensu lato has not been critically reevalu- ated. De J d morphological delimitation of the ‘integrifolium complex’ has been accepted in national checklists (e.g., Kartesz 1999), but Cronquist (1980) treated both geographic segments as A. integrifolium with- out formal recognition of infraspecific taxa. Other floristic accounts have dealt SIDA 21(4): 2015-2021. 2005 2016 BRIT.ORG/SIDA 21(4 = © Astranthium robustum be aa ra eo Ast thi ort g et «o Astranthium integrifolium Fic. 1.G ic distributi ifolium, A. ciliatum, and A. rebustum. S 1 solid circl d triangles are from pets le ny me Ont BRIT TEX. LL, and VDB.0 lished (D tion 1977; Smith 1988; USDA, NRCS 2001). Populations of A. iliatum ee occur in Mexico (Nuc uevo Leon and Tamaulipas). with only one or the other of the two taxa. Accounts for Oklahoma and Arkan- sas (e.g., Taylor & Taylor 1994; Smith 1994) also have referred to A. integrifolium without finer taxonomic distinction, while those for Texas and the Great Plains have recognized infraspecific variants (e.g., Correll & Johnston 1970; Barkley 1986; Diggs et al. 1999). A reevaluation of Astranthium integrifolium sensu lato indicates that the o previously recognized geographic segments are allopatric and distinguished by vegetative, floral, and cypselar features. Consistent with taxonomic ranks of other Astranthium species (see comments below), the two taxa are treated here at specific rank. TAXONOMY 1. santero ciliatum (Raf.) Nesom, comb. nov. Bellis ciliata Raf., New Fl. N. Amer. 24. 1837. Astranthium integrifolium (Michx.) Nutt. var. ciliatum (Raf.) Larsen, Ann. Mis- souri Bot. Gard. 20:35. 1933. Astranthium integrifolium (Michx.) Nutt. subsp. ciliatum (Ral.) De Jong, Michigan State Univ. Mus. Publ. Biol. Ser. 2:504. 1965. LECTOTYPE (De Jong 1965): USA. NESOM, TAXONOMIC REVIEW OF ASTRANTHIUM INTEGRIFOLIUM 2017 = TEXAS. AUSTIN Co.: San Felipe de Austin, 1835, T. Drummond II. 221 (NY website photol, as “Bellis integrifolia Michx.”; ISOLECTOTYPES: K, NY, P, PH BellisciliataR iflora Raf., New Fl.N. Amer. 2:25. 1837. Ast Nutt. var. pin aes Sida 2:348. 1966. NEOTYPE (selected hee) USA. TEXAS. Fannin Co., 4 mi N of Bonham, sandy ditch bank, 10 Jun 1945, L.H. Shinners 7842 (SMU)). Rafinesque noted that “it [presumably ‘Bellis ciliata’] has alsoa var. triflora, with leaves all acutish and only 3 [ray] flowers. The folioles of the perianthe are lanceolate acuminate in all the sp. not linear as Hooker says, the seeds are obovate pubescent.” Because Rafinesque noted that his new species, Bellis ciliata, occurs “in Texas and probably extending to Louisiana and Arkansas,” it seems reasonable to infer that Ben eat var. arlons aso} was epee ona moras collection, I f B.ciliata. +h pee | perhaps from Drummo De Jong (1965) also ie this taerences in enane that the type locality ae var. triflora was “Tex e indication of the type, and no type material of var. as. ” Rafinesque how triflora has been anne or cited in et literature. Small capitula with relatively few and small ray floret son in Astranthium integrifolium and A. ciliatum, a [have seen no plants with fewer than ikely Te roduced late in the sea- —_ six florets. Plants of the neotype were described as producing 6-8 ray florets. It seems that the capitulum observed ae described by Rafinesque had lost several ray florets during collecting, mounting, or han Astranthium integrifolium Moe vat. Serene Larsen, Ann. Missouri Bot. Gard. 20:36. 1933. Type: UNITED STATES. TEXAS. Matagorda Co.: Matagorda, sandy prairies, 5 Mar 1914, EJ. Palmer 4855 (HOLOTYPE: MO!). Shinners i Pere noted that the presence of ate clusters of leaves reflects early growth stages of a single plant. — rosu 2. Astranthium integrifolium (Michx.) Nutt, Trans. Amer. Philos. Soc., ser. 2, 7:312. 1841. Bellis integrifolia Michx., Fl. Bor. Amer. 2:131.1803. TYPE: UNITED STATES. [TENNESSEE] “Cumberland,” A. Michaux s.n. (HOLOTYPE: P, fide De Jong 1965). De Jong (pp. 434-435) in- ferred from historical accounts that Michaux probably made the collection in June in the vicinity of Nashville. Michaux himself (1803) noted that the species occurred “a rivulorum et in collibus umbrosis Tennassée.” “Cumberland” is the only ie refer- € ence on the holot Distinctions between Astranthium integrifolium and A. ciliatum are in the fol- lowing contrasts. Plants fibrous-rooted; basal and lower cauline leaves 3-6 cm long * 7-22 mm wide; involucres 3.5-6 mm high; ray corollas (6-)8-17 mm long; cypselae (1.4-)1.6-2 (-2.2) mm long X 0.9-1.1 mm wide, surface minutely papillate-pebbly with linear striations barely discernible, glabrous or sometimes sparsely glochidiate-pubescent near the apex, or 7 northern Kentucky and West Virginia) glochidiate-pubescent over the whole s Astranthium integrifolium Plants slender- see rarely fibrous-rooted; basal and lower cauline fees 5-4 (-5) cm long X 3-11(-14) mm wide; involucres (2—-)2.5—4.5 mm high; ray corollas (4-)6-10(-12) mm long;cypselae 1-1.6 mm long X 0.6-0.8 mm wide, surface with minute longitudinal striae but otherwise nearly smooth, not papillate-pebbly, sparsely to densely glochidiate-pubescent from base to apex Astranthium ciliatum The morphological distinction of Astranthium integrifolium and A. ciliatum corresponds with their geography, as mapped in Figure 1. The two are essentially completely separate in distribution: A. ciliatum occurs west of the Mississippi 2018 BRIT.ORG/SIDA 21(4) River, A. integrifolium to the east. The disjunct outlier in Holmes Co., Missis- sippi (Woodson and Anderson 1555, MO), is typical A. integrifolium, the outlier in Mississippi Co., Arkansas (Pyle 669, TEX!), is typical A. ciliatum. Astranthium integrifolium was first reported for West Virginia by Duppstadt (1992), without citation of vouchers. Details are given below. The limited, disjunct distribution and the occurrence primarily along roadsides suggest that this extended population system might be of recent origin, per- haps by accidental dispersal from Kentucky, but the habitats appear to be oth- erwise natural. ~— Collections examined: WEST VIRGINIA. Barbour Co.: along Pleasant Creek Public Hunting Area road, extending 1.5 mi, both sides of road into Taylor Co., 26 May 1991, Bush s.n. (W VA); Pleasant Creek hing Area, roadside, 6 Jun 1991, Clarkson s.n. W VA); along Co. Road 10, between Hwy blic Hunting Area; area of Prunus, Hunting & Fis 119 and Tygart Lake, N side of Pleasant Creek in Pleasant Creek Pu Liriodendron, Acer, Cornus, and Crataegus, with much invasive Rosa multiflora and Lonicera maachti, Astranthium locally abundant in grassy habitats on roadsides, roadbanks, and adjacent fields, 3 Jun 2002, Nesom FW154 (BRIT, GH, KY, NCU, OS, TENN, TEX, UARK, US, WVA). Taylor Co.: along Pleasant Creek Public Hunting Area road, extending 1.5 mi, both sides of road into Barbour Co., 2 Jun 1991, Baer s.n. (WVA). DISCUSSION De Jong (1965, p. 510) observed that “The two subspecies lof Astranthium integrifolium] are separated from one another by quantitative characters and may be recognized throughout their respective ranges. The occasional failure of a single character is compensated for by other characters. The key differ- ences between the two subspecies hold true when they are grown in the green- house under uniform conditions.” Nevertheless, De Jong noted that in Arkan- sas, subsp. ciliatum “overlaps” and “hybridizes” with subsp. integrifolium (p. 474 and p. 505). “The number of specimens which are thought to be putative hybrids is relatively numerous, but not enough specimens are available from different localities to assess the pattern of variation accurately” (p. 511). “The recognition of subspecies rather than varieties is prompted by the considerable ranges of subsp. integrifolium and subsp. ciliatum and the relatively narrow zone of intergradation” (p. 510). Intergrades between Astranthium integrifolium and A. ciliatum were cited by De Jong (1965) from Arkansas (e.g., Demaree 16912-SMU!, Harvey 45-MOl, SMU, Engelmann 129-MO), Oklahoma (not seen), and Missouri (e.g., Bush 7534- MOI, Steyermark 22642-MO)). He did not specify the nature of the intergrada- tion, but from sheets annotated by him, it can be inferred that this was prima- rily an interpretation of root morphology and general vigor (stem height and leaf size). The ‘intergrades’ cited from these states (all within the range of A. ciliatum, as recognized here) tend to be fibrous-rooted rather than taprooted, but rare plants from Texas (e.g., Gonzales Co., Turner 371I-SMU; Harrison Co., Orr 182-SMU) also are weakly fibrous-rooted; all of these are interpreted here NESOM, TAXONOMIC REVIEW OF ASTRANTHIUM INTEGRIFOLIUM 2019 as population variants rather than intergrades, because they belong with A. ciliatum in involucral height, ray corolla length, and especially in cypselar size, surface morphology, and vestiture. The same is true for two Arkansas collec- tions cited by De Jong as A. integrifolium subsp. integrifolium (Hot Springs Co., Soulard s.n.-MO!, Washington Co., Harvey s.n-MO!) and for Missouri collections annotated as “aff. subsp. integrifolium’ (e.g., Palmer 39297-MO], Palmer 39483- MOl, Steyermark 10393-MO!). Some Arkansas collections cited by De Jong as in- termediate between the subspecies were annotated by him simply as “A. integrifolium subsp. ciliatum” (e.g., Bush 929-MO), Palmer 5992-MO)). Robust plants of Astranthium ciliatum are similar in habit to A. integrifolium, and depauperate plants of A. integrifolium are similar in habit to A. ciliatum. The Arkansas collections interpreted by De Jong as “subsp. integrifolium” or as reflecting hybridization were made around Hot Springs (Hot Springs Co.) and Fayetteville (Washington Co.). These plants are gener- ally taller than average for the species (up to 32 cm tall) and have leaves that range larger, and some have fibrous roots, but in involucral size, ray length, and cypselar morphology, they belong with A. ciliatum. Cypselae of plants from Arkansas and Missouri also range longer (1.1-1.6 mm) than in Texas and Okla- homa (1-1.2 mm). Whether the larger sizes of these plants might reflect con- vergence or an ancestral similarity with A. integrifolium sensu stricto is not clear. Even though various features overlap in variation, features of cypselar morphology (vestiture and epidermal surface) provide consistent distinction between the two taxa, especially with recognition of the disjunction in their geographic ranges, and I have not seen any plant that could be regarded as in- termediate. Insummary, the present study finds that there is no unequivocal evidence for HYP RCIEauOn peevecn pant of aor ae and A. ciliatum. In face tl their allopatri t unity for genetic interchange, and differences in cypselar morphology suggest a isolation is complete. CONSISTENCY IN TAXONOMIC RANK Morphological distinctions between Astranthium integrifoliumand A.ciliatum are relatively small, but their pattern of relationship and treatment at specific rank are analogous and consistent with the taxonomy of species pairs (as rec- ognized by De Jong 1965) of Astranthium found in Mexico. The two are essen- tially identical in chromosome number (2n = 8) and chromosome morphology (De Jong 1965). Their northern distributions and morphological similarity sug- gest that they have an evolutionary sister relationship. De Jong (p. 523) noted that A. robustum (2n = 6) “is related to A. integrifolium’ ... but “the species also shows resemblance to A. condimentum and A. orthopodum and has the same chromosome number and karyotype as these two species.” The relationship of Astranthium orthopodum (BL. Rob.) E. Larsen and A. 2020 BRIT.ORG/SIDA 21(4) condimentum De Jong parallels that of A. integrifolium and A. ciliatum. They are similar to each other in chromosome number (2n = 6) and chromosome morphology and probably are evolutionary sister taxa. Each has a substantial geographic range, but they are allopatric in distribution. They are morphologi- cally separated primarily on the basis of root characters: plants of A. condimentum are annuals from a slender taproot; plants of A.orthopodum are biennials or short-lived perennials from a fibrous-rooted caudex, sometimes with short rhizomes or basal offsets. Additionally, there are small and overlap- ping differences in stem orientation and leaf shape and size. Astranthium splendens De Jong (2n = 18) and A. beamanii De Jong (2n = 24) both apparently have a base chromosome number of x = 3 and also prob- ably are evolutionary sister taxa. They are sympatric but grow at different el- evations and are otherwise distinguished on the basis of cypselar vestiture (and apparently nothing else): cypselae of A. splendens are glochidiate-hairy over the whole surface while those of A. beamanii are glabrous or sparsely glochidiate-hairy only near the apex. ACKNOWLEDGMENTS Loans of specimens from KY, MO, WKU, and WVA, help of TEX-LL staff dur- ing a recent visit, help from Donna Ford- Werntz (W VA) and Bill Grafton in the investigation of the West Virginia plants, and review comments of George Yatskievych are gratefully acknowledged. REFERENCES Barktey, I.M. 1986. Astranthium. In: Great Plains Flora Association. Flora of the Great Plains. Univ. Press of Kansas, Lawrence. P. 886. Cuester, E.W., B.E.Worroro, and R. Krat. 1997. Atlas of Tennessee vascular plants. Volume 2. Angiosperms: Dicots. Misc. Publ. 13, Center for Field Biology, Austin Peay State Univer- sity, Clarksville, TN. Corrett, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner Cronauist, A. 1980.Vascular flora of the southeastern United States, Vol.|. Asteraceae. Univ. of North Carolina Press, Chapel Hill. De Jon, D.C.D.1965.A systematic study of the genus Astranthium (Compositae, Asteraeae). Michigan State Univ. Mus. Publ. Biol. Ser. 2:429-528. Diccs, G.M., Jr., B.L. Liescome, and RJ. O’Kennon. 1999, Shinners and Mahler's illustrated flora of north central Texas. Sida, Bot. Misc. No. 16. DuppstaoT, W.H. 1992. Updates on the vascular flora of West Virginia: 8. Proc. West Virginia Acad. Sci.64:15. Great PLAINS FLORA ASsociATION . 1976. Atlas of the flora of the Great Plains.lowa State Univer- sity Press, Ames. KarTesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the NESOM, TAXONOMIC 2021 vascular flora of the United States, Canada, and Greenland. In: Kartesz, J.T. and C.A. Meacham. Synthesis of the North American Flora, North Carolina Botanical Garden, Chapel Hill. Larsen, E.L. 1933. Astranthium and related genera. Ann. Missouri Bot. Gard. 20:23-44. MicHaux, A. 1803. Flora Boreali-Americana.Levrault, Paris. SHINNERS, L.H. 1950. Notes on Texas Compositae-VI. Field & Lab. 18:156-159. Smith, E.B. 1988.An atlas and annotated list of the vascular plants of Arkansas (ed.2). Dept. of Botany & Microbiology, Univ. of Arkansas, Fayetteville. Smith, E.B. 1994. Keys to the flora of Arkansas. Univ. of Arkansas Press, Fayetteville. Taytor, RJ. and C.E.S. Tayior. 1994. An annotated list of the ferns, fern allies, gymnosperms and flowering plants of Oklahoma (ed. 3). Biology Dept. Herbarium, Southeastern Okla- homa State Univ., Durant. USDA, NRCS. 2001.The PLants Database, Version 3.1 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490, USA. 2022 BRIT.ORG/SIDA 21(4) Book REVIEW LYNNE RICHARDS and RONALD J. Tyr. 2005. Dyes from American Native Plants: A Practical Guide. (ISBN 0-88192-668-X, hbk.). Timber Press Inc. 133 S.W. Second Ave, Suite +50, Portland, OR 97204-3527, US.A. (Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $29.95, 340 pp., 155 color photos, 6" x 9" 7 itil ] c 4 } th tal US. Introductor CHIAPLETS This book covers 158 natural dye plants that are native cover the history of natural dyes, natural dyeing processes, snd SUP] okies needed to use natural dyes at home. The authors tested 158 native dye plants using five different mordants and two different dyeing sults are carefully catalogued in seven detailed chapters arranged by color: purple, processes. The re ludes a table of dye plants, dye process, red, green, yellow, orange, brown, and black. Each chapter in mordant, and the color produced, along with chips of each co A field guide at the end of the book includes color photographs of live plants, common and book. This guidebook to natu- a or. scientific names, and a short description of each ae included in the ral dyes is an excellent reference book for economic botanists, textile artists, and crafters interested in natural ne —Marissa Oppel, Herbarium Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 76012-4060, U.S.A. SIDA 21(4): 2022, 2005 SFUDIES OF NEOTROPICAL COMPOSITAE 1 NOVELTIES IN CALEA, CLIBADIUM, CONYZA, LLERASIA, AND PLUCHEA John F. Pruski Garden Missouri Botanica ‘Oo Notes PO. St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT The combinations caled auneditcranea (Vel! lab sa lea platylepis) and Calea triantha ( Vell.) Pruski, (syn: Cal ies | ified | in by Velloso Tees A ieee of Meyeria hispida DC: is designated. A key to the ee centering about Calea myrtifolia is given. Clibadium arriagadae Pruski (Heliantheae) from Ecuador is named as a new segregate of Clibadium pentaneuron. Clibadium arriagadae is the same taxon as represented by the invalid Clibadium zakii. Clibadium arriagadae differs from C. pentaneuron by leaf blades ea 3- or 5- ae monte or near base (vs. subpalmately or plinerved from above blade base) and hirsute (vs A lectotype (BM-CLIFF folio page 405, Coma oe is designated lor Conyz bifronsL.@ cee bifrons L., Inuleae), and this name Pluchea, is one ae the flora of the New World. The eomibinaden Conyza popayanensis een) rs cASIETEAE), repaeing the mesilate Cay a ungiC (Benth.) Cuatr, non. Pers., is propose € ) validated fora Boliv- 1asasynonym of ian species cially described in tribe hianiciene and Llerasia lucidula is L. macrocephala. Philip Miller's Conyza baccharis is lectotypified and is an earlier name for Pluchea rosea. The combination Pluchea baccharis (Mill.) Pruski (Pl ucheeae) is made for this coastal plain species, which occurs from eastern North America southward into Nicaragua. RESUMEN € proponen las combinaciones Saea meditemanea (Vell) Pruski ee Calea platylepis) y Calea ne Vell.) Pruski, (sin.: Calea hist ae) para dos especies que se lectotipifican aqui mediante ilustraciones de Velloso. Se designa un oot para ae hispida DC. Se ofrece una 1 epee es clave a las sopecies proximas a Calea myrtifolia. Clibadium arriagadae Pruski (Heliantheae) de Ecuador mo un nuevo segregad ode Clibadium a Clibadium arriagadae es re eee | el t por el nombre invalido Clibadium zakii adium arriagadae difiere ec. pentaneuron por los limbos de las hojas palmatinervias con 3 6 nervios desde de la base o cerca (vs. subpalmatinervias o triplinervias desde mas arriba de la base) e hirsutas (vs. estrigosas) abaxialmente. Se designa un lectotipo (BM-CLIFF folio pagina 405, Conyza 3) para one bifrons L. (= Inula bifrons L., Inuleae), y este nombre, por hab Pluchea, se excluye de la flora del Nuevo Mundo. Se propone la donibinceion Cone popayanensis (Hieron.) ee eae pe pecuaaae a i “ ue Conyza uliginosa (Benth.) Cuatr, non. Pers. Se valida } Y Prick] (A : Jo RAlixza | it rt te originalmente en la tribuMutisieae, : lucidulaset inoénimo de L. macrocephala Conyz se lectotipifica y es un nombre anterior para Pluchea rosea. Se hace a combinacioén Pluchea Beeches (SL ) Pruski (Plucheeae) para esta especie de la Hanura costera, que aparece desde el Este de Norte América hasta Nicaragua en el Sur. SIDA 21(4): 2023-2037. 2005 2024 BRIT.ORG/SIDA 21(4) The purpose of this note is to validate names in Calea L., Clibadium F Allam. ex L., Conyza Less., Llerasia Triana, and Pluchea Cass. needed in various floris- tic works of Neotropical Compositae being done at the Missouri Botanical Gar- den, and to lectotypify Conyza bifrons L., which is excluded from the flora of the Americas. — CALEA Jose Velloso prepared Flora fluminensis, a landmark flora for the environs of Rio de Janeiro, Brazil in 1790, but died in 1811 prior to its publication. The text was printed in 1825 (Velloso 1825) and distributed in 1829. The text, however, remains incomplete, treating species corresponding to icones published only in volumes I-8 (8 pro parte). No text is available for species illustrated volume 8 (pro parte), nor for those in volumes 9-11. The Compositae comprise all of vol- ume 8 and part of volume 10, but text is available for about only the first half of the 164 icones (Velloso 1827) published in volume 8. The 11 volumes of figures are dated 1827 (Velloso 1827), but were distributed only in 1831. Thus, 1831 is taken as the date of validation of species represented only by these diagnostic icones, but without corresponding text. It is the plates of two such Velloso names (Figs. 1 & 2) that are clearly identifiable with two South American species of Calea (Heliantheae; syn. Neurolaeneae), new combinations for which are made below. No type specimens of these two names are known to exist, and conse- quently the illustrations are designated as the lectotypes. ee Calea mediterranea (Vell.) Pruski, comb. nov. (Fig. 1). Buphthalmum mediterraneum Vell, Fl. Flumin. (cones) 8 t. 135, 1827 [1831]. Lectotype (designated here): t. 135, Vell. FI. Flumin. (cones) 8. 1827 [1831]. Caled platylepis Schultz-Bip. ex Baker in Martius, FL. bras. 6(3):267. 1884. Lectotypification from among the dozen or so syntypes is deferred Distribution and ecology.—Calea mediterranea (Vell.) Pruski isa xylopodial sub- shrub flowering from October to April. It occurs in the Brazilian planalto south- wards into Paraguay and northern Argentina. Calea mediterranea is a member of Caled sect. Haplocalea (Less.) Pruski (Pruski 1998 sub Calea platyle pis), and is closely related to C. cymosa Less., the type of the section. This section is characterized by generally whorled leaves and umbelliform capitulescences. Calea mediterranea diflers from C. cymosa Less. by narrower, pubescent, pinnately veined (vs. broader, nearly glabrous, 3- or 5-plinerved) leaves and by lanceolate, pubescent (vs. ovate, glabrous) outer phyllaries. Additionally, the lower most whorl of leaves in C. mediterranea is often reduced, as illustrated by Velloso (1827), a feature aed ie of C.cymosa. The radiate capitula of C. mediterranea and C. cymosa distinguish each of them from the otherwise closely related C. hassleriana C hod. and C. ret lauler: Gard- ner (syn. Ichthyothere ternifolia Baker), which have discoid capitula. PRUSKI, STUDIES OF NEOTROPICAL COMPOSITAE 2025 Syno: Polvo: Super IY BUPHCT HAL MUM MUR IDICIUEIRIR ANU MI (Tab. 155) Fic. 1. Lectotype of Buphthalmum mediterraneum Vell. [= Calea mediterranea (Vell.) Pruski], from Velloso, Fl. Flumin. (Icones) 8: t. 135. 1827 [1831]. 2026 BRIT.ORG/SIDA 21(4) Sy Nig’. Poly Q: Superl. AS TER WRAAR WT UYS ¢Tablaxo.) PRUSKI, STUDIES OF NEOTROPICAL COMPOSITAE 2027 Calea triantha ( Vell.) Pruski, comb. nov. (Fig. 2). Aster trianthus Vell, Fl. Flumin. (cones) 8: t. 120. 1827 [1831]. Lectotype (designated here): t. 120, Vell. Fl. Flumin. (Icones) 8.1827 [1831]. Meyeria hispida DC,, Brods aa ee Calea hispida ee) pas in Martius, Fl. Bras. 6(3):261. 884. LECTOT O PAULO, campis editis, Nov. 1 1833, Lund 866 (LECTOTYPE: G-DC [IDC raiestiche 800. 975.II1.3], ISOLECTOTYPES: C-4, S). The d to the lower right of the G-DC lectotype also has a small twig of H. Imp. Bras. 405 mounte Lund collection. [Lectoparatype: Herb. Imperial Brasil (probably Vauthier) 405: G-DC (frag- ment of a sheet from P), F (fragment of a sheet from P), P-2. Distribution and ecology—Calea triantha (Vell.) Pruski is a subshrub to shrub endemic to Brazil, where it is centered in the state of Parana. It is known to flower from December to April. Calea triantha is one of 13 species of the C. myrtifolia DC. species group (sensu Pruski 1984; Pruski & Urbatsch 1988) of Calea section Meyeria (DC) Benth. & Hook.f. Pruski and Urbatsch (1988) provided a key to the then-known members of this group. Their key is revised herein, incorporating the above new synonymy and C. semirii Pruski & Hind, which was described subsequently (Pruski & Hind 1998). KEY TO THE SPECIES CENTERING ABOUT CALEA MYRTIFOLIA (CALEA SECT. MEYERIA) Calea marginata 1. Leaves glabrous, entire, margins thickened (Sdo Paulo and Parana, Brazil) S.F. Blake 1. Leaves glabrous to hispid, entire to serrate, margins not thickened. eaves lanceolate, ca. 6-12 cm long, venation parallel, ca. 3-7-veined (Goias, Brazil) Calea nervosa Barroso 2. Leaves lanceolate to cordate, less than 6.5 cm long, venation pinnate to trinervate. pitula one per branch. 4. Leaves Moen ee foveolate below (Sao Paulo and Parana, Brazil) Calea Baas (DC.) Baker Calea ilienii Malme Calea monocephala Dusén 4. Leaves elliptic-ovate, smooth below. 5. Peduncle ca. 8-15 cm long (Parana and Santa Catarina, Brazil) 5. Peduncle ca. 1.5-6 cm long (Parana, Brazil) 3. eee cymose. eaves whorled. 7. Leaves ee four per node, essentially sessile, smooth, pinnat duncles 3-15 cm long; ray corolla limb 5-13-nerved; outer phyllaries and leaves green below; pappus scales . in length (Distrito Federal and Goias, Brazil) ea nected ihe & Urbatsch . Leaves generally Oise per node, Solty © =e rugulose, trin peduncles 0.5-6 cm long; ray lla lirr nerved;outer phyllaries leaves rust-colored below, less Lait green; pappus scales slightly to greatly unequal in length (Minas Gerais, Brazil). 8. Leaves adaxially glabrous or nearly so; involucres campanulate to hemispherical; outermost phyllaries at least apically herbaceous, about as long as the next ser ies; ray COrO ollas yellow, tube long, limb 12-14.5 mm long, commonly abaxially glandular; disk e; pe- N 2028 BRIT.ORG/SIDA 21(4) corolla tube shorter than the throat; cypselae 2.5—3.2 mm long, wit! 1-3 pappus scales often much longer than the others ____ Calea ioeroodeps Pruski & Urbatsch 8. Leaves adaxially hispidulous to sparsely pilose; involucres turbinate to cylindrical; outermost t phyllaries mostly scarious, usually much shorter than the next series; llas pale yellow, tube 3.6-< orollas 4.5mm long, limb 7-8. mm aioe sbavielly eglandu ar diskeoraila tube about as-long-as the throat; cypselae 3.8-4.8 mm long, pappus scales gen- erally slightly unequal ____ Calea semirii Pruski & Hind 6. Leaves opposite 9. Leaves ovate, Aiepiciiouie to hispid, serrate, basally cordate, paee shorter than 2.5 cm (Minas Gerais south to Santa Catarina, Brazil:s | hispida) Calea ie triantha (Vell.) Pruski 9, Leaves elliptic to elliptic-ovate es to seer to serrate, basally cuneate, longer than 2.5 10. Leaves to 3.5 cm wide, shiny, serrate, glal ‘foli phyl- laries serrulate (P ea Calea chodatii Hassler 10. Leaves less than 2.5 cm wide, somewhat shiny or not, entire to ser- rulate, glandular or puberulent; fo aoe Cuter Bee entire. 11. Leaves narrowly elliptic; capitula ca ay lla limb 8-10-nerved; disk corolla lobes ae and 15 mm; egeseee glabrous (Rio Grande do Sul, Brazil and Uruguay) Calea kristiniae rUSKI . Leaves elliptic to icon ave capitals & ca. 35- HOWeleg fey corolla limb 5(—7) 1.5m cypselae pubescent on angles. 12. Leaves entire, ca. 1.5 cm wide (Minas Gerais south to Rio Grande do Sul, Brazil) ____ Calea a oie (DC.) Baker 12. Leaves serrulate, to ca. 2.5 cm wide (coastal S40 Paulo sou to Rio Grande do Sul, Brazil) Calea Tsien Baker ’ CLIBADIUM Arriagada (1995) provided an overview of Clibadium (Compositae: Heliantheae), including full synonymy and a key to species. In this overview. Arriagada reduced to synonymy all four northern South American names of Clibadium proposed by Robinson (1992). A monograph of Clibadium has now been published (Arriagada 2003), and again all names in Clibadium proposed by Robinson (1992) are listed as synonyms. lagree with Arriagada (1995, 2003) that C. pentaneuron SF. Blake includes the synonymous C. funkiae H. Rob; C. laxumS.F Blake includes C. alatum H. Rob.;and C.glabrescens S.F Blake includes C. napoense H. Rob. Thus, three validly described species in Robinson (1992) are encompassed within more widespread taxa, these originally described by Blake in the 1920s and 1930s. A fourth name (C. zakii) in Robinson (1992) was based on Zak & Jaramillo 2881 (MO, US) but without “the single herbarium” housing the holotype being “specified.” This name is consequently invalid (Art. 37.6 of the Code, Greuter et al. 2000) and does not exist nomenclaturally. This plant belonging to Clibadium section Clibadium is validated here with a new PRUSKI, STUDIES OF NEOTROPICAL COMPOSITAE 2029 type collection and a new epithet honoring my friend Dr. Jorge Arriagada, the monographer of the genus. Clibadium arriagadae Pruski, sp. nov. (Fig. 3). Type: ECUADOR. Cotopaxt road between Quevedo & Lacacunga, 76 km E Quevedo, 0°57'S, 79°O1'W, 2300 m, 5 Apr 1983, Croat 55804 (HOLOTYPE: MO; ISOTYPE: QCNI ] : d ] A C. pentaneuron alfinis, sed laminae | vel pinnativenosae), rotundata vel truncata (non cuneata), et subtus hirsuta (non strigosa) diversa. Shrubs to ca. 2 m tall; stems sometimes vining, subterete to subhexagonal, hir- sutulous. Leaves simple, opposite, petiolate; petioles 0.8-3.5 cm long, hirsute; blades broadly ovate, 4-15.5 cm long, 2-11 cm wide, stiffly chartaceous, pal- mately 3- or 5-veined from or near base, secondary and tertiary reticulation prominent, base rounded to truncate, margins serrulate, apex acute to acumi- nate, the adaxial surface scabrid, hirsutulous, the abaxial surface hirsute, eglandular. Capitulescence terminal, many-headed, loosely corymbiform pan- iculate, branches 2-14 cm long, hirsute, ultimate branching t1 ichotomous. Ca- pitula 10-12-flowered, disciform, shortly pedunculate, 4-5 mm tall; involucre hemispherical; phyllaries ca. 3-seriate, subequal to weakly graduated, stiffly chartaceous, weakly 3-5-veined adaxially, apically hirsutulous, otherwise gla- brous, toca. 4.5mm long, 2-2.5 mm wide, outer ones pyriform, apically acute to acuminate, mid-series and inner ones ovate, apically obtuse to rounded; recep- tacle convex-conical, to ca. 1 mm broad, weakly paleate; paleae lanceolate, to ca. 3mm long, weakly conduplicate; peduncles 1-2 mm long, terete, glabrous to hirsutulous, one-bracteolate, bracteole lanceolate, 1-2 mm long, hirsute. Mar- ginal florets uniseriate, pistillate, 5 or 6, mostly included within involucre; co- rolla ca. 2 mm long, tubular, cream-colored, apically pilosulose with non-glan- dular trichomes, minutely ca. 3-lobed, style branches ca. 1 mm long, weakly exserted. Disk florets functionally staminate, 5 or 6, mostly included within involucre; corolla broadly funnelform, ca. 2.5mm long, cream-colored, 5-lobed, lobes deltoid, erect, 0.5-0.9 mm long, pilosulose with non-glandular trichomes, anthers generally included, to ca. 1.7 mm long, dark greenish to black, append- age elongated but not greatly sculptured, basally short-sagittate, filaments ca. 0.3mm long; style undivided, apex often exserted from corolla; ovary rudimen- tary. Cypselae oblong, flattened, 2-2.5 mm long, apically papillose, otherwise glabrous or nearly so, epappose. Paratype: ECUADOR. Botivar: Carretera Chillanes-Bucay, en la hacienda ‘ head del Sr. Gonzalo Gomez, 1°55'S, 79°05’ W, 2100 m, 10 Sep 1987, Zak & Jaramillo 2881 (F, MO, US). Distribution and ecology—This species is known only from the Pacific drain- age slopes of the Andes in Bolivar and Cotopaxi, Ecuador. It has been collected in flower in April and September from 2100-2300 meters elevation. Clibadium arriagadae differs from C. pentaneuron by leaf blades mostly palmately 3- or 5-veined from or near base (vs. subpalmately or plinerved from 2030 BRIT.ORG/SIDA 21(4) ° ° . nh Lt . : : . Fic. 3. Cibad: ,} Ah R hy £ I h / \ 1 | kK | Jt £hLil«ad e- (Photographs of the holotype, Croat Nn * £ y VETIGUIUTT TOIT Very 55804, MO). PRUSKI, STUDIES OF NEOTROPICAL COMPOSITAE 2031 well above blade base), these rounded (vs. cuneate) basally and hirsute (vs. stri- gose) abaxially. The types of the C.funkiae H. Rob,, C. pileorubrum Cuatrec.,, C. sarmentosum Cuatrec., and C. scandens Cuatrec. each have leaves strigose abaxially and these four names are accepted as synonyms of C. pentaneuron, as proposed by Arriagada (1995, 2003). The stems of C.arriagadae are hirsutulous throughout, thus differing from those of C. pentaneuron (Ecuador and Colom- bia), which are sometimes str Arriagada (1995, 2003) ad C. funkiae (Antioquia, Colombia) to syn- onymy of C. pentaneuron. At one point, | thought that the prominent resin ducts in the phyllaries some material from Antioquia and in Forero et al. 2279 (MO; Choc6, Colombia near border with the northern limits of Valle, Colombia) could be used to distinguish this material from the generally more southern C. pentaneuron. However, the collection from Chocois near the center of distribu- tion of C. pentaneuron, and other material of C. pentaneuron from Antioquia lacks the prominent phyllary resin ducts. Thus, there seems to be no meaning- ful morphological features that one could use to split the Colombia material of C. pentaneuron into more than a single taxon. Indeed, this was the observation of Arriagada (1995, 2003), the monographer who reduced C. funkiae to syn- onymy. Moreover, no case for geographic separation of C. funkiae as distinct from C. pentaneuroncan be made. Thus, | recognize C. pentaneuroninaslightly narrower concept than does Arriagada (1995, 2003), and segregate only Clibadium arriagadae from it. The morphological distinctions among relatives of C. pentaneuronare pro- vided below in the key to species, which modifies couplet #27 of Arriagada (1995, 2003) KEY TO THE SPECIES CENTERING ABOUT CLIBADIUM PENTANEURON 27. Abaxial leaf surface hirsute; leaf blade generally palmately veined from or near base (Ecuador) Clibadium arriagadae Pruski 27. Abaxial leaf surface rally strigose; leaf blade pinnately veined or if plinerved then from well above ies 27.1 Most phyllaries apically acute to ere capitula 24—28-flowered; marginal florets 9-13; disk florets ca.15 (Ecuador)__—sCilibaddium manabiense H. Rob. 27.1 Most phyllaries apically obtuse to ac ett 10-14-flowered; marginal florets 5 or 6; disk florets 5-8 (Colombia and Ecuador) Clibadium pentaneuron S.F. Blake CONYZA Conyza bifrons L. (= Inula bifrons L., Inuleae) is lectotypified upon material from the Old World, and this name is thus excluded from the flora of the New World. The new combination C. popayanensis (Hieron.) Pruski (Astereae) from the Andes is proposed to replace the illegitimate C. uliginosa (Benth.) Cuatr. non Pers. Although C. primulifolia (Lam.) Cuatrec. (which includes as a synonym 2032 BRIT.ORG/SIDA 21(4) C.chilensis Spreng. the type of Conyza) was transferred to Erigeron L. by Greuter (2003), I recognize Conyza at the generic rank. Inula bifrons L., Sp. PL. (ed. D112 46. (ras Lee TOTYPE (designated by Anderberg, Taxon 47:363. 1998): EUROPE: “Habitat in Italia, Galloprovincia, Pyrenaeis,” sin. coll. (LINN 993.11 [IDC mi- crofiche 177. 577.115). Conyza bifrons L. var, bifrons, Sp. PL. 861.1753. Pluchea bifrons (L) DC, Prodr. 5:45]. 1836, LECTO- Type (designated here): EUROPE: “Habitat in Pyrenaeis, ce sin. coll. (BM-CLIFF folio page 405, Conyza 3 [barcode BM00064704 3], eee aph MO Conyza bifrons var. flosculosa L., Sp. PL. 862. 1753. LECTOTYPE coe ited by Reveal, Taxon 47:358. 1998): EUROPE: “Habitat in Dyce Canada,” sin. a (BM-SLOANE vol. 96, page 26). Conyza bifrons var. radiata L., Sp. Pl. 861.1753. Lectotype (designated by Anderberg, Taxon 47:358. 1998): t. 127 as “Conyza pyraenaica folits primulae veris” in Hermann, Parad. Bat., 127. 1698 Distribution and ecology.—Inula bifrons isa summer flowering herb to | m tall It occurs from the Pyrenees of southern France eastwards into Romania and Bulgaria (Tutin et al. 1976). Linnaeus (1753) named Conyza bifrons L.and two varieties of it, giving the locality of “Habitat in Eanes Canada” for all three names. Later, Linnaeus (1763: 1207) treated C. bifrons as being solely American and represented by Plukenet plate 87 figure + (1705), thought to have been drawn from Canadian material, whereas simultaneously Linnaeus proposed the heterotypic Inula bifrons L.(1763: 1236), with similar auriculate-clasping leaves, for the European elements. Pluchea bifrons(L.) DC. (Plucheeae) was misapplied to material from the Americas by Candolle (1836), as noted by Godfrey (1952), who used the name P foetida(L.) DC. for American plants formerly called P. bifrons. Gray 1884: 225) noted that the Plukenet plate was drawn from material collected in Europe, not Canada; thus no original material of either C. bifrons L. or I. bifrons L. is from the Americas. Conyza bifrons L., however, has not previously been lectotypified (C. Jarvis, pers. comm.). Linnaeus (1763) restricted the concept C. bifrons (1753) to plants he thought to be American, thus potential for misapplication of this name to plants from the Americas remains. Because Linnaeus (1753, 1763) cited a poly- nomial from Linnaeus (1737), C. bifrons L. is lectotypified here by a specimen in the Clifford herbarium, this specimen referable taxonomically to I. bifrons L. (Inuleae). Conyza bifrons L. is thus excluded formally from the flora of the New World, in agreement with Godfrey (1952) and Tutin et al. (1976). The name C. uliginosa (Benth.) Cuatrec., used for a northern Andean herb (e.g., Aristeguieta 1964; Cuatrecasas 1967; Jorgensen & Leon-Yanez 1999), is an illegitimate later homonym of C. uliginosa Pers., Synops. 2:427. 1807. A new combination based on the senior synonym given by Cuatrecasas (1969) is thus proposed. — Conyza popayanensis (Hieron.) Pruski, comb. 1 nov. pS ron popayanensis Hieron., Bot. Jahrb. Syst. 28:586. 1901. Type: COLOMBIA. CAUC amo de Guanacas, Andium centralium PRUSKI, STUDIES OF NEOTROPICAL COMPOSITAE 2033 popayanensium, 3000-3500 m, Aug. no year given, Lehmann 7962 (HOLOTYPE: B, destroyed, photograph sub Field neg. #14855: MO; LECTOTYPE (designated by Cuatrecasas, Webbia 24:217. 1969): K; ISOTYPES: FI, P. US [photograph: MO)). Erigeron uliginosus Benth., Pl. Hartw. 204. 1845, as “uliginosum.” Conyza uliginosa (Benth.) Cuatrec., Webbia 24:216. 1969, non Pers. 1807. TYPE: ECUADOR: PICHINCHA: In uliginosis ad Hacienda de Chisinche sub Volcan IIliniza (as “monte Ilinissa” in see ue), 1842, Hartweg 1131 (HOLOTYPE: K; ISOTYPES: G [photograph sub F neg. 28634: MO], NY [photograph: MO}, P. wh! Hlacionce de C pisInche: isa anew kms NE of Hliniza, thus resin in Prov. Pichineher Bot. Jahrb. Syst. 28:586. 1901, as “columbiana.” eee Siiadies var. columbiana (Hieron.) Cuatrec., seen 9:5. 1963. COLOMBI CUNDINAMARCA: In silvis montanis densis locis pumidTs supra Sibaté, 2800 m, 3 Feb — Lehmann 2535 (HOLOTYPE: B, destroyed; ISOTYPE: Erigeron bonariensis var. meridensis Cuatrec., Trab. Mus. Nac. Ci. Nat., Ser. Bot. 33:132. 1936. LEC- TOTYPE (chosen from among syntypes by Cuatrecasas, Webbia 24:216. 1969): VENEZUELA. vee Serra Nevada de Mérida, s.d., Moritz 1373 (LECTOTYPE: P). “Moritz 1373” was Rotcites specifically in the protologue, but rather only indirectly by name ee of “Sch. Bip schedam.” It seem best to accept the lectotypification of Cuatrecasas (196€ Distribution and ecology.—Conyza popayadnensis (Hieron.) Pruski occurs from 2500-4400 meters elevation in the Andes of Venezuela, Colombia, Ecuador, and Peru. Conyza popayanensis is a branched perennial herb with sessile leaves and a generally dense corymbiform capitulescence with peduncles generally much shorter than 5mm. The capitula have pubescent subequal long-triangular phyl- laries witha broad central colored portion and broad stramineous margins, and the marginal florets have entire or nearly so corolla limbs generally about 0.5 mm long. Cuatrecasas (1969) noted that one of the two plants on the destroyed Berlin holotype had an open capitulescence. Nevertheless, this plant on the de- stroyed holotype has weakly pubescent leaves typical of this species. Colombian Conyza uliginosa var. hirsuta (Hieron.) Cuatrec. [syn.: Erigeron uliginosus var. hirsutus Hieron., Bot. Jahrb. Syst. 28:587 1901, as “hirsuta”] was recognized by Cuatrecasas (1969). This taxon resembles C. popayanensis, but has more densely pubescent herbage, an open capitulescence, peduncles to 20 mm ong, narrower phyllaries, and marginal florets with sometimes deeply bifid corolla limbs often to about | mm long. Because I have not seen type material of this name, I decline to synonymize it or to recognize it at the species rank. If further study shows that C. uliginosa var. hirsuta deserves specific recognition, it should be noted that Chinese C. hirsuta L. blocks the transfer to Conyza of this varietal name. —y LLERASIA The below combination is provided for a Bolivian species of Compositae tribe Astereae, originally described as a species of tribe Mutisieae. Llerasia ee antes (Rusby) Pruski, comb. ney Se macrocephala Rusby, Descr. r. PL.162.1 abrera, Notas Mus. La Plata, Bot. 15:41. 2034 BRIT.ORG/SIDA 21(4) 1950. TYPE: BOLIVIA. LA PAz: North Yungas, Unduavi, 3300 m [as “3000 m” in protologue], Nov 1910, Buchtien 3080 (HOLOTYPE: NY; ISOTYPE: US [holotype of Haplopappus lucidulus S.F Blake). Haplopappus lucidulus S.F Blake, Amer. J. Bot. 4:114. 1927 (as “Aplopappus”). Llerasia lucidula (S.F Blake) Cuatrec., Biotropica 2:43. 1970. Type: BOLIVIA. LA PAz: North Yungas, Unduavi, 3300 m, Nov 1910, Buchtien 3080 (HOLOTYPE: US; IsoTYPE: NY [holotype of Moquinia macrocephala Rusby). Distribution and ecology.—Llerasia macrocephala (Rusby) Pruski is a vining shrub occurring from 2500-3300 meters elevation in Bolivia. Because of similar leaf surfaces occasionally closely tomentose abaxially, species of Vernonieae (especially those of Piptocarpha) and discoid species of Mutisieae (especially those of Gochnatia H.B.K.) are occasionally confused. For example, Badillo 1994), Pruski (1997), and Sancho (1999) treated Piptocarpha upatensis V.M. Badillo as a species Gochnatia. Stifftia axillaris G.M. Barroso & Vinha, described by Barroso and Vinha (1970) as a species of Mutisieae, was recognized by Robinson (1979) as a species of Piptocarpha (Vernonieae). Simi- larly, Moquinia macrocephala, described by Rusby (1920) as having leaf sur- faces closely sntose abaxially and asa species of Mutisieae (in the Gochnatia generic alliance), is here treated as a member of tribe Astereae. Rusby (1920) described Buchtien 3080 (NY) as Moquinia macrocephala (Mutisieae). Later, Blake (1927) noted that at US Buchtien 3080 was filed as an undetermined species of Gochnatia, but actually belonged to tribe Astereae, where he described it as Haplopappus lucidulus. Blake (1927) treated most of the ll species of Llerasia as Haplopappus [sub “Aplopappus’| sect. Diplostephioides (Benth. & Hook. f.) S.F Blake, whereas Cuatrecasas (1970) res- urrected Llerasia (tribe Astereae) from synonymy of Haplopappus. Ina pollen review, Wodehouse (1929: figure 8) noted that by spiny pollen Moquinia macrocephala stands apart from taxa of the Moquinia-Gochnatia plexus. Cabrera (1971) treated most Moquinias under Gochnatia, but excluded Gochnatia macrocephala (Rusby) Cabrera from Gochnatid. Lagree with Blake (1927), Wodehouse (1929), and Cabrera (1971) that Buchticn 3080 belongs to Astereae rather than to Mutisieae. Here | provide the combination Llerasia macrocephala (Rusby) Pruski, which replaces the synonymous L. lucidula (S.F. Blake) Cuatrec., the latter recognized by Cuatrecasas (1970). PLUCHEA Britten (1898) noted that Conyza baccharis Miller, partly characterized by au- riculate-clasping leaves with broad serrulate blades and reddish florets, is con- specific with Pluchea bifrons (L.) DC. sensu Candolle 1836). Godfrey (1952), however, recognized the white-flowered North American populations formerly called P. bifrons as P.foetida (L.) DC. Conyza bifrons L., as lectotypified above on European material, indeed is a heterotypic synonym of European Inula bifrons L. Godfrey (1952) also segregated most of the reddish-purple flowered popula- PRUSKI, STUDIES OF NEOTROPICAL COMPOSITAE 2035 tions formerly called P. bifrons as P. rosea R.K. Godfrey, which subsequently has been widely recognized (e.g., Cronquist 1980; Nesom 1989; Arriagada 1998). The lectotype of Conyza baccharis has very densely pubescent outer phyllaries and florets with reddish corollas. The earlier C. baccharis is thus conspecific with P rosea, and the needed new combination for Flora Mesoamericana and Flora North America is proposed herein, this updating the earlier identification of C. baccharis by Britten (1898). Pluchea baccharis (Mill.) Pruski, comb. nov. Conyza baccharis Mill, Gard. Dict. (ed. 8) ees no. 16. 1768. LECTOTYPE (designated here): MEXICO: CAMPECHE: “Grows naturally at mpeachy,” sin. coll. (BM [barcode BM000833507], photograph MO). Material grown in the ee garden by Philip Miller is not extant. The Mexican material was presumably gath- ered by Robert Millar, who also sent Conyza no. 15 from “Campeachy” to Philip Miller. ? Baccharis viscosa Walter, Fl. Carol. 202. 1788, hom. illegit., non Lam. 1785. TYPE: U.S.A. unknown, not seen in BM puieronene of Walter's herbarium. Walter’s description partly re “Varietates, floribus albis, et s rubris.” 1 do not know of this name being lectoty oiled but if it were to be a on the reddish-flowered material it would seemingly belong here in synonymy. Pluchea rosea ae ied Elisha Mitchell Sci. Soc. 68:266. 1952. TyPE: U.S.A. FLORIDA. La lake shores, vicinity of Eustis, 16-31 May 1894, Nash 758 (HOLOTYPE: GH; IsoTYPES: F MO, NY, ke Co. Distribution and ecology.—This is a coastal plain species occurring (see Godfrey 1952; Nesom 1989; Arriagada 1998) in the southeastern United States (North Carolina, South Carolina, Georgia, Florida, Alabama, Mississippi, Louisiana, and Texas), the Bahamas, Cuba, Mexico, and Mesoamerica (Campeche, Quintana Roo, Belize, Honduras, Nicaragua). Pluchea baccharis (Mill) Pruski is very similar and most closely related to P. foetida (L.) DC. (Cronquist 1980; Nesom 1989; Arriagada 1998), from which it differs most notably by pinkish or reddish (vs. generally white) corollas. Pluchea baccharis also tends to have a less dense capitulescence, shorter and more densely pubescent phyllaries, and narrower capitula than does P. foetida. Villasenor (1989) noted that the report by Sousa and Cabrera (1983) of P. foetida as occurring in Quintana Roo was based on a misdetermination. A collection from Veracruz, Mexico is the sole Mexican or Central American collection of P. foetida reported by Nesom (1989), and the material from Veracruz seen by me is referred here to P. baccharis. Pluchea baccharis is also similar to P. longifolia Nash, P. mexicana (RK. Godfrey) G.L. Nesom (described as a var. of P. rosea), and P. yucatanensis G.L. Nesom. Pluchea baccharis differs from P. longifolia by smaller capitula and smaller leaves, from P mexicana by abaxially sessile-glandular (vs. stipitate glandular) leaves, and from P. yucatanensis by abaxially hirsute and glandular (vs. solely sessile-glandular) leaves. Godfrey (1952) listed the illegitimate and non-typified Baccharis viscosaas a synonym of P foetida (L.) DC. 2036 BRIT.ORG/SIDA 21(4) ACKNOWLEDGMENTS l appreciate the reviews of Jorge Arriagada (SCL) and Guy Nesom (BRIT), who kindly read this manuscript for SIDA. lam very grateful that Lucia Kawasaki (F) verified that the abaxial leaf surfaces are indeed strigose on the holotypes of both Clibadium sarmentosum and Clibadium scandens. 1 would also like to thank Anna Balla (F), Charles Jarvis (BM), and Michael Nee (NY) for help in various aspects of this work. Frederick Keusenkothen (MO) is thanked for tak- ing the photographs of the Velloso illustrations. REFERENCES ArisTeGuieta, L. 1964.Compositae. In: T. Lasser, ed. Flora de Venezuela, vol. 10.Instituto Botanico, Caracas.Pp. 1-941. ArriAcana, J.E. 1995. Key to species of Clibadium (Compositae, Heliantheae) with notes on synonymy and morphological variation. Rev. Acad. Colomb. Cienc. 19(74):453-461. Arriacaba, J.E. 1998. The genera of Inuleae (Compositae; Asteraceae) in the southeastern United States. Harvard Pap. Bot. 3(1):1-48 Arriacana, J.E. 2003. Revision of the genus Clibadium (Asteraceae, Heliantheae). Brittonia 55:245-301. Bapitto, V.M. 1994 [1995]. Enumeracion de las Compuestas (Asteraceae) de Venezuela. Revista Fac. Agron. (Maracay) 45:1-191. Barroso, G.M. and S.G. pa ViHNA. 1970. Stifftia axillaris, uma espécie nova de Compositae. Loefgrenia 44:1-2. Buake, S.F.1927.The section Diplostephioides of Aplopappus. Amer. J. Bot. 14:107-115. Britten, J. 1898. The Conyzas of Miller’s dictionary (ed. 8). J. Bot. 36:51-55. Caprera, A.L. 1971.Revision del género Gochnatia (Compositae). Revista Mus. La Plata, n.s., Bot. 12(66):1—160. Canpoite, A.P.be.1836.Prodromus systematic naturalis regni vegetabilis, vol.5.Treuttel and Wurtz, Paris. Cronauist, A. 1980. Asteraceae. Vascular flora of the southeastern United States, vol. 1. The University of North Carolina Press, Chapel Hill. Cuatrecasas, J. 1969. Prima flora Colombiana. 3. Compositae-Astereae. Webbia 24:1-335. CuaTrecasas, J. 1970. Reinstatement of the genus Llerasia (Compositae). Biotropica 2: 39-45. Goorrey, R.K. 1952. Pluchea, section Stylimnus, in North America. J. Elisha Mitchell Sci. Soc. 68:238-271 + four plates. Gray, A., 1884. Caprifoliaceae - Compositae. Synoptical flora of North America. 1(2):1-474. Greuter, W. 2003. The Euro + Med treatment of Astereae (Compositae) - generic concepts and required new names. Willdenowia 33:45—47. Greuter, W., J.McNeit, FR. Barrie, H.M., Burpert, V. Demoutin, T.S. Fircueiras, D. H. Nicouson, PC. SiLva, J.E. Skoc, P. Trenane, NJ. Turtano, and D.L. Hawkswort (eds.). 2000. International code of botanical nomenclature (Saint Louis Code). Adopted By The Sixteenth International Botanical Congress St. Louis, Missouri, July - August 1999. Regnum Veg. 138. PRUSKI, STUDIES OF NEOTROPICAL COMPOSITAE 2037 JORGENSEN, PM. and S. Leon-YAnez (eds.). 1999. Catalogue of the vascular plants of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard. 75:i—viii, 1-1182. LINNAEUS, C. 1737. Hortus Cliffortianus. Amsterdam. Linnaeus, C. 1753. Species plantarum. Stockholm. Linnaeus, C. 1763. Species pantarum. Ed. 2. Stockholm Nesom, G.L. 1989. New species, new sections, and a taxonomic overview of American Pluchea (Compositae: Inuleae). Phytologia 67:158-167. PLukenet, L. 1705. Amaltheum botanicum. London. Pausi, J.F. 1984. Calea brittoniana and Calea kristiniae: Two new Compositae from Brazil. Brittonia 36:98-103. Pruski, J.F. 1997. Asteraceae. In: J.A. Steyermark, PE. Berry, and B.K. Holst, eds. Flora of the Venezuelan Guayana, vol. 3. Missouri Botanical Garden, St. Louis. Pp. 177-393. Pruski, J.F. 1998. Novelties in Calea (Compositae: Heliantheae) from South America. Kew Bull. 53:683-693. Pruski,J.F.and DJ.N. Hino. 1998. Two new species of Calea (Compositae: Heliantheae) from Serra do Grado Mogol and the surrounding area, Minas Gerais, Brazil. Kew Bull. 53: 695-701. Pruski, J.F. and L.E. Ursatscu. 1988. Five new species of Calea (Compositae: Heliantheae) from Planaltine Brazil. Brittonia 40:341-356. Rosinson, H. 1979. New species of Vernonieae (Asteraceae). Ill. Additions to Piptocarpha. Phytologia 44:300-306. RosiNsoNn, H. 1992. Four new species of Clibadium from northern South America (Asteraceae: Heliantheae). Phytologia 73:149-154. Rusey, H.H. 1920. Descriptions of three hundred new species of South American plants: with an index to previously published South American species by the same author. Privately published, New York. SANCHO, G. 1999. Gochnatia calophylla - a synonym of Gochnatia oligocephala (Gardner) Cabrera (Asteraceae, Mutisieae). Willdenowia 29:235-237, Sousa S.,M.and E.F.Casrera C. 1983. Listados floristicos de México. II. Flora de Quintana Roo. Instituto de Biologia, UNAM, Mexico City. Tutin, [.G., V.H. Heywooo, N.A. Buraes, D.H. Vacentine, S.M. Watters, and D.A. Wess (eds.). 1976. Flora Europaea, vol. 4. Cambridge University Press, Cambridge. Vettoso, JM. ba. 1825 [1829]. Florae fluminensis. typographia nationali, Rio de Janeiro. Vewoso, JM. ba. 1827 [1831]. Florae fluminensis (lcones). A. Senefelder, Paris. VILLASENOR, J.L. 1989. Manual para la identification de las Compositae de la Peninsula de Yucatan y Tabasco. Techn. Rep. Rancho Santa Ana Bot. Gard. 4:iii + 1-122. WobeHouse, R.P. 1929. Pollen grains in the identification and classification of plants. IV. The Mutisieae. Amer. J. Bot. 16:297-313 + 2 plates. a 2038 BRIT.ORG/SIDA 21(4) Book REVIEWS C.M. Menzeiand G.K. Waiter (eds.). 2005, Litchi and Longan: Botany, Production, and Uses. (ISBN 0-85199-696-5, hbk.). CABI Publishing, 875 Massachusetts Avenue, 7th floor, Cambridge, MA 01239, U.S.A. (Orders: 617-395-4056, fax 617-354-6875, email cabinao@cabi.org, wwwcabi-publishing org/bookshop). $140.00 hbk., 336 pp., 72 color, 7" x 9 3/4' Litchi and longan fruit are extremely important economic plants throughout Asia and are popular in Asian cuisine. Pacific Rim countries are the biggest producers of litchi and longan fruit, but the plants are cultivated and used all over the world. This book is edited by two experts from the Queensland Department of Primary Industries and Fisheries and covers information on ae many cultivars of litchi and longan fruit, including Litchi yur (Sapindaceae). Written by experts in the field, this review ultis 1, horticulture, and production. chinensis Sonn.and Di ig book s the research a science of nie and longan e editors include chapters on litchi and longan taxonomy, ae | ders, pests, irrigation, flowering, harvesting, diseases, and fruit maturation. This isan excellent resource r researchers who study and litchi and longan. Beautiful color propagation, fruit disor- for growers, horticulturists and othe and black-and-white photographs are included.—Marissa Oppel, Herbarium Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 76012-4060, U.S.A, BARBARA Boswortu. 2005. Trees: National Champions. (ISBN 0-262—2592-2, hbk.). The MIT Press, 55 Hayward Street, Cambridge, MA 02142-1315, U.S.A. (Orders: 617-253-5643 800-405-1619, mitpress-orders@mit.edu, http:// mitpress.mit.edu). $39.95, 169 pp., b/w photos throughout, 12" « 91/2" A collection of panoramic views of remarkable trees seen through the eyes of an artist. “Currently, there are 889 national champions and cochampions among the 826 eligible species in the United States.” The book pictures 70 of these, including natives and non-natives, as black & white — taken by Barbara Bosworth from 1991 to 2004, using an 8 x 10 camera. Tree champions are judged on an index of circumference, height, and crown spread, and some of the a aphs aoe obvi ious eis the biggest of its species, but its isolation behemoths. For others, the viewer is aware that the tre and stark contrast with surroundings underlie the sense of beauty and awe conveyed byt the photo- suy Nesom, Botanical Research Institute of graph. So many of these individuals are lone survivors. Texas, Fort Worth, TX, 76012-4060, U.S.A. SIDA 21(4): 2038. 2005 UNA NUEVA ESPECIE DE GUAREA (MELIACEAE) PARA COSTA RICA Alexander Rodriquez Instituto Nacional de Biodiversidad (INBio), Apdo. 22-3100, Santo Domingo, Heredia, COSTA RICA arodrig@inbio.ac.cr RESUMEN Se describe e ilustra Guarea subsessilifolia, una nueva espe: endémica del cerro Turrubares en el pacifico central de Costa Rica, ademas, se comenta su relaci6n con otras especies del género ABSTRACT Guarea subsessilifolia, a new and endemic species from cerro Turrubares in the central pacific of Costa Rica, is described and illustrated and relationships with related taxa are discussed Guared es uno de los géneros mas grandes de la familia Meliaceae en América tropical y fue establecido por Linneo en 1753 bajo el nombre Guara, aunque en 1771 el mismo Linneo corrigié su escritura a Guarea (Coronado 2003 Este género ha sido estudiado taxonémicamente en varias ocasiones. Entre los primeros aportes importantes encontramos el presentado por Casimir de Candolle (1878) quien en una monografia completa para la familia Meliaceae reconocié 2 secciones, EuGuarea (actualmente Guarea) con 67 especies y la seccion Ruagea (actualmente reconocido como un género distinto) con tres especies (Coronado 2003). Casi LOO anos tr rieron para que la familia fuera nuevamente revizada y fueron Pennington y B.T. Styles (1981) quienes elaboraron una nueva monografia de la familia para el neotropico. En esta revision se reconocen 35 especies de Guarea para América tropical, se indica la presencia de otras 5 especies en Africa tropical y solamente 10 especies fueron reconocidas del trabajo de Casimir de Candolle, ademas, se reporto para Costa Rica la presencia de 7 especies. También, algunas floras locales han sido aportes regionales importantes en la revision de este género, tal es el caso de Flora de Guatemala (Standley & Steyermark 1946), Flora de Panama (Smith 1965) y Flora de Nicaragua (Pennington & Styles 200). Recientemente, Coronado (2003) en suestudio relacionado con el complejo de Guarea glabra Vahl entre México y Panama, indicé la presencia de 10 especies para Costa Rica, 3 de las cuales actualmente inéditas, G. arcuata Coronado, G. gentryi Coronado y G. zarceronensis Coronado y distintas a la especie aqui descrita El género Guarea se caracteriza por estar constituido de arboles 0 arbustos, con hojas alternas, compuestas, generalmente pinnadas y con una yema terminal SIDA 21(4): 2039-2044, 2005 2040 BRIT.ORG/SIDA 21(4) de crecimiento intermitente, por su condicidn dioica, flores unisexuales por aborto, aunque pareciendo perfectas, caliz cupuliforme, pétalos libres, en la mayoria de los casos val vados, estambres con filamentos completamente unidos y formando un tubo cilindrico, con disco nectarifero intraestaminal, ovario 2- 13 locular, loculos con 1-2 6vulos, frutos capsulares, 2-10(-14) valvados y finalmente por mostrar loculos con 1-2 semillas, las cuales se encuentran rodeadas por una sarcotesta carnosa, roja o anaranjada. -as recientes exploraciones en la vertiente pacifica de la cordillera de Tilaran, en Monteverde y en el cerro Turrubares localizado en el pacifico cen- tral de Costa Rica ha producido el hallazgo de ésta notable especie de Guarea cuyos caracteres difieren de los taxa anteriormente reportados para el género (Pennington et al. 1981; peste 2003). A continuacién se presenta la descripcion de esta especie insolita. — — Guarea subsessilifolia Al. Rodr, sp. nov. (Figs. 1, 2). Tirpo: COSTA RICA. SAN Jost: Turrubares, San Luis, Faldas del cerro Pelon, 09°48'55'N, 84°28'48"W, 1190 m, 19 May 2005 (11s), Rodriguez se OTIPO: INB; ISOTIPOS: BRIT, CAS, CR, DUKE, GH, MEX, MEXU, MO, NY, TEX, UC, US, USJ, W). ta di Ab omnibus congeneribus folia subsessilia vel brevipetiolata, foliola proxima re Arbusto 2.5-4 m de altura, dioico. Tronco con corteza interna rosada; tallitos glabrescentes a cortamente hirsutos en ramitas terminales, con o sin esparcidas lenticelas. Hojas pinnadas, con una yema terminal de crecimiento intermitente; subsésiles 0 peciolos hasta 0.7 cm de largo, subteretes a levemente aplanados en el lado adaxial, ferrugineo-puberulentosa hirsutulos, glabrescentes con la edac raquis |-17 cm de largo, subterete a canaliculado en el lado adaxial, ferrugineo- puberulentoa hirsutulo, glabrescente con la edad; foliolos 1-10 pares, opuestos, subsésiles o peciolulos hasta 2 mm de largo y 1.25-1.75 mm de ancho, teretes: lamina 1.5-13 cm de largo y 0.8-4 cm de ancho, oblongo-elipticos, elipticos a ovados, foliolos proximales marcadamente reducidos, 3-8.5 mas cortos que los distales, base cuneada a obtusa, en ocasiones levemente oblicua, apice acuminado oagudo, 5-14 pares de venas secundarias, concolora y verde-grisacea a gris-rojiza al secar, haz y envés puberulentos a esparcidamente pubescentes sobre el nervio principal, glabrescentes con la edad, eglandulosos. Inflorescencias terminales 0 axilares en ramitas terminales, solitarias, tirsos 0 cimas, erectas; masculinas con 8-35 flores, 2-6 cm de largo y 1.5-3. cm de ancho, pedunculo 0.1-2 cm de largo, ferrugineo-puberulento a hirsutulo, glabrescente con la edad, raquis 0.5-4 cm de largo, ferrugineo-puberulento a hirsutulo glabrescente con la edad; femeninas con 4-13 flores, 1-3 cm de largo y 1-2 cm de ancho, pedunculo 0.2-0.5 cm de largo, ferrugineo-puberulento a hirsutulo, glabrescente con la edad, raquis 0.3-0.7 cm de largo, ferrugineo-puberulento a hirsutulo, glabrescente con la edad. Flores estaminadas con pedicelos 2-4 mm de largo, articulados, esparcidamente ferrugineo-puberulentos, 1-2 bracteolados, iu RODRIGUEZ, TA RICA 2041 Fic. 1. Guarea subsessilifolia Al. Rodr. (Rodriquez 9613). A. Habito. B. Base de la hoja. C. Inflorescencia masculina. D. Inflorescencia femenina. E. Flor femenina. F. Corte longitudinal de flor femenia. G. Corte transversal del ovario. H. | i Jinal de fl li L. Antera Anterodio. I. Fruto. J. Flor masculina. K. C bractéolas 0.5-1 mm de largo; caliz 1-1.5 mm de altura, ciatiforme, verde a pardo hacia la parte distal, esparcida y diminutamente pubescente, lobulos 4, 0.2- 0.35 mm de largo; corola con 4 pétalos, 5-5.5 mm de largo y 1.25-1.5 mm de ancho, blanco-rosados, aunque rosados en botones florales, pice agudo, lado externo esparcida y diminutamente pubescente, interno papilado hacia el apice; tubo estaminal 4-5 mm de largo y 1.5-2 mm de ancho, apice irregular te dentado, lado externo papilado; anteras 8, 0.75-1 mm de largo y 0.3-0.45 mm de ancho, oblongas a oblongo-obovadas; gin6foro 1-1.25 mm de largo y 0.75-0.9 mm de ancho, glabro, disco nectarifero 0.4-0.5 mm de largo y LI-L3mm de ancho, ovario 0.7-0.85 mm de largo y 1-1.2 mm de ancho, con esparcidos y largos tricomas, pistilo sobresaliendo levemente sobre el tubo estaminal, estilo 2.25-2.5 mm de 2042 BRIT.ORG/SIDA 21(4) largo, glabro, estigma 0.3-0.45 mm de largo y 0.6-0.75 mm de ancho. Flores pistiladas con pedicelos 1.5-4 mm de largo, articulados, esparcidamente ferrugineo-puberulentos, 1-2 bracteolados, bractéolas 0.5-1 mm de largo; caliz 0.75-1 mm de altura, ciatiforme, verde a pardo hacia la parte distal, esparcida y diminutamente pubescente, lobos 4, 0.2-0.4 mm de largo; pétalos 4, 5-5.5 mm de largo y L.7-2 mm de ancho, blanco-rosados, aunque rosados en botones florales, apice agudo, lado ee esparcida y diminutamente dgieaaaue interno papilado hacia el apices staminal 4-5 mm de largo y 2-2.5 mm de ancho, apice irregularmente dented: lado externo papilado; anterodios 8, 0.75- Imm de largo y 0.3-0.45 mm de ancho, oblongos a oblongo-obovados; gindéforo 0.5-0.6 mm de largo y 0.75-0.9 mm de ancho, glabro, disco nectarifero 0.3-0.5 mm de largo y 1.3-1.5 mm de ancho, inconspicuo; ovario 4 locular, 0.7-0.85 mm de largo y 1.3-1.5 mm de ancho, sericeo, un dvulo por léculo; pistilo similar en largo al tubo estaminal, estilo 1.6-L8 mm de largo, glabro, estigma 0.25-0.3 mm de largo y 0.6-0.7 mm de ancho. Frutos 1-1.5 cm de largo y 1-2 cm de ancho, globosos, lisos, base obtusa, apice obtuso a subtruncado, rojizos al madurar, glabrescentes a esparcidamente pubescentes, (3-)4 loculado, pericarpo 0.5-] mm de grosor. Semillas | por loculo, 0.6-0.8 cm de diametro, subrodeadas por una sarcotesta rojo-anaranjada. Fenologia.—Flores en Mayo y frutos en Abril y Noviembre. Distribucion.—Endémica de Costa Rica, conocida en la vertiente pacifica de la cordillera de Tilaran, en Monteverde y en el pacifico central sobre el cerro Turrubares, en bosques htmedos a elevaciones entre 850-1600 m. Guarea subsessilifolia se reconoce y distingue de otras especies por su habito arbustivo y por sus hojas subsésiles o con peciolos muy cortos y por sus foliolos proximales conspicuamente reducidos (en ocasiones caedizos); hojas hasta con LO pares c¢ pistiladas con el ovario sericeo, 4 loculado y con un 6vulo por léculo, finalmente por sus pequenos y globosos frutos, (3-)4 loculares, con una semilla por léculo, con valvas lenosas y delgadas. pam e foliolos, inflorescencias cortas y escasamente floreadas, flores A pesar de que las diferencias florales entre los individuos estaminados y pistilados no es evidentes, se encontraron diferencias significativas entre los individuos de ambos sexos. Los individuos masculi inflorescencias conspicuamente mas largas y con mayor numero de flores, el caliz, el gindloro y el estilo son ligeramente mas largos, ademas, el ovario y el disco nectarifero son mas angosto y menos pubescente que los individuos femeninos. Esta especie se podria relacionar con G. glabra Vahl, G. guidonia(L.) Sleumer, G. jamadicensis Proctor, G. pubescens (Rich.) A. Juss. y G. macrophylla Vahl al considerar conceptos taxondomicos usados por Pennington (1981) que incluyen ovario (3-)4 loculado, pubescente, con un ovulo por loculo, corola pequena, nunca alcanzando 12 mm de largo, frutos globosos, pequefios, no sobrepasando 2.5. cm de diametro, lisos, con ausencia de costillas o lenticelas evidentes, RODRIGUEZ 2043 pericarpio delgado, menos de 3 mm de grosor e inflorescencias cortas. Sin em- bargo, estas especies se distinguen facilmente por mostrar peciolos evidentes y claramente diferenciados. También, con base en el trabajo de Coronado (2003) y utilizando los mismos conceptos taxondmicos adoptados por Pennington (1981), dentro del complejo de G. glabra Vahl la especie podria relacionarse con G. bullata Radlk. G.chiricana Standl. y G. petensis Coronado, sin embargo, estas especies se distinguen igualmente por mostrar peciolos evidentes y claramente diferenciados. Esta especie es notablemente rara, se conoce tnicamente de 2 localidades, el cerro Turrubares y Monteverde, regiones en las cuales las poblaciones observadas son muy reducidas. Es probable que esta especie se encuentre en alto riesgo de extincion, mas considerando que la regiones especificas de recolecta nose encuentran en unaestricta categoria de proteccion y corresponde a sitios con alto uso del suelo en actividades como ganaderia y agricultura. En los bosques de esta region, se observo que G. subsessilifolia crece simpatricamente con G. kegelii Turcz., ademas, otras especies comunes en el sitio son Croton megistocarpus Gonz. Ram. & Poveda (Euphorbiaceae), Chrysophyllum brenesii Cronquist (Sapotaceae), Daphnopsis folsomii Barringer & Nevling (Thymelaeaceae), Hirtella triandra Sw. (Chrysobalanaceae), Krugiodendron acuminatum Gonz. Ram. & Poveda (Rhamnaceae), Piper subfuscum C. DC. (Piperaceae), Hoffmannia psychotriifolia (Benth.) Griseb. (Rubiaceae), Lippia cardiostegia Benth. (Verbenaceae), Picramnia antidesma Sw. (Gimaronpnee. Rondeletia calycosa Donn. Sm. (Rubiaceae), Siparuna gesnerioides (Kunth) A. DC. (Monimiaceae), Irophis mexicana (Liebm.) Bureau (Moraceae), entre otras. Etimologia.—El epiteto latino subsessilifolia hace alusion a la presencia de hojas casi sésiles. Parativos. COSTA RICA. Puntarenas: 2.5 km ocste de Monteverde sobre la carretera a L eae Los Llanos, 10°18'N, 84°50'W, 1200 m, 6 Feb 1997 (rs), Haber 12028 (CAS, CR, GH, INB, MO, NY, US); 3.5 km oeste de Monteverde sobre la carretera a Lagarto, Los Llanos, 10°18'N, 84° 50'W, ae m, 6 Feb 1997 (frs), Haber 12109(BM, CAS, CR, DUKE, GH, INB, MEXU, MO, NY, TEX, US). San José: Turrubares, Faldas del cerro Bares, Zona Protectora Cerros de Turrubares, 09°47'30"N, 84°28'30"W, 1600 m, 6 Nov 1990 (frs), Zuniga 376 (CR, INB, MO); San Luis, Faldas del cerro Pelon, 09°48'55°N, 84°28'48"W, 1190 m, 7 Oct 2004 Us), Rodriguez 9344 (CAS, CR, GH, INB, MO, NY, US), Rodriguez 9614 (BRIT, CAS, CR, UKE, GH, INB, MEXU, MO, NY, TEX, US, W); Zona Protectora Turrubares, Cuenca del rio Grande de Tarcoles, 09°47'N, 84°29'W, 850-1100 m, 6 Abr 1993 (frs), Grayum 10551 (NB). AGRADECIMIENTOS El autor desea agradecer cordialmente a Maria T. German Ramirez y a un revi- sor anonimo por la revision y los comentarios al articulo, a Silvia Troyo por su magnifica ilustracion y finalmente a John Pruski por su ayuda en las diagnosis del latin. 2044 BRIT.ORG/SIDA 21(4) REFERENCIAS Coronabo, |.M. 2003. Systematic revision and multivariate analysis of the Guarea glabra Vahl (Meliaceae) complex from Mexico to Panama. Thesis for master of science de- gree. University of Saint Louis, Missouri. De Canpotte, C. 1878. Meliacées. Monogr. Phan. 1:399-758. PENNINGTON, T.D., B.T. Stytes, y D.A.H. Taytor. 1981.A monograph of Neotropical Meliaceae. Fl. Neotrop. Monogr. 28:1-470. PENNINGTON, T.D.y B.T. Sty.es. 2001.Meliaceae. En: Stevens, W.D,, C. Ulloa, A. Pool, y O. Montiel, eds. Flora de Nicaragua, Tomo Il. Monogr. Syst. Bot. Missouri Bot. Gard. 85:1419-1430. Smity, C.E. 1965. Meliaceae. Flora of Panama. Ann. Missouri Bot. Gard. 52:55-79, STANDLEY, P.C. y J.A. STevermMaRK. 1946. Meliaceae. Flora of Guatemala. Fieldiana, Bot. 24(5): 444-468 DIOSPYROS TORRESII (EBENACEAE): A NEW BLACK ZAPOTE FROM TROPICAL MEXICO Mitchell C. Provance Andrew C. Sanders Depar tment of Bota 'y al 1d Plant Sciences Herbarium (UCR) College of Nat | and Agricultural Sciences University of California, Riverside University BiGalRernia Riverside Riverside, California 92521-0102, U.S.A. a California 92571-0102, U.S.A andrew.sanders@ucr.edu mitchtel eine. ABSTRACT new Species of black eapers from eae Mexico is described, and photos of the holotype are provided. D =. Provance & A.C. Sanders is based on collections from two localities rE in north and central Oaxaca. This species appears to belong to a group of closely related American Diospyros with fruits that are dark-colored and sweet at maturity. Key Worps: black zapote, Diospyros i, Ebenaceae, Mexican persimmons, systematics, taxonomy, Tehuacan-Cuicatlan allege ee negro RESUMEN Se rdesene una nueva ioc del México pea y se pienen fotos ce apie S oe ion de -. Provance & A.C. Sanders se | S | norte y centro de Dae Se cree que esta especie pertenece a un grupo cercanamente relacionado de Diospyros de América que tienen frutos de color oscuro y que son dulces cuando maduran. A monograph of the Mexican Diospyros (Ebenaceae) is the current focus of stud- ies by the first author of this paper. During the course of reviewing thousands of collections of American Diospyros, it has become clear that there are several undescribed taxa in Mexico. This paper describes one such species based on collections from Oaxaca, Mexico, that do not belong to any currently recog- nized species. A comprehensive key to the Mexican species of Diospyros has not been published since that by PC. Standley (1924). Such a key and classifica- tion will soon be published by the first author in connection with his current work on the genus in Mexico. The new species described here is best considered a member of what we informally recognize as the Diospyros rosei Complex. The flowering sepals of this complex tend to have vermiform glandular hairs at their apices. The sepals of female flowers tend to be strongly accrescent and are 5-8 in number. The species of this group produce medium size fleshy fruits that are dark-colored and sweet at maturity, and may contain from 10 to lo ovules. In addition to the species described here, this group includes: Diospyros rosei Standl., Diospyros palmeri Eastw., Diospyros californica 1.M. Johnst., Diospyros oaxacana Stand. Diospyros sonorae Stand. Diospyros texana Scheele, Diospyros riojae Gomez-Pompa, Diospyros conzattii Standl., Diospyros rekoi SIDA 21(4): 2045-2050. 2005 2046 BRIT.ORG/SIDA 21(4) StandL, and Diospyros xolocotzii Madrigal & Rzed. Some South American spe- cies may belong to this group as well. However, sufficient material from that continent was not available for investigation. We have evaluated the holotype of D. morenoi A. Pool, a species recently described from Nicaragua (Pool 1997). In our opinion, this material is consistent with D. rekoi, a species for which we have seen material from Mexico and EFI Salvador. SPECIES DESCRIPTION Diospyros torresii M.C. Provance & A.C. Sanders, sp. nov. (Fig. 1). Type. MEXICO. OAXACA: Mpio. Santiago Texcalcingo: 11 km al E de Teotitlan del Camino carr. a Huautla de Jiménez, lapprox. 18° 11'N, 97° 02'W], 1710 m, 17 Mar 1985, R. Torres C.& M.A. Martinez 6636 (HOLOTYPE: MO; ISOTYPE: CHAPA) Frutices vel arbores 4 m alta; laminae lanceolatae vel ovatae, petioli decurrentes; petala ovalia vel as meas ac eaaag introrsa secus ee oe eae um, Calbyces 5-7 partiti sepala atropurpurei ubi re ati. 8 | Shrubs or trees 4 m tall, probably facultatively deciduous; old stems often pul- verulent, rarely smooth, dark reddish-brown to dark gray; current years stems with stiff, erect, reddish hairs densely covering surface, occasionally retrorse, some slightly wavy, densely appressed at the shoot apex; lenticels common, fill- ing tissue yellowish to off-white; bud scales convex, ovate, densely appressed reddish hairy; petioles 2-4 mm long, both sides densely erect white hairy, some- times clavate glandular hairy, convex below, + flat and minutely V-grooved above, distal half minutely winged along the margin, sometimes winged the entire length; mature leaves entire, alternate, chartaceous, lanceolate to ovate, 3-6 cm long including the petiole, 2-3 cm wide, blade abruptly decurrent on the petiole, mostly grayish in herbarium material, base rounded, obtuse or acute, margin revolute, apex obtusely rounded, sometimes acutely rounded, below sparsely to moderately erect white hairy, hairs sometimes slightly wavy, some- times clavate glandular hairy, minutely papillate, usually minutely black gland- dotted, above irregularly wavy to + rugose, glabrate to minutely erect white hairy, sparsely and minutely papillate, sometimes sparsely clavate glandular hairy; laminar extrafloral nectaries abaxial, minute, roundish, mostly raised, probably green in living material; venation + brochidodromous; midrib promi- nent below, sub-terete, densely to very densely erect white to reddish hairy, above flush or barely raised, rare —_— y slightly impressed, yellowish, sparsely to densely erect hairy, less hairy distally, often becoming glabrate, often clavate glandular hairy near the petiole; 2° venation 5-7 major lateral veins on each side of mid- rib, below raised, above flush with surface or barely raised, very obscure; 3° venation reticulated below raised or not, visible or not, above impressed, of ten obscure, sometimes not visible; new leaves strigillose below, especially along the midrib, clavate glandular hairy, above minutely hairy, the hairs mostly erect, sometimes appressed, clavate glandular hairy; female flowers solitary on young PROVANCE AND SANDERS, A NEW SPECIES OF DIOSPYROS FROM TROPICAL MEXICO 2047 C. Provance & A.C. Sanders. Holotype, &. Torres C. & M. A. Marti 36 (MO) with detail of leaf fae and female flower (indicated by the arrow). 2048 BRIT.ORG/SIDA 21(4) stems; flowering pedicels 2 mm long; pedicel bracts 2, sub-opposite, 4-7 mm long, 0.75 mm wide, linear, densely minute erect hairy, densely clavate glandu- lar hairy; fruiting pedicels 5-10 mm long, + L5 mm wide, somewhat stout, densely but minutely erect hairy, apex hat-shaped; calyx 5-7-merous, sinuses rounded, acute; calyx tube cupulate, exterior sparsely to moderately minutely appressed hairy; flowering sepals 5-6 mm long, + 2 mm wide, sparsely hairy, the hairs containing a reddish-brown compound, exterior moderately to sparsely appressed hairy, sparsely and minutely clavate glandular hairy, apically dark glandular vermilorm hairy, interior densely but minutely clavate glandu- lar hairy, with low to moderate numbers of simple hairs, especially towards base; fruiting sepals accrescent in fruit, 1o-17 mm long, 5-6 mm wide, thick, + spreading, lance-ovate to elliptic, apices acutely-rounded, venation obscure, minutely hairy on both sides, sparser towards the apex, minutely black gland- dotted on both sides; female corolla 3 mm long; corolla tube 1.25 mim long, 2.0- 2.5mm wide, exterior densely appressed hairy, interior glabrous; lobes 5, + 1.75 mm long, oval to quadrate, exterior densely appressed hairy, inside glabrous, involute, more so on the right margin than the left; ovary globose, smooth, sparsely minute hairy; styles 5, one third of length of ovary, connate, becoming distinct distally; stigmas 5, short, labiate; staminodes none seen; male flowers unknown; fruit a berry, ellipsoid, + + cm long, 3 cm wide; mesocarp fibrous, | orange in herbarium material; exocarp + 2 mm thick, hypodermis moderately thick, sclerified, epidermis minutely blistered, glabrous, atropurpureous in dried material. Distribution and Ecology.—To our knowledge, Diospyros torresii has so far Oaxaca, Mexico, however, the type was col- only been collected in the state o lected within ca. 2 km of the Puebla state line, and we believe that it probably occurs in that state also. The type collection was made in oak woodlands of the Sierra Mazateca west of Huautla. The condition of the material from the Sierra Mazateca leads us to believe that D. torresii is facultatively deciduous. Both sheets of Torres 6636 demonstrate ripe fruit, new female flowers and relatively few mature leaves, but there are a number of young shoots with very young leaves. The paratype was collected from the Tlacolula Valley, but unfortunately is without specilic ecological data. Etymology.—This species is named in honor of Rafael Torres Colin, an au- thority on Bauhinia (Fabaceae) and Oaxacan floristics, and also the collector of the type. — PARATYPE: MEXICO. Oaxaca: Mpio. Villa Diaz Ordaz: Barranca of Diaz Ordaz, NW of Mitla, 1700 m, I] Feb 1966, M.& A. Kirkby 2739(NA) DISCUSSION Diospyros torresii is currently known [rom two localities in Oaxaca. The mate- rial used to describe this new species was previously identified as D. palmeri PROVANCE AND SANDERS, A NEW SPECIES OF DIOSPYROS FROM TROPICAL MEXICO 2049 Eastwood (R. Torres C. & R Cedillo T. 6636) or D. aequoris Standley (M. & A. Kirkby 2739). Morphologically it most resembles Diospyros oaxacana Standley. Diospyros riojae Gomez-Pompa and Diospyros conzattii Standley. It can easily be distinguished from these species provided fertile material is available. In fact, even though identification of sterile Diospyros material is sometimes difficult, in the case of D. torresii sterile material should not pose a major problem. The paratype is sterile but we are confident that this material represents the new species. Vegetative features that distinguish D. torresii from D.conzattii include its smaller, usually thicker leaves and shorter petioles. In addition, the lamina of D. torresii is abruptly decurrent on the petiole, and decurrent for a shorter distance than in D. conzattii, whereas the leaves of D. conzattii are typically tapered basally and often decurrent on the petiole for nearly the entire length. The leaves and petioles of D. oaxacana are velutinous to densely pilose, often on both sides, while in D. torresii they are glabrous or have only sparse, minute, erect hairs. Furthermore, the leaves of D. oaxacana tend to be oval to obovate versus the lanceolate to ovate leaves of D. torresii. The leaves of D. riojae tend to be larger than those of the new species, but more importantly, the 2°and 3°veins as viewed from the adaxial surface of the leaves are raised and contrast sharply with the lamina. In D. torresii the 2° and 3° veins are scarcely visible adaxially. An interesting detail concerning the type locality is that itis only about 20 km from Coxcatlan, Puebla, and other archaeological sites that have yielded evidence of early agriculture in Mexico (e.g. Smith 1965; Eubanks 2001). Most close relatives of D. torresii are known to produce fruits that are edible or even highly desirable. Diospyros oaxacana has been recorded asa useful plant in the Tehuacan-Cuicatlan Valley (Casas et al. 2001), and D. conzattii has been re- corded as a useful plant in Chinantec-speaking communities of the Oaxacan highlands in the District of Cuicatlan (Lipp 1971). Given the large size of the ripe fruits, their probable edibility, and the nearness of this species to some of the oldest agricultural sites in the New World, it may be worthwhile to reevalu- ate the identification of putative Diospyros digyna seeds associated with nearby archaeological sites (e.g. Smith 1965; Callen 1965). Similarly, herbarium vouch- ers, if they exist, associated with ethnobotanical reports of Diospyros usage by nearby indigenous populations (Lipp 1971) should also be reviewed. Few specimens of this new taxon are known to us, and although this makes it tempting to recommend formal protection for D. torresii, we first recommend focused collecting in Oaxaca before such action is taken. ACKNOWLEDGMENTS The authors of this paper would like to express their gratitude to Brett Provance for preparation of the Latin diagnosis, Jeffrey Ross-Ibarra for translating the abstract into Spanish, Victor Steinmann for reviewing a very preliminary manu- script and subsequently making many useful suggestions, and also the anony- 2050 BRIT.ORG/SIDA 21(4) mous reviewers for there recommendations. We would like to thank all of the herbaria that have loaned material for the revision of Mexican Diospyros, but especially CHAPA, MO and NA for the loan of specimens that brought this new species to our attention. We would like to thank E.O. Plummer for all of his technical assistance. REFERENCES Caiten, E.O. 1965. Food habits of some Pre-Columbian Mexican Indians. Econ. Bot.19: 335-343. Casas, A., A. VALIENTE-BANuerT, J.L. Viveros, J. CABALLERO, L. Cortes, P. DAvita, R. Lira, and |. Ropricuez. 2001. Plant resources of the Tehuacan-Cuicatlan Valley. Econ. Bot. 55:129-166. Eusanks, M.W. 2001.The mysterious origin of maize. Econ. Bot. 55:492-514. Lire, FJ. 1971. Ethnobotany of the Chinantec Indians, Oaxaca, Mexico. Econ. Bot. 25: 234-244, Poot, A. 1997. Diospyros morenoi (Ebenaceae), a new species from Nicaragua. Novon 7: 189-190 Smith, C.E., Jk, 1965. The archeological record of cultivated crops of New World origins. Econ. Bot. 19:323-334. STANDLEY, P.C. 1924. Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23(4):1124—1129. STAUROCHILUS LEY TENSIS A PHILIPPINE SEGREGATE OF STAUROCHILUS FASCIATUS (ORCHIDACEAE: AERIDINAE) Eric A. Christenson 4503 21st Avenue West Bradenton, Florida 34209, U.S.A. orchideric@juno.com ABSTRACT The new combination Staurochilus leytensis (Ames) E.A. Christenson (Orchidaceae) is made. RESUMEN Se hace la nueva combinacion Staurochilus leytensis (Ames) E.A. Christenson (Orchidaceae). Oakes Ames and his associates at Harvard University accepted extremely broadly defined species during the first half of the 20th century and in the pro- cess reduced many taxa to synonymy. Vandopsis leytensis Ames was treated as a synonym of Staurochilus fasciatus by Ames and Quisumbing (1932). This species is here recognized as distinct and a new combination is published in Staurochilus. Staurochilus fasciatus (Rchb/f.) Ridl, J. Linn. Soc., Bot. 32:350. 1896. Trichoglottis fasciata Rchb£., Flora 55(9):137. 1872; Stauropsis fasciata (Rchb-f.) Benth., Index Kewensis 982. 1885. TyPE: “Hinterindien,” collector unknown (HOLOTYPE: W). Distribution.—Thailand, Laos, Kampuchea. Vietnam, Peninsular Malaysia, Sumatra, Borneo (following Seidenfaden 1988) Staurochilus leytensis (Ames) E.A. Christenson, comb. nov. Basiony: eee leytensis Ames, Orchid. 5:222. 1915. Type: THE PHILIPPINES: Leyte, Dagami, 60 m, I¢ 1912, CA. Wenzel 14 (LECTOTYPE: AMES; ISOLECTOTYPE: NY!, designated by Seidenfaden 1988) Ames and Quisumbing (1932) illustrated S. leytensis (as Stauropsis fasciata (Rchbf.) Benth.) with photographs, black and white drawings, and colored drawings without noting its distinctive features. Distribution.—Staurochilus leytensis has been collected on Agusan, Leyte, Quezon, Rizal and Sorsogon at elevations of 60-800 m (Valmayor 1984, as Trichoglottis fasciata Rchb£.). While these records need to be reexamined, there is no reason to believe that any of them represent a far disjunct population of true Staurochilus fasciatus which is native to Southeast Asia, adjacent Indone- sia, and reportedly Borneo. While S. fasciatus and S. leytensis are clearly sister species, they are amply me 'An image of the isolectotype is available online at nybg.org SIDA 21(4): 2051-2052. 2005 2052 BRIT.ORG/SIDA 21(4) distinct and geographically quite isolated from each other. In S. fasciatus the leaves are V-shaped in cross section and ligulate, the lateral sepals are falcate- incurved (*bowlegged”), the petals are flat, the large lateral lip lobes lie in the same plane as the midlobe, and the sepals and petals are densely marked. In contrast, S. leytensis has leaves that are [lat and proportionately broader, diver- gent lateral sepals, incurved petals, shallowly suberect-incurved lateral lip lobes, and different floral markings. The species can be distinguished in following key: Leaves ligulate, V-shaped in cross-section, up to 12 x 2.5 cm; lateral sepals incurved- falcate such that the three sepals form a tall isosceles triangle; petals flat and held rigidly at 180°, large lateral lip lobes lie in one plane together with the midlobe Staurochilus fasciatus Leaves oblong-elliptic, flat with only a depressed midvein, up to 10 x 3 cm; latera sepals strongly divergent such that the three sepals form an equilateral triangle; petals incurved yielding a shallowly cupped flower, petals of S. /eytensis appear to be more narrowly clawed than those of S. fasciatus but more specimens are needed to quantify this difference, lateral lip lobes shallowly erect-incurved and do not lie in the same plane as the midlobe Staurochilus leytensis While both S. fasciatus and S. leytensis have pale yellow sepals and petals with transverse brown bars, the markings are different. In S. fasciatus the bars are thick and often coalesce toward the segment apices forming nearly solid brown patches. The bars of S. le ytensis are narrower, cover significantly less of the sur- face, and do not appear to coalesce into solid patches. Onan historical note, when Ridley (1896) first described Staurochilus with S. fasciatus as the sole species he stated that “It is commonly stated in horticul- tural books that this isa native of the Philippines. | have not seen any thence.” It appears that he was on the right track after all and that the species does not occur in the Philippines. — REFERENCES Ames, O. and E. QuisumBinc 1932. New or noteworthy Philippine orchids, Il. Philippine J. Sci. 47:197—220,+ 29 plates. [Stauropsis fasciata p. 214] Riotey, H.N. 1896. The Orchidaceae and Apostasiaceae of the Malay Peninsula. J.Linn. Soc., Bot. 32:213-416. [Staurochilus p.351] SEIDENFADEN, G. 1988. Orchid genera in Thailand: 14. Fifty-nine vandoid genera. Opera Bot. 95:1-398. [Staurochilus pp. 88-96] Seibenraden, G.and J.Woop 1992. The Orchids of Peninsular Malaysia and Singapore. Olsen & Olsen, Fredensborg. Vatmayor, H.L. 1984. Orchidiana Philippiniana. Eugenio Lopez Foundation, Metro Manila. ESTUDIOS EN LAS APOCYNACEAE NEOTROPICALES X VIL: UNA REVISION DEL GENERO GALACTOPHORA (APOCYNACEAE: APOCYNOIDEAE) J. Francisco Morales Instituto Nacional de Biodiversidad (INBio) Santo Domingo de Heredia, COSTA RICA RESUMEN Las especies cee genero nina WGodeee ae aot ADO s son ae en una monogratia lo una nueva especie, G. tw) dos nuevos sinonimos. Se bindan claves ee iones, ilustraciones y especimenes examinados. ABSTRACT +4 y The species of the genus Galactophora Woodson (Apocynaceae: A lina mono- graph. Six species are accepted, including a new sets G. angustifolia and two new synonyms are Sil proposed. Keys, description s and specimens examined are provided. Galactophora (Apocynaceae, Apocynoideae) es un pequenio género de 6 especies, originalmente descrito por Woodson (1932) y distribuido en el SE de Colombia, Venezuela, Pert, Bolivia y Brasil, que se puede reconocer facilmente por su habito usualmente erecto o suberecto (raramente escandente), hojas con la lamina coridcea o subcoriacea, usualmente revolutas marginalmente, sin coléteres en el nervio central adaxialmente, inflorescencias cortamente racemosas 0 cimas reducidas, terminales o subterminales, usualmente con pocas flores, sépalos sin coléteres en la base de la cara adaxial, anteras aglutinadas a la cabeza estigmatica, cabeza estigmatica con cinco crestas 0 proyecciones longitudinales, restringidas a la base, y la ocasional presencia de pelos glandulares en tallos, inflorescencias y sépalos. La presencia de otros géneros en Sur América con caracteres morfoldgicos similares no es comutn, pudiéndose confundirse solamente con algunas especies de Mandevilla y Macrosiphonia con habito erectoe inflorescencias terminales. Sin embargo, estos géneros se pueden separar con facilidad por la presencia usual de coléteres, ya sea en el nervio central de las hojas adaxialmente, asi como en la base de la cara adaxial de los sépalos. Por otro lado, aunque Prestonia erecta (Malme) J.F Morales también tiene habito erectoe inflorescencias terminales, la presencia de una corona anular alrededor de la boca en esa especie la separa al instante, asi como hojas con la lamina foliar membranacea. Siguiendo con la revision de géneros de las tribus Echiteae y Mesechiteae (sensu Endress & Bruyns 2000), una monografia del género Galactophora es SIDA 21(4): 2053-2079. 2005 2054 BRIT.ORG/SIDA 21(4) presentada a continuacion. Para tal fin, la mayoria de colecciones tipo fueron revisadas, asi como las colecciones depositadas en los principales herbarios de Europa, Norte América y del N de S América (Colombia, Ecuador, Pert y Bo- livia). El esquema utilizado aca sigue el empleado anteriormente en otras revisiones y monografias de la misma serie (e.g., Morales 2002, 2003). Caracteristicas morfologicas notables Hojas.—Las hojas son opuestas, aunque raramente hojas verticiladas puedan ser encontradas en algunos especimenes, careciendo de coléteres en el nervio cen- tral (adaxialmente). Las laminas son invariablemente coridceas, con los margenes usualmente revolutos al secar y es comun que las venas secundarias apenas estén impresas en ambas superticies, mientras que las venas terciarias raramente estan impresas (e.g., G. schomburgkiana Woodson). Pubescencia.—La pubescencia de tallos, hojas e inflorescencias (cuando presente) es tipica de Galactophora y esta compuesto por pelos unicelulares, lati te rigidos y usualmente de color oscuro al secar, con la parte distal y apical, conspicuamente engrosada y glandular (Fig. 1). Este tipo de pubescencia no esta presente en el resto de los miembros de las tribus Mesechiteae ni Echiteae, a pesar de la alta variabilidad de patrones de pubescencia presentes en géneros como Mandevilla. Sol: teen M. pachyphylla Woodson, pelos cortos, bul bosos y a veces levemente uncinados estan presentes, pero nunca como los descritos en Galactophora. En forma tradicional y continua, la pubescencia ha sido utilizada por varios taxonomos para separar multiples taxones, entre ellos De Candolle (1844), Muller (860) y Woodson (1933, 1935, 1936). Sin embargo, la variacion intraespecifica en forma general en las Apocynaceae puede ser muy alta y este caracter debe ser usado en la medida de lo posible con precaucion y en conjuncion con otros caracteres morfoldgicos. De esta manera, tomando en cuenta la relativamente escasez de cantidad de material disponible en el siglo 18 yen la primera mitad del siglo 19, muchas especies fueron separadas basados en rangos extremos de pubescencia, sin especimenes intermedios que permitieran delinir la conexion entre estos extremos. El relativamente alto numero de material disponible hoy en dia, ha demostrado que varias de esas especies solo representan variaciones extremas de la pubescencia dentro de un mismo taxon y han debido ser sinonimizadas en monografias o revisiones recientes (e.g., Fallen 1983; Hansen 1985; Leeuwen berg 1994 a; Morales 1997, 1998, 1999, 2002, 2003; Williams 1998). De esta manera, la misma situacion se ha presentado en Galactophora, donde G. crassifolia y G.calycina han sido separadas basicamente por el indumento presente en la primera y ausente en la segunda (Woodson 1936; Morillo 1995). Tanto los ejemplares tipo como el resto de colecciones examinadas son idénticos en su morfologia general y se encuentran una serie de especimenes con grados intermedios de pubescencia, lo que impediria aun mas el pensar en reconocer estas dos formas a nivel de variedad. MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2055 Fic. 1. Detalle de | 1 1 1,1 ay oe pee ee | or yt ( 5 ef & Bangi 4627 INB). tOL/, Esta variabilidad de la pubescencia en este género, reafirma nuevamente laalta variabilidad intraespecifica en las Apocynaceae neotropicales de ese caracter. Inflorescencias.— Woodson (1936) originalmente describi6 auc las inflorescencias en este género eran subumbeliformes o escorpioidales y corimbosas, pero el andlisis y estudio de este caracter ha determinado que en realidad las inflorescencias son cortamente racemosas 0 reducidas a cimas (ocasionalmente con apariencia subumbeliforme), algunas veces con solo 1 6 2 flores. En forma usual tienen pocas flores,con un rango de 1a5 por inflorescencia, aunque ocasionalmente se encuentran grupos de hasta 1] flores. Las bracteas florales son siempre escariosas y no se reporta la presencia de bracteolas a lo largo del pedicelo. Cdliz.—El caliz presenta cinco sépalos, los cuales pueden ser totalmente glabros o presentar en diferentes grados de densidad, el wotee indumento de Galactophora. La presencia 0 ausencia del indumento no es determinante para separar especies, pero la longitud de los sépalos en algunos casos puede ser util para separar algunos taxones. Sin embargo, la mas notable e importante diferencia respecto a tratamientos monograficos previos se refiere a la presencia léteres en la base de la cara adaxial de los sépalos. Woodson (1936) reporto 2056 BRIT.ORG/SIDA 21 4) que los sépalos tenian “several to many indefinitely distributed squamellae,” es decir, que albergaban una serie de coléteres fusiformes, pequenos y diminutos, similares a los presentes en otros géneros de las Echiteae, como por ejemplo, Peltastes. Sin embargo, después de examinar numerosos sépalos de diferentes especimenes de todas las especies, es claro que Galactophora no tiene coléteres, tal y como fueron descritos e ilustrados en la monografia de Woodson (1936). La ausencia de coléteres fue previamente notado por Morillo (1995), quién en su tratamiento de Galactophora para la Flora de la Guyana Venezolana, cito que los sépalos, al parecer, carecian de coléteres en la base interna. En forma usual, los coléteres estan presentes en casi todos los miembros de las Echiteae y Mesechiteae, con la excepcion de Laubertia y Rhabdadenia. Corolas.—En forma general, es comun que las corolas presenten cinco linales externas que corren desde la base del tubo hasta el margen de los lobulos (Fig. 2). Aunque en material fresco las crestas son apenas evidentes, son mucho mas evidentes en material seco. Este tipo de crestas no estan presentes en ningun otro miembro de las Mesechiteae o Echiteae y es un costillas ocrestas longit caracter bastante distintivo que nos ayuda a reconocer con facilidad las especies de este género. Las partes de la corola mencionadasa través de las descripciones siguen a Morales & Fuentes (2004 b). Ovario y foliculos.—En todas las especies, los dos carpelos se hayan fusionados postgenitalmente en forma basal, separandose ligeramente por encima del nectario. Esto provoca que los foliculos, casi en forma invariable, se hallen fusionados en el area basal proximal al caliz en un area inferior a 2(-2.5) cm de su longitud, para luego separarse de manera definitiva, continuando de forma divergente, continua y rigida. Este tipo de fusion basal, da una falsa apariencia de que los foliculos fueran estipitados (Fig. 3). Este caracter no esta presente en el resto de las Mesechiteae 0 Echiteae y es bastante distintivo para el género. Notas en la clasificacion intragenérica El género Galactophora fue descrito en forma relativamente reciente por Woodson (1932), por lo que no fue incluido como tal en clasificaciones infragenéricas previas (e.g., Schumann 1895). En forma posterior, Pichon fue el primero en proponer una extensa clasificacion de las Apocynaceae (1948 a, b, c; 1949; 1950 a, b) , clasificando a Galactophora en la subfamilia Echitoideae (“Echitoidees”), tribu Parsonsieae (“Parsonsiées”), junto con la mayoria de los miembros de las Echitoideae sensu Woodson (1935, 1936), excepto Mandevilla y sus géneros satélites (e.g., Allomarkgrafia, Mesechites). Adicionalmente, en su clasificacion de las Echotoideae, Pichon (1950 a), ubicé a Galactophora dentro de una subtribu propia (Galactophorinae), basado en algunos caracteres de la corola y frutos. Posteriormente, Leeuwenberg (1994 b) propuso otro sisterma de clasificacion, tratando a Galactophora dentro de la tribu Echiteae subtribu MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2057 Fic 2 Niotall dol +#ill INB). B. G. sh J O° Alot | age fate NB) ~ =] Echiteae, donde tanto Mandevilla, Echites y los géneros relacionados, fueron tratados en la misma subtribu. Sin embargo, en forma general, las tribus propuestas por Leeuwenberg son artificiales (Sennblad et al. 1998) y confusas, debido a la ausencia de una explicacién de los criterios utilizados en su delimitacion. Ahora bien, Endress & Bruyns (2000) propusieron la mas reciente clasificacion de las Apocynaceae, incluyendo por primera vez Asclepiadaceae, tal y como fue sugerido por analisis moleculares previos (e.g,, Judd et al. 1994; Sennblad & Bremer 1996). En el sistema de Endress & Bruyns (2000), Galactophora fue tratado en la tribu Mesechiteae, junto con Mandevilla y sus géneros relacionados. La inclusion de Galactophora dentro de esta tribu, fue basado probablemente por la presencia de las cinco proyecciones o costillas basales de la cabeza estigmatica. Sin embargo, sus hojas carecian de coléteres en la superficie adaxial del nervio, caracter comun al resto de géneros en esa tribu, con la excepcion de Secondatia, otro género que fue excluido de esa tribu en forma preliminar por Morales (2003), basado en las caracteres de la cabeza estigmatica. Las relaciones intergenéricas de los miembros de las Mesechiteae 2058 BRIT.ORG/SIDA 21(4) Fic. 3 Natall Do [eg Bee ee lanl £1. | fal n tf, 42 Ali o 1, DIRT MOB Guillen & Roca 2853, \NB. han sido confusas, debido a los caracteres algunas veces relativamente débiles usados en su separacion. En forma reciente, Simoes et al. (2004), realizaron un estudio molecular y morfologico para probar la monofilia de la tribu Mesechiteae, donde se confirm6 que Galactophora debe ser excluido de la tribu Mesechiteae, al igual que Secondatia. Por lo tanto, dado que su ubicacion tribal actual es incierta, no se puede comentar en forma apropiada las relaciones intergenéricas de Galactophora. TRATAMIENTO SISTEMATICO Galactophora Woodson, Ann. Missouri Bot. Gard. 19:49. 1932. Tivo: Galactophora crassifolia (Mull. Arg.) Woodson Hierbas arbustivas, raramente escandentes o lianas; las ramitas tiernas algunas veces aplanadas o anguladas, usualmente teretes 0 subteretes con la edad, con secrecion lechosa o acuosa, glabras, glabrescentes a variadamente glandular- pubescentes, los coléteres inter e intrapeciolares, fusiformes e inconspicuos. Hojas opuestas, raramente verticiladas, sésiles, subsésiles a pecioladas, peciolos eglandulares, sin coléteres a lo largo de la costa adaxialmente, pero con MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2059 inconspicuos coléteres en los axilas; laminas glabras, glabrescentes a variadamente glandular-pubescentes, coriaceas a subcoriaceas, eglandulares, sin coléteres en el nervio central adaxialmente, usualmente variadamente revolutas, la venacion secundaria y terciaria usualmente inconspicua, mas raramente levemente impresas en alguna de las caras de la hoja. Inflorescencias racemosas 0 cimas reducidas, terminales o subterminales, usualmente con pocas flores (1 a 11), glabras, glabrescentes a variadamente glandular-pubescentes, pedunculadasa sésiles 0 subsésiles, bracteas escariosas, inconspicuas, bracteolas ausentes; sépalos 5, esencialmente iguales, algo imbricados basalmente, escariosos, subfoliaceos, o folidceos, sin coléteres en la base de la cara interna: corola infundibuliforme, glabra, glabrescente a variadamente glandular- pubescente exteriormente, sin corona anular 0 corona epiestaminal (apéndices coronales libres) interiormente, usualmente con cinco crestas o costillas longitudinales externamente, distribuidas desde la base del tubo hasta la base de los lobulos, el limbo dividido en SCO OE ues, estes waliscamerte nervados longitudinalmente, con aestivacion dextrorsa; estam inco, insertos dentro del tubo, incluidos, los filamentos cone ene 2 indumento infraestaminal usualmente presente; anteras os y aglutinadas endos puntosa la cabeza estigmatica, las bases estéril ,carpelos dos, unidos apicalmente al estilo, fusionados Sent en su region basal, usualmente hasta la altura del nectario; cabeza estigmatica con cinco proyec costillas longitudinales basales, el resto relativamente cilindrico, con el apice algo engrosado; 6vulos numerosos, multi-seriados, dispuestos en una placenta axilar y biseriada; nectario anular, entero, subentero o irregularmente lobulado, mucho mas corto que el ovario. Foliculos 2 apocarpicos, teretes a subteretes, continuos, fusionados en su parte basal y luego separados y divergentes entre si, glabros, glabrescentes 0 variadamente pubescentes, dehiscentes a lo largo de la sutura ventral; semillas numerosas, secas, truncadas apicalmente, comosas en el apice micropilar, usualmente rugosas, mas raramente casi lisas. Género neotropical con 6 especies, distribuido principalmente en el SE de Colombia, Venezuela y Brasil, con una especie extendiéndose hasta Pert y el NE de Bolivia. CLAVE PARA LAS ESPECIES DE GALACTOPHORA 1. Sépalos 15-25 mm de largo 3.G. crassifolia 1. Sépalos 4-12.5 mm de largo. 2. Parte iBtevieree rn tubo de ge corola de 19-31 mm de largo,la boca de (18-)23-32 fe venacion secundaria y terciaria impresa adaxialmente; distribucién geografica en tepuis entre 750-2000 m .G, segue 2. Parte inferior del tubo de la corola de 5.5-14 mm de largo, la boca de 5-16 m diametro; venacién secundaria levemente o no impresa adaxialmente, ie 2060 BRIT.ORG/SIDA 21(4) venas terciarias no i be distribucion geografi ysak y Areas diversas entre 100- 1200 m. 3, Laminas foliares de 8.5-15.5(-17) x (4-)5-8 cm, ovadas a ovado-elipticas, levemente cordadas basalmente 2.G.colellana IOCVETNMEMILE C 2. ov foliares de 1.7-6.2 x 0.5-2.4.cm, as a< elipticas, obtusas a redondeadas basalm Cul 4. la con g base cordada, sésiles o ces 1.G. angustifolia 4. Hojas con la base cuneada, redondeada a “obtusa, pecioladas o si cortamente erie la base nunca cordada. 5. Inflorescencias con 5 a 11 flores, pedunculo de 7-42 mm de largo; sépalos de 4-6 mm de largo; parte superior del tubo de la corola de 15 16mm de longitud 3. HGyPUleneNs 5. Inflorescencias con | a 2 flores, pedunculo menos de 2 mm de largo sépalos de 8-11 mm de largo; parte superior del tubo de la corola de 23-31 mm de longitud 4.G. pumila 1. Galactophora angustifolia J.[° Morales, Sp n0¥ wv. (Fig. 4). ve Cc ‘OL ounle. CAQUETA: parque nacional natural Chiribiquete 19 Nov 1992 (fD, M. Velayos, J. Cardiel, J. Pedrol & M. felleria 6319 GOLOTIPO: MA: ee P fotogratia, INB) AG. pulchella Woodson, cui alfinis, loliis cordatis, sessilis vel subsessilis et 1-1.4 cm latis differt. idos, glabros o glabrescentes pelos glandulares (cuando Habito desconocido; tallos subteretes a teretes, so e inconspicuamente glandular-hirsutulos, los presentes) Negros, coléteres inter e > intrapeciolares i inconspicuos, ca. | mm de largo. Hojas sésiles a subsésiles, el peciolo menos de | de largo; lamina 6.8-9 x |- 1.6 cm, angostamente ovadas a angostamente ovado-elipticas, agudas apicalmente, cordadas basalmente, coriaceas a subcoriaceas, glabras o glabre- scentes y con inconspicuos pelos esparcidos abaxialmente, levemente revolutas marginalmente o no revolutas, la venacion secundaria levemente impresa abaxialmente, algunas veces inconspicua, las venas terciarias no impresas. Inflorescencia terminal, glabrescente, con mas de 6 flores, pedunculo 5-6 mm de largo, pedicelos 2.5-3 mm de largo, bracteas 1-15 x 0.5-1 mm, escariosas; sépalos 5-6 = I-1.3 mm, angostamente ovados a angostamente ovado-elipticos, acuminados, subfoliaceos, glabrescentes a muy esparcidamente glandular- hirsutulos exteriormente; corola de color desconocido, glabrescente exteriormente, con solo pelos distribuidos en las lineas externas de la corola, el apice del boton floral cortamente acuminado; parte inferior del tubo 6-7 1L.5- 2mm, la parte superior 14-15 mm de largo, angostamente conica, 3-4 mm en diametro en el orificio; lobulos 7-8 x 4-5 mm, angostamente obovados; anteras 4-4.2 mm de largo, glabras, las bases estériles cortamente acuminadas, cabeza estigmatica 1.5-1.8 mm de largo; ovario 2-2.6 mm de largo, glabro; nectario 0.8- | mm de largo, entero a subentero. Foliculos desconocidos Distribucion, habitat y ecologia.—Endémica a Colombia al Departamento de Caqueta, donde es conocida unicamente de la localidad tipo, creciendo en — Ee, MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2061 2.5 mm 33mm L ! 5 com = 4mm C Simm Fie Gal A au gee apy 1 6279 P) A. Ramit infl 195. B. Sénalos. vista adaxial. mostrando | i lét C. Antera. vista dorsal. D. Cab ti 4tica. E. Nectario y ovario J 2062 BRIT.ORG/SIDA 21(4) vegetacion en bordes de quebradas, en elevaciones de 645 m. Especimenes con flores han sido recolectados en Noviembre. Galactophora angustifolia es una especie muy distintiva, que se podria confundir con G. pulchella, pero se diferencia con facilidad por sus hojas sésiles o subsésiles, con el peciolo inferior a 1 mm de largo, hojas angostamente ovadas o angostamente ovado-elipticas, con la lamina |-1.6 cm de ancho y con la base cordada, asi como la corola con la parte inferior del tubo de 6-7 mm de largo. Tanto el habito como el color de las flores de esta especie son aun desconocidos, debido a que ambos caracteres no son especificados en la etiqueta del holotipo. _— en) 2. Galactophora colellana Morillo, Anales Jard. Bot. Madrid. 48:27. 1990. (Fig. . Tipo: VENEZUELA. AMAZONAS: Rio Negro, Cerro de la Neblina, NNE del Pico Phelps, 15 Mar 1984 ({D, Liesner 16644 (HOLOTIPO: VEN; ISOTIPOS: INB, MO, NY). A Arbusto erecto hasta 1.3 m de altura; tallos aplanados en ramitas jovenes, subteretes a teretes cuando viejos, sdlidos o levemente huecos, esparcidamente glandular-hirsutulos, los pelos glandulares negros; coléteres inter e intra- peciolares inconspicuos, ca. | mm de largo. Hojas sésiles a subsésiles, el peciolo 1-1.5 mm de largo; lamina 8.5-15.5(-17) x (4-)5-8 cm, ovadas a ovado-elipticas, obtusas, agudas, agudo-mucronuladas a obtusas apicalmente, levemente cordadas basaliente, coriaceas a subcoriaceas, muy esparcida y diminutamente papilado-puberulentas en ambas superficies, algo revolutas marginalmente, la venacion secundaria levemente impresa, algunas veces inconspicua, las venas terciarias no impresas. Inflorescencia terminal, diminutamente puberulenta, esparcidamente glandular-setosa, con 5 a 8 flores, pedunculo 14-28 mm de largo, pedicelos 5-8 mm de largo, bracteas 1-1.5 x 0.5 mm, escariosas; sépalos 4-5 x 2-2.5 mm, ovados, acuminados, subfoliaceos, muy esparcidamente glan- dular-hirsutos exteriormente; corola blanca, muy esparcida e inconspicuamente glandular puberulenta a glabrescente exteriormente, el apice del boton floral agudo; parte inferior del tubo 13-14 x 15-2 mm, la parte superior 15-16 mm de largo, angostamente conica, 6-7 mm en diametro en el orificio; lobulos 9-10 x 4-5 mm, angostamente obovados; anteras 4.5-5 mm de largo, glabras, las bases estériles cortamente acuminadas, cabeza estigmatica 1.5-1.8 mm de largo; ovario 2-2.5 mm de largo, glabro; nectario 0.8-1 mm de largo, entero a subentero. Foliculos desconocidos. Distribucion, hdbitat y ecologia.—Esta especie se encuentra distribuida en Colombia, Venezuela y Brasil, donde crece en vegetacion arbustiva enana, sabanas arbustivas y afloramientos rocosos en elevaciones de 400-800 m. Especimenes con flores han sido recolectados en Marzo, Julio, Septiembre y Octubre. Galactophora colellana es una especie que se puede confundir con G. crassifolia y G. schomburgkiana por el tamano y forma de sus hojas, pero que se fees MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2063 4onm 4cm 2mm 2mm 4mm B. Sénalo. vista adaxial. C. Antera. vista I ' Fic.5. Galactophora colellana (Liesner 16644, INB).A.Ramit dorsal. D. Cabeza estigmatica.E. Nectario y ovario. 2064 BRIT.ORG/SIDA 21(4) separa lacilmente por sus flores con la parte inferior del tubo de la corola de 13- 14 mm de largo (vs. 19-32 mm), asi como sépalos de 4-5 mm de largo (vs. 5-25 mm). | dos. COLOMBIA. Amazonas: rio Caqueta, La Pedrera, 2 Oct 1952 (11), Schultes & Cabrera 17673 (US). VENEZUELA. Amazonas: Atabapo, Cerro Huachamacari, 5 Nov 1988 ({1), Liesner 25878 (MO). BRASIL. Amazonas: Sao Gabriel do Cachoeira, Morro dos Seis Lagos, Lago do Dragao, Oct 1987 (fl), Farney et al. 1723 (NY); Rio Negro, Morro do Seis Lagos, sin fecha (fl), Weber & Knob 1719 (NY). ce ae et Sone wen |. Arg.) Woodson, Ann. Missouri Bot. Gard. 19:50. 1932. (Figs. 3, 6). Amblyanthera crassifolia Mull. Arg. FL Bras 6(1):143. 1860. Rhodocalyx ee OF Arg.) Miers, Apocyn. S. Amer. 139. 1878. T1po: BRASIL. AMAZONAS: cerca de San Carlos, Rio Negro, 1853-1854 (£1, fr), Spruce 3136 (HOLOTIPO: K; IsOTIPOs: B (destruido, foto F neg 38739), BM Coto, INB), BR, CGE, G (foto F neg 26870), K [2 laminas], NY, PW). Dipladenia calycina Huber, Bol. Mus. Goeldi 7:113. 1913, nom. nud Dipladenia calycina Huber ex Ducke, Arch. Jard. Bot. Rio de Janeiro 3:247. 1922. Galactophora calycina (Huber ex Ducke) Woodson, Ann. Missouri Bot. Gard. 19:50. 1932. Tipo: BRASIL. PARA: rio Mapuera, Trombetas, al NE de los rapidos de Taboleirinho, 12 Dic 1907 (1, fr), Ducke 9124 (HOLOTIPO: MG, foto en INB). actophora magnifica Woodson, Ann. Missouri Bot. Gard. 19:382-383. 1932, syn. nov. Tipo: BRASIL. MATO GROSSO: Juruena, Abr 1909 (ED), Hoehne 1759 (HOLOTIPO: US). ae Ga Arbusto erecto (0.5-)1-2.5 m altura, algunas veces escandente y mas raramente una liana; tallos conspicuamente aplanados y angulados en ramitas jOvenes, subteretes con la edad, solidos, densamente glandular-hirsutos en ramitas jovenes, esparcidamente glandular-hirsutos a erapies ones): o glabros en tallos viejos, los pelos glandular es Negros, coléteres i Inte1 peciolat es inconspic UOS, 1mm o menos de largo. Hojas sésiles a subsésiles, el peciolo 1-2 mm de largo; lamina (2.5-)3-11 x 2.1-8 cm, ovadas, ovado-elipticas, obtuso-mucronuladas a emarginado-muc ronuladas api almente _cordadas a subc ordadas b asalmente, I coriaceas a subcoriaceas, glabras a or adaxialmente, diminuta y esparcidamente papiladasa glabras 0 g] baxial algunas veces con puntuaciones negras, var cea revolutas marginalmente, la venacion secundaria levemente impresa, algunas veces inconspicua, las venas terciarias usualmente no impresas. Inflorescencia terminal a subterminal, esparcidamente glandular-puberulenta,con | a5 flores, pedanculo 8-23 mm de largo, pedicelos 10-16 mm de largo, bracteas 0.5-1 « 0.5-1 mm, escariosas: sépalos 15-25 x 4-6 (-7) mm, ovados, angostamente ovados a ovado-elipticos, agudos 0 acuminados foliaceos, esparcidamente hirsutulos a glabrescentes; corola con el tubo crema, blanco a blanco-rosado, los lobulos rosados, muy esparcidamente glandular- puberulentas a glabrescentes, el apice del boton floral agudo; parte inferior del tubo 22-32 x 2.5-3 mm, la parte superior 24-32 mm de largo, conica, 11-16 (-24) mm en diadmetro en el orificio; lobulos 24 x 18 mm, angostamente obovados; anteras 7-8 mm de ie glabras, las bases estériles cortamente acuminadas, cabeza estigmatica 1.9-2.2 mm de largo; ovario 2.9-3.2 mm de largo, MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2065 5mm 4 3mm 2em Fic.6. Galactopt ifolia (Guillén & Roca 2853, |NB).A. Ramit fl y frutos.B. Sépalo, vista adaxial C. Antera, vista dorsal. D. Cab tig Atica. E. Nectario y ovario. F. Semilla 2066 BRIT.ORG/SIDA 21(4) glabro; nectario 1-1.2 mm de largo, leve e irregularmente pentalobulado. Foliculos LO-17 x 0.3-0.5 cm, continuos, glabros, glabrescentes 0 variadamente glandular-puberulentos; semillas 3-4 mm de largo, glabras, coma 19-27 mm de largo, crema a blanco-crema. Distribucion, habitat y ecologia.—Se encuentra en Colombia, Venezuela, Brasil, Peru y Bolivia, donde crece en sabanas, bosques enanos en formaciones de arena blanca y en afloramientos rocosos con vegetacion arbustiva enana, en elevaciones de 50-600 m. Especimenes con flores han sido recolectados en todos los meses del ano. Frutos se reportan de Junio a Febrero. Nombres comunes.—cupia blanco (Venezuela, Amazonas); bejuco de carbon (Venezuela, Amazonas). Galactophora crassifolia es la especie mas comutn y amp distribuida, asi como la mas facil de reconocer dentro del género por la longitud de sus sépalos, los cuales varian de 15 a 25 mm de largo (vs. 5-12.5 mm ene resto de los taxones). Es comun que esta especie sea una hierba erecta 0 un arbusto pequeno, pero también se conocen especimenes que reportan su habito en forma de liana (e.g., Liesner 3415, MO; Guillén & Roca 2853, INB Galactophora pacing es remue a la sinonimia. (sc mene la Unica diferencia en los ej ipod s especies es la presencia del indumento de pelos Slandularest enG.c al id y laausencia de los mismos en G. calycina, ~~ lamente — siendo similares en la longitud de las corolas, sépalos, anteras y demas caracteres. Esto fue comentado en forma previa por Morales y Fuentes (2004 a), quienes Sugirieron la sinonimia de G. calycina. En el tiempo de la monografia de Woodson (1936), estas especies eran conocidas por solo 17 colecciones (G. crassifolia con7 y G.calycina con 10),de las cuales solamente 9(7 y 2 resperctiva- mente) fueron revisadas por Woodson. El numero evidentemente mayor de material disponible hoy en dia, ha permitido conocer mejor la variacion morfologica intraespecilica de esta especie y demilitar mas claramente sus limites reales. Por otro lado, varias estados intermedios son encontrados, con colecciones con muy escasos pelos, lo que impide atin mas la segregacion de ambas especies. Es importante recalcar que la pubescencia por si sola en un caracter bastante variable dentro de las tribus Echiteae y Mesechiteae, donde existen géneros y especies con una alta variabilidad morfologica en ese caracter y que por lo tanto, la separacion de taxones por Unicamente esa caracteristica es insostenible. Basado en los comentarios y conclusiones anteriores, G. magnifica es también reducida a la sinonimia, pues no existen diferencias con la coleccion tipo de G. crassifolia, excepto por la ausencia del indumento en la primera. Especimenes examinados. COLOMBIA. Amazonas: rio Apaporis, raudal de Jirijirimo, | Sep 1986 (1), Bernal et al. 1243 (COL); Leticia, rio Caqueta, Santa Isabel, 9 Abr 1994 (1), Cardenas & Gangi 4627 (COAH, INB, MO); rio Caqueta, 25 Jul 1977 (ED, Ferndndez-Péres 20121 (COAH, COL); rio Caqueta, Araracuara, sabana de la Angostura, 21 Dic 1951 (1), Garcia-Barriga & Schultes 14149 (COL, US); MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2067 Araracuara, rio Caqueta, 7 Ago 1977 (f1), Idrobo 8932 (COL); Araracuara, 8 Ago 1977 (fr), Id robo 8965 (COL); rio mee Araracuara, Nov 1982 (f1), Idrobo et al. 11266 (COL), Idrobo et al. 11465 (COL); Araracuara, rio Caqueta, 6 Sep 1959 (f1), Maguire et al. 44167 (INB, NY, U), Araracuara, balcon del diablo, 16 ae 1984 (f1), Palacios et al. 458 (COAH), 18 Feb 1986 (fr), Palacios & Plazas 855 (COAH), 3 Mar 1986 (fr), Palacios & Plazas 1168 (COAH), 5 Mar 1986 (f1, fr), Palacios & Plazas 1220 (COAH); Araracuara, 21 Sep 1990 (f1), Restrepo & Sdnchez 11(COAH, MO), rio Caqueta, Ser aes 1 Abr ns (fl), Sastre & Reichel 5190 (COL, P); rio Apaporis, Raudal de Jerijerimo, Mar 1951 (fD, Schultes 12109 (COL, GH, MO); rio Apaporis, Cachivera de Jirijirimo, 11 Jun 1951 CD, Schultes rs Cabrera 12368(COL, MO, NY, U, US); rio Apaporis, Cachivera de Jirijirimo, 7 Jul 1951 (fr), Schultes & Cabrera 12968 (COL, GH, MO, US); rio Apaporis, Cachivera de Jirijirimo, 16 Sep 1951 (fD, Schultes & Cabrera 14060 (COL, MO, US); NE de San Carlos de Rio Negro, 9 Abr 1984 (£D, Stein a we) Amazonas-Vaupés: Rio a Jirijirimo, Nov 1951 1D, Garcia-Barriga 13752 (COL, NY, | o Apaporis, 28 Feb 1952 ({1), > van der Hammen 154 (COL); rio Apaporis, raudal eee 27 Nov 1951 D), Schultes & ae 14626(COL). So pes NO de Araracuara, 6 Nov 1992 (fl), Arbelaéz & Sueroque 314 (HUA); Solano, rio Caqueta, Araracuara, 15 Nov 1992 (1), Arbelaéz & Matapi 347 (COAH, HUA): Solano, Araracuara, 10 Nov ean ae 426 (COAH, HUA); Santa Isabel, Sabanas del Solarte, 4 Dic 1996 (fr), Arbelaéz et al.626 (COAH, HUA, U), tie nacional natural Chiribiquete, serrania e, 26 Nov 1992 (f1, fr), Barbosa & Rueda 8063 (MA); Araracuara, rio Caqueta, balcon del diablo, - ~~ 1993 (1), Cardenas et al. 4121 (COAH): arioiaes 13 Nov 1991 (f1), Duivenvoorden e al. 2743 (COAH); Araracuara, Balcon ae diablo, 13 Dic 1983 (f1. fr), Forero & Pabon 9815 (COAH, COL, MA); sierra del Chiribiquete, ref rdo, 22 Nov 1992 (f1), Franco et al. 4264 (COL, MA): sierra de Chiribiquete, 13 Dic 1991 (ED, GCateano etal. 2252(COl ,MA); Araracuara, Balcon del Diablo, 25 Jan 1989 (fl), Gentry & Sdnchez 65166 (MO). Guainia: serrania de Naquén, nel 24 Jul 1992 (£1), Cortés et al. 129 (COAH); Puerto Inirida, 15 Ago 1975 (fl), Garcia-Barriga 20879 (COL, F US), Cano Colorado, La Esperanza, 15 Abr 1993 (fl), Madrindn & Barbosa 1057 (COL, GH, MO); caserio de Santa Rita, rio Guainia, 15 Oct 1977 (£1), Pabon et al. 355 (COL); Puerto Colombia, rio Negro, 27 Oct 1977 (£1), Pabon et al. 427° (COL). Vaupés: rio Vaupés, cachoeira de Yurupari, 10 Nov 1943 (fD), Allen 3163 (COL); serrania de Taraira, al NO de raudal de la Libertad, 4 Ago 1993 (fD, Cortés & Rodriguez 788 (COAH, OL); 5 Ago 1993 (fD, Cortés & Rodriguez 805 (COL), Mita, entre Timbo y Bogota Cachivera, 7 Jul (f1), Galeano et al. 95 (COL); Mitu, Mandi, rio Vaupés, 29 Jul 1993 ({D, Galeano et al. 1135 (COAH, COL): rio Vaupés, Miriti, 24 Nov 1993 (fl), Galeano et al. 1873 (COAH, COL); rio Kubiya, eas 30 Jun 1958 (£1), Garcia- ae et al. 16037 (COL): Cerro de Chiribiquete, rio Macaya, 17 Jai 4(f1), Gutiérrez & Schultes 684 (MEDEL); rio Negro, San Felipe, 13 Nov 1952 (f{D, Humbert se a i cano Cubiyu, eres Indigena 2 Sabana, 26 Abr 1993 (fr), Madrindn et al. 1109 (COL); Mita, rio Vau 28 Oct (fr), Mejia et al. 2678 (COL); rio Piraparana, tributario del rio Apaporis, 6 Sep 1952 (ED), Sc a i eee 17228 (US), 18 Sep 1952 (fL), Schultes & Cabrera 17508 (US), rio Negro, San Felipe, 25 Oct 1952 (fl, fr), Schultes et al. 18003 (MO, US); rio Vaupés, entre Mitt y Javareté, 14 May 1953 (£1), Schultes & Cabrera 19381 (COL, MO, U, US); rio Vaupés, Raudal de Yurupari, Nov 1951 (f1, fr), Schultes & Cabrera 19713 (MO, U); Mitt, rio Parana-pichuna, 2 Jul 1975 (st), Za rucchi 1364 (COL); Mitu, rio Kuubiyt y rio Parana-Pichuna, 7 Jul 1975 (f1, fr), Zarucchi 1395 (COL, GH, K), 9Jul 1975 (f1, fr), Zarucchi 1407 (COL, GH, K); Mitu, sobre el rio Vaupés, Circasia, 13 Sep 1976 ({D, Zarucchi 2030 (COL, GH, k): ie rio Vaupés, entre rio Ti y rapidos de Mandi, 23 Sep 1976 ({1), Zarucchi 2116 (COAH, COL, GH, k, 1S); Mita y cercanias, 11 Nov 1976 (£1, fr), Zarucchi 2213 eves GE 1, K, MO, US); Mitu, rio Kubiyu, on 1979 oe eu 2463 2 ae COL, GH, MA, MO, U, W. LA. Amazonas: Atures, N de Santa Rosa de Ucata, ne 1992 - fr), Berry et al. 5197 (MO); a a del rio Temi, i 1995 (fl, fr), Berry et al. 5587 (MO); Bana, S de Yavita, 28 Nov 1995 (fl), Berry et al. 5709 (MO); San Carlos de Rio Negro, 3 Abr 1978 (£1, clad k 6587 (QCNE); Santa Cruz, rio Atabapo, 9 Set 1960 ({1), Foldats 3804 (NY, VEN); cerro Yapacana, rio Orinoco, Abr 1931 (fD), Holt & Blake 716 (US), Atures, Valle del Cano Camani, O de Cerro Morrocoy, 10 Oct 1979 (1), Huber 4636 (MO, MYE NY, VEN, Z); Rio Negro, carretera San Carlos-Soano, 16 Sep 1980 (fl), Huber et al. 5658 Kw eS, 2068 BRIT.ORG/SIDA 21(4) (COL, K, MO, NY, VEN); E de San Carlos de Rio Negro, 13 Nov 1977 (f1, fr), Liesner 3415 (MO): Bana, NE de San Carlos de Rio Negro, 26 Nov 1977 (f1, fr), Liesner 3903 (MO); Rio Negro, faldas de Cerro Aracamuni, Oct 1987 (LD, Liesner & Carnevali 22403 (MO); Rio Negro, NE de San Carlos de Rio Negrc 1987 (fl, fr), Liesner & Carnevali 22881 (MO); Cerro Yavita, rios Atabapo y Orinoco, 19 Oct 1950 (i D. ne 29300 (NY); rio Guainia, E de Maroa, 25 Nov 1953 (fl, fr), Maguire et al. 36399 (kK, NY, US); rio Guania, E de Maroa, 6 Oct 1957 (f1), Maguire et al. 41711 (COL, NY, P, W), 7 Oct 1957 (£1), Maguire et al. 41739 (NY, US); N de Puerto Colombia, Rio Guainia, 12 Oct 1957 (fl), Maguire et al. 41842 (NY); Rio Negro, San Carlos do Rio Negro, caminoa Solano, 21 Abr 1984 ([)), Plowman 13708 (F. WAG): N de sees rio Sopapo, 10 Oct 1988 (1), Romero & Gudnchez 1629 (GH, NY); Ede Santa Rosa de Ucata, 23 Oct 1989 ({D, Romero & Melgueiro 2234 (GH, MO, NY); Rio Negro, San Carlos, 15 Dic 1947 (£D, Schultes : Lopez 9372 (MO); Rio Orinoco, cerca de Fernando de Atabapo, 25 Abr 1954 ({D, Silverio LO (MO, NY, P); Rio Negro, San Carlos de Rio Negro, 23-29 Jul 1982 ({1), Stergios & Aymard 4266 (PORT, US), cerca de San Carlos de Rio Negro, 1984 (f1), Stergios & Aymard 7725 (MO, PORT), entre sabana Esmeralda y base SE de Cerro Duida, 22 Ago 1944 (IL), Steyermark 57840 (FE MO); Sabanita Morocoto, rio Orinoco, 30 May 1959 (f1, fr), Wurdack & Adderley 42683 (NY); Atabapo, rio Cuchaken, Oct 1989 (£1), Velasco 763 (MO, PORT). PERU. Madre de Dios: rio Health, Santuario Nacional de las Pampas del Health, 14 Jun 1992 (1, fr), Castillo & Foster 6910 (F USM), BRASIL. Amazonas: rio Uatuma, ltapiranga, 18 Ago 1979 ({]), Cid et al. 475 (NY); carretera Manaus-Caracarai, 20 Feb 1979 (f1, fr), Coélho et al. 940 (BM, INPA); Airao, entre Manaus-Caracarai, 27 Mar 1979 (fD, Coélho et al. 1395 (BM, COL, INPA); Ponta Negra, cerca de Manaus, 20 Nov 1910 (f1), Ducke 11197 (MG, loto en INB); Manicoré, camino a Humaita, 24 Abr 1985 (1), Ferreira 5798 (INPA, K, NY, US, WAG); entre Vaupés y rio Arary, Cachoeira Uapuhy, 2 Nov 1945 (fL), Froes 21314 (NY); carretera Manaus~Ponta Negra, cerca de Ponta Negra, 6 Abr 1974 (f{D, Gentry & Prance 11218 (MO); Presidente Figueiredo, Campina das Pedras, Jun 1985 (f)), Huber & Texeira 10679 (NY, WAG); rio Uaupés, camino a Cova do Diabo, 18 Nov 1997 (fr), Kawasaki 199 (NY); carretera Manaus-Caracarai, Igarapé Lages, 10 May 1974 (fD, Nelson & Lima 21094 (NY), carretera Manaus-ltacoatiara, rio Urubu, 4 Abr 1967 (f1), -rance et al. #749 (INPA, K, NY, US); rio Urubu, 29 May 1968 (IL), Prance et al. 4871(COL, K, NY, US); Manaus-Itacoatiara, LO Jun 1968 (f1, fr), Prance et al. 5125 (NY, US); entre Manaus y Caracarai, 9 May 1974 (f]), Prance et al. 21045 (NY, U, US, Z); camino a Terra Preta, 1 Jul 1975 (f1), Prance 23531 (NY): margen del rio Araca, cerca de Serrinha, 25 Jul 1985 (fr), Prance et al. 29762 (NY, WAG): Manaus, Ponta Negra, 22 Jun 1961 {D, Rodrigues & Lima 2863 INPA, NY): carretera Manaus- ae SJul 1968 ({1), Rodrigues et al. 8503 (INPA, US); Presidente Figueiredo, Represa de Balbina, 4 Jul 1986 (f1), Thomas et al. 5324 (NY, WAG); Manaus, Jul 1900 (f{1D), Ule 5175 (G, MG, foto en INB): entre enn Caracarali, 14 Sep 1979 EL, fr), Zarucchi et al. 2565 (MO, NY); E de Humaita, carretera do Estanho, 27 Sep 1979 (ED, Zarucchi et al. 2592 (INB, MO, NY); S de Borba, 22 Jun 1983 (fl), Zarucchi et cf pe CINPA, K, NY); Axinim, rio ener 7 Jul 1983 (ED, eke . (NY); Presidente ele carretera Manaus-Caracarai, 9 Ago 1983 (fl, fr), Zarucchi et al. 3 Y).Mato G Reserve do Cabagal y Chapada dos Parecis, 26 1995 D, Hatsch bac . et ac Bente US). Para: ena carretera Santarém-Cuiaba, Serra do Cachimbo, 25 Abr 1983 (f1), es etal. 935 (INPA, NY); Alto Tapajos, rio Cururt, Il Feb 1974 (fl, fr), Anderson 10768 (COL, K, MO, NY, US, Z); Oriximina, rio Mapuera, 30 Jun 1980 (£D, Davidson & Martinelli =u (NY); Faro, 21 Ago 1907 (1, fr), Ducke 8434 (G, foto F neg. 26809, MG, foto F neg. 45951, P. US): Faro, Ago 1907 (fl), Ducke 8695 (U); Faro, 17 Ene 1910 CEL, fr), Ducke 10477 (MG, foto en INB); c le Infiry, 12 Feb 1910 (f1), Ducke 10686 (G, foto F neg. 26869, MG, foto en INB); Ariramba, 4 Dic 101 10 GD, Ducke 11343 (MG, foto en INB): Trombetas, Mariapixy, 17 Dic 1910 ({)), Ducke 11943 (MG, oo en oe Itapecuru, 30 Jul 1912 (FD, Ducke 12090 (MG, foto en INB); Tapajos, rio Cururt, 25 Jul 1959 (f1, fr), Eeler 1034 (MG, NY), 28 Ene 1960 (f1, fr), Egler 1188 (MG, NY), Oriximina, rio Mapuera, campina de Trés Thlas, 25 Nov 1987 (fr), Farney et al. 2008 (NY); Oriximina, rio Mapuera, 30 Jun 1980 ({1.fr), Ferreira et al. 1202 (INPA, MO, NY, US, WAG): Oriximina, rio Mapuera, 19 Ago 1986 ({1), Ferreira et al. 7802 (INPA, K, MO, NY, US, WAG); cuenca del ~~ C5 ~ MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2069 rio Trombetas, rio Mapueira, 6 Jun 1974 ({1), Campell et al. 22535 (INPA, K, MO, NY, U, US, Z): Infiry, 12 Feb 1910 (f{D, Ducke s.n. (EF MG); Serra do Cachimbo, 15 Dic 1956 ()), Pires et al. 6289 (NY); entre a Santarém, ase 1977 (ED, Prance et al. 25006 (INB, INPA, K, NY, US). Rond6énia: Costa Marques, 5 Mar 1987 (fl), Nee 34491 (NY) BOLIVIA. La Paz: cae de, Luisita, 9 Feb 1988 (fl), Haase fee Santa Cruz: Velasco, entre Florida y Bella Vista, 4 Nov 1994 (fl), Guillén et al. 2545 (INB, MO, USZ); Velasco, Localidad El Refugio, 18 Ene 1995 (f1, fr), Guillén & Roca 2853 (INB, MO, USZ): ee parque nacional os Kempff Mer- cado, Pampa Grande de Bella Vista, 11 Ago 1995 (£1), Guillén et al. 3908 (INB, MO, U 4. near aad nse (cares Woodson, Ann. Missouri Bot. Gard. 19:51-52. 1932. (Fi RASIL. AMAZONAS: Cano Pimicheiro, Jun 1854 (fD, Spruce 3718 (HOLOTIPO: K; ISOTIPOS: P ae F neg. 38740), W). Ga actophora petiolata Markgr., Notizbl. Bot. Gart. Berlin-Dahlem 14:129. 1938, syn. nov. TIPO. RASIL. AMAZONAS: sae uruicuriary y Negro, 26 Feb 1936 (fl, Ducke 30109 (HOLOTIPO: RB; ISOTIPOS: G, K, MG, P, U, U Arbusto erecto, 0.5-2 m de altura; tallos aplanados en ramitas jovenes, teretes a subteretes cuando viejos, sdlidos, usualmente glabros; coléteres interpeciolares inconspicuos, menos de 1 mm de largo. Hojas con el peciolo de 1-5 mm de largo; lamina 2 DOD 13-2 6-2 Oem sCHDU casaaneosamentc: elipticas, raramente I te, coriaceas, cuneadas ba ealinenie glabrasa g ambas superticies, usualmente revolutas marginalmente, la venacion secundaria levemente impresa en ambas caras 0 inconspicua, las venas terciarias usualmente no impresas. Inflorescencia ter- minal a subterminal, diminutamente puberulenta, con unos pocos y esparcidos pelos glandulares, con 5a 11 flores, pedtnculo 7-42 mm de largo, pedicelos 7- 10 mm de largo, bracteas 1-1.5 x 0.5-0.9 mm, escariosas; sépalos 4-6x 1-1.5 mm, muy angostamente ovados a linear-ovados, acuminados, escariosos, esparcida a moderadamente glandular puberulentos, algunas veces algo glabrescentes cuando viejos; corola rosado palido o blanco-rosadas, el ave tee basalmente, con varios pelos glandulares exteriormente, algunas entes cuando viejas, el apice del boton floral angostamente agudo; parte inferi ior del tubo 5.5- 7 x 15-2 mm, la parte superior 15-16 mm de largo, angostamente cénica, 5-7 mm en diametro en el orificio, l6bulos 11-17 x 7-10 mm, obovados; anteras 4-5 mm de largo, glabras, las bases estériles angostamente agudas, cabeza estigmatica 1.1-1.4 mm de largo; ovario 2.1-2.5 mm de largo, glabro; nectario 0.8-1 mm de largo, levemente pentalobulado. Foliculos 7-12 x 0.2-0.3 cm, continuos, glabros; semillas 6.5-7.5 mm de largo, glabras, coma 17-23 mm de largo, crema. Distribucion, habitat y ecologia.—Restringida a Venezuela y el N de Brasil, donde crece en vegetacion de sabanas y bosques arbustivos enanos en zonas de arena blanca, en elevaciones de 100-200 m. Especimenes con flores han sido recolectados entre Enero y Junio y en Septiembre. Especimenes con frutos han sido recolectados en Septiembre y Octubre. Galactophora pulchella se puede confundir con G. pumila, pero se separa ovado- elipticas, obtusasa 1A} As 2070 BRIT.ORG/SIDA 21(4 2.5 mm 2.5mm oa 4 mm Sem fem 3mm Fic. 7. Galactophora pulchella (Froes 12558, INB). A. Ramita con flores y frutos. B. Sépalo, vista adaxial. C. Antera, vista dorsal. D. Cabeza estigmatica. E. Nectario y ovario. F. Semilla. MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2071 por sus inflorescencias con mucho mas flores (5-11 vs. 1-2), con los pedunculos presentes y conspicuos, usualmente de mas de 7 mm de largo (vs. ausente 0 in- ferior a 2 mm de largo). Las diferencias entre las colecciones tipo de Galactophora pulchella y G. petiolata son minimas y la nica diferencia permisible son las hojas mas angostas en el tipo de la ultima especie, siendo concordantes en el resto de los caracteres mortfoldgicos. s. VENEZUELA. Amazonas: Atures, Cano Ucata, aluente del rio Orinoco, 20 ‘ids 1992 (fl), Berry et al. 5155 (INB, MO); Atabapo, Cucurital de Caname, Cano Caname, 30 Abr-1 May 1979 ({1), Davidse et ae 16996 (INB, MO); Rio Negro, rio Siapa, ; = 1981 (LD, Huber & Medina 5765 (MO, MYENY, U, VEN); Alto rio Orinoco, base NO del Cerro Yapacana, 16 Mar 1953 ({D, Maguire & Wurdach 34481L(NY); sees rio Pacimoni, 7 Feb 1954 (fl), Maguire et al. 37595 (NY); Casiquiare, Cano Hechimoni, rio Siapa, 9 Feb 1954 ({1), Maguire et al. 37609 (MO, NY, US); rios Pacimoni-Yatua, Casiquiare, Piedra Arauicaua, 28 Sep 1957 (fr), Maguire et al. 41631 (K, NY, US). BRASIL. Amer ctes: Dies iNEgrOy. Vaupés, Serra do Tucano, 10 May 1942 ({D, Froes 12558 (F, NY); Igarapé Tibuiari, Vaupés, 22 Nov 1987 (fl), Kawasaki 247 WAG 5. Galactophora pumila Monachino, Mem. New York Bot. Gard. 10:126-127, fig. 16 a-b. 1958. (Fig. 8). Tivo: VENEZUELA. AMAZONAS: Rio Guainia, Sabana el Venado, banco del Cano Pimichin, 14 Abr 1953 (FD), Maguire & Wurdack 35563 (HOLOTIPO: NY; ISOTIPO: Hierba erecta hasta 0.3 m de altura; tallos aplanados y algunas veces levemente alados en ramitas jOvenes, teretes a subteretes cuando viejas, sdlidos, muy esparcida e inconspicuamente puberulentos, glabrescentes con la edad; coléteres interpeciolares inconspicuos, menos de | mm de largo. Hojas con el peciolo 0.5- 1.5mm de largo; lamina (1.7-)2.1-3.8 (0.6-)0.8-L7 cm, angostamente elipticas, obtusas “ peponaeacs® apo algunas veces muy inconspicuamente adas a ebtieas basalmente, coriaceas, glabras, algunas veces con puntuaci te, revolutas marginalmente, la venacion secundaria leer ee impresa adaxialmente a mas comunmente inconspicua, venas terciarias usualmente no impresas. Inflorescencia terminal, glabrescente, con 1 a 2 flores, pedunculo inconspicuo 0 ausente, pedicelos 8- 13.5mm de largo, bracteas menos de 1 mm, escariosas, sépalos 8-1] x 14-2 mm, angostamente ovados a linear-ovados, acuminados, escariosos, glabrescentes; corola purpura, rosado-ptrpura a rosada, con unos escasos pelos glandulares antes de la antesis, usualmente glabrescentes cuando viejas, el apice del boton flora agudo; parte inferior del tubo 10-11 x 15-2 mm, la parte superior 23-31 te conica, 14-16 mm en diametro en el orificio; lobulos 14- 19 x 8- 16 mim, abovedee, anteras 4.8-5.2 mm de largo, glabras, las bases estéril das, cabeza estigmatica 1.9-2.2 de largo; ovario 1.6-2.1 mm de largo, glabro, nectario 0.4-0.5 mm de largo, usualmente muy leve e irregularmente lobulado. Foliculos 85-15 x 0.2-0.3 cm, continuos, glabros; semillas desconocidas. mm de largo, cénicaa 2072 BRIT.ORG/SIDA 21(4) 6mm =< > Imm 4cm 4cm 3mm 1 mm Fic. 8. Eeaalctophiora pane (A-F de ee INB; . i _— artige sis on. A. ses B. Sépalo, vista adaxial, t d ra, vista dorsal. E. Cabez estigmatica. F. Nectario y ovario.G. Foliculos. MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2073 Distribucion, habitat y ecologia.—Conocida en las partes bajas de la cuenca amazonica en Colombia y Venezuela, donde crece en sabanas y vegetacion arbustiva asociada a afloramientos rocosos en elevaciones de 100-400 m. Especimenes con flores han sido recolectados en Abril, Junio, Agosto y Noviembre. Galactophora pumila es una especie muy distintiva que se reconoce con facilidad por su pequefio tamano, raramente excediendo los 30 cm de altura, hojas angostas y pequenas, in reducidasa1 62 flores, y corolas con los lobulos relativamente pequenos, raramente excediendo los 19 mm de largo. Especimenes examinados. COLOMBIA. Guainia: Puerto Inirida, 14 Ago 1975 (f]), Garcia-Barriga 20841 (COL, F, US); Puerto Huesito, sabanas del Alto de La Cruz, 18 Ago 1975 (fr), Garcia-Barriga 20893 (COL, US); Cacagual, rio Atabapo, 13 Set 1957 (fl), Maguire et al. 41434 (COL, NY, US); Puerto Inirida Cacahual, rio ne 25 Nov 1998 (f1, fr), Man 659 (COAH). VENE . Amazonas: Santa Cruz, rio Atabapo, 9 Set 1960 (fl), Foldats 3824 (NY, VEN): Atabapo, mci tee de Canaripo, bajo rio ae 30 May 1978 (£D, Huber 1934 (NY), Atabapo, Laguna de Yagua, 24 Ago 1978 (fl), Huber 2559 (NY); Casiquiare, sabana al O de Pimichin, 25 Ago 1978 (fl), Huber 2594 (INB, MO, MYE NY, VEN); Atabapo, Guarinuma, rio Atabapo, 25 Ago 1978 (£D, Huber 2664 (NY); Atabapo, base del Cerro Cucurito, 8 Dic 1978 (fD, Huber & Tillet 2953 (NY); rio Atabapo, al O de Cacagual, 19 Nov 1953 (f{D, Maguire et al. 36274 (NY); rio Guainia, Sabana El Venado, 23 Nov 1953 (fr), Maguire et al. 36358 (NY, US); rio Guainia, cano Pimichin, Sabana El Venado, 10 Oct 1957 (fl), Maguire et al. 41809 (NY); rio Atabapo, Cano Cumare, San Fernando de Atal ape ns 1959 (f1), Wurdack & Adderley 42773 (G, K, NY, U, US); Sabana Manacal, rio Atabapo 12 Jun 1959 (fl, fr), Wurdack & Adderley 42951 (NY, W). ou. oo 6. Galactophora schomburgkiana Woodson, Ann. Missouri Bot. Gard. 19:50-51. 1932. (Fig. 9). Tipo: GUYANA: Datos perdidos (f1, fr), Schomburgk s.n. (HOLOTIPO: K). Galactophora schomburgkiana var. megaphylla Monac., Mem. New York Bot. Gard. 10:127, 129. 1958. Tipo. VENEZUELA. AMAZONAS: Cerro Duida, rio Cunucunuma, 22 Nov 1958 (fr), Magu- ire et al. 29706 (HOLOTIPO: NY, ISOTIPOS: F (foto F neg. 51104), US). Hierba erecta, 0.5-2 m de altura; los tallos teretes a subteretes, algo aplanados en ramitas jovenes, sdlidos, diminuta y moderada a esparcidamente glandular- puberulentos, los pelos glandulares negros, glabrescentes con la edad; coléteres interpeciolares inconspicuos, ca. | mm de largo o menos. Hojas con el peciolo 2- 7(-9) mm de largo; lamina (3.5-)4-8.5 _ 1.9-)2.8-5 cm, ovadas, angostamente elipticas a ovado-elipticas, obtusas a redondeado-mucronuladas apicalmente, cordadas a subcordadas basalmente, coriaceas, esparcida y diminutamente glandular- puberulentas en ambas caras, algo revolutas marginalmente, las venas terciarias y secundarias cops yevanicnte cee en ambas caras. Inflorescencia terminal r-puberulenta, con 3a 5 flores, pedtinculo 1.4-50 mm de largo, pedicelos 8-15 mm de largo, bracteas 1-3_ 0.5- 1 mm, escariosas; sépalos 5-11(-12.5) _ 2-2.5 mm, angostamente ovados a angostamente ovado-elipticos, acuminados, escariosos a subfoliaceos, muy diminuta y esparcidamente glandular-puberulentos, algunas veces algo glabrescentes cuando viejos; corola purpura, rosado-ptrpura a rojizo-rosado, 2074 BRIT.ORG/SIDA 21(4) 5mm 6mm 4cem 1 3 2mm A “ {cm Fic.9. SUC aoa 18557,M0).A.Ramit fl f vista dor Nec y B. Sépalo, vista adaxial. C. Antera, tario y ovario. F. Semilla. MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2075 muy esparcidamente glandular puberulentas exteriormente, algunas veces glabrescentes cuando viejas, el apice del botén floral agudo; parte inferior del tubo 19-31 x 3-3.5 mm, la parte superior 25-42 mm de largo, conica, (18-)23- 32 mmen diametro en el orificio; lobulos 23-30 x 22-32 mm, obovados; anteras 7.2-8 mm de largo, glabras, las bases estériles muy cortamente acuminadas, cabeza estigmatica 1.9-2.2 mm de largo; ovario 2.8-3.1 mm de largo, con varios pelos glandulares; nectario 1-1.2 mm de largo, irregularmente pentalobulado. Foliculos 9-14 x 0.5 cm, continuos, diminuta y densa a moderadamente glan- ular-puberulentos; semillas 5-6.5 mm de largo, glabras, coma 20-25 mm de largo, canela o amarillo-canela. Distribucion, hdbitat y ecologia.—bosques enanos y ventosos, areas de vegetacion en lajas graniticas, sabanas y bosques enanos y mezclados con pastizales en elevaciones de 750-2000 m. Floracion ocurre de Octubre a Junio. Especimenes con frutos se han recolectado entre Marzo y Junio, Noviembre y Diciembre. Galactophora schomburgkiana puede confundirse con G. crassifolia, pero se puede separar por sus hojas cortamente pecioladas, con peciolos de 2-4.5 (-6.5) mm de largo (vs. O-L5 mm de longitud), hojas con la venacion terciaria usualmente impresa en ambas caras (vs. no impresa) y sépalos mas cortos, de 5-12.5 mm de largo (vs. 15-25 mm). La variacion intraespecifica extrema e intergradada en el tamano de las hojas presentes en algunos miembros de las Apocynoideae, no permite reconocer la segregacion de variedades basado en ese variable caracter, maxime si no existen en forma adicional, caracteres morfologicos discontinuos para separarlas. Por lo tanto, G. schomburgkiana var. megaphylla es reducida a la sinonimia. Especimenes examinados. VENEZUELA. Amazonas: Cerro de la Neblina, NNO de Pico Phelps, 12 Feb 1985 (f1), Boom & Weitzman 5757 (INB, MO, NY); Serrania Part, rio Part, 31 Ene 1951 (fl), Cowan & Wurdack 31085 (NY), 2 Feb se oe Conan & einai k aie. 4 Feb 1951 £1), Cowan & Wurdack 31203 (E NY, US); Atures tnia Piaroa, 6 Nov 1984 (fD), Guanchez & Melgueiro 3296 (WAG); Atabapo, cerca de Salto Lo os iene tributario de rio Iguapo, 11 Mar 1985 (f1, fr), Liesner. uae rio Negron cima del Cerro Aracamuni, 25 Oct 1987 (f1, fr), Liesner & Carnevali = 22411(MO). Boli le Auyantepui Sep 1937 (L 1), Cardona 112 (US), Piar, Auyan- tepui, rio Churum, 27 Mar 1987 (£D), Del ascio 13199 (MO, VEN): Gran Sabana, 28 Mar-12 Abr 1988 ({D, Herndndez et al. 1(1), Hernandez et al. 24 (P); Piar, cima del Auyan-tepui, rio Churum, 30 Mar 1987 (fl, fr), Holst 3807 (MO, NY, U, US); Piar- oe area del rio Uaiparu, NNO de Ikabaru, 18 Feb 1986 (fr), Huber & Fernandez 11338 (MO, NY, WAG); E de El Paujil, O de Santa Elena, rio Las Ahallas, 30 Oct 1985 (fl), Liesner 19167 (MO); S de El Paujil, Moricahl, 8 Nov 1985 (£D, Liesner 1971] (MO); rio Kamarang, NO de San Rafael, 26 Mar 1952 (fl), Maguire 33622 (NY, US); Kavanayén, 4 Abr 1952 (f1, r), Maguire 33745 (NY); falda SO del Ptari-tepui, 18 Dic 1952 ({D, Maguire & Wurdack 33822 (FE NY), 17 Dic 1952 (fl), Maguire & Wurdack 33855 (NY); laldas SE del Ptari-tepui, 10 Nov 1944 ({]), Steyermark 59975 (F MO), base del Carrao-tepui, 4 Dic 1944 (f1, fr), Steyermark 60845 (F), Auyan tepui, rio Churtn, 4 May 1964 (f1), Steyermark 93359 (NY, , PUS, VEN), 12 May 1964 (f1, fr), Steyermark 93755 (FN EN); Auyan-tepui, Guayaraca, 18 May 1964 ({[D, Steyermark 94184 (NY); quebrada El Cajon, E de Icabarti, 18 Dic 1978 (fl), Steyermark 117817 (MO, VEN); Auyan-tepui, Dic 1937-Ene 1938 (£1), Tate 1134 (NY) 2076 BRIT.ORG/SIDA 21(4) GUYANA. Region Cuyuni-Mazaruni, Holitipu, 7 Feb 1996 ({1, fr), Clarke 1040(P. US): Kavanayen, 28 May 1946 (£1, fr), Lasser hg laminas], NY, VEN), montanas Pakaraima, cerro Aymatoi, 17 Oct L981 (EI, fr), Mass et al. 5825 (U, WAG); Cuyuni-Mazaruni, al O de cima del Holitipu, 31 May 1990 (1, fr), McDowell & Gopaul 3015 (C AY, F, RP MO, NY, P, U, US). APENDICE 1: LISTA NUMERICA DE TAXAS ACEPTADOS Galactophora angustifolia J.F. Morales Galactophora pulchella Woodson Galactophora colellana Morillo Galactophora pumila Monachino Galactophora crassifolia (MUll.Arg.) Woodson Galactophora schomburgkiana Woodson APENDICE 2: INDICE A NOMBRES EN EL TRATAMIENTO SISTEMATICO Amblyanthera crassifolia (MUull.Arg.) Woodson crassifolia Mull.Arg. (= G. crassifolia) magnifica Woodson (= G. crassifolia) Dipladenia petiolata Markgraf (= G. pulchella) calycina Huber ex Ducke (= G. crassifolia) pulchella Woodson Galactophora pumila Monachino angustifolia J.F. Morales schomburgkiana Woodson calycina (Huber ex Ducke) Woodson (= G. —Rhodocalyx crassifolia) crassifolius (Mull.Arg.) Miers (= G. calycina) colellana Morillo APENDICE 3: INDICE A EXSICADOS Allen, P., 3163 (3). Ducke, A., 5.n. (3); 8434 (3); 8695 (3); 10477 (3); Amaral, |. et al., 935 (3). 10686 (3); 11197 (3); 11343 (3); 11943 (3); Anderson, W., 10768 (3). 12090 (3); 30109 (4). Arbelaez, M. 426 (3). Duivenvoorden, J. et al., 2743 (3). Arbelaez, M.& J. Matapi, 347 (3). Egler, W., 1034 (3); 1188 (3). Arbelaez, M.& F. Sueroque, 314 (3). Farney, C., 1723 (2); 2008 (3). Arbelaez, M. et al., 626 (3). Fernandez-Pérez, A., 20121 (3). Barbosa, C.& J. Rueda, 8063 (3). Ferreira, C.,5798 (3). Berry, P. et al.,5155 (4); 5197 (3); 5587 (3); 5709 Ferreira, C. et al., 1202 (3); 7802 (3). (3). Foldats, E., 3804 (3); 3824 (5). Boom, B.& A.Weitzman, 5757 (6). Forero, E.& M. Pabon, 9815 (3). Cambell, D.et al., 22535 (3). Franco, Pet al., 4264 (3). Cardenas, D.& G. Gangi, 4627 (3). Froes, R., 12558 (4); 21314 (3). Cardenas, D. et al, 4121 (3). Galeano, M. et al., 95 (3); 1135 (3); 1873 (3); 2252 Cardona, F., 112 (6) (3); Castillo, J.& R.Foster,6910 (3 Garcia-Barriga, H., 13752 (3); 20841 (5); 20879 (3); mo Cid, C.A.et al. 475 (3). 20893 (5). Clark, H., 6587 (3). Garcia-Barriga, H.& R. Schultes, 14149 (3). Clarke, D., 1040 (6). Garcia-Barriga, H. Et al., 16037 (3). Coélho, L. et al., 940 (3); 1395 (3), Gentry, A.& G. Prance, 11218 (3). Cortés, R. & J. Rodriguez, 788 (3); 805 (3). Gentry, A.& M. Sanchez, 65166 (3). Cortés, R. et al., 129 (3). Guanchez, F.& E. Melqueiro, 3296 (6). Cowan, C. & J. Wurdack, 31085 (6); 31135 (6); Guillén, R. & V, Roca, 2853 (3). 31 203 (6 ). Guillén, R. et al, 2545 (3); 3908 (3). Davidse, G. et al., 16996 (4). Gutiérrez, G.& R. Schultes 684 (3). Davidson, C.& G. Martinelli, 10606 (3). Haase, R., 827 (3). Delascio, F., 13199 (6). Hatschbach, G. et al., 63903 (3). MORALES, UNA REVISION DEL GENERO GALACTOPHORA Hernandez, S. et al., 1 (6), 24 (6). Hoehne, F,, 1759 (3). Holst, B., 3807 (6). Holt, £.& E. Blake, 716 (3). Huber, O., 1934 (5); 2559 (5); 2594 (5); 2664 (5); 4636 (3). Huber, O. & A. Fernandez, 11338 (6). Huber, O.& E. Medina 5765 (4). Huber, O.& Texeira, 10679 (3). Huber, O.& S.Tillet, 2953 (5). Huber, O.et al., 5658 (3) Humbert, H., 27444 (3). Idrobo, J, 8932 (3); 8965 (3). Idrobo, J. et al, 11266 (3); 11465 (3). Kawasaki, M., 199 (3); 247 (4). Lasser, T., 1806 (6). Liesner,R.,3415 (3); 3903 (3): 16644 (2);18551 (6 19167 (6); 19711 (6); 25878 (2). am Liesner, R. & G. Carnevali, 22403 (3); 22411 (6); 22881 (3). Maas, P. et al., 5825 (6). Madrinan, S.& C. Barbosa, 1057 (3). Madrinan, S. et al., 1109 (3). Maguire, B., 29300 (3); 33622 (6); 33745 (6). Maguire, B. & J. Wurdack, 33822 (6); 33855 (6); 34481 (4); 35563 (5). Maguire, B. et al., 29706 (6); 36274 (5); 36358 (5): 1434 (5); 739 (3); 41809 (5): 36399 (3); 37609 (4); 37595 (4): 4 41631 (4); 41711 (3); 41 41842 (3): 44167 (3). Marin, C.,659 (5). McDowell, T.& D. Gopaul, 3015 (6). Mejia, A., et al., 2678 (3) Mora, L.& T.Van der Hammen, 154 (3). Nee, M., 34491 (3) Nelson, B. & Lima, 21094 (3). Pabon, M.et al., 355 (3);4278 (3). — 2077 Palacios, P.& B. Plazas, 855 (3); 1 Palacios, P.et al., 458 (3) Pires, M.et al.,6289 (3). Plowman, T., 13708 (3) Prance,G.et al.,4749 (3);4871 (3);5145 (3 (3); 23531 (3); 25006 (3); 29762 (3). Restrepo D.& M. Sanchez, 11 (3) Rodrigues, W. & Lima, 2863 (3). Rodrigues, W.et al., 8503 (3). Romero, G.& F.Guanchez, 1629 (3). Romero-Castaneda, R.& E. Melgueiro, 2234 (3). Sastre, C. & H. pe 5190 (3). Schomburgk, R., s.n. (6) Schultes, R., 12 ae Schultes, R. & ae 12368 (3); 14060 (3); 14626 (3); 17228 (3); 17673 (2); 19381 (3); 19713 (3). Schultes, R.& F. Lépez, 9372 (3). Schultes, R. et al., 18003 (3). Silveiro, J., 10 (3) Spruce, R., 3136 (3);3718 (4). Stein, B., 1492 (3). Stergios, B.& G. Aymard, 4266 (3); 7725 (3). Steyermark, J., 57840 (3); 59975 (6); 60845 (6); 3359 (6); 93755 (6); 94184 (6); 117817 (6). Tate, G., 1134 (6). Thomas, W. et al., 5324 (3). Ule,£.,5175 (3). Weber, A.& A. Knob, 1719 (2). Wurdack, J. . L. He 42683 (3); 42773 (5); 168 (3); 1220 (3). 21045 pubes 12968 (3); 17508 (3); — a Velasco, J., - (3). ayos, M.et al.,6319 (1). Zarucchi, J., 1364 (3); 1395 (3); 1407 (3); 2030 (3); 2116 (3); 2213 (3); 2463 (3); 2979 (3). Zarucchi,J.et al., 2565 (3);2592 (3): 2847 (3);3235 (3). AGRADECIMIENTOS Se agradece a los siguientes herbarios por el envio de material en préstamo o por permitir el uso de sus colecciones; ALCB, ASE, BHCB, BM, CAY, CEN, CGE, COAT COLsGV Re Ese. GOAL, @G.G-BOIS: G-De Gi rib, HUA, IN Pacis LPB, MEDEL, MG, MO, MOL, NY, P QCNE, RB, SPE US, USM, USZ, VEN, VIC, W, Z. Se agradece la autorizacion del Missouri Botanical Garden (MO) y a James Solomon, por la fotografia de Galactophora crassifolia aqui utilizada. Un espe- cial reconocimiento al Herbario del Museo Paraense Emilio Goeldi (MG) en Belém, Para, Brasil y en especial a lone Bemerguy, por el envio de fotografias del holotipo y paratipos de Dipladenia calycina. También quiero agradecer a Luci- 2078 BRIT.ORG/SIDA 21(4) lle Allorge, por permitirme el accesoa las colecciones de los herbarios del Museo de Historia Natural de Francia (P), asi como a Bruno Wallnofer (W) en Vienna, Austria y a Paul e Hiltje Maas por las facilidades brindadas para la visita de varios herbarios en Holanda. Otras personas que colaboraron con facilidades en el trabajo 0 logistica en diferentes herbarios en Sur América fueron Julio Betancur (COL), Ricardo Callejas y Javier Francisco Roldan (HUA), Stephan Beck (LPB), Alvaro Cogollo JAUM), Homero Vargas (QCNE), Asuncion Cano (USM) y Alfredo Fuentes (USZ). Asimismo, quiero agradecer la ayuda de Rodolfo Vasquez y Rocio Rojas por facilitar mis visitas a Lima, Peru. a REFERENCIAS De Canpoite, A. 1844. Apocynaceae. En: A de Candolle, ed. Prodromus systematis naturalis regni vegetabilis, Treuttel & Wurtz, Paris. Paris. 8:317-489. Enpress, M.E. y Bruyns P. 2000. A revised classification of the Apocynaceae s.|. Bot. Rev. 66: 1-56 FALten, M.1983.A taxonomic revision of Condylocarpon (Apocynaceae). Ann. Missourl Bot. Gard. 70:149-169. Hansen, B.F. 1985. A monographic revision of Forsteronia (Apocynaceae). Ph. D. disserta- tion, Univ. of South Florida, Tampa. [University Microfilrns International, Ann. Arbor]. Jupp, W., W. Sanoers, y M. Donochue. 1994. Angiosperm family pairs: preliminary phyloge- netic analyses. Harvard. Pap. Bot.5:1-51. LeeuwenperG, AJM. 1994a. A revision of Tabernaemontana Two. The New World species and Stemmadenia.The Royal Botanic Gardens, Kew, Richmond, England LeeuwenserG, A.J.M. 1994b. Taxa de the Apocynaceae above of the genus level. Agric. Univ. Wageningen Pap. 94:45—60. Moritto, G.N. 1995. Galactophora (Apocynaceae), En: J.A.Steyermark et al.,eds. Flora of the Venezuelan Guayana. Timber Press; Missouri Botanical Garden, Portland; St.Louis, U.S.A. 2:498, 501-504. Morates, J.F. 1997. A synopsis of the genus Prestonia (Apocynaceae) section omentosae in Mesoamerica. Novon 7:59-66. Morates, J.F. A synopsis of the genus Mandevilla (Apocynaceae) in Mexico and Central America. Brittonia 50:214—233. Morates, J.F. 1999. A synopsis of the genus Odontadenia (Apocynaceae) en A. J. M. Leeuwenberg (ed.), Series of revisions of Apocynaceae XLV. Bull. Jard. Bot. Nat. Belg. 67:381-477. Morates, J.F. 2002. Studies in Neotropical Apocynaceae Il:A revision of the genus Fernaldia. Rhodora 104:186—200 Morates, J.F. 2003. Studies in Neotropical Apocynaceae Ill: A revision of the genus Secondatia, with discussion of generic classification. Candollea 58:305-319. Morates, J.F.y A. Fuentes. 2004a. Estudios en las Apocynaceae Neotropicales V: una nueva especie, nuevos reportes y nueva sinonimia en las Apocynaceae de Bolivia. Sida 21: 165-174. MORALES, UNA REVISION DEL GENERO GALACTOPHORA 2079 Monrates, J.F. y A. Fuentes. 2004b. Estudios en las Apocynaceae Neotropicales VIII: nuevas especies de Mandevilla (Apocynoideae, Mesechiteae) para Peru y Bolivia, con notas sobre la morfologia floral en corolas infundibuliformes. Candollea 59:167-1 74. Mutter Arcoviensis, J. 1860. Apocynaceae.En:C.F.P. von Martius, ed. Flora Brasiliensis Munchen, Wien, Leipzig. 6(1):1-180. PicHon, M. 1948a. Classification des Apocynacées. |. Carissées et Ambelaniées. Mém. Mus Natl. Hist. Nat., Sér.B., Bot. 24:111-181. Pichon, M. 1948b. Classification des Apocynacées. V. Cerbérroidées. Not. Syst. Paris 13: 212-229. PicHon, M. 194 8c. Classification des Apocynacées. XIX. Le rétinacle des Echitoidées. Bull. Soc. Bot. France 95:211-216. PicHon, M. 1949. Classification des Apocynacées. IX. Rauvolfiées, Alstoniées, Allamandées et Tabernaémontanoidées. Mém. Mus. Natl. Hist. Nat.27:153-251. PicHon, M. 1950a. Classification des Apocynacées. XXV. Echitoidées. Mém. Mus. Natl. Hist. Nat., Sér.B., Bot. 1:1-143. PicHon, M. 1950b. Classification des Apocynacees. XXVIII. Supplément aux Plumérioidées. Mem. Mus. Natl. Hist. Nat., Sér.B., Bot. 1:145-173 ScHUMANN, K.M. 1895. Apocynaceae. In: A. Engler & K. Prantl, eds. Die natutrlichen Pflanzenfamilien Vol 4(2):109-189. SENNBLAD, B. y B. Bremer. 1996. The familial and subfamilial relationships of Apocynacecae and Asclepiadaceae evaluated with rbcL data. Pl. Syst. Evol. 202:153-175. SENNBLAD, B., M.E. ENoress, y B. Bremer. 1998. Morphology and molecular data in phylogenetic fraternity: the tribe Wrightieae (Apocynaceae) revisited. Amer. J. Bot. 85:1 143-1158. Simoes, A., M. Enpress, T. vAN DER Niet, L. KinosHita, y E. Conti. 2004. Tribal and intergeneric rela- tionships of Mesechiteae (Apocynoideae, Apocynaceae): evidence from three noncoding DNA regions and morphology. Amer. J. Bot. 91:1409-1418. Wituiams, J.K. 1998.A revision of Thenardia Kunth (Apocynaceae, Apocynoideae). Lundellia Woopson, R.E. 1932. New or otherwise Noteworthy Apocynaceae of Tropical America II. Ann. Missouri Bot. Gard. 19:45-76. Woopson, R.E. 1933. Studies en the Apocynaceae IV. The American genera of Echitoideae XXVI. Ann. Missouri Bot. Gard. 20:605—/90. Woopson, R.E. 1935. Studies in the Apocynaceae. lV. The American genera of Echitoideae XXVI. Ann. Missouri Bot. Gard. 22:153-306. Woopson, R.E. 1936. Studies in the Apocynaceae. lV. The American genera of Echitoideae. Ann. Missouri Bot. Gard. 23:169-438. 2080 BRIT.ORG/SIDA 21(4) Book REVIEW David More Gllustrations) and Joun Waite (text). 2002. The Hlustrated Encyclo- pedia of Trees. (ISBN 0-88192-520-9, hbk.). Timber Press Inc. 133 $.W. Sec- ond Ave, Suite +50, Portland, OR 97204-3527, US.A. (Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $79.95, 800 pp., water color paintings, 6" X 9" This book is the mnmed ate aed of the 2004 Collins Tree Guide by Owen Johnson (text) and David More 1 the previous Sida issue [21(3):1322]. The illustrations are ex tremely accurate a de ancins aE se is remarkable little duplication of illustrations between the two. My comment about the HarperCollins volume also holds for the Timber Press one: “For those interested in cultivated trees in the U.S.A, this is a prime reference.” Anyone interested in trees will find iti interesting and usetul (and that’s a broad recommendation). t tl leaf of The Illustrated Encyclopedia are good: “Years ago, artist David More ff. seth iueell the ae of observing and meticulously painting the trees found in Britain, France, Ger- lative or introduced, growing wild or cultivated in many, and the Low Countries—common or rare arboreta, ae Sud gardens. Deciduous trees were painted in the dead of winter as well as in full leaf; he also painted precise details of leaves or needles, bark, blossoms, fruit, nuts and cones. This enormous undertaking required years of study, travel, sketching, painting, and repainting until he was satisfied with each leaf on each page. The results are to be found in this extraordinary book. Well over a thousand species are shown together with as many cultivars. John W hite’ s accompanying | and other text describes the native range of each species, the date of its introduction intoc key facts. The trees to be found in British and European landscapes are mostly those to be found in North America as penn surprising, since so many popular trees cultivated in Europe are in fact American natives. Thus, this s will prove as useful and inspiring in the Western Hemi- sphere as in the lands of its aan "Guy Nesom, Botanical Research Institute of Texas, Fort Worth, TX, 76012-4060, U.S.A. SIDA 21(4): 2080. 2005 LECTOTYPIFICATION AND A NEW COMBINATION IN MATELEA (APOCYNACEAE: ASCLEPIADOIDEAE) FOR AN ENDEMIC HISPANIOLAN VINE Alexander Krings Herbarium, Department of Botany North Carolina State University Raleigh, North Carolina 27695-7612, U.S.A. Alexander_Krings@ncsu.edu ABSTRACT A new combination in Matelea is proposed: Matelea domingensis. RESUMEN Se propone una nueva combinction in Mateled: Matelea domingensis. Critical study of West Indian specimens of subtribe Gonolobinae (Apocynaceae: Asclepiadoideae) has resulted in the need for a new combination for an endemic Hispaniolan vine: Matelea domingensis (Alain) Krings, comb. nov. BAsiony: Gonolobus domingensis Alain, Moscosoa 3:46. 1978. TyPE: REPUBLICA DOMINICANA: trepadora de 50-60 cm de largo; flores verde amarillentas; sobre rocas, al pie de un farallon, estribo sur del Isabel de Torres, Puerto Plata, alt. 750 m, 16-17 Aug 1975, Alain & Perfa Liogier 23780 (LECTOTYPE, here designated: JBSD!) The critical character defining placement in Gonolobus Michx.—laminar dor- sal anther appendages (Woodson 1941; Rosatti 1989; Stevens 2001)—is lacking, although mistakenly attributed to the species by Alain Henri Liogier (loc. cit., 1994). When pressed, the apically bilobed, staminal coronal segments (Cs sensu Liede & Kunze 1993; Kunze 1995) of the single prominent open flower of the type specimen were flattened in such a manner to perhaps superficially ap- pear as dorsal anther appendages (Cd sensu Kunze 1995) (Fig. 1, A). However, close scrutiny, as well as study of an additional flower on the type (pressed side- ways), reveals that the ‘appendages are in fact erect staminal coronal segments (Fig. 1, B; Fig. 2). Other characters that have been used to refer taxa to Gonolobus include winged follicles and the absence of glandular hairs (see Woodson 1941). Follicles are unknown for Matelea domingensis, but this character appears to be moot. Few fruit collections were appz ly available to Woodson (1941) and more recent analysis has shown the character not to be useful in generic de- limitation (Krings, unpubl.). Glandular hairs, although thought characteristic SIDA 21(4): 2081-2085. 2005 2082 BRIT.ORG/SIDA 21(4) Fic. 1. Flowers ae the oes of mareles aeninaeists (Alain) Krings (Alain & Perfa Liogier 23780, JBSD): A, Openly pressed flower, g I gments arching p OvEFINE gynostegium; B, sideways pressed flower, showing an apically bilobed t staminal | ).Cs = staminal coronal segment; Pe = petal; Se = sepal. Scale in millimeters. KRINGS MALELCA 2083 Nine \ os re P en Co Cs bie 0 ¢ A ieriea ne: 's (Alain) Krings (based on Alain & Perfa Liogier 23780 /JBSD).C lum:€ staminal coronal segment. of Matelea Aubl. by Woodson (1941), are also without circumscriptional value in the Gonolobus-Matelea question, being present in both the type of Gonolobus Michx. (ie., G. suberosus (L.) R. Br.) and numerous species lacking dorsal anther appendages (Rosatti 1989). From study of Gonolobus species in the West Indies and the southeastern United States (including the type; see Krings & Xiang 2004), it appears that characters useful for the recognition of Gonolobus ss. in- clude the combined presence of dorsal anther appendages and cordate leaf bases, although at least G. pubescens Griseb., G. stellatus Griseb., and G. stape- lioides Desv.ex Ham. have rounded to cuneate leaf bases. A cushion-like gyno- stegial corona of fused staminal and interstaminal segments that is more or less prostrate, rather than erect, is also nearly ubiquitous among West Indian and southeastern United States Gonolobus taxa, including G. suberosus, G. martinicensis Decne, G. stellatus, and G. stephanotrichus Griseb. lt appears ab- sent in G. jamaicensis Rendle, although additional material is needed for fur- ther study. A reticulate corolla, as found in M. domingensis, does not occur in West Indian or Southeast United States taxa bearing dorsal anther appendages (these referable to Gonolobus). Reticulate corollas however, are common in sev- eral West Indian taxa that bear winged follicles but lack dorsal anther append- ages (see Krings 2005a, b). On-going research aims to resolve the relationships among these taxa using molecular data. Recent progress has shown monophyly for the Gonolobinae (Liede-Schumann et al. 2005), however, with the inclusion 2084 BRIT.ORG/SIDA 21(4) of only five Matelea and Gonolobus taxa, intra-subtribal jeans were not a focus of the study and were not resolved. The emergence of ‘Matelea gonocarpa’ (type for Gonolobus) within a clade of other Gonolobus taxa supports prior treat- ment of the taxon in Gonolobus (see Rosatti 1989; Krings & Xiang 2004). Until a better resolution is achieved and rather than maintaining two internally very polymorphic genera, taxa lacking laminar, dorsal anther appendages, includ- ing M. domingensis, appear best treated in Matelea. Some discrepancies exist between the protologue and the type specimen for Matelea domingensis. Two syntypes were cited in the protologue: “Alain & Perfa Liogier 13780 (SDM, NY)”, “A.& P Liogier & N. Melo 23348(SDM).” As cited, “SDM” is not an official Index Herbariorum acronym, rather an abbreviation for the herbarium of the Jardin Botanico Nacional Dr. Rafael M. Moscoso in Santo Domingo, Dominican Republic or JBSD. However, on the sheet at JBSD, the col- lection number is typed as 23780. The rest of the label information on the sheet is consistent with the protologue. A handwritten determination on the label reads: “Gonolobus domingensis Alain, sp. nov.” The collection number “13780” is also cited by Liogier (1994), although the collector recently indicated that it should be 23780 as on the specimen label (Liogier, pers. comm.). Specimens were — requested as well from NY, however, G. domingensis does not appear to be part of their collections. The additional syntype “A.& P. Liogier & N. Melo 23348 (SDM)” was not included ina loan from JBSD and it remains unclear whether it is extant. Considering that no other specimens could be found, Alain & Perfa Liogier 23780 (JBSD) is here designated lectotype for Gonolobus domingensis Alain. ACKNOWLEDGMENTS | thank the curators and staff of the following herbaria for access to their col- lections or loans of specimens: BH, BM, BOLO, BREM, BRIT, C, CGE, DUKE, E, F PLP LAS Pe PG enn, GOR HAC rab, BGT). bs), oily M, MICH, MIN, MISS, MO, NCU, NO, NY, O, OK, OXF, P, PH, RSA, S, TENN, TEX, U, UC, UNA, US, USCH, USE WILLI, WU, Z. Lalso thank the curators and staff of the following herbaria for searching their collections for West Indian Gonolobinae material, although finding no representation: BG, BKL, BR, BUF COLO, CR, FLAS, IA, ISC, LD, MSU, NEU, NSW, UPS, TUR. The thoughtful re- view of a previous version of the manuscript by Alain Liogier and Mary En- dress is gratefully acknowledged. REFERENCES Krincs, A.2005a. Notes on the Matelea bayatensis-correllii-tigrina complex (Apocynaceae - Asclepiadoideae - Gonolobinae) in the Greater Antilles and Bahamas. Sida 21: 1525-1533. KRINGS MATELEA 2085 Krinas, A. 2005b. A new combination in Matelea (Apocynaceae - Asclepiadoideae) for an endemic Jamaican vine. Sida 21:1515-1517. Krines, A.and Q.-Y. Xianc. 2004. The Gonolobus complex (Apocynaceae - Asclepiadoideae) in the southeastern United States. Sida 21:103-116. Kunze, H. 1995. Floral morphology of some Gonolobeae (Asclepiadaceae). Bot. Jahrb. Syst 117:211-238. Liebe, S.and H. Kunze. 1993.A descriptive system for corona analysis in Asclepiadaceae and Periplocaceae. PI. Syst. Evol. 185:275-284. Liebe-SCHUMANN, A. Rapini, D.J.Goyper, and M.W.CHase. 2005. Phylogenetics of the New World subtribes of Asclepiadeae (Apocynaceae — Asclepiadoideae): Metastelmatinae, Oxypetalinae, and Gonolobinae. Syst. Bot. 30:184-195. Liocier, A.H. 1994. La flora de la Espanola.VI.Familia 178.-Asclepiadaceae. Universidad Central del Este 50, ser.27:11-49. Rosati, T.J.1989.The genera of suborder Apocynineae (Apocynaceae and Asclepiadaceae) in the southeastern United States. J. Arnold Arbor. 70:443-514. Stevens, W.D. 2001. Flora de Nicaragua: Asclepiadaceae. Monogr. Syst. Bot. Missouri Bot. Gard. 85:234-270. Woopson, R.E. 1941. The North American Asclepiadaceae. Ann. Missouri Bot. Gard. 28: 193-244. 2086 BRIT.ORG/SIDA 21(4) Book NOTICE Grorcr W. Cox. 1999. Alien species in North America and Hawaii: Impacts on natural ecosystems. (ISBN 1-55963-680-7, pbk.). Island Press, 1718 Connecticut Avenue, N.W,, Suite 300, Washington, DC 20009, U.S.A. (Orders: Univer- sity of Chicago Distribution Center, 110305. Landley Ave., Chicago, IL 60628, U.S.A.; 800-621-2736, custserv@pres.uchicago.edu). $45.00, 387 pp.,6" x 9" itly organized, easily read- able, laden with interesting information and interpretation, a truly remarkable overview. Hopefully, most biologists have known of this book for several years. Tig Introduction 1. The threat of exotics: Biotic pollution 2. North American invaders: the invited and uninvited 3. A brief history of invasions: Human history — an exotic’s perspective Part Il. Regional Perspectives +. The eastern seaboard: Exotics discover America 5. West coast bays and estuaries: swamping the natives 6. Northern temperate lakes: Chaos along the food chain 7, Western rivers and streams: Pollution that won't wash away 8. Eastern forests: The dark side of forest biodiversity 9. Florida and the Gulf lowlands: Hostile ecosystem takeovers 10. Plains and intermontane grasslands: Exotics at home on the range : Western floodplains: Disturbing the disturbance regime The Pacific states: Mediterranean mixing pot 13. Hawaiian Islands: Exotics in the islands of Eden Part IIL. Biotic Perspectives 14. Exotic game and fish: Addiction to game and fish introduction 15. | lomegrow n exotics: Natives out of plac 16. Human domesticates and associates: on best friends and closest associates slays IV. ans: RCTepECHVES 17 ty structure: Biodiver sity bombs 18.1 Exotics and ecosystems impacts: Changing the way nature works 19. Exotics and evolution: Assimilation or conquest? Part V. Policy Perspectives 20. Living with exotics: The ecological economics of exotics 21. Exotics and public policy: Are all exotics undesirable? SIDA 21(4): 2086. 2005 PALAEOANTHELLA HUANGIIGEN. AND SP. NOV, AN EARLY CRETACEOUS FLOWER (ANGIOSPERMAE) IN BURMESE AMBER George Poinar Jr. Kenton L.Chambers Department of Zoology Department of Botany and Plant Pathology Oregon State University Oregon State University Corvallis, Oregon 97331, U.S.A. Corvallis, Oregon 97331, U.S ABSTRACT Palaeoanthella aoanel ar & ay nov, is described from an me) Cretaceous ee flower in Burmese amber. The racterized by small, staminate f S cup-shaped perianth of 8 fused ase ae een in one series with 8 equal eee nee sta- mens ina single whorl, more or less alternating with the tepals, and having 2 lobed, 4-locular an- thers opening by longitudinal slits. Adjacent pollen ani to have originated from the anthers is inaperturate with finely ridged and grooved exines. The fossil shows possible affinities with the eumagnoliid angiosperm family Monimiaceae. Kry worps: Burma, fossil, fossilized resin, Monimiaceae, Myanmar RESUME Paleoanthella huangii gen. et ee nov, est ae a pa tird’une fleur du Ciera aus ieur as l ambie de Birmanie. | staminée - cupule, forme de BoE sane fusionnés en une série simple et elect pee ou moins avec ah deux lobes et quatre ey s'ouvrant par des fentes longitudinales eae Le polenat trouvé a cété des anthéres es provenant de -ci; il est inaperturate, avec des exines finement carénées et rainurées. Ce fossile montre des affinités possibles avec les Angiospermes Eumagnolides de la famille Monimiaceae. INTRODUCTION A new genus and species of angiosperm with possible affinities with the fam- ily Monimiaceae is described from Early Cretaceous Burmese amber. Since the Early Cretaceous was a period of early angiosperm diversification, all speci- mens from this time period are extremely important in establishing a mini- mum age for the appearance of various floral characters. While only a single staminate specimen is available for study, it is well preserved and presents an interesting arrangement of staminal features. MATERIALS AND METHODS The piece of amber containing the flower weighs 2.8 gm and is more or less trapezoidal in outline, with a greatest length of 25 mm, greatest width of 19 mm and greatest depth of 7 mm. The flower is situated about 5 mm under the surface of the amber. Due to the presence of insect fossils adjacent to the flower, SIDA 21(4): 2087-2092. 2005 2088 BRIT.ORG/SIDA 21(4) the amber could not be re-polished further. Examination and photographs were made with a Nikon stereoscopic microscope SMZ-10 R at 80x anda Leica Wild M3Z stereoscopic microscope at +00x. Pollen grains in the amber adjacent to the flower were photographed with a Nikon Optiphot microscope at 600~. Amber from Burma occurs in lignitic seams in sandstone-limestone de- posits in the Hukawng Valley, southwest of Maingkhwan in the state of Kachin (26°20'N, 96°30E). Nuclear magnetic resonance (NMR) spectra of amber samples taken from the same locality as the fossils indicated an araucarian (possibly Agathis) source of the amber (Lambert & Wu, unpublished data, 2002). Palynomorphs from the amber beds where the fossil originated have been as- signed to the Upper Albian of the Early Cretaceous (97-110 million years ago) (Cruickshank & Ko 2003); however, since the amber is secondarily deposited, the age could be older. DESCRIPTION The flower is approximately 1 mm in diameter. The conspicuous anthers are situated in a whorl at the edge of the receptacle. Since the flower is funnel- shaped, it was difficult to obtain a photo from the top with all its features in focus. Pollen grains attached to the anthers and tepals and in the amber adja- cent to the flower indicate that the flower was in anthesis when it entered the resin. The pollen grains illustrated here are considered to have originated from the flower since 1) they are the same size as those on the anthers, 2) they are adjacent to the flower, and 3) a search through the rest of the amber matrix did not reveal any grains similar to those adjacent to the flower. Palaeoanthella Poinar @ Chambers, gen. nov. Type sprcirs: Palacoanthella huangii Poinar S& Chambers, sp. nov. Unisexual; staminate flowers small, actinomorphic, perianth cup-shaped, united, bearing 8 lobes (tepals) arranged in one series; tepals connate below, separate above, low, rounded at tip; receptacle bearing 8 equal, subsessile sta- mens ina single whorl; stamens unappendaged, alternating with tepals; anthers - locular, opening lengthwise by marginal slits; pistillate flowers unknown. — Palaeoanthella huangii Poinar @ Chambers, sp. nov. (Figs. 1-2). Typr: MYANMAR (BURMA): KACHIN: northern Myanmar, Amber mine in the Hukawng Valley, SW of Maingkhwan, (26°20'N, 96°30'E), Jul 2004, Chialang Grand Huang, Burmese-97 (HOLOTYPE: male flower deposited in the collection of Chialang Grand Huang, Edison, New Jersey 08820, ] AL) Single staminate flower: dimensions of 945 um across shorter axis and 1094 jum across longer axis (the difference between the axes is because the tepals on one axis have been eaten by an herbivorous insect); perianth cup-shaped, 34 jum long, externally hispidulous; stamen length, 270-338 um; stamens subsessile, the four visible filaments ranging from 27-40 ym in length; anthers 270-338 «um in length, basifixed, opening by lateral slits; center of receptacle PALAEOANTHELLA HUANGII CRETACEOUS FLOWER IN BURMESE AMBER 2089 Fig. 1. A. Palaeoanthella huangii in Burmese amber. Scale bar = 217 Lim. Arrow shows a complete tepal; arrowhead shows tepal damaged by a micro-herbivore. B. Two pollen grains adjacent to P. huangii. Scale bar = 15 |1m. C. Single pollen grain adjacent to P huangii. Scale bar = 15 tm. 2090 BRIT.ORG/SIDA 21(4) eats a seep Siang > PO ee oP .7 $ehis Sissies? t9°9 ee © + »F Fic 7 Siecckl if] Dal halla b giti ie L Scale bar = 500 Lum. 7 filled with spongy tissue; pollen grains inaperturate, nearly spherical, 17-20 jim in diameter, exine finely ridged and groove Etymology.—Genus name from the Greek “palaios” for ancient, old, the Greek “anthos” for flower and -ella as a diminutive ending for small. Species named after Chialang Grand Huang, who loaned this valuable specimen for study. — DISCUSSION Itis impossible to assign this flower toa present day family with certainty. How- ever, it does possess some characters [small, unisexual, actinomorphic flowers, cup-shaped monochlamydeous perianth, subsessile stamens, reduction in size of tepals; four-locular anthers opening by 2 lateral longitudinal slits (in stami- nate flowers witha relatively open floral cup)] found in the family Monimiaceae (Melchior 1964; Hutchison 1966; Endress 1980; Philipson 1986, 1993). Several POINAR AND CHAMBERS, PALAEOANTHELLA HUANGII, AN EARLY CRETACEOUS FLOWER IN BURMESE AMBER 2091 extant genera of the Monimiaceae have similar features. Mollinedia has uni- sexual flowers with a cup-shaped floral base. While there are numerous sta- mens in this genus, the filaments are very short and the anthers open length- wise by slits (Perkins 1901). Another genus is Kibara with small unisexual flowers in a hemispherical cup with 8 lobes arranged in 4 series, and 4 sta- mens. However the anthers open by a single apical slit (Perkins 1901). No extant members of this family have 8 stamens arranged in a single whorl, with the number of stamens equal to the number of tepals and more or less alternating with the tepals. Pollen of the Monimiaceae varies considerably in size and shape, ranging from spherical to ellipsoidal and from 10 to 50 microns in diameter (Money et al. 1950). Erdtman (1966) describes the grains as usually nonaperturate, 2(-3)- sulcate or oligofor(aminoid)ate, tenui-exinous. The exine can be thick or thin and can appear as granular, finely pitted-reticulate, spinuliferous, ridged or grooved (Money et al. 1950; Sampson 1993). Acolpate pollen with ridged and grooved exines similar to those adjacent to the fossil occurs in the extant genus Tambourissa (Money et al. 1950). It is difficult to identify any defined structures in the center of the flower that might represent stamen appendages, nectaries, or vestigial carpels. The occurrence of the Monimiaceae in the Lower Cretaceous would be con- sistent with the primitive status ol this family as determined by morphologi- cal and molecular findings. Based on their analysis of the plastid matK gene sequences of various angiosperms, Hilu et al. (2003) placed the Monimiaceae, together with the rest of the Laurales, in the informal group eumagnoliids, which together with the Chloranthales and monocots, form a sister group to the eudicots. The separation of Monimiaceae sensu stricto from the related fami- lies Sipat unaceae, Gomot tegaceae,a nd Atherospet mataceae is supported by the molecular studies of Renner (1999) based on data from six plastid genome re- gions. The Monimiaceae occur in warm temperate to tropical areas of the south- ern hemisphere and enter the Eurasian mainland in Malaysia and Thailand. There are no records of this family in Burma today. The chewed tepals on P huangii are evidence of herbaceous insect activity, possibly by a beetle or moth larve a. The tip of one of the anthers also shows bite marks, indicating indiscriminat li flower parts. The disturbance caused by the herbivore ‘could have dislodged the flower and caused it to fallin the resin. Several insects groups appear to be involved in the pollination of members of the Monimiaceae. A species of thrips (Thrips setipennis) was reported as the sole pollinator of Wilkiea huegeliana in an Australian subtropical rainforest, with both male and female flowers serving as brood sites for thrip larvae (Williams et al. 2001). In Ecuador, members of the genus Siparuna are pollinated by gal midges of the family Cecidomyiidae. These insects deposit their eggs in the male flowers where the larvae presumably feed on the tissues (Feil 1992). Both gall — 2092 BRIT.ORG/SIDA 21(4) midges and thrips occur in Burmese amber and representatives of these groups could have pollinated P huangii. A gall midge is preserved near the fossil f lower. Since the amber mines are located on the Burma Plate, which is part of Laurasia (Mitchell 1993), P huangii can be considered of Old World origin. ACKNOWLEDGMENTS The authors thank Chialang Grand Huang for loaning this specimen for study, Marcos Kogan for the use of his Leica Wild microscope, Jean-Francois Voisin for providing the résumé, and Peter K. Endress and Roberta Poinar for helpful comments on earlier versions of this paper. REFERENCES CRUICKSHANK, R.D.and K.Ko.2003. Geology of an amber locality in the Hukawng Valley, north- ern Myanmar. J. Asian Earth Sci. 21:441-455. Enpress, PK. 1980. Ontogeny, function and evolution of extreme floral construction in Monimiaceae. Plant Syst. Evol. 134:79-1 20. Erotman, G. 1966. Pollen morphology and plant taxonomy. Angiosperms. Hafner Publishing Company, New York. Fei, J.P. 1992. Reproductive ecology of dioecious Siparuna (Monimiaceae) in Ecuador: a case of gall midge pollination. J.Linn. Soc. Bot. 110:171-203. Hitu, K.W.et al. 2003. Angiosperm phylogeny based on matk sequence information. Amer. J.Bot. 90:1 758-1776. HUTCHINSON, J. 1966.The genera of flowering plants (Angiospermae). Vol. 1. Dicotyledones, Clarendon Press, Oxford. Metcuior, H. 1964. A.Engler’s Syllabus der Pflanzenfamilien. Vol.2, Angiospermen, Gebruider Borntraeger, Berlin MitcHett, A.H.G. 1993. Cretaceous-Cenozoic tectonic events in the western Myanmar (Burma)-Assam region. J.Geol. Soc. London, 150:1089-1102. Money, L.L., LW. Baitey, and B.G. Swamy. 1950. The morphology and relationships of the Monimiaceae. J. Arnold Arbor. 31:372-404. Perkins, J.and E. Gita. 1901. Monimiaceae. In: A. Engler, ed. Das Pflanzenreich, lV. 101.Verlag von Wilhelm Engelmann, Leipzig. Pp. 1-122. PHiuipson, W.R. 1986. Monimiaceae. Flora Malesiana Part 1, 10:255-326. PHiuipson, W.R. 1993. Monimiaceae. In: K. Kubitzki, J.G. Rohwer, and V. Billrich, eds. The fami- lies and genera of vascular plants. Il. Springer-Verlag, Berlin. Pp.426-437. Renner, S.S. 1999. Circumscription and phylogeny of the Laurales; evidence from molecu- lar and morphological data. Amer. J. Bot. 86:1301-1315. Sampson, F.B. 1993. Pollen morphology of the Amborellaceae and Hortoniaceae (Hortonioideae: Monimiaceae). Grana 32:154-162. Wituams, G.A.,P. Adam, and L.A.Mounp.2001.Thrips (Thysanoptera) pollination in Australian subtropical rainforests, with particular reference to pollination of Wilkiea huegeliana (Monimiaceae).J. Nat. Hist. (London) 35:1-21. A NEW COMBINATION IN STENOTUS (ASTERACEAE) Caleb A.Morse R. fee McGregor Her bar ium Uni nett of Kansas yee West nstant Avenue Lawrence, Kansas 6604 > U.S.A. A new combination is necessitated by the forthcoming treatment of the genus Stenotus Nutt. (Asteraceae) for the Flora of North America. The combination Stenotus lanuginosus (A. Gray) Greene var. andersonii (Rydb.) was attributed to H.M. Hall by Kartesz (1994a, 1994b), and has been perpetuated in other publications (International Plant Names Index 2004; USDA, NRCS 2004). However, Hall’s combinations were published as subspe- cies in the genus Haplopappus Cass., not varieties in Stenotus. Recent treatments, including the one prepared by the present author for a forthcoming volume of the Flora of North America project, have regarded North American sections of Haplopappus sensu Hall as distinct genera. Therefore, a new combination for this well-marked taxon is necessitated. ee boris aie oe Si ok Greene var. andersonii (Rydb.) C.A. Morse, com sie Bull. Torrey Bot. Club 27:615. 1900. Aplopappus an i ne Montana Coll. Agric Sci Sitid Bot. L100. 1905. oplosanery tars andersonii H.M. Hall, Publ. Carnegie Inst. Wash. 389:172. 1928. Haplopappus lanuginosus var. anc Cronquist, Univ. Wash. Publ. Biol. 17(5):219. 1955. Type: U.S.A. MONTANA: Belt Mountains, Anderson 3561 (HOLOTYPE: NY, UC [fragment] ersonil 886, Tonestus linearis A. Nelson & J.F Macbr, Bot. Gaz. 56:478. 1913. TypF: U.S.A. IDAHO: Payette Na- tional Forest, 1912, G.B. Mains J]. V-6 (HOLOTYPE: RMI ISOTYPE: RMD. ACKNOWLEDGMENT I thank Kanchi Gandhi for discussion of this issue. REFERENCES Hatt, H.M. 1928. The genus Haplopappus: A phylogenetic study in the Compositae. Carn- egie Institute of Washington Publication 389. Kartesz, J.T. 1994a. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland (2nd Ed.) Volume 1 — Checklist. Timber Press, Portland. Kartesz, J.T. 1994b. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland (2nd Ed.) Volume 2 - Thesaurus. Timber Press, Portland. THE INTERNATIONAL PLANT Names Index. 2004. Published on the Internet http://www.ipni.org [accessed 2 Dec 2004]. USDA, NRCS. 2004. The PLANTS Database, Version 3.5 (http://plants.usda.gov/). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. [accessed 2 Dec 2005] SIDA 21(4): 2093. 2005 2094 BRIT.ORG/SIDA 21(4) Book REVIEW E.G. Bosrov and N.N. Tzvevev (eds.) STANWYN G. SHETLER (General Scientific ed.). 2004. Flora of the USSR: Alphabetical Indexes to Volumes I-XXX (trans- lated from Russian). (ISBN not given). Smithsonian Institution Libraries, Washington, D.C. $99.50 (hbk.), 241 pp., 2 maps, 6 1/4" x 91/2". “The Alphabetical Indexes are cumulative indexes to all 30 volumes of the F = hs 1@ USSR ae together constitute the thirty-first and final volume of this monumental work. T } (7 mainly of separate indexes to the scientific (Latin) names of (1) the eae and Q) the ae ‘and species in the Flora, but it also includes a list or index of the authors of the Latin names, alphabetized by the standard abbreviations used for them in the text, and summaries of other he used abbreviations” (from the Scientific Editor's Preface). Synonyms for genera are included but not for species. Two maps are at the very back: “Floristic regions of he USSR” a foldout) and “Regions used to indicate general distributions of species in the ‘Flora of the L The whole set (in Russian) was “initic ated under the add vision and chief editorship of Acade- mician V.L. Komarov.” The original put ere issued 1934-1960; English translations appeared 1968-2002, begun by the Israel ae - Scientific Translations and [inished by the Amerind Pub- lishing Co. in New Delhi. The Asteraceae, recognized by some as highly significant, are published as volumes 25 through 30; they are now complemented by a treatment (translated to English) of the family for the “Flora of Russia: The European part and bordering regions” (vols. VIL in 2002 and VIII in 2003). Itisextremely useful to have an English-readable flora for this huge part of the earth. Flora Europed and various detailed treatments for Australia are generally comparable, but these are only now being approached in kind by the Flora of China and Flora of North America (north of Mexico).— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 21(4): 2094, 2005 PHARUS PRIMUNCINATUS (POACAE: PHAROIDEAE: PHAREAE) FROM DOMINICAN AMBER George Poinar Jr. Emmet J. Judziewicz Department of Zoology Department of Biology Oregon State University University of Wisconsin-Stevens Point Corvallis, Oregon 97331, U.S.A. Stevens Point, Wisconsin 54481, U.S.A. poinarg@science.oregonstate.edu emmet.judziewicz@uwsp.edu ABSTRACT A grass ae with a eyes a Saas uncinate floret in Dominican amber is described as the new species Pharus primuncinatus Judz. & Poinar (Poaceae: Pharoideae: arene The fossil can be fern from all extant eae aie of the genus by the small (9.9 mm long), curved female floret; relatively short glumes; and the uncinate portion of the floret Se downwards to the middle or slightly below the middle of the lemma. TI relative may be the Mesoameri- n P. mezii. This is the earliest record of a fossil grass that can be assigned to an extant genus; the earliest record of a member of the basal subfamily Pharoideae; and the only known fossil Pharus spikelet Key worps: fossil grass, Pharus, P. latifolius, P mezii, P primuncinatus, Poaceae, Pharoideae, Phareae, Dominican amber RESUMEN En base a una ia aes fosil con un flésculo cilindrico, sigmoide y uncinado se describe una nueva ecie, Pharus primuncinatus Judz. & Poinar (Poaceae: Pharoideae: Phareae), del ambar de la on lica nn ae Se diferencia de todas las especies del genéro existentes por su espiguilla pistiiade perienas ee mun de lareo) saeeayh ee curvado; glumas pean warne nite cortas; y ha ael apice. | [ es evpdsblameat P. mezii de América Central. me es el a graminea [osil que puede asignarse a un género existente; el registro mas antiguo del cere de la subfamilia Pharoideae; y el unico fésil de una espiguilla de Pharus. INTRODUCTION Grass fossils are rare. Those that have been reported mostly occur in sedimen- tary deposits and their identification at the generic level is problematic. How- ever, amber has remarkable preservative qualities for both plant and animal fos- sils (Poinar 1992) and a number of angiosperm remains identifiable to extant genera occur in Dominican Republic amber (Poinar & Poinar 1999). Domini- can amber originated from resin of the extinct tree, Hymenaea protera Poinar (Leguminosae: Caesalpinioideae) that thrived some 20-40 million years ago in Hispaniola (Poinar 1991; Poinar & Poinar 1999). One especially interesting fos- sil in this amber was a spikelet belonging to the grass genus Pharus P. Browne which was associated with mammalian hair (Poinar @ Columbus 1992). SIDA 21(4): 2095-2103. 2005 2096 BRIT.ORG/SIDA 21(4) Pharus was long placed in the Bambusoideae Judziewicz 1987; Watson & Dallwitz 1992), but recent molecular evidence indicates that it is much more basal than the bamboos and indeed is the second earliest-diverging basal clade in the Poaceae (Clark & Judziewicz 1996; Judziewicz et al. 1999; Grass Phylog- eny Working Group 2001). Daghlian (1981) considered at least 11 reported macrofossils as true fossil grasses but cautioned against assigning fossil pollen grains to the Poaceae since other members of the Poales have similar pollen types. In his review of paleoagrostology, Thomasson (1987) listed 11 published reports of fossil grasses, but determined only three of these to be undoubted representatives of the Poaceae. While the oldest-known unequivocal fossil grass is from the Paleocene / Eocene boundary in Tennessee (Crepet & Feldman 1991), bam busoid-like mono- cots extend back into the Early Cretaceous (Poinar 2004), thus the origin of the family may be quite ancient. The present study describes the fossil female Pharus spikelet reported earlier (Poinar G Columbus 1992). MATERIALS AND METHODS The small triangular piece of amber (7 gm, 37 mm longest length x 25 mm longest width, 15 mm thick) containing the fossil came from La Toca mine, situ- ated between Santiago and Puerto Plata in the Cordillera Septentrional of the northern portion of the Dominican Republic. Dating of Dominican amber is still controversial with the latest proposed age of 20-15 mya based on foramin- ifera Uturralde-Vinent & MacPhee 1996) and the earliest as 45-30 mya based oncoccoliths (Cépek in Schlee 1999). A range of ages for Dominican amber may be likely since the amber fossils are associated with turbiditic sandstones of the Upper Eocene to Lower Miocene Mamey Group (Draper et al. 1994). Domini- can amber is secondarily deposited in sedimentary rocks, which makes a defi- nite age determination difficult (Poinar @ Mastalerz 2000). Observations and photographs were made with a Nikon stereoscopic microscope SMZ-10 R and Nikon Optiphot TM at magnifications up to 600x. RESULTS The following characters of the fossil place it in the genus Pharus: A single- flowered female spikelet with et short, glabrous glumes and an inrolled, cylindrical, tapering, 7-nerved lemma covered with uncinate (hooked) macrohairs. — Order Poales, Family Poaceae, Subfamily Pharoideae (Stapf) L.G. Clark & Judz., Tribe Phareae Stapf Pharus primuncinatus Judz. & Poinar, sp. nov. (Figs. 1-5). Type: DOMINICAN RE- PUBLIC: La Toca amber mine located between Santiago and Puerto Plata in the Cordillera Septentrional of the northern portion of the Dominican Republic. A female spikelet depos- 2097 Fic.1. Snikelet of PF i j in Domini ber. Bar = 860 uum. 2098 BRIT.ORG/SIDA 21(4) Fics. 2-5. PF j D ber (elongate strands in all phot 9 f lian hair) 2. Tip g bearded portion (deli 1 with ) 1 with unci hairs. Bar = 36 jum. 3. Upper portion of | howing range in size of uncinat hairs. Bar = 278 tum. 4, Lower portion of lemma and t gl ( ),Not I pill f f | Bar = 450 Lim 5. Detail of j hai e one strand f lian hair (arrow) enclosed by the plant macrohairs. Bar = 56 Lum. 2099 ited in the George Poinar amber collection (accession # M-]-4) maintained at Oregon State University, Corvallis, OR 97331. Fossil female spikelet 9.9 mm long (without pedicel) and 2.3 mm wide sub- tended by a short pedicel (0.93 mm long), consisting of two glumes anda single floret. First glume 5.2 mm long; second glume 3.3 mm long (the tip bent back and could have been damaged, thus it may have been longer); lemma approxi- mately twice the length of the glumes, somewhat sigmoid, tapered apically into a beak, 7-nerved, with strongly inrolled margins; with a discrete area, begin- ning approximately 2/5 from the base of the lemma and extending all the way to the tip, densely covered with uncinate macrohairs, the hairs extending from the tip 1/3 of the length of the lemma on the ventral side (3.5 mm from apex), but over half the length of the lemma (5.4 mm from the apex) on the dorsal side; macrohairs 0.22-0.56 mm in length; basal portion of the lemma covered with numerous small papillae (Fig. 4). Currently, seven extant species of Pharus are recognized; they range from Mexico and Florida to Argentina and Uruguay. Using the keys provided by Judziewicz (1987, 1991), the fossil falls into a group of three species with curved to sigmoid female lemmas: P latifolius L., P mezii Prodoehl, and P. vittatus Lem. Pharus vittatus has female lemmas 19-26 mm long, much longer than those of the fossil, which hasa mature lemma 9.9 mm long. The extant species P. latifolius and P. mezii (Table 1, Fig. 6) have smaller female spikelets than P vittatus, but the floret (9.9 mm oe ang 2.3mm wide) and first glume (5.2 mm long) of P. primuncinatus are signi ly shorter yet wider than those of P. latifolius|flo- tets (12=)13-17(= 19) m mm long, 1-2.3 mm wide, first glume 9-13 mm long] and P mezii [florets 10.5-13.5 mm long, 0.9-1.8 mm wide, first glume 5.3-7.3(-8) mm long]. Also, the female floret of the fossil is bearded with uncinate macrohairs over the majority of its exposed (ie., not covered by the glumes) surface, while in P latifolius and P mezii the beard is restricted to the apex of the floret (Fig. 6). Etymology.—The specific epithet primuncinatus is from the Latin “first unci- nate one,” alluding to the first fossil record of hooked macrohairs on a spikelet. DISCUSSION Presently, four species of Pharus occur in Hispaniola, including P. lappulaceus AublL, P latifolius, P parvifolius Nash, and P virescens Doell. Pharus mezii, per- haps the closest relative of the fossil species, is a Mesoamerican and northern South American species of seasonally dry forests. The morphological evidence does not exclude the possibility that P. primuncinatus is directly ancestral to both P latifolius and P. me zii. The manner by which the female spikelet arrived in amber has already been cited as the first example of epizoochory in the fossil record (Poinar & Columbus 1992). This condition is obvious from the number of mammalian hairs associated with the spikelet, one of which is still attached to the lemma 2100 BRIT.ORG/SIDA 21(4) Taste 1. Comparison of Pharus primuncinatus to two closely related species. Pharus latifolius Pharus primuncinatus sp nov. Pharus mezii Distribution Widespread in wet Fossil (Eocene to Miocene) Seasonally dry forests, orests in the rom Hispaniola mostly along the Neotropics, including Pacific coast from Hispaniola Mexico to Panama; rare in dry forests of Colombia, Venezuela, and Ecuador; not in the West Indies Female floret (12-)13-17(-19) 99 10.5-13.5 length (mm) Female floret 1-2.3 2.3 0.9-1.8 width (mm) Female floret: 75-12 43 7.5-10 Length/width ratio Female first glume 9-13 5.2 5.3-7.3(-8.0) length (mm) Female first glume: 0.6-0.8(-0.9) ca.0.5 0.5-0.6 Female floret ratio Distribution of Restricted to Found over most of the Restricted to apical uncinate hairs apical 10-20%; exposed surface 15-25%; on the female rarely sparse hairs (33-60%) rarely sparse hairs elsewhere by the uncinate macrohairs (Fig. 5). These hairs were identified as the guard hairs of a carnivore (Poinar & Columbus 1992). There is circumstantial evidence that many species of mammals disperse the adhesive female florets of Pharus species, as attested by such local common names of P. latifolius and P lappulaceus Judziewicz 1987: 296-297, 387-388) as “fruta de perro” [fruit of the dog] (Cuba), “riz chien” [dog rice] (French Guiana), “alcusa” [dog in Quichual] (Ecuador), “barba de puma” and “barba de tigre” [beard of the jaguar] (Peru), “yagua-arroz [jaguar rice] (Paraguay), “barba de paca” [beard of the paca, a large rodent] (Brazil), and “barba de macuco” [beard of the mon- key] (several Brazilian collections). The genus may apparently be dispersed by birds as well; common names include “pega pollo” [stick-to-the-chicken] (Do- minican Republic), and herbarium specimens with feathers adhering to the inflorescence have been observed. The present report is the earliest record of a fossil grass that can be assigned toan extant genus, the earliest undoubted record of a member of the basal sub- family Pharoideae and tribe Phareae, and the only described fossil spikelet of a member of the Pharoideae. In 1986, the second author also examined a plastic 10 mm 2102 BRIT.ORG/SIDA 21(4) cast of a leaf blade of an undoubtedly pharoid grass (presumably Leptaspis R. Br, sister genus of Pharus), from mid-Miocene (12 million years old) volcanic ash northwest of Lake Baringo, Kenya (ca. 1°N, 36°F) kindly provided by Chris- tine Kabuye and Bonnie FE Jacobs; a duplicate cast is on deposit at the Univer- sity of Wisconsin-Madison herbarium (Judziewicz 1987; Clark & Judziewicz 1996). ACKNOWLEDGMENTS The authors thank J. Travis Columbus for originally identifying the fossil grass and lending assistance to this study, and thank him and Lynn G. Clark for their helpful reviews. REFERENCES Ciark, L.G.and EJ. Juoziewicz. 1996. The grass subfamilies Anomochloideae and Pharoideae (Poaceae). Taxon 45:641-645 Crepet, W.L. and G.D. Fetoman. 1991. The earliest remains of grasses in the fossil record. Amer. J. Bot. 78:1010-1014. DacHuan, C.P.1981.A review of the fossil record of Monocotyledons. Bot. Rev.47:517-555. Draper, G., P. Mann P., and J.F. Lewis. 1994. Hispaniola. In: Donovan, S. and T.A. Jackson, eds. Caribbean geology: an introduction. The University of the West Indies Publishers’ As- sociation, Kingston, Jamaica. Pp. 129-150. Grass PHYLOGENY Workinc Group. 2001.Phylogeny and subfamilial classification of the grasses (Poaceae). Ann. Missouri Bot. Gard, 88:373-457. ITURRALDE-ViNENT, M.A. and R.D.E.MacrHee. 1996.Age and paleogeographic origin of Domini- can amber. Science 273:1850-1852. Juoziewicz, E.J. 1987. Taxonomy and morphology of the tribe Phareae (Poaceae: Bambusoideae). Ph.D. dissertation. University of Wisconsin, Madison Jubziewicz, E.J. 1991 [as 1990]. Poaceae (Gramineae). In: Gorts-Van Rijn, A.R.A, ed. Flora of the Guianas. Series A, fascicle 8, Koeltz Scientific Books, Koenigstein. Pp. 1-727. Juoziewicz, E.J., L.G. Clark, X. LONDONO, and M.J. Stern. 1999. American Bamboos. Smithsonian Institution Press, Washington, DC. POINaR Jr., G.O. 1991. Hymenaea protera sp. nov. (Leguminosae: Caesalpinioideae) from Dominican amber has African affinities. Experientia 47:1075-1082 Poinar Jr, G.O. 1992. Life in Amber. Stanford, CA: Stanford Univ. Press. POINAR Jk., G.O. 2004. Programinis burmitis gen. et sp. nov., and P laminatus sp. nov., early Cretaceous grass-like monocots in Burmese amber. Austral. Syst. Bot. 17:497—504. Poinar Jr, G.O. and J.T. Corumsus. 1992. Adhesive grass spikelet with mammalian hair in Dominican amber: First fossil evidence of epizoochory. Experientia 48:906—908. PoInar Jk. G.O.and R. Poinar. 1999. The Amber forest. Princeton University Press, Princeton. Poinar Jr. G.O. and M. Mastacerz. 2000. Taphonomy of fossilized resins: determining the biostratinomy of amber. Acta Geol. Hispanica 35:171-182. 2103 ScHtee, D. 1990. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk., Ser.C, 28:1-100. THOMASSON, J.R.. 1987. Fossil grasses: 1820-1986 and beyond. In: Soderstrom, T.R., K.W. Hilu, C.S.Campbell,and M.E. Barkworth, eds. Grass systematics and evolution. Washington, Smithsonian Institution. Pp. 159-167. Watson, L. and M.F. Datiwitz. 1992. The grass genera of the world. CAB international: Wallingford. 2104 BRIT.ORG/SIDA 21(4) BOOKS RECEIVED Jose M. MANZANARES. 2002. Jewels of the Jungle: Bromeliaceae of Ecuador. Part I: Bromelioideae. (ISBN 9978-42-547-0, hbk.). Imprenta Mariscal, Quito Ecuador. (Orders: Betty Patterson, e-mail bromeliad@airmailLnet). $120.00 plus $5 p&h (USA, please request shipping for overseas), 1-240 pp., color figures and line drawings, 9" x 1] 3/4", Jost M. MAnzz :s with collaboration of W. TILL, E.Goupa, and E. Patterson. 2005. Jewels of the Jungle: Bromeliaceae of Ecuador. Part II: Pitcairnioideae. (ISBN 9978-42-022-4, hbk.). Imprenta Mariscal, Quito Ecuador. (Orders: Betty Patterson, e-mail bromeliad@airmail.net). $120.00 plus $5 p&th (USA, please request shipping for overseas), 241-544 pp., color figures and line draw- ings, 9" x 11 3/4" | lla} According to the author s, volumes Iland IV (The Tillandsioideae) will probably be published in three years. Joe E. Hottoway (edited by AManpA NFiLL). 2005. A Dictionary of Common Wild- flowers of Texas & the Southern Great Plains. (ISBN 0-87505-309-X, pbk.). TCU Press, TCU Box 298300, Fort Worth, Texas 76129, U.S.A., (Orders: wwwors.tcu.edu/). $29.95, 178 pp., b/w line drawings, 7" x 10". STEPHEN FREER (translator). 2003. Linnaeus’ Philosophia Botanica. (ISBN 0-19-856934- 3, pbk.). Oxford University Press. (Orders: Oxford University Press, 2001 Evans Road, Cary, NC 27513, U.S.A., 800-451-7556, 919-677-1303 fax, wwwoup.com). $89.50, 402 pp., b/w figures, 7 1/2" x 9 3/4", Paperback edition, 2005. Linnaeus’ Philosophia Botanica (The Science of Botany) was first published in 1751. SIDA 21(4): 2104. 2005 NOTES ON LIBERTIA (IRIDACEAE: SISYRINCHIEAE) IN SOUTH AMERICA Peter Goldblatt Marcela Celis BA. co Curator of African Botany Instituto de Ciencias Naturales our Botani a arden Universidad Nacional de Colombia PO. AA. 7495 Bogotd St. Louis, Missourt 63 om USA. COLOMBIA peter.goldblatt@mobot.org ymceelisp@.unal.edu.co ABSTRACT Comparison of collections of J Libertia colombiana (Foster 1939) from Colombia, Ecuador, and Peru show that the Bolivian L. } Foster 1946) cannot be upheld as a separate species. We provide an expanded description of L. colombiana and establish its renee across the Andean paramo as far south as Bolivia, noting the first collections for Ecuador and Peru. \ l id | € to the South American species of Libertia, which also occurs in New Guinea, New Fenane: and eastern Australia where there are some six species currently recognized, and possibly three more awaiting descrip- tion, making a total for the genus of 13 species. RESUMEN Mediante comparaciones de ejemplares de Libertia boliviana (Foster 1946) y L. ee (Foster 1939) de Colombia, Ecuador y te se propone ee . boliviana no puede ser especie aparte. P de I i. como una mbiana y determinamos su rango de dissibucion en los paramos andinos hasta le sur de Bolivia, al igual que las primeras colecciones para Ecuador y Pert. Asi mismo, incluimos una clave para las especies suramericanas de Libertia que también se pn ae en Nueva Cree Nuere cane y el este de Australia, en donde se reconocen cerca de seis espec de descripcion para un total de 13 especies para Fi género. A member of tribe Sisyrinchieae of Iridaceae subfamily Iridoideae (Goldblatt 1990), Libertia is one of two genera of the family with species in Australasia and South America, the other being Orthrosanthus (Goldblatt @ Henrich 1987; Goldblatt 1990). Libertia is recognized mainly by the outer tepals being smaller than the inner, and often much smaller, and more or less green while the inner tepals are usually white, but blue in L. sessiliflora. Like other species of Sisyrinchieae, the filaments are partly united and the style divides at the top of the filament column into three slender, diverging branches. A relatively un- specialized member of the tribe, Libertiad appears to have no other synapomor- phies except for a unique basic chromosome number, x = 19 (Goldblatt & Takei 1997). Australasian species include L. paniculata (R. Brown) Sprengel and L. pulchella(R. Brown) Sprengel, in Australia and New Guinea and four currently recognized in New Zealand, L. grandiflora (R. Brown) Sweet, L. ixioides (Foster SIDA 21(4): 2105-2112. 2005 2106 BRIT.ORG/SIDA 21(4) [.) Sprengel, L. micrantha A. Cunningham, L. peregrinans Cockayne et Allan. Three more were recognized for New Zealand by Blanchon (1998) in an unpub- lished Ph. D. thesis, L. cranwellide, L. edgariae and L. mooreae. Of the 14 species of Libertia described from South America (Index Kewensis), only five are gen- erally recognized, three in Chile (Skottsberg 1928, 1953; Munoz 1906; Rodriguez & Marticorena 2000), and one each for Bolivia and Colombia (Foster 1939, 1945). New collections made in the Colombian paramo (Celis 2000) show that Libertia is represented there by one relatively uniform species, L. colombiana R.C. Foster that was described in 1939. The first collections of Libertia from Ecuador and Peru, made in 1998 and 1975 respectively, fall within the range of variation for L. colombiana. The presence of L. colombiana in Ecuador had not been established when Jorgensen and Leon-Yanez (1999) published their check- list of the flora of Ecuador and Libertia was not included in a checklist of the flora of Peru (Brako & Zarucchi 1993). A second species, Libertia boliviana, was distinguished from L. colombiana (and from the closely related Chilean L. tricocca Philippi) by its smaller and fewer flowers and in having the filaments free to the base (rather than partly united) (Foster 1946). New collections from Bolivia show that Foster's distinc- tion was incorrect. The filaments are united in the proximal half in the type (Buchtien 701, GH), and in two additional collections (Table 1). Morever, there is no significant difference in the size and number of the flowers per inflores- cence unit in specimens from Bolivia and those from Colombia and Ecuador. In fact, we can find no character separating L. colombiana from Bolivian collec- tions of Libertia (Table 1). We thus unite L. colombiana and L. boliviana. The poor documentation of L. colombiana from Bolivia (three collections), Ecuador (one collection), and Peru (one collection) may indicate its rarity in these coun- tries or may be due to its inconspicuous flowers. Until recently, there was alsoa paucity of records from Colom bia. now remedied by specialist collecting there. e immediate relationships of Libertia colombiana are evident ly with the Chilean L. tricocca which is broadly similar in general appearance, modest stat- ure, and flowers with long pedicels. Libertia tricocca is readily distinguished by its narrower leaves, mostly 2-3 mm wide, of firm texture with thickened midribs and margins, its normally a smaller stature, seldom exceeding 20 cm, and short rhizome to 2 cm long. Flowers of L. tricocca also have shorter inner tepals, ca. 4.5 mm long, and the filaments united for ca. one fourth their length (versus inner tepals 6.5-7 mm long and filaments united for about half their length in L. colombiana). The remaining species of South American Libertia are the robust, large- flowered plant known as L. chilensis(Molina) Gunckel (also known by the later name L. formosa Graham) and the remarkable L. sessiliflora (Poepp.) Skottsb. (syn. L. caerulescens Kunth & Bouche). This last species has blue flowers borne i 2107 Taste 1. Comparison of quantitative characteristics of Libertia colombiana and L. boliviana. Character L. colombiana L. boliviana Height 20-40 cm 30-39 cm Rhizome length (2)5-18 cm 7.5-9cm Cauline leaves 6.5-28 cm X 3-6 mM 10-26.5 cm X 4-5 mm Flowering stem length 17-34cm 10-15 ¢ Cauline leaves 2.7-11 cm X 1-3 mm 6cm X 2mm Peduncles (of rhipidia) 2-4.5.cm X* 1mm 2-4.5cm X 1mm Rhipidial spathe length Outer 9-13 mm Outer 7-9 mm Inner 6-10 mm Inner 6-7 mm Pedicel length (0.3-)1.2-2 cm 0.7-2.5 cm Flowers ee rhipidium 2-3(-4) 2-3 Color flowers white white Outer nee length 4mm 3.8-4 mm Inner tepal length 7mm 5mm Filament length 1.8-2 mm, united in lower half 2mm, united in lower half Anthers —1.3 mm, subasifixed 1 mm, subasifixed Ovary 1-2.5 X 1.5-2.8 mm 2.3 * 2.2mm Style 0.8 mm Imm Style branch length 2mm 2mm Capsule 4X 4-45 mm 4x 5mm Seed length 1mm 1mm in sessile rhipidia (the cymose inflorescence units of many Iridaceae) on a straight, unbranched flowering stem. We include an emended description of Libertia colombiana below,a key to the South American species, and outline the complex synonymy of the South American species following Rodriguez and Marticorena (2000). Libertia colombiana R.C. Foster, Contr. Gray Herb. 127:44. 1938. (Fig. 1). Tye: OLOMBIA: Risaralda (“Caldas”), rio San aac abajo del Cerro Tatama, 2600-2800 m, 7 Sep 1922, Pennell 10357 (HOLOTYPE: GH} ISOTYPE: Libertia boliviana RC. Foster, Contr. Gray Herb. 1614. 1946, syn. nov, TYPE: BOLIVIA: La Paz, Region Andina, 3200 m, Nov 1910, O. Buchtien 701 GIOLOTYPE: GH). Evergreen, often tufted herb 20-40 cm high, with a creeping rhizome up to 18 cm long. Leaves 6.5-28 cm X 3-6 mm, in two ranks, sword-shaped, apex attenu- ate, the margins ciliate, sometimes conspicuously so toward the apex. Flower- ing stem 10-34 cm long, subterete, usually branched, bearing cauline leaves progressively smaller above, 2.7-11 cm long. Inflorescences rhipidia, terminal on the main and secondary branches, or sessile in axils of branches 2-4.5 cm long, 2-4-flowered; spathes subequal, the outer 7-13 x 0.4-2 mm, slightly longer than the inner (5-10 X 0.4 mm), lanceolate, apex slightly curvate, with mar- ginal cilia conspicuous toward the apex. Flowers on pedicels mostly 12-25 mm 2108 BRIT.ORG/SIDA 21(4) Fic. 1. Libertia colombi R.C. Foster. A. Habit. B. Detail of leaf.C.S {style | hes_D Inner tepal. E. Side view of outer tepal. F. Capsule. G. Seed. (/drabo 3997). 2109 long, well exserted from the spathes, subtended by a bract 1-6 mm long, white; tepals unequal, the outer whorl smaller than the inner, basally connate for 0.3- 0.5 mm, the outer 3.8-4 X 2.2 mm, elliptical-oblong, obtuse, cucullate, vena- tion acrodromus-parallelodromous, with 5 veins conspicuous, the inner 6.5-7 x 3-3.2 mm, spathulate-ovate, obtuse, venation dendroid. Filaments 1.8-2 mm long, partially united in the lower half for 0.8-1 mm; anthers subasifixed, 1-13 mm long. Ovary 1-2.5 mm long x 1.5-2.8 mm diameter, spheroidal to ellipsoid; style dividing just beyond the top of the filament column into three branches, each 2 mm long, terminally stigmatic. Capsules 3-4 x 4-5 mm, subglobose, borne on pedicels 1.5-3 cm long; seeds ca. 36 in each capsule, 1 mm long, rounded, surface rugose, reddish brown. Distribution.—Andean southern America mainly in paramo, 2200-3900 m, fairly common in Colombia in the Vertiente Caucana, Vertiente Magdalenense, and Vertiente Occidental, and evidently local and rare in Ecua- dor, Peru, and Bolivia. Additional specimens exami ined: BOLIVIA. La Paz: provincia Nor ee 4.7 km al NE (abajo) de Chuspipata, 16°17'S - 67°47' W, 2800 m, 11 Nov 1987, seeial aed ee Sud Yungas, 1.3 km al oeste de Unduavi, 16°18’S - 67°55'W, 3400 m, 12 1987, Sol omon, oe 17418 (MO). COLOMBIA. Antioquia: Abriaqui, Parque Nacional Natural Las oe 6°37'2 N - 76° i. W, 29 Abr 1990, Sey neal ai ); Andes, vereda La Siria, 6°37.2 N - 76°18.2 W, 2200 m, 03 Mar 1995, Sanchez, D. 416 DEL). Cauca: Paez, Cordillera central, cabeceras del rio Palo, quebrada del rio Lopez y eae a Duende, 2°57.3'N - 76°9.45'W, 3400-3450 m, 03 Dic 1944, C 18944 (GH, VALLE): Inza, alrededores de la Laguna de Cusiyaco, 2°33'N - 76°12'W, 3017 m, 07 Oct 1951, Idrobo 3991(COL). Quindio: oe Camino finca Servia-Valle Chiquito, 4°13'N - 75°48'W, 2700-3900 m, se 1990, Vélez, M. 2086 (HUQ); Génova, Vereda alto San Juan, finca La Caucasia, paramo, 4°13'N - 75°48'W, 3200-3500 m, . Dic 1995, Vélez, M.6534 (HUQ). Risaralda: Santuario, Cerro Tatama, 5° oe ee W, 3200-3400 m, 08 Sep 1922, Pennell 10474 (GH); Santuario, Vereda Las Colonias, 400 m arriba del campamento, 5°2.6'N - 76°3.4'W, 2910 m, 02 Feb 1983, Torres 1495 (COL). Tolima: Ibagué, Parque Na- cional Natural los Nevados, parte alta del rio Toche, principalmente en la margen derecha del rio, °36'N - 75°23'W, 3200 m, 29 Jun 1985, Barbosa, C. 3556 (FMB); Ibagué, ee ee del Nevado del Tolima, del Rancho hacia la Cueva, 4°36'N - 75°23'W, 2900-3100 m, 08 A Jaramillo 5122 (COL). Valle: Jamundi, Los Farallones de Cali, cerca a las cuevas a los Osos, ao aN - 76°50'W, 3600 m, 26 Ago 1991, Calderon 44A (COL). ECUADOR uragua: Banos Canton, Parque Nacional Llanganates, faldas del Cerro Negro, valle de Los meats cae 78°15'W, 3500 m, 11 Oct 1998, Vargas H. et al. 2738 (MO). PERU. Cuzco: near Machu Picchu, along old Inca path to Cuzco, Dec 1975, Rafinski sn. (KX). REVISED KEY TO LIBERTIA IN SOUTH AMERICA ]. Flowers pale blue, sessile or with un a less than 3 mm long; ovary ca. 4 L. sessiliflora . Flowers white, outer tepals green at ie . ieee (7-)10-25 mm long; ovary 1-2.5 mm long; capsules subglobos 2. Stem more or less straight, with ste a. except ie terminal; flowers 4—- 10 per rhipidium; inner tepals 10 X 6.5 mm; filaments 6 mm long, anthers 2.5 mm long; style branches 5 mm lon L. chilensis 2. Stem flexuose, rhipidia pedicellate except in axils of branches; flowers 2-4 per rm long; capsules ellipsoid, 7— oO pee 2110 BRIT.ORG/SIDA 21(4) rhipidium; inner tepals 4.5-7 X ca. 3 mm; filaments 1.8-2.5 mm long, anthers 1-1.3 mm long; style branches 2(-3) mm lon 3. Plants less than 20 cm high; creeping rhizome up to 2 cm long; leaves 1-3 mm wide, firm-textured with thickened midribs and margins; inner tepals 4.5 mm long; filaments united for ca.one fourth their length; capsules 2 X 3mm L.tricocca 3. Plants 20-40 cm high; creeping rhizome (2-)5-18 cm long; leaves 3-6 mm wide, without thickened midribs and margins; inner tepals 6.5-7 mm long; filaments united for ca. half their length; capsules 4 x 4-5 mm L. colombiana SYNONYMY OF THE SOUTH AMERICAN SPECIES (EXCLUDING LIBERTIA COLOMBIANA) 1. Libertia chilensis (Molina) Gunckel, Rev. Chil. Hist. Nat. 31:87. 1927. Strumaria chilensis Molina, Sagg. Stor. Nat. Chili ed. 2:130. 1810. Type: unknown-—the identity of the basionym is convincingly demonstrated by Gunckel (1927) to be the plant better known as L. formosa R. Grah. based on an analysis of the protologue. Libertia Lunes Grah., Edinb. N. Phil. J. 1833:383. Oct 1833. TyPE: CHILE [as Chili, Cape Horn|: imported by J. Anderson and cultivated first at the Clapton Nursery, London, and then in E ane Scotland; no preserved specimen known. Bielauons aeeite in Edwards's 34) serve to Botanical Register (Lindley 1833) and in Curtis's Botanical identify the species and may be ose as ty ay material as hey were grown toni the same stock originally collected in Chil tivated in London and then in Edinburgh. Accord- ing to Lindley (1833) the plants were collected by J. Anderson on the coast of Chiloe Island. We designate the illustration in Curtis's Botanical Magazine as lectotype. Libertia crassa R. Grah., Edinb. N. Phil. J. 1833:383. Oct. 1833. TYPE: CHILE [as Chili, Cape Horn|: imported by J. Anderson and cultivated first at the Clapton Nursery, London, and then in Edin Libertia elegans Poepp., Fragm. Syn. PL. Chil. 3. 1833. Roterbe elegans (Poepp.) Steud. in Lechler, PL Chil. Exsice. no. 569, comb. inval. non publ. Type: CHILE: near Valdivia, without date, Lechler sn. (SOTYPE: K). Libertia grandiflora Phil, Bot. Zeit. 14:648. 1856, nom. illeg. non L. grandiflora (R. Br) Sweet, Hort Brit. b498. 1826. Type: CHILE: Juan Fernandez, collector and date not known (ISOTYPE: SGO). dsurgh, Scotland; known only from the description. Notes.—We preter not to designate lectotypes for Libertia elegansand L. grandi- flora because there may be additional material unknown to us. Libertia chilensis Klotzsch mss was included by Baker (1877) in the syn- onymy of L. elegans Poepp. and is not a valid name. It therefore does not invali- date the combination L. chilensis (Molina) Gunckel. The plant referred to in the literature as Libertia ixioidesC. Gay (Fl. Chil. 6:31. 1854) is incorrectly attributed to that author. Gay merely called a Chilean plant by this name, citing Forster fil. and Sprengel as authors of the basionym and combination in Libertia respec- tively, ie, L. ixioides (Forster f.) Spreng., which is a New Zealand species. Like- wise, the plant called Libertia ixioides Klatt lin Mart. FL. Bras. 2:530, pl. 68, fig. 2. 1871] was not intended as a new species and L. ixioides Spreng. was cited in the text as the source of the name. This is the same New Zealand species, and was presumably cultivated in Brazil where there are no native species of Libertia. 2111 Libertia macrocarpa Klatt, Linnaea 31:384 (1861-1862) is sometimes cited as a synonym of L. chilensis and a possible type has been located at K. This sheet is annotated “Chile, Valparaiso, cultivated at Hort. Gl.” and the collector is not recorded. The specimen is a New Zealand species of Libertia, perhaps L. ixioides, which it matches in the oboviod capsules. 2. Libertia sessiliflora (Poepp.) Skottsb., Nat. Hist. Juan Fernandez & Easter Is- land 2:778. 1928. Sisyrinchium sessilifl orum Poepp., Notiz. Geb. Natur-Heilk. (ed. L. F Se ae 1829; et Fragm. Syn. PI. 2.1833. Tekel sessiliflora (Poepp.) Kuntze, Revis. Gen. 2:703. 1891. TyPE: CHILE: without precise locality, Sep, Poeppig 283 (HOLOTYPE: B, not seen extant, MO, photo!). Sisyrinchium sessiliflorum Hook. & Arn., Bot. Beechey Voy. 1:47. 1830, nom. illeg. non S. Sen rT BaD Mee ies eae Concepcion, Beechey s.n. (probable HOLOTYPE: K!). il é, Linnaea 19:382. 1847. Type: CHILE: Valparaiso, Lagunillia, cultivated in Berlin, May 1845, without collector (HOLOTYPE: B, not seen but extant; MO, photo!). Note.—It is not clear whether the name Sisyrinchium sessiliflorum J.D. Hook. & Arn., Bot. Beechey Voy. 1:47.1830 (listed as valid, for example, in Index Kewensis) was intended as a new species or merely the use of Poeppig’s epithet, published a year earlier but not cited. While it seems prudent to assume the latter, thus simplifying the nomenclature of this species, the Code of Botanical Nomencla- ture requires citation of an author or the name must be treated as a new species. 3. fwene triccoca Phil, Linnaea 29:63. 1857-1858. Type: CHILE: Valdivia, without date, hilippi 944 (SYNTYPES: B, K!, SGO, not seen; MO, photos!), near Tomé (environs of Concepcion at K), mien date Germain s.n. (possible SYNTYPES: K!, SGO, not seen; MO, photo). ACKNOWLEDGMENTS We are grateful to the Instituto de Ciencias Naturales at the Universidad Nacio- nal de Colombia and The Missouri Botanical Garden, specially Colombia Project and their coordinators Olga Marta Montiel and Rosa Ortiz-Gentry for the given facilities to do this investigation. We thank to Henry Arellano by for his illus- tration, to Rodrigo Bernal for his comments about the synonymies and to the curators and staffs of COL, FMB, GH, HUQ, K, MEDEL, MO and VALLE for al- lowing us to study their collections. REFERENCES Baker, J.G. 1877. Systema Iridearum. J. Linn. Soc. Bot. 16:61-180. Baker, J.G. 1892. Handbook of the lrideae. Saat Bell & pons Soo BLANCHON, D. 1998. The genus Libertia (S tion and evolution. Doctoral Dissertation, School of pecs Sciences, eee of Auckland, New Zealand. Brako, L. and J.L. ZaruccHi. 1993. Catalogue of the flowering plants and gynmosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45 Cetis, M. 2000. lridaceae. In: O. Rangel, ed. Colombia Diversidad Bidtica Ill, La regién de vida 2112 BRIT.ORG/SIDA 21(4) paramuna. Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia, Bogota, Colombia. Pp. 256-260. GraHam, R. 1834. Libertia formosa. Beautiful Libertia. Curtis's Bot. Mag. 61: plate 3294. Gunckel, H. 1927.Notas botanicas. Rev. Chilena Hist. Nat. 31:86-89. Foster, R.C. 1939. Studies in the lridaceae. 1.Contr. Gray Herb. 127:33-48. Foster, R.C. 1946. Studies in the flora of Bolivia. 1. lridaceae. Contr. Gray Herb, 161:3-19. Gotostart, P. 1990. Phylogeny and classification of lridaceae. Ann. Missouri Bot. Gard. 77: 627. Gotostart, P. 1998. lridaceae. In: K. Kubitzki, ed. Families and genera of flowering Plants Springer Verlag, Heidelberg [with contributions from J.C. Manning and P. Rudall]. 2: 295-335. Gotostatt, P. and J.E. HenricH. 1987. A review of the New World species of Orthrosanthus Sweet (lridaceae). Ann. Missouri Bot. Gard. 74:577-582. JorGENSEN, PM. and S.Leon-YANez. 1999.Vascular plants of Ecuador. Missouri Botanical Garden Press, St. Louis, Missouri Linoey, J. 1833. Libertia formosa. Handsome Libertia. Edwards's Bot. Register 6: plate 1630. Ropricuez, R.and C. Marticorena. 2000. Commentarios taxonomicos en lridaceae Chilenas. Gayana Bot. 57(2):169-179. SKOTTSBERG, C, 1928. The natural history of the Juan Fernandez Islands. A supplement to the pteridophytes and phanerogams of Juan Fernandez and Easter Island. Almqvist & Wiksells, Uppsala. ARISTIDAE ELUDENDAE: ARISTIDA HITCHCOCKIANA (POACEAE)—A VALID SPECIES? Kelly W. Allred Range Science eal um Department of A & Range Sciences New Mexico State oe Las Cruces, New Mexico 88003, U.S.A kal ee ABSE RAGE The original descriptions and prologues, type specimens, and available material of Aristida hitchcockiana Henrard and A. appressa Vasey were compared. Based on morphology, distribution, and habitat, there seems to be no reason to recognize A. hitchcockiana as a valid species, and it is synonymized within A. appressa. RESUMEN ] ] 4] ] ] y demas material disponible de Aristida enon and Heneard y A appre ssd Vasey. B Ensancose en la morfologia, la distribucion geogratica, y el habitat, p vara reconocer como especie valida a A. hitchcockiana. Por (seaeee se la sinonimizaa A appressa Aristida hitchcockiana was described in 1927 by J.T. Henrard, in his monumen- tal “A Critical Revision of the Genus Aristida” (Henrard 1926, 1927, 1928). His new species was described and illustrated as having long narrow panicles, sub- equal glumes, the second truncate or emarginate, and unequal awns (Fig. 1). In his earlier work on the North American species of Aristida, A.S. Hitchcock in- cluded the type-to-be of A. hitchcockiana within A. arizonica (Hitchcock 1924). Accordingly, Henrard called attention to the shorter spikelets and awns, and truncate, emarginate glumes of A. hitchcockiana as compared to A. arizonica. The new species was known only from the original description and from the type collection at Las Sedas, Oaxaca, México (Hitchcock 1935), and largely ig- nored or not relevant to subsequent works on Mexican Aristida until Beetle’s Las Gramineas de México (Beetle 1983), in which he recorded the species from seven states in México. 1 do not know to which plants he applied the name, only that the illustration is at variance with both Henrard’s and Beetle’s own de- scriptions, in that it shows an open panicle with spreading branches and pedicels, acuminate glumes, and long equal awns. In these publications and in Henrard’s prodigious “A Monograph of the Genus Aristida” (Henrard 1929, 1932, 1933), A. hitchcockiana was compared most closely to A. arizonica, from which it was distinguished by having longer glumes, lemmas, and awns, but which belied its more obvious similarities. SIDA 21(4): 2113-2118. 2005 2114 BRIT.ORG/SIDA 21(4) Fic. 1. Spikelet, 2nd glume apex, and callus of Aristida hitchcockiana [from H 4 (1929)] A comparison of Aristida hitchcockiana with A. appressa Vasey (Fig. 2), based on original descriptions, type material, and specimens, failed to reveal any morphological characters by which to distinguish them (Table 1). For Henrard, the single most diagnostic feature of A. hitchcockiana was the trun- cate and shortly awned apex (the “curious tip”) of the second glume, clearly illustrated in the original description and relied upon solely and explicitly in the keys of his Monograph (Henrard 1932, p. 237). This feature is seen clearly in the type, but even there, some of the second glumes are acutish and not very truncate. Examination of nearly 100 specimens of A. appressa showed a nearly complete gradation from glumes with truncate and shortly awned apices, througha series with progressively longer awns and more narrow apices, to the other extreme of glumes with long awns, acuminate apices, and noticeable lat- eral setulae. The truncate apex can be easily overlooked or obscured when the glume is folded or rolled. Perhaps this has lead to the characterization of the glumes of A. appressa as being acute or acuminate (Henrard 1932; Beetle 1983), ALLRED, ARISTIDA HITCHCOCKIANA—A VALID SPECIES? 2115 Cc 4 | | ee 4 lt £ Ao akt ft re u 4/1037) Fic.2 F when in fact obtuse to truncate glume apices are frequently encountered in that species as well. In addition, I could find no correlation of this condition (a truncate apex) toany other feature. Both species develop long internodes (longer than the sheaths), a line of hairs across the collar (represented by a rim in older material, or glabrate), noticeably long hairs above the ligule on the upper sur- face of the blade, and lateral awns shorter than the central awn. Henrard (1929) called particular attention to the “very curious crisp pu- bescence” of the culms, panicle branchlets, and lower glumes of Aristida appressa. This is caused by a scaberulous type of pubescence, usually in lines along the ridges or nerves of the afore-named parts. It is well-expressed in the type of A. appressa, but much less so in many other specimens, and can com- monly be absent. This same scaberulous pubescence is found in the type of A. 2116 BRIT.ORG/SIDA 21(4) Taste 1. Comparison of Aristida hitchcockiana and A. appressa. Feature Aristida hitchcockiana, type Aristida appressa Culm height (cm) 75 40-120 Peduncle length an 10-40 Internode pubescence — Middle sheath cna (cm) Sheath pubescence Ligule length (mm) Auricle region Collar Longest blade length (cm) Blade width when flat (mm) Blade involution Blade margin Blade upper pubescence Panicle length (cm) su wer branch length (cm) Pulvini First glume length (mm) First glume apex Second glume length Second glume apex Glume pubescence mm) Lemma length to awns (mm) Lemma vestiture Lemma beak Callus length (mm) Central awn length (mm) Lateral awn length (mm) abou to scaberulous one ciliolate, with hairs to 2.8mm scaberulous in a line eo —N flat, ee to be rolled due a6 hic kened puberulent/scaberulous with scattered long hairs near ligule 17-24 absent 8-10 acute truncate to acute scaberulous on the nerves 11-12 glabrous in the lower part, scaberulous in the upper slightly exceeding the glumes glabrous to puberulent 4-11 glabrous to scaberulous flaring, ciliolate to glabrate, often with hairs to 3 mm glabrous to scaberulous in a convolute when young, flat & curling when oe slightly thicken eee eae with scattered long hairs near ligule 14-30 5-12 absent 5-12 truncate to acuminate 6-12 truncate to aul glabro lous on the nerves 7-13(-16) glabrous in the lower part, scaberulous in upper equal to much exceeding the g 0.6-1 15-22 7-17 hitchcockiana,on culm internodes, branchlets, and the | ower glumes. This fea- ture has the same degree of inconsistency as the truncate glume apex: ranging from noticeably present to absent. Henrard’s holotype came from Las Sedas, Oaxaca, México (approximately 100 km northwest of the city o Pr Oaxaca on highway 131, at N17.3523° W96.9444°), collected by Charles L. Smith sometime in 1894. The village was a former rail- road station, and sits at about 2100 m ina pine-matorral transition zone, a com- mon elevation and habitat for Aristida appressa. Aristida appressa is found in ALLRED, ARISTIDA HITCHCOCKIANA—A VALID SPECIES? 2117 all the surrounding states (Chiapas, Veracruz, Puebla, Guerrero), as well as in Oaxaca itself. The basis for Aristida hitchcockiana seems to be simply one end of a single line of variation. Its recognition is partly an artifact of observation: noticed ina few plants where the glumes are not folded or rolled, but overlooked in other plants where the glume apices are obscured. It lacks any distinctive morphol- ogy, distribution, or habitat. For these reasons, the name A. hitchcockiana Henrard is subsumed without recognition under the older name A. appressa Vasey, as detailed below. Aristida appressa Vasey, Contr. US. Natl. Herb. 1(8):282. 1893. Type: MEXICO. JALIsco: Guadalajara, 1886, E. Palmer s.n. (HOLOTYPE: US-745676}, ISOTYPES: L fragm., W). — hitchcockiana Henrard, Meded. Rijks-Herb. Leiden No. 54A:233-234. 1927. TYPE: XICO. OAXACA: Las Sedas, 1894, CL. Smith 918 (HOLOTYPE: US-991670)). Selected sy ined (of 76 total): COSTA RICA. Guanacaste: 4 km W of the Inter-Ameri- can Hwy on the road to es 320 m, 24 Oct 1968, R. Pohl 11324 (F). EL SALVADOR. Chalatenango: along Hwy 4, 4 km SSE of La Palma, pine forest, 950 m, 1] Jun 1970, R. Pohl 11890 (F). GUATEMALA. Chimaltenagno: near a Pixcayo, oak and ae forest, 1650-188 m, 3 Feb 1939, P. Standley 64492 Huehuetenango: about Laguna de Ocubila, Huehuetenango, 1900 m, 7 Jan 1941, P. Standley 827 5 (F). HONDURAS. Comayagua: half-way cate ee ik and Villa San Antonio, 25 Jan 1936, W. Archer 3841 (US). El Paraiso: grassy pine forest of Cuesta Galeras road to Guinope, 1400 m, 24 Nov 1970, A. Molina R. 25913 (US). Morazan: open hillside, = Clara Creel, Rio Yeguane Valley, 800 m, 17 Dec 1946, L. Williams 11259 (G); region of Las Mesas, steep pine-wooded ee ied m, 14 Oct 1951, J. Swallen 10740 (US). MEXICO. Chiapas: steep rocky slope with Que long Mexican Hwy 190 in the Zinacantan paraje of rae 3500 [t, 17 Aug 1965, D. Bette 11869 (F); 11 km W of Tuxtla Gutierrez in rolling hills, 24 Oct 1973, F Gould 14433 (NMCR). Distrito Federal: Villa Guer- rero, 21 Oct 1960, T. Tateoka 1139 (US). Guerrero: Manchon, 26 Sep 1936, G. Hinton 9593 (MO). Guanajuato: about 6 km east of Guanajuato, rocky soil on an eroded hillside, 17 Oct 1952, FE. Sohns 312 (US); km 40, Guanajuato City-Dolores Hidalgo, 15 Sep 1946, E. pie eee (US); Sta. Cruz de Jventino Rosas rumbo a Guanajuato, | Sep 1981, A. Beetle 7 . México: 1.7 km W of Tejupilco on Mexican Hwy 134 towards Temascaltepec, 1430 m, 6 Oct oo a Peterson 11058 (NMCR). Oaxaca: Las Sedas, in 1894, Chas. L. Smith 918 (US-type); 4.8 km e of Mitla on Mexican Hwy 179 to- wards Ayutla, gentle slopes near small creek and cultivated fields, 1760 m, 14 Sep 1990, P. Peterson 9861 (NMCR); NE of Oaxaca, 13 km N of El Punto on Mexican Hwy 175 and 10.8 km S of Guelatao, 1970 m, 17 Sep 1990, P. Peterson 9936 (NMCR); 151 km SW of Oaxaca on Mexican Hwy 190, oak woods, 1100 m, 15 Oct 1976, J. Brunken 369 (MO); from ip to San Ildefonso de Villa Alta proper, on open and rocky mountain slopes, 29-30 Oct 1944, J. Vera Santos 3543 (US), savanna near Revolucion Mexicana, 800 m, 3 Nov 1981, D. Breedlove ee (G). Puebla: near Cholula, around the Cholula pyra- mid and Church Nuestra Senora de la Remedios, 14 Dec 1972, A. Beetle 2276 (MO). Veracruz: 3 km al N de Chacalapa, 400 m, 26 Sep 1965, J. Rzedowski 21211 (F) ACKNOWLEDGMENTS Many thanks to the curators of E G, MO, US, and WIS, who kindly supplied material for study, and to PM. Peterson and S.L. Hatch for their helpful com- ments as reviewers. David Lee Anderson generously rendered the abstract in Spanish. Thanks to Jesus Valdés-Reyna for additional information about Las Sedas. 2118 BRIT.ORG/SIDA 21(4) REFERENCES Beetle, A.A. 1983.Las Gramineas de México, vol.1.COTECOCA [Comision Técnico Consultiva de Coeficientes de Agostadero], México, D.F., México Henrard, J.7. 1926. A critical revision of the genus Aristida. |. Meded. Rijks-Herb, 54:1-220. Henrard, J.7. 1927. A critical revision of the genus Aristida. Il. Meded. Rijks-Herb. 54A: 221-464. Henrard, J.T. 1928. A critical revision of the genus Aristida. Ill. Meded. Rijks-Herb. 54B: 465-701. Henrard, J.T. 1929. A monograph of the genus Aristida. 1. Meded. Rijks-Herb. 58:1-137. Henrarb, J.T. 1932.A monograph of the genus Aristida. Il. Meded. Rijks-Herb. 58A:157-325. Henrarb J.T. 1933.A monograph of the genus Aristida. Index. Meded. Rijks-Herb. 58B:I—XIl. HitcHcock, A.S. 1924. The North American species of Aristida. U.S. Natl. Herb. 22:517-586. HitcHcock, A.S. 1935. Aristida. North American flora 17:376-406. A LECTOTYPE FOR STACHYS FLORIDANA (LAMIACEAE) John B. Nelson Daniel B. Ward Department of Biological Sciences Department of Botany University of South Carolina University of Florida Columbia, South Carolina 29208, U.S.A. Gainesville Florida 32611, U.S.A. ABSTRACT coy Dy Fa | ] aes Mees | j sy a Ce ] fl a Cc} ] In the absence of f ex Benth. in DC. (1848) from material not clearly cited by Bentham but probably in his possession at time of publication. Key Worps: Stachys, Lamiaceae, lectot ] ABRISS Aufgrund des Fehlens von Originalmaterial wird fur Stachys floridana Shuttlew. ex Benth. in DC. (1848) ein Lektotyp aus Belegen ausgewahlt, die Bentham zwar nicht zitiert hat, die aber wahrscheinlich zur Zeit des Publikation in seinem Besitz waren. A review of the genus Stachys (Lamiaceae) in the southeastern United States (Nelson 1981) left unresolved some difficult but noncritical nomenclatural de- tails. One of the most vexatious of these remaining inquiries has been the search for a suitable type and type locality for a widespread southeastern species, Stachysfloridana. Since no single specimen was designated by its author, George Bentham, which can be interpreted as a holotype, our intent here is to docu- ment the steps taken in this search and its ultimate resolution by choice of an appropriate replacement, a lectotype. Bentham (Prodr. 12:478, 1848) published Stachys floridana using a name coined by Robert J. Shuttleworth for specimens ollected for him by Ferdinand Rugel. See Geiser 1948, for a brief cca of Rugel.) neue published his new Stachys by first indicating the source of the name, “(Shuttlew! pl. Rug mss.).” He then provided a 41-word diagnostic description based upon materials “Ad Tampa Bay Floridae.” As indicated by the further notation “(h. Torr! Rtigell!),” Bentham’s material apparently consisted of two collections, a specimen from the herbarium of John Torrey anda second from Rugel. Bentham concluded his presentation by providing a 21-word supplemental description that applies to a commonly observed, full-sun variant. The following relevant collections have been located, presented here in (ap- parent) order of date of collection: 1. Unknown collector 72: Handwritten label bearing “72” and reading “Stachys floridana Shuttl.,” “Tampa Bay [Hillsborough County] Florida,” and “Torrey 18[?]6.” Specimen seen: K (photo only). The hand is undetermined. SIDA 21(4): 2119-2124. 2005 2120 BRIT.ORG/SIDA 21(4) 2. Rugel s.n.: Printed labels from the Shuttleworth herbarium, reading “Stachys floridana Shuttl. n. sp. Ad margines agrorum, prope Tallahassee [Leon County], Florida, legit Rugel, Mai 1843.” Specimens seen: BASBG, BM, K, MIN, MO, NY, USCH, Z [3]. Since Shuttleworth sold and distributed sets of Rugel’s 1843 collections (Geiser 1948), additional specimens may exist. 3. Rugel 176: Labels headed “From The United States National Herbarium,” bearing the printed words “Florida—F. Rugel: 1842-1849. Ex Herb. Mus. Brit.,” a handwritten “176,” and the plant name. Specimens seen: BM, F MO [2], NY, US. Apparent duplicates of this collection are at FLAS and NCU, but accompanied by labels printed with the words “Ex Herbario Musei Brittanici,” and modern typescript indicating the specimen to be Rugel 176, from Lake Monroe [Volusia County], Florida, collected June 1848 (cited imprecisely as from “near Jackson- ville” in Nelson 1981). Since Bentham indicated he had seen two collections of his new species, no holotype can be cited. It would indeed bea straightforward matter to designate the Tampa Bay collection as lectotype if it were not for a series of uncertainties that time and continued investigation have not resolved. The collector and date of collection of the Tampa Bay specimen remain obscure. Since correspondence and specimen exchanges between Alvan Went- worth Chapman of Apalachicola, Florida, and John Torrey, New York, and be- tween Torrey and George Bentham are well known, our first assumption was that Chapman was either the collector or responsible for transmitting the col- lection. Certainly Chapman knew the plant; his Flora of the Southern United States (1860) recorded the species as occurring in “Middle and South Florida.” But the date borne by the specimen is too early for Chapman’s involvement. The unclear digit, as shown by a photograph, is most likely a “2”, a date of 1826 would have required Chapman, born in 1809, to have been 17 at the time of col- lection. Even if the digit were a “3,” as seems marginally possible, Chapman is unlikely to have been the collector or communicator; he did not move to Florida (to Quincy, Gadsden County), asa young physician, until 1835 (Barnhart 1921). Other early Florida collectors were considered as the possible source. The Seminole Wars, which began in 1818, resumed in 1835 and continued intermit- tently until the 1840s, made the Florida peninsula an inhospitable place for travel and collection. Yet the military forces sent into the state permitted a few soldier-botanists to reach areas otherwise inaccessible and unknown. Of those whose duties brought them into central Florida, at least three served at Fort Brooke, modern Tampa, or reasonably would have passed through that area to duty stations elsewhere. Gilbert White Hulse, an army surgeon, was based at Fort Brooke beginning in 1836, and sent specimens to Torrey (Ewan 1971). Bradford Ripley Alden, a West Point officer, arrived at Fort King, near present- day Ocala, in 1832 or 1833 and is also known to have sent specimens to Torrey. Melines Conkling Leavenworth, a doctor also stationed at Fort King, collected NELSON AND WARD 2121 widely in the northern peninsula during the 1830s. Other botanical explorers in pioneer Florida are also known (Wunderlin et al. 2000). But no records present clear evidence as to the identity of this first collector of Stachys floridana. We have been unable to discover either at K, within the Torrey herbarium (now NY), or in other herbaria, any further early collections of Stachys floridana from the Tampa Bay region. It is not apparent why Torrey, who is indicated as the source of the Tampa Bay collection now at K, would have parted with his only specimen. Though the handwriting cannot be confirmed as Bentham’, neither is it clearly that of Torrey. The possibility thus remains that Torrey trans- mitted a unicate of little perceived value, and that the “Iampa Bay” designation was added by someone other than its collector. But whoever the collector and the place of collection, Bentham’s notation of “Torrey”—and the absence of other Torrey specimens available to Bentham—strongly suggests that the collection now marked “72” was among the material in Bentham’s possession at time of description. Rugel’s Tallahassee collection of Stachys floridana is also imperfectly docu- mented. It is probable his 1843 specimen reached Bentham (at K) as part of a set purchased from Shuttleworth; Rugel’s collections of that year apparently were distributed in no other way (Geiser 1948). These sets may have been available for purchase as early as 1844—at a price of 24 Swiss francs per 100 (per “cen- tury”)—well before publication in 1848. Why Bentham failed to indicate his knowledge of the 1843 specimen is unclear; “ad Tampa Bay” may have seemed sufficient to encompass also Tallahassee. But Bentham did refer to Rugel as a source, and it appears likely that at least one of the specimens collected by Rugel near Tallahassee in May 1843, which consist of a mixture of shade and full-sun variants, was among the original material upon which Bentham based his de- scription. The Tallahassee specimen available to Bentham at Kew Gardens (K) is ona sheet bearing two collections [Fig. 1]. Two excellent plants of Stachys floridana on the right side of the sheet have been mounted alongside an unrelated Stachys sp., apparently from Chiapas, Mexico. The pathway of Rugel 176 is similarly unclear. The more detailed labels ac- companying this collection at FLAS and NCU~as from Lake Monroe, Florida— suggests that BM may have retained (and distributed to other herbaria) only an abbreviated version of the information available from the original labels. The year of collection, 1848, as stated by the FLAS and NCU specimens, is the same as publication of the name. Since the date is in typescript and of recent applica- tion, it may merely reflect an archivist’s assignment of the publication date to accompany the older printed label. If indeed this were the year of collection, there would have been scant opportunity for timely transmittal of those speci- mens from Rugel to Shuttleworth and then to Bentham. Moreover, there is no annotation or other marking on the specimen to indicate that Bentham (at K) 2122 BRIT.ORG/SIDA 21(4) fr Stachys Floridana Shuttl. a. sp. jp omaen i be J L Two plants, ee | Ae Pe od 1° i) 1 Pee , lallahassee, Florida, May 1843. ys Sp., app ly from Chiapas, Mexico.) NELSON AND WARD, 2123 had occasion to view the holdings at BM. The bulk of the Shuttleworth her- barium was not acquired by the British Museum until 1877 (Geiser 1948), and if Rugel 176 were part of that transfer it would of course not have reached En- gland until long after Bentham’s publication. Thus none of the early collections are fully satisfactory choices as lecto- type for Bentham’s Stachys floridana. Yet none of the three collections exam- ined can be categorically excluded. Fortunately, the morphological uniformity of this species, as represented by these collections, is such that no taxonomic consequence will follow the selection of one rather than another. But Rugel 176, though the collector's name was cited by Bentham, is clearly a marginal choice as lectotype; it seems unlikely that the specimen at BM, if seen by Bentham, was available to him until after his publication of the name. The second possible lectotype, from Tampa Bay, was probably in Bentham’s hands prior to publication, but it is of uncertain provenance and without known duplicates, and is of tenuous quality upon which to base a name. We have chosen the third possibility, the specimen from Tallahassee, Florida, as our lectotype: Stachys floridana Shuttlew. ex Benth. in DC,, Prodr. 12:478. 1848. Type: US.A. FLORIDA: [Leon Co.| Tallahassee, May 1843, Rugel s.n. (LECTOTYPE, designated here: K, two plants, right half of sheet [Fig. 1], ISOLECTOTYPES: BASBG, BM, MIN, MO, NY, USCH, Z; (USCH is a mixture, bearing a flowering stem of Physostegia in addition to the Stachys). — Even though Bentham made no reference to material from Tallahassee, the low probability of his having seen Rugel 176, from Lake Monroe, gives a correspond- ingly high probability to him having seen Rugel s.n., from Tallahassee; that col- lection may reasonably be interpreted as the “Rugel” specimen which he cited. The Kew Gardens collection is representative of acommon full-sun form of the plant and will, we trust, serve to anchor future understanding of this name. ACKNOWLEDGMENTS We are grateful to the curators and staff of the above-mentioned herbaria for the loan of specimens or gift of photographs for this study, especially Ray M. Harley (K), Roy Vickery (BM), and Thomas A. Zanoni (NY); to Hartmut Hilger (BSB) for composition of the German abstract; and to Loran C. Anderson (FSU), Fred R. Barrie (F), Victoria C. Hollowell (MO) for their helpful suggestions in review of our manuscript. REFERENCES BARNHART, J.H. 1921. 1n:W. Kimball, Reminiscences of Alvan Wentworth Chapman. J. New York Bot. Gard. 22: 1-11. BENTHAM, G. 1848. Labiatae. In: A.P. de Candolle, Prodromus Systematis Naturalis Regnum Veg. 12:478. CuHapman, A.W. 1860. Flora of the southern United States. Ivison, Phinney & Co., New York. 2124 BRIT.ORG/SIDA 21(4) Ewan, J. 1971.In:R.W. Long and O. Lakela, A flora of tropical Florida. Univ. of Miami Press, Coral Gables. GrISER, S.W. 1948. Biographical note on Dr.Ferdinand Rugel, American botanist.Field & Lab. 16:113-119. Newson, J.B. 1981. Stachys (Labiatae) in southeastern United States. Sida 9: 104-123. Wunoeatin, R.P., B.F. Hansen, and J.Beckner. 2000. Botanical exploration in Florida.In:Wunderlin and Hansen, Flora of Florida, vol. |. Univ. Press of Florida, Gainesville. TAXONOMY OF THE SYMPHYOTRICHUM (ASTER) SUBULATUM GROUP AND SYMPHYOTRICHUM (ASTER) TENUIFOLIUM (ASTERACEAE: ASTEREAE) Guy L.Nesom Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT North American and Central American taxa of Symphyotrichum sect. Oxytripolium (S. subulatum sensu lato and S. so utns sensu lato) have been Eneatee at pet Specnle and varietal rank. As interpreted HEX I g specific rank th f; ] taxa, including the South American native S. SaaTIOUN which is oe primarily as a waif in the U.S.A. A key is provided to the annual taxa under consideration and summaries of synonymy are given for each. State distribution records ted for the following: S. subulatum—Arkansas, Nebraska; S. divaricatum—New Mexico; S. bahame aie S.expansum—Florida, Oklahoma; S. squamatum—Alabama, California ae Louisiana, Tex RESUMEN Los taxa de Norte América y América Central de Symphyotrichum sect. Eek hen m(S. subulatum sensu lato y S. le CLuTyS sensu pat) se han trati eet tanto con rango esf varietal. Tal como se 1 I 2 i 1 indican que el » iaerepiids para los cinco taxa rice incluyendo la ne Sur Americana S. “a es m, ne se cita principalmente como una planta abandonada en U.S.A. Se ofrece una clave es anuales en epiecocen y se hacen resumenes de las sinonimias para cada uno de ate Sele dist los de los siguientes: - subulatum—Arkansas, Nebraska; S. divaricatum—Nuevo México; S. bahamense—Georgia,; S. expansum—Florida, Oklat homa: s. squamatum—Alabama, California, Florida, Texas. Seven taxa of Symphyotrichum sect. Oxytripolium (DC.) Nesom (Symphyotri- chum subg. Astropolium (Nutt.) Semple) comprise Symphyotrichum (Aster) subulatum (Michx.) Nesom sensu lato and Symphyotrichum (Aster) tenuifolium (L.) Nesom sensu lato. Six of these taxa are native primarily to North America and Central America (including the Antilles and Bahamas); one is native to South America. One or several of them occur as cosmopolitan weeds, but iden- tifications need to be reexamined for accuracy and consistency. Sundberg (1986, 2004, 2005) has followed a broad species concept, emphasizing putative in- tergradation among the taxa (see comments below), and treated S. subulatum as a single species with five varieties and S. tenuifolium with two varieties. All seven of these taxa are treated here (and in Nesom 1994) at specific rank; bases for the taxonomic decisions are differences among the taxa in morphology, ge- ography and ecology, chromosome number, self compatibility, and sterility in SIDA 21(4): 2125-2140. 2005 2126 BRIT.ORG/SIDA 21(4) natural and experimental hybrids. Information on reproductive biology, chro- mosome numbers, and hybridization is from Sundberg (1986). The observations and considerations here are predicated on the initial study and sorting of the North American oxytripolioid taxa by Sundberg (1986, 2004), who found the larger patterns in a taxonomically difficult complex and pro- vided detailed information regarding typification. My disagreements with Sundberg are primarily in asssignments of rank, based largely on interpreta- tion of data, as | mostly agree with his delimitation of taxa. While I have seen plants in the field and studied a large number of specimens, Sundberg collected this group widely and for his dissertation research had on hand several thou- sand specimens from various herbaria. The lesser intensity of the present analy- sis and commentary, however, does not invalidate the conclusions. The key pro- vided below is based on Sundberg’s dissertation study (1986) but has been modified as | worked through collections. Hopefully, the present overview will supplement that of Sundberg’s FNA treatment (2005) in facilitating more ac- curate identifications of these taxa. ss Documentation is provided for various distribution records, which have not been given in Sundberg’s dissertation or publications. His distribution maps (1986) were small-scale and did not show U.S.A. counties. Some points of the present discussion were made earlier in brief (Nesom 1994). Annual taxa—Symphyotrichum subulatum sensu lato A map compiled by Sundberg (1986) shows that in their native (New World) ranges, the five annual taxa are discrete in geographic distribution, each almost completely allopatric with the others. In those with partially contiguous ranges, he indicated in text that intermediates occur in relatively small areas, but in- termediates were not shown on the map. The taxa are morphologically distinct although by relatively small differences. Symphyotrichum squamatum and S. bahamense are tetraploids (2n = 20), while the other taxa are diploids (2n = 10). Sundberg reported naturally occur- ring intermediates between (1) S. bahamense [2n = 20] and S. subulatum [2n = 10], (2) S. bahamense [2n = 20] and S. expansum [2n = 10], and (3) S. divaricatum 2n = 10Jand S. expansum [2n = 10]. His own study, however, provided evidence regarding internal reproductive isolation among these taxa. “Artificial hybrids produced in the greenhouse among these [five] varieties are highly sterile” (Sundberg 1986, p. 63). He obtained plump achenes (presumably those that were germinable) only from crosses between S. bahamense-S. divaricatum, S. bahamense-S. expansum, and S. expansum-S. squamatum, and each of these pair- ings was between a diploid and tetraploid. A photo in Sundberg (1986) shows 15 mitotic chromosomes of a triploid artificial hybrid between Symphyotrichum bahamense and S. expansum. — Notwithstanding the significance of naturally occurring intermediates to NESOM, TAXONOMY OF THE SYMPHYOTRICHUM SUBULATUM GROUP 2127 Sundberg’s view of the variation patterns, he did not report the occurrence of a naturally occurring triploid plant among the 86 natural populations of annual oxytripolioid taxa from which he made chromosome counts. Nor, apparently (judging from his vouchers at TEX), did he make a chromosome count of a plant suspected of being a natural hybrid of a diploid-tetraploid parental cross. Semple (1992) noted that of 6908 cl ted for North Ameri- can asters (mostly yap yemncnam) and goldenrods, oe 8 (0.12%) were trip- loid. This suggests that intergradation may not be as prevalent as Sundberg surmised, if it can be inferred from Semple’s data that triploids survive at a very low frequency. Sundberg (2004, p. 906) pointedly summarized his rationale for treatment of these taxa at infraspecific rank: “The varieties intergrade morphologically where their distributions approach one another.” | have been unable to corrobo- rate this implied ubiquity of intergradation, certainly not to the extent that would suggest treating all taxa asa single species. The annual taxa appear to be essentially discrete in morphology at their points of geographic contact and overlap. Tendencies for overlapping variation in one or a few morphological fea- tures, as described by Sundberg (2004), are not necessarily the result of intergra- dation, which characteristically is understood to imply the existence of a zone of morphological intermediacy with continous gene exchange. Discontinuities in morphology imply the existence of reproductive isolation. Sundberg’s sen- tence immediately following the one above suggests that his view of “intergra- dation” reflected a broad interpretation of that process: “This [intergradation] may be the result of past hybridization events and limited gene flow across reproductive barriers.” Even with recognition that reproductive isolation exists among the annual oxytripolioid taxa, morphological differences often are subtle. Infraspecific variability and parallel variation, especially within Symphyotrichum divaricatumand S. bahamense, produce individuals that might be misidentified without an understanding of the morpho-geographic patterns. Differences among the diploid taxa, however, are clearer, and the tetraploid S. bahamense apparently is reproductively isolated from the three closely related diploid taxa with which it is contiguous-sympatric. The species concept underlying the present analysis emphasizes biological discontinuities. Annual plants of sect. Oxytripolium adventive in Australia and various Pacific islands have mostly been identified simply as Aster subulatus (eg., Walker 1976; Harden 1992; Jones 1999), although Soejima and Peng (1998) re- ported the occurrence of two taxa (as A. subulatus var. subulatus and A. subulatus var. sandwicensis) in Taiwan. Smith (1991) included A. subulatus for Fiji, noting that it probably existed only as a ballast waif. Naturalized plants from other parts of the world have been identified as A. squamatus, e.g. Europe (Tutin et al. 1976), Russia (Tamamschyan 1959), and Zimbabwe (Mapaura & Timberlake 2128 BRIT.ORG/SIDA 21(4) 2004). Where two or more of these taxa may co-occur as adventives in regions outside of their native range, observations and perspective of the present com- mentary suggest that they will remain morphologically discrete. For example, S.squamatum, S. bahamense,and S.e xpansum in characteristic mor phology have been recorded from Japan (see bel ow), The name Aster exilis Elliott (Sketch Bot. S.Carolina 2:344. 1823) has often been applied to these plants, but as noted by Shinners (1953), a type has never been located and Elliott’s description may well have applied to some form of Symphyotrichum dumosum. With heads on the upper branches “in racemes on peduncles two to four lines long,” ray florets “twice as long as the involucrum,” and occurring “in the western districts of Georgia,” the plants that Elliott de- scribed could hardly be any of the annual taxa considered here. So eae subulatum (Michx.) Nesom, Phytologia 77:293. 1994. Aster subulatus Michx., Fl. Bor.-Amer. 2:111. 1803. Symplyol trichumsubulatum var. subulatum (sensu Sundberg 2004). TYPE: U.S.A. “PENNSYLVANIA Aster subulatus var. euroaster Fernald & Griscom, Rhodora 37:183. 1935. TYPE: U.S.A. VIRGINIA. Aster subulatus var. obtusifolius Fernald, Rhodora 16:61. 1914. Type: CANADA. NEW BRUNSWICK. Aster ensifer Bosserdet, Taxon 19:250. 1970. TypF: U.S.A. MASSACHUSETTS 2n = 10. Self-compatible. Primarily outer coastal plain of the Gulf and Atlantic coasts of Canada (New Brunswick) and the eastern U.S.A. (Texas, Louisiana, Mississippi, Alabama, Florida-northeastern counties disjunct to the western panhandle region, Georgia, South Carolina, North Carolina, Virginia, Maryland, Delaware, New Jersey, Pennsylvania, New York, Connecticut, Rhode Island, Massachusetts, New Hampshire, Maine): coastal salt and brackish marshes, depressions between sand ridges, spoil banks along canals, shorelines near the coast. Also in inland marshes and saline areas of various states (Arkansas, Ne- braska, Illinois, Indiana, Ohio, Michigan, and Ontario). Semple et al. (2002) noted that the species may have been introduced into Ontario only after salt mining began in the region. It was first collected in Michigan in 1914 at a salt mine and “survives now along well salted highways” (Voss 1996). In Illinois, it is “adven- tive along highways, rapidly spreading in ne. HI.” (Mohlenbrock 2002). Label data and photos of herbarium collections made in eight counties of the Chi- cago region (V Plants 2005) indicate that the plants there grow mostly in ditches and road shoulders. Collections from south-central Arkansas (citations below) are from an area apparently polluted by salt from oil drilling. — — First reports for Arkansas. Union Co.: 5 mi S of Calion, sandy oil spill barrens, 8 Oct 1988, Sundell 8794 (VDB); edge of bare vegetation-less area in salty runoff area from oil wells beside Union Co. Rd. 25, L mi N of Urbana near a branch of Richmond Creek, 22 Oct 1987, Thomas 103,102 (NLU); salty runoff area beside small stream just F of Lawson and S$ end of Ark. 129, area graded in attempt to clean up runoff from oil wells, 22 Oct 1987, Thomas 103,117 (NLU); salty area from oil well runoff deside branch of Mill Creek, 2 mi N of Old Union and Ark. 15, 7 Oct 1988, Thomas 107,952 (NLU, TEX); Lmi N of Urbana, salty area along a branch of Richmond Creek, beside Union Co. Rd. 25,7 Oct NESOM, TAXONOMY OF THE SYMPHYOTRICHUM SUBULATUM GROUP 2129 1988, Thomas 108,025 (NLU, VDB); sandy soil in oil field N of Ark. 335 along E ae of uae ook 2 mi E of Norphlet, 15 Sep 1989, Thomas 112,871 (NLU). First reports for Nebras W of Lincoln, Oak Lake, plant very common along saline shore, 7 Oct 1974, er ae ca NEW? artificial mend aa Creek, N of eer campus, Shildneck C-14017 (TEX). ymphyotricl in Lancaster County (e.g., Shildneck C-14016, TEX). de subulatum usually is distinctive in its heads in a dense, elon- yramidal-paniculate arrangement (or corymbiform in small plants with — few heads), relatively long involucres, phyllaries without a distinct apical green zone, ray florets 1.5-2.5(-3) mm long and coiling back distally in 1/2-1 coils, disc florets 4-10(-13), accrescent pappus, and typical salt marsh habitat (the only one of the annual taxa adapted to saline substrate). Axillary heads sometimes mature as sessile to subsessile, as is characteristic of S. bahamense, but other features of S. subulatum establish its identity. It perhaps forms triploid hybrids with S. bahamense (fide Sundberg) but apparently is more completely isolated from the other annual taxa (see comments under S. divaricatum). SymP OO ae caeaaaaies (Nutt.) Nesom, Phytologia 77:279. 1994. Tripolium aricatum Nutt., Trans. Amer. Philos. Soc. 2, 7:296. 1841. Aster divaricatus (Nutt.) Torrey & i ee FLN. ee 2:163. 1841 (not Aster aia icatus L.). TYPE: U'S.A. MISSISSIPPI “inundated banks of the Mississippi,” collected by Thomas Nuttall, probably in December 1811, in the vicinity of Natchez, Mississippi, or around New Orleans, Louisiana (Graustein 1967). ASCE Suu ligulatus Shi Field & Lab. 21:159. 1953. $ ee eee awe var. 1 Sida 21:907. 2004. TYPE: US.A. TEX EX heer neomexicanus Wooton & sae ea US. Natl. Herb. ea on Type: U.S.A. NEW MEXICO (see citation and comments belo 2n=10.Self-incompatible.Common in the south-central U.S.A. (Texas, Oklahoma, Kansas, Nebraska, New Mexico, Arkansas, Louisiana, Mississippi, Alabama, Kentucky (fide Clark et al. 2005), Tennessee, Missouri (in the southeasternmost two counties), apparently spreading eastward (e.g., Virginia, Nesom 2000) and expected toappear elsewhere along the Atlantic coastal plain. Mexico (Tamaulipas southward to the vicinity of Tampico in Veracruz, northern Coahuila, and Chihuahua, in the area of Cd. Chihuahua, Cd. Delicias, and Meoqui). Figure 1. Disturbed habitats, often moist (but usually not wet), sand, loam, and clay, com- mon and often extremely abundant along roadsides and ditches and in lawns; in the drier Great Plains region, it occurs on lake shores, marsh and playa mar- gins, a es and flats. Sometimes flowering into February. D Jew M Chaves Co.: Roswell, 3800 ft, Aug 1900, Earle & Earle 327 (US, holotype of Aster neomexicanus; NMC isotype). Eddy Co.: Carlsbad Springs, Standley 40329 (US). Guadalupe Co.: Los Esteros Creek, Ischaikowsky 401 (ARIZ). The collections from Eddy and Guadalupe cos. were recorded by Sundberg on exsiccatae lists filed in herbarium TEX. Symphyotrichum divaricatum is distinct from the other annual taxa in its rela- tively long and conspicuous ray florets and in its tendency to produce heads in 2130 BRIT.ORG/SIDA 21(4) a diffuse arrangement (vs. sessile to subsessile axillary (S. bahamense), distally clustered (S squamatum, S. expd nsum), or densely elongate, pyramidal-panicu- late (S. subulatum). Before production and maturation of axillary heads, the aspect of young plants of S. divaricatum may resemble that of S. bahamense. Heads in small plants of S.divaricatum and in plants from Mississippi and Ala- bama often are produced in a corymbiform arrangement, more characteristic of S. expansum, but the larger heads, long-acuminate phyllaries, and much longer ray corollas indicate their identity. Sundberg (2004) noted that Symphyotrichum divaricatum is “the least vari- able taxon” [among the annual oxytripolioids], but I observe that it is markedly variable at least in head size (inner phyllaries (4-, 4.5-)5-5.5(-6.5) mm long) and in height (plants (3-)20-100(-200, 300) cm tall). A collection from Hidalgo Co., Texas (Cory 51331, SMU), was noted by its collector to be of plants up to 3 meters tall, “the largest aster plant | have ever seen.” Plants in lawns will continue to produce small heads even after being mowed to about 3 centimeters in height. The combined geographic range of Symphyotrichum divaricatum, S. bahamense, and S. expansum is roughly doughnut-shaped, with the Gulf of Mexico as the hole—each of the three taxa occupies a major portion of the cir- cumference. As noted below, S.divaricatum and S. expansum are slightly, inter- mittently sympatric at the extremities of their ranges in west Texas and adja- cent Mexico (Fig. 1). The geographic ranges of S. bahamense and S.divaricatum approach each other but apparently do not make contact—the easternmost portion of the range of the latter is in southern to central Alabama, while the former reaches its westernmost point in Bay, Gulf, and Franklin cos., Fla.,in the central panhandle region (Fig. 1). Symphyotrichum bahamense and S.expansum are sympatric in southernmost Florida. The geographic range of Symphyotrichum divaricatum closely approaches that of S. subulatum in places along the Gulf Coast. Plants of S. divaricatum even grow to terrestrial edges of marsh and deeper water along the coast, but habitats of the two taxa are distinct and they appear to be completely repro- ductively isolated. — l ] }; : ] Representative coastal localities for 5 ly approaching habitats of 5. subulatum). Alabama. Mobile Co.: B: srleshig Park, brackish moist can 22 Oct 1969, Kral 38290 (VDB); E of Theodore in Deer River area, sandy open dock area (Navy) along Mobile Bay, 25 Oct 1999, Kral 89064 (NLU, VDB). Louisiana. Vermilion Par.: Redfish Point, W side of Vermilion Bay, vicinity USL field station, scattered in marsh (brackish) Aha: near Symphyotrichum subulatum], 28 Oct 1961, Ree 26(VDB). Texas. Jefferson Co.: 3.5 mi SW of Port Arthur, moist places of coastal flats, 18 Nov 1945, oe 50949 (SMU). 10 mi W of S; sbine - on a y 87, sand above intertidal zone, 19 NY 1968, Mahler 5175(SMU). Locality for s on label) with S.divaricatum. Mississippi. es Co.: eS vic. eee Sof a ou- ise St.and Washington Ave., heavily disturbed fill area, oo a with oyster fragments, growing within 994, A 3 1d 2170 (V DI ) ao NY, UD}. 100 yds of Aster subulatus var. ligulatus, 5 Nov See further comments following S. expansum. NESOM, TAXONOMY OF THE SYMPHYOTRICHUM SUBULATUM GROUP 2131 b oN & Symphyotrichum bahamense fo) Fic.1.G hic distrit f5 | ict divaricatum. Al tl t h of S. bahamense in the ro panhandle and the western approach an sympatty of 5. popu U. 2 A. record as 5. divapeatinn are from BRIT-SMU, MO, NLU, NMC, OKL, TEX-LL, VDB, a 8 Rect. for 5, eg aneumin in eo are se from Te LL and from various other herbaria, accumulated ina yet un- HICEe ereae. Records for S. TIT alu VDB. Symphyotrichum bahamense (Britton) Nesom, Phytologia 77:276. 1994. Aster ahamensis Britton, Bull. Torrey Bot. Club 41:14. 1914. Aster subulatus var. bahamensis (Brit- ton) Bosserdet, Taxon 19:249. 1970. Type: BAHAMAS: GRAND BAHAMA Aster subulatus var. elongatus Bosserdet ex Jones & Lowry, oe ser. - S20: ee (not S } erdet, Taxon 19:250. 1970, nom. invalid., without designation of a ichum ongatum (Bosserdet ex Jones & Lowry) Sundberg, ae 21: 907. 2004. TYPE: ete var. € U.S.A. FLORIDA. 2n = 20. Self-compatible. From the eastern portion of the Florida panhandle throughout most of peninsular Florida to the Keys (43 counties recorded for Florida in this study); also in a few localities of coastal Georgia and in the Bahama Islands (including the type). Hispaniola: Santo Domingo, 25 Oct 1929, Ekman 13918 (LL); Alain (1962) included S. bahamense in the flora of Cuba. Ja- pan: Chiba Pref., Futtsu City, abandoned rice paddy, 8 Oct 1985, Enomoto s.n. (TEX). Ditches and depressions, pond edges, edge and upper part of salt marsh, fresh water marsh, fields, grassy roadsides, lawns, disturbed sites, woods edges. Documentation for occurrence in Georgia. Glynn Co.: ca. 0.4 mi S of Eend of bridge of Jekyll Island, 2132 BRIT.ORG/SIDA 21(4 = upper part of salt marsh, 25 Oct 1975, Duncan 29660 (VDB); ca. 1.7 mi NW of St. Simons, higher part of salt marsh, 17 Sep 1971, Duncan 23665 (VDB); exposed, low roadside at Super 8 Motel, 23 Oct 2001, McNeilus 01-339 (NLU). McIntosh Co.: ca. 2.4 mi due N of southern tip of Sapelo Island, marshy area, usually fresh water, just back of narrow strip of oaks along Duplin River, 14 Oct 1956, Duncan 20635 (BRIT, TEX). Symphyotrichum bahamense is characterized by its distinctive arrangement of heads (at first at ends of long, bracteate branches, then produced and maturing as axillary and nearly sessile or on very short lateral branches, commonly ap- pearing secund to subsecund) and ray corollas mostly 2-3.5(-4) mm long, with blue to purple laminae coiling back in 2-3 coils. The ray corollas are shorter than in S. divaricatum and the disc florets fewer. According to Sundberg (1986), intermediates between Symphyotrichum bahamense and S. subulatum “occur sporadically” in Florida on northern ex- tremities of the range of the former (Duval Co. and along the coast of the pan- handle region). He noted (2004, p. 907) that “Intergradation [with S. subulatum is demonstrated in the compactness of the capitulescence and the number of disk and ray florets.” Such putative intermediates do not appear to be common, however, and Fl’s would be triploid and sterile (Sundberg 1986). There is no speci- men at TEX indicated to be such a hybrid. Putative intergradation between Symphyotrichum bahamense and S. expansum in southern Florida was noted by Sundberg (2004, p. 907) to be evi- denced by “individuals of [S. expansum]... more robust (to 1.5 m tall) than else- where and [with] the ligules... often pink, instead of white.” If any of these pu- tative intermediates are S. bahamense-expansum hybrids, the Fl’s would be triploid and sterile. Plants of S. bahamense from southern Florida have a ten- dency for early heads to develop on relatively shorter peduncles in a corymboid arrangement; these have the general appearance of S.expansum but can be iden- tified as S. bahamense by their larger involucres and larger, blue to purple ray florets (examples: Hendry Co: Brass 33406, USF; Levy Co: Semple 3966, USF), and the later (axillary) heads tend to be sessile or short-pedunculate, more like typical S. bahamense. The couplet below give details of differences. Heads at first at ends of long, bracteate branches, then produced and maturing as axillary and nearly sessile or on very short lateral branches, commonly on one side of the main stem and appearing secund to subsecund, in paniculiform arrange- ments; inner phyllaries 5-6.5 mm long; ray florets in 2-3 series, laminae blue to purple, (2-)2.5-4 mm long and 0.2-0.4 mm wide (dried), coiling back in 2 or more coils; disc florets 11-23 Symphyotrichum bahamense Heads usually corymbiform to thyrsiform in arrangement (borne primarily on distal branches, distally clustered); inner phyllaries 4—5.5 mm long; ray florets in 1(-2) se- ries, laminae white to light pinkish or slightly blue, 2-3 mm long and 0.1-0.3 mm wide (dried), remaining straight or coiling back in 1-2 coils; re florets (6-)8-15 mphyotrichum expansum Sundberg (1986) suggested that the tetraploid eae a m bahamense may have had an alloploid origin, with parents the diploids S. divaricatum and either NESOM, TAXONOMY OF THE SYMPHYOTRICHUM SUBULATUM GROUP 2133 S.expansum or S. subulatum. Such an origin would account for at least some aspects of morphological intermediacy in S. bahamense. Symphyotrichum expansum (Poepp. ex Spreng.) Nesom, pee e) 77:281.1994. Erigeron expansus Poepp. ex Spreng., Syst. Veg. 3:518. 1826. TYPE: CUB — Aster inconspicuus Less., Linnaea 5:143. 1830. Aster exilis var. eles (Less.) Hieron., Engl. ahi 29:19. 1900. TYPE: CUBA: homotypic with E a multiflorus Hook. & Arn., Bot. Beechey Voy. 87. 1832. TYPE: U.S.A. HAWAIL Synonymy fide Jones (1984). qeigorea supalatan ies? DC. var. pea oa Nee Gen. sp. Aster. 157, 286. 1833. g, Sidi a 21:907. 2004. TYPE: U.S.A. HA AWAIT olectou pitied by sates (2004): homotypic with Ad divaricatus var. sand wicensis Tripolium subulatum (Michx.) DC. var. cubense DC,, Prodr. 5:254. 1836. Aster sul ulatus var. cubensis (DC.) Shinners, Field & Lab. 21:16]. 1953. Type: CUBA: homotypic with E yn expansus Aster divaricatus (Nutt.) Torr. & A. Gray var. sandwicensis A. Gray, Proc. pee Acad. Arts 7:173. 807. Aster a (A. Gray) Hieron., Bot. Jahrb. Syst. 29.20, 1901. Aster subulatus var. sandwicensis (A. Gray) A.G. Jones, Brittonia 36:465. 1984. TYPE: U.S.A. HAwalt as lectotypified baone: (1984). ie (1984) lectotypified ee cist ane interpreted it to represent the taxon identified here as S. squamatum. S in 1986, but later (2004) decided that the type is correctly identified as S. expansum, in the sense of the present study. Aster fee orus Nutt. var. gracilis Benth. ex Hemsley, Biol. Centr. Amer. Bot, 122. 1881. TYPE: RICA. Synonymy fide fen oe (1986). Aster pe Elliott var. australis A. Gray, Synopt. Fl. N. Amer. 1(2):203. 1884. Aster subulatus var. australis (A. Gray) Shinners, Fi & Lab. 21:158. 1953. TyPE U.S.A. HAWaIt homotypic with A. divaricatus var. sand wicensis. Aster madrensis M.E. Jones, Contr. Western Bot. 12:43. 1908. TYPE: MEXICO. CHIHUAHUA. 2n = 10. Self-compatible. Moist or wet places, southwestern USA (Texas, New Mexico, Oklahoma, Arizona, California, Nevada, Utah), Florida (southernmost counties and other scattered localities), Mexico (all states, including southern Tamaulipas, Nuevo Leon, Coahuila, Chihuahua, and Sonora), Central America (Guatemala, Belize, Nicaragua, Costa Rica), Antilles Jamaica, Hispaniola-D.R.). Hawaii. Japan: Okayama Pref., Kasaoka City, on newly reclaimed land at Kasaoka Bay Polder, 14 Sep 1984, Enomoto s.n. (TEX). First report for Oklahoma. Cimarron Co.: along a small creek ca. 7 mi E of Kenton, 25 Sep 1976, J. Taylor 23717 (BRIT). Cimarron County is the western extremity a the Oklahoma panhandle, rela- tively close to the Texas panhandle localities in ae ane tchi cite ow Disjunct localities in Texas. Hartley Co.: sandy g Punta de Agua Creek een Romero and Middle Water, in water of eee 9 Oct 1964, Correll 30339 Ct SMU) Pc oueon Co.: Lake Meredith Natl. Rec. Area, Spring C 1 fishi marsh area, NE side (immedi ately ‘dowenstream) of Sanford By Vik in water of ditch beside marsh 20 Septem- ber 2002, Nesom & O’h 853(BRIT). Mason Co.: 5 air mi NNW of Mason, 2.1 mi N of jet. of Hwy 29 and Hwy 398, then 2.4 mi NW on dirt road, 24 Sep 1999, Singhurst 8248 (TEX). Real Co.: Dry Creek, 0.1 mi S of (downstream from) mouth of Javelina Creek, ca. 800-1000 ft. N of Dry Creek Rd. (Lost Canyon Rd.) from a point 4.0 roadmiles E of its jet. with St. Rt. 55 at Barksdale, elev. ca. 1650 ft., W shoreline of pond, 3 Oct 1998, Carr 17771 (TEX). Val Verde Co.: Pecos River at Highway 90, S of the high bridge of Hwy 90, along the E bank of the Pecos, locally abundant, 9 Nov 1999, Henrickson 22624 (TEX). 2134 BRIT.ORG/SIDA 21(4) Representative documentation for Florida. Collier Co.: Vic. of Naples, S of town, common in marshy ditch, 9 Oct 1962, Cooley 9028 (USF). Dade Co.: Perrine, empty, oolitic lot, 19 Sep 1973, Kral 51893 (VDB). Lake Co.: Eustis-Trout Lake Nature Center, | Oct 199], Daube nmire s.n. (USF). Mo Co.: Big Pine Key, Sands subdivision, 8 Sep 1981, Brumbach 9729 (BRIT, USF-2 sheets); Marathon Key near intersection of Hwy | with 37th Street, roadside fill, 11 Nov 1983, Sundberg 2327 (TE X)and 2328 (TEX); K ie L ie 0.2 mS of Tavernier Creek along Hwy 1, roadside fill, 1 1983, Sundberg 2329 (TE *X).O osa Co Eeli in Air Force Bz Ase, Prassy area around 1 pond j ust ee Bl vd, along 7th St. 21 Nov on Wil ee 11929 (USF). Symphyotrichum expansum is recognized by its relatively small heads distally clustered ina corymbiform to thyrsiform arrangement and short (but still coil- ing at maturity), whitish to pinkish or light blue ray florets about as long or slightly shorter (in coiled form) than the pappus. Among the annual taxa, it is the most geographically widespread and elevationally diverse. In the western U.S.A. and Mexico, typical S. expansum occurs at 1O0-1650(-1950) meters; from Central America to Florida, it rarely grows at more than 10 meters. The range of Symphyotrichum expansum apparently slightly overlaps that of S. divaricatum in southeastern New Mexico, western Texas, and adjacent Mexico. For the most part, the two are clearly distinct, and attempts by Sundberg (1986) to cross these two diploid taxa produced 0-3% plump achenes, almost all of which were inviable. Ina yet unpublished floristic study in the Texas pan- handle region (Hutchinson Co.), typical S.expansum has been observed in close proximity, without intermediacy, to typical S.divaricatum: the latter isan abun- dant colonizer in the sandy clay at many sites of the fluctuating shoreline of Lake Meredith (e.g., Nesom & O’Kennon 689, as cited above), while S.expansum was observed in the muck of a wet ditch and marsh margin at only one area immediately below the dam (Nesom & O’Kennon 853, BRIT). In Presidio Co., Texas (Big Bend Ranch State Natural Area), the two taxa have been observed and collected in close proximity, without evidence of intergradation: S. divaricatum (Worthington 22636, TEX, UTEP) and S. expansum (Worthington 22637, TEX, UTEP). Worthington noted by annotation that he observed two species of ‘aster’ in BBRSNA. Pringle apparently observed two co-occurring entities in Chihuahua, on the “wet banks of the Sacramento River [vicinity of Cd. Chihuahual, 13 Sep 1886”: Pringle 751 (LL) is S. divaricatum while Pringle 750 (LL) is S.expansum. Even though it appears that some degree of reproductive isolation exists between Symphyotrichum divaricatum and S. expansum, Sundberg (2004, p. 906) noted that “Populations intermediate in ligule length and width occur in trans-Pecos Texas, parts of New Mexico (including the type of Aster neomexicanus, collected in Chaves Co.), Arizona, and Chihuahua, Mexico.” My observations corroborate the existence of plants with longer and slightly wider rays, which also are blue to purple, in contrast to the smaller, white to pink rays of S. expansum. Most of these occur where the two species are sympatric and apparently are relatively uncommon, compared to the parents. Such putative NESOM, TAXONOMY OF THE SYMPHYOTRICHUM SUBULATUM GROUP 2135 intermediates are similar to S. expansum in their small heads (inner phyllaries mostly 4-4.5mm long) all strongly distally disposed on wiry peduncles. Because of its relatively large ray corollas, the Chaves Co. collection is identified and mapped here as typical S.divaricatum,, although in habit it resembles S.expansum } } livaricat ? ansum). U.S.A. Texas. Brewster Co.: 3.5 Collecti ined (S S of Marathon, infrequent in naned at Pena Blanca spring, 21 Oct 1946, me nock 46587 (SMU, TEX). Jeff Davis Co.: gravel and sand bars of Limpia Creek near Ft. Davis, 8 Oct 1926, Palmer 23123 (TEX). Pecos Co.: roadside along irrigation ditch near Farm Road 1053, 1/2 mi N of Imperial, chromosome g 1967, Watson 147 (TEX); ca. 5 mi W of Fort Stockton along [H-10, moist ditch number n=5, 20 Au 2160(TEX). Presidio along frontage road S of freeway, chromosome number n=5, 26 Aug 1983, erg Co. infrequent at sa near Rex Ivy's Lodge above La Jitas, 2200 ft, 24 Sep 1961, Warnock 18163 (TEX). Reeves Co.: 285, S of BECO, — ae 1941, Strandtman s.n. (TEX). Mexico. Chihuahua: Delicias, along ne 10 Oct 1957, Knol MU) The broad distributions and distinct ene of Symphyotrichum divaricatum and S. expansum, their overlap and co-occurrence in a relatively small zone of sympatry, and the relatively few putative intermediates are taken here as rationale for treating both of them at specific ran Symphyotrichum squamatum (Spreng.) Nesom, Phytologia 77:292. 1994. nee za sq a Spreng., Syst. Veg, 3:515. 1826. Aster squamatus (Spreng.) Hieron., Bot. Jahrb. Syst. 29:19. 1901. Conyzanthus squamatus (Spreng.) Tamamsch., Fl. U.R.S.S. 25: ve 1959, ea ae subulatum var. squamatum (Spreng.) Sundberg, Sida 21:908. 2004. TYPE: URUGUAY. MONTEVIDEO. Erigeron semiamplexicaule ee Reise 1:31]. 1834. TYPE: ?. Synonymy fide Cabrera (1978). meres ci Colla, M Reale Accad. Sci. Torino 38:14, pl. 25.1835. Aster asteroides ~ la) y, Mem. Torrey Bot. Gu 4.213. 1895. TYPE: CHILE: Synonymy fide Sundberg (1986 ee berteroana Phil,, Linnaea 28:737. 1836. Type: CHILE: Synonymy fide Sundb cs Tripolium conspicuum Lindley ex DC, Prodr. 5.254. 1836. Aster bangii Rusby [nom. nov], Mem. Torrey Bot. Club 4:213. 1895. ae CHILE: Synonymy fide Sundberg (1986). Aster linifolius Griseb., Abhand. Konigl. Gesellsch. Wissens. Gottingen 24:178. 1879. TYPE: ? Syn- onymy fide Cabrera (1978). Aster ae ae Ann. New York Acad. Sci. 7:139. 1893. TYPE: PARAGUAY: Synonymy fide Sundberg (1986). Tripolium moelleri Phil, Anal. Univ. Chile 87:403. 1894. Aster oe (Phil.) Reiche, Anal. Univ. Chile 109:338. 1901. Type: CHILE: Synonymy fide Cabre Tripolium oliganthum Phil. Anal. Univ. Chile 87:403. 1894. ae ie LE: Synonymy fide Cabrera (1978). — Erigeron depilis Phil, Anal. Univ. Chile 87:417. 1894. TYPE: CHILE: Synonymy fide Cabrera (1978). Aster barcinonensis Sennen, Bull. Acad. Int. Geogr. Bot. 23:242. 1914. Type: SPAIN: Synonymy fide Sundberg (1986) 2n = 20. Self-compatible. Native to South America and apparently widely dis- tributed there: rare in California and the southeastern U.S.A. (Alabama, Florida, Louisiana, Texas), apparently mostly as a waif, usually on or near beaches and ballast dumps. Naturalized in Australia), Japan), Iraq(), Africa), France(), and probably other regions of the world. Noted by Britton (1914) to occur on Ireland Island and Boaz Island, Bermuda. 2136 BRIT.ORG/SIDA 21(4) Documentation for U.S.A. occurrences. Alabama. Mobile Co.: sandy W end of Sener Island, 18 Oct 1973, Tayorand Taylor si ites Dauphin Island, Itasca PI. near Iberville Dr, roadside near dunes, 9 Aug 1965, Deramus D752 (VDB); Battleship Park, by Mobile-Baldwin Co. causeway, abundant on moist brackish sands, 22 Oct 1969, Kral 38282 (NLU). California. Kern Co.: S of Greenfield, intersec- tion of Cottonwood Rd and Buena Vista Rd, along roadside ditch, 16 Apr 1983, Sund berg 2093, “this population n = 10 pairs” (TEX). Florida. [Escambia Co.]: Pensacola, waste ground, 27 Jul 1899, Curtiss 6497 (GH [fide Shinners 1953], USF); Franklin Co.: Apalachicola, ballast weed, Jul 1897, Chapman s.n. (MO [fide Shinners 1953], SMU). Louisiana. Orleans Par.: weedy areas ae streets N of New Orleans Convention Center from Howard Street W to elevated hwy in New Orleans, 10 Nov 1991, Thomas 126,773 (NLU). Texas. [Galveston Co.]: “Galveston, sandy beaches, damp sands along the streets, 8 Aug 1902, J. Reverchon 3319” (MO, US-2 sheets); the MO collection was cited by Shinners (1953) as Aster pear var. australis; it was identified as Symphyotrichum squamatum by Sundberg (1986) and Nesom (19¢ annette squamatum is recognized by its corymbiform to thyrsiform arrangement of heads (borne primarily on bracteate distal branches and dis- tally clustered), inner phyllaries 5-5.5 mm long, with sharply delimited apical green zones, and ray florets numerous (21-28(-38)) with filiform, erect (non- coiling) corollas shorter (13-2 mm long) than the mature pappus. It is the only one of the taxa treated here that is not a North American native: its evolution- ary relationship to the others may be correspondingly distant. Natural hybrid- ization has not been reported between S. squamatum and any other taxon. KEY TO THE ANNUAL TAXA |. Heads usually dense in an elongate, pyramidal-paniculate arrangement inner phyl- laries 6-7 mm long, wae apices linear-acuminate, distal margins often inrolled/ involute, green zone of phyllaries narrowly lanceolate, usually extending the entire length of the ye 1s bases short or absent; pappus accrescent, 4—5.5 mm long at maturity and usually ae than coiled ray corollas; habitats wet, saline __ Symphyotrichum subulatum 1. Heads corymbiform to thyrsiform, diffusely paniculate, or secund to subsecund and paniculiform arrangements or at the tips of long, bracteate branches; inner phyllar- ies 4-6.5 mm long, phyllary — seus to acuminate, distal margins inrolled/in- volute or not, green zone of phylla late to elliptic, chartaceous bases usu- ally conspicuous; pappus not accrescent, 3.5—4(-5) mm red at maturity, longer or shorter than ray corollas; habitats moist to wet, rarely 2. Phyllary tips appressed, acute, flat, inner phyllaries 7 er lanceolate, dis- tinctly demarcated, apical green zone, proximal 1/2-1/3 white-chartaceous; ray floret laminae erect, often involute along the edges (curling inward lengthwise), rarely coiling back distally (if so, then only ca. 1/2 coil), usually shorter than mature pappus; disc florets (3-)7- = sympnyetichum squamatum 2. ae) tips Ris linear-acuminate, distal ins oft ,inner yllari wly lanceolate, often weal apical green zone, tite: areca 1s bases short, ca. 1/3-1/2 the iecath of the gy ray floret laminae not involute along edges, usually coiling back distally in 1-4 or more coils, usually as long or longer than mature pappus; disc florets nee 5,11-23, or (20-)33-45(-50) - | [ae tn +} oe . } . Al 3, Heads usually OPYIMIDITOTIN) t \ arrangement (WOME PHIMdllly ON NESOM, TAXONOMY OF THE SYMPHYOTRICHUM SUBULATUM GROUP 2137 distal branches, distally clustered); inner phyllaries 4—5.5(-6) mm long, phyl- lary apices acute to abruptly short-acuminate or long-acuminate, distal mar- gins inrolled/involute or not; ray florets in 1(-2) series, corollas 2-3 mm long, laminae 0.1-0.3 mm wide (dried), white to light pinkish or slightly blue, coil- ing back in 1-2 coils or less commonly remaining straight; disc florets (6-) 8-15 Symphyotrichum expansum Head arrangements diffusely paniculiform to pyramidal-paniculiform to corymbiform or secund to subsecund and paniculiform; inner phyllaries 5— 6.5 mm long; phyllary apices long-acuminate, distal margins usually inrolled/ involute; ray florets in 1-3 series, corollas 2-7 mm long, lamina m wide (dry), white to blue or puple, coiling back in 2-4 or more eee disc flo- rets 11-23 or (20-)33-45(—50). Heads often at ends of long, bracteate branches, axillary heads usually m turing on elongate lateral branches, the whole arrangement sen diffusely paniculiform to pyramidal-paniculiform, or heads more distally disposed and the arrangement corymbiform to thyrsiform; ray florets in 1 series, co- rollas mostly 4-7 mm long, laminae 0.4-0.8 mm wide (dry), blue to white, coiling back 3-4 or more times; disc florets (20-)33-45(—50); south-central U.S.A., extreme northwestern Mexico Symphyotrichum divaricatum Heads at first at ends of long, bracteate branches, then produced and ma- turing as axillary and nearly sessile or on very short lateral branches, com- monly on one side of the main stem and appearing secund to subsecund, in paniculiform arrangements; ray florets in 2-3 series, ne mostly 2- 3.5(-4) mm long, laminae 0.2-0.4 mm wide (dry), blue to purple, coiling back in 2-3 coils; disc florets 11-23: Florida, coastal Georgia, Bahamas Symphyotrichum bahamense bw > > Perennial taxa—Symphyotrichum tenuifolium sensu lato Symphyotrichum tenuifolium sensu stricto and S. bracei (S. tenuifolium var. aphyllum) are diploid, rhizomatous perennials endemic to coastal and near- coastal habitats. The former occurs mostly in marshes of western Cuba, the Bahamas, and the west coast of southern and central Florida; S. tenuifolium occurs in marshes along the Gulf coast from Texas to panhandle Florida and then along the Atlantic coast from northeastern Florida northward as far as Rhode Island, New Hampshire, and Maine. The key by Sundberg (2004) sepa- rates the two perennial taxa in a number of features. Each of them has been treated at specific rank by regional botanists (Cronquist 1980; Wunderlin 1982, 1998: Wunderlin & Hansen 2004), but Long (1970), Long and Lakela (1971), and Sundberg (1986, 2004) have regarded S. braceiasa variety within a more broadly conceived species. In the initial description of var. aphyllum, Long (1970, p. 4D) noted that it is “connected by intermediate forms” to var. tenuifolium and is “a West Indian-Florida population segregate of the more northern|[S. tenuifolium]” Sundberg (1986, 2004) observed that the two taxa intergrade where their ranges overlap along the Gulf Coast of northern and central peninsular Florida, from Taylor to Pinellas counties, where “almost all specimens are intermediate,” sug- gesting that parental forms apparently are absent or rare and that gene flow is 2138 BRIT.ORG/SIDA 21(4) continuous. In contrast, | find that typical S. bracei occurs northward well into the range of typical S. tenuifolium (e.g, Hernando Co, S. bracei, Sundberg 2305, TEX; Citrus Co., S. bracei, Schmid A-6, USF, Taylor Co., S. bracei, Godfrey 61659, BRIT). Sundberg (2004) scored Godfrey 61659 (FSU) as typical of S. bracei ex- cept for root structure, which is lacking on the BRIT duplicate. Species-rank concepts of Cronquist, Wunderlin and Hansen, and the present study emphasize the distinctive morphologies of the two taxa, their mostly allopatric ranges, and the apparent hybridization and intermediacy that occurs within only a relatively small area of overlap. Long and Sundberg have emphasized the zone of intermediacy as rationale for treating these two taxa as geographic segments of a single species. Whichever point of view is adopted, treatment of these at specific rank may be more subjective than for the annual taxa, where reproductive isolation apparently is stronger. Symphyotrichum tenuifolium (L.) Nesom, Phytologia 77:293. 1994 (1995). Aster tenuifolius L., Sp. PL. 2:873. 1753. Type: U.S.A. “in America septentrionale Symphyotrichum bracei (Britton ex Small) Nesom, Phytologia 77:276. 1994. Aster bracei Britton ex Small, Fl. Miami 190, 200. 1913. Type: BAHAMAS. NEW PROVIDENCE. Aster tenuifolius var. aphyllus R.W. Long, Rhodora 72:40.1970. Symphyotrichum wien ium(L.) Nesom var. dphyllum (R.W. Long) Sundberg, Sida 21:905. 2004. TyPF: U.S.A. FI ACKNOWLEDGMENTS Dick Wunderlin, Bruce Hansen, and Kanchi Gandhi made the critical interpre- tation, adopted here, of the validation and correct authority for Aster subulatus var. elongatus. Dick Wunderlin and John Semple made various helpful comments and observations. lalso am grateful to the staff of TEX-LL for their help and to NLU, NMC, and USF for loans of specimens. REFERENCES ALaIn, H. 1962.Flora de Cuba (Compositae) 5:175-313. Britton, N.L. 1914. Studies of West Indian plants—V. Bull. Torrey Bot. Club 41:1- Casrera, A.L. 1978.Flora de la Provincia de Jujuy, Republica Argentina. Coleccion Cientifica del INTA, Tomo XIll, Parte X: Compositae. Buenos Aires. Ciark, R.C.,R.L. Jones, . Weckman, R.L. THompson, J.W. THieret, Kentucky Stare NATURE Preserves Com- Mission, and K. Freeman. 2005. State records and other noteworthy collections for Ken- tucky. Sida 21:1909-1916. Cronauist, A. 1980. Vascular flora of the southeastern United States, Vol. |. Asteraceae. Univ. of North Carolina Press, Chapel Hill. Graustein, J.E. 1967. Thomas Nuttall, naturalist: Explorations in America, 1808-1841. Har- vard Univ. Press, Cambridge, Mass Harden, GJ. (ed.). 1992. Flora of New South Wales. Vol. 3. New South Wales Univ. Press, Kensington, NSW, Australia. — NESOM, TAXONOMY OF THE SYMPHYOTRICHUM SUBULATUM GROUP 2139 Jones, A.G. 1984. Nomenclatural notes on Aster (Asteraceae)—Ill. The status of A. sandwicensis. Brittonia 36:463-466 Jones, A. 1999. Aster, p. 266, in Wagner, W.L., D.R. Herbst, and S.H. Sohmer. Manual of the flowering plants of Hawai'i (rev. ed.), Volume 1. Univ. of Hawai'i Press and Bishop Mu- seum Press, Honolulu, Hawaii. Lone, R.W.and O.LakeLa.1971.A flora of tropical Florida. Univ. of Miami Press, Coral Gables, Florida. Lone, R.W. 1970. Additions and nomenclatural changes in the flora of southern Florida-l. Rhodora 72:17-46. Maraura, A.and J. TimBervake (eds.).2004.A checklist of Zimbabwean vascular plants. South- ern African Botanical Diversity Network Report No. 33.SABONET, Pretoria and Harare. MOoHLENBROCK, R.H. 2002. Vascular flora of Illinois. Southern Illinois Univ. Press, Carbondale and Edwardsville. Nesom, G.L. 1994. Taxonomic overview of Aster sensu lato (Asteraceae: Astereae), empha- sizing the New World species. Phytologia 77:141-297. Nesom, G.L. 2000. Noteworthy collections from herbarium NCU [Apiaceae, Asteraceae, Geraniaceae, Rhamnaceae]. Castanea 65:80-83. Sempte, J.C. 1992.A geographic summary of chromosome number reports for North Ameri- can asters and goldenrods (Compositae: Astereae). Ann. Missouri Bot. Garden 79: 9 Semete, J.C., $.B. Hearp, and L. Brouittet. 2002. Cultivated and native asters of Ontario (Compositae: Astereae). Univ. Waterloo. Biol. Ser.41:1-134 SHINNERS, L.H. 1953. Notes on Texas Compositae-IX. Field & Lab. 21:155-162. Smith, A.C. 1991. Flora Vitiensis nova: A new flora of Fiji (Soermatophytes only). Vol. 5. Angiospermae: Dicotyledones, families 170-186. Pacific Tropical Botanical Garden, Lawai, Hawaii. Soeima, A.and C.-l. Pene. 1998. Aster (Compositae). In: Huang, T.C., ed. Flora of Taiwan (ed. 2). Vol. 4, Angi ,Dicotyledons [Diapensiaceae—Compositae]. Editorial Committee po | of the Flora Semin National Taiwan Univ. Taipei. Pp. 848-868. SunpserG, S.D. 1986. The systematics of Aster subgenus Oxytripolium (Compositae) and historically allied species. Ph.D. dissertation, Univ. of Texas, Austin. SunpeerG, $.D. 2004. New combinations in North American Symphyotrichum subgenus Astropolium (Asteraceeae: Astereae). Sida 21:903-910. Sunpsers, S.D. 2005. Symphyotrichum (in part).In:Flora of North America Editorial Commit- tee, eds, 1993+. Flora of North America North of Mexico. Vol.21.New York and Oxford. Tamamscuyan, S.G. 1959. Conyzanthus. In: B.K. Schischkin, ed. Flora of the USSR [translated from Russian, 1999]. Smithsonian Institution Libraries, Washington, D.C. Pp. 174-177 [originally 185-188]. Tutin, T.G., V.H. Heywoop, N.A. Burces, D.M. Moore, D.H. Vacentine, S.M. Watters, and D.A. Wess. 1976. Flora Europaea, Vol.4:Plantaginaceae to Compositae (and Rubiaceae). Cambridge Univ. Press, Cambridge. 2140 BRIT.ORG/SIDA 21(4) Voss, E.G. 1996. Michigan flora. Part Ill: Dicots (Pyrolaceae—Compositae). Cranbrook Insti- tute of Science Bull.61 and Univ. Michigan Herbarium, Ann Arbor. V Piants. 2005.V Plants: A virtual herbarium of the Chicago region. The Morton Arbore- tum, Field Museum of Natural History, and Chicago Botanic Garden. Accessed May 2005. Wacker, E.H. 1976. Flora of Okinawa and the southern Ryukyu Islands. Smithsonian Institu- tion Press, Washington, D.C Wunberuin, R.P. 1982. Guide to the vascular plants of central Florida. Univ. Presses of Florida, Tampa. WuNbertin, R.P. 1998. Guide to the vascular plants of Florida. Univ. Press of Florida, Gaines- ville. Wunberuin R.P. and B.F. Hansen. 2004. Atlas of Florida vascular plants (internet). [S.M.Landry and K.N.Campbell (application development). Florida Center for Community Design and Research.] Institute for Systematic Botany, Univ. of South Florida, Tampa. Accessed April 2005. PAPPUS VARIATION IN NORTH AMERICAN ASTERS. |. DOUBLE. TRIPLE AND QUADRUPLE PA PPUS IN SYMPHYOTRICHUM AND RELATED ASTER GENERA (ASTERACEAE: ASTEREAE) John C. Semple and Jennifer L.A. Hood Department of Biology University of Waterloo Waterloo, Ontario, CANADA N2L 3G1 jcsemple@sciborg.uwaterloo.ca ABSTRACT The pappus traits of 84 taxa of Canadanthus, Ampelaster, Psilactis, and ei m (subtribe Symphyotrichinae) and 14 other species of asters were examined. Most species of Symphyotrichinae had a pappus consisting of three whorls; 1) a secondary inner whorl of a few ee 40-85% the length of the inner primary whorl, 2) a primary outer whorl of many bristles with tapering ends about 90-95% the length of the inner primary whorl, and 3) a primary inner whorl of many bristles with very weakly to strongly clavate ends. In addition, a few species of Symphyotrichum have a ves- tigial secondary outer whorl of a single short bristle only about 10-30% the length of the primary inner whorl. In some species of Symphyotrichum the secondary inner whorl was difficult to distin- guish from the primary outer whorl or possibly was absent. In a small sample of species of Aster, en rid, Eurybia, Eucephalus, Galatella, Herrickia, lonactis, Oclemena, and Oreostemma, the appus was either miple or quadraple: The short secondary outer whorl was present in most species sae ae 1 to very few bristles or al fruits. In some cases, the secondary inner whorl of mid length tapering bristles was present in some individuals in a species but not in others. The quadruple pappus with strongly clavate inner bristles appears to be plesiomorphic for the North American Clade of the tribe Astereae. In single species samples of Crinitaria, Linosyris, and Felicia the pappus differed from that of other species examined RESUMEN = examinaron las caracteristicas ck los yates de 84 taxa de Canada nthus, Ampelaster, Psilactis, y y otras 14 especies de compuestas. La mayoria de ie acnecies de Symphyotrichinae denen un vilano de tres ie 1) un verticilo secundario interno de pocas sedas con el 40- ae de la longitud gel venue interno prima io, 2) un verticilo externo primario con I d | baer 95% de la onic ch ee : | | a Ig peciesd a richum tienen un hala peed eee | : j Je] 10-30% del verticilo interno palma, es un AAlys | ue ee ] | alongit Id ¢ VETLICIiO interno primario. En alguna especies de Sane el verticilo es interno fue dificil de diferenciar 2 primario externo o posiblemente estaba ausente. En una pequena muestra de especies de Aster, ae Eurybia, aoe Suiatelld Herrickia, longs Oclemena, y el mma, a Vv jilano era triple ) CoD L ] a | ] a | oO c oF 5 J = ] ] -] = th } enalgunos [rutos. ca algunos casos, e presente en algunos individuos a una especie pero no en otros. El vilano cuadruple con las sedas SIDA 21(4): 2141-2159. 2005 2142 BRIT.ORG/SIDA 21(4) interiores fuert te clavadas parece ser plesiomorfico para el clado Norte Americano de la tribu Astereae. En una sola muestra de especies de Crinitaria, Linosyris, y Felicia el vilano diferia de las otras especies examinadas. INTRODUCTION The symphyotrichoid aster genera Canadanthus Nesom, Ampelaster Nesom, and Symphyotrichum Nees have long been reported to have a simple pappus with non-clavate bristles, either as species of Aster L. (e.g., Gray 1884; Fernald 1950; Cronquist 1955, 1968a, 1980, 1994) or recently as species of Canadanthus, Ampelaster, and Symphyotrichum (Nesom 1994, 2000; Semple et al. 2002). In contrast, some other North American asters historically treated at times in other genera or in Aster sensu lato have been reported to have a double (Eucephalus Nutt., Sericocarpus Nees; Gray 1884; Cronquist 1955) or a triple pappus (Doellingeria Nees, Cronquist 1968, 1980; Nesom 1994; Semple et al. 2002). The putatively double pappus consisted of a short whorl of outer bristles and a much longer whorl of inner bristles, while the putatively triple pappus had a short outer whorl and two long inner whorls, the outer slightly shorter and tapering and the inner bristles clavate. Species of Solidago L. historically were also con- sidered to have a simple pappus (e.g., Gray 1884; Fernald 1950; Cronquist L968b, 1980; Nesom 2000; Semple et al. 1999). However, Hood and Semple (2003) dem- onstrated that nearly all species of goldenrods had a double pappus with two primary whorls of long bristles (the outer shorter and tapering, the inner clav- ate) and at least some species had an additional secondary outer whorl of a few very short bristles. That is, a genus thought to have a simple pappus in fact had a triple pappus like that reported for Doellingeria (synonym: Aster sect. Triplopappus Torr. & A. Gray; Torrey and Gray 1841; Semple et al. 2002). Our discovery that goldenrods had a double (or rarely triple) pappus raised the pos- sibility that other genera of Astereae thought to have a simple pappus might also be double or triple. This paper on pappus variation is one ina series being prepared by the first author’s lab to report the findings of investigations of ae traits, including pappus variation in the North American Clade of the raeae (sensu Noyes & Rieseberg 1999). The pappus whorl terminology pro- — by Hood and Semple (2003; secondary outer whorl, primary outer whorl, and primary inner whorl) is used throughout this paper with the addition of new label for a fourth whorl not seen in Solidago, the secondary inner whorl of tapered mid length bristles. MATERIALS AND METHODS A preliminary survey was undertaken to examine under the dissecting micro- scope the pappus bristles of one or two specimens of representative species of the sections and subsections of Symphyotrichum. Subsequently, a more rigor- ous survey was conducted involving 80 taxa and one hybrid of Symphyotrichum SEMPLE AND HOOD, 2143 and 4 species of other genera in the subtribe Symphyotrichinae Nesom and 14 species of other North American and Eurasian aster genera listed in Table 1. Observations were made using a dissecting scope (LO-70X) ora compound light microscope (20-400). The degree of the clavateness of bristle tips was deter- mined using the 0-4 rankings described in detail in Hood and Semple (2003). At least five different fruits from each species were observed under the dissecting microscope at a maximum of 70; most observations were made at 30-40. For the most part, observations were made on specimens in the WAT Herbarium, but additional material on loan from BRIT, CAN, GH, and NY (Holmgren et al. 1990) was also examined to expand the number of taxa. In addition to several methods used in evaluating pappus features listed by Hood and Semple (2003), assessment of the presence or absence of any short secondary outer whorl lin- ear to scaly bristles and other pappus features were a tematically under- taken. Thus, Table 1 has five columns of observational data plus a column for additional comments, while only four columns were included by Hood and Semple (2003). Observations on the compound microscope were made from slides prepared as follows. For each species, two ray floret and two disc floret cypselae (with co- rollas or without corollas still attached) were mounted in Cytoseal'™™ 60 (low viscosity) mounting medium under a cover slip. Observations at 100-400 on the compound microscope were made similarly to the observations under the dissecting scope at 30-70. Observations made on the two kinds of scopes were compared and any discrepancies were resolved by re-examining specimens. Digital photomicrographs were taken using a Nikon CoolPix 990 camera manually held against the ocular lens of either the dissecting or compound microscope. Pictures were taken of specimens under the compound light mi- croscope with either below stage or above stage lighting. Final digital illustra- tions were made using CorelDraw 12® from digital images edited with Corel PhotoPaintl2® (Corel Corp.). RESULTS AND DISCUSSION Vice The pappus of the majority of species of the Symphyotri consists of three whorls (Figs. 1-4): 1)a secondary inner whorl of a few intermediate length, fine, tapering bristles usually about 50-70% of the length of the primary inner whorl, 2) a primary outer whorl of tapering, non-clavate bristles that were generally 5-10% shorter than the inner whorl, and 3)a primary inner whorl of very weakly to strongly clavate tipped bristles. The phylogenetically more basal taxa of the Symphyotrichinae Canadanthus modestus (Lindl.) Nesom (Figs. 1A-G), Ampelaster carolinanus( Walt.) Nesom (Figs. IH-K), and Almutaster pauciflorus (Nutt.) Love & Love (Figs. 2A-E) are all diploids (2n = 18) with x = 9 chromo- somal base number (Brouillet et al. 200la, b; Semple et al. 2002). These have a triple pappus with no secondary outer whorl of short bristles or scales observed. Taste 1. Pappus variation in subtribe Symphyotrichi S f ict and related genera. Clv, degree of clavateness of inner bristles (0 = not clavate to 4 = strongly clavate); Clv-Tap, clavate alternating with tapered bristles (primary inner sad outer whorls, y = yes, — not eulousy 50); Lou, primary outer bristles shorter than primary inner bristles; Ovrlp, degree of overlapping of bristles at base (O= not observed; i=slight overlap; 2 p);2"4-Out, evidence fora he i outer whorl of very short scaly-bristles. Taxon Clv Clv-Tap Lgth Ovrip 2°4-Qut Comments SYMPHYOTRICHINAE Nesom Canadanthus Nesom (x = 9) .modestus 1 y y 2 n 27 inner bristles 60% of 1° inner Ampelaster Nesom (x = 9) A. carolinianus 2 y y 0) n 2°° inner, few, 50% of 1° inner Almutaster Love (x = 9) A. pauciflorus 1 y y 0 n 2” inner 50-60% of 1° inner Psilactis A. Gray (x = 9,4, 3) is (x = 4) ] y y 0 n heterocarpic, ray fruit epappose; 2™7 inner bristle 40-60% of 1° inner Symphyotrichum Nees subg. Chapmaniani (Semple) Semple (x = 7) S. chapmanii 2 y y 2 y 2°° inner, few, 50-60% of 1° inner; possible 2" outer whorl bristle, one seen, ane 30% of 1° inner subg. Symphyotrichum sect. Symphyotrichum (x = 8) subsect. Symphyotrichum series Symphyotrichum S. novi-belgii Le) | ak c. = | 2° inner, very few, 70% of 1° inner S. retroflexum 4 y y 2 y 2” inner bristles, few , 70-8% of 1° inner; possible 2°° outer bristle, 30% of 1° inner, only seen on one fruit S. robynsianum 0 - y 0) n 2° inner, few, 70-80% of 1° inner series Punicei (House ee S. elliottii 2 y y 1 n 2°° inner, few, 70-80% of 1° inner wa prLZ (p)LZ WOIS/9YO'LINE Taxon Clv Clv-Tap Lgth Ovrlp 2"4_Out Comments S. prenanthoides 0 - y 0 2”? inner, few, 60-80% of 1° inner 5. puniceum 0 - y 2 n 2"? inner bristles, few, 40-70% of 1° subg. Symphyotrichum sect. Symphyotrichum (x = 8) subsect. Occidentales (Rydb.) Nesom (Foliacei) chilense y y n 2"? inner, few, 75-80% of 1° inner S.eatonii 2 y 0 n 2° inner, few, 75-80% of 1° inner S. foliaceum 3 y y 0 n 2° inner, very few, 80% of 1° inner S.jessicae 2 y y ) n 2" inner, very few, 70-80% of 1° S. spathulatum 3 y y 0 n 2" inner, very few, 70-80% of 1° S. subspicatu 3 y n 2” inner bristles not seen subg. Symphyotrichum sect. Symphyotrichum (x = 8) subsect. Heterophylli (Nees) Semple series Cordifolii (G.Don in Loudon) Semple S malum ] y y 1 n 2” inner bristles, few, 80% of 1° S. ciliolatum 0 y y 0 n 2° inner bristles, few, 70-80% of 1° S.cordifolium 1 y y 0 n 27 inner bristles, few, 70% of 1° S.drummondii | y y 0 n 2% inner bristles, few, 70% of 1° S. shortil 1 y y ] n 2°¢ inner bristles, few, 70% of 1° S.texanum 1 y y 0 n 2°? inner bristles, few, 70-80% of 1° S.undulatum 2 y y 0 n 2° inner bristles, few, 80% of 1° S. urophyllum 0 y y 0 n variable bristle lengths; 2"° inner bristles, few, 60-70% of 1°; 1° outer 85-90% of 1° inner series Concinni (Nees) Semple 2 y y 0 n 2°¢ inner bristles, few, 70-80% of 1° S. oolentangiense =) y y 2 n 2° inner bristles, few, 70-80% of 1° S.attenuatum 0 - y 0 n 2™ inner bristles 60-70% of inner subg. Symphyotrichum sect. Symphyotrichum (x = 8) subsect. Dumosi (Torr.& A. Gray) Nesom S. boreale 0 - y 0 n 2° inner bristles, few, 60-70% of 1° S.dumosum var.dumosum 1 y y 0 n 2°° inner, O-few, 70-80% of 1° inner var. strictior 2 y y 0 n 2°° inner, very few, 70% of 1° inner ‘QOOH ONY 31dW3S SPL? TABLE 1. (continued) Taxon Clv Clv-Tap Lgth Ovrlp 2"4-Qut Comments subg. Symphyotrichum sect. Symphyotrichum (x = 8 S.eulae 3 S.lanceolatum subsp. hesperium var. hirsuticaule var. interior var. lanceolatum var. latifollum S. lateriflorum var Jateriflorum ar. angustiflolium S. nahanniense S.praealtum var. angustior var. praealtum S.ontarione S. tradescanti _welshii S. parviceps S.porteri NNN ast et ls RF 1 subg. Symphyotrichum sect. Symphyotrichum (x = 8 S.depauperatum 0) 0 2 patio ) subsect. Dumosi (Torr.& A. Gray) Nesom (continued) colon. como OOM. Linnaeus, en su reconocimiento original ,escogid “colonum” como epiteto. Esta palabra, del latin ‘colonus,” es un Oseinaye. En si, bajo el Cédigo Inte acien a, de Nonencatina Botanica, debe conservar su ortogtalia original, euglania que sea el géner fermenting, ) ae | d Asi pues el 7 la ecnecie ec F. } oO L oO rv Link. INTRODUCTION The Old World grass known as Jungle Rice,a member of the genus Echinochloa (Gramineae), is now found in tropical and temperate areas throughout the world. Though in ancient times and into the early 20th century it served as an edible grain, it is now best known asa secondary forage for cattle or as a troublesome weed of wet soils. It was given scientific recognition in the mid-18th century, first as a species of Panicum, then as a member of the newly formed genus Echinochloa. In these two genera it has borne only a single specific epithet. Yet over the decades different authors, with about equal frequency, have given this epithet two spellings—colona and colonum. But few have attempted to explain their chosen spelling. And none have provided adequate justification of which spelling is correct. HISTORY Jungle Rice—a common name of relatively recent origin—has been known from ancient times. The grains have been found in the intestines of mummies of early Egypt, where their amount and purity make it “most probable that the plant SIDA 21(4): 2171-2183. 2005 2172 BRIT.ORG/SIDA 21(4) was cultivated as a cereal” (Tackholm & Drar 1941). Its use as a foodstuff in India continued to be documented into the 20th century (Gamble 1928; Bor 1968). The plant was recorded and illustrated by at least three pre-Linnaean authors: Plukenet (1692), Sloane (1696, 1707), and Ehret (1748). When Linnaeus (1759) published Panicum colonum he transcribed the epithet wholly in lower case. His diagnosis of the new species read: P spiculis alternis secundis muticis ovatis scabris, rachi teretiuscula. He referred to the il- lustrations of both Sloane (1707) and Ehret (1748). His basis may have been a specimen (LINN 80.23; Savage 1945) now in the Linnaean Herbarium obtained from the Irish physician, Patrick Browne, who returned to England in 1756 af- ter a stay in Jamaica (Stafleu 1971); the sheet bears a “Br” in Linnaeus’ hand. He may also have seen material in the Jamaica collections of Sir Hans Sloane whom he visited in 1736 (Stearn 1957:110), and was certainly familiar with the plate in Sloane's (1707) book. The plant’s relatively distinct morphology, which matches the Sloane and Ehret plates, has made a secure linkage with Linnaeus’ diagno- sis and name; no later author has questioned the accuracy of their application to the grass known as Jungle Rice. Panicum is a genus of great size, recently estimated (Mabberley 1996) to contain more than 500 species. It would be even larger if certain groups of spe- cies had not been removed as deserving of independent generic rank. Echinochloa was among the first of these distinctive groups to be given generic standing, by Beauvois (1812), to contain the familiar Barnyard-grass, FE. crusgalli (L.) Beauv. Beauvois at the same time formed a second generic segregate, Oplismenus. Kunth (1816) then made a transfer of Panicum colonum, to form Oplismenus colonus (L.) HBK. and Link (1833) published the now-universally accepted combination Gf not spelling), Echinochloa colona (L.) Link. Floristic botanists were slow to accept the new combinations. Influential writers throughout the 19th century—Hooker (1897) and Cooke (1908) in India; Grisebach (1864) in the West Indies; Nash (in Britton & Brown 1898), Chap- man (1897), and Mohr (1901) in North America—continued toemploy Panicum colonum. [Nash (1898), under Britton’s editorial edict that all species must have common names, is the apparent originator of the now widely used “Jungle Rice.”] Perhaps Merrill (1923) was the last important author to retain Linnaeus’ P colonum. Other than its initial formation, the first significant use of Linnaeus’ epi- thet in the segregate genus Echinochloa appears to have been by Nash (in Small’s “Flora of the Southeastern United States” 1903), who chose to follow Link (1833) in forming the epithet as “colona.” Nash seized a second opportunity to pro- mote this spelling by his authorship of the grasses in Britton and Brown (1913), expanding the usage into northeastern North America. The practice received further approbation by Hitchcock (1909) in Cuba; by Stapf (in Prain 1920) and Hutchinson and Dalziel (1936) in Africa; by Gamble (1928) in India; by Hitch- — WARD, A CASE Ur LONUM 2173 cock Gn Small’s “Manual of the Southeastern Flora” 1933); and by Rozhevits and Shishkin (1934) in Russia. But opposition began to arise to the burgeoning use of colona. Hitchcock (who in his earlier works had employed colona) noted (1913): “Dr. E.L. Greene called attention to the fact that the specific name is not an adjective, and sug- gested that it is probably a genitive plural. The word appears to be contracted from colonorum, genitive plural of colonus,a husbandman or a colonist. Dr. J.A. Nieuwland has kindly searched Latin authorities and verifies this conclusion, though there appears to be no direct authority for the word colonum.” Wiegand (1921)—other than Hitchcock, the only author expressing an opin- ion who had devoted significant time to the taxonomy of Echinochloa—ob- served: “Hitchcock, following Greene, has called attention to the fact that the name colonum is not an adjective and hence should not be declined.” And Bor (1960), perhaps irritated by the use of colona without explanation by other au- thors addressing the flora of India, brusquely commented: “The correct form of the specific epithet is colonum, a contraction of colonorum, and not colona.” Argument in defense of colona was slow to appear. Clayton (in Hepper 1968) may have been the first: “The declension of the epienct depends on whether it is regarded as a noun or an adjective. Lexic differ, but the adjectival use was acceptable to those of Linnaeus’ own time. ” This ealannon was expanded by Cope Gn Nasir & Ali 1982): “The epithet is sometimes treated as the irregu- lar genitive plural of a noun (of the farmers’) and spelt colonum. However, there seems no reason to depart from the adjectival form familiar to botanists; though not in the purest classical tradition, its use was sanctioned by lexicographers of Linnaeus’ own time.” Though citing as his authority an author who had taken the contrary view, Michael (2003) justified colona: “Hitchcock (1913) consid- ered that ‘colonum’ was a non-declining contraction, but dictionaries of Linnaeus time treated it asa declining adjective. Because Linnaeus was the first to name the species (as ‘Panicum colonum), it seems best to follow the practice considered correct in his day; hence ‘E. colona’.” Only these six authors have been found who expressed a justification for their use of either Echinochloa colona or E. colonum. The many others either held no opinion or gave none in their floristic writings. The three most detailed North American studies—Hitchcock 1920; Wiegand 1921; Gould et al. 1972—all used colonum, though only Hitchcock and Wiegand provided justification of the spelling. Two world-scale compilations of plant names (Uphof 1968; Mabberley 1996) pointed in opposite directions. Two comprehensive listings of plant names for temperate North America (Shetler & Skog 1978; Kartesz 1994) similarly differed in their spelling of the epithet. A recent and influential in- ventory of plants of economic importance worldwide (Wiersema & Leon 1999) chose colona. A cursory survey of floristic authors addressing Echinochloa has shown 2174 BRIT.ORG/SIDA 21(4) that in the 20th century (and into the 21st) there is rough equivalence to the two positions; 36 have used E. = while 43 have used E. colonum. Auth hol loyed Echinochl A lo-Rodriguez (1996), Allen (1992), Balick, Nee aa ene (2000), Britton and Brown (1913), cee and Renvoize (1982, 1986), Clewell (1985), Dassanayake et al. (1994), Diggs et al. (1999), Duncan and Kartesz (1981), Gamble (1928), Gibbs Rus- sell et al. 1991), Gould (1975), Green (1985), Hepper (1968), Hitchcock (1909), Howard (1979), Hutch- inson and Dalziel (1936), Jones et al. (1997), Maire (1952), Michael (2003), Nasir and Ali (1982), Prain (1920), Robinson and Fernald (1908), Rozhev ; hishkin (1934), she (1903, 1933), Stace (1997), Stevens et al. (200), Thulin (1995), ce et al. (2003), Wagner et al. 1990), Watson and Dallwitz (1992), Wolford and Kral (1993), Wunderlin es ee etal. i Authors who have employed Echinochloa colonum: Adams (1972), Backer (1968), Blomquist (1948), Bor (1960, 1968), Britton and ee 11920), Correll and Correll (1982), Correll and Johnston (1970), Davis (1985), Fernald (1950), Gleason (1952), Gleason and Cronquist (1963), Godfrey and Wooten (1979), Hall 1978), Hatch et | (1990), Hite ae oe 1920, 1931, 1935, 1936), Hitchcock and Chase (1917, 1951), Hodge (1954), Li et al. (1978), Maheshwari (1967), Proctor (1984), Pulle (1966), Radford etal. (1968), Rechinger (1964, 1971), Robyns and Tournay (1955), Saldanha ae scene 976), Shouliang (1990), Srivastava (1976), Standley (1937), Steyermark (1963), Swallen (1955), Tackholm and Drar (1941), Terrell (1977), Tutin et al. 1980), Walker (1976), Wiggins (1980), ae (1976). But correct orthography, as in other more obvious niches of plant taxonomy, is not governed by popular vote, but by conformation to codified rules. Though rules are difficult to understand, sometimes treacherous to follow, they are the nly path to consistent usage. DISCUSSION Orthography is “the art of spelling words according to accepted usage” (Ran- dom House 1979). In taxonomic parlance, “accepted usage” is defined by the rules of the International Code of Botanical Nomenclature (Greuter et al. 2000). In the present instance, the determination of whether colona or colonum is cor- rect is decided by interpretation and application of Article 23, the Names of Species. It is critical to determine whether the word was first used as an adjec- tive, or as a noun. Resolution of these alternatives requires that there be under- standing of the origin of the word and its use in the naming of Jungle Rice. Colonus was a term used in the late Roman Empire for a worker who was bonded to the farmland of a wealthy landowner; though technically nota slave, the worker was not free to seek employment elsewhere. (This practice later be- came the coerced labor of the middle-age feudal system.) The word colonus isa second declension Latin noun; it is masculine. It is often translated as “free- born serf,” or at times as “husbandman,” a now-obsolete term surviving only as “animal husbandry,” the care and raising of agricultural animals. Occasionally it is read as “farmer” or as “colonist,” in recognition of the modern inapplicabil- ity of the original meaning. Latin is a highly inflected language (Stearn 1983), that is, the ending of each word indicates the case, number, and gender. Colonus is the nominative WARD, A AASL VE ECHIN 2175 singular, colonum the accusative singular, coloni the genitive singular, colonorum the genitive plural, etc. Were a Roman to observe, “The colonus kicks the horse” (or equus, also a second declension noun), he would say, “Colonus equumcalcitat.” Were the horse to do the kicking, the expression would be, “Equus colonum calcitat.” If the horse kicks more than one person, “Equus colonos calcitat.” In common practice the word colonus had no feminine ending, that is, there appears to have been no widely used colona in the Latin language (Lewis & Short 1879). That spelling appears to be recorded only twice in the ancient writ- ings (by Ovid). Just as in “horse” (where “mare” indicates the female), the femi- nine gender most often would have been expressed by a separate word or disre- garded entirely. Similarly, there cannot be a neuter colonum in the nominative; the structure of the language may appear to permit it, but the concept of a “neu- ter” worker would be without meaning. Moreover, there is no adjective colonus (or colona or colonum) in the Latin language. It seems most improbable that “dictionaries of Linnaeus’ time treated it as a declining adjective” (Michael 2003). Indeed, were colonus treated as an adjective, the word would be unintelligible when translated into English (free- born serfish”?). The statements by the six authors who gave reasons for their use either of colona or colonum need examination. All were brief, some cryptic, and some misleading or erroneous. The three who spoke for continued use of colonum (Hitchcock 1913; Wiegand 1921; Bor 1960) clearly understood the word to be a noun (the wording of Hitchcock and of Wiegand: “not an adjective”). They erred, perhaps, in that each seemed to assume his readers would properly interpret this fact to require retention of the original spelling. Two spoke of the wor colonum being a contraction of colonorum; this remark is unneeded in that, while colonorum is available (the genitive plural of colonus), colonum is itself a perfectly good form (the accusative singular), thus requiring no “contraction.” Nieuwland, as quoted by Hitchcock (1913), raised a further detail, that “there appears to be no direct authority for the word colonum.” There indeed seems to be no documented colonum in classical Latin (Lewis & Short 1879). Nieuwland’s point may be that he believed proper taxonomic style calls for use only of Latin words known to be recorded in surviving Latin writings. Yet, once recognized as a second declension noun, the word colonus implies appropriate spellings in other number and case. The three authors who spoke for changing the spelling to colona (Clayton 1968; Cope 1982; Michael 2003) are more difficult to understand. The claim that lexicographers “of Linnaeus’ own time” accepted colona (Clayton, paraphrased by Cope and Michael) is made without documentation, and no such lexico- graphic treatment has been seen (the usage by Ovid perhaps excepted). Most significantly, all three state or indicate that they believe the word colonus may 2176 BRIT.ORG/SIDA 21(4) be treated as an adjective and thus altered in spelling to agree with the associ- ated genus, perhaps without comprehending that such action must carry them outside the parameters of acceptable Latin. Linnaeus did not employ the epithet elsewhere than in 1759. The word colonus, however spelled, appears to be found in post-Linnaean technical bo- tanical literature only in application to the grass described by Linnaeus (Google, Nov 2003). There are thus no guiding examples of its use with other genera by other authors. The pathway is indirect by which Linnaeus probably came to use this word for his epithet. The references he cited lack the word: colonum does not appear in the phrase-names accompanying the plates of Sloane (1707) nor Ehret 1748), nor in Sloane’s (1696) more extensive text. But Sloane (1696)—though this pub- lication was not cited by Linnaeus—referred to a still-earlier publication: Plukenet (1692). There, under a drawing that may be the first illustration of Jungle Rice, and accompanied by a phrase name (Gramen paniceum minus, spica divulsa) cited by Sloane, Plukenet noted his plant to be “Pestis Coloni,” or “plague of the farmer.” Plukenet’s work was well known to Linnaeus, and it is most probable that this phrase (Coloni, here, in the genitive singular) was the inspiration for his selection of “colonum.” APPLICATION TO BOTANY Classical Latin, of course, is not the same as botanical Latin (Stearn 1983). The use of Latin as an international language, a practice of the past 250 years, is relatively rigid, with many words given precise meanings unknown to the Ro- man writer or scholar. These meanings may originate, not with their classical use, but with the application toa botanical situation, as determined by the Re- cent author who needs a special term for a special structure. If the term is employed, not just for descriptive purposes, but for a botani- cal name, the author's latitude is without limit. It is generally recognized that good style encourages an author to use a term or combination of terms, from Latin or Greek, that closely track classic usage. But, encouragement aside, there is no requirement in the Code (Greuter et al. 2000) that the word (or words) used in forming a name be appropriate, or that it be spelled correctly, or that it have any meaning whatsoever. A provision of the Code (Art. 23.1; Greuter et al. 2000) would appear to re- strict this latitude: “The name of a species is a binary combination consisting of the name of the genus followed by a single specific epithet in the form of an adjective, a noun in the genitive, or a word in apposition...” Colonum, though accusative, is used in apposition. But the near-simultaneous permission (Art. 23.2) that an epithet “may even be formed arbitrarily” does allow, by modern rules, deviation from strict nominative structure for a word used in apposition. WARD, A CASE Vr 2177 Thus Linnaeus, though writing far in advance of the modern rules, still falls within their parameters. It was common practice for Linnaeus (1753, et seq.) to select as the epithet for hisnew name a word pre-existing in the medieval botanical literature. Many of these words, perhaps most, were adjectives. But others were nouns—known as substantives—and are carried over unchanged into modern botanical usage. Until the mid 20th century many authors indicated the substantive origin of epithets by retaining a capital letter at the beginning of each epithet so formed. Now, though capitalization of substantives is still permitted, the majority of authors de-capitalize epithets, giving uniformity to the structure of names, but obscuring the history and the original usage of the epithet. Nouns used for epithets are treated differently from adjectives. Adjectives must agree (in case, number, and gender) with the genus to which they are at- tached, and this agreement is indicated by the requisite change in spelling. In contrast, the Code (Art. 23.5; Greuter et al. 2000) mandates that a noun retains its own gender and ending irrespective of the gender of the generic name. Anexample lies near at hand, of a noun used as an epithet and transferred, without change in spelling, to a genus of another gender. Linnaeus (1753) also described and named the plant now commonly known as Barnyard-grass; he termed it Panicum Crusgalli. “Crusgalli” is literally translated as “chicken’s leg,” but is usually interpreted to mean “cock’s-spur.”) When transferred to Echinochloa by Beauvois, it became E. crusgalli (in modern, preferred usage). Though Linnaeus did not indicate the source of this epithet, his use of a capital initial letter designated it asa substantive. No juent author has attempted to treat it as an adjective and adjust the original spelling so as to agree with the gender of the new genus. Linnaeus’ Panicum colonum was received differently. In the masculine ge- nus Oplismenus, Kunth (1816) recorded it as O. colonus. In the feminine genus Echinochloa, Link (1833) stated it to be E. colona. The arguments so weakly pre- sented for treating colonum as an adjective would perhaps have been strength- ened had their proponents noted that the originators of these segregate genera had done so also. Had Clayton (1968) referred to botanists rather than lexicog- raphers, he would have been accurate in his observation that “adjectival use was acceptable to those of Linnaeus’ own time.” The judgments of Kunth and of Link, however, are still just judgments of later authors, no different from those of the many still later authors who chose E. colona. Only the action by the original author, Linnaeus, could potentially carry decisive weight. It is unknown why Linnaeus (1759) chose “colonum’ as the spelling of the epithet for his new species. He was, of course, assigning the new entity to the genus Panicum, a genus he had formed earlier 1753) and which he had treated 2178 BRIT.ORG/SIDA 21 = 4) as neuter (as indicated both by the ending of the word (-um) and by the ending of adjectival epithets he placed thereunder). Since Linnaeus both wrote and spoke Latin (Stafleu 1971:83), he cannot be thought of as making a beginner's error, that is, he would have known full well that the nominative was colonus and that the word, carried into botanical usage, would normally retain its nomi- native spelling unchanged. Too, if he obtained his epithet from the brief usage by Plukenet (1692), he knew the word to be a noun. Yet his use of lower case for the initial letter of colonum indicates he thought of the word as formed differ- ently from other substantives. Two alternatives are offered. Perhaps Linnaeus did understand the word to be a noun and chose the accusative, or colonum, for reasons of euphony, for smooth combination with its assigned genus Panicum. Or perhaps Linnaeus chose to disregard its meaning asa noun and saw it only asa sequence of letters which could be treated as an adjective and declined to agree with its genus. CONCLUSION The second of these possibilities is untenable. One cannot break away from the certainty that Linnaeus would have recognized the word was a noun and must have intentionally chosen the accusative, colonum, so that it would follow smoothly his genus Panicum. His choice of colonum is within the practices of the 18th century and the language of the modern Code. His preference for the harmonious colonum rather than the discordant colonus in no way negates its status as a noun. His decapitalization of the initial letter is stylistic and imma- terial. Noargument seems convincing that Linnaeus thought of the word as an adjective. Though Linnaeus, by creating the new name, had the option of se- lecting for its epithet whatever word he wished, his choice of a word that is a noun removes the power of later authors to treat it as an adjective. As a noun whose spelling is unchanged in whatever genus it may be placed, the name formed by Link in 1833 must be read as Echinochloa colonum. ACKNOWLEDGMENTS [thank J. Richard Abbott (FLAS), Mary Barkworth (UTC), William J. Dress (BH), Mark A. Garland (PIHG), Walter S. Judd (FLAS), and Dan H. Nicolson (US) for their useful and important comments and suggestions, Christine M. Housel (ABT) for the Hispanicized abstract, and Emily W. Wood and colleagues (GH) for access to critical pre-Linnaean publications. REFERENCES Acevebo-Ropriauez, P. 1996. Flora of St. John, U.S. Virgin Islands. New York Botanical Garden, New York. [522-E£. colona] Apams, C.D. 1972. Flowering plants of Jamaica. Univ. of the West Indies, Mona. [186-E. colonum] WARD, A LADSE UF ECHINOCHLOA COLONUM 2179 Auten, C.M. 1992. Grasses of Louisiana, 2nd ed. Cajun Prairie Habitat Pres. Soc., Eunice. [98:. colona] Backer, C.A. 1968. Flora of Java. Groningen, Netherlands. [3:556-E. colonum] Batick, M.J., M.H. Nee, and D.E. AtHa. 2000. Checklist of the vascular plants of Belize. New York Botanical Garden Press, New York. [187-E. colona] Beauvois, P. 1812. Essai d'une Nouvelle Agrostographie 53, 161. Biomauist, H.L. 1948. The grasses of North Carolina. Duke Univ. Press, Durham. [185-E. colonum] Bor, N.L. 1960. Grasses of India, Burma and Ceylon. Pergamon Press, London. [308-F. colonum| Bor, N.L. 1968. Flora of Iraq. Ministry of Agric., Baghdad. [9:479-E. colonum] Britton, N.L.and A. Brown. 1898. Illustrated flora of the northern United States, Canada and the British Possessions, 1st ed.Grasses:G.V. Nash. New York Botanical Garden, New York. [3:496-Panicum colonum] Britton, N.L.and A. Brown. 1913. \llustrated flora of the northern United States, Canada and the British Possessions, 2nd ed. Grasses: G. V. Nash. New York Botanical Garden, New York. [1:134-E. colona] Britton, N.L.and C.F. MittspauGu. 1920. The Bahama flora. New York. [26-E. colonum] CuHapman, A.W. 1897. Flora of the southern United States, 3rd ed.Cambridge Botanical Sup- ply, Cambridge, Mass. [587—Panicum colonum] Crayton, W.D. and S.A. Renvoize. 1982. Flora of tropical East Africa: Gramineae. Royal Botani- cal Garden, Kew. [557-559-E. colona] Crayton, W.D. and S.A. Renvoize. 1986. Genera graminum: Grasses of the World. Royal Bo- tanical Garden, Kew. [289-E. colona] Crewett, A.F. 1985. Guide to the vascular plants of the Florida panhandle. Florida State Univ. Press, Tallahassee. [139-E. colona] Cooke, T. 1908. Flora of the Presidency of Bombay. Taylor and Francis, London. [931—Pani- cum colonum] Corrett, D.S. and H.B. Corrett. 1982. Flora of the Bahama Archipelago. Cramer, Vaduz, Ger- many. [132-EF. colonum] Corrett, D.S. and M.C. Jounston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner. [183-E. colonum] Dassanayake, M.D, E.R. Fosserc, and W.D. Ciayton. 1994. Flora of Ceylon. Amerind Publ. Co., New Delhi. [8:181-E. colona] Davis, PA. 1985. Flora of Turkey. Univ. Press, Edinburg. [9:145-E. colonum] Diccs, G.M., B.L. Liescoms, and RJ. O’Kennon. 1999. Shinners & Mahler's illustrated flora of north central Texas. Botanical Research Inst. of Texas, Fort Worth. [1266-E. colona] Duncan, WH. and J.T. Kartesz. 1981.Vascular flora of Georgia. Univ.of Georgia Press, Athens. [15-E. colona] Euret, G.D. 1748. Plantae et Papiliones Rariones. Heidelberg, Germany. [t. 3, f.3] Fernato, M.L. 1950. Gray's Manual of Botany, 8th ed. American Book Co., New York. [223-E. colonum] 2180 BRIT.ORG/SIDA 21(4) Gamete, J.S. 1928. Flora of the Presidency of Madras. Adlard and Son, London. [3:1776-E. colona] Gises Russett,G.E., L. Warson, M. Koekemoer, L. SMook, N.P. Barker, H.M. ANDERSON, and M.J. DALLwitz. 1991. Grasses of southern Africa. National Botanic Garden, Pretoria. [1 19-E. colona] GLEASON, H.A. 1952. Illustrated flora of the northeastern United States and Canada. Lan- caster Press, Lancaster, Pennsylvania. [1:233-F.colonum] Gteason, H.A. and A. Cronauist. 1963. Manual of vascular plants of northeastern United States and adjacent Canada.Van Nostrand Co., Princeton, New Jersey.[114-E£.colonum] Goprrey, R.K.and J.W. Wooten. 1979. Aquatic and wetland plants of the southeastern United States. Univ. of Georgia Press, Athens. [1:155-E. colonum] Goutp, FW. 1975. The grasses of Texas. Texas A. & M. Univ. Press, College Station. [538-F. colona] Goutb, F.W., M.A. Au, and D.E. FairsrotHers. 1972. A revision of Echinochloa in the United States. Amer. Midland Naturalist 87:36-59. [56-E. colonum] Green, J.W.1985.Census of the vascular plants of western Australia, 2nd ed. Dept.of Agric., Perth. [32-E. colona] GreuTer, W.,J.McNeit, ER. Barrie, H.M. Burvet, V. DEMOULIN, T.S. FiticuieRas, D.H. Nicotson, P.C. Siva, J.E.Scoe, P. TREHANE, and N.J. Turtano. 2000. International code of botanical nomenclature. Saint Louis Code. Koeltz, Germany. GriseBacH, A.H.R. 1864. Flora of the British West Indian Islands. Reeve & Co,, London. [545- Panicum colonum] Hatt, D.W. 1978. The grasses of Florida. Univ. of Florida, Ph. D. diss. Gainesville. [338-E. colonum] Hatcu, 5.L., K.N.GANbHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Experiment Station, College Station. [183-E£. colonum] Heprer, FN. 1968. Flora of West Tropical Africa, 2nd ed.Grasses:W.D. Clayton. Crown Agents, London. [3:439-440-E. colona] HitcHcock, A.S. 1909. Catalogue of the grasses of Cuba. Contr. U.S. Nat. Herb. 12:183-258. [213-E£. colona] HircHcock, A.S.1913.Mexican grasses in the United States National Herbarium.Contr. U.S. Nat. Herb. 17:181-—389.[256-257-E. colonum] HircHcock, A.S. 1920. The North American species of Echinochloa. Contr. U. S. Nat. Herb. 22:133-153.[150-E. colonum] HitcHcock, A.S. 1931. North American Flora. New York Botanical Garden, New York. [17(4):311-E. colonum] HitcHcock, A.S. 1935. Manual of the grasses of the United States, 1st ed. U.S. Dept. Agric., Washington. [692-EF. colonum] HitcHcock, A.S. 1936. The genera of the grasses of the United States. U.S. Dept. Agric. Bull. 772.(247-E. colonum] HitcHcock, A.S.and A. CHase. 1917. Grasses of the West Indies. Contr. U.S. Nat. Herb. 18:261- 471.[345-E. colonum] WARD, A CASE OF ECHIN HLO h N 2181 HitcHcock, A.S. and A. CHase. 1951. Manual of the grasses of the United States, 2nd ed. U.S. Dept. Agric., Washington. [711-E. colonum] Hooce, W.H. 1954. Flora of Dominica, B.W.|. Lloydia 17:1-238. [1:131-E. colonum] Hooker, J.D. 1897. Flora of British India. Reeve & Co., London. [7:32—Panicum colonum] Howarb, R.A. 1979. Flora of the Lesser Antilles. Grasses: F. W. Gould. Arnold Arboretum, Ja- maica Plain, Mass. [3:125-E. colona] HutcHinson, J. and J.M. Datziet. 1936. Flora of West Tropical Africa, Ist ed. Crown Agents, London. [2:558-E. colona] Jones, S.D.,J.K. Wipre, and PM.Montcomery. 1997.Vascular plants of Texas. Univ. of Texas Press, Austin. [243-E. colona} Kartesz, J.T. 1994. A synonymized checklist of the vascular flora, 2nd ed. Timber Press, Portland, Oregon. [438-E. colona] KUNTH, C.S. 1816. Nova genera et species plantarum. [1:108-—Oplismenus colonus] Lewis, C.T. and C. SHort. 1879.A Latin dictionary. Clarendon Press, Oxford. Li, H., T. Liu, T. HUANG, T. Koyama, and C.E. DeVou. 1978. Flora of Taiwan. Grasses: C. Hsu. Epoch Publ. Co., Taipei. [5:552—E. colonum] Link, J.H.F. 1833. Hortus Regius Botanicus Berolinensis. Berlin. [2:209-E. colona] Linnaeus, C. 1753. Species plantarum. Stockholm. Linnagus, C. 1759. Systema naturae, 10th ed. Stockholm. [2:870-—Panicum colonum] Massertey, D.J. 1996. The plant-book, 2nd ed. Cambridge. [248-EF. colona] ManesHwarl, P.1967.The flora of Delhi. Gossain & Co., Calcutta. [393-F. colonum] Maire, D.R. 1952. Flore de l'Afrique du Nord. Lechevalier, Paris. [1:311-E. colona] Merritt, E.D. 1923.Enumeration of Philippine plants. Dept. of Agriculture and Natural Re- sources, Manila. [1:62—Panicum colonum] MicHaet, PW. 2003. Flora of North America. Oxford Univ. Press, New York. [25:398-400-F. colona] Monr, C.1901.Plant life of Alabama.U.S. Dept. Agric., Washington. [358—Panicum colonum] Nasia, E. and S|. Au. 1982. Flora of Pakistan, no. 143. Grasses: T. A. Cope. Islamabad. [196- 197-E. colona]. PLukeneT, L. 1692. Phytographia. Londini. [tab. 189, fig. 5] Prain, D.1920.Flora of tropical Africa. Grasses: O. Stapf. Reeve & Co.,London.[9:607-E. colona] Proctor, G.R. 1984. Flora of the Cayman Islands. Royal Botanic Garden, Kew. [190-191-€. colonum]. Putte, A. 1966. Flora of Surinam. Netherlands. [1(1):409-E£. colonum] Raororo, A.E., H.E. AHtes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. Grasses: A. E. Radford. Univ. of North Carolina Press, Chapel Hill. [132-E£. colonum] RecHINGeR, K.H. 1964. Flora of lowland Iraq. Grasses: N. L. Bor. Cramer, Weinheim, Germany. [51-E. colonum] RECHINGER, K.H. 1971.Flora lranica. Grasses: N.L. Bor. Akademische Druck, Graz, Austria. [70:479- E.colonum] Rosinson, B.L. and M.L. Fernaco. 1908. Gray’s New Manual of Botany, a handbook of the 2182 BRIT.ORG/SIDA 21(4) flowering plants and ferns, 7th ed. Grasses: A. S. Hitchcock. American Book Co., New York. [118-E. colona] Rosyns, W.and R. Tournay. 1955.Flore des Soermatophytes du Parc National Albert. Institut des Parcs Nationaux du Congo Belge, Bruxelles. [3:92—EF. colonum] RozHevits, R.Y.and B.K. SHISHKIN. 1934. Flora of the U.S.S.R. National Science Foundation, Wash- ington. [2:33-E. colona] SALDANHA, C.J. and D.H. Nicotson. 1976. Flora of Hassan District, Karnataka, India. Amerind Publ. Co., New Delhi. [726-E. colonum] Savace, S. 1945, A catalogue of the Linnaean Herbarium. London SHETLER, S.G.and L.E. Skoa. 1978. A provisional checklist of species for Flora North America (revised). Missouri Botanical Garden, St. Louis. [163-F. colonum] SHOULIANG, C. 1990. Flora Republicae Popularic Sinicae. Science Press.[10(1):252-F.colonum] Stoane, H. 1696, Catalogus plantarum quae in Insula Jamaica. Londini. [p. 30] Stoane, H. 1707. A voyage to the Islands Madera, Barbados, Nieves, S. Christophers and Jamaica. London. [t. 64, f. 3] Smatt, J.K. 1903. Flora of the southeastern United States, 1st ed. Grasses: G. V. Nash. New York. [84-E. colona] Smait,J.K.1933.Manual of the southeastern flora. Grasses: A. S. Hitchcock. New York. [83—E. colona] Srivastava, T.N. 1976. Flora of Gorakhpurensis. Today and Tomorrow, New Delhi. [363-E. colonum] Stace, C. 1997. New flora of the British Isles, 2nd ed. Cambridge Univ. Press, Cambridge. (911-E. colona] Starteu, FA. 1971. Linnaeus and the Linnaeans. Utrecht, Netherlands. STANDLEY, PC. 1937. Flora of Costa Rica. Field Museum of Natural History, Chicago. [18:75-E. colonum} Stearn, W.T. 1957. An introduction to the Species Plantarum and cognate botanical works of Carl Linnaeus. Bartholomew Press, Great Britain. Stearn, W.T. 1983. Botanical Latin, 3rd ed. David & Charles, London. Stevens, W.D.,C. Uttoa, and A. Monteit. 2001. Grasses: R.W. Pohl and G. Davidse. Flora of Nica- ragua. Missouri Botanical Garden Press, St. Louis. [3:2034-F. colona] STEYERMARK, J.A. 1963. Flora of Missouri. lowa State Univ. Press, Ames. [233-E. colonum] Swatten, J.R. 1955. Flora of Guatemala. Field Museum of Natural History, Chicago. [2:112-E. colonum] TackHotm, V. and M. Drar. 1941. Flora of Egypt. Fouad Univ., Cairo. [1:446-EF. colonum] Terrell, E.E.1977.A checklist of names for 3,000 vascular plants of economic importance. U.S. Dept. Agric. handbook 505. Washington. [47-E. colonum] THULIN, M. 1995. Flora of Somalia. Grasses: T. A. Cope. Royal Botanic Gardens, Kew. [4:223-F. colona] Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Botanical Research Inst. of Texas, Fort Worth. [805—E. colona] WARD, A LAE UI 2183 Turin, T.G., V.H. Heywooo, N.A. Burces, D.M. Moore, D.V. VALENTINE, S.M. Watters, and D.A. Wess. 1980. Flora Europaea. Cambridge Univ. Press, Cambridge. [5:262-E. colonum] UpxHor, J.C.T. 1968. Dictionary of economic plants, 2nd ed. Cramer, New York. [193-E. colonum] Waaner, W.L., D.R. Herest, and S.H. Soumer. 1990. Manual of the flowering plants of Hawaii. Grasses: P. J.O’Connor. Univ. of Hawaii Press, Oahu. [2:1535-E. colona] Waker, E.H. 1976. Flora of Okinawa and the Southern Ryuku Islands. Smithsonian Institu- tion Press, Washington. [211-E. colonum] Warson, L. and M.J. Datiwitz. 1992. The grass genera of the world. C. A. B. International, Cambridge. [341-E. colona] Wiecano, K.M. 1921. The genus Echinochloa in North America. Rhodora 23:49-65. [53-E. colonum] Wiersema, J.H. and B. Leon. 1999. World economic plants: A standard reference. CRC Press, Boca Raton, Florida. [195—E. colona] Wicains, |.L. 1980. Flora of Baja California. Stanford Univ. Press, Stanford. [942-E. colonum] Worrorp, B.E. and R. Krat. 1993. Checklist of the vascular plants of Tennessee. Botanical Research Inst. of Texas, Fort Worth. [19-E. colona] Wunoeatin, R.P. 1998. Guide to the vascular plants of Florida. Univ. Press of Florida, Gaines- ville. [106-E. colona] ZANGHERI, P. 1976. Flora Italica. Grasses: A. B. Cattarini. Padova, Milan. [1:910-E. colonum] Zutoaca, F.O., O. Morrone, G. Daviose, T.S. Ficcueiras, PM. Peterson, R.J. Sorena, and E. Juoziewicz. 2003.Catalogue of New World grasses. Smithsonian Inst.,Washington.[3:215-E.colona] 2184 BRIT.ORG/SIDA 21(4) BOOK NOTICE NEAL K. VAN ALFEN, GEORGE BRUENING, AND WILLIAM O. Dawson (eds). 2005. Annual Review of Phytopathology: Volume 43, 2005. (ISBN 0-8243-1343-7, hbk; ISSN 0066-4286). Annual Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews.org, 800- 523-8635, 650-493-4400, 650-424-0910 fax). $183.00 (U.S.A.), $188.00 (Int’L), 694 ppt 26 Contents of Volume 43 of Annual Review of Phytopathology: Being at the ls Place, at the Right Time, for the Right Reasons—Plant Pathology, Robert K. Webster Kenneth Frank Baker—Pioneer Leader in Plant Pathology, R. James Cook Replication of i and Ilarviruses: Role es Goal Protein, John F Bol Resistance of Cotton Towards Xanth rv. malvacearum, E. Delannoy, B. R. Lyon, P. mey, A. Jalloul, J. F eee L. ee allen M. Fss senberg, M. Nicole Plant etre A Threat to Global Food Security, Richard N. Strange, Peter R. Scott Vi ee Viroid-Host nn ae tions, Ricardo Flores, Carmen Hernandez, A. Emilio Martinez de Alba, rio Jose-At io D os pasate [Pl Margery L. Daughtrey, D. Michael Benson A lth Jrnamental I tants, ce ero antH The Biology of ite iifesl ans . Is Center of Origin, Niklaus J. Griinwald, Wilbert G. Flier Plant Pathology and RNAi: A Brief History, John A. Lindbo, William G. Doughtery nine Mechanisms of Defense Against Biotrophic and Necrotrophic ae ns, Jane G Lipids, Lipases, and Lipid-Modifying Enzymes in Plant Disease Resistance, Jyoti Pathogen Testing and Certification oy Vitis and Prunus Species, Adib ene aa K. Uyemoto, li lazebrook Deborah A. Golino, Giovanni P M Mechanisms of Fungal Speciation, a . Kohn Phytophthora Ramorum: Integrative Research and Management of an Emerging Pathogen in Califor- nia and Oregon Forests, David M. Rizzo, Matteo Garbelotto, Everett M. Hansen nd] ] tati Rj t ] D is J Exploiting Chinks in the Plant’s Armor: Evolution and a of Geminiviruses, Maria R. Rojas, Charles Hagen, William J. Lucas, Robert L. Gilber Molecular Interactions Between Tomato and the Leaf ae Pathogen Cladosporium fulvum, Susana Rivas, Colwyn M. Thomas oe of ey eee in Filamentous Fungi, Jae-Hyuk Yu, Nancy Keller z ‘tions, Anna E. Whitfield, Diane E. Ullman, Thomas L German Tospovirus Hemipterans as Plant Sen Isgouhi Kaloshian, Linda L. Walling RNA Silencing in Productive Virus Infections, Robin MacDiarmid Signal Crosstalk and Induced Resistance: Straddling the Line Between Cost and Benefit, Richard M. Bostock Genetics of Plant Virus Resistance, Byoung-Cheorl Kang, Inhwa Yeam, Molly M. Jahn Biology of Plant Rhabdoviruses, Andrew O. Jackson, Ralf G. Dietzgen, Michael M. Goodin, Jennifer N. : & ragg, Min Den SIDA 21(4): 2184. 2005 ICBN CLARIFICATION NEEDED: USE OF RANKS Guy L.Nesom and Barney Lipscomb Botanical Research Institute of Texas 509 Pecan Stre Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT The current (2000) International Code of Botanical Nomenclature is open to divergent interpreta- tion regarding the use of ranks. Article 4.1 outlines secondary ranks to be used between the principal ranks of family ane species and below Species tle 4.2 states Stliat ranks ao by “sub-” (termed ee rank, immediately subsidiary to principal or second- ary eae are used to increase the number of anke to a “greater number” than formed in 4.1. Some axonomists, in contrast, apparently Be these Articles sug} that tertiary rams Dey be used cet reference to secondary ranks in species without varieties or forms). Vinee fevmularions are offered for Articles 4 and 5 that may more clearly express the intent of the Code: Formutation | if the intent is to mandate that tertiary ranks between family and species, and below species, be used a in conjunction with sec- ondary ranks, ForMULATION 2 if the intent is that tertiary ranks may be used without reference to secondary ranks RESUMEN El actual Codigo Internacional de Nomenclatura Botanica (2000) esta abierto a interpretaciones diversas respecto al uso de los rangos. El Articulo 4.1 esboza los rangos secundarios para usar entre los rangos principales de familia y especie, y especie e inferiores. El Articulo 4.2 establece que los rangos con el prefijo “sub-” (Ilamados aqui rango “terciario”, inmediatamente siguientes en secuencia 1 Pe Meee ee a ; , 6 E rE a 6 aun “numero mas grande” que los formados en 4.1. Algunos taxonomos, por el contrario, interpretan aparentemente estos Articulos de modo que los rangos terciarios pueden usarse sin referencia a los rangos secundarios (ej. subgénero en un género sin secciones 0 series; subespecies en especies sin variedades 0 formas). Se ofrecen formulaciones alternativas para los Articulos 4 y 5 que pueden expresar mas claramente la intencion del wee cone ILACION 1 si la intencién es *s de ey que los rangos terciarios entre familia y especie, pecie, } g } : f ; yi secundarios, FoRMULACION 2 si la intencion es que | g iarios p los rangos secundarios. The Articles of the International Code of Botanical Nomenclature (Greuter et al. 2000, the “Saint Louis Code”) are “mandatory” rules (Preface, p. vii), and they are generally carefully and rigorously followed by taxonomic botanists. Such nomenclatural prescriptions are intended to provide a stable method of nam- ing and to avoid creation of superfluous names. Valid publication must be in accordance with the Articles. e 2000 Code is open to divergent interpretation regarding the use of ranks and associated implications for valid nomenclatural practice. As Articles 4] and 4.2 are written, ranks in 4.2 (i.e., ranks in addition to those in 3.1 and 4.1) SIDA 21(4): 2185-2191. 2005 2186 BRIT.ORG/SIDA 21(4) are used ina classification after associated ranks in 3.1 and 4.1 are used, ie., rank subgenus is used in a genus after there are named sections and/or series in the classification; similarly, rank subspecies is used in a species in which varieties and/or forms already are in use. Current practice, however, is inconsistent re- garding which ranks must be used and which ranks are optional. We place the following observations and suggestions on record with the hope that they may lead to clarification of this part of the Code. Rules pertaining to sequence and relative order of ranks are found prima- rily in Articles 3, 4,and 5. A closely related pair of these rules—Articles 4.1 and 4.2—is the focal point of apparent ambiguity. Article 4.1. “The secondary ranks of taxa in descending sequence are tribe (tribus) between family and genus, section (sectio) and series (series) between genus and species, and variety (varietas) and form (forma) below species.” — Article 4.2. “If a greater number of ranks of taxa is desired, the terms for these are made by adding the prefix sub- to the terms denoting the principal or secondary ranks. A plant may thus be assigned to taxa of the following ranks (in descending sequence): regnum, subregnum, divisio or phylum, subdivisioorsubphylum, classis, subclassis, ordo, subordo, familia, subfamilia, tribus, subtribus, genus, subgenus, sectio, subsectio, series, subseries, species, subspecies, varietds, sub va rietas, fo rma, subforma.” [bold added] Article 4.1 outlines secondary ranks to be used below principal ranks. Article 4.2 states that ranks prefixed by “sub” and immediately subsidiary in sequence to principal or secondary ranks are used to increase the number of ranks (pre- sumably for a “greater number” than formed in 4.1). Recommendation 26A.2 appears to be consistent with the intent of the ICBN in using the “sub” ranks in conjunction with ranks provided in 3.1 and 4.1: Recommendation 26A.2 “A subspecies not including the type of the correct name of the species should, where there is no obstacle under the rules, be givena name with the same final epithet and type as a name of one of its subordinate varieties.” Article 3 specifies the principal ranks “in descending sequence” as dom, division or phylum, class, order, family, genus, and species. Articles 4.1 and 4.2 deal with secondary ranks (4.1) and “tertiary ranks” (4.2—those pre- fixed by “sub;” although the phrase “tertiary rank” is not used by the ICBN, it is a useful one and apparently consistent with the intent of the Code). Article 5.1 emphatically fixes the relative order of ranks. ~— — King- Article 5.1. “The relative order of the ranks specified in Art. 3. and 4 must not be altered (see Art. 33.7 and 33.8).” Examples in clarification in 33,7 indicate that 1) principal ranks must be assigned in relative order (e.g., species may not con- tain genera); NECAM AND TIDCCANR USE OF RANKS 2187 2) secondary ranks must be used within the principal rank to which they are subsidiary (e.g., section must be used within the rank of genus); and 3) asecondary rank can be subsidiary only toa secondary rank earlier in rela- tive order (e.g., forms cannot be divided into varieties). Based on the 4.2 sequence and relative order of ranks, although not explicitly given in example by the Code, varieties cannot be divided into subspecies. Va- rieties can be clustered within subspecies rank. Changes instituted in the 1994 Code The structure of Articles 4.1 and 4.2 in the 2000 (Saint Louis) Code was first instituted in the 1994 (“Tokyo”) Code (Greuter et al. 1994), which divided Ar- ticle 4.1 of the 1988 Code (Greuter et al. 1988) into two parts (4.1 and 4.2). The newly structured Articles 4.1 and 4.2 remained unchanged in the 2000 Code and were not suggested for modification by the Vienna botanical congress for the forthcoming 2006 Code (Fred Barrie, pers. comm.). The 1988 Code has the following: Article 3.1. “The principal ranks of taxa in ascending sequence are: species (spe- cies), genus (genus), family (familia), order (ordo), class (classis), division (divisio), and kingdom (regnum). Thus, except for some fossil plants (see Art. 3.2), each species is assignable to a genus, each genus toa family, etc.” Article 4.1. “If a greater number of ranks of taxa is required, the terms for these are made either by adding the prefix sub- to the terms denoting the ranks or by the introduction of supplementary terms. A plant may thus by assigned to taxa of the following ranks (in descending sequence): regnum, subregnum, divisio, subdivisio, classis, subclassis, ordo, subordo, familia, subfamilia, tribus, subtribus, genus, subgenus, sectio, subsectio, series, subseries, species, subspecies, varietas, subvarietas, forma, subforma.” No distinction in the 1988 Code was made among ranks below principal ranks. The phrase “secondary ranks” (referred to in the 1988 Code as “supplemen- tary terms”) was first introduced in the 1994 Code. The two proposals for change relating to 1988 Article 4.1 (Silva 1993; Greuter @ McNeill 1993) were rejected by general vote (McNeill 1993) but were referred to the Editorial Committee, which adopted them in slightly modified form (Greuter, McNeill, & Barrie 1993). In the original proposal by Silva for modification of Article 4.1 1993, p. 186), identification of secondary ranks was done with the intention, at least in part, that “proliferation of ranks by use of a prefix should be restricted to... prin- cipal and secondary ranks.” In discussion of the proposals at the nomenclature sessions (prior to the Editorial Committee meeting), Rapporteur-général Greuter noted the following, regarding what was to become Article 4.2:“What Silva had attempted, and perhaps partly achieved, was to bring a coherent logic into the hierarchy of ranks—where hierarchy meant, not the taxonomic hierarchy but a classification of ranks by their importance” (Greuter et al. 1993, p. 40). 2188 BRIT.ORG/SIDA 21(4) Difference in Interpretation of the 2000 Code The wording of Articles 4.1 and 4.2 in the 2000 Code directly implies that ter- tiary ranks are used between family and genus, between genus and species, and below species (secondary ranks are available in each area) only after use of an immediately preceding secondary rank (in descending sequence). Between kingdom and family (where secondary ranks do not exist), tertiary ranks im- mediately follow principal ranks. None of the Code notes or examples perti- nent to 4.1, 4.2, or 5.1 (and 33.7 or 33.8, as pointed to by 5.1), however, provides explicit clarification regarding this. In alternative interpretation and in prac- tice, and apparently contrary to the 2000 Code, subfamilies are used ina classi- fication without use of tribes, subgenera without sections, and sul species with- out varieties. This difference in interpretation may exist because of a disparity between Articles 4.1 and 4.2: tertiary ranks precede the secondary ranks in “relative or- der” and “descending sequence” but because tertiary ranks are used to increase the number of ranks beyond those provided by secondary ranks, secondary ranks precede tertiary ranks in order of use. If the Code does not intend to man- date this order of use in ranks, then the separation of 4.1 from 4.2, coupled with the wording of +.2 (Ifa greater number of ranks of taxa is desired”), is stated incorrectly or at least is misleading. = Alternate formulations for clarification In clarification of the problem discussed here, modifications of the 2000 (Saint Louis) Code are suggested. Two alternate formulations provide a contrast be- tween what appear to be different interpretations of the Code. Article 3.1 (un- modified from the 2000 code) is included within both alternatives. If modifications are necessary for the 2012 ICBN in regard to points con- sidered here, a formal proposal in Taxon will be required. Because of the expanse of time between now and the next Code version, because we are not taking a position of advocacy, and because what the Editorial Committee intended in 1993 is not clear to us, we offer this commentary asa beginning point of discussion. FORMULATION | If the intent of Articles 4 and 5is to mandate that tertiary ranks be used only in conjunction with secondary ranks between family and genus, between genus and species, and below species, then we suggest that the following better ex- press the intent of the Code. Article 3.1 (unmodified from 2000 code). The principal ranks of taxa in descending sequence are: kingdom (regnum), division or phylum (divisio, phy- lum), class (classis), order (ordo), family (familia), genus (genus), and species (species). Thus, each species is assignable to a genus, each genus toa family, etc. Article 4.1. A plant may be assigned to taxa of the following ranks (in USE OF RANKS 2189 descending sequence): regnum, subregnum, divisio or phylum, subdivisio or subphylum, classis, subclassis, ordo, subordo, familia, subfamilia, tribus, subtribus, genus, subgenus, sectio, subsectio, series, subseries, species, subspe- cies, varietas, subvarietas, forma, subforma. Article 4.2. Secondary ranks of taxa are tribe (tribus) between family and genus, section (sectio) and series (series) between genus and species, and vari- ety (varietas) and form (forma) below species. Terms for tertiary ranks of taxa are made by adding the prefix “sub-” to the terms denoting the principal and/ or secondary ranks. Tertiary ranks are added if a greater number of ranks of taxa (beyond secondary ranks) is desired. Article 4.3. Further ranks may also be intercalated or added if a greater number of ranks of taxa (beyond tertiary ranks) is desired , provided that confu- sion or error is not thereby introduced. Article 5.1. The relative order of the ranks specified in Arts. 3 and 4 must not be altered (see Art. 33.7 [the examples following 33.7 would be better placed here] and 33.8). The sequence of use of ranks between the principal ranks fam- ily and species and below species is secondary (in descending sequence), then tertiary. Tertiary ranks follow the principal and/or secondary ranks from which they are derived. Any of the tertiary ranks may be omitted without altering the relative order; the secondary ranks series and forma may be omitted without altering the relative order. Note a.—Use of the rank of tribe precedes use of subfamily; use of the rank of section precedes use of subgenus or series; use of the rank of variety pre- cedes use of subspecies or form. Note b—A genus may be included in a family without reference to a tribe or toa subfamily (omission of one or both of the ranks between genus and fam- ily does not affect the relative order of ranks). If Formulation 1 were adopted, a date might be set beyond which the rules would apply—in order to avoid chaotic invalidity of names at tertiary rank. Or, proposal of a name at tertiary rank prior to an appropriate name at secondary rank might be set to automatically establish the secondary rank. FORMULATION 2 If the intent of Articles 4 and Sis that tertiary ranks may be used without refer- ence to secondary ranks, then we suggest that the following better express the intent of the Code. This formulation returns to the less restrictive nature of the 1988 Code, in which no distinction in use was made between secondary and tertiary ranks (as they are termed here). Here, there is no problem with subspe- cies as sole infraspecific rank within a classification or with subgenus as sole subdivision of a genus. Article 3.1 (unmodified from 2000 code). The principal ranks of taxa in descending sequence are: kingdom (regnum), division or phylum (divisio, 2190 BRIT.ORG/SIDA 21(4) phylum), class (classis), order (ordo), family (familia), genus (genus), and species (species). Thus, each species is assignable to a genus, each genus to a family, etc. Article 4.1 (same as in FORMULATION 1). A plant may be assigned to taxa of the following ranks (in descending sequence): regnum, subregnum, divisio or phylum, subdivisio or subphylum, classis, subclassis, ordo, subordo, familia, subfamilia, tribus, subtribus, genus, subgenus, sectio, subsectio, series, subseries, species, subspecies, varietas, subvarietas, forma, subforma. Article 4.2. Secondary ranks of taxa are tribe (tribus) between family and genus, section (sectio) and series (series) between genus and species, and vari- ety (varietas) and form (forma) below species. Terms for tertiary ranks of taxa are made by adding the prefix “sub-” to the terms denoting the principal and/ or secondary ranks. Article 4.3 (same as in FORMULATION 1). Further ranks may also be interca- lated or added if a greater number of ranks of taxa (beyond tertiary ranks) is desired, provided that confusion or error is not thereby introduced. Article 5.1. The relative order of the ranks specified in Art. 3 and 4 must not be altered (see Art. 33.7 [the examples following 33.7 would be better placed here]and 33.8). Any of the secondary or tertiary ranks may be omitted without altering the relative order, but use of tertiary ranks must follow the principal or secondary ranks from which they are derived. Note a.—A genus may be included in a family without reference to a tribe or toa subfamily (omission of one or both of the ranks between genus and fam- ily does not affect the relative order of ranks); then rank of subgenus may be used within a genus without reference to sections; the rank of subspecies may be used within a species without reference to varieties. Ranks of Taxa in Relation to Biology The rank of subspecies sometimes is said to apply to a taxon more “species- like” than a variety and for this reason should precede “variety” in relative or- der of rank. Intraspecific population systems, however, like species themselves, vary continuously in degree of differentiation and reproductive isolation, and if varieties and subspecies both are treated as morpho-geographic taxa, then a biological distinction between the two ranks is arbitrary. We agree with Fred Barrie (pers. comm.) that the ICBN “legislates the names and relative order of ranks, not the taxonomic concepts attached toa given rank nor the [biological] conditions under which it is appropriate to use one over another.” The discus- sion here of the ICBN structure and intent are detached from considerations of the importance or biological significance of ranks. ACKNOWLEDGMENTS Preparation of this discussion was precipitated by Ken Chambers, who brought to our attention another Code issue (included in an early version of the manu- USE OF RANKS 2191 script)—that issue is less complex and apparently will be resolved by the ICBN Editorial Committee before publication of the Vienna Code. We appreciate the comments of Tom Lammers, Gerry Moore, Rich Rabeler, Dick Wunderlin, and especially those of Fred Barrie (early manuscript version) and John Strother, which considerably sharpened the clarity of concepts and presentation. REFERENCES Greuter, W. et al. (eds.). 1988. International code of botanical nomenclature. Regnum Veg. 118. Koeltz Scientific Books, Konigstein, Germany. GreuTer, W. et al. (eds.). 1994. International code of botanical nomenclature (Tokyo Code). Regnum Veg. 131. Koeltz Scientific Books, Konigstein, Germany. GreuTer, W. et al. (eds.). 2000. International code of botanical nomenclature (Saint Louis Code). Regnum Veg. 138. Koeltz Scientific Books, KOnigstein, Germany. Greuter, W. and J. McNeil. 1993. Synopsis of proposals on botanical nomenclature—To- kyo 1993. A review of the proposals concerning the International Code of Botanical Nomenclature submitted to the XV International Botanical Congress. Taxon 42: 191-271) GreuTer, W., J.McNEILL, and FR. Barrie. 1993.Report on botanical nomenclature—Yokohama 1993. Englera 14:1-265. McNeitt, J. 1993. Preliminary mail vote and report of Congress action on nomenclatural proposals. Taxon 42:907-922. Siva, PC. 1993.(290-320) Thirty-one proposals mainly concerning editorial matters. Taxon 42:185-190. 2192 BRIT.ORG/SIDA 21(4) BOOK NOTICES ELIZABETH Losos and Ecsert GILES LEIGH, JR. (eds.). 2004. Tropical Forest Diversity and Dynamism: Findings from a Large-Scale Plot Network. (ISBN 0-226- 49346-6, pbk.). The University of Chicago Press, 1427 E. 60" Street, Chicago, IL 60637, U.S.A. (Orders: 1-800-621-2736, 773-660-2235 fax, wwwypress. uchicago.edu). $38.00, 645 pp., b/w f come graphs, 6" x 9". }: The book, Tropical Forest Diversity and Dynamism, i parts: 1) Introduction; 2) The Whole is Greater Than the Sum of the Plots; 3) Habitat Specialization and Species ae in Forest Dynamics Plots; 4) Local Variation in Canopy Disturbance and Soil Structure; 5) The Diver- sity of Tropical Trees: ha aie 6) The Diversity of Tropical Trees: The Role of Pest on 7) = Forest aa) namics Plo ‘This book grew out of a symposium organized by the Center for Tropical Forest Science (CTFS) and held in August 1998 at the Smithsonian Institution in Washington, DC.” Patrick B. Durst, Curis Brown, HENryYLITO D. TAcio, and Mtyuk! IsHikAWA. 2005. In Search of Excellence: Exemplary Forest Management in Asia and the Pacific. (ISBN 974-7946-68-8, pbk.). Food and Agriculture Organization of the United Nations Regional Office for Asia and the Pacific and Regional Community Forestry Training Center for Asia and the Pacific, Bangkok, Thailand. (Orders: Patrick B. Durst, Senior Forestry Officer, FAO Regional Office for Asia and the Pacific, 39 Phra Atit Road, Bangkok 10200, Thailand; 66-2- 697-4000, 66-2-697-4445 fax, email Patrick.Durst@fao.org). Price not given, 404 pp., color photos, graphs, maps, 7" X 10" “The pubbestion eee hts a abil of nian eenent approaches that have proven particularly = llenges. Thus, it reaches out to foresters, policy-makers, plan- ners and anyone interested in the future of forestry in Asia and the Pacific.” SIDA 21(4): 2192. 2005 HERBARIUM ACRONYM CORRECTION FOR IPOMOEA SEAANIA (CONVOLVULACEAE) HOLOTYPE: SIDA 21(3). 2005 Richard Felger Daniel F. Austin Drylands Institute Arizona-Sonora Desert Museum PMB 405, 2509 N. Campbell Ave. 2021 N. Kinney Road Tucson, Arizona 85719, U.S.A. Tucson, Arizona 85743, U.S.A. In a recent paper we described Ipomoea seaania as a new arborescent morning glory endemic to southern Sonora (Felger and Austin 2005). On page 1296 we list the location of the holotype as “UA,” the informal local Tucson “acronym” for the University of Arizona. The correct herbarium designation is ARIZ; that is where the holotype is deposited. The line should read: “Felger with Robert S. Devine 85-301 (HoLotyee: ARIZ.” REFERENCE Feicer, R.S.and D.F. Austin. 2005. /pomoea seaania, a new species of Convolvulaceae from Sonora, Mexico. Sida 21:1293-1303. SIDA 21(4): 2193. 2005 2194 BRIT.ORG/SIDA 21(4) BOOK NOTICES University of California Press James Lazett. 2005. Island: Fact and Theory in Nature. (ISBN 0-520-23277-1, hbk.). University of California Press, Berkeley, CA 94704, U.S.A (Orders: Califor- nia Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A., 609-883-1759, 609-883-7413 fax: ww W.UCpI ess.edu). $49.95, 382 pp. 40 color plates, b/w figures, 7" X 10". F rom the main r—“He presenise com pelling evidence that high levels of natural biodiversity under- | lly with espe to high-level consumers such as humans.” Island: Fact and Theory in Natine is broken up int shapters: 1) Ways of Looking at Diver- sity; 2) How to Count Snakes—and Other Things; 3) Putting es sae 4) Great Guania and the Isles of Yesteryear, 5) The Cast, 6) Nature and Man. The author notes in the introduction that two major themes run through the book. “They are noel! its mRUSIE os its bl an oe ase makes for stability, and that is good for us...Second, be war ‘ol las.” rf oO Puitip W. Runpet and Rosert GustaFson. 2005. Introduction to the Plant Life of Southern California: Coast to Foothills. (ISBN 0-520-23616-5, pbk.). Uni- versity of California Press, Berkeley, CA 94704, U.S.A (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, US.A., 609-883-1759, 609-883-7413 fax; www.ucpress.edu). $18.95, 316 pp., 311 color photos, 4 1/2" x 71/4" Highlights from the back cover—‘Features 327 color ey vividl ” illustrating the major plant communities, Describes more than 300 plant species; C f Santa Barbara, Ventura, Los Angeles, Orange, San Diego, western Riverside, and San Bernardino, as w ‘well as the Channel Islands; bucludes a list of parks and other public areas for viewing southern California's plant communities.” mmon and Latin names are provided for all plants. Exce — lent color photographs illustrate the plane communities discussed as well as species of plants that dominate the communities. SIDA 21(4): 2194, 2005 MISCELLANEOUS CHROMOSOME NUMBER REPORTS FOR POA (POACEAE) IN NORTH AMERICA Robert J.Soreng Department of Botany eg Museum of Natural History mithsonian Institution ee DC 20013-7012, U.S.A. sorengr@si.edu ABSTRACT The following 66 eae numbers and vouchers for 24 species of Poa L. are reported from Canada, México, and the US.A. P. abbreviata subsp. pattersonti, 2n = 42,2n=42,2n=42;P.alpina,2n = 28+II, One S2 O50) 2n= a 2n=42,2n =56; P. arctica subsp. aperta, 2n = 98+I; subsp. arctica, 2n = 56; subsp. arctica (longipila form), 2n = 56-59, 2n = 80, 2n = 88; P atropurpurea, 2n = 28; P. bigelovii, 2n = 28+I; P cusickii subsp. pallida, 2n = 56+II; P fendleriana subsp. albescens, 2n = 28+II, 2n = 56; subsp. fendleriana, 2n = 56 (4X), 2n = 59, 2n = 58-60, 2n = 58-64; subsp. longiligula, 2n = 56, 2nca.56; P. glauca subsp. glauca, 2n = 56, 2n = 56-58; subsp. rupicola, 2n = 48, 2n = 48-50, 2n = 54-56, 2n ca. 100; P interior, 2n = 42, P laxa subsp. banffiana, 2n = 84; P. leptocoma 2n = 42 (2X); P. lettermanii, 2n = 14; nine 2n = 42; P. nervosa, 2n = 28; P occidentalis, 2n = 14 (2X), 2n = 28; P. paucispicula, 2n = 42; P. piperi, 2n = 28; P. pratensis subsp. alpigena, 2n = 56+III; P. reflexa, 2n = 28 (4X); P. secunda subsp. aa 2n = 63; subsp. secunda, 2n = 84-88+IL P sierrae, 2nca. 58: P. strictiramea, 2n = 28-29, 2n= 29+11, P. supina c.v. SUPERNOVA, 2n = 14; P. tracyi, 2n = 28 a 2n = 28+]; P. unilateralis subsp. pachypholis, 2n = 42; and P.unilateralis subsp. unilateralis, 2n = 84. RESUMEN c y testigos de 24 especies de Poa L. de Canada, México, y E.E.U.U:: P. abbreviata subsp. i la 2n = 42; P. alpina, 2n = 28+II, 2n = 32+] (2X), 2n = 4041, 2n = 42, 2n= 56; P. arctica pase aperta, 2n = 98+; subsp. arctica, 2n = 56; subsp. arctica (forma longipila), 2n = 56- 59, 80, 2n 8; eae eo P. bigelovii, 2n = 28+1; P.cusickii subsp. pallida, 2n =56+I; esas sien albescens, 2n = 28+H1, 2n ee subsp. fendleriana, 2n = 56 (4x), 2n = 59, 2n =58- 8-6 4; subsp. longiligula, 2n = 56, 2n ca. 56; P glauca subsp. glauca, 2n = 56, 2n = 56-58; subsp. rpc 2n= 48, 2n = 48-50, 2n = 54-56, 2nca. ce P interior, 2n = 42; P. ae banffiana, 2n = P. leptocoma 2n = 42 (2X), P lettermanii, 2n = 14; P. napensis, 2n = 42; P. nervosa, 2n = 28; P. oe 2n=14 (2x); P. paucispicula, 2n = 42; P. piperi, 2n = 28; P. ratnssubsp ae oe = 56+II1; P reflexa, 2n = 28 (4%); P secunda i juncifolia, 2n = 63; subsp. secunda 4-88+II1; P sierrae, 2nca. 58; P strictiramea, 2n = 28-29, 2n = 29+I], P supina c.v. SUPER eee Ma ee cyi, 2n=28 (5X), 2n=28+1; P unilateralis subsp. pachypholis, 2n = 42; y P. unilateralis subsp. unilateralis, 2n = 84. The Flora of North America North of México (Morin et al. 1993) editorial policy requires that chromosome numbers be independently published prior to being reported in the treatments. Therefore, | am reporting a series of chromosome counts here for Poa that are unreported or only mentioned with partial voucher SIDA 21(4): 2195 — 2203. 2005 Taste 1. Voucher information for chromosome counts in the genus Poa that are new or mentioned with no or only partial voucher information in Soreng (1985, 1990, 1991a, 1991b, 1993, 1998) and Soreng and Hatch (1983). R/S = RJ. Soreng, RWS = R. W Spellenberg. Taxon County & State Specific location, date, collection no. & herbarium Chromosome no. (2n), and notes Poa abbreviata subsp. pattersonii (Vasey) A.Léve, D. Love & B.M. Kapoor Poa alpina L. Poa arctica subsp. aperta (Scribn. & Merr.) Soreng U.S.A. Colorado: Colorado: Montana: CANADA. Alberta: Alberta: U.S.A. Colorado: Colorado: Wyoming: Wyoming: U.S.A. Colorado: Clear Cr.Co.: Rocky Mts., Mt. Evens top, SSE of Georgetown ca.13 km,3 Aug 1984, RUS, R. Bayer, M. Dunford & G.L. Stebbins 2555 (US) Summit/Park Co. boundary, Rocky Mts., Tenmile Range, North Star Mt., Hoosier Ridge W of Hoosier Pass, 2 Aug 1984, RJS,R. Bayer, M. Dunford & GL. Stebbins 2548 Deer Lodge Co.: Anaconda-Pintlar Wilderness, Mt. Tiny, above Storm Lk.,6 Aug 1980, RJS & RWS 1165-2 (US) US — Banff N.P, ca. 100 km N of Banff on hwy 93,E slopes of Mt. Peyto, S of Peyto Lk.,N of Bow Lk., 28 Jul 1980, RJS & RWS 1018 (US) Plateau Mt., between Mt. Livingston and Mt. Burke, ca. 67 km due N of Colman, 2 Aug 1980, R/S & RWS 1705 (US) Sagauche Co.: San Luis Mts.,N slope of Baldy Chato, off Big Meadow Rd. FR 790,17 Aug 1980, RJS & RWS 1406-a (US) Sagauche Co.: San Luis Mts.,N slope of Baldy Chato, off Big Meadow Rd. FR 790, 17 Aug 1980, RJS & RWS 1406-b (US) Park Co.: Beartooth Pass, E summit, 8 Aug 1980, R/S & RWS 1213-5 (US) Sublett Co.: Little Sheep Mt., NW of N end of Green Lakes ca.6 km, 10 Aug 1980, RJS & RWS 1290 (US) Sagauche Co.:San Luis Mts.,N slope of Baldy Chato, off Big Meadow Rd., FR 790, 17 Aug 1980, RJS & RWS 1412-a (US) ) 42 (Soreng 1991b, with partial voucher) information 42 (Soreng 1991b, with partial voucher information) nN = 21, from pollen division (Soreng 1991b, with partial voucher information) 42 (new) 56 (new) 28+1I (new) 32+1 (new) 40+I (new) 32+1 (new) 99 (Soreng 1985, without voucher) 9617 (p)L? VaIS/DHO'LINE Tas_e 1. (continued) Taxon County Specific location, date, collection no. Chromosome no. (2n), & State & herbarium and notes Poa arctica R.Br. CANADA. Kananaskis Prov.P, at Kananaskis Summit (Highwood Pass), 56 (new) subsp. arctica Alberta: near Mt. Arethusa ca.67 km S of Seebe and Hwy 1,on Hwy 40, on W side, 31 Jul 1980, R/S & RWS 1094 (US) SA. Pitkin Co. (W of Lake Co. line?), Rocky Mts., Sawatch Range, 88 (new) Colorado: Independence Pass, 15 Aug 1980, RJS & RWS 1397 (US) Montana: Deer Lodge Co.: Anaconda-Pintlar Wilderness, Mt. Tiny, 80 (new) above Storm Lk.,6 Aug 1980, R/S & RWS 1180 (US) Montana: Glacier Co.: Glacier N.P, Pigan Pass, 4 Aug 1980, RUS & 56-59 (new) RWS 1142 (US) Poa atropurpurea Scrion. U.S.A. San Bernardino Co.: Baldwin Lake, 1985, R/S 2632 (US) 8 (Soreng 1993, without California: voucher) Poa bigelovii Vasey & U.S.A. New Lincoln Co.:White Mts., NE of Sierra Blanca, below 28+1 (Soreng 1985, without Scribn. Mexico: Monjeau L.O., 16 Jun 1981, R/S 1584t (US) voucher) Poa cusickii subsp. pallida Park Co.:NE of Gardner 10 km, E of Jardine, Jun 08 1984, 56+ll (Soreng 1991a, with Soreng Montana: RIS 2453-a (US) voucher, but location incomplete and number erroneously reported as RJS 2456) Poa fendleriana subsp. MEXICO. Sierra Madre Occidental, Creel, near air strip, 15 Apr 1984, 56 (new) albescens (Hitchc.) Soreng Chihuahua: RIJS & RWS 2309 (US) Sonora: 5 km NW of Cananea on microondas road N from road to 28+I1, with inversion bridge Sonora, 19 Mar 1982,RJS & RWS 1780-5 (US) & fragments poreng 1985, without voucher ) Poa fendleriana (Steud.) U.S.A. New Catron Co.: Sheridan Gulch, 21 May 1983, R/S & D. 56, with multivalents (new) Vasey subsp. fendleriana Mexico: Ward 2125 (US) Mexico: Dona Ana Co.: Organ Mts., W side, below and E of Baylor 56 (new) Pk., 10 Feb 1984, RJS & R. Neilson 2190-b (US) VOd YO4 SLUOd3Y IWOSOWOYH) ‘NINOS L612 Taste 1. (continued) Taxon Coun Specific location, date, collection no. Chromosome no. (2n), & State & herbarium and notes Mexico: Grant Co.: Black Range, 19 km NW of Mimbres, D. Ward 58-64 (new) 81-04 (NMC) Mexico: Lincoln Co.:White Mts., Montgomery Biological Research 56 (new) Station, 8 km N of Ruidoso, 18 Apr 1981, RJS 71580 (US) Mexico: Sandoval Co.: Sandia Mts.,W base, Juan Tabo Picnic Area, 58-60 (new) NE of Albuquerque, 6 Jun 1983, R/S & RWS 2172 (US) Mexico: Socorro Co.: San Mateo Mts.,21 Mar 1984, R/S 2303 56 (new) [no voucher] MEXICO. Sierra Madre Occidental, 7 km E of Tomachic, 14 Apr 1984, n= 2841, mitotic, pistillate Chihuahua: RIS 2306 (US) plant (new) Poa fendleriana subsp. Apache Co.: Chuska Mts., 6.7 km NE of Lukachukai, on 56 (new) longiligula (Scribn.& Arizona: BIA-13,9 Jun 1983, RIS & RWS 2177 (US) T.A. Williams) Soreng Wyoming: Park Co.. Mammoth Hot Springs, 08 Jun 1984, R/S 2454 (US) N Ca, 28, mitotic (new) Poa glauca Vahl subsp. CANADA Plateau Mt., between Mt. Livingston and Mt. Burke, 67 km 56-58, meiosis irregular Alberta: due N of Colman, 1 Aug 1980, R/S & RWS 1098-3 (US) (new) U.S.A. New Taos Co.:Wheeler Peak, ridge 3.3 km N of peak, 0.4 km S of 56, multivalents and laggers Mexico: Frazer Mt., 19 Aug 1980, RJS & RWS 1454-1 (US) common (new) Poa glauca subsp. rupicola U.S.A. Pitkin Co.: Rocky Mts., Sawatch Range, Independence Pass, Ca. 100 (new) (Nash) W.A.Weber Colorado: 15 Aug 1980, RIS & RWS 1372-18 (US) yoming: Park Co.:Clay Butte Look-Out., ca. 2 km W of Beartooth 54-56, multivalents (new) Lk.,8 Aug 1980, RJS & RWS 1221-2 (US) Wyoming: dito, R/S & RWS 1221-5 (US) 48-50, multivalents (new) Wyoming: Sublett Co.: Top of Little Sheep Mt., NW of N end of Green 48 (new) Lakes ca.6.25 km, 10 Aug 1980, R/S & RWS 1299-6 (US) Poa interior Rydb. U.S.A. Sagauche Co.:San Luis Mts.,N slope of Baldy Chato, off Big 42 (new) Colorado: Meadow Rd. FR 790,17 Aug 1980, RJS & RWS 1422-a-3 (US) 8617 (p)LZ VdIs/9¥O'LINA Tasie 1. (continued) Taxon Specific location, date, collection no. & herbarium Chromosome no. (2n), and notes Poa laxa Haenke subsp. banffiana Soreng Poa leptocoma Trin. — Poa lettermanii Vasey Poa napensis Beetle Poa nervosa (Hook.) Vasey s.str. (excluding Poa wheeleri Vasey) Poa occidentalis Vasey Poa paucispicula Scribn. & .Merr Poa piperi Hitchc. Poa pratensis L. subsp. alpigena (Lindm. Hiitonen ae CANADA or U.S.A.: California: USA. Oregon: U.S.A. New Mexico: New Mexico: New Mexico: CANADA. Alberta: California: U.S.A. Alaska: Glacier Co.:Glacier N.P, Pigan Pass, 4 Aug 1980, RUS & RWS 1137 (US) Glacier Co.:Glacier N.P, Pigan Pass, 4 Aug 1980, RJS & RWS 1148-4 (US) Summit Co.: Mt. Murdock E of Bald Mt. Pass, Hwy 150, 12 Aug 1980, RJS & RWS 1347-2 (US) Napa Co.: Calistoga, S end of landing strip W of Lincoln Ave., 27 May 1986, RJS 2926 (US) Marion Co. Silver Cr. Falls S.P, Winter Falls, 6 Jun 1986, RJS 2960 (US) Rio Ariba Co.: SW of Coyote, Puerco C.G.,,ca.33 km NW of Los Alamos, 15 Aug 1978, RJS & S.L. Hatch 48 (US) Otero Co.: Sacramento Mts.,ca. 8.3 km ENE of Cloudcroft S of NM-244 on CR-7, Dec early 1978, RJS 1236 (US) Otero Co.:Cloudcroft, $.L. Hatch-2222 (TAES) Banff N.P, ca. 100 km N of Banff on hwy 93,£ slopes of Mt. Peyto, S of Peyto Lk., N of Bow Lk., 28 Jul 1980, RIS & RWS 1016 (US) Del Norte Co.: Off hwy 199 0.6 km on Patrick Cr.Rd above the Middle Fork of the Smith Rv., 2 Jun 1986, RJS 2950 (US) Nome, Jul 1983, G.L. Stebbins A-3107! (US) 84 (Soreng 19916, with partial voucher information) 42 (Soreng & Hatch 1983) 42 (new) 14 (A. Love, pers.com., letter ca. 1982, reported by, Soreng 1991a, voucher unknown) 42 (Soreng 1991a, with partial voucher and location) 28 (new) 4 (Soreng & Hatch 1983) 8 (Soreng & Hatch 1983) 14 (Soreng & ee 1983, count by S.L 42 (Soreng g faa 983, reported as Poa leptocoma) 28 (Soreng 1990, 1993, without voucher) 56+lll, ee det.as Poa arctica YOd YOd S1YOd7Y IWOSOWOUHD ‘ONIYOS 6617 Taste 1. (continued) Taxon County Specific location, date, collection no. Chromosome no. (2n), & State & herbarium and notes Poa reflexa Vasey & Scribn. U.S.A. New Taos Co.:Wheeler Pk. La Cal Basin, ca. 1.7 km NNW 8 (Soreng & Hatch 1983) Mexico: of peak, 19 Aug 1980, R/S & RWS 1478-4 (US) Utah: Summit Co.: Mt. Murdock E of Bald Mt. Pass, Hwy 150, 8 (Soreng & Hatch 1983) 11 Aug 1980, R/S & RWS 1336 (US) Wyoming: Park Co.: Clay Butte Look Out., ca. 2 km W of Beartooth 28 (new) Lk.8 Aug 1980, R/S & RWS 1227 (US) Wyoming: Sublett Co.:S side of Little Sheep Mt., NW of Green Lakes 28 (Soreng & Hatch 1983) ca.5 km, 10 Aug 1980, R/S & RWS 1260-3 (US) Poa secunda subsp. Lander Co.:Toiyabe Range, E of Austin ca. 13 km on 63 (Soreng 1991b, with juncifolia (Scribn.) Soreng Nevada: Hwy 50,1 Jul 1980, R/S 827 (US) partial voucher information) Poa secunda J.Pres| SA. Glacier Co.:Glacier N.P, Siyeh Pass Trail, 4 Aug 1980, 84-88+Il (Soreng 1991b, subsp. secunda Montana: RIS & RWS 1135 (US) with partial voucher information) Poa sierrae T. Howell U.S.A. Eldorado Co.: Deep Canyon, N. Fork of American River, ca. 58 (new) California: E of Colfax off hwy 80 ca. 2 m,ca.0.8 km NE of river crossing of lowa Hill-ColfaxRd., 30 May 1986, RIS &G.L. Stebbins 2931 (US) Poa Strictiramea Hitche. MEXICO. Sierra Madre Occidental, W of San Jose Babicora, C. El n= 14+I, mitosis (Soreng Chihuahua: Diablo Pass, 2 km W on road to Madera, 13 Apr 1984 1991a, with voucher and RIS & RWS 2304-a (US) partial location) Chihuahua: dito, RJS & RWS 2304-b (US) N= 14-15+ll, mitosis (new) Poa supina Schrad. cv. U.S.A. Cultivated from commercial seed, 2000, R/S & J. n=7, mitosis J. Cayouette, SUPERNOVA Maryland: Cayouette 5950-b (US) unreported) Poa tracyi Vasey U.S.A. New Bernalillo Co.: Sandia Crest, rim N of Tram, 16 Jul 1981, 28 (Soreng & Hatch 1983) Mexico: RJS & K. Gadzia 1642 (US) New Mexico: Colfax Co.: WNW of Raton, Raton City Park, 16 Aug 1978, 28 (Soreng & Hatch 1983) RJS & S.L. Hatch 64 (US) 0072 (pL? VOIS/9¥O'LINa TABLE 1. (continued) Taxon County & State Specific location, date, collection no. & herbarium Chromosome no. (27), and notes Poa unilateralis subsp. n L in . PUCI YH! MOTD (Piper } Soreng Poa unilateralis Scribn. subsp. unilateralis New Mexico: New Mexico: New Mexico: New Mexico: New Mexico: U.S.A. Washington: U.S.A. Oregon: Colfax Co.: NW of Raton, Raton City Park, 31 May 1979, RIJS 266 (US) Colfax Co.: N of Raton, John Mayer's Ranch, down canyon from Raton Pass on side of Bartlet Mesa, E side of US-25, 31 May 1979, RJS 267 (US) Colfax Co.; Raton Ranch, NW of Raton ca. 7.5 km, 3 Jun 1979, RIS 272 (US) Colfax Co.: ie Ranch, NW of Raton ca. 12 km, 3 Jun as RJS 274 (US) ncoln Co. Sierra Blanca, circ below the Peak, 10 Jul 1982, he (US) Pacific Co. waco, RWS & D. Southerland 1522A (NMC) Curry Co.: 3.3 km S of Gold Beach on serpentine road cut, Buena Vista Waysides, 300 ft above the ocean, 22 Jun 1949, J. Clausen 2151 (CAS) 8 (Soreng & Hatch 1983 er 28 (Soreng & Hatch 1983 mo 28 (Soreng & Hatch 1983) 8 (Soreng & Hatch 1983) 28+4+1 (new) 42, R.W.St TT g unt (Soreng 1998, with partial voucher information) 84, as Poa unilateralis, J. Clausen unpublished (Soreng 1991a, with voucher but no location) ¥Od YOd S1Y0d7Y IWOSOWOUH) ‘ONAHOS LOzz 2202 BRIT.ORG/SIDA 21(4) information in Soreng (1985, 1990, 1991a, 1991b, 1993, 1998) and Soreng and Hatch (1983). Table 1 includes full specimen citations and herbaria (acronyms follow- ing Holmgren et al. 1990) where the vouchers are deposited for all of my previ- ous and new reports. My own counts were done between 1978 and 1988. Meth- ods for the chromosome preparations were given in Soreng and Hatch (1983). In addition, vouchers and/or notes of a few counts done by other botanists/ authors that have not been reported previously are included. Jacques Cayouette provided his new chromosome count of P. supina from the recently introduced (in North America) cultivar SUPERNOVA. The count reported in Soreng (1991a) for P lettermanii was mentioned to me ina letter by A. Love, ca. 1982. This re- port is interesting as it raises to three the number of diploid species in the New World (the others are P. occidentalis and P. pseudoabbreviata). Verification of the count for P lettermanii is needed since I only have the correspondence record. | found an unpublished report for P unilateralis subsp. unilateralis of 2n=84 ona herbarium specimen at CAS, the count likely done by the collector of the specimen, geneticist Jens Clausen. Myers (1947) reported a count by Stebbins of 2n = 42, presumably for the typical subspecies (as P unilateralis subsp. pachypholis is rather local, more recently published, and restricted to the coast of NW Oregon and adjacent Washington). Richard W. Spellenberg made a count of 2n = 42 for P unilateralis subsp. pachypholis. Although emphasis has switched away from cytogenetic comparisons of species to DNA analyses in Poa (Gillespie & Soreng 2005; Soreng 1990), it is important to have an understanding of the cytogenetic history of taxa in order to interpret results of other analyses, and to be able to locate vouchers and know where they were collected. Of the 66 counts listed in Table 1, 34 are unreported elsewhere. The base chromosome number in the genus Poa is x = 7, and the counts reported here generally correspond to multiples of seven, but unbal- anced sets of chromosomes were frequently encountered. Roman numerals given after numbers (i.e; 2n = 28+I1) represent unpaired chromosomes in the metaphase or anaphase of meiosis, or unbalanced numbers in mitosis or later stages of meiosis. Although supernumary or B chromosomes have been reported frequently in Poa, no attempt was made to distinguish unbalanced chromotin of this type from fragments resulting from irregular meiosis, etc. Most of the counts reported here conform to numbers reported by other authors for the same taxa. Poa sierrae (2n = 28) is the only taxon reported here for the first time. In taxa with previously reported counts, other than my own, and disregarding the extra chromotin, the only the previously unrecorded numbers in any taxon re- ported here are; 2n = 56 in P.fendleriana subsp. albescens, and 2n = 48, 48-50, 54-56, and ca. lOO in P glauca subsp. rupicola. This work continues to show the pattern in Poa of few diploid taxa, numerous taxa with low, fairly stable tetra- and hexaploid numbers, other taxa with higher eupolyploid series, and taxa — — SORENG, CHROMOSOME REPORTS FOR POA 2203 with eupolyploid peaks connected by dysploid series of numbers (Hiesey & Nobs 1982; Stebbins 1950). ACKNOWLEDGMENTS Most of these chromosome counts were done durning my Masters degree re- search under Stephen L. Hatch and then Kelly W. Allred, and PhD reseach un- der Richard W. Spellenberg, at New Mexico State University. My appreciation is extended to these advisors, to Gerrit Davidse and Lynn J. Gillespie for their helpful reviews of the manuscript, and to Jacques Cayouette for allowing me to report his new chromosome count. REFERENCES Gittespie, L.J. and RJ. Sorenc. 2005. A phylogenetic analysis of the bluegrass genus Poa based on cpDNA restriction sited data. Syst. Bot. 30:84-105. Hiesey, W.M. and M.A. Nos. 1982. Experimental studies on the nature of species VI: Inter- specific hybrid derivatives between facultatively apomictic species of bluegrasses and their responses to contrasting environments. Publications of the Carnegie Institute of Washingon. Vol. 636. Houmeren, PK., N.H. Houmeren, and L.C. Barnett. 1990. Index herbariorum. Part |.:the herbaria of the world. New York Botanical Garden, Bronx. Monin, N.R. et al. (eds.). 1993 forward. Flora of North America north of Mexico.Oxford Uni- versity Press, New York. Myers, W.M. 1947. Cytology and genetics of forage grasses. Bot. Rev. 13:319-421. Soren, RJ. 1985. Poa L.in New Mexico, with a key to middle and southern Rocky Moun- tain species (Poaceae). Great Basin Naturalist 45:395-422. Sorena,R.J.1990.Chloroplast DNA phylogenetics and biogeography in a reticulating group: study in Poa (Poaceae). Amer. J. Bot. 77:1383-1400. Soren, R.J.1991a. Systematics of the Poa L.,group“Epiles” (Poaceae). Syst. Bot. 16:507-528. Sorenc, RJ. 1991b. Notes on new subspecific taxa and hybrids in North American Poa (Poaceae). Phytologia 71:390-413. Sorene, RJ. 1993. Poa L.1n:J.C. Hickman, ed. The Jepson manual: higher plants of California. University of California Press, Berkeley. Pp. 1284-1291. Soren, R.J. 1998. An infrageneric classification for Poa in North America, and other notes on sections, species, and subspecies of Poa, Puccinellia, and Dissanthelium (Poaceae: Poeae). Novon 8:187-202. SorenG, R.J.and S.L. Hatcu. 1983. A comparison of Poa tracyi and Poa occidentalis. Sida 10: 123-14): Steains, G.L. 1950. Variation and evoluton in plants. Columbia Univeristy Press, New York. 2204 BRIT.ORG/SIDA 21(4) Book Notices University of California Press Gary Gricos, Kiki Parscu, and Laurer Savoy. 2005. Living with the Changing Cali- fornia Coast. (ISBN 0-520-24447-8, pbk.). University of California Press, Berkeley, CA 94704, U.S.A (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A., 609-883-1759, 609-883-7413 fax; www.ucpress.edu). $24.95, 540 pp., b/w photographs 6" x 9" d The first edition of this book titled Living with the California Coast, was published in 1985. In the author's words, the first part of the book provides the reader “with some basic ue iuet on how the shoreline works, the processes and hazards that occur here, things t before buying or building, options in hazardous locations, and how policies and me eee influence our response.” In the second part of the book, “the authors and coastal geologists familiar with specific regions de- scribe these individual areas [coastline areas son the Oregon anne to Mexico], including what we know about their geology, hazards, and histories Norman Myers and JeNnirer Kent (eds.). Foreword by Eowarp O. Wi:son. 2005. The New Atlas of Planet Management. (ISBN 0-520-23879-6, pbk.). Uni- versity of California Press, Berkeley, CA 94704, U.S.A (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, US.A., 609-883-1759, 609-883-7413 fax; www.ucpress.edu). $39.95, 304 pp., color photos, graphs, drawings, 91/4" x 12 1/2". Authors’ comments about this book.—‘This is li atlas. It maps and analyses a living planet atacr iag pone in its ssiscee aac one pees, own, See to disrupt and exhaust its life-support em rf And it proposes that we have the chance to redirect our course, er Gaon caretakers of our future. e New Atlas of Planet Management is a first approach to this challenging task. It organizes the mass of available environmental data, statistical predictions, and other conflicting opinions and solutions into a simple, coherent structure. A is divided into seven sections: Land, Oceans, Elements, Evolution, Humankind, Civilization, and Managen of these is considered from three per- apace Potential resources, Crises, and sanagernent alternatives. ritical area o concern and to wel? first, what it has to offer; second, where, how, and wh i i ow we might set about bana things right, by ee a = of alternative pe re than a structure fora Book this analytical for | | management. We hope it will spur the rising global deb f pros] *—Nor man Myers and Jennifer Kent. This book is extremely well-illustrated with extensive captions describing the issues, the re- sources, the crises or even possible solutions. to pli anet SIDA 21(4): 2204. 2005 RELATIONSHIPS BETWEEN PLANT FOLKLORE AND ANTITUMOR ACTIVITY: AN HISTORICAL REVIEW! Richard W. Spjut? World Botanical Associates Bakersfield, California 93380-1145, U.S.A. www.Worldbotanical.com ABSTRACT The National cancers Institute's (NCD necord of plants that have shown significant inhibitory effect 960-1974, was man with species and genera in ete on medicinal folklore, including poisonous plants, to determine whether their percent- ages of active plants were ies icantly greater than those a at random (10.4%). The percent active species in medicinal and/or poisonous references in general were found to be 1.4 to 2.6 times greater, whe ie numbe be diff ferent kinds of medicinal uses appear related to Ss ee data ] of li plants were screened more thoroughly because of their wide- er occurrence. The best coneladont is seen with poisonous plants, Saat medicinal plants that suggest a moderate to effect; their percentages of active species were nearly three (29.3%, anthelmintics) to ae ane (45.7%, arrow and homicidal poisons) greater than plants screened at random. Selection of plants based strictly on use in folk medicine would on! ben- efit new tart ene scene PECeTalns, whereas i in the long-term t effective to all of f plants ay ale: since sue common medicinal species wel be collected irregardless. A era that have not been exhaustively studied, especially to species with econ uses ‘that eae toxicity or are considered poisonous. RESUMEN El registro de plantas del Nanonal CARE. Tsintite (NCD ne hades que ne mostrado un efecto inhibidor significativo en ompararon con a y especies que aparecen en referencias de meena papules incl d oe venenosas, para determinar en que medida \ ignifi Asal rae | = : ic ee | hoger ] a las investigadas al azar (10. 496). E El /o venenosas en general se encontr6 que era del 4 a 2.6 veces mayor, mientras que el numero y diferentes tipos de usos meine. qeancen. relacionados con Gai geograficos de Sale que cmipien indi ue) pl su I i las ae venenosas, incluyendo las plantas 'A summary of the data in this paper was presented at the Society for E ic B Symposium on Plants And Cancer held in Baltimore, August 1975. An alternate cage was ae hed in Cancer earn ees in August 1976 (Spjut & Perdue, Vol. 50, 8:979-985). Left out Medicinal Plants of The Philippines, genera with geographically disjunct uses of meacinal uate and activity according to the tumor systems employed. 2USDA Agricultural ees ae Medicinal Plant Resources Laboratory, Beltsville, MD 20705. The NCI termi- ated their ARS in 1982. Spjut left the USDA in March 1997.World Botanical A Bakersfield, CA 93380-1145. SIDA 21(4): 2205 — 2241. 2005 2206 BRIT.ORG/SIDA 21(4) a ae ] f, -| aT A f. ] CCEILLCTICI r activas nue e cerca de tres i 3%, _antihelminticos) a cuatro veces (45.7%, venenos para flechas y ] q uc [ a seleccion de plantas basada estrictamente en el uso en medicina popular probablemente s seria beneficiosa para los nuevos programas de investigacion, mientras que a largo término, parece tener un costo efectivo mayor la investigacion sistematica de una diversidad de plantas ane eponinis: Ni ve las ee ses comunes pueden colectarse en cualquier parte. Una estrat en géneros que no hayan sido estudiados ephauisnivamnenté y eae en eee con usos sid venenosas. medicinales que indiquen t | INTRODUCTION The USDA Agricultural Research Service (ARS) was a major supplier of plant samples for the National Cancer Institute (NCI) Cancer Chemotherapy Screen- ing Program from 1960-1982. The objective of this program was to identify novel chemical structures produced by plants that would be useful in treatment of cancer. Two major discoveries of novel anticancer drugs from this period were taxol (Wani et al. 1971), isolated from stem-bark of Taxus brevifolia Nutt. (Taxaceae), initially collected in Washington, August 1962, followed discovery of confirmed antitumor activity in KB Cell Culture (KB), July 1964 (NCICPAM, 1977), and camptothecin (Wall et al. 1966), isolated from Camptotheca acuminata Decne. (Nyssaceae), based on fruit samples collected in September 1961 from a USDA Plant Introduction Station in Chico, California, and reported to have confirmed antitumor activity in L-1210 Leukemia (LE), July 1962 (NCI CPAM 1977). Semi-synthetic derivatives of compounds from both species are currently employed to treat various cancers (Cragg et al. 1996). The commer- cial development of these anticancer drugs, however, did not occur until the 1990s. In 1986, the NCI re-developed its biodiversity screening program of natu- ral products (Boyd 1992; Cragg et al. 1996; Newman et al. 2003); however, the acquisition of plant samples for the NCI screen was suspended in 2004. In August 1975, a symposium on “Plants and Cancer” was held in Balti- more, MD at the Annual Meeting of the Society for Economic Botany. The con- tributors included many scientists actively involved in the NCI search of new anticancer drugs from plant products who had agreed, in advance, to provid- ing a research contribution. My assigned study was “Plant Folklore: A Tool for Predicting Sources of Antitumor Activity? Other contributed papers were “Pro- curement of Plant Materials for Antitumor Screening” (Perdue 1976), “Prepara- tion of Plant Extracts for Antitumor Screening” (Statz & Coon 1976), “Bioassay of Plant Extracts for Anticancer Activity” (Abbott 1976), “Isolation and Chemi- cal Characterization of Antitumor Agents from Plants” (Wall et al. 1976), “Types of Anticancer Agents Isolated from Plants” (Hartwell 1976), “Distribution of Anticancer Activity in Higher Plants” (Barclay & Perdue 1976), “Novel Plant- Derived Tumor Inhibitors and Their Mechanisms of Action” (Kupchan 1976), “Pharmacology of Antitumor Agents from Higher Plants” (Sieber et al. 1976), — SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2207 and “Plant Products in Cancer Chemotherapy” (Carter 1976). These and others were published collectively in Cancer Treatment Reports, edited by Robert E. Perdue, Jr, and Jonathan L. Hartwell (Vol. 60, No. 8, 1976). Upon investigating the relationships between antitumor activity and plant folklore, I felt that plants used in folklore were not going to lead to discovery of novel compounds any more than a systematic sampling of the world’s plant diversity based on taxonomy, the approach that had been in practice 14 years. Therefore, in order to show this, the most common medicinal uses of plants, and also poisonous plants, would need to be investigated. During the course of the study, the results on the NCI active species found in literature on medicinal and poisonous plants, in comparisons to those screened at random, raised more questions than could be answered, including the one originally proposed. The Spjut and Perdue (1976) paper excluded much data in another manuscript that had been completed and peer reviewed. After nearly 30 years, the unpublished data still seem relevant to present day studies in ethnobotany and pharmacology, particularly the relationship between antitumor activity and folklore indicating plant toxicity; therefore, this paper will focus on that relationship, including also data from Spjut and Per- due (1976). Another important relationship involves the multiple uses for a large number of widely distributed species; their impact on the apparent correlation between antitumor activity and medicinal folklore will be di d. Addition- ally, Spjut (1985) reviewed the random screen methodology in detail with refer- ence to unpublished data on The Philippine medicinal plants; these data will be presented in this publication. MATERIALS AND METHODS Literature Surveys.—This paper deals with data compiled from literature and the NCI plant screening program prior to 1977. Folklore and plants in this study were limited to literary sources for evaluating medicinal uses and poisonous effects of higher plants in man and animals. Included are plants believed to have medicinal or poisonous properties, and the scientific literature dealing with active chemical agents in confirmed poisonous and medicinal plants. Botani- cal data and the references cited, including the nomenclature of plants, are not updated since this paper was prepared and last reviewed in July 1976; however, in regard to pharmacological data on compounds that were isolated, more re- cent references are provided. Fight compendia on medicinal and poisonous plants were employed to identify which of their genera and species were active in the NCI program: Har- din & Arena (1974), Hartwell (1967-1971), Kingsbury (1964), Krochmal & Krochmal (1973), Quisumbing (195), Train et al. (1957), Webb (1948), and Weiner (1972). One of these, Quisumbing (1951), was further utilized to determine whether a specific medicinal use was more closely correlated with antitumor 2208 BRIT.ORG/SIDA 21(4) activity. Because antitumor activity appeared to correlate with a wide variety of medicinal uses, additional data from Quisumbing (1951) were compiled and analyzed in regard to multiple uses of plants as related to their geographical distribution. Additionally, we (Spjut & Perdue 1976) prepared our own compila- tion on plants used as (1) anthelmintics, (2) fish poisons, and (3) arrow, ordeal and homicidal poi to determine whether there wasa correlation between antitu- mor activity and plant toxicity in contrast to medicinal plants in general. Active species.—An active species is defined as one represented by one or more extracts having shown a significant inhibitory effect in any tumor sys- tem used in the NCI preliminary screen; these were primarily KB Cell Culture (human epidermoid carcinoma of the nasopharynx, KB, 1960-1982), P-388 Leu- kemia (PS, 1968-82), Lewis Lung Carcinoma (LL, 1962-66), Walker Carcinoma 256 (WA, 1966-69), Sarcoma 180 (SA, 1956-62), Adenocat cinoma 755 (CA, 1956- 62) and L-1210 Leukemia (LE, 1956-71) (Abbott 1976; Geran et al. 1972: Hartwell 1976; Suffness & Douros 1979). The NCI provided a print-out of their active spe- cies for this study; additionally, another printout indicating tumor systems for the confirmed active species was consulted (NCI CPAM 1977) Active agents have included a broad spectrum of compounds (Hartwell 1976), some of which were precluded from further screening (e.g., tannins, phy- tosterols) by changes made in the extraction procedure and tumor assays (Hartwell 1976); thus, the NCI screen evolved to become more selective in iden- tifying active candidates for drug development by eliminating classes of com- pounds not considered useful for treating cancer (Hartwell & Abbott 1969). During the 1960s, tannins—in aqueous extracts from a wide variety of plants— were frequently active in WA, but also in CA, LL and SA tumors; a total of 164 species, representing 7.7% of all active species (2,127) in this study were tannin actives (Barclay & Perduel976; Hartwell 1976). Later, tannins were extracted out belore testing, while tumors insensitive to tannins were subsequently em- ployed (Hartwell 1976). Consequently, many variables are represented in the definition of an active species, such as differences in extraction procedures, quantity and kind of tumor syst employed, parameters that define activity from testing extracts, and whether specific plant parts screened correspond to those employed in folklore. Nevertheless, it is felt that all plants regarded active by the NCI from 1960-1976 are valid for making comparisons with folk uses of plants. Comparisons between the NCI active species and those in the literature considered taxonomic synonyms and closely related species when known. For instance, the NCI active species, Thalictrum polycarpum (Torr) S. Wats., based ona sample collected and identified by A.S. Barclay from southern California in 1962, was not found in the literature reviewed to have medicinal or poison- ous reports; however, this species could be interpreted asasynonym of T fendleri Engelm. (Munz 1959), one that was reportedly used in medicine by the Indian SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2209 Tribes of Nevada (Train et al. 1957). Based on taxonomy, I. polycarpum is con- sidered a medicinal plant. Active genus. Comparisons were also made at the genus level; however, the size of the genus varies—from just one species (e.g., Camptotheca) to more than 1,000 species (e.g., Euphorbia), Willis 1922) had determined that 47% of all genera are monotypic, 17% have two species, 8% have three species, and the re- maining 28% have four or more species. An active genus is one with one or more active species. Because most genera have more than one species (53%), the per- centages of active genera will be higher than active species. Also, when more than one species in an active genus is reportedly used medicinally and/or poi- sonous, the relationship between antitumor activity and folklore will appear closer, or lie between the percentages of active genera and active species. Random Screen. The rationale of the NCI screen has been to regard any species as a potential source for novel anticancer drugs; thus, screening of plants has been considered random. In practice, however, collecting was not purely random. One reason is that it is not possible to collect every plant species en- countered in the field, because the quantity of dry weight needed may not be practical to obtain. Another is that geographic sampling has not been uniform for political and economic reasons. The number of genera and species screened and active in the NCI program was determined by A'S. Barclay for the symposium on “Plants and Cancer” at the Society for Economic Botany meeting in Baltimore, August 1975. His data accounted for all species and genera screened by the NCI—up to the end of 1974, taking into consideration synonyms and samples that the NCI acquired not only from the USDA, but from all contractors. His tabular summary is repro- duced here, Table 1 (Barclay & Perdue 1976). The percentages for active genera, 26.0, and species, 10.4, are the bases for making comparisons to those in folklore references; however, it must be kept in mind that the numbers for active species and genera are cumulative; ie., they do not represent the actual frequency at which activity occurs. This is because some species have been screened more than once, or have included more plant parts than others, thus, have had more opportunity to show activity—also keep- ing in mind that the NCI screen has become more selective over time. GENERAL SURVEYS The NCI computer record of active plant species was compared with species and genera cited in indices or texts of eight compendia to determine which have shown antitumor activity (Table 2). With two exceptions, active species were 1.4 to 2.6 times more frequent in references on medicinal and/or poison- ous plants than in plants screened at random, while results with active genera were more consistent—at nearly double that of the random screen. The greater variation at the species level for medicinal plants is partly due 2210 BRIT.ORG/SIDA 21(4) Taste 1. NCI overall screening data for vascular plants (1960-1974). Number Screened Number Active % Active Genera 4,716 |, 225 26.0 Species 20,535 2,127 10.4 to many species not screened, in contrast to higher percentages of genera screened. For one reference, Quisumbing (1951), it was determined that 626 of the 855 species were tested; thus, instead of the 16.4% active of those recorded (855), 22.4% of those species actually screened (626) were active—nearly double that of the random screen. In regard to the wide ranging values seen for poisonous plants, the lower percentage of 9.2% active species in Webb (1948) seems related to many species that are suspected to cause poisoning of livestock. When data from the same reference was restricted to species that were reported to be poisonous and also used medicinally, the percent active species was notably higher, 18.9%. These data suggest that plants, both poisonous and used medicinally, are more likely to show antitumor activity than those strictly used medicinally. Also, data from other references (Kingsbury 1964; Hardin & Arena 1974) had more plants con- firmed to be poisonous, which in Hardin and Arena (1974) were restricted to those taken internally (Spjut & Perdue 1976). The higher percentages of active species (21.5%, 41.1%) and genera (56.4%, 66.4%) in these references on poison- ous plants indicate that toxicity is a factor in the apparent correlation between antitumor activity and plants generally used in medicinal folklore. ACTIVE PLANTS ACCORDING TO NUMBER AND KINDS OF MEDICINAL USES Quisumbing (1951), in his Medicinal Plants of the Philippines, provided species indices for 116 different categories of therapeutic uses and for 111 different kinds of specific diseases, a total of 227 different medicinal applications from which 90 were selected on the basis of 19 or more species being listed to determine whether antitumor activity was more closely correlated witha particular thera- peutic effect (Appendix I, 62 medicinal applications) or specific disease (Ap- pendix II, 28 medicinal applications). What we found, however, was a broad correlation with all medicinal applications (Appendix I, 11). This broad correla- tion appears related to a large number of widely distributed species for which many have probably been screened more than once by the NCI, while a correla- tion between antitumor activity and toxicity is also evident. These relation- ships will be made apparent in the data and discussion that follow. Quisumbing (1951), in reporting on 855 species in 580 genera and 143 families of vascular plants in The Philippines, did not limit his review to medicinal uses within The Philippines. He also drew on literary sources outside The Philippines. SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2211 Tasce 2.Number and percent of active genera and active species for medicinal and poisonous plants in eight selected references. Reference Genera Genera %Genera Species Species % Species Listed active active Listed active active Medicinal Plants Krochmal (1973) 207 131 63.3 251 67 26.7 Quisumbing (1951) 580 271 46.7 855 140 16.4 Train et al. (1957) 142 77 54.2 214 32 15.0 Webb (1948) 398 228 57.3 529 87 16.5 Weiner (1972) 285 156 54.7 388 73 18.8 Poisonous Plants Hardin & Arena (1974) 113 13 66.4 141 58 41.1 Kingsbury (1964) 282 159 56.4 488 105 21.5 Webb (1948) 433 211 48.7 760 70 oe Poisonous Plants used Medicinally Webb (1948) 229 153 66.8 196 37 18.9 Plants Used Against Cancer Hartwell (1967-1971) 1,201 480 46.5 2729 314 17.3 (1,033 (tested) (1,815 (tested) tested) tested) Thus, many plants not known to be used medicinally in The Philippines were included so long as the plant occurred there, a practice not uncommonly em- ployed by many ethnobotanists in other geographic studies of medicinal plants. Nevertheless, the result is that there are many widespread species represented. This is evident in part by finding that 8% of all species in Quisumbing (1951) are endemic to The Philippines, based on geographical data he also provided; thus, 92% of the species in Quisumbing (1951) extend beyond The Philippines. The distribution of endemic species according to the number of uses is shown in Figure 1. Among 110 species in Quisumbing (1951) listed for only one medicinal application, 25% were found to be endemic to The Philippines, fol- owed by a sharp decline for those reported under multiple applications—15% for plants listed under two medicinal applications, 8% for three medicinal ap- plication—to none found under nine or more medicinal applications. It is cer- tainly not surprising to find that narrower geographically distributed species have fewer medicinal reports. However, the extent to which medicinal species are reported for many differ- ent uses is perhaps not fully realized by many ethnobotanists. The 808 species listed, among the 90 medicinal applications selected from Quisumbing (1951), ac- counted for a whopping, 5,843 species entries (meaning that many of the 808 species are used for more than one application), the distribution of which is shown in Figure 2. As an average, 50% of the species reported under any one 2212 BRIT.ORG/SIDA 21(4) ENDEMIC SPECIES and MEDICINAL APPLICATIONS % Endemic Species l J 16+ Medicinal Applications Fic 1 ip} Ty Th Pe aH - is L £ Ace on ££ A777 J 1 1 - Nn L ae 41 c1 Th “ L £ J 1 Fe lL ~ L £ L VUILL/ VI y . + . mT 1£, J 1 CO, A ta Tha Dhil f, in Fig I I y used for only ry 5 species with two medicinal applications, 15% demic, etc., t lemics for species reported to | es 1 iH = CC | 4 la L J heat | 4 fanc.1\ g (1951) medicinal application were also found under 11 or more other medicinal appli- cations. The extent of the widespread occurrence for many of the medicinal plants reported by Quisumbing (1951) is further evident by percent species screened according to the number of uses recorded, Figure 3, and the fact that relatively few species were actually collected from The Philippines. Some of the medici- nal applications in the higher multiple use categories were combined to obtain a more equitable number of species for each category. The results show, as one might expect, a definite correlation between the number of uses and percent species screened, increasing from 45% for species with only one medicinal ap- plication, to 99% for those with 16 or more medicinal applications. Plants were procured largely from the United States, Australia, New Zealand, Fiji, Taiwan, India, Turkey, Ethiopia, Kenya, Tanzania, South Africa, Ghana, Mexico, Panama, Colombia, Brazil, and Peru. Small numbers of collections were also obtained from other countries; see also procurement map in Perdue (1976). For the 90 selected medicinal applications from Quisumbing (1851), 626 species in 531 genera were found to have been screened of which 140 species SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2213 Species SPECIES and MEDICINAL wll APPLICATIONS 12 34 65 6 7 8 9 1011 1213 141516 171819 20 21 22 23 24 25 26 27 28 29 30 31 Medicinal Applications A 2 i *1 4 £ ‘4 Ph L .< 1° iH 4 £. J LON Ack. i: J Fic.2 of hat included 808 species in Quisumbing (1951) ). The number of species for each number of medicinal applications decreases from 110 ee used for just one apingore to one species, Artemisia vulgaris L., cited under 31 different medicinal applications. (22.4%) in 265 genera (49.7%) were active (Appendix IIL); additionally, 40% of the 140 active species were found to have 12 or more medicinal applications. One medicinal application with notably high percentages of active species and genera was plants used against hemorrhoids, 35.3% (24) of the 68 species and 72.1% of the 61 genera. Are plants used for treatment of hemorrhoids more closely correlated with antitumor active plants than plants used for other purposes? Statistically, the distribution of active genera and species for the medicinal applications in Quisumbing (1951) might be expected to follow a bell-shaped curve distribution in which there will be higher than average as well as lower than average percentages of active species (and genera). The categories with higher percentages of active species would also be expected to have more widely distributed species based on data presented in Figures 1-3 and the absence of plant collections from The Philip- pines as already indicated. Indeed, among 68 species listed by Quisumbing (1951) for plants used against hemorrhoids, 75% (51) were reported for 11 or more other medicinal applications, which included 23 of the 24 active species. Therefore, it cannot be | are more likely to show antitumor activity than plants used for other purposes. 1 that plants used fora partic ular remedy suchas hemorrhoids 2214 BRIT.ORG/SIDA 21(4) SPECIES TESTED and MEDICINAL APPLICATIONS % Species Tested 96 99 100 12 3 4 5 6 7 8 9 10 11 12-15 16+ Medicinal Applications Fic 3.P tnt H 14 n AICI SE H ae iH L € eH 1 " . £, Ag H iH 1°. ON anf 997 Ait, 1: n i hu And hj (1001) Th numerical rarenory: of medielnal uses of plants is shown to increase from 45% ck species repore to have just one 16 or more different medicinal application On the other hand, one might argue that the use of plants for many medicinal remedies by one or more cultures should constitute strong evidence for discover- ing biological activity. At the species level, however, cultural diffusion might ex- aggerate and multiply reports (Watson 1983), whereas medicinal reports based on disjunct occurrences of closely related species in genera may appear more valid, depending, however, on the size of the genus and number of medicinal species reported. The following six cases exemplify how folklore may appear in one case to have strong validity, while in other instances appears inconclusive. 1) Brucea (Simaroubaceae) is a small paleotropical genus of 6 species with B.antidysenterica in Africa and B. javanica (L.) Merr. in southeast Asia that have reportedly been used for treating skin diseases, dysentery, tapeworm, and cancer (Burkhill 1935; Chopra et al. 1956; Dalziel 1937; Hartwell 1967-1971. Quisumbing 1951; Watt & Breyer-Brandwijk 1962; Webb 1948). Anticancer ac- tivity has been identified in both species and one other, B. guineensis G. Don, found only in west tropical Africa without any reported use. The anticancer compound, bruceantin (Kupchan et al. 1973), isolated from B. antidysenterica, has undergone preclinical studies as a potential drug for cancer chemotherapy. It was found to be toxic in human application; however, derivatives of related SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2215 compounds are still being investigated for cancer chemotherapy (Cuendet & Pezzuto 2004; Mata-Greenwood et al. 2001). 2) Colubrina (Rhamnaceae) includes one widespread species, C. asiatica (L.) Brongn. eight species of spotty distribution in the Old World, one in India, three in Indonesia and four in Madagascar, plus about 22 species distributed in tropical and subtropical America Johnston 1971). Colubrina asiatica has been used as an abortifacient and for treating skin diseases (Quisumbing 1951). Spe- cies of Colubrina in the West Indies and Mexico have been used as an anthelm- intic and for treating dysentery and skin diseases (Standley 1922-1926). Anti- cancer activity has poe identified in six New World species, but not in C. asiatica. An d, colubrinol (Wani et al.1973), isolated from C. texensis (Torr. & Gray) A. Gray, is related to maytansine, which has undergone clinical studies for cancer chemotherapy as discussed below. Colubrina californica, a closely related species to C. texensis, has also shown similar activity, but no medicinal reports could be found for these species. 3) Maytenus (Celastraceae) is a large pantropical genus of 150 or more spe- cies with relatively few species reported for medicinal purposes. One species in South America, Maytenus ilicifolia Mart. Ex Reiss., has been employed for treat- ing a variety of ailments such as peptic ulcers, dyspepsia, gastralgia, enteritis, cystitis, insomnia, nervousness, acne, hemorrhoids, dysentery, and cancer (Hartwell 1967-1971; Morton 1968). In Mexico, M. phyllanthoides Benth. has been employed as a remedy for scurvy and toothache (Standley 1922-1926), and M. pseudocasearia Reiss. has been used to treat dysentery (von Reis Altschul 1973). In East and South Africa, four or five species have been used medicinally as remedies for amoebic dysentery, diarrhea, colic, malaria, epilepsy, “madness,” colds and cancer (Harington 1969, Watt & Breyer-Brandwijk 1962). Anticancer activity has been identified in 21 of 31 Maytenus species screened. An ansamacrolid, maytansine (Kupchan et al. 1972), isolated from several African species, underwent clinical trials for cancer chemotherapy. This was discon- tinued because of toxicity; however, there is renewed interest in derivatives of maytansinoids, which are less toxic (Bander et al. 2003; Larson et al. 1999). 4) Ficus (Moraceae) is a very large pantropical genus, “800 species (Airy Shaw 1973), and many Ficus species are employed medicinally for a variety ot purposes throughout the tropics. Seventeen species had shown antitumor ac- tivity; yet, none have yielded compounds for clinical studies. 5) Fritillaria (Liliaceae) has about 85 species distributed in temperate re- gions of the northern hemisphere (Airy Shaw 1973). In China, species of Fritillaria are used for a wide variety of ailments that include cancer (Hartwell 1967-1971; Steinmetz 1962). In Europe and the Himalayas of India, several species have been used against asthma and tuberculosis (Steinmetz 1962). The NCI has screened species from Southeast Asia, Europe, and the United States; none have shown activity. 2216 BRIT.ORG/SIDA 21(4) 6) Thamnosma (Rutaceae) is a small genus of 8 species with a spotty dis- tribution: southern Africa, Arabia, Socotra and the southwestern United States (Airy Shaw 1973). Africans have smoked plants of T. africana Engl. to relieve chest conditions (Watt & Breyer-Brandwijk 1962). A decoction of the stems of I. montana Torr. & Frem. has been used by Native American tribes of Nevada for colds and as a tonic (Train et al. 1957). Both species have been screened by the NCI; neither was active. It is apparent from these six cases that an objective analysis is difficult. Subjectively, one might weigh small genera (Brucea) more than large genera (Ficus), similar medicinal uses as opposed to different uses—among different cultures, spotty distribution as seen for species of Brucea and Thamnosma, over continuous distribution as in the case of Ficus, and to the kinds of medicinal applications, especially cancer (e.g., Brucea, Fritillaria, Maytenus) as opposed to treating colds (e.g, Thamnosma). In Ficus it might appear significant that many species are used medicinally in folklore; however, of 174 species of Ficus screened by the NCI, only 9.8% were active, which is slightly less than that of the random screen (10.4%). In the case of Fritillaria, however, there is no corre- lation evident due to lack of activity. PLANTS USED AGAINST CANCER Hartwell (1967-1971) compiled a record of more than 3,000 species of plants reported in folklore for treating cancer and other symptomatic conditions such as warts and tumors. The vascular plants included 2,725 species representing 1,201 genera and 185 families. An estimated two-thirds of the species and 86% of the genera were screened for antitumor activity based on sampling of four families (Fabaceae, Liliaceae, Rubiaceae, Rutaceae: Spjut & Perdue 1976); it was not practical to compare all 2,725 species in Hartwell against the record of 20, 225 species screened, as was done for the NCI record of 2,127 active species of which 314 active species were found in Hartwell (1967-1971). Thus, an extrapo- lated result is provided, indicating 17.3% active species and 46.5% active genera for those screened and used against cancer (Table 2). The percentages of active species and active genera found in Hartwell’s (1967-1971) record of plants used against cancer are comparable to that seen in the general references on medicinal plants (Table 2). It should be realized that the greater the number of species included ina study like that of Hartwell (1967- 1971), the greater the number of species that will be represented with relatively narrower ranges in geographical distribution; thus, the impact of the more thor- oughly screened, widely distributed species, will be less. The 1.7 fold increase in active species and the L.8 fold increase in active genera over the random screen in Hartwell’s (1967-1971) plants used against cancer is perhaps a more realistic assessment of the relationship between plants used in medicinal folklore and those that have shown antitumor activity in the NCI screen. SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2217 RELATIONSHIPS BETWEEN ANTITUMOR ACTIVITY AND MEDICINAL PLANTS, Toxic PLANTS, AND POISONOUS PLANTS General Surveys.—The percentages of active species in the general surveys (Table 2) indicated that poisonous plants, including those with medicinal uses, appear more likely to show antitumor activity than medicinal plants in gen- eral. Antitumor activity among the different therapeutic uses (Appendix 1) were also evaluated for evidence of a correlation with plant toxicity; for example, a plant used as an emetic will likely induce a stronger physiological reaction, which could also be more harmful if taken in excess, than a plant taken asa stimulant. Ina further review of the 62 medicinal applications in Quisumbing (1951, Appendix 1), ten were selected as representative of two therapeutic use categories: (1) five that represent a weak-to-moderate effect—stimulant, alter- ative, diaphoretic, aperient, and laxative—and (2) five that appear to exert a moderate-to strong physiological effect—purgative, cathartic, abortifacient, anthelmintic, and emetic. A comparison of the percentages of the active spe- cies in the two categories (Table 3) show that the percentages of active species are all higher in the moderate-to-strong category, suggesting, therefore, that plants with medicinal uses associated with possible toxic side affects are more likely to show antitumor activity than medicinal plants in general. Plants Used as Anthelmintics.—Plants used as anthelmintics—those taken internally by humans for helminth infestations such as tapeworm, roundworm, guinea worm, elephantiasis and shistosomiasis—are included in Table 3 as an example of a medicinal application where one may expect a moderate to strong reaction in using a plant product that results in the expulsion or destruction of parasitic worms. Thus, from this perspective, the 30% active species of the 150 species listed in Quisumbing (1951) would appear to have a closer correlation with antitumor activity when compared to the 22.4% active species for all me- dicinal plants in that same reference, besides the less frequent active species among those therapeutic uses that imply a weaker physiological effect (Table 2, 3, Appendix LD). Nevertheless, an independent review of the literature was conducted to determine which species are reported as anthelmintics—because of Perdue's observation on such plants in Ethiopia that were also active in the NCI screen (Spjut & Perdue 1976). Recorded were 668 species in 457 genera and 128 fami- lies of which 482 species in 433 genera were screened. The active species, and the bioassay(s) in which they were active, are indicated in Appendix IV, a com- plete list of plants used as anthelmintics for this study with references to each species is available at www.worldbotanical.com. Of those tested, 29.3% of the species and 52.2% of the genera were active. The 29.3% active for anthelmintic species is nearly three times that of the 2218 BRIT.ORG/SIDA 21(4) Taste 3. Antitumor activity as related to potency of therapeutic effect: selected medicinal applica- tions from Quisumbing (1951). Therapeutic Use Percent Active Species Weak to Moderate in Effect Stimulant 148 Alterative 23.4 Diaphoretic 23.1 Aperient 22.5 Laxative 20.6 Moderate-To-Strong In Effect Purgative 25:7 Cathartic 25.9 Abortifacient 27.9 Anthelmintic 30.0 Emetic 3251 random screen, and is clearly higher than that seen in general references on medicinal plants (Table 1), in particular the 22.4% found for all Medicinal Plants of the Philippines (Quisumbing 1951). These data support the finding that me- dicinal plants with indication of toxic side effects, such as the case with anthelmintics, are more likely to show biological activity, than medicinal plants in general. Plants Used as Fish and Arrow Poisons.—As with anthelmintics, we com- piled separate lists for plants used as fish and arrow poisons that also included ordeal and homicidal poisons (Spjut & Perdue 1976). These data can be found at www.worldbotanical.com; in this publication, only the active species with ref - erence to the tumor assay are listed, Appendix V, VI. The results, presented in Table 4, show that the percent active species among those tested was 38.6% for plants used as fish poisons and 45.7% for plants used as arrow, homicidal and/or ordeal poisons. Plants used as poisons are obviously more toxic than those generally used for medicinal purposes, which are not employed for lethal purposes, but still can be deadly if taken in excess. One might also expect fish poisons to be some- what less harmful than arrow poisons, because fish poisons are used to capture fish for consumption in which the fish are often only stunned, whereas arrow poisons are intended to kill. Data on antitumor activity that correlates with these differences (Table 4) are seen as another example of a correlation between plant toxicity and antitumor activity. The correlation that is evident between poisonous plants and antitumor activity led to further evaluation in regard to the type of tumor activity, be- cause activity in poisonous plants was suspected as largely occurring in the KB Cell Culture, a bioassay that is sensitive to cytotoxic agents (Hartwell 1976). SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2219 Taste 4. Antitumor activity in poisonous plants. Poisons Genera tested %Genera active Species tested %Species tested active Fish 158 65.8 145 38.6 Arrow, Ordeal, 60 75.0 70 45.7 & Homicidal Data in Appendix III, IV, V, and VI, which indicate tumor system of activity with their percentages of active species and genera, confirmed this. These data are summarized in Table 5. The percentages of KB active species were found to be 6.7% for medicinal plants in The Philippines (Quisumbing 1951), 11.4% for anthelmintics, 20.7% for fish poisons, and 30.0% for arrow poisons, in contrast, for example, to activity in the WA assay that was 8.5%, 8.3%, 8.3%, and 7.1%, respectively. Clearly, there is correlation between antitumor activity and plant toxicity based on the KB assay and folklore data. DISCUSSION AND CONCLUSIONS Selective approaches to screening plants for antitumor activity have been con- ducted previously by taxonomy (Belkin & Fitzgerald 1953b), by taxonomy and medicinal use such as anti-malarial plants in the Amaryllidaceae (Fitzgerald et al.1958), and by specific medicinal or poisonous applications such as plants used as cathartics, diuretics and pesticides (Belkin et al.1952a; Belkin & Fitzger- ald 1952b, 1953c). These and other similar experimental studies were limited to screening against Sarcoma 37. It is interesting to note that in the case with plants used as cathartics, nearly half of the species tested were active. This might be compared to another study by the same authors using the same bioassay in screening “miscellaneous plants” in which they found only 14% active (Belkin & Fitzgerald 1953a); a comparison that is analogous to the “random screen” in the present study. One important discovery relating to these investigations came from the medicinal use of a root extract of May-apple, Podophyllum peltatum L. (Berberidaceae), known as “podophyllin.” Hartwell (1960, 1976) indicated he had investigated podophyllin and samples of May-apple because of their use against cancer by practitioners in the United States and by the Penobscot Indians of Maine. Records for such use were found to date back to 1849; additionally, in Louisiana May-apple was used to treat venereal warts or as an “escharotic,” dat- ing back to 1845 (Hartwell 1960). Podophyllotoxin and two peltatins were iso- lated and found to be highly active in Sarcoma 37 (Hartwell & Shear 1947). Hartwell (1976) commented that the development of podophyllotoxin as a po- tential drug was complicated by toxicity, but also indicated “there is reason to hope that chemical derivatives may be developed which with will eliminate this disadvantage.” “Etoposide” and “teniposide” are semi-synthetic derivatives 2220 BRIT.ORG/SIDA 21(4) Tase 5, Comparison of general and specific folk uses of plants with percentages of active species according to antitumor assay. Folklore Use KB PS WA LL SA Medicinal Uses in General (Quisumbing 1951) 6.7 8.2 8.5 1.1 3.5 Anthelmintics 11.4 9.5 8.3 2.1 5.6 Fish Poi 20.7 97 8.3 4] 8.9 Arrow & Homicidal Poisons 30.0 18.6 7.1 1.4 1.4 currently in use as drugs to treat small-cell lung cancers, testicular cancer, car- cinoma, and lymphomas (Moraes et al. 2002). Their development, known also as “VM-26" and “VP-213,”came from 4demethylpodophyllotoxin that was found in a Himalayan species, Podophyllum hexandrum Royle (Hartwell 1976). Advocates of promoting folklore as the tool for discovery of biologically active compounds must recognize that there are a large number of widely dis- tributed species that are frequently reported for use in medicines, and have al- ready been chemically investigated. Examples of these, which have shown an- titumor activity, are candlenut (Aleurites molucanna[L.] Willd), custard apples (Annona reticulata L., A. squamosa L.), star fruit (Averrhoa carambola L.), cab- bage (Brassica olearacea), paradise-flower (Caesalpinia pulcherrima [L.] Sw), Indian laurel (Calophyllum inophyllum L.), safflower (Carthamus tinctoris L.), Madagascar periwinkle (Catharanthus roseus [L.] G. Don), coconut (Cocos nucifera L.), coffee (Coffea arabica L.), taro (Colocasia esculenta [L.] Schott), sun- flower (Helianthus annuus L.), Indian heliotrope (Heliotropium indicum L.), beach morning glory (Ipomoea pes capre [L.] R. Br), mango (Mangifera indica L.), China-berry (Melia azedarach L.), oleander (Nerium oleander L.), avocado (Persea americana Mill), peach (Prunus persica L.), pomegranate (Punicagrana tum L.), bracken fern (Pteridium aquilinum [L.] Ku ie bran erove (Rhizophora mangle L.), castor bean (Ricinus communis L.), nightshade (Solanum nigrum L.), teak (Tectona grandis eae yellow oleander (Thevetia peruvidana [Pers.] K. Schum.) (Tables | and 2 in Spjut 1985; Buckingham 1993-2005; USDA 1980), and most other species in Quisumbing (1951) that were found to be active in the NCI screen (Appendix IID. Uses for many of these active species date back to the early domestication of plants (Zohary & Spiegel-Roy 1975), a time when there was lack of concern for intellectual property rights or ownership that, for the most part, has evolved only since the last decade (Lesser 1997). Hartwell (1960) noted that cancer rem- edies can be found as early as 1500 B.C. in the Ebers papyrus of Egypt, that “plant remedies for cancer are described in ancient Chinese and Hindu medical writ- ings,” that “the record continues unabated through the Graeco-Roman period and the Christian and Arabian-Middle Ages to modern times,” and that “the SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2221 roster of the hundreds of medical, pharmacological and botanical works rec- ommending herbal treatments for cancer reads like a summary of the great names in the history of medicine.” | have further suggested that the various uses for many of the widespread species (e.g, Appendix IIL) are the result of cultural diffusion; thus, any indigenous ownership claim(s) for a particular use for a particular plant remedy cannot be easily substantiated. Cultural diffu- sion may also explain many medicinal uses for a species within a relatively narrow geographic area, as evident with plants used by Indian Tribes of Ne- vada (Train et al. 1957). Although the occurrence of anticancer activity among plants used as folk- lore remedies, when compared with that for plants tested at random, suggests that folklore could be a useful tool for predicting sources of anticancer activity, there are also costs that have to be taken into consideration in trying to selec- tively pursue such plants (Hartwell 1976). A field team can randomly collect as many as 60(-100) samples ina day from 10-30 species (Perdue & Hartwell 1969), whereas a more selective approach, as I have experienced with recollections of active plants, would yield only 1-2 samples per day. Thus, a random field col- lection could generate 1-3 new active leads each day, whereas it would require 2-3 days to obtain a similar result in a selective approach. It might be added that this folklore study was based on reports in literature. Obtaining such in- formation directly in the field would cost even more. On the other hand, it is also evident from the data presented in this study that many of the alleged medicinal species would be collected in a random (biodiversity prospecting) screening program—because of their widespread occurrence. Furthermore, a biodiversity (random) type of approach undertaken systematically is not only less expensive, but will also yield novel compounds from plants not reported in folk literature (e.g., camptothecin from Camptotheca acuminata, Perdue et al. 1970), and provide a scientific foundation for identifying chemotaxonomic, ecological and other relationships of pharmacological value. Random collec- tions can also include medicinal and/or poisonous plants in the collection strat- egy, the focus of which might be on genera that are clearly indigenous or en- demic toa collection area, and would likely yield novel compounds. The NCI screen involves more than just identifying leads such as the 2,127 active species reviewed in this study; other steps in drug development include isolating and identifying the active compounds, pharmacological evaluation of the active compounds, and clinical evaluation for treating cancer in three phases (Goldin et al. 1974). Criteria for clinical consideration during the 1970s had included activity in a panel of tumor systems such as the L-1210 Leuke- mia, KB Cell Culture, P-388 Leukemia, new Lewis Lung tumor, and B16 Mela- noma (Goldin et al. 1974; Hartwell 1976). Compounds from only ~1% of the 2,127 active species had reached clinical evaluation—Table 1 in Hartwell (1976). Sev- enteen of 21 genera in Hartwell (1976, Table 1) were identified as having less of — 2222 BRIT.ORG/SIDA 21(4) a taxonomic relationship to each other among the compounds of clinical inter- est (Acer, Brucea, Camptotheca, Caesalpinia, Cephaelis, Cephalotaxus, Colchi- cum, Fagara, Heliotropium, Holacantha, Maytenus, Ochrosia, Stereospermum, Taxus, Thalictrum, Tripterygium, Tylophora). With exception to Camptotheca and Holacantha, these genera were found to have species reported in the litera- ture as poisonous. Holacantha, a genus of two species, has a very limited distri- bution in southwestern North America, thus, the lack of medicinal reports for this genus is not unexpected, although a closely related genus, Castela, includes species used in folk medicine (Standley 1922-1926). Similarly, Camptotheca, a monotypic genus of limited distribution in China, lacks reports on medicinal use except for one general reference on a herbarium specimen “drug plant” F A. McClure 6546 at AA (Perdue et al. 1970). Of the remaining genera, all except Cephalotaxus, Ochrosia, Tripterygium and Tylophora have species reportedly used against cancer or cancer like symptoms (Hartwell 1967-1971). It might be noted that nearly all active compounds in these plants were discovered from screening in the KB Cell Culture (Hartwell 1976). The correla- tion between anticancer activity and plant use indicative of toxicity might in- dicate that future screening of plant extracts could place more emphasis on bioassays that can detect cytotoxicity, such as the KB assay (Perdue 1982: Spjut S Perdue 1976); however, KB activity alone will not lead to development of a new anticancer drug, as evident for plants used as arrow poisons, in which 21% of the active species are strictly KB actives. Many of these plant poisons belong to genera in the Apocynaceae and Asclepiadaceae whose activity is largely due to cardenolides, steroid lactones that have not demonstrated much in vivo ac- tivity (Hartwell 1976, Table 15). Poisonous plants in two other families, Cucurbitaceae and Datiscaceae, have yielded only cucurbitacins, triterpenes that are toxic without in vivo activity (Hartwell 1976, Table 10; Cassady & Suffness 1980). Additionally, many other species of poisonous plants are in the Euphorbiaceae in which P-388 Leukemia activity was more frequent, but the compounds were largely phorbol esters (Suffness & Douros 1979). Such com- pounds are known to be tumor-promoting (Farnsworth et al. 1976), while also inactive in other antitumor assays (Suffness & Douros 1979; Cassady & Suffness 1980); however, one non-tumor promoting phorbol ester was found to have po- tential for treating AIDS (Gustafson et al. 1992). Nevertheless, the extent to which plant genera include species reported in folklore to be poisonous, and also used in medicine, especially against cancer, certainly deserve further study. The potential for discovery of novel chemothera- peutic agents would appear greater when geographical evidence indicates simi- lar uses in different cultures as earlier described for Brucea and Maytenus, while Hartwell (1967-1971) also mentioned that Heliotropium indicum and other spe- cies of this genus have been reported in folklore for treating cancer in scattered regions of the world. Thus, the relationship between anticancer activity and SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2223 folklore appears more meaningful and less coincidental when there is this kind of support from taxonomic and geographic data. Future screening might focus on genera that have yet to show activity. A good example is Fritillaria, a genus reportedly rich in alkaloids with highly toxic species that are used for medici- nal purposes, including cancer (Steinmetz 1962). One of the most useful drugs in the chemotherapy of acute childhood leu- kemia (and other cancers), is vincristine from the periwinkle, Catharanthus roseus (L.)G. Don., one of the many widely distributed species used in folk medi- cine. This discovery resulted not from a search for antitumor activity, but was incidental to a search for compounds with hypoglycemic activity. The plant was under investigation in two different laboratories because of its folk use asa remedy for diabetes (Carter 1976). These facts, and the apparent correlation in this paper between various uses of medicinal plants and antitumor activity, suggest that antitumor activity should be looked upon as just one kind of bio- logical activity that probably correlates well with a broad spectrum of other kinds of biological activity. There is a growing interest in natural products as food additives and as al- ternative medicines, partly promoted by an awareness and need for biodegrad- able natural products to replace synthetic chemical compounds that increas- ingly contaminate our environment Jacobson 1989). Where new kinds of biological activity are sought, such screening programs can benefit not only by taking into consideration folkloric uses of plants, but also the massive amount of data generated by the NCI random screen, suchas the many novel antitumor agents that have been reported. Therefore, one would hope that the NCI con- tinue screening of natural products. The byproducts of this program are in- valuable as many compounds, undoubtedly, will find use in other therapies if they cannot be used to treat cancer. A case in point is recollections of antitumor active plants from which small amounts were funneled to Martin Jacobson at another ARS laboratory in Beltsville, MD who apparently found good insecti- cidal activity in many of the NCI active plants, e.g, Arnica chamissonis Less. ssp. foliosa (Nutt.) Maquire (USDA ARS Medicinal Plant Resources correspon- dence; data recorded for requests of recollections by active species and geo- graphical location; www.worldbotanical.com; see also Jacobson 1989). Finally, there is one aspect of the folk medicine that cannot be compared with the NCI’s random method of searching for potential anticancer drugs. In folk medicine, prescriptions may include a combination of two or more plants, and/or other substances. This is especially common in Chinese medicine (American Herbal Pharmacology Delegation 1975). The separate ingredients of a prescription may not show activity, but one may speculate on whether there isa synergistic effect with combined materials as often seen in drug combina- tion therapies. 2224 BRIT.ORG/SIDA 21(4) APPENDIX I. ANTITUMOR ACTIVITY IN QUISUMBING (1951) PLANTS ACCORDING TO THERAPEUTIC PROPERTIES Therapeutic Number of Number of % of Number of Number of % of Property Species Species Species Genera Gener enera i Active Active Listed Active Active Abortifacient 6] 17 27.2 58 63 74.) Alexipharmic 20 8 40.0 20 16 80.0 Alterative 47 in| 23.4 45 27 60.0 Anthelmintic 150 45 30.0 132 88 66.6 Antiarthritic 25 5 20.0 22 16 727 ntiasthmatic 83 22 26.5 74 49 66.2 Antibechic 121 22 18.2 99 57 57.6 Antibilious 27 8 29.6 25 14 56.0 Antiblennorrhagic —-110 28 25.5 98 53 53.5 Anticatarrhal 36 8 22.2 34 20 58.8 Anticephalagic 96 23 24.0 89 50 56.2 Anticolic 7\ 18 25.4 69 46 66.6 Antidiabetic 35 8 22.9 31 22 71.0 Antidiarrhoetic 156 39 25.0 136 85 62.5 Antidyspeptic 60 15 25.0 54 34 63.0 Antidysenteric 177 43 243 150 86 57.3 Antiherpetic 26 9 34.6 25 13 52.0 Antimalarial 50 13 325 37 25 67.6 Antinephritic 23 3 13.0 22 7 31.8 Antineuralgic 22 5 22-7 2] eZ 57.1 Antiodontalgic 56 15 26.8 51 33 64.7 Antipyrotic 29 5 17.2 29 18 62.1 Antirheumatic 167 40 24.0 140 80 57.1 Antiscabious 77 17 22.) 67 43 64.2 Antiscorbutic 38 10 26.3 35 19 54.3 A tic 42 10 23.8 39 25 64.1 Antispasmodic 49 15 30.6 46 30 65.2 Antisyphilitic 37 10 27.0 34 18 52.9 tiveno 50 9 18.0 46 22 478 Aperient 40 9 22.5 38 19 50.0 Aperitive 27 7 25.9 25 14 56.0 Aphrodisiac 48 9 18.8 47 27 574 Astringent 174 42 24.1 156 94 60.3 Carminative 92 iia 12.0 80 44 55.0 Cathartic 27 7 25.9 24 18 75.0 Demulcent 64 11 17.2 59 33 55.9 Depurative 39 10 25.6 36 21 58.3 Diaphoretic 91 21 23.) 85 50 58.8 Digestive 27 8 29.6 25 16 64.0 220 53 24.) 18] 107 59.1 Emetic 78 25 32.] 74 52 70.3 SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2225 APPENDIX I. (CONTINUED) Therapeutic Number of Number of % of Number of Number of % of Property Species Species Species Genera Genera Genera i Active Active Listed Active Active Emmenagogue 132 34 25.8 119 72 60.5 Emollient 77 22 28.6 69 42 60.9 Expectorant 54 11 20.4 50 64.0 Febrifuge 222 53 23.9 191 112 58.6 Galactagogue 26 7 26.9 23 14 60.9 Hemostatic 36 8 22.2 35 19 54.3 Laxative 63 13 20.6 62 36 58.1 Lithotriptic 27 4 148 27 15 55.6 24 6 25.0 20 13 65.6 Pectoral 40 14 35.0 39 27 69.2 Poultice 218 41 18.8 178 85 478 Purgative 105 27 25e, 85 59 69.4 Refrigerant 53 6 11.3 48 29 60.4 Rubefacient 38 iis 34.2 35 24 68.6 S 3] 5 16.1 27 13 48.1 Stimulant 108 16 148 89 50 56.2 Stomachic 145 34 23.4 125 76 60.8 176 32 18.2 155 84 54.2 Tonics (bitter) 34 10 29.4 33 23 69,7 Vesi 22 5 22.7 19 13 68.4 Vulerary 82 13 15.9 76 35 46.1 APPENDIX II. ANTICANCER ACTIVITY IN QUISUMBING (1951) PLANTS ACCORDING TO SPECIFIC DISEASES Medicinal Number of Number of % of Number of Number of % of Species Species Species Genera Genera Genera is ti Active Listed Active Active Abscess 22 7 31.8 22 17 TAS Alopecia 26 5 19.2 26 13 50.0 Amenorrhoea 29 5 Veh. 27 18 66.7 Anasarca 57 7 29.8 54 37 68.5 Aphthae 57 17 29.8 54 37 68.5 Bronchitis 39 8 20.5 36 22 61.1 Cholera 29 6 20.7 2f 16 59.3 Constipation 30 10 33.3 28 19 67.9 Ears, Affections of 36 8 22.2 32 19 59.4 Eczema 24 10 41.7 22 16 27 Eyes, Affections of — 40 9 22.5 39 22 56.4 Furuncles 65 16 24.6 63 37 58.7 Gingivitis 19 6 31.6 19 14 veo 2226 BRIT.ORG/SIDA 21(4) APPENDIX II. (CONTINUED) Medicinal Number of Number of % of Number of Number of % of Species Species set ies Genera Genera Genera Listed Active Active iste Active Active Hemoptysis 26 3 11.9 26 11 423 Hemorrhoids 68 24 35:3 6| 44 72.) Indigestion 20 4 20.0 19 13 68.4 Jaundice 32 10 31.3 31 18 58.1 Leprosy 34 g 26.5 34 18 52.9 Liver Diseases 43 10 23:3 39 21 53.8 Menorrhagia 23 4 17.4 22 17 pe Nervous Diseases 50 16 32.0 48 34 70.8 Ophthalmia 21 9 429 21 15 714 Skin Diseases 123 29 23.6 105 55 52.4 Throat Diseases 57 16 28.1 49 34 69.4 Tinea 37 12 324 30 19 63.3 Tuberculosis 47 10 21.3 44 25 56.8 Ulcers 120 26 21.7 113 65 57:5 Wounds 128 26 20.3 111 69 62.2 APPENDIX III. ANTITUMOR ACTIVE SPECIES IN QUISUMBING (1951) MEDICINAL PLANTS OF THE PHILIPPINES Species Tumors PS WA LL SA CA Other ADI PIRCAIOS k; 1 1 Zid era (Roxb.) Benth. ] Aegis oes (L.) Willd. 1 ‘a cathartica L. ] pened scholaris (L.) R. Br. 1 rphophallus ee 1 nie NICOBS nacr»varda; 4 | L 1 | Wight & Arn 1 naxag Q luz A. Gray ] Monona muricata L, 1 eenene reticulata L. ] 1 ] ] MIU SGUGITTO Le ation: aie mph. ex Pers.) Lesch. see air flava (L.) Merr. 1 Argemone mexicana L. | ae. curassavica L. 1 1 Averrhoa bilimbi L. ] A } hn! al. 1 SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2227 APPENDIX ILI. (CONTINUED) Species Tumors KB WA LL SA CA Other — = Bacopa monniera (L.) Wettst. Barringtonia asiatica (L.) Kurz 1 Bauhinia malabarica Roxb. 1 R havin diffirea L Brassica olearacea L. 1 Bryophyllum pinnatum Kurz Caesalpinia ee (L.) Sw. 1 ] D1 Calotropis gigantea ( ander ex ee nf. Calophyllum inophyllum L. 1 ] Canna indica L. 1 on ite le, 1 diosnerm| ahabijm L. ] — = — Carthamus tinctorius L. 1 1 Cassia a alata L. ] Ca ict t+ identalis L. 1 ns siamea Lam. 1 Casuarina equisetifolia L. Catharanthus roseus (L.) G.Don 1 Ceiba pentandra (L.) Gaertner 1 1 Celastrus pees Willd. Celosia argentea L. 1 1 Centella asiatica (L.) Urban 1 Cerbera manghas L. 1 Cestrum nocturnum L. 1 Ch ate Burm. f. ] — = Clerodenrdon fragans R. Br. ] Cocos nucifera L. ] Coffea arabica L. 1 Corchorus olitorius L. 1 Cordia dichotoma Forst. 1 Crateva religiosa Forst. f. 1 Crescentia cujete L. 1 Roxb.) R. Br.ex Lindley Cryptostegia grandiflora Cyperus rotundus L. ] Datura mete! L. ] Derris trifoliate Lou 1 iospyros discolor mile 1 ] Dodonaea viscose (L [=) Jacq. ] Dregea volubilis (L.f) Benth. ] ex Hook. f. Duranta repens L. 1 2228 BRIT.ORG/SIDA 21(4) APPENDIX III. (CONTINUED) Species Tumors KB WA LL SA CA Other Elephantopus scaber L. 1 1 Elephantopus mollis Kunth Entada phaseol oides (L.) Merr. 1 Erythrina variegata L. | | Erythroxylum coca Lam. 1 Flag llaria indica L. Gloriosa superba Grangea maderaspatana Poir. Graptophyllum pictum Griff. Hedychium coronarium Koenig. ] Heliqanthrire ANN Ww — = Hern Andina ; | ] IC ICG MG, Homonoia ae ja Lour. ] Hyptis suaveolens (L.) Poit. Jatropha c Jatropha seer L. Jussiaea erecta L. ] Justicia procumbens L. 1 Kalanchoe laciniata - DC. 1 pl peat indica 1 nsium SQINESCUD ae 1 L I vara L. 1 Reveaena ae al. 1 mle fponcun ag 1 unasia amara Blanc ] Mallotus oie 1 (Lam.) Muell.-Arg. Mangifera indica L. 1 Manilkara zapota (L.) D. Royle Melia Fe CAL. 1 1 1 Melia dubia Cav 1 sais umbellata (L.) Hall. f. i a aus ja) Dy — nigra L. | AA th ; lAh] L 1 ss ‘indicum Mill. analoy» piliforn (| L.) ] —_— — Se y L. 1 spalum scrobiculatum L. | Passiflora foetida L. | Pedil o tithymaloides (L.) Po SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2229 APPENDIX III. (CONTINUED) Species Tumors KB WA LL SA CA Other P americana Mill. 1 Ph ites australis (Cav.) 1 Tie: ex Steudel aes S pe jana L. ] 1 (L.) Liebm. 1 Piper umbellat uml. 1 eee lobium saman (Jacq.) ] nth. ee ica granatum L. ] Quassia amara L 1 1 Quisqualis indica L ] Rhinacanthus nasutus Kurz ] Ricinus communi 1 ] Rubia cordifolia L. 1 ae virosa (Roxb. 1 Willd.) Baillon Pee cuneiformis Blanco 1 1 1 LE Senecio scandens Buch. Ham. 1 Setaria palm ee ieee Stapf ] Sida i 1 Solanum nigrum aL. 1 ] ] Solan verasa 7 jumt. 1 ] nets con le ] Streblus asper Lour. ] ] ] mae a pandacaqul ] Tamarindise indicts. | . 1 Tectona grandis L. 1 Tephrosia purpurea (L.) Pers. ] leniagit ia ca pe ] ] ] Theva a peruviana (Pers) ] ] Schumann lia asiatica (L.) Lam ] T lantas Merr. & Rolfe ] Trema orientalis (L.) Blume 1 Trianthema portulacastrum L. ] Vernonia cinerea (L.) Less. ] pogcongs eros (Blanco) Merr. 1 ricana Us ] Total # Active: 140 42 51 a3 7 Pap 3 Screened: 626 Percent Active: 22.4% 6.71% 8.15% 8.47% 1.12% 3.51% 0.48% 2230 BRIT.ORG/SIDA 21(4) APPENDIX IV. PLANTS USED AS ANTHELMINTICS THAT HAVE SHOWN ANTITUMOR ACTIVITY Species Tumors KB PS WA LL SA CA Other Abrus precatorius L. 1 1 pas Siepenane DC. 1 a rgifoli 1 (Hochst) Lk. E. Coda Acokanthera oppositifolia 1 (Lam.) L.E.Codd Afrormosia latiflora (Benth. 1 Agrostemma githago L. 1 Ailanthus altissima (Mill.) 1 Swingle g Alangium salviifolium (L. f.) 1 1 | Wangerin Alerjrites molicanna (L.) Willd. ] Alstonia scholaris (L.) R. Br. 1 Ambrosia artemisiifolia L. 1 Anacardium occidentale L. ] Annona glabra Annona muricata L. Annona reticulata L. Annona senegalensis Pers. nnona squamosa L Apocynum androsaemifolium L. An Vieliiaal ata bh; ee a ee ee An, dyte qi Ait, R. Meyer ] Arcangelisia flava (L.) Merr. 1 Aristolochia indica L. 1 > wn al igs) = QD nA Se s ja) nn nn D xs ma S = Barringtonia eee (L.) Kurz 1 Ps variegat 1 ersama ee pea 1 | 1 1 ae arborea S, Wats. ] Boerhavia diffusa L. 1 1 Brassica olearacea L. 1 Bridelia micrantha 1 1 1 Brucea antidysenterica 1 1 (Hochst.) Baillon Brucea javanica (L.) Merr. 1 Calocarpum sapota (Jacq.) Merr. SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2231 APPENDIX IV. (CONTINUED) Species Tumors KB PS WA LL SA CA Other Calophyllum inophyllum L. ] 1 Calotropis gigantean (L.) 1 Dryander ex Aiton f. 1 Calotropis procera (Aiton Dryander ex Aiton f. Canavalia cathartica Thouars 1 Capparis deciduas (Florsk.) ] =a Edgew. Carissa edulis Vahl ] Cassia alata L. 1 Cassia quriculata L. ] Cassia occidentalis L. ] Catharanthus roseus (L.) G.Don 1 Celosia argentea L. 1 ] Citrullus lanatus (Thunb.) Masf. Cl qt (Willd.) ] Hook. f.ex Benth. Clausena excavata Burm. f. ] — Clerodendrum indicum (L.) ] O. Kuntze Clerodendrum phlomoides L. f. ] Cocos nucifera L. 1 Coix lachryma-jobi L. ] Cordia dichotoma Forst. 1 Cornus florida L. ] Croton macrostachyus Hutch. ] ] 1 Del. Croton megalocarpus Hutch. ] Cyperus rotundus L. 1 Cryptostegia grandiflora (Roxb.) 1 R. Br.ex Li Cyp ip di q| lus L. 1 Datura mete! L. 1 Dichroa febrifuga Lour. 1 Dicran pteris linegris (Bi irm. f.) ] Underw. Dodonaea viscosa Jacq. ] Dryopteris filix-mas (L.) Schott ] Ekebergia capensis Sparrm 1 Elephantopus scaber ] ] Embilia schimperi ] Entada phaseoloides i Merr. ] Erythrina variegata L. 1 ] Erythrophleum suaveolens 1 (Guill. & Perr.) Brenan BRIT.ORG/SIDA 21(4) 2232 APPENDIX IV. (CONTINUED) Species Tumors KB PS WA LL SA CA Other Ficus sterrocarpa Diels 1 Gloriosa superba L. 1 Hp] fey Ait 4], L. ] ] Helenitum hoopesii A. Gra ay 1 Hippomane mancinella 1 Holarrhena pubescens 1 ] Buch.-Ham.) Wall. Jatropha curcas L. 1 Juglans hele L. 1 Lhaininenise< NIN] a ] ] jussiae suffruticosa L. | Lansium d ti orrea 1 Liriodendron tulipifera L. | Laat hi; t7 PR b. ] Maesa lanceolata Forsk. 1 1 sen jae (Lam.) ] uell.-A Manaiferq hg lal L. ] Maprounea africana Muell.-Arg. 1 Maytenus senegalensis 1 1 (Lam.) Exell Melia pear L. | 1 1 Melia dubia Cav. 1 Morus nigra L. ] Myrica cerifera L. 1 Myrsine africana L. | Nauclea latifolia Sm. 1 Nicotiana glauca Grah. 1 feciak ia agli (Forsk.) Chiov. 1 | phyealls peruviana a ] 1 Phytolacca americana L. | (Schumach.) Milne-Redh. Pinus palustris Mill. 1 1 Pinus taeda L. 1 Piper umbellatum L. | Plectranthus blumei (Bent.) | Launert mend rubra L. 1 hyvllitm pelt It] mL. ] Prunus persica (L.) Batsch. 1 FV eas ae nian L. 1 (L.) Kuhn. 1 = SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2233 APPENDIX IV. (CONTINUED) Species Tumors KB PS WA LL SA CA Other Punica granatum L. 1 Quassia amara L. 1 1 Quisqualis indica L. ] ia pulchra Gilg & 1 1 hellenb. Rhizophora mangle L. 1 Rhus typhina L. ] ] Salvia officinalis L ] Securidaca eneneduncuat 1 Fresen. Silents engearaiele Lk 1 1 1 LE sphacranths ah atcan us L. ] on Anas henninasi) Gila 1 GING Tagete minuta L. 1 Amarin ndus indicus L. ] Tanacetum vulgare L. ] ] Tectona grandis L. ] Tephrosia purpurea (L.) Pers. ] Tephrosia vogelii Hook. f. bs ip hy {5 ty, qa (I hen Trema orientalis (L.) Blume ] on id ale Vahl ] Urti ca aa legit 1 Vernonia amygdalina Del. 1 \ ; ; q (I L.) Le \ Pui Drake ] oe a roan Sond. ] Total # Active: 141 55 46 40 10 DY. 2 Screen ed: 482 Percent Active: 29.3% 11.41% 9.54% 8.30% 2.07% 5.60% 0.41% 2234 BRIT.ORG/SIDA 21(4) APPENDIX V. ANTITUMOR ACTIVE PLANTS USED AS FISH POISONS Species Tumors KB PS WA LL SA CA Other Acacia albida Del. 1 i Acokanthera oppositifolia (Lam,) L.E.Codd Ad, ; bh R If. f ] Dall. |. se ae corniculatum (L.) 1 Bla ee al (Spach) Nutt. 1 Agave americana L. ] 1 1 MS aceon procera i Benth. 1 1 on me arvensis L. 1 namirt ee & Arn. 1 Annonar muricata L. 1 na squamosa L. ] neces curassavica ] ] arring nia asiatica (L.) Kurz Caesapini ne (L.) Sw. 1 hyllum inophyllum L. 1 aa alata L. Cerbera manghas L. ] Chlorogalum pomeridianum 1 (DC.) Kunth. Cleistanthus collinus Benth. Croton sylvaticus L. Cucumis ficifolius A. Rich. Datisca cena (Presl.) Baillon Datura mete! L. ] Derris ce Lou 1 iospyros maritima line | Dodonaea viscosa Jacq. 1 Eremocarpus setigerus (Hook.) 1 — Benth. Euphorbia esula L. 1 1 horbia hyberna L. Fagara macrophylla (Oliv.) Engl. Fluggea leucopyrus W a 1 es yaar curcas L. 1 Leucaena leucocephala (Lam.) Dewit. SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2235 APPENDIX V. (CONTINUED) Species Tumors KB PS WA LL SA CA Other Lonchocarpus urucu Killip & 1 mit av philippensis (Lam.) 1 uell.-Arg. ie ia pee ty 1 1 1 Millettia ferruginea (Hochst) 1 Bak. Mundulea sericea (Willd.) 1 A.Chev. Pergularia an (Forsk.) Chiov. 1 1 Persea americ Mill. ] Phyllanthus a iensis 1 Muell.-Arg. Piscidia erythrina L. Pleiogynium solandri Engl. Sapindus saponaria L. Stephania abyssinica (Dillon & A. Rich.) Walp. faxus paccata | ES 1 aDC ee ae ae Tephios ia pulpated (L.) Pers. 1 Te ephrosi Ta) vogelii i 100k. fi ] Thevetia ies iana (Pers.) Schum. 1 1 Verbascum phlomoides L. ] ] ae globosa (Blanco) Merr. 1 Total: 56 3 Species 30 14 12 6 12 0 2 Screened: 1 Percent active: an 6% 20.69% 9.66% 8.28% 4.14% 8.28% 0.00% 1.38% 2236 APPENDIX VI. BRIT.ORG/SIDA 21(4) ANTITUMOR ACTIVE PLANTS USED AS ARROW, HOMICIDAL, AND/OR ORDEAL POISONS Species Tumors SA CA Other Abrus precatoriu Acokanthera ees ia Stapf Acokanthera oblongifolia (Hochst.) L.E cl kanthera oppositif ee Acokanthera schimperi (A. DC.) Schweinf. AA, ; fp RQ Balf. f, sla alt ag oe (Dennst.) Nicols Peal toxicaria Fup ex Pers.) Lesch. Boophone disticha H nhyllim inonhy Se ok procera (Aiton) Dryander ex Aiton f. Canthium comprosoides F.Muell. Cassine crocea (Thunb.) Kuntze Cerbera mangas L. Cheiranthus cheri L. Derris trifoliata Lour. Erv thy, hlo africa 7 to (Benth.) Harms bh rlpalak | } Tremaut ex Kotschy Fagara macrophylla (Oliv.) Engl. Glor riosa Supe erba L. } uu UTIHIUIe mancinella L. Jatropha curcas L. la ; rl ty) Correa Lophopetalum javanicum (Thunb,) Kuntze Lunasia amara Blanco Parkia filicoidea Welw. ex Oliv. Rauvolfia mombasiana Stapf Securidaca longipenduculata Fresen. trophanthus courmontii Franch. tronhanthys hispi us DC , Tephrosia vogelii Hook. f. Thevetia peruviana (Pers.)Schum. Total: 32 Active Species Screened: 70 Percent active: 45.7% 30.00% 18.57% 7.14% 1.43% 1.43% SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2237 ACKNOWLEDGMENTS I thank Gordon Cragg and three anonymous reviewers for their critical review and comments and Susan Spjut for her editorial suggestions. REFERENCES Airy SHaw, H.K. 1973. J.C. Willis’ dictionary of flowering plants and ferns (ed.8), Cambridge Univ. Press, London. Assor,B.J. 1976. Bioassay of plant extracts for anticancer activity.Cancer Treat.Rep.60:1007— 1110. AMERICAN HerBaL PHARMACOLOGY DELEGATION. 1975. Herbal pharmacology in the People’s Re- public of China. National Academy of Sciences, Washington, DC. Banoer, N.H., D.M. Nanus, M.I. Micosky, L. KosTakoGLu, S. VALLABAHAJOSLUA, and S.J. GOLDSMITH. 2003 Targeted systematic therapy of prostrate cancer with a monoclonal antibody to pros- trate-specific membrane antigen. Semin Oncol 30:667-677. Barctay,A.S.and R.E. Perbue, Jr. 1976.The distribution of anticancer activity in higher plants. Cancer Treat. Rep.60:1081—-1113. Bevin, M.,D.B. Firzceraco and N.Feux. 1952a.Tumor-damaging capacity of plant materials. II. Plants used as diuretics. J. Nat. Cancer Inst. 13:741-744. Beckin, M.and D.B. Fitzcerato. 1952b. Tumor-damaging capacity of plant materials. |. Plants used as cathartics. J. Natl. Cancer Inst. 13:139-155. Beckin, M.and D.B.Fitzcerato. 1953a. Tumor-damaging capacity of plant materials. V. Miscel- laneous Plants. J. Natl. Cancer Inst. 14:607-625. Bevin, M.and D.B. FitzcerAto. 1953b. Tumor-damaging capacity of plant materials. IV. Coni- fers. J. Natl. Cancer Inst. 13:895-903. Betkin, M.and D.B. Fitzcerato. 1953c. Tumor-damaging capacity of plant materials. Ill. Plants used as pesticides. J. Natl. Cancer Inst. 13:889-893. Boyp, M.R. The future of new drug development. In J.E. Neiderhuber, ed. Current Therapy in Oncology, B.C. Decker, Inc., Philadelphia. Pp. 11-22. BUCKINGHAM, J. 1993-2005. Dictionary of natural products. Chapman and Hall/CRC Press, 1993 and hardbound supplements, 1994-1997, and CD ROM supplements, published at 6 month intervals. BurkHitt, LH. 1935. A dictionary of the economic products of the Malay Peninsula. 2 Vols., London. Cassapy, J.M.and M. Sureness. 1980. Terpenoid antitumor agents. In J.Cassady and J. Douros, eds. Anticancer agents based on natural product models, Academic Press, New York. Pp. 201-269 Carter, S.E.and R.L. Liuneston. 1976. Clinical evaluation of drugs from plants. Cancer Treat. Rep. 60:1141-1156. CHopra, R.N.,S.L. Nayar, and |.C.CHopra. 1956. Glossary of Indian medicinal plants. Council of Scientific and Industrial Research, New Delhi. Craae, G.M., JE. Simon, J.G. Jato, and K.M Snaper. 1996. Drug discovery and development at 2238 BRIT.ORG/SIDA 21(4) the National Cancer Institute: Potential for new pharmaceutical crops. In: J. Janick, ed. Progress in New Crops, ASHS Press, Arlington, VA. Pp. 554-560. Cuenbet, M.and J.M. Pezzuto. 2004. Antitumor activity of bruceantin:an old drug with new promise. J. Nat. Prod.67:269-272. Daiziet, JM. 1937.The useful plants of West Tropical Africa. Crown Agents for the Colonies, London. FARNSWORTH, N.R., A.S. BinceL, H.H.S. Fone, A.A. SALEH, G.M. CHRISTENSON, and S.M. SaurFereR. 1976. Oncogenic and tumor-promoting spermatophytes and pteridophytes and their active principles. Cancer Treat. Rep. 60:1171-1214 FirzGerAto, D.B., J.L. Hartwett and J. Leirer. 1958. Tumor-damaging activity in plant families showing antimalarial activity. Amaryllidaceae. J. Natl. Cancer Inst. 20:763-774. Geran, R.E., N.H. Greenserc, N.M. Macoonato, A.M. Schumacher, and BJ. Abbott. 1972. Proto- cols for screening chemical agents and natural products against animal tumors and other biological systems. Third ed. Cancer Chemotherapy Rep., Part 3(3):1-103. Gotoin, A.,S. Carter and N. Manet. 1974. Evaluation of antineoplastic activity: requirements of test systems. In O. Eichier, A. Farah, H. Herken, and A.D. Welch, eds. Handbook of ex- perimental pharmacology, new series, Springer Verlag, Berlin and New York. Pp. 12-32. Gustarson, K.R., J.H. CARDELLINA II, J.D. McMaron, R.J. Gutakowski, J. IsHitoya, Z. SZALLASI, N.E. Lewin, P.M. Bumpers, O.S. Weistow, J.A. Beutier, R.W. BuckHelr, Jr, G.M. Craca, PA. Cox, J.P Baber, and M.R. Boyo. 1992. A nonpromoting phorbol from the Samoan medicinal plant Homalanthus nutans inhibits cell killing by HIV-1.J.Med. Chem. 35:1978-1986. Hardin, J.W. and J.M. Arena. 1974. Human poisoning from native and cultivated plants. Second ed., Duke Univ. Press, Durham, NC. Harincton, J.S. 1969. Herbal Davis remedy, mixture—Maytenus heterophylla, Sanseviera sp. and Scutia myrtina, a written comm. to J. Hartwell at the NCI from the Head of The Cancer Research Unit of the National Cancer Association of South Africa, South Afri- can Institute for Medical Research. Hartwett, J.L. 1960. Plant remedies for cancer.Cancer Chemother. Rep. 7:19-24. Hartweit, J.L. 1967-1971. Plants used against Cancer. A survey. Lloydia 30:379-436, 1967 and ten additional installments ending with 34(4):286-437, 1971. Hartwell, J.L. 1976. Types of anticancer agents isolated from plants. Cancer Treat. Rep. 60:1031-1067. Hartwett, J.L.and MJ. SHear. 1947.Chemotherapy of cancer:Classes of compounds under investigation and active components of podophyllin. Cancer Res. 7:716. Hartwett, J.L. and BJ. Assorr. 1969, Antineoplastic principles in plants: Recent develop- ments in the field. In S. Garattini, A. Goldin, F Hawking and LJ. Kopin, eds. Advances in Pharmacology and Chemotherapy, Vol. 7, Academic Press, New York and London. Pp. 117-2 Jacosson, M. 1989. Focus on phytochemical pesticides. Vol.|, The neem tree.CRC Press, Inc., Florida. JoHNsTON, M.C. 1971. Revision of Colubrina. (Rhamnaceae). Brittonia 23:2-53. SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2239 Kincspury, JM. 1964. Poisonous plants of the United States and Canada. Prentice Hall, Englewood Cliffs, NJ. KrocHMal, A.C. 1973.A guide to the medicinal plants of the United States. Quadrangle, the New York Times Book Co, New York. KUPCHAN, S.M., Y.Komopa, W.A. Court, GJ. THOMAS, R.M. Smith, A. Karim, C.J. Kitmore, R. C. HALTIWANGER, and R-F. Bryan. 1972. Maytansine, a novel antileukemic ansa macrolide from Maytenus ovatus. J. Amer. Chem. Soc. 94:1354—-1356. KuUPCHAN, S.M, R.W. Britton, M.F. Siccter, and.M.F. Siecet. 1973. Bruceantin, a new potent anti- leukemic simaroubolide from Brucea antidysenterica. J. Org. Chem. 38:1 78-179. KupcHAN, S.M. 1976. Novel plant-derived tumor inhibitors and their mechanisms of action. Cancer Treat. Rep. 60(8):1 115-1126. Larson, G.M., B.T. SHANEBERGT and T. SNEDEN. 1999. Two new maytansinoids from Maytenus buchananii. J. Nat. Prod. 62:361-363. Lesser, W.H. 1997. International treaties and other legal and economic issues relating to the ownership and use of genetic resources. In: K.E. Hoagland and A.Y. Rossman, eds. Global genetic resources: access, ownership, and intellectual property rights. Associa- tion of Systematics Collections, Washington, DC. Pp. 31-50. Mata-Greenwooo, E., J.F. Dacuste, PA. Grieco, J. Dou, J.D. McCuesney, J.R. Menta, A.D. KINGHORN, and J.M. Pezzuto. 2001. Novel esters of glaucarubolone as inducers of terminal differentia- tion of promyelocytic HL-60 Cells and inhibitors of 7,12-dimethylbenz[aJanthracene- induced preneoplastic lesion formation in mouse mammary organ culture. J. Nat. Prod. 64:1509-1513. Moraes, R.M., H. Lata, E. Beoir, M. MAgeoot, and K.CusHmAn. 2002. The American mayapple and its potential for podophyllotoxin production. In:J. Janick and A.Whipkey, eds. Trends in New Crops and New Uses. ASHS Press, Alexandria, VA. Pp. 527-532. Morton, J. 1968. A survey of medicinal plants of Curacao. Econ, Bot. 22:87-102. Morton, J. 1974. Folk Remedies of the Low Country.E.A. Seemann Publ. Co., Miami. Munz, PA. 1959. A California Flora. Univ. California Press, Berkeley and Los Angeles (3"¢ Printing). NATIONAL Cancer Institute (NCI). 1977. Confirmed Plant and Animal Materials (CPAM) Re- ports. A Computer Printout, Drug Evaluation Branch (unpublished), Developmental Therapeutic Program, Division of Cancer Treatment, dated 30 September 1977, plant materials, 606 p.; animal materials, ~100p. Newman, D., G.M. Cracc, and K.M. Snaver. 2003. Natural products as sources of new drugs over the period 1981-2002. J. Nat. Prod., 66 (7), 1022-1037. Perbue, R.E., Jr. 1976.Procurement of plant materials for antitumor screening. Cancer Treat. Rep. 60:987-998. Peroue, R.E., Jr. 1982. KB Cell Culture: |. Role in discovery of antitumor agents from higher plants. J. Nat. Prod. 45:418-426. Peroue, R.E., JR. and J.L. Hartwett. 1969. The search for plant sources of anticancer drugs. Morris Arbor. Bull. 20(3):35-53. 2240 BRIT.ORG/SIDA 21 = 4) Peroue, R.E., Jr., R.L. Smit, M.E. Watt, J. Hartwett, and B. Asgorr. 1970. Camptotheca acuminata Decaisne [Nyssaceae] Source of Camptothecin, an Antileukemia Alkaloid, USDA Tech. Bull. 1415. QuisumBine, E. 1951, Medicinal plants of the Philippines. Tech. Bull. 16, Philadelphia, Dept. Agric. and Nat. Res., Manila. Reis ALTSCHUL VON, S. 1973. Drugs and foods from little-known plants. Harvard Univ. Press, Cambridge, MA. Sieger, S.M., J.A.R. Meao, and R.H. Avamson. 1976. Pharmacology of antitumor agents from higher plants. Cancer Treat. Rep. 50:1127-1139. Spyut, R.W.and R.E. Peroue, Jr. 1976. Plant folklore: A tool for predicting sources of antitumor activity? Cancer Treat. Rep. 60:979-985. SpyuT, R.W. 1985. Limitations of a random screen: Search for new anticancer drugs in higher plants. Econ, Bot. 39:266-288. STANDLEY, PC. 1922-1926. Tress and shrubs of Mexico. Contr. U.S. Natl. Herb., Washington Govt. Printing Office, Vol. 23. Starz, D.and F.B. Coon. 1976. Preparation of plant extracts for antitumor screening. Cancer Treat. Rep. 60:999-1005. Sureness M.and J.Douros. 1979. Drugs of plant origin.In V.T. DeVita and H. Busch, eds.Meth- ods of Cancer Research Vol. 16:73-126. STEINMETZ, E.F. 1962. An effective anticancer drug? Fritillaria cormus and its possibilities for the cure of other stubborn diseases. Quarterly J. Crude Drug Res. 2:285-298, Train, P,, J.R. HenricHs and W.A.. ArcHer. 1957. Medicinal uses of plants by Indian Tribes of Nevada. Contributions toward a flora of Nevada, No. 45. Revised ed. with summary of pharmacological research by W.A. Archer. A series prepared through the cooperative of the National Arboretum and the Plant Introduction Section, USDA, ARS, Beltsville, MD. Unite States Department oF Acricutture (USDA). 1980. Combined list of genera extensively screened or completed (GESOC) and materials collected 6 or more times. Revised Feb. 14 (unpublished, available at www.worldbotanical.com), Watt,M.E.,M.C.Wani, CE. Cook,and KH. Paumer. 1966.Plant antitumor agents.|. The isolation and structure of camptothecin, a novel alkaloidal leukemia and tumor inhibitor from Camptotheca acuminata. J. Amer. Chem. Soc. 88:3888-3890. Watt, M.E.,M.C. Want, and.H. Taytor. 1976.Isolation and chemical characterization of antitu- mor agents from plants.” Cancer Treat. Rep. 60:1011—1030. Want, M.C.,H.L. Taytor, M.E. Watt, P. Coccon, and A.T.McPuxait. 1971.Plant antitumor agents. VI. The isolation and structure of taxol, a novel antileukemic and antitumor agent from Taxus brevifolia. J. Amer. Chem. Soc, 93:2325-2327. Wan, M.C.,H.L. Taytor, and M.E. Watt. 1973. Plant antitumor agents: colubrinol acetate and colubrinol, antileukaemic ansa macrolides from Colubrina texensis. Chem.Commun.P. a Watson, A.M.1983.Agricultural innovation in the early Islamic world. The diffusion of crops and farming techniques, 700-1100. Cambridge Univ. Press. SPJUT, HISTORICAL REVIEW OF PLANT FOLKLORE AND ANTITUMOR ACTIVITY 2241 Watt, JM. and M. Brever-BRANWUK. 1962. The medicinal and poisonous plants of southern and eastern Africa (2nd ed).E and S Livingstone, Ltd., Edinburgh and London. Wess, L.J. 1948. Guide to the medicinal and poisonous plants of Queensland. Bull. No. 232, CSIRO, J. J. Gourley, Govt. Printer, Melbourne, Australia. Weiner, M.A. 1972. Earth medicine-earth foods (plant remedies, drugs, and natural foods of the North American Indians). MacMillan Publ. Co. Wits, J.C. 1922. Age and area. Cambridge Univ. Press. Zouary, D. and P. Spiecet-Roy. 1975. Beginnings of fruit growing in the Old World. Science 187:319-327, 2242 BRIT.ORG/SIDA 21(4) Book NortICES Timber Press JAMES CULLEN (text) with photographs by Despite Waite and Friepa Curistie. 2005. Hardy Rhododendron Species: A guide to Identification. (ISBN 0-88192- 723-6, hbk.). Published in association with the Royal Botanic Garden Ed- inburgh and Timber Press Inc. 133 S.W. Second Ave, Suite +50, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $49.95, 496 pp., color photos, 7" x 9", peg Oy era | “eR +} th : ct : } ,-} 1. J | ee | ] PUPLISHLET Scomments, rOr mark reference provides the keys to the accurate ideutiteston of the nearly 300 dhodédendion spe- cies widely in cultivation. An extensive introduction places the work in context as it examines the history of Rhododendron aoe ries and g hes a full survey of plant structures throughout the d many species. Species are lis ) that similar species occur close to each other in the text. These are fully described, nti complete citations of previous references and notes n the occurrence of wild-origin specimens in cultivation. Beautiful photographs include close-up ee ol flower and leaf, microscope images sof leaf surfaces, and easy-to-use diagnostic keys, mak- ing this the indisputable volume for plant identification. A milestone in the identification of rhodo- >n. and t dendrons, this will become an essential reference for botanists, nurse rym Each taxonomic entry includes a description of one or more of the following: Synonymy, paileee tration, Habit, Leaves, Vegetative Buds, Inflorescence Buds, Calyx, Corolla, Nectary, Stamens, Pollen, Ovary, Style, Stigma, Fruit, Seeds, Distribution, Flowering, a Zone, and additional notes. Taxonomic keys are provided for subgenera, sections Piet Oupoi_r and Nori Kincsspury. 2005. Timber Press Pocket Guide to Bulbs. (ISBN 0-88192-740-6, hbk.). Timber Press Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $34.95, 175 pp., color photos, 9 1/2" x 111/2" Publisher comments.—‘This is a must-have book for eceigies urban ee andseaDe architects, and avid gardeners. The authors offer a planting philosophy as . for instance, questions of scale and movement such as positioning entrances and paths, creating connective lay- outs, using plants as barriers and as conduits, and maintaining the designed space as plants grow” SIDA 21(4): 2242. 2005 ETHNOBOTANICAL REPORT FROM MANGROVES OF PICHAVARAM, TAMIL NADU STATE, INDIA K. Venkatesan V. Balakrishnan Department of Botany Department of Botany Annamalai University Annamalai Univ ersity Annamalainagar — 608 002 Annamalainagar — 608 002 INDIA K.C. Ravindran V. Devanathan| eee of Botany Loe in Botany malai University niversity of Madras Peace ee 608 002 Guindy Craps INDIA Chennai — 600 005, INDIA ABSTRACT Ethnobotany deals with the study of the natural and traditional interrelationships between man and plants. Mangroves are one of the primary — eatures of coastal areas and play a significant role in coastal ee Mauer a areas are economically exploited, epee for caaaee and fishery products. A bout other a ts h d from pec mangroves, is fee freely available. ih his paper reviews the utilization of the mangroves as source of medicinal pega for the pichavatatn coastal People. Af HonstiG suEvey was gundentaisen and 11 A list of pant species along with the plan Be art/s used and their mode of application reroute to be efficacious for diff RESUMEN La Etnobotanica trata del estudio de las interrelaciones naturales y tradicionales entre el hombre y las plantas. Los manglares son una de las caracteristicas primarias de las areas costeras y juegan un papel importante en el desarrollo costero. Las areas de mangl ar son i anay Sou cenen epeelaigite para madera y pronucies pesqueros. Un 4 manglares, esta menos ote Este article revisa el uso re ee panelicse como heains de ies medicinales por los habitantes costeros de near aram. Se moe un di ico y se encontraron LI especies vegetales etnomedicinales de Pic es avaram. Se aporta una usta ae especies a as con la parte o partes usadas y su ied de uso para que sean efi INTRODUCTION Mangroves are assemblages of trees and shrubs that grow in the intertidal re- gion of tropical and subtropical coastlines, in areas where river water mixes with sea water. Mangroves have two components, mangrove forests and their hennai '¢ | ti I } liffmail CAS inB indy Campus, U 600 005, India. SIDA 21(4): 2243 - 2248. 2005 2244 BRIT.ORG/SIDA 21(4) associated water bodies. A group of woody trees and shrub that can grow well in saline water and logged condition constitute the forest component (Selvam ®& Karunakaran 2004). Pichavaram is in the North-East coastal part of Tamil Nadu State and lies between Lat.11° 27' N; Long. 79° 47' E. The Pichavaram mangrove wetland is located in the northern extremity of the Cauvery delta, near the mouth of the Coleroon River. Its total area is about 1,350 ha, with its many small islands being colonized by thirteen true man- grove species. Remote sensing data indicates that nearly 54% of the mangrove forest of Pichavaram (total forested area 700 ha, excluding water bodies, sand dunes etc.) was in a degraded state in 1986. Analysis of the remote sensing data of 1996 showed that the mangrove forest cover has increased by about 60 ha compared to the area in 1987, which is mainly due to the restoration efforts being undertaken collaboratively by the Tamil Nadu Forest Department and M.S. Swaminathan Research Foundation along with the participation of the local communities. Pichavaram mangrove receives fresh water mostly during the northeast monsoon season from October to November. It is reported that 13 species are present in the mangrove wetlands, Tamilnadu; these include two species of Rhizophora in Pichavaram, which are regarded as endangered spe- cies, a fact that underlines the importance of the Pichavaram mangrove froma biodiversity stand point. Currently there is an increasing desire to unravel the centuries old secrets of traditional medicines. In India, ethnobotanical studies witha good scientific base have appeared in last 1-5 decades (Chopra et al. 1956; Jain 1963, 1965; Jain & Borthakur 1980; Bhargara 1981; Yadav @ Patil 2001). MATERIALS AND METHODS An ethnobotanical survey of the forest people of Pichavaram mangrove was un- dertaken during the 2001-2002. Through repeated interactions and participa- tory rural appraisal (PRA), details on the eeeere ay of me plants used by the folk population were gathered with vill 1s described by Jain (1983). According to Jain (1987) enples: of lants were collected, identified and voucher specimens were deposited in the Department of Botany, Faculty of Sci- ence, Annamalai University, Annamalainagar, Tamilnadu. The collected plants were identified using the flora of the Presidency of Madras (Gamble & Fischer 1915-1935) This paper describes the most important medicinally used plants. RESULTS The traditional knowledge system in Tamilnadu State is fast eroding. There is an urgent need to record all ethnobotanical information among the diverse eth- nic communities before the traditional culture is completely lost. As of result of the present study the ethnobotanical use of 11 plant species belonging to 8 VENKATESAN ET AL., ETHNOBOTANY OF PICHAVARAM MANGROVES, INDIA 2245 families have been reported from the Pichavaram mangrove area. A range of preparations are used to treat diseases. The most popular medicinal prepara- tions are plant extract, decoction, paste and juice. Information on botanical name, herbarium number, local name, family, diseases cured/uses, parts used and mode of application are given in the Table 1. DISCUSSION AND CONCLUSION There is an increasing demand for the production of healthcare medicines and cosmetic items from plant origin based on the ancient knowledge in folk rem- edies. Plant sources are being identified for further investigation for their phar- macological properties and efforts are also being made to investigate the ac- tivities of isolated individual constituents. Eleven species of mangroves were identified as medicinally important for the treatment of snakebite, skin diseases, dysentery, urinary disorders as well as for birth control and as blood purifiers. Communities (Irulars and Meenavars) living close to and interacting with the Pichavaram mangrove wetlands have gained a unique cognitive understand- ing of the medicinal values of the plants within the mangroves. Experience has led to a rich knowledge system that is reflected in the native classifications of mangrove wetlands. The role of ethnobotanical studies is of crucial importance as some mi- raculous medicines for incurable diseases are known to the local communities As knowledge acquired through the ages is usually passed on from generation to generation, it is necessary to popularize the identity and utility of these me- dicinal plants. Management systems are losing their value because of various factors. Any assistance to protect these traditional systems will win the confidence of the local people; this in turn will be the first step toward the success of commu- nity-based mangrove conservation and management. The collection and docu- mentation of traditional knowledge on ethnomedicines is important for find- ing new, effective safe drugs for a range of ailments. However, to give more support to the above findings, further investigation is needed for the isolation of the active principles and for the pharmacological evaluation of different medicinal plant species. The estimation of the number of endangered medicinal plants is rising day by day Jain 1992). Medicinal plants are threatened due to over exploitation, shifting cultivation, deformation and environmental pollution (Karuppusamy et al. 2002). The conservation of medicinal plants not only saves local lives but also improves the socio economic condition of the people where the plants ex- ist. The following strategies have been suggested for the conservation of me- dicinal plants. Taste 1. Ethnobotanical report from mangroves of Pichavaram, Tamil Nadu State, India. Latin Name/Family Local Name Disease Cured/Uses Parts Used Mode of Application (Voucher Specimen) Acanthuls ilicifolius L., Kazhimulli Snakebite Fruits Crushed fruits are made into a dressing canthaceae (AUB 102) and applied as a dressing to the snake bite. Kidney stones Whole plant The whole plant is boiled in water and the patient drinks the final solution, half of a glass each time until the signs and symptoms of the kidney stones disappear. Skin diseases, small Whole plant A whole plant paste is applied to the affected pox, health promotion, body for skin diseases. The juice is orally given detoxification and ulcer for other diseases. Avicennia alba Blume Vellaikandal Birth control Resin Resinous substances are mixed with water and Beebe (ave 108) given orally. A Narikandal Bone pain, urinary Leaves The leaves are boiled in water and the extract Avicenniaceae nes 110) disorders, bronchial is given orally asthma, stomach disorders and detoxification guiera cylindrica (L.) Blume, Pannukkuchi Tumor inhibitors Leaves ee contain alkaloids. A leaf paste is applied Rhizophoraceae (AUB 532) constipation o the tum Whole plant i. whole pre is boiled in water and the solution is drunk twice daily after meals Clerodendron inerme Gaertn Peenarisangu Pain, jaundice, Leaves A leaf paste is used Verbenaceae (AUB 428) washing dishes ae infected wounds, anti-inflammation and itching skin Sap of leaves Leaf extract Leaf sap is used Leaf extract and paste is used. 9b77Z (p)LZ VaIS/9¥O'LINA Taste 1. (continued) Latin Name/Family (Voucher Specimen) Local Name Disease Cured/Uses Parts Used Mode of Application [= ; I hon Euphorbiaceae (AUB 483) /pomoea pes-caprae (L.) Sweet, Convolvulaceae (AUB 325) Rhizophora mucronata Lam., Rhizophoraceae (AUB 372) Rhizophora apiculata Blume, Mullen ae 333) ane ee ee ae 238) Xylocarpus granatum K.D. eni Meliaceae (AUB 267) Thillai Adappankodi or Kuthiraikulambu Kanthal Kandal Kozhikali or Seethavavazham Somundri Toothache Latex Anti-inflammation Roots iene and swelling Whole plant Leaves at dise av Fatigue Seeds Diarrhea, nausea and Bark vomitin g Diarrhea, vomiting, ameobiasis Bark and stop bleeding h ltches Whole plant ash ysentery Bark Illumination of hair Oil Diarrhea and cholera Bark A latex extracted from leaves and bark is applied to the toot Fresh root material is used. A whole plant extract is used. A leaf paste is use Crushed seeds are used A bark extract is given orally. A bark extract is given orally. The plant ash is applied to the skin. A bark extract is used. Oil extracted from the seeds is applied to the hair. A bark decoction is given orally. VIGNI‘SJAQHONYW WYYVAVHDId 40 ANVLOSONHL] “Tv 13 NVSILVNIA Lyez 2248 BRIT.ORG/SIDA 21(4) a) Control over exploitation b) Establishment of mangroves c) Cultivation of rare and endangered medicinal species d) Awareness creation of the utility and conservation of medicinal plants to local communities ACKNOWLEDGMENT The authors wish to express their gratitude to K.P Chellappan, Dean, Faculty of Science (retired) and R. Panneerselvam, Professor and Head, Dept. of Botany, Annamalai University, for their encouragement and help in various ways. Two anonymous reviewers greatly improved the manuscript. REFERENCES BHarcava, N. 1983. Ethnobotanical studies of the tribes of Andaman and Nicobar islands, India. Onge. Econ. Bot. 37:1101- Cuopra, R.N., S.L. Nayar, and 1.C. CHopra. 1956. Glossary of Indian medicinal plants. Council of Scientific and Industrial Research, New Delhi, India. Gamete, J.S.and C.F.C. FiscHer.1 915-1935. Flora of the Presidency of Madras.London (issued in Il parts: 1-7 by Gamble, 8-11 by C.E.C. Fischer), vols. (1-3) Calcutta. Jain, S.K. 1963. Studies in Indian ethnobotany-less known uses of fifty common plants from tribal areas of Madhya Pradesh. Bull. Surv. India 5:223-226. JAIN, S.K. 1965. Medicinal plant lose of the tribals of Bastar. Econ. Bot. 19:236-250. JAIN, S.K. 1983. Ethnobotany in India — An overview. BSI, Howrah, India. Pp. 1-37. Jain, S.K. 1987.A manual of ethnobotany, Oxford Publishers, Jodhpur, India. Jain, S.K. 1992. Ethnobotanical aspects of Indian tropical ecosystems. In: K.P. Singh and J.S. Singh, eds. Tropical ecosystem, ecology and management. Wiley Eastern Ltd., New Delhi, India. Pp.507-511. Jain, S.K.and S.K. BortHakur. 1980.Ethnobotany of the mikirs of India. Econ. Bot. 34:264-272. KARUPPUSAMY, S., K.M. RAJASEKARAN, and N. KarMeGam. 2002. Medicinal plant resources of Dindigul district of Tamil Nadu. J. Natcon 14(1):151-158. Sewvam, V.and V.M. KARUNAKARAN. 2004. Ecology and biology of mangroves.M.S. Swaminathan Research Foundation, Taramani, Chennai: 7. Yaoay, S.S. and H.S. Patit. 2001. Traditional medicines and healthcare system of tribals of Satpuda Region, Maharashtra State. Plant Archives 1(1&2):111- PINGUICULA VULGARIS (LENTIBULARIACEAE) AND ITS USES IN NORWAY Torbjorn Alm Department of Botany Troms@ Museum University of Troms@ N-9037 Troms@, NORWAY ABSTRACT Pinguicula vulgaris isa common species in Norway, and well known in folk tradition, not least for its reputed use to curdle milk. Most vernacular names recorded so far, e.g. the widespread tettegras or tettegress (“rennet grass” or “thickening grass”) reflect this use. The fat leaves have also found some sin folk medicine, e.g. to treat wounds and ringworms, and in folk bac medicine, mainly to -| treat sore teat. There are also a few records of P. vulgaris being in magic, and in children’s games. £ c NORWEGIAN SUMMARY /NORSK SAMMENDRAG Pinguicula eeu er vennee i Norge, og godt ene) i folketeaaiS et ikke minst for sin virkning f.eks. det vidt utbredte tettegras eller tettegress. Bladene har ee anne en viss aumendalse i foleete en bla. til a behandle sar og ringorm, og i folkelig veterinaermedisin, seerlig som en kur for sare spener. I noen fa ilfeller har P. vulgaris ogsa tjent som vernerad, i magi og i barnelek. INTRODUCTION Of the three Pinguicula species found in Norway, P. vulgaris L. is by far the most widespread. It is found almost throughout the country, and in many different environments, from near sea level to 1570 m a.s.l. in interior southern Norway (Elven 1994:559). P. alpina L. is more demanding in terms of habitats, preferring calcareous substrates, and the tiny P villosa L. is restricted to Sphagnum hum- mocks on oligotrophic mires in interior SE Norway and northeastern North Norway (Alm 2000). All three species share the typical characteristics of the genus Pinguicula: a rosette of insect-trapping leaves, secreting a viscous, enzymatic fluid in the presence of small insects or other prey (Casper 1966; Heide 1912; Heslop-Harri- son & Heslop-Harrison 1980, 1981; Heslop-Harrison & Knox 1971; Legendre 2000; Warming 1886). P. vulgaris belongs to the subgenus Pinguicula (Casper 1966; Legendre 2000; cf. Jobson et al. 2003). Christen (1961) studied the species from a pharmacological point of view. In Scandinavian folk tradition, the leaves of Pinguicula vulgaris have been used as tettegress, “thickening grass” or “rennet grass,” to treat or preserve milk, but how this was done, the characteristics of the product, and the mechanism SIDA 21(4): 2249 — 2274. 2005 2250 BRIT.ORG/SIDA 21(4) behind are disputed (Brondegaard 1971). The use for dairy products in particu- lar has attracted the attention of numerous authors, e.g. Bergsaker (1982), Brondegaard (1951, 1971), Evjen (1986), Forsén (1966), Gisler (1749), Gunnerus (1774), Jaeger-Lejrvik 1959), Olsen-Sopp (1912), Rank (1960, 1971), and Weisaeth (1990). Hoeg (1974) provides much information on the ethnobotany of P vul- garis in Norway. Larsson (1988) carried out an extensive survey of vernacular names in the Nordic countries (see also comments in Hansson 1990). Wix (1995, 1996) provided some notes on traditional uses in Scandinavia, but the language barrier kept him from checking more than a few references. —y SCOPE AND SOURCES This paper aims at a comprehensive review of ethnobotanical traditions related to Pinguicula vulgaris in Norway. It should be noted that P alpina may also have served as tettegress, although all sources that identify the species (and not only the genus) mention only P. vulgaris (e.g. Bergsaker 1982; Bjorndal 1949; Brondegaard 1971; Gunnerus 1774; Hoeg 1974; Kirkevoll 1940; Sortdal 1981; Tonning 1773), or note that the plant used had blue flowers (e.g. Klonteig 2000 and seven letters in NFS Manum). Several voucher specimens confirm the iden- tification as P. vulgaris, e.g. in thel8th century herbarium of J.-E. Gunnerus (in TRH), the 1837-39 herbarium of the linguist Ivar Aasen (Lid 1941), and some vouchers in TROM. Throughout Norway, P vulgaris or tettegras/tettegress is well known in folk tradition. Numerous authors mention its use in milk. Much useful information is found in the vast collection of Norwegian ethnobotanical data assembled by Ove Arbo Hoeg in the period 1925 to 1973. However, his large volume (Hoeg 1974) makes little use of previous publications. In addition to data from about seventy publications pr on Nor- shival data have been incor bala here, mainly from NEG (Norsk etnologisk gransking/Norwegian ethnological survey), NFS (Norsk folkeminnesamling/Norwegian folklore collection) and NOS (Norsk ordbok, seddelarkivet/Norwegian dictionary, card archive). Furthermore, some data have been excerpted from my own ethnobotanical records, mainly from North Norway; these are referred to as EBATA + year and record number. Informants are not identified here; transcripts and some recordings of the original inter- views are stored at the Department of Botany, Tromso Museum (TROM). Un- less otherwise stated, all citations have been translated from Norwegian. lat wegian traditions, s Vernacular names No record of any Norse name for Pinguicula vulgaris seems to exist; the species is not mentioned by Heizmann (1993). The oldest surviving record, Marie sko (“Mary’s shoe”), was made at Bergen in 1599, and is found in the diary of Sivert Grubbe (Rordam 1873). It isan unusual name (see Table 1), although a couple of ALM, ETHNOBOTANY OF PINGUICULA VULGARIS IN NORWAY 2251 other vernacular names referring to the Virgin Mary are known. They are obvi- ously younger than the Christianization of Norway,and must have been coined after AD 1000. Most Norwegian vernacular names for Pinguicula vulgaris reflect its use for making tettemelk or “thickened milk” (Fig. LD. Tette may be translated as “ren- net” (otherwise known as kjase or lope in Norwegian), but the etymological meaning or root is “(make) thick, thight, compact” (Bjorvand & Lindeman 2000; Torp 1919). A straightforward tettegras or tettegress (“rennet grass” or “thicken- ing grass”), with some dialectal variations, predominates over large areas (Table 1). Exceptionally, it may also occur in place-names, e.g. Tettgrasmyra in Trysil (Kvernbekk 1979:64). Kjasegras/gress “rennet grass”) reflects similar use. Sev- eral names on melhk-, mjelk- and mjalk- (“milk”) refer to the use in dairy prod- ucts. Other vernacular names refer to the fat and/or slimy appearance of the leaves, e.g. feitgras, “fat grass” (Tonning 1773:4), sleipgras, “slippery grass” (Hoeg 1974:473), slimgras, “slime-grass” (Tonning 1773:4), sapeblomme, “soap flower” (Halvorsen 1988:188), and the North Sami vuodjalasta “butter leaf”). Melkekors “milk cross”) and other names containing kors or kross (“cross”) obviously refer to the leaf rosette. According to Brondegaard (1971), it may have reminded people of the cross-shaped lower end of traditional churning sticks, but available records would rather suggest that people thought that the leaf rosette looked like a cross or star. Pinguicula vulgaris in pastures Pinguicula vulgaris is hardly a very frequent pasture-grass for cattle or sheep. In some areas, the plant was considered harmful, by adversely affecting milk. People believed that milk would become thick if the cows consumed P. vulgaris, as recorded at Tolga, Trysil and Asnes in Hedmark (Furuset 1980:126; Reichborn- Kjennerud 1922:87; NFS Manum), @yer in Oppland (NEG 69:15278), Tjome in Vestfold (NEG 69:14491), Marnadal in Vest-Agder (NEG 69:15544), Jolster in Sogn og Fjordane (NEG 69:14404), and other stations in southern Norway (Hoeg 1974:491). In Gausdal (Oppland), Vestad (1984:60) was told that “the milk could become thick if the cows ate much of this grass.” At Trysil in Hedmark, people had noted that milk products could become thick and sour during the summer. In such cases, they believed that the cows had grazed tettgubbe Cthickening old man”), according to Furuset (1980:126) either P. vulgaris or Drosera spp. In Troms, N Norway, people claimed that such effects of Pinguicula vulgaris occurred mainly in the autumn (Alm 1983:393). Some believed P vulgaris to be poisonous, as reflected in the vernacular name sprengeras “burst grass”) in Central Norway (Hoeg 1974:492). It was said to be harmful for horses, sometimes for cattle or sheep. Strom (1762:112) noted that “afew” people at Sunnmore in W Norway considered P. vulgaris as harmful to sheep, though he does not specify in which way. A similar belief is reported 2252 BRIT.ORG/SIDA 21(4) Fic. 1.1n N gi f , Pil 9 uct based on a P vulgaris culture, “Tjukkmjalk,” byR ieriet, i le nationwide. | Iso the fi food product in N y with a “controlled origin” | J\ ehtel 1 i 1 4 “aL fod J+4° ra . L at VISCOUS Uldalt UlGl hal adel 1 nary (rennet-t fed | / from Telemark by Wille (1786), but people there also claimed that sheep avoided the plant. Farmers in Valdres, interior SE Norway, believed that it caused a liver disease in sheep (Kirkevoll 1940:174). At Singsas in Sor-Trondelag, Central Nor- way, P. vulgaris was believed to cause colic in cattle (Hoeg 1974:492). In other areas, no such negative effects on grazing animals were known. In his topo- graphical description of Gudbrandsdalen in SE Norway, Hiorthoy (1785) noted that “Whether this herb is harmful to the sheep, as stated by Mr. Strom, is not known here.” Folk medicine In Norwegian folk tradition, Pinguicula has found some, but restricted use for medicinal purposes. It was mostly used externally. A decoction of the leaves in water could be used to remove lice from children, and to promote the growth of fair hair: “When the leaves are boiled in water, and the children’s heads are washed with it, lice are purged, and the hair grows, and also gets a yellow colour” (Tonning 1773:5). Mohr (1786:152) noted similar use. In Hallingdal (Buskerud, SE Norway), an ointment was made by boiling Taste 1. Alohabetical list of Norwegian vernacular names for Pinguicula vulgaris (original spelling, if different from present-day Norwegian, is indicated). Mu- nicipalities (Communes) are given is possible. “Numedal area” and similar records indicate vernacular names that may derive from several municipalities within the given area. Norwegian English translation Area and source Adam og Eva Adam and Eve Troms: Kvaenangen (EBATA ee Blastjern Blue star Nord-Trendelag: Nordli (Haeg 19 ) Feitegras Fat grass Norway, unspecified (Reichborn- eect 1922:87) Feitgras Fat grass Norway, unspecified (Tonning 1773:4, as Feit-Gras) Feitgress Fat grass Norway, unspecified (Viborg 1793:15, as Feitgraes; Hornemann 1806:20, as Feitgrees) Feittstjerna Troms: S@rreisa (Haeg 1974:493) logfang Fly-catcher Nordland: Vefsn (Lundestad 1992:35; Oksendal 1977:99, 1993:110) Geitablom Goat flower Hordaland: Kvinnherad (Hoeg 1974:493) Geitmjolk Goat milk So@r-Trandelag: Selbu (Haeg 1974:493) Giftgr. Poison grass Se@r-Trondelag: Raros (NFS O.A. Hoeg) Sater Goat-fat-flowers Se@r-Trendelag: Bjugn: Stjorna (Hoaeg 1974:493) Gjetslek Goat's lick Hedmark: Tolga (Haeg 1974:493) Gjoketunge Cucoo’s tongue Hedmark: Elverum (Ha@eg 1974:493) Istegras Curdle Buskerud: go (NEG 69:1451 L Kinnekross Butter bucket cross Hordaland: Fusa; Sund: H florden; Sogn og Fjordane: Aurland; Balestrand (H@eg 1974:493): Nordland: Steigen (H@eg 1974:493) Kjzesegras Rennet grass Norway, unspecified (Reichborn-Kjennerud 1922:87); Telemark: Notodden: Bolkesj@ (Djupedal 1959:65), Seljord (Ross 1895:397), Tinn (Klonteig 2000:83) Kjaesegress Rennet grass Telemark (Wille 1786:122, as Kjaese-Grees) Kjerringkjeft Old woman's mouth Telemark: Vinje (H@eg 1974:493) Kjokkmjalkgras Korstroll ukors Maria taregress Marie sko Maritetta Thick-milk-grass Cross-troll Cow cross Mary's tear grass Mary’s shoe Mary’s rennet S@r-Trandelag: Soknedal (Haeg 1974:493) Ser-Trondelag: Holtalen: Alen (Haeg 1974:493) Hordaland: Fusa: Halandsdal (Haeg 1974:493) Nordland: Rana (unpublished note by A.Blytt 1870, as Mariee Taaregrees) Hordaland: Bergen (Rerdam 1873:405, diary note by Sivert Grubbe, July 6, 1599) Hordaland: Ulvik (H@eg 1974:492) AVMHON NI SIXYSTNA VINDINONId 10 ANVLOGONHLI ‘W1¥ £S7Z Tale 1. (continued) vS7Z Norwegian English translation Area and source Melkekors Milk cross Norway, unspecified (Viborg 1793:15; Hornemann 1806:20):; Sogn og Fjordane: Vagsey: Nordre Vagsay (Haeg 1974:493); Mare og sdal: Sunnmagre area (Strom 1756;fol. 66a, as Melcke-Kaarset; Strom 1762:111,as Mzelke-Kors) Melkekross Milk cross Sogn og Fjordane: Lzerdal (NFS Manum) Mjelkekors Milk cross Sogn og Fjordane: Farde, Naustdal (H@eg 1974:493), Jalster (NEG 69:14362): Mare og Romsdal: Sunnmare (Aasen 1860:10) Mjelkgras Milk grass Hordaland: Sund (Haeg 1974:493) Mjelkgress Milk grass Nordland: Grane (H@eg 1974:493) Mjelkkross Milk cross Hordaland: Sunnhordland area (Haeg 1974:493) Mjeltekors Milk cross Norway, unspecified (SchUbeler 1888:184); More og Romsdal: @rsta: Vartdal (Hoeg 1974:493) Mjolkegras Milk grass Hedmark: Eidskog, Os (Hoeg 1974:493) Mjolkekors Milk cross Hedmark: Eidskog (Fjellstad 1966:171);Vest-Agder: Farsund: Lista; Hordaland: Fusa, Kvam, Strandbarm, Stord, Tysnes; Sogn og Fjordane: Balestrand, Bremanger, Heyanger: Lavik, Leikanger, Laerdal, Sogndal, Vik; More og Romsdal: Heroy (H@eg 1974:493) Mjalkekross Milk cross Sogn og Fjordane: Hayanger (NFS Manum);M g Romsdal: Sunnmere area (Aasen 1860:10) Mjolkrot Milk root Se@r-Trondelag: Selbu (Hoeg 1974:493) Mjaltekross Milk cross Norway, unspecified (SchUbeler 1 pene Myrbukk Mire buck Sogn og Fjordane: Jalster (Haeg 19 ) Myrstjerne Mire star eee Hadsel, Tjeldsund (NFS O. . om Troms: Berg (NEG 69:14465); Finnmark: Alta (NEG 69:14465). Orm(e)gras Worm grass Telemark: Nissedal; More og Romsdal: Halsa (Haeg 1974:493) Ringormblomst Ringworm- flower Nordland: Sortland (H@eg 1974:493); Troms: Troms@ (NFS O.A. Hee Ringormgras Ringworm-grass Oppland: Lillehammer: Faberg (Haeg 1974:493); Telemark: Nissedal (NFS O.A. H@eg); Nordland: Steigen, Sortland (H@eg 1974:492-493): Troms: Skanland, Sarreisa (Hoeg 1974:493) (p)LZ VIS/DYO'LIYA Table 1. (continued) Norwegian English translation Area and source Ringormgress Sapeblomme Sinagras Skaleblom Sleipgras Slimgras Slimgress Smerkross Snigleblom Sniglegras Sprenggras Tetgras Teettegress Teettgras Tetegras Tetta Tettagras Tette Tetteblad Tetteblomst Ringworm grass Soap flower Sina grass Cup flower Slippery grass Slime- Slime-grass Butter cross Snail flower Snail grass Burst grass Thickening grass Thickening grass Thickening grass Thickening grass Thickening grass Thickening grass Rennet Thickening grass “Rennet” Thickening leaf Thickening flower North Norway, unspecified (NFS Gade-Gran 49, as ringormgres) Telemark: Vinje (Halvorsen 1988:198) Rogaland: Sandnes: Hetland an 974:493) Aust-Agder: Bykle (Haeg 1974: Nordland: Brannay: Velfjord ie 1974:493) Norway, unspecified (Tonning 1773:4, as Sliim Gras) Norway, unspecified (Viborg 1793:15,as aM Hornemann 1806:20, as oe pace! based on Tonning 17 Sogn Fjordane: Aurland, Luster: Hafslo (Haeg mee oe ie sae Vags@y: Nordre Vagsay (Haeg 1 se More og Romsdal: Vags@y: Nordre Vags@y (Haeg 19 Hedmark: Tolga (Reichborn-Kjennerud 1922:87); Sor- cane Holtalen: Haltdal, Alen, Melhus, Midtre Gauldal: Singsas, Tydal (H@eg 1974-493) Nordland: Hattfjelldal (unpublished note by H. Christiansen) Nordland: Rana (Heltzen 1834/1981:63); Troms: Troms@ (Solvang 1924:28) Nordland: Rana (Heltzen 1834/1981:63); Troms: Troms@ (Solvang 1924:28) Troms: Troms@ or Lyngen: Ullsfjord (NEG 69:1 7443); Finnmark: Ser-Varanger (NFS O.A. Haeg 90) Nordland: Vega (Engen 1975 Oppland: @stre Toten (H@eg 1974:492) Hordaland: Ulvik wigs alana Astlandet (H@eg 1974:47 |: Fi Sj (NEG 69:14326), Karmoy: lorvastad (NEG 69: 14655), Suldal (Sandvik 1 991:283) (Hoeg 1974:492) Hordaland: Hardanger area (NFS Manum) Hedmark: Folldal (NFS Manum), Stange (NFS Manum); Oppland: Lillehammer (NFS Manum); Akershus: Skedsmo (NFS Manum); Hordaland: Hardanger (NFS Manum); Troms: Troms@ (EBATA 2001:7) AVMYON NI SIYVOTNA VINDINONId 40 ANVLOGONHL] ‘W1¥ $S2Z Taste 1. (continued) Norwegian English translation Area and source Tettegras Thickening grass Norway, unspecified (Aasen 1860:10;Ramus 1715:270); Hedmark: Eidskog (Fjellstad 1966:171), Elverum (NEG 69:14267), Folldal (NEG 69:14554), Grue (NFS Manum), Tynset (NEG 69:14262), Asnes: Hoff (NEG 69:14372); Oppland: Begndal (Hagen 1950:281), Fron Jenshus 1986:181), Gausdal (Vestad 1984:60; NEG 69:15596), Lillehammer (NFS Manum); Sor-Aurdal (NFS Manum), S@r-Fron (NFS Manum),Vestre Slidre (NOS, note by G. Kirkevoll), Valdres area (Kirkevoll 1940:174; Lineikro 1975:53; NOS, note by O. Hegge), @yer (NEG 69:15278); Astfold: Spydeberg (NEG 69:14290); Buskerud: Hemsedal (NEG 69:14904), Hol (NOS, note by G. Sollien), Nore og Uvdal (NEG 69:14398), Numedal area (Flatin 1918:56), Ringerike (NFS Manum); Vestfold: Andebu (NOS, note by O. Bravoll & O. Berg), Telemark: Krager@ (NFS Manum), Porsgrunn: Brunlanes (NEG 69:14363), Tinn (Klonteig 2000:78); Aust-Agder: Birkenes: Herefoss (NEG 69:14380), Risor: Sonderled (NEG 69:14279): Valle (NEG 69:14307), Amli: Gjavdal (NEG 69:14289); Vest-Agder: Kvinesdal (NEG 69:14375), Vennesla (NEG 69:14370); Rogaland: Forsand (NEG 69:14391), Gjesdal (NEG 69:20503), Hjelmeland: Fister (NEG 69:14344), Ha: Nerbe, Sandnes: Hagsfjord (NOS, note ioe Hannaas), Time (NEG 69:14366), Vindafjord: Imsland (NEG 69:15141); daland: Askoy: Lavik (NOS, note by T. Hannaas), Bamlo (NEG 69:14392), ae (NEG 69:15226), Lindas (NEG 69:14477), Os (NOS, note by T. Hannaas), Ullensvang (Skre & Skre 1974:53), Voss (NEG 69:14284); Sogn og Fjordane: Askvoll (NOS, note by T. Hannaas), Jalster (NEG 69:14404), Stryn (NOS); Mare og Romsdal: Averay: Kvernes (NEG 69:14320), Fraena (NEG 69:14259), Hareid (Bjarndal 1949:121), Sunndal: @ksendal (NEG 69:14954), Vanylven (NEG 69:14885, 19514), Volda (NEG 69:14302), Alesund: Borgund (NEG 69:15171); Ser-Trandelag: Agdenes (NEG 69:14294), Hemne (NEG 69:14387, 14400), Holtdlen: Haltdalen (NEG 69:14282), Midtre Gauldal: Singsas, Soknedal and Storen (NEG 69:14508, 14300, 14281), Selbu (Evjen 1986:63), Snillfjord (NEG 69:17287); Nord-Trondelag: Levanger: Skogn (NEG 69:14579), Lierne: Nordli 9577 (p)LZ VIS/SYO'LINS TaBLe 1. (continued) Norwegian English translation Area and source Tettegres Tettegress Tettegubbe Tettemjalkgras Thickening grass Thickening grass Thickening old man Thickened-milk grass (NEG 69:14345), Steinkjer: Soarbu (NEG 69:14330), Verdal (NEG 69:14384); Nordland: unspecified (NEG 69:14356), Bindal (NEG 69:14368), Grane (NEG 69:14274), Hattfjelldal (NEG 69:14312; NOS, note by Hallfrid Christiansen), Bronnoy:Velfjord (Strompdal 1938:73), Vefsn (NEG 69:16462), Melay (NEG 69:14468), Beiarn (NEG 69:14621), Skjerstad (NEG 69:21821), Boda: ee (NEG 69:14468), Lofoten area (Blix 1971:218-219), Vestvagay (NOS, no Hallfrid Christiansen); Troms: Harstad (Alm 1983:393), Berg (Haeg fot 492) Telemark: Skien (NFS Manum) Norway, unspecified (Gunnerus 1772:20, as Teette-Greess; Hornemann 1 806:20, as Tettegrees; Schubeler 1888:184, as Tettegraes; Tonning 1773:4, as Tette-Gres; Viborg 1793:15, as Tettegrzes); Oppland: Gudbrandsdalen area pes ee Tette-Grees), Vestre Toten (NFS Manum); Buskerud: Kongsberg (G.T. Holm 1750's manuscript, printed in Haeg 1940:95, as Teetegraess): Sogn og Fjor ate Sunnfjord area (Arentz 1802:87, as Tette-Graes); Mare og Romsdal: ee area (Krogh 1813:289, as Teettegraes), Sunnmare area (Strom 1756-fol. 66a, as Tette-Greesset, 1762:111,as Tette-Grees); Sor-Trandelag: Meldal (NFS Manur), Selbu (NEG 69:21925), Nord-Trandelag: Grong (NEG 69:14263), Verdal (Gunnerus 1768:81, as Teette-Grees); Nordland: Hattfjelldal (NFS Manum), Beiarn (NEG 69:14359) Hamaray (EBATA 2005:70), Anday (EBATA 2005:52); Troms: Harstad (Alm 1983:393; NEG 69:14547), Dyray (NEG 69:22808), Torsken (EBATA 1984:6), Balsfjord (NEG 69:20627), Troms@ (EBATA 2005:44, 50), Lyngen or Tromsg: Ullsfjord (NEG 69:1 7443), Storfjord (Nilsson & Johansen 1994-49), Kafjord (NEG 69:16667), Skjervay (EBATA 2005:45), Kvaenangen (NEG 69:20974): Finnmark: Hammerfest (EBATA 2001:1, 2005:2), Kvalsund (EBATA 2005:13), Masoy (EBATA 1994:1), Lebesby (NEG 69:22155), Sor-Varanger (annotated voucher specimen by A.B.Wessel in TROM) Akershus: Skedsmo (NFS Manum); Troms: Troms@ (EBATA 2005:43) Nord-Trondelag: Meraker (Haeg 1974:493) AVMHON NI SIUWSINA VINDINONId JO ANVLOGONHLI ‘WIV LS? TABLE 1. (Continued) Norwegian English translation Area and source Tetteplante Tettgras Tettgubbe Tettmelkplante tplante Tettskjergull Ti ras Tjettgras Tjettgras Tjettgress Tussebetgras Ystegras Thickening plant Thickening grass Thickening old man Thickened-milk plant Thickening plant Thickening fragile gold Thickening grass “Rennet” Thickening grass Thickening old man Thickening leaf Thickening grass Thickening grass Thickening grass Gnome-bite grass Water violet Lapwing fat Curdle grass Vestfold: Tjame (NEG 69:14491); Tel k: Drangedal (NFS Manum); Troms: Lenvik (EBATA 2004:23) Hedmark, widespread (Haeg 1974:492), Elverum (@vrebo@ 1939:4), Asnes (NFS Manum); Oppland, widespread (H@eg 1974: ie, pall (Solberg 1937:229); More og Romsdal: Tingvoll (Sortdal 1981:22; NEG S sa ac (Rise 1947:51); Nord-Trandelag: Nerey: Foldereid (NEG 69:1 4309) Hedmark: Elverum (Hoeg 1974:492; NFS Manum); Trysil (Furuset 1980:126, name also used for Drosera spp.); Akershus: Skedsmo: Lillestram (H@eg 1974:492): Sar- Trondelag: Orkdal: Geitastrand (H@eg 1974:492); Troms: Bardu (H@eg 1974:492) Nord-Trondelag: Steinkjer (NFS Manum Hedmark: Rena (NFS Manum) Se@r-Trendelag: Meldal (Haeg 1974:493) @stlandet, unspecified (Haeg 1974:492); Rogaland: Ha: Nzerb@ (NEG 69:14367) Norway, "rare" (Haeg 1974:492 Hordaland: Odda (NEG 69:22415) Se@r-Trendelag: Selbu (H@eg 1974:492; NFS Manum) Ser-Trandelag (Weiszeth 1990:84) Nord-Trandelag (H@eg 1974:492), Steinkjer: Beitstad (NEG 69:15202) and Egge (NEG 69:14295) Se@r-Trandelag (Weiszeth 1990:84) Nord-Trondelag (Haeg 1974:492), Grong: Hauan (NEG 69:14280) a 2 lela (Mehlum 1891:397; Reichborn-Kjennerud 1922:87) je: Mosli (H ee 1974:493) Nerane ne (NEG 69:14261) Buskerud: Kradsherad niin 1976:993) 8577 (p)LZ VaIS/9¥O"LINE ALM, ETHNOBOTANY OF PINGUICULA VULGARIS IN NORWAY 2259 the plant, and used to treat what people considered to be tussebitt “gnome bites”), usually infected wounds (Mehlum 1891:397; Reichborn-Kjennerud 1922:87). Hoeg (1974:492) noted that the leaves were used for wounds, eg. in Modalen (Hordaland, W Norway) and Rana (Nordland, N Norway). A slightly more frequent medicinal use was to cure ringworms, recorded at Lillehammer (Hedmark, SE Norway), Sortland (Nordland), Sorreisa, Berg, and perhaps Troms@ (Troms), the four latter all in N Norway (Hoeg 1974:492; Reichborn-Kjennerud 1922:87, 1941:56, NFS Gade-Gren 49). The mode of use was simple: “The root [rosette] leaves were used for ringworms. They rubbed the leaves around the sick part.” (NFS O.A. Heeg). In Faberg, SE Norway and Rana, N Norway, the leaves were used to treat warts. They have also served as a cure for eczema in Troms (Hoeg 1974:492) and in Porsanger, Finnmark (EBATA 2005:84), N Norway. Folk veterinary medicine Locally, Pinguicula vulgaris has found some use in folk veterinary medicine, again mainly as an external ointment. Some used it to treat wounds (Heeg 1974:492; Kirkevoll 1940:174). According to the latter author, it was mixed with linseed oil in Valdres (Oppland, SE Norway). P. vulgaris was also used as a cure for sore teat. Sami herdsmen used the leaves of Pinguicula to treat sore teat in reindeer (Gunnerus 1772; Tonning 1773:5), and Norwegian farmers used it for cows in Lerdal and Vik (Sogn og Fjordane, W Norway), and for cows and goats in central Norway (Hoeg 1974:492; Weiseth 1990:84). The record from Lzrdal is instructive: (..) this is Melkekrossen ‘the milk cross” |. It grows on wet rocks and in damp places. At home, we used to boil it with some kind of fat. It was used to anoint the teat of cows when they were sore or cracked. This was a good, old advice which we used when | was at the summer farm at home in Sveretfjorden. Per- haps it is still used.” (NFS Manum, letter dated 28 September 1958). In Etnedal (Oppland) and Hegeland (Aust-Agder), Hoeg (1974:492) re- corded local use of P. vulgaris to calm down cows who had already mated. Ac- cording to Storaker (1928:63), it was also used to cure some kind of “bone dis- ease” in cattle. Calendar A wide range of plants have served as calendar marks in Norway, e.g. to indi- cate when the harvest could start. Pinguicula vulgaris is not an important one, but has found at least local use, a tradition first noted by Hans Strem in his 1756 diary: “When Tette-Gresset or Melcke-Kaarset has sprouted, one uses this as a sign, that the cattle are fed [will find sufficient pasture] and may, without dan- ger, be let out to feed on the grass.” (Strom 1756:fol. 66a, cited from Standal et al. 1997:143). 2260 BRIT.ORG/SIDA 21(4) Hoeg (1974:492) noted a similar tradition, ie. that the cows could survive outdoors when the rosettes of P. vulgaris appeared, at some stations in Western Norway. In a few cases, flowering is suggested as the marker, but this gives an unlikely, late date. Apotropaic and magical uses In parts of western Norway, P. vulgaris is known as mjolkekross (“milk cross”) and similar names. The plant was placed in the milk bucket the first time the cows were milked outdoors in spring (Hoeg 1974:491-492); the same tradition applied to Potentilla erecta (L.) Rauschel. In both cases, the practice served mainly as an apotropaic, based on a kind of similarity magic: putting the “fat” leaves of Pinguicula vulgaris or the yellow flowers of Potentilla erecta into the milk bucket should ensure a good yield of fat and yellow butter. A fine account is available from Hoyanger in Sogn og Fjordane, W Norway: “I know this plant well. (...) It was called mjolkekross. In spring, during the first evening the cows were milked outdoors, we had to burn bueld [“farm or cattle fire”]. That is, we collected wood and juniper (sprake) to makea fire. While it was burning, the cows should be milked, and in the milk bucket, there had to be a fine mjolkekross. This should ensure a good yield of milk during the sum- mer. I was told so by an old dairy maid when | accompanied her while she was milking the cows.” (NFS Manum, undated [1958] letter). The observation had been made some 45 years earlier, i.e. about 1913, when the female informant had visited the neighbouring farm and repeatedly par- ticipated in the “bueld” ritual. Her great-grandmother had done the same thing, but kept it secret—as is often the case with such magic rites (additional letter from the same female informant in NFS Manum). Exceptionally, P. vulgaris has also served other magical purposes. At Ringerike (Adal) in SE Norway, people believed that if the plant was laid under the pillow for the night, the girls would dream of their coming husband (NFS Manum). Children in Dalsfjord (Volda, W Norway) believed that finding much P. vulgaris meant they would recover all their sheep when the pasture season was over in the autumn (Hoeg 1974:492). Children’s games Pinguicula vulgaris is hardly an attractive plant for children, though it may at least arouse their curiosity. The only record of any use in children’s games de- rive from the island of Seiland in Finnmark, N Norway: “As children we used the rosette as soap. It was somewhat slippery and slimy and felt like handling soap.” (EBATA 2005:2). Use in dairy products According to widespread lore in Norway, the leaves of P vulgaris were used to treat milk, which although turning sour, would still retain a better taste than if ALM, ETHNOBOTANY OF PINGUICULA VULGARIS IN NORWAY 2261 left untreated, and also gain a desired “thick” quality. (Bergsaker 1982; Grude 1945:106; Hovdhaugen 1971:34; Sandvik 1991; Schtbeler 1888:185; records in NEG, NFS, and NOS). Haeg (1974:490) collected information on such use from 83 municipalities in Norway. However, descriptions of the actual process of pre- paring such milk differ widely in folk tradition, as do the ascribed qualities or characteristics of the product. Several aspects need to be clarified: how “tettemelk” was made, the product and its characteristics, why it was made, and, finally, geographical distribution and time-line. Preparation “Thickened” milk could be made in several different ways. Most frequently, it was made by adding a small amount of tette or rennet, often just a spoonful, as astarter culture. This tette could, however, derive from various sources. At least in the 19th and 20th century, rennet from the belly of slaughtered calves was the most frequently used source of rennet in Norwegian peasant societies. It contains a proteolytic enzyme and various milk bacteria, and its ability to pro- duce curdled milk is well documented (cf. Wix 1995, 1996). Contrary to this, the effect of tette made from Pinguicula vulgaris is disputed. Ramus (1715:270), Gunnerus (1772), Tonning (1773) and other 18th century au- thors found no reason to doubt that tettegress could be used to prepare thickened milk. An early account is found in the 1756 diary of Hans Strom, in a section de- tailing the plant lore of Kvamsoya in Sande, More og Romsdal, W Norway: “This herb is also placed in the milk which by this shall gain a fine taste, or perhaps become thicker.” (Strom 1756:fol. 66a, cited from Standal et al. 1997:143). In his Flora norvegica, also a rich source of plant-lore, Gunnerus (1772:20) included only a very short note on the ethnobotany of P. vulgaris: “Norv. Tatte- Grass (qvia adhibetur lacti hyperboreo parando).” A much more detailed ac- count of the way tettemelk was prepared is found in his 1774 treatise on dairy products in Norway: “Thick sour milk is much used for food in Norway, and when prepared for this purpose, one mostly uses Tatte” (...) “to make it thick, by which it acquires a better taste. For Ta@tte is used the well-known, so-called Tatte-Gras (Pingvicula vulgaris), of which the leaves are put ina dish of fresh milk, which thereafter thickens and becomes so sticky, that it may be drawn out in long threads. Subsequently, this Tawtte will pass its quality on to other milk, into which a spoonful is mixed, just as has already been noted by Mr. von Linné in his Flora lapponica n. XI. p.10. litt. 3.de lacte compacto hyper boreo, and this thick milk, which in this way passes on to further milk, into which it is mixed, the same character, has from this property acquired the name: Tatte-Melk.” (Gunnerus 1774:142-143). Tonning (1773) added some interesting details, including observations on the time of year for such practice, and the supposed economic benefits: 2262 BRIT.ORG/SIDA 21(4) “In some places in Norway, when the summer is at its warmest, and the milk will not easily curdle, but rather usually becomes sour too quickly, the peasant wives places the fat and slimy leaves of this herb in the sieve, through which the fresh milk is passed, which in two or three days gives the so-called Taet-Melk (Lac hyperboreum). This kind of milk, which thereby becomes so thick and viscous, that it may be drawn out in long strings, is very economic in numerous rural households, since it may be mixed either with fresh milk, or with water, and thereby greatly increased [in quantity].” ... “Such Tat-Melk is not known to be used in other places than in Norway and Sweden.” (Tonning Lf (3). At Gudbrandsdalen in SE Norway, Hiorthoy (1785) recorded the following procedure: “This grass is collected and placed in the milk vessels, so that the milk should more rapidly curdle and become thick. A spoonful of such milk is sub- sequently used for other milk vessels, since it has the same effect as the grass itself” lie. served as a starter culture] (Hiorthay 1785:99). Klonteig (2000) provides a detailed account of the procedure used at Tinn in Telemark, SE Norway. He also noted that P vulgaris had to be collected in spring, while it was growing. “They rubbed clean wooden cups, troughs or buckets with this leaf rosette lof P. vulgaris]. Then they poured fresh milk into the cup. It is left standing until the milk thickens and gets suitably sour. It should not be left standing for so long that the milk could be drawn out as long threads. The tette fungus [sic] prevented the milk from getting mouldy or rotten, so that it stayed fresh for a long time.” (Klonteig 2000:43; further comment on the time of year p. 78). Related procedures, ie. sieving fresh milk through Pinguicula leaves, or pouring milk into a container with such leaves, are frequently mentioned in folkloristic and ethnobotanical literature. Evjen (1986:63) claimed that at least locally in Trondelag, the plant part used was the crushed roots, not the leaves. A record from Jolster in Sogn og Fjordane, W Norway, confirms that the roots were sometimes included, and provides some additional details on the collec- tion of plant material: “They put their fingers down at the plant [base], below the root, and then extracted both the root and the plant, washed it and put it at the bottom of the container.” (NEG 69:14362) In most cases, only the leaves were used. Some specify that they had to be thoroughly cleaned, others that washing the plant should be avoided, as re- corded in Kvaenangen, Troms (N Norway): “At Valan, two old females told me that one had used tettegres (...) to make rennet. When using the plant, it should not be washed. The root was cut off and insects stuck in the slime were removed; the rennet was in the slime.” (NEG 69:20974) ALM, ETHNOBOTANY OF PINGUICULA VULGARIS IN NORWAY 2263 It should be noted that almost all ethnobotanical traditions included here apply to the Norwegian majority population, living in what was traditionally a faring society. Sorensen & Olsen (1981:28-29) mention similar use of P. vul- garis as a substitute rennet among the Finnish ethnic minority of SE Norway, also with an agriculture-based way of living. Contrary to this, Pinguicula plays a minor role in Sami ethnobotany. Those that were involved in traditional rein- deer herding would have little use for it (except to treat sore teat in reindeer), the fat reindeer milk hardly needs rennet to become thick, although Angelica archangelica L. was often added, according to some authors (e.g. Kuoljok 1971:58) as a kind of rennet. A single example of Sami use of Pinguicula vulgaris as a rennet substitute may be quoted, from Seiland at the coast of Finnmark, N Nor- way, where people based their living on fisheries and small-scale agriculture. Referring to the 1950's or later, its use was described as follows: “We collected these stars [the leaves], washed them, and poured warm, fresh milk over them. It was left standing for a day or so, and then it became thick- ened milk.” “It was like a pudding in the bowl. When you took a spoonful, the hole remained.” Only fresh milk was considered suitable: “When they ceased having cows, they tried with milk they had bought. But it did not work, (...) it did not turn into true thickened milk.” (EBATA 200L)) A frequently used, alternative source of rennet in Norwegian folk tradition was terrestrial snails, e.g. the large, black Arion ater L., and, according to the descriptions given, several other species as well. Accordingly, such snails were knownas tettegubbe (‘thickening old man”) or similar terms (numerous records in NEG 79): identical names have been recorded for Pinguicula vulgaris (Table 1). Hoeg (1974:490) commented on this tradition, but expressed some doubt if snails had really been used in milk. Such use is, however, well documented from the western, central and northerns parts of Norway (Alm 1983:393, 1985:41-42; Bjorndal 1949:121; Blix 1971; Evjen 1986; Fjellstad 1966:171, Hovdhaugen 1971:34; Weiszeth 1990; NFS Gade-Gron 150; numerous records in NEG). At least in the north, this tradition survived well into the 20th century. In Troms, | have re- peatedly been told the names of persons who had used snails for this purpose (e.g. EBATA 1978:26, 2005:45). My own mother had been served tettemelk in her youth, but did not like it, in particular because one—according to local lore— could get an unpleasant surprise when the bottom of the bucket became vis- ible. Leaves of Pinguicula were not objected to by anyone, but snails certainly were. In some cases, snails may have been more commonly used than Pinguicula, leading toa folk belief that the effect of the latter was due to snails having rested on the leaves, depositing slime (Alm 1983:393). In More og Romsdal, W Norway, snails were considered the “very best” source of tette (Bjorndal 1949:121). A few records also show that such use was known, at least locally, by the Finnish and Sami ethnic minorities, e.g. at Porsanger in Finnmark, N Norway (EBATA 1992:11). The note of Bjorndal (1949) is interesting in specifying that three different — 2264 BRIT.ORG/SIDA 21(4) kinds of rennet were used at Hareid in More og Romsdal (W Norway)—and that an apotropaic precaution was added when preparing the milk: “(...) to get thickened milk, they either used rennet from previously prepared thickened milk, or tettegras (Pinguicula vulgaris), or, the very best: an ordinary black snail. The rennet should be applied to the bottom of the bucket with the fingers, not with a spoon or other utensils, and always in the shape of a cross. If snails were used, they were first wrapped in linen towels and then placed two by two asa cross. This was done to ward off evil.” (Bjorndal 1949:121). The product and its characteristics Pinguicula-based thickened milk was usually made from fresh milk, without heating or boiling it. When tette had been added to the milk, the mixture was stored in a modestly warm place, often on a special shelf. It should not be too warm, in which case the product would turn sour, nor too cold. The final prod- uct, Le. fine tettemelk (thickened milk), should be fresh, not very sour, taste well, and be so viscous that it formed a rope from the spoon when retrieved from the container Jaeger-Leirvik 1959). Though descriptions of tettemelk vary, most (eg. Sorensen & Olsen1981:29: Sortdal 1981; NFS Manum) agree that it was more vis- cous or ropy than milk made from ordinary rennet. As to the rennet or tette itself, whether derived from calves or from Pin- guicula, people knew that it could be stored for long periods. Weisaeth (1990.78) noted that people at a farm in Trendelag had kept the same culture for more than 40 years. His experiments also showed that tette made from P. vulgaris was robust, and could be stored in frozen condition. More frequently, tette was stored in a desiccated form (Loras 1978:8), e.g. by letting it dry in a glazed cup Jaeger-Leirvik 1959:46), in a towel (Storen 1919), ina wooden container (Weiseeth 1990:76), or on a piece of wood, which was put in a bag and placed in the food stores (Ambjorgrud et al. 1965). When needed again, it was soaked in fresh milk or in a mixture of warm water and fresh milk (Weisaeth 1990:76). Why was it made? In the old Norwegian society, milk often formed a large part of the diet. It could be the major constituent of several daily meals, especially in inland areas, less so at the coast (Gren 1942:82). In addition to the culinary aspects noted above, there were two main reasons for preparing tettemelk—related to economy and storage. In the past, fresh or sweet milk was little used in Norway (Gren 1942:83:. Opedal 1940:55). Cream was usually removed to make butter, most of which was sold to allow some cash income. Tettemelk could be prepared from both fulland skimmed milk. In both cases, the milk sugars (lactose) were transformed to milk acid (Weiszeth 1990:83). Due to its thick character, tettemelk had a greater ability to make people feel well fed (Loras 1978:7). It could also be mixed with water and consumed asa drink to quench thirst. ALM, ETHNOBOTANY OF PINGUICULA VULGARIS IN NORWAY 2265 In the past subsistence economy, cattle were frequently inadequately fed during the winter, especially towards spring as fodder stores ran out. As a re- sult, cows would only produce milk for a restricted period of year, mainly in summer (Jeger-Leirvik 1959:46). Preparing curdled milk was important in terms of allowing milk products to be stored. The various “thickened” milk prod- ucts (bearing a wide variety of vernacular names in Norwegian) could easily be stored for a long time, depending on storage conditions and the time of year. Storing for weeks, months or half a year or more is frequently mentioned in the NEG records. Jeger-Leirvik (1959:47) made a much more modest claim, that tettemelk could be stored for about one week. Rennet was also used to prepare kjellermelk, ve. “cellar milk,” which could be stored for several months in summer (Gron 1942:84). Freshly sieved, warm milk was mixed and boiled with 1/3 of water, cooled to body temperature, and transferred toa barrel with rennet. This procedure was repeated daily until the barrel was full. The mixture was stirred frequently, until turning sour. Only skimmed milk was used (Storen 1919). Contrary to tettemelk, “cellar milk” was boiled before transfer to the buckets where it was stored (Weiszeth 1990:78). The product could be stored for about one year (Gron 1942:84; Olsen-Sopp 1912). In parts of Norway, large containers of milk prepared in this way were stored for winter use, e.g. in wooden buckets holding up to 300 litres at Malselv in Troms, N Norway (Seeter 1926:234). Geographical distribution and time-line Utilization of Pinguicula vulgarisas a rennet substitute—and vernacular names that suggest such use (Table 1)—are known from most of Norway, including Ostlandet/SE Norway (Fjellstad 1966:171; Flatin 1918:56; Hagen 1950:281; Halvorsen 1988:198; Kirkevoll 1940:174; NFS Manum), Vestlandet/W Norway (Aasen 1860:10; Arentz 1802:87; Bjorndal 1949:121; Lundberg 1998:253-254; Skre & Skre 1974:53; Strom 1762:111-112; NFS Manum), Trendelag (Evjen 1986:63; Weiszeth 1990:84; NFS Manum) and North Norway (Alm 1983:393; Blix 1971:218- 219; Jenssen 1982:44; Solvang 1924:28; Strompdal 1938:73; NFS Manum); numer- ous records in Haeg 1974 and NEG may be added to this list. Preparation of tettemelk with P. vulgaris seems to have survived longer in the central and northern parts of Norway than in the south. In the latter area, such use may have been uncommon already in the 19th century. In his large Norwegian dictionary, the linguist Ivar Aasen explained tettegras as “A herb, which was previously used to make ‘Tette” (Aasen 1873; cf. Gron 1942:83).Con- trary to this, some of those who contributed to the NEG records, mostly in 1959- 1960, had first-hand experience of such use. In fact, tettemelk may still be used in parts of Norway. According to Weiszeth (1990:78) it was in daily use at some farms in Trondelag, Central Norway in the late 1980's. Thickened milk based on P. vulgaris is now marketed as a commercial product by a dairy at Roros in 2266 BRIT.ORG/SIDA 21(4 Ser-Trondelag, Central Norway (Ola Arvid Feragen, pers. comm. 2005), avail- able throughout Norway. Although common, the practice of using Pinguicula vulgaris as a substi- tute rennet was not universally known. From Telemark in SE Norway, Wille (1786:122) noted that it was “used here only very rarely instead of rennet.” At Kredsherad in Buskerud, SE Norway, Merch (1976:993) noted that “Ystegras (tettegras) is not known to have been used for making thickened milk.” Asan alternative to Pinguicula vulgaris, a few sources mention similar use of Drosera leaves, e.g. in Rogaland (NOS) and Trondelag (Weiszth 1990). Hoeg (1974) does not mention such use of Drosera species, but he recorded some ver- nacular names, similar to the most frequent ones for Pinguicula vulgaris, that would suggest it. The effect on milk: folk belief or reality? In the early 20th century, Olsen-Sopp (1912) carried out an experimental study of tettemelk and how it could be made. According to him, Pinguicula-based cultures failed to produce true thickened milk. The product did have a ropy character, but an evil smell. Nilsson (1950), who carried out a microbiological study in Sweden, came to similar conclusions. Experiments at a dairy labora- tory in Rogaland (SW Norway) in the 1970s also failed to produce a satisfying product (Bergsaker 1982). Ambjorgrud et al. (1965) considered the effect of Pin- guicula to be little more than mere superstition. Lords (1978), in her thesis on dairy products, carried out new experiments with Pinguicula vulgaris as a source of rennet, but concluded that it was useless. On the other hand, it is easily proven that sieving milk through Pinguicula leaves gives it a ropy character, instantly yielding at least a small amount of a highly viscous, jelly-like product. According to my own experiments, this works even with pasteurised milk, but yields only a small amount, whereas Weiseth (1990:80) succeeded only with fresh milk. Several informants cited in Heeg (1974) noted that one should use fresh, still warm milk directly from the cow. Even in ethnobotanical literature, the effect of P vulgaris on milk is dis- puted. Hoeg (1974:490) suggested that it was nothing but an example of the doctrine of signatures, ie. that people inferred the alleged ability to make milk “thick” from the fat leaves of Pinguicula. This was refuted by Weiszeth (1990), based both on his own experiments and 20th century tradition in his home district of Trendelag, Central Norway. According to him, Pinguicula could in- deed be used to make thickened milk. In his experiment, 10-15 fresh leaves of Pinguicula sufficed to turn one litre of sweet milk into “a fine and good, thick tettemelk” (Weiszeth 1990:76). In Sweden, Larsson (1988) claimed to have suc- ceeded in using a Pinguicula-based culture, but no details are given on the pro- cedure; some data are provided by Alm and Larsson (1983). Until recently, however, documentation in terms of milk characteristics, ALM, ETHNOBOTANY OF PINGUICULA VULGARIS IN NORWAY 2267 chemistry and the bacteria involved has been weak. A recent study by Haug (1996) succeeded in producing thickened milk using Pinguicula vulgaris. An excellent culture was derived from plant material collected at Tromse. And old rennet culture from Roros in Central Norway, supposedly originally from Pin- guicula vulgaris, also yielded a satisfying product. Both these cultures contained strains of Lactococcus lactis subsp. cremoris and Leuconostoc mesenteroides subsp. dextranicum. However, plant material (Pinguicula leaves) from three other Norwegian sites failed to yield thickened milk. Haug (1996) suggested that the leaves had been collected too late in the season, since some ethnobotanical records cited by Heeg (1974) indicated that plants should preferably be collected early in the season, while they were growing. This is partly contradicted by the success of the Tromso material, which was collected in late summer (August). At present, knowledge of bacteria present on Pinguicula leaves is limited, and it is certainly possible that only some plants house species and strains suitable for rennet. This would explain some of the past confusion as to whether Pin- guicula works or not—and the wide range of results obtained, from complete failures to excellent cultures. Comparison with the use of Pinguicula vulgaris in other areas Pinguicula vulgaris is widely distributed in the northern hemisphere, includ- ing Europe and the northern parts of North America, but absent in most of Asia (Hultén & Fries 1986). Despite this, it plays a much more prominent role in folk tradition in the Old than the New World. Vernacular names reflecting its fat, slimy leaves are widely distributed in Europe, e.g. in Italy, the Nether- lands, and in the German-speaking countries (Brondegaard 1951:958; Marzell 1977:762-763; Schtbeler 1888:185), and in much of NW Europe. Very little in- formation on any use of Pinguicula species is available from North America. For P. vulgaris, Moerman (1998:403) only notes that the Owekeeno of Canada kept dried roots as a good luck charm. Such use is also known from Europe. In Scotland, the plant had some reputation as an apotropaic, protecting cows and milk from witches and other evil influences (Darwin 1996:128; Grigson 1955:312; Milliken & Bridgewater 2004:163-164; Vickery 1995:56). Just as in Norway, the leaves of P vulgaris have been used to cure wounds in other parts of Europe, e.g. in Germany and Great Britain; other Pinguicula spe- cies were used for the same purpose in Spain (Brondegaard 1961:959; Grigson 1955:312-313). As noted above, the main use of P vulgaris in folk veterinary medicine in Norway was to treat sore teat. This cure is also known from the Great Britain (Allen & Hatfield 2004; Grigson 1955) and the Alps, where Bauhinus (1650-51) noted that herdsmen used it for the same purpose. The belief that P vulgaris could cause harm to livestock, known from parts of Norway, was widespread in Sweden (Larsson 1988, map 26), and is also known from France (Brondegaard 2268 BRIT.ORG/SIDA 21(4) 1951:958) and Great Britain (Grigson 1955:312). In Scotland, it was considered to make the milk of grazing cows disagreeable and stringy, and was reputed to cause disease, e.g. liver fluke infestation, in sheep (Milliken & Bridgewater 2004:66, 122, 248). Thickened milk has been much used in Eurasia, especially in alpine areas where summer farms or transhumance prevailed (Rank 1971). Before the ad- vent of refrigerators and other modern technology, it was an important way of storing milk. Vernacular names and other traditions suggesting the use of P vulgaris as a rennet substitute are widely distributed in Europe, especially in the NW, including Norway, Sweden (Larsson 1988), and Great Britain (Grigson 1955:312). Numerous Swedish sources mention such use, including Linnaeus (1737:10), who provided a detailed description of its use among Swedish set- tlers in the country’s northern part. Vernacular names suggesting use in dairy products are also known from the Faroes (Brondegaard 1971:80; Svabo 1959:156) and in Iceland (Brondegaard 1971:80; Hjaltalin 1839: Mohr 1786; Nilsson 1988:155; Olsson 1961:118-119; Schtibeler 1888:185), both areas largely settled by people of Norwegian ethnic origin. Some vernacular names recorded in Scotland, e.g. on Orkney and the Shetland islands, may derive from Norse settlers, but other Scots and Gaelic names suggest that the tradition was known to the Scots as well. P vulgaris was used as rennet for cheese in Lanarkshire (Darwin 1996:128; Milliken S Bridgewater 2004:65; Vickery 1995:56). A few names of similar origin are known in the German, French and Finnish languages (Brondegaard 1971:80, Rank 1960:60). Marzell (1977:766) mentioned the use of Pinguicula as a rennet substitute from Karnten in Austria. In summary, numerous authors have carried out experiments with Pin- guicula vulgaris in milk, with widely different results. Experiments and folk tradition agree that the leaves (or their proteolytic enzymes) do have some e fect on milk; ie. by making it stringy. Most laboratory (e.g. Lords 1978; Nilsson 1950; Nilsson & Nilsson 1958; Olsen-Sopp 1912) experiments with Pinguicula have failed to produce thickened milk, at least of a quality suitable for food and storage, and Rank (1960) suggested the folk use of—or belief in—Pinguicula vul- garis as a substitute for rennet was an example of similarity magic, based on a comparison of the viscous, thickened milk and the slimy leaves of the plant. The recent study of Haug (1996) convincingly demonstrated that P vulgaris may be used to make thickened milk, but also that some Pinguicula-based cul- tures failed to do so. This may explain some of the past confusion as to whether Pinguicula works or not. sh ACKNOWLEDGMENTS Anne-Marie Wiersholm provided access to and help with the archives at NFS/ Norsk folkeminnesamling. Ola Arvid Feragen at Rerosmeieriet supplied infor- mation on their use of a Pinguicula culture for thickened milk. The referees ALM, ETHNOBOTANY OF PINGUICULA VULGARIS IN NORWAY 2269 provided many useful comments. Laurent Legendre, University of Western Australia, helped with some literature. REFERENCES AasEN, |. 1860. Norske Plantenavne. Budstikken 2:9-37. Aasen, |. 1873. Norsk Ordbog. Ed. 2. Mallings Boghandel, Christiania. A.ten, D.E.and G. Hatrieto. 2004. Medicinal plants in folk tradition. An ethnobotany of Brit- ain & lreland. Timber Press, Portland, Oregon. Avm, L.and |. Larsson. 1983. Fran forntid till framtid. Den nordiske tatmjdlken—en produkt med gamla anor. Nordisk mejeriindustri 1983(6):396-399, Aum,T. 1983. Trollbaer og tortengress. Folkeminne fra Klatran og Sarvikmark. Om planter og plantenavn. Halaygminne 16:373-396. Aum, T. 1985. Kongro og kokkelur. Folkeminne fra Klatran og Servikmark. Om dyr og dyrenavn. Haloygminne 17:40-60. Atm, T. 2000. Flora of North Norway: Pinguicula villosa L. (Lentibulariaceae). Polarflokken 24:191-205. Amps@rGRuD, O., |. Borke, J. JANSEN, and E. Moe. 1965. Tette. In: Norsk mat. Cappelen, Oslo. Pp. Arentz, H. 1802. Fortseettelse af: Beskrivelse over Sandfjord i det nordre Bergenhusiske Amt. Topogr. J. Norge 8(29):1-144. BauHINus, J. 1650-1651. Historia plantarum universalis. Yverdun. BercsAker, J. 1982. Tettegras og andre plantar i gamalt mjalkestell. Meieriposten 1982(18):482, 484, 490. Buorvano, H. and F.O. Linpeman. 2000. Vare arveord. Etymologisk ordbok. Novus, Oslo. BuoaNDAL, M. 1949. Segn og tru. Folkeminne fra More. Norsk Folkeminnelags Skr.64:1—-199. Bux, D. 1971. Nordnorsk mat. Haloygminne 13:215-223. Bronvecaard, V.J. 1951.Primitiv lobe. Meelkeritidende 1951:178-181. Branvecaaro, VJ. 1961.Vibefedt (Pinguicula) i folkemedicinen. Farm. Tid. 71(51):957-962. Bronpecaaro, V.J. 1971.Primitiv lobe. Svenska Landsmal och Svenskt Folkliv 94:75-90. Casper, S.J. 1966. Monographie der Gattung Pinguicula L. Bibl. Bot.127-128:1-209 + 16 pl. Curisten, K. 1961. Beitrag zur Pharmakochemie und Pharmakologie des gemeinen Fettkrautes (Pinguicula vulgaris L.). Die Pharmazie 16:92—102. Curisten, K. and T. Gorponorr. 1960. The active principle of Pinguicula vulgaris. Arzneimittelforschung 10:560-561. Darwin, T. 1996. The Scots herbal. Plant lore of Scotland. Mercat Press, Edinburgh. Duupeoat, R. 1959. Brev til lvar Aasen om norske plantenamn. Blyttia 17:61—-66. E.ven, R.(ed..) 1994. Johannes Lid & Dagny Tande Lid:Norsk flora. Ed.6. et norske samlaget, Oslo. Encen, B. 1975.Vegamélet. Eit oversyn med hovudvekt pa kvantitet og klusilar. Ei synkronisk og diakronisk gransking. Thesis, University of Oslo. Ewen, G. 1986. Tettemelk/tjukkmelk. Arb. Sparbu Historielag 1986:63-64. 2270 BRIT.ORG/SIDA 21(4) Fyetistao, L.M. 1966.1 grendom. Folkeminne fra Eidskog Ill. Norsk Folkeminnelags Skr. 98: Fiatin, T. 1918,.Gamalt fraa Numedaal Ill. (Tilleggsbok til Norsk folkekultur 4). Erik Gunleikson, Risor. Forsen, R. 1966. Die Langmilch (Pitkapiimd). Meijeritieteellinen Aikakauskirja 26(1). Furuset, O. 1980. Jordbrukstradisjonar fra Bjorsetgrenda. Arb. Glamdalen 39:1 15-130. Gister, N. 1749.Beskrifning om Tatmjdlkens tillagning. Kongl. Svensk. Vetensk.-Acad. Hand. 10:11-14. Gricson, G.1955. The Englishman's flora. Phoenix House, London. Gron, F. 1942.0m kostholdet i Norge fra omkring 1500-tallet og opp til var tid. Skr. Norske Vidensk.Akad. Oslo. Il. Hist.-Fil. Kl. 1942(4):1-264. Grube, D. 1945. Mat- og kjakenstell. Fram husmorskule, Oslo. Gunnerus, J.E. 1772. Flora norvegica. Pars posterior. Hafniaze (Copenhagen). Gunnerus, JE. 1768. Om nogle Norske Planter. Kongel. Norske Videnskabers Selsk. Skr. 4: 81-86. Gunnerus, J.E. 1774. 0economisk Afhandling om alle de Maader, hvorpaa Melken nyttes i Norge. Kongel. Norske Vidensk. Selsk. Skr. 5:26-152. Hacen, A. 1950.Mer omkring gamle plantenavn fra Valdres. 2. Tidsskr. Valdres Historielag 6:27 2-283. Hatvorsen, R. 1988.En del av kulturarven. Plantenavn i Vinje-dialekten. Blyttia 46:195-198. Hansson, A. 1990.Om tatmjolk, tatgras, surmjdlk och skyr. Svenska Landsméal och Svensk Folkliv 1989:88-103. Hauc, |. 1996. Bakteriologiske og teknologiske aspekter vedr@rende produksjon av tettemelk. Thesis, Norwegian Agricultural University. Heive, F. 1912.[The structure and biology of arctic flowering plants.] Lentibulariaceae (Pin- guicula). Medd. Gre@enland 36:441-481. HeizMann, W.1993. Does der Erie enemems in Altwestnordischen. (Reallexikon der germanischen A g gsbdnde, Band 7). Walter de Gruyter, Berlin - New York. Hewrzen, |.A.1834/1981.Ranens beskrivelse. Manuscript, printed as:Ranens Beskrivelse 1834 by Rana museums- og historielag/Lofotboka. Hestop-Harrison, Y. and J. Hestop-Harrison. 1980. Chloride ion movement and enzyme se- cretion from the digestive glands of Pinguicula. Ann. Bot. 45:729-731. Hestop-Harrison, Y. and J. Hestop-Harrison. 1981. The digestive glands of Pinguicula: struc- ture and cytochemistry. Ann. Bot.47:293-319. Hestop-Harrison, Y.and R.B.Knox. 1971.A cyto-chemical study of the leaf-gland enzymes of insectivorous plants of the genus Pinguicula. Planta 96:183-211. HiortHoy, H.F. 1785. Physisk og Ekonomisk Beskrivelse over Gulbransdalen Provstie | Aggershus Stift i Norge. Farste del. Kiabenhavn HJAuTALIN, O.J. 1830. [slenzk grasafraedi. Ke@benhavn Hae, O.A. 1940.G.T. Holms liste over plantenavn fra Numedal og Sandsveer i 1750-arene. Nytt Mag. Naturv. 80:89-107 ALM, ETHNOBOTANY OF PINGUICULA VULGARIS IN NORWAY 2271 Hgec, O.A. 1974. Planter og tradisjon. Floraen i levende tale og tradisjon i Norge 1925- 1973. Universitetsforlaget, Oslo - Bergen - Troms@. HorNemann, J.W.1806.Forse@g til en dans} onomisk Plantelaere.Et Priisskrivt.Andet Oplag, foreget med norske og holstenske Planter. Kabenhavn HovpHaucen, E. 1971.Dei gode henders gjerning. Bondekvinnenes liv og lagnad i dei siste hundre ara. Landbruksforlaget, Oslo. Huttén, E.and M. Fries. 1986. Atlas of North European vascular plants north of the Tropic of Cancer. Koeltz Scientific Books, Koenigstein JaceR-Leirvik, P. 1959. Tettmelk og settmelk i ytre Namdal. Norsk Folkemus. Arb. 13.43-50. JensHus, G. 1986. Fronsmalet. Fron historielag, Vinstra. Jenssen, @. 1982. Leirfjord bygdebok. Szerbind |. Helgelandsmalet i Leirfjord. Sprak og sprakkultur der Vefsn og Rana moter @yan. Leirfjord bygdeboknemnd. Josson, R.W., J. PLayrorb, K.M. Cameron, and V.A. Atsert. 2003. Molecular phylogenetics of Lentibulariaceae inferred from plastid rps16 intron and trnL-F DNA sequences: Impli- cations for character evolution and biogeography. Syst. Bot. 28:157-171. KirKevoLt, G. 1940.Plantor og plantenamn fra Valdres, serleg fra Vestre Slidre. Tidsskr.Valdres Historielag 4:165-176. Ktonteic, O. 2000. Fjelloygder i attersyn. Fra det gamle Tinn i Telemark. Forlaget Grenland, Porsgrunn. Krocn, J.A. 1813. Efterretninger om Provstiet Nordfjord i Bergens Stift i Norge. Topogr- Stat. Saml. Kongel. Selsk. Norges Vel 2(1):1- 291. Kuouok, K.E. 1971, Fada och nédféda. Hur manniskan anvande vildmarkens tillganger. LTs forlag, Stockholm. Kverneekk, PG. 1979. Navnekulturen i Trysil. Arb. Trysil 1:57-66. LaGeRBERG, T., J. HoumBoe. and R. NoroHacen. 1957. Vare ville planter, vol. 6(1). Johan Grundt Tanum, Oslo. Larsson, |. 1988. Tatmjdlk, tatgras, surmjdlk och skyr.En datorstodd ordgeografisk studie 6ver nordiska ord rorande aldre tiders mjolkhushallning. Stockholm Studies in Scandi- navian Philology. New Series 18:1-236. Lecenore, L. 2000. The genus Pinguicula L. (Lentibulariaceae): an overview. Acta Bot. Gall.147(1):77-98. Lip, J. 1941. lvar Aasens herbarium. Nytt Mag. Naturvitensk. 82:5 7-80. Lineikro, U. 1975. Utnyttinga av naturen i det gamle bygdesamfunnet. Tidsskr. Valdres Historielag. Arb. 1975:51—54. Linnagus, C. 1737.Flora Lapponica. Salomo Schouten, Amsterdam. Lords, B.1978. Dyrkingsbetingelsene og lagringstidens betydning for kvaliteten av tettemelk. Thesis, Norwegian Agricultural University. Lunpeers, A. 1998. Karm@ys flora. Fagbokforlaget, Bergen. Lunpestap, H. 1992: Ordsamling fra Vefsn. Smaskr.Vefsn Mus. 5:1-197. Marzett, H. 1977.Worterbuch der deutschen Pflanzennamen. Vol. 3.S. Hirzel Verlag, Stut- tgart - Franz Steiner Verlag, Wiesbaden. Mentum, A. 1891. Hallingdal og Hallingen. Lyche, Drammen. 2272 BRIT.ORG/SIDA 21(4) Mivuken, W. and S. Briocewarer. 2004. Flora Celtica. Plants and people in Scotland. Birlinn, Edinburgh. Moerman, D. 1998. Native American ethnobotany. Timber Press, Portland, Oregon. Monr, N. 1786. Forsag til en Islandsk Naturhistorie. Kigabenhavn Marcu, A. 1976. Kradsherad. Bind Ill. Bygdehistorie. Kradsherad kommune, Noresund. Nicsson, A.B. and L. JOHANSEN. 1994. Mattradisjoner i Skibotn. Menneske og miljo i nord- Troms. Arb. 1994:44-49. Nitsson, G. 1950. Nagot om bakteriefloran i den svenska tatmjdlken. Svensk. Mejeritidn. 1950:411-416. Nitsson, G.and R. Nisson. 1958. Studies concerning Swedish ropy milk. The antibiotic quali- ties of ropy milk. Arch. Mikrobiol. 31:191-197. Zxsenvat, H. 1993. [Lokale namn pa plantar]. Flog-fangar.Far etter fedrane.Arb.Vefsn, Grane og Hattfjelldal 1994:11 @ksENDAL, K. 1977. Plantenamn fra Vefsn. Arb. Helgeland 1977:94-102. Otsen-Sopp, O.J.1912.Tdtte, die urnordische Dauermilch und verwandte Milchsorten, sowie ihre Bedeutung ftir die Volksernahrung. Erste Serie. Centr. Bakteriol. Parasitenkunde Abt. 1l 33:1-54. Otsson, A. 1961. Koagulerad mjdlk i dldre tiders hushallning. Rig:1 13-122. Opepat, H. 1940. Makter og menneske.Folkeminne fra Hardanger IV. Norsk Folkeminnelags Skr. 46:1-192. Ovresa, L. 1939.Gamle plantenavn i Elverum. @stlendingen 39(7):4. Ramus, J. 1715. Norriges Beskrivelse. Kiobenhavn RANK, G. 1960.Vilken roll har tatorten spelat i mjdlkhusshallningen? Folktro kontra vetande. Folk-Liv 24—25:65-75. RANk, G. 1971. Verbreitungsverhaltnisse einiger Milchprodukte im eurasischen Raum. Ethnol. Scand. 1971:120-126 ReICHBORN-KJENNERUD, |. 1922.Vare folkemedisinske laegeurter. Kristiania. REICHBORN-KJENNERUD, |. 1941.Var gamle trolldomsmedisin. Ill. Vidensk.-Akad. Oslo. II. Hist-fil. KI. Skr. 1940(1):1-221. REICHBORN-KJENNERUD, |. 1944.Var gamle trolldomsmedisin. lV. Vidensk.-Akad. Oslo. II. Hist-fil. KI. Skr. 1943(2):1-263. Rise, O.J. 1947. Oppdalsboka. Historie og folkeminne.|. Johan Grundt Tanum, Oslo. Roroa, H.F. 1873. Sivert Grubbes dagbog. Danske Mag. 4. Rk. 2:361-406. Ross, H. 1895. Norsk ordbog. Tilleeg til “Norsk Ordbog" af Ivar Aasen. Cammermeyer, Christiania. SANDvik, S. 1991. Suldalsmdlet. Malleera, ord og vendingar. Norsk bokreidingslag, Bergen. SaTER, |. 1926. Maalselvdalen, Maalselv og O@verbygd herreder. Oslo. ScHUBELER, F.C. 1888. Viridarium norvegicum. Norges Veextrige. Vol. 2. Universitetsprogram, Christiania. Skre, B. and |. Skre (eds.) 1974. Hordaland og Bergen i manns minne. Daglegliv ved hundrearsskiftet. Det norske samlaget, Oslo. ALM, ETHNOBOTANY OF PINGUICULA VULGARIS IN NORWAY 2273 SoverG, R. 1937.|s-gud: dikt pa Dovremal. Aschehoug, Oslo. SoLvaNG, J. 1924.Gamal tru og skikk i bygda. Halaygminne 1:23-32. Sorensen, R.and J.B. Ovsen. 1981. Finnskogene. Gyldendal norsk forlag, Oslo. Sortoal, K.K. 1981. Laekjerader. Tingvoll, Nordmegre. Talatrosten 1981:22-23. STANDAL, R., K. AaLBeRG, K., and T. Aarset. 1997. Hans Stram: Annotations Boog over de Merkvaerdigheder som udi Syndmars Fogderie forefindes indrette[t] Anno 1756. Skr. Haram Kulturhistoriske Lag 41:|-XXX1, 1-311. Storaker, J.T. 1928. Naturrigerne i den norske Folketro. (Storakers samlinger IV). Norsk Folkeminnelags Skr. 18:1-292. Storen, K. 1919. Melkebruket. In: @degaard, N., ed.: Landbruksboken. Vol. 2. Aschehoug, Kristiania. Pp. 891-896. Strom, H. 1756.Annotations Boog over de Merk som udi Syndmers Fogderie forefindes indrette Anno 1756. [Handwritten manuscript (128 double folio pages printed and annotated in Standal et al. 1997] Strom, H. 1762. Physisk og Oekonomisk Beskrivelse over Fogderiet Sandmar, beliggende i Bergens Stift i Norge. Forste part. Sorge. Stromppal, K. 1938. Gamalt fra Helgeland Il. Norsk Folkeminnelags Skr.40:1-112. Svaso, J.C. 1959. Innberetninger fra en Reise i Farge 1781 og 1782. Selskabet til Udgivelse af faeraske Kildeskrifter og Studier, Kkabenhavn Tonnina, H. 1773. Norsk medicinsk og oeconomisk Flora.|. Kigbenhavn. Tore, A. 1919.Nynorsk etymologisk ordbok. Aschehoug, Kristiania Vestap, J.P. 1984. Gausdalsmalet. Gausdal. VisorG, E. 1793. Forsag til systematiske danske Navne af indenlandske Planter forfattet til Brug for Lerlingerne ved den Kongelige Veteri kol Vickery, R. 1995. Dictionary of plant-lore. Oxford University Press, Oxford—New York, Warmine, E. 1886.0m nogle arktiske veexters biologi. Bihang, Kongl. Svensk. Vetensk.-Akad. Handl. 12, Afd. III(2):1-40. Weisaty, G. 1990. Nytt og gammelt om tettegras, tette og tettemelk. Vare Nyttevekster 85:76-87, Witte, HJ. 1786. Beskrivelse over Sillejords Praestegield i @vre-Tellemarken i Norge tilligemed et geographisk Chart over samme. Gyldendals forlag, Kkabenhavn. Wise, N.1779.Physisk, oeconomisk og statistisk Beskrivelse over Spydebergs Preestegjeld og Egn i Aggerhuus-Stift udi Norge. Christiania. Wx, L. 1995. The traditional uses of Pinguicula in food. Part 1.Uses and geographical dis- tribution. Intern. Pinguicula Study Group Newsl.6:11-15. Wix, L. 1996.The traditional uses of Pinguicula in food.Part 2.The use of Pinguicula vulgaris in the preparation of tatmjalk. Intern. Pinguicula Study Group Newsl. 7:16-20. Le { j a t Kigbenhavn ARCHIVAL SOURCES EBATA: my own ethnobotanical records,with year and running number (e.g.EBATA 2004.7), mostly from interviews; original material at Dept. of Botany, Troms@ Museum. 2274 BRIT.ORG/SIDA 21(4) NEG (Norsk etnologisk gransking / Norwegian ethnological survey). Answers to ques- tionnaire No.69,0n milk and milk products, distributed 1959. NFS (Norsk folkeminnesamling / Norwegian folklore collection): (a) NFS Gade-Gren, questionaires on Norwegian folk medicine, distributed 1911; (b) NFS O.A. Haeg: origi- nal material of O.A. Haeg; (c) NFS Manum: answers to a 1958 query by S.Manum in “Magasinet for alle” about vernacular names and uses of Pinguicula vulgaris. NOS (Norsk ordbok, seddelarkivet / Norwegian dictionary, card archive). NOTES ON LOUISIANA BOTANY AND BOTANISTS, 1718-1975 Joseph A.Ewan Edited by pele a Panels Department of Ecology Biology e University Tula New Orleans, Louisiana 70118-5698, U.S.A. Bditor: s note.—The present paper was written ca. 1992 while the author, formerly at Tulane Univer- ity, was Research Associate at the Missouri Botanical Garden, St. Louis. It was intended for a volume of essays oe botanical artist Margaret Stones, but the collection was never published and the manuscript was abandoned. Before the intended submission, Ewan asked me to make corrections and suggest modifications, which I did. With those few and minor changes made, Ewan's notes 0 Louisiana's botanical history seemed to me well worthy of publication as a stand-alone nes tion. For Jos eph A. Ewan (1909-1999) see AS. Bradburn, Sida 19:219-222 (2000); LJ. Dorr, Taxon 49:107- 111 (2000); c = Walt, Plant Sci. Bull. 46:14 (2000). For Ewan bibliography, see LJ. Dorr and D. Holland, Arch. Natl. Hist. 27:307-334 (2000). I thank A.S. Bradburn, LJ. Dorr, E.G. Sundell, and J.W. Thieret for reviewing the manuscript and offering valuable suggestions. A concern for history has always been regarded as a mark of a civilized man.He who knows only his own generation remains always a chi — These two maxima, the first from historian Louis B. eee the ae from college president George Norlin, read from sides of the same coin: Pp erspective and from the wide-angled lens of history. We review the lives and botany of a select few of Louisiana’s naturalists- artists-collectors who have left records times only sketchily, in books, letters, and dried plant specimens from 1718 to 1975. This essay is only an introduction to the history of botanical exploration in the state. Well-known figures are mentioned, but barely remembered persons will be particularly noticed on the premise that search warrants may be posted. My annotated Bibliography of Loui- siana botany, listing 384 entries to the year 1950, is the base of this review.' A supplement to the bibliography by Eric Sundell brought the coverage to 1975 with an additional 433 titles, chiefly of recent taxonomic revisions. The first naturalist to visit Louisiana was Antoine Simon Le Page du Pratz, who as a traveling architect and engineer observed the flora and fauna He arrived 25 August 1718 reporting that New Orleans existed “only in name.” Though du Pratz died in 1775, ina real sense I met him through the enthusiasm of Stanley Clisby Arthur and his bookseller, Joseph Harmanson, by the publi- cation of acomposite altered English translation of du Pratz’s History of Louisi- ana based on the London edition. This New Orleans imprint of 1947, impor- tant for its annotations and a topical index, had a press run of only 600 copies. Du Pratz had gathered notes of plant and animal names and uses from his early contacts with his slave girl of the Chitimache tribe, purchased soon after his SIDA 21(4): 2275 — 2296, 2005 2276 BRIT.ORG/SIDA 21(4) arrival. Louisiana natural history began when she whacked an alligator on the snout as it approached his campfire. From crocodiles to cockroaches, which he remarked are relished by house cats, his narrative is interesting. Five chapters report on forest trees and other botanical subjects. Although he mentions about fifty trees, no specimens collected by him are known. He noticed mangroves as he entered the delta,’ stating that they were “very common all over America.” If you believe you know our flora, try identifying every tree in du Pratz’s forest. Benjamin Smith Barton, Philadelphia naturalist-physician, loaned his copy of du Pratz’s “Louisiana” to Meriwether Lewis in 1803. Lewis returned the book four years later after its trip to Oregon. * Du Pratz stayed in Louisiana sixteen years, sometime on the Bayou St. John, at an Indian village above New Orleans, and at Natchez. His Histoire was written from memory after his return to France. Opinions differ: biohistorian Elliot Coues, though he edited Lewis and Clark’s travels and others, did not produce an annotated edition. Coues concluded du Pratz’s “matter is very wild and of no account.” I suggest his isa clue book, to be verified. Perhaps the best known of early French travelers on the Mississippi River was Pierre Charlevoix (1682-1761), ° Jesuit missionary-historian whose visit in 1720 was first published in 1744, but not Englished until John Shea published its six volumes between 1866 and 1872. Charlevoix cited black vomit, Ilex vomitoria, cassina,’ then known by the Indian name apalachina, which altered into Apalachicola, the present Florida river and town. Understandably it was plants then used by Man, black vomit, sweet gum, and wax myrtle, that entered Charlevoix’s text.8 Before plants were collected and kept in cabinets, they were more or less carefully observed. From this description what do you suggest was Bossu’s plant: “a tree that grows in Louisiana that bears a fruit similar to a banana that the natives call hasseminier. Savages cover their cabins with the bark and also use itto make little boxes called cassot, in which to put fruits.”° Bossu also says the bark was used to make a trumpet. What tree was it? Asimina? Jean-Bernard Bossu (1720-1792), son of a surgeon, was in wars in Italy, moved to the marines, left for Louisiana in 1750 but then was assigned to Illinois country. He lived with the Akancas or Quapaw Indians near the mouth of the Arkansas River, staying there for seven years. Bossu returned to Louisiana in 1758, then for a third time as a private citizen, 1770-1771. The early travels of Bossu were first translated into English by a German, Johann Reinhold Forster,'? who later would ship with Capt. Cook on his Sec- ond Voyage. Samuel Derris Dickinson in 1982 provided a fully annotated ren- dition of Bossu’s New Travels of 1770-1771.!! As had Charlevoix, Bossu mentions the plants related to man’s use, but with more detail: cassine, Ilex: candleberry, Myrica cerifera; ginseng, Panax quinquefolius; red laurel, Persea borbonia; white EWAN, NOTES ON LOUISIANA BOTANY AND BOTANISTS, 1718-1975 2277 laurel, Magnolia virginiana (2); sassafras, Sassafras albidum, sweet gum or copala, Liquidambar styraciflua, palmetto, Sabal minor, “American tea,” Ceanothus americanus; and yucca, Yucca sp. Scarcely noticed in the history of our plant introductions is Gaillardia, known in the South as blanket-flower. In 1786 a six-page paper was published by the Paris Academy of Sciences, the first scientific paper devoted solely toa Louisiana genus. Fougeroux de Bondaroy described Gaillardia pulchellaas cul- tivated from Louisiana seed in France by one d’Essales.!* Whether a cultivated voucher was preserved I do not know. Gaillardia was a sensation when it flow- ered in Europe. Three botanists, Lamarck, Buc’hoz, and l’Héritier raced its de- scription into print in 1788. With the happy exception of Philip Miller’s Chelsea gatherings, garden-grown specimens are dubiously labeled or lost.'’ The intense competition among nurserymen to seize the market was not peculiar to France; it will inevitably complicate the effort to fix types of such discoveries as Gail- lardia. The effort to improve the wild plant and the resulting cultivars further add to the botanical bouillabaisse. Where was that Gaillardia seed collected? | suggest on the Opelousas prairies. We know William Bartram’s oak-leaf hydrangea hereabouts. The descrip- tion of his Florida experiences with bellowing alligators at the Alachua savan- nah, dramatically sketched in action, are classic. These drawings and impor- tant albums of plant specimens, preserved in the Natural History Museum, London, were reported with annotations in 1968.'+ That Bartram suffered great pain from an injury and eye infection meant that his “plan of peregrinations” through Louisiana had to be curtailed, and he collected no specimens. Never- theless he had pressed westward to the Pearl River country, stayed four or five weeks at the seat of an Englishman named Rumsey on what Bartram called “Pearl Island,” which was wiped away by hurricane years ago. He recuperated under the roof of “friendly” Rumsey whose fruit trees had reached “the utmost degree of perfection.” His eyes “having sufficient strength to endure the open daylight” he set off from Pearl Island westward in a “handsome large boat with three negroes to navigate her.” Using today’s maps we trace Bartram’s route through the Rigolet’s, along Lake Pontchartrain’s north shore, across the mouth of the Tangipahoa River through Pass Manchac to Lake Mauripas, to Amite River and an outpost on the Mississippi River. Turning north on the river he lodged at the plantation of William Dunbar,” a Scottish gentle-man naturalist whose other plantation near Natchez is better known. He traveled in a “neat cypress boat with three oars” to Pointe Coupee on the west side of the river, then on as far as Port Hudson, admiring the White Cliffs. He finally reached White Plains, an isolated prairie, by horseback, 27 October 1775. Bartram was impressed by sassafras with straight trunks up to 40 or 50 feet. There he saw “great and beau- tiful whooping cranes.” He then retraced his route in Louisiana, his eyes still — —_— 2278 BRIT.ORG/SIDA 21(4) painful, and returned to Carolina. If you would follow his journey through the southern states, pick up the “naturalist’s edition” of Travels by Bartram’s Boswell, Francis Harper. !© Although Claude Robin was a student of Jussieu, and the author of the first description of the Louisiana flora, it is Rafinesque who translated and re- vised Robin’s work, publishing it as Florula Ludoviciana. Claude César Robin!” visited Martinique and Santo Domingo en route to Louisiana. Details of his trav- els in Louisiana are vague, but he wrote long detailed plant descriptions, noted folk names and uses which amounted to 238 pages of his Voyage (Paris, 1807). Gaillardia is described but without comment on its dramatic introduction twenty years earlier. Robin’s text deserves critical study. A three-column table comparing Robin’s, Rafinesque’s, and what we think their sometimes cryptic phrases mean, would be welcome indeed. '® Though we have yet to discover any herbarium records of Robin’s, we value his contribution. Incidentally, our black locust, Robinia,commemorates Jean Robin, a 16th century French botanist, not Claude Robin. An actor in the widescreen drama of Audubon’s life, though he is barely mentioned in Audubon’s writings, was the young artist Joseph Robert Mason.!® Mason is unnoticed in botanical history for his plant backgrounds. He was born in Cincinnati in 1807 and wasa pupil at thirteen in Audubon’s drawing class in Cincinnati in the spring of 1820. The instructor must have been impressed with the lad’s talents for he took him with him on the flatboat trip to New Orleans. “As the cumbersome ark drifted downstream,” to borrow Marshall Davidson's words, Audubon was busy shooting birds and sketching for his planned Birds of America. Meanwhile Mason was collecting samples of the plants he would incorporate with the bird drawings, often later added to Audubon’ originals. They arrived at Bayou Sara, West Feliciana Parish, in January 1821. Eight days later they were in New Orleans where Audubon added bird drawings to his portfolio, meanwhile making portraits of townspeople to meet his expenses. But when Audubon received an offer from a wealthy plantation owner's wile, Lucretia Pirrie, to teach her daughter drawing, he accepted and they set off for Oakley Plantation near St. Francisville. It was there that Joseph Mason would make most of his Louisiana drawings. For example, Mason drew loblolly pine, Pinus taeda,and Audubon signed the drawing “James Pirrie’s plantation, Loui- siana, July 10, 1821. Plant J.R. Mason.”2° To my knowledge this was the only instance when Audubon recorded Mason’s part in his work. Mason drew Mag- nolia grandiflora in fruit on 5 October 1821, but Audubon did not indicate on the drawing that it was Mason’s art. I like Mason’s jessamine, Gelsemium sempervirens. His red-flowered Iris fulva is no match for Margaret Stones’, but it is an interesting record for the species about twenty years after it was first collected by Aloysius Enslen who was then gathering for Prince Lichtenstein.”! Audubon wrote to his wife Lucy in 1822, carefully distinguishing Mason’s talent EWAN, NOTES ON LOUISIANA BOTANY AND BOTANISTS, 1718-1975 2279 from his own, that Joseph Mason “draws flowers better than any man probably in America.” Understandably Mason chafed under Audubon’s unwillingness to recognize his artistic talents in what was already Audubon’s grand plan. On 23 August 1822, Mason left Audubon’s employ at Natchez and returned to Cin- cinnati. Whether it was Mason who later worked in Philadelphia for William PC. Barton has not been confirmed. Joseph Mason deserves a study and a census of his Louisiana drawings that will, however, be nettled with confusion, partly on dates, but perennially on attribution. ane ee American Heritage edition of Audubon’s watercolors with Marshall D luction, and assistance from the late Harold W. Rickett of New York Botanical Garden, will be your best source for the investigation. 23 A familiar spring umbellifer about New Orleans is chervil, as the genus Chaerophyllum is known in Europe; ours is C. tainturieri described by Will- iam Jackson Hooker in 1835 from Louis Tainturier’s specimen. ** I found some hints about the Tainturier family in New Orleans’ St. Louis cemetery no. 1, aisle 3, that suggest they came from Santo Domingo; no one has put all the dates and doubts into a believable account. S.W. Geiser suggests that Tainturier was pro- fessor of mathematics at the College d’Orleans, located at the corner of Hospi- tal and St. Claude streets in New Orleans. Eight letters survive in the Hooker correspondence, the last, 18 April 1836, informs of the dispatch of some plants collected “at 60 miles above New Orleans and which, in great part, are different from those which were sent twelve years ago.” This Tainturier essay-in-the-wait- ing would assist the systematic botanist, and gratify the historian: botanists do not live alone. The College d’Orleans, founded in 1811, represented French in- fluence as Louisiana’s first institution of higher learning. One commentator, however, remarked that the Creole cared little for schooling, and so the college “sank in a sea of troubles” and was closed in 1826.7? The natural sciences have always flourished with the hand of what Mark Catesby called “encouragers” or the patrons. Little known are the agents who forwarded the collections, held the mail, the true confreres. Joseph Barabino, who kept a small apothecary shop at 144 Old Levee Street, was the agent for Lesueur and Say.*° The French historian of natural history E.T. Hamy left us an account of Barabino: a zealous naturalist who was especially interested in entomology, but whose equipment was quite Ee wrnied belay his new friend with advice, furnished him with pins, cork sheets, etc., etc. The drugstore was the gathering place of several devotes of natural — It was near the nee mianikee and everything that ge in the vicinity of the city was brought there. Lesueur dreamed of making it the center of a society similar to that of Philadelphia. Barrabino spoke several languages and correspondence with foreign countries could be carried on with ease.” Barabino collected naturalia for Bory de Saint-Vincent and for the Lyceum of Natural History in New York from the year 1825.28 When the paleontologist of Philadelphia’s Academy, Samuel George Morton, named the Cretaceous fossil 2280 BRIT.ORG/SIDA 21(4) Inoceramus barabini he remarked “I name this species in memory of the many favors I have received from my friend Joseph Barabino, Esq., of New Orleans.”?° Lesueur’s portrait of Barabino in the archives of New Harmony bears a note by Robert J. Usher, librarian of the old Howard Memorial Library at Lee Circle: “he died while quite young from fever contracted in the swamps in which he was searching for ferns.” The year was 1834. At Oxford University’s Botany Department I was shown the portrait of Charles Giles Bridle Daubeny (1795-1867), *° who kept a pair of organ-grinder’s monkeys at the gate to the Botanic Garden next door to where he lived, in the Gatehouse. These lively guards set off an alarm at unwanted visitors. Daubeny, the son of a rector at Stratton in Gloucestershire, a graduate of Winchester and Magdalen College, was intended for a medical career, but his classes at Edin- burgh aroused his interest in geology, especially in volcanology. After travels on the continent he began, when twenty seven, to teach chemistry at Oxford, and two years later was made Professor of Botany in 1834. He visited Canada and the United States, touring Louisiana 5-28 March 1838. His Journal ofa tour (1842) in an edition of 100 copies “for private circulation,” accounts for why so few know Daubeny’s American sojourn. New Orleans, Opelousas, “Lake Chicou” come into his narrative but it was a lingering winter that year, and so his plant notes are fewer than for other states he visited. In Daubeny’s Popular geography of plants (1855) he comments that magnolias “must be seen in America before we can form any conception of their splendour. A petted Magnolia nailed up against a south wall in an English garden, gives a very poor idea of the magnifi- cent trees to be seen there, sometimes 90 feet in height... whilst the profusion of their large white blossoms, just delicately tinted, is beautifully contrasted by the background of shining dark green leaves.” *! In a letter to WJ. Hooker, Daubeny wrote “I am distracted by too many objects,” and that Hooker would find scant botanical notices in his book. ** Daubeny’s Journal deserves, indeed invites with its honest commentary, a rebirth by some publisher, though some of us may not agree when on leaving New Orleans he said “I never left a large city with less regret.” Thomas Drummond wrote his patron William Jackson Hooker at Glas- gow from New Orleans on the 5th of January 1832 “I take this opportunity of a vessel going direct for the Clyde to forward what collections I have made dur- ing the past season.” * Born in Perthshire, Scotland, Drummond fell into the orbit of William Jackson Hooker, then Professor of Botany at Glasgow. Perhaps lowly mosses had encapsulated their friendship. Though Professor Hooker re- ported on Drummond’ field work in America in the pages of the Journal of Botany soon after his letters arrived, to read the eleven closely packed letters of Drummond, now preserved at Kew, is to sense the high tide of botanical explo- ration. Drummond was collecting plants (and other naturalia) for subscribers whom Professor Hooker had contacted on his behalf. On the 20th of May 1832 EWAN, , 1718-1975 2281 Drummond wrote “I have been extremely busy” and asked Hooker to negotiate only eight or ten sf for each species. To keep up anticipations, however, Drummond often mentioned plants he had collected, for example, Acacia, Al- lium, Crinum, and Zizania. Drummond told Hooker, “you frequently find a single specimen & probably don’t find it again so that it is impossible to have all the collections [of a single number] full.”? At the close of his 1832 season in Louisiana, Drummond hoped he would be able to proceed north from Covington to Natchez through the pine woods north of Lake Pontchartrain where he found what he called “a few pretty plants,” namely two species of Rhexia and two of Sabatia. The extreme barrenness of this country, however, disappointed him. He mentioned that he had found Drosera brevifolia and Pinguicula lutea. Before Louisiana, Drummond had been with Sir John Franklin in the Ca- nadian Arctic, and then in the Canadian Rockies collecting both mosses and flowering plants. Yes, this was the “Drummond” of Drummond's phlox, named by Hooker? His phlox was collected in southeastern Texas, grown from seed, the flowers described by Hooker as “brilliant rose-red or purple varying exceed- ingly on different individuals in intensity.” Drummond, however, did not live to admire his introduction. Hooker regretted that although the phlox “bids fair to be a great ornament to the gardens of our country [it must] serve as a fre- quent memento of its unfortunate discoverer.” Drummond’ plan after explor- ing Florida had been to sample the exciting Cuban flora, but his last days in the spring of 1835 will never be known. The British consul at Havana sent Drummond's death certificate to Professor Hooker. Geiser wrote “had [Drummond] made Texas his permanent home the history of Texas botany would have been written very differently,” and that he was a man of “tremen- dous physical energy, of persistence ... forgetful of self ... it seems an unneces- sarily cruel fate that kept [Drummond] from bringing to completion his work in Texas.” * An international enterprise founded by a physician and a professor-parson of the German town of Essingen related to a botanist of Louisiana. A Natural History Traveling Society, or the Unio Itineraria of Wurttemberg, was founded by Dr. Ernst Steudel and Professor Christian Hochstetter, both of Essingen. The Unio Itineraria collected specimens from correspondents: William Darlington of West Chester, Pennsylvania, and the French naturalist Jean Louis Berlandier, who collected in Texas, were among the members. By exchange-sale, specimens were distributed to cabinets especially in Europe. Dr. Joseph C. Frank,*’ born in Rastadt, who had published a local flora of his native town in 1830, was de- puted by the Society to travel and collect in the United States. According to Stuckey, Dr Frank botanized in southwestern Ohio and the Unio Itineraria dis- tributed sets of his plants, 100 sheets for 11 florins each. These were an impor- tant source of information on 19th Century Ohio flora. The Kentucky botanist 2282 BRIT.ORG/SIDA 21(4) Dr. Charles Short says the Grand Duke of Baden commissioned Frank to collect and investigate the flora of the southern states, but his Louisiana stay was brief.*® He and his wife were stricken with yellow fever, and Frank died two months later in New Orleans in November 1835. He was fifty three. His wife returned to Germany with his collections, and they were distributed in 1836 by Unio Itinerarid. Josiah Hale, born in Virginia and a private pupil of Rafinesque, graduated at Transylvania with a medical degree in 1822.°° He then moved to Port Gibson, Mississippi, an important shipping center where he practiced medicine and collected the local flora. He took off two years for poor health to botanize in Louisiana until 1825 when, as a physician in residence, he moved to Josiah Johnson's plantation twenty miles from Alexandria and continued to collect plants. Hale’s first letter to John Torrey in 1838 began his association with Torrey and Gray.*° They proposed the genus Halea in 1842, but forty years later Gray decided it was indeed a species of Tetragonotheca. Occasionally a Hale label would catch Gray’s eye, for example, “Ulmus crassifolia Nuttall. Grows in swamps, subject to inundation, Red River - flowers late in Sept. and ripens fruit in Oct. It wi il be seen by the present splecimen] that the expression ‘ramis teretibus’ does not universally apply.” ! Hale’s plants were not numbered, and so after Charles Short and others had divided the original specimens and ex- changed a portion, the origin “Louisiana” was often all that accompanied the specimen. The keen interest in botanical exchanges of this era may be seen, for ex- ample, as when Hale wrote to Torrey, “at the request of my friend Dr. Leaven- worth, | have put up & shipped on the brig Mary Ann, Capt. Wade, a box of specimens of plants growing in the neighborhood of Alexandria.” +? On another occasion Hale wrote to George Engelmann from Canton, Mississippi, “some weeks ago | put on board the steamer Woodruff, at New Orleans, for St. Louis, a small box of specimens of plants for Dr. Mead of Augusta, IIL, directed to your care, by his instructions.” ? Hale took early retirement, married, and invested in local enterprises. When his fortune of 100,000 dollars plunged to 10,000 dollars, he moved to New Orleans to begin private practice again. During these six years in New Orleans he joined Riddell and others to found the New Orleans Acad- emy of Sciences on | April 1853, and was elected the first president. Heart trouble set in in January 1856, and Hale died 21 July. In the Academy’s Minute Book is written, Dr. Hale “has, perhaps done more to make known the peculiarities of the flora of Louisiana than all others taken together. *# Hale's particular inter- ests were grasses and sedges, two groups not enamored of by his friend Riddell. Elsewhere I have taken the historical heights reached by two American botanists for whom two 14,000-foot peaks in Colorado have been named, Gray’s and Torrey’s, for they dominated nineteenth century systematics. + In the middle decades so many novelties were being discovered in the Great West that EWAN, NOTES ON LOUISIANA BOTANY AND BOTANISTS, 1718-1975 2283 they nearly ran out of names of botanists to commemorate. William Marbury Carpenter*? did not collect in Colorado or California, but the stunning genus Carpenteria, of which there is but a single species, named by Torrey, commemo- rates an endemic saxifragaceous shrub of the southern Sierra Nevada. Carpen- ter was born in West Feliciana Parish 25 June 1811, about twelve miles from St. Francisville. The Carpenters had lived in Louisiana since 1773, and on his mother’s side, the Marburys since 1795. At eighteen Carpenter was admitted at West Point as a cadet, but with a rheumatic heart he returned home just before graduation. Soon after leaving West Point he accepted a professorship of Natu- ral Science at Centenary College, Jackson, Louisiana, then called College of Loui- siana, teaching botany, geology, “and some other things.” He botanized widely in Louisiana on vacations, occasionally with students. In 1832, for example, he collected around Opelousas. Carpenter wrote to John Torrey, “You will perhaps find some of my state- ments respecting the size of plants in the south, as inclining to extravagance. If, however, you have traveled in our delta, you will at once know that it is not exaggeration. For example, in vol. 1. p. 260 Fl[ora] Nlorth] Almerica] you state as follows of the Berchemia volubilia |*supple-jack” of the bayous, Berchemia scandens] climbing to the height of 12 or 15 ft. In Louisiana, trees exceeding 100 ftin height are sometimes completely covered by it,” and “I believe that vines of it are common here which would measure more than 200 ft, and would havea circumference near the root, of 6 to 9 inches.” It is not recorded when Carpenter collected his M.D. degree, but he prac- ticed about Jackson and from 1842 taught materia medica at the Medical School of Louisiana in New Orleans. Charles Lyell, British geologist, visited New Or- leans, and we have his story: “Dr. William Carpenter, although in full practice asa physician, kindly offered toaccompany me [to examine the geology around Balize, in the Mississippi delta] and his knowledge of botany and geology, as well as his amiable manners, made him a most useful and agreeable compan- ion.” #8 They had carried Charlevoix’s maps of the passes, published in 1743, and had found them remarkably accurate. Before his death at thirty-seven years Dr. Carpenter published on geology in Silliman’s Journal. Perhaps biographical details are buried in Benjamin Silliman’s papers at Yale. Fewer than one hundred Carpenter specimens now survive in the Tulane University herbarium. They were once part of the New Orleans Academy of Sciences collection assembled by the physician-botanists Riddell and Hale. Who was the leading botanist of Louisiana in the Nineteenth Century, who engaged the important collectors Josiah Hale and William Carpenter to coop- erate with him in what might have become the first synopsis of the state flora? John Leonard Riddell. Riddell will remain a riddle. More writers have discussed Riddell than any other figure in our story, yet no full biography of this man 2284 BRIT.ORG/SIDA 21(4) with details of his various enterprises has ever appeared, though Karlem Riess came close.*? Perhaps the very incubus of records—twenty eight manuscript diaries at Tulane—have hampered the effort. “At the time of his death, 1865, [Riddell] was considered by many to be the foremost American scientist.” After botanical instruction from Amos Eaton, whom we must admit was one of the most colorful figures in our history of science, *! Riddell was actively botanizing in Ohio, and trying to sell his bound book-like herbaria to citizens as well as to teachers in female academies. He advertised in the Marietta news- paper that he was willing to collect plants for sale: dried specimens for conver- sation pieces! In 1833 he began corresponding with John Torrey and sent him herbarium specimens. His 1l6-page Synopsis of the flora of the western states published in Cincinnati in 1835—no small synopsis—was the basis for his bo- tanical activities in Louisiana after his arrival in New Orleans in 1836 to teach chemistry officially, and natural history actually at the Medical College of Loui- siana. In many ways we are reminded of Rafinesque. Besides Riddell’s catalogue of plants growing spontaneously in Franklin County, Ohio, his six-page “Geo- logical ramble ... near Cleveland,” and his abstract on Oscillatoria structure, by 1847 he was also lecturing at the People’s Lyceum of New Orleans on “Orrin Lindsay's plan of aerial navigation with a narrative of his explorations in the higher regions of the atmosphere and his wonderful voyage around the moon.” Riddell’s lecture was printed, and it has been suggested that he anticipated H.G. Wells by some half-century. Remember that Riddell’s invention of the first prac- tical binocular microscope was noticed in the eleventh edition of the Encyclopaedia Britannica. Riddell served as Federal postmaster of New Orleans during Jefferson Davis's Confederate years. When the Confederate postal system was started | June 1861, prepayment of all postage in cash was demanded. It was then that Riddell circumvented Confederate action by issuing “provisional stamps” and “fractional currency” in denominations of one cent to five dollars—all to facili- tate the Federal cause.” Riddell was undoubtedly a spy for the Union. Small question but that Riddell’s botanical work had to be set aside. And where is the largest collection of Riddell’s specimens to be consulted? Not in the United States, but at the Natural History Museum, London. Through A.H.G. Alston’s efforts it had acquired from a provincial museum in the Midlands a set of 320 specimens, mostly of Louisiana plants (although the bound volume is titled “Ohio”!). Paris had 290 Riddell sheets, acquired with the Durand herbarium,? from which Riddell specimens were distributed. ** The Fielding Herbarium at Oxford University contains about 70 specimens, possibly acquired by Prof. Daubeny.” On 27 March 1838 Daubeny met Riddell on his American tour, and though he recorded his New Orleans visit in his Journal, strangely he does not mention Riddell. In this country Tulane University probably has the largest series, roughly EWAN, NOTES ON LOUISIANA BOTANY AND BOTANISTS, 1718-1975 2285 125 sheets, these surviving from the old New Orleans Academy of Sciences her- barium. Gray Herbarium, Torrey Herbarium, Philadelphia Academy, Darlington Herbarium at West Chester, Missouri Botanical Garden, and Smithsonian Institution, all have Riddell specimens. His conflict with Gray, the leading figure in American botany at the time and the only botanist to win a head in the Hall of Fame in New York City, made history. In 185] Riddell com- pleted a synoptical account of the plants of Louisiana which he submitted to the Smithsonian Institution for publication. Joseph Henry sent the manuscript to Asa Gray for consideration, a customary practice. Though no correspondence relative to Gray's rejection has ever been located, the evidence is found in the Gray Herbarium. Gray scissored and inserted pages from Riddell’s manuscript into the copy of Synopsis of the flora of the western states which Riddell had presented to him. There will also be found neatly folded manuscript pages in packets on herbarium sheets of the respective species described in Riddell’s manuscripts. ° Handicapped by residence in a city which Lyell characterized as gripped by cultural paralysis, plagued by two attacks of yellow fever in 1837 and 1853, weakened by the animosities of his colleagues, bound by high costs of local printing and by protracted delays in mail deliveries with his botanical corre- spondents, Riddell labored under impressive difficulties. Yet, as James Cassedy expressed it, “he combined luck, native talent, and audacity to achieve consid- erable scientific success in antebellum United States.” °8 Granted, his native tal- ents were less than those of an Engelmann or a Chapman, two practicing phy- sicians who achieved more substantial botanical successes. For all its imperfections Riddell’s Flora would have stabilized the knowledge of Louisi- ana botany in the mid-nineteenth century. “The scientific man is always on the road, never at journey’s end,” as T.D.A. Cockerell wrote, “we necessarily work with incomplete materials and more or less inadequate tools. We have to build on foundations often poorly established, and no matter how clever or industrious we may be, posterity will have to re- vise and correct much of what we have done. So true is this, that it is easy to become discouraged, and many do fall away and give up the quest.” Americus Featherman collected plants in Louisiana from 1858 until 1875. He was born in 1822 in Oettingen, a county in Germany, and came to the United States when seventeen years of age. According to John Hendley Barnhart he studied medicine at Paris, practiced medicine in Missouri and studied and prac- ticed law.°° He published three reports on agricultural botany while teaching in the Louisiana State University between 1869 and 1872. For some reason he returned to Europe in 1875 and lived in Paris. Over one hundred of his plant specimens are reported to be in the Paris herbarium. Asa Gray disposed of twelve proposed new species described from Louisiana by Featherman. One, Sabatia oligophylla, Featherman illustrated in water color.°! 2286 BRIT.ORG/SIDA 21(4) — The New Orleans World's Industrial and Cotton Centennial Exposition of 1884 brought together botanists with their exhibits. John Gill Lemmon and his wife Sara Allen Plummer Lemmon exhibited Pacific Coast conifers, pressed ferns, and wild flowers.°* George Vasey submitted grasses, and Joseph Finley Joor, who had lived in Texas for a decade, exhibited the woods of Texas. Joor, born on the Comite River, Parish of East Baton Rouge, graduated from the New Orleans School of Medicine, served as Quarantine Surgeon at Ship Island Sta- tion, and practiced at Thibodaux.® Facing poor health he moved to the prairies of Texas, first at Harrisburg, then at Birdston. While at Birdston he wrote to George Engelmann for some plant identifications: “I have now on hand speci- mens of 100 doubtful plants, including several Cruciferae, a Claytonia, a Callirrhoe.” °* He listed a dozen genera, all evidence of his botanical acuity. Joor was appointed Assistant Commissioner for Texas to prepare exhibits of woods and Texas plants for the Exposition. It was Paul Tulane’s gift of $10,000 to establish a natural history collection, shortly after his million dollar educational endowment in 1884, that brought Joor to the Tulane University of Louisiana. He began his assistant curatorship by arranging the 120 mounted birds, small-case habitat groups of chimpanzee, platypus, Kodiak bear, etc., and salvaging the herbarium including collections of Hale, Riddell, and Carpenter which had suffered from neglect in the New Orleans Academy of Sciences. He was appointed Professor of Botany in 1889 though without teaching duties. Tulane President Johnson urged Joor to go to Avery Island when a new shaft in the salt mines was exposing fossils and arti- facts. Joor reported that the Mcllhennys “most agreeably and hospitably received me.” © Fossils collected there were divided between Tulane and Mr. Edward Avery Mcllhenny; among them were two mastodon teeth, bones of equus, and of agiant sloth. For five years Professor Joor, no longer practicing medicine, cared for the Tulane Museum, continuing to collect plants mostly on his lim- ited free time, and corresponded with botanists, among them George Vasey, A.W. Chapman, and William Trelease of the Missouri Botanical Garden. The early Joor specimens from the New Orleans Academy of Sciences were not among those purchased by the Garden in 1897.°° Joor died at the age of forty-four. His daughter, Harriett, who taught art at Newcomb College and who retired to Lafay- ette, Louisiana, asked if | would accept her father’s letters on behalf of Tulane. Gladly I did. Those 103 pieces, including a postcard from Asa Gray, now in Tu- lane archives, relive Joseph Joor’s enthusiasm for plant study. Perhaps the best known Louisiana botanist of the 1890's was the clergy- man Abbé Augustus Barthélemy Langlois, born in Charanay, Loire, France, 24 April 1832. He attended the “Grand Seminary” of Lyons, and after 1855 the Semi- nary of Cincinnati, Ohio, and was ordained in 1857. He was appointed rector of Plaquemines Parish, which extended 110 miles along the Mississippi River. On his arrival at his new home he found the bloody cassock of his predecessor who —_ EWAN, , 1718-1975 2287 had been murdered when called out during the night. Pastor Langlois served thirty years at Pointe a la Hache, amassing a reference library and comprehen- sive collections—his large herbarium went to Catholic University of America, but was later dispersed to several other institutions. °’ In 1887 Langlois moved to St. Martinville in the Teche country. That year he published what he called his provisional Catalogue of the Mississippi Delta flora. °° In it he enumerated nearly 1200 seed plants, 650 fungi, 96 mosses, and 29 hepatics. Lichens were not listed, but over 200 numbers of them had been collected by the time of his death in 1900. Langlois’ fourteen letters to E.L. Greene from 1894 to 1897 tell of his “very happy voyage to Europe” in 1896 when he examined the herbaria at Geneva. “I only discovered at Boissier’s Herb. my Louisiana lichens partly determined by John Mueller of Argau before his death.” © He had also seen the “famous bryologists” Capt. Ferdinand Renauld and Jules Cardot. Langlois’ nephew, rec- tor at Breaux Bridge, Louisiana, wrote after his father’s death that Langlois’ li- brary of “at least 300 volumes” included the thirty-six volumes of Job Bicknell Ellis’s North American fungi.”° According to Saccardo’s Sylloge fungorum there are only 50 sets of Ellis in the world. Shirley Tucker has published a gazetteer of Langlois collection sites and a bibliography.” What’s in a name? How about Bush? How about a “country storekeeper and botanist” as the New York Times headlined its story.’* When Benjamin Frank- lin Bush, born in Columbus, Ohio, 21 December 1858, was seven, his mother moved him to Independence, Missouri.’? The Missouri Pacific Railroad had just been opened from St. Louis to Kansas City, and so Benjamin and his mother rode the first train. Passenger pigeons were in the woods, Carolina parakeets (“paroquets” could be caught with a coat or hat), and prairie chickens were abun- dant. The Bushes moved to Courtney, sixteen miles east of Kansas City in the early 1890s. In Courtney he sold shoes, overalls, plugs of tobacco, and groceries. Yet Bush was able to prepare herbarium specimens. Thousands of sheets are in leading herbaria—Index Herbariorum lists twenty-seven herbaria: the Gray Herbarium has 3400, and the National Herbarium over 5000. His early contacts with Asa Gray and George Engelmann launched his traveling career, and Wil- liam Trelease’s contacts were critical. In 1899 Bush began a decade of explora- tion for the Arnold Arboretum with Professor Sargent relying on his field knowl- edge to ferret out the hawthorns then under pursuit. Ernest Jesse Palmer, who wrote a portrait of Bush, met him in 1900. Both botanists had collected in Loui- siana. It is Bush’s article in Sargent’s Garden and Forest in 1897 about his search for corkwood, a shrub or small tree to thirty feet, that still challenges.’* Cork- wood, Leitneria floridana, discovered in swamps in southeast Missouri, was described by the Apalachicola botanist A.W. Chapman, friend of Asa Gray and John Torrey. Before Bush’s discovery Leitneria had been known only from its first collection in Florida, and a dubious Drummond specimen from southeast 2288 BRIT.ORG/SIDA 21(4) Texas. Trelease sent Bush to search Bayou Goula, White Lake, Louisiana, but though he “examined thoroughly” the cypress swamps he failed to find any Leitneria. He considered the “largest development” of Leitneria to be in the “big Lake, in se. Mo.and ne. Arkansas,” where shrubs were “about twelve feet in height with stems nearly four inches in diameter.” Who will find the first Leitneria in Louisiana? Reginald Wodehouse Somers Cocks took his M.A. at Trinity College, Cam- bridge, with first honors in classics.’” He must have been influenced especially by Kew-trained curator at the Cambridge Botanic Garden, Richard Irwin Lynch, indeed a linchpin. Lynch had raised the garden’s collection almost to that of Kew for its “botanicks.”’° Cocks first arrived in Canada in 1890, but soon came to Louisiana. There are specimens at Tulane dated “Feliciana, March 1892,” and by 1898 he was active in the Louisiana Society of Naturalists. By May 1906 he was corresponding with E.L. Greene, then at the Smithsonian, inquiring about Langlois’ localities, and by December he had sent a “small package of plants” for Greene’s attention. “I was very often able to get help from Dr. Mohr and Fa- ther Langlois” Cocks wrote, “but since their death there has been no botanical investigator anywhere in these parts.” / The year 1908 was a flowering for Professor Cocks, for two reasons: he met Harvard dendrologist Charles Sprague Sargent, and after one year at Louisiana State University he accepted the Ida Richardson Chair of Botany at Tulane. This chair had been created by the wife of the Dean of the Medical School, Tobias Gibson Richardson.’* Cocks was already familiar with the Tulane herbarium, citing records in his paper in the Society of Naturalists’ Proceedings as early as 1900.’ It is the Sargent-Cocks circulation that was so significant to our botany, the systole and diastole that kept specimens in motion and diagnosed. I was instructed in Louisiana botany by the over 300 letters written by Charles Sar- gent to Professor Cocks, “my companion in annual journeys of exploration through the forests of Louisiana,’®° from 1908 until 1926 when their lives and letters ceased. These letters are alive with the pursuit of hawthorns, oaks, and hickories, which Sargent conjectured he had missed in the first edition of his Manual (1905). For example, Sargent wrote Cocks, “in August 1901 Bush col- lected at Minden a sterile branch of a distinct-looking” species of Crataegus subgenus Crus-galli, adding a tease: “Is not Minden a place to explore?” ®! Sometimes the “Cock’s spur” is not always on the hawthorn. Sargent: “Brit- ton says the... Nutmeg Hickory is called Bitter Walnut in Louisiana. This must be a fake story for in the first place the kernal [sic] is not bitter, and secondly is so rare in Louisiana that it cannot have secured a popular name.” Sargent wrote: “Thank you again for all your kindness to me during my visit to Louisiana. | never had a better week or saw finer or more interesting trees, and we must make another trip together before long.”*? After that 1910 letter they had many trips together. Some of you may know the house at the corner of Carrollton and a _—d EWAN, NOTES ON LOUISIANA BOTANY AND BOTANISTS, 1718-1975 2289 Freret streets built in 1849 by Nathaniel Wilkinson in Gothic cruciform de- sign, the house snuggled behind a curtain of trees and shrubs. When Charles Low lived there Professor Sargent was house guest on his visits to New Orleans. When I came to New Orleans in 1947, Percy Viosca was a local watchword. He had supplied teaching materials for biology classes, was with the State De- partment of Conservation, Curator of Reptiles, etc., in the State Museum, New Orleans, and was a President of the Louisiana Academy of Sciences. “Louisiana's first major herpetologist probably was Percy Viosca, 1892-1961.”®* He published a benchmark paper in the journal Ecology on “Louisiana wetlands and the value of their wildlife and fisheries resources.” ® He had given a radio talk sponsored by the Smithsonian Institution, this published in Scientific Monthly.®° Viosca trumpeted Louisiana in 1933 in “a handy reference for tourists ... and nature lovers generally,” published in New Orleans.*” This is still a very “handy” guide for tracing names now hushed by progress, and the folding map of landform-vegetation by L-E. Boesch is useful. But it was Viosca’s careful study of Louisiana irises that calls our wider attention. Edgar Anderson, that zealous researcher at the Missouri Botanical Garden, gave Viosca front-page recognition in his book Introgressive hybrid- ization published in 1949.58 Introgressive hybridization was a botanical topic that in a decade translated the genetics of hundreds of American plants into printer's ink. Viosca’s study of Louisiana irises had soberly reduced the pano- ply of described “species” proposed by J.K. Small and EJ. Alexander into what Viosca interpreted as but a few freely interbreeding species. There were ninety- five taxa in Small’s Manual of 1933. Edgar Anderson had known irises in the field while collecting and researching the genus Tradescantia with Robert Woodson. Paul Percy Viosca wasa field naturalist who knew the sound of frogs, of birds in fleeting migration, and the non-migratory snakes we met in the bay- ous. | recall my visit to his bedside when he imprinted a thought: “When I have so much knowledge of the fauna and flora, why should this cancer snuff out my life?” He was seventy-two when he died a few weeks later® Versatile Clair Alan Brown, associated with Louisiana State University from 1926 for forty-two of his seventy-nine years, was first a forester and a mycolo- gist studying wood-rotting fungi? Later he wrote on Louisiana’s fossil pollen record, her paleogeologic history, revegetation after flood waters, what plants grow on Indian mounds and the extent of middens, weeds in rice fields, and mushroom poisoning among cattle. He contributed nine chatty essays to Loui- siana Conversation Review. His Louisiana trees and shrubs introduced our woody plants to hundreds of young biologists and vacationers.*! He liked ferns and found their soil requirements interesting, co-authoring with Donovan Correll a book on ferns of the state. Their attractively illustrated guide, pub- lished in an edition of only 500 copies, considers where and why ferns grow in a region we often think of as fern-poor. 2290 BRIT.ORG/SIDA 21(4) Clair was a smiling botanist, optimistic, and thoroughly social. When he married Clara Douglas, serials librarian at the University in 1963, his interest in petrified wood joined her enthusiasm for lapidary handicraft. Clair knew this state. | remember how he warned me ona field trip along Thompson Creek to watch for quicksands. His handy manual Wild flowers of Louisiana, illus- trated with his own full-color photographs, will be a record of our flora of 1972 a hundred years from now.” The life so short, the craft so es to learn, Th assay so hard, so sl q Gin Chaucer, Parlement of Foules, ca. 1382 ACKNOWLEDGMENTS My thanks to so many librarians, archivists, and biologists who have aided my efforts in tracing our history. If this review is rewarding, thank my wife, Nesta,°* my companion in zealous research. NOTES 1. Joseph Ewan,"A bibliography of Louisiana botany,” SouthW. Louisiana J. 8 (1967 [1968}) 2-83 (hereinafter cited as Ewan, Bibliog.) Eric Sundell,“Supplement 1951-1975," Tu- lane Stud. Zool. Bot. 21 (1979) 3-66. Naturalists of the Old South, planned on the model of Rocky Mountain naturalists (Denver, 1950) but never published, amounts to seven pamphlet boxes of 5x8 cards, recording quotations from correspondence, notes from archives, museum records, holographs from S.W. Geiser, etc. (hereinafter cited as Old South)."Historical sketch of Louisiana botany” by R.S.Cocks, Proc. Louisiana Soc. Naturalists (1897-1899) 69-84 (cited as Cocks, Sketch).”Some notes on the botanical history of Louisiana” by RJ.Usher, Home Gardening 1 (6)(1941) 12-13, 20-21: (7) 12-13, 19-20, and (8) 12-13. Ferns and fern allies of Louisiana (Baton Rouge, 1942) by CA. Brown and D.S. Correll (cited as Brown and Correll, Ferns). John Francis McDermott published two symposia with relevant chapters: J. Ewan,"Scientist on the frontier,” Research opportunities in American cultural history (Lexington, 1961) 81-101, and "French naturalists in the Mississippi Valley,” French in the Mississippi Valley (Urbana, 1965) 159-174 (cited as Ewan, French naturalists). Roster of 70 persons in J. Ewan, “Letters from Charles Sprague Sargent to Reginald Somers Cocks, 1908-1926," J. Arnold Arbor. 46 (1965) 1-44, 122-159, 324-361,41 1-444 (cited as Sargent-Cocks letters).J.Ewan, “Historical problems for the working taxonomist,” Taxon 18 (1969) 194-203 (cited as Ewan, Problems). . Antoine Simon Le Page du Pratz, History of Louisiana, Joseph G. Tregle, ed. (Baton Rouge, 1975). Dictionary of Louisiana biography, Glenn R. Conrad, ed. (Lafayette, 1988) (cited as DLB). Cocks, Sketch 69-80. Ewan, Bibliog. 8.Ewan, French naturalists 164. T. Becket, London, 1774 ed. (New Orleans [1947]) 223. Black mangrove, formerly Avicennia nitida Jacq., now A. germinans (L.) L.See E.L. Little Jr, U.S.D.A. Agric. Handb. 541 (1979) 59. NO aie EWAN, NOTES ON LOUISIANA BOTANY AND BOTANISTS, 1718-1975 2291 4. PR. Cutright,“Lewis and Clark and Du Pratz,” Bull. Missouri Hist. Soc. 21 (1964) 31-35. Quarter of a millenium, Edwin Wolf Il and Marie Elena Korey, eds. (Philadelphia, 1981) fig. 90. p.111.The copy now in the Library Company, Philadelphia. For B.S. Barton see J. Ewan and N. Ewan, Benjamin Smith Barton, keystone naturalist and physician in Jeffersonian America (Missouri Bot. Garden Press, St. Louis, 2005, in press). Elliot Coues, Birds of the Colorado Valley (Washington, 1878) 582. . Charles E.O’Neill, Charlevoix’s Louisiana, selections from the History and the Journal (Ba- ton Rouge, 1977). Useful documented account of Pierre FX. de Charlevoix, 1682- 1761, in B. Boivin,“La flore du Canada en 1708," Provancheria 9 (1977) 223-297, DP. Penhallow, Trans. Roy. Soc. Canada 5 sect.4 (1888) 51-52. Ewan, Bibliog. 8. . AH.G. Alston and R.E. Schultes, “Studies of early specimens and aspects of Ilex vomitoria," Rhodora 53 (1951) 273-280. . Economics of wax myrtle awaits study. Ewan, French naturalists 166.Ewan, Bibliog. 76. . Samuel Dorris Dickinson, New travels in North America by Jean-Bernard Bossu, 1770- 1791 (Natchitoches, La., 1982) 117. . Johann Reinhold Forster, transl. Travels through that part of North America formerly called Louisiana, by M. Bossu (London, 1771) 2 vols. Plant identifications by Forster, 1:347-355, to which is added Forster's “Catalogue of the known plants, shrubs, and trees in North America’as they relate to works of Catesby, Kalm,and Gronovius. Ewan, Bibliog. 9. . See note 9 for Dickinson, Bossu. For synopsis of Bossu editions see Ewan, Science 139 (1963) 478-479. Auguste Denis Fougeroux de Bondaroy, Hist. Acad. Roy. Sci. Mém. Math. Phys. (Paris) 1786 (1788) 5. Ewan, Bibliog. 10. . Hazel le Rougetel, Chelsea gardener, Philip Miller 1691-1771 (Portland, Ore., 1990) 182. 14. J.Ewan, William Bartram, botanical and zoological drawings, 1756-1788 (Philadelphia, 1968) 154-167. . William Dunbar, 1749-1810,"Scottish-born scientist and Mississippi planter,”took up his plantation near Natchez in 1792. Dunbar befriended Bartram’s friend, Alexander Wilson. See Clark Hunter, Life and letters of Alexander Wilson (Philadelphia, 1983) 101, 373,and Wilson’s letter, 358. . Francis Harper, Travels of William Bartram, naturalist’s edition (New Haven, 1958).From this Harper classic, a paper-bound edition with background essay by Robert McC. Peck (Peregrine Smith, Salt Lake City, 1980) is reliable for maintaining original pagi- nation of the 1791 edition. . For Claude César Robin, 1750-post 1807, see F. Monaghan, French travelers in the United States, 1765-1932 (New York, 1961) 81. . CS. Rafinesque, Florula Ludoviciana (New York, 1817) is an abridged and augmented translation of Robin's“ Du ™ \O © co) — — N oats, Ww Gn OV ™N (ee) s “Flore Louisianaise” in his Voyage dans Iintérieur de la Louisiane, de la Floride occidentales ... 1802 [to] 1806 (Paris, 1807) 3:325-538. See J.Ewan, editor's introduction, reprint Rafinesque, Florula Ludoviciana (New York, 1967) i-xl. . Essential reference: Stanley Clisby Arthur, Audubon, an intimate life of the American \O 2292 BRIT.ORG/SIDA 21(4) woodsman (New Orleans, 1937) 273, the basis of Irving T.Richards,“Audubon, Joseph R.Mason,and John Neal,"Amer Lit. 6 (1934) 122-140, by correspondence with Arthur. Current study: Lois Elmer Bannon and Taylor Clark, Handbook of Audubon prints (Gretna, La., 1991) 57, Mason's backgrounds, 91-108. Francis Herrick, Audubon the naturalist (New York, 1917) 2:69 refers to Thomas Sully’s report of Mason working in Philadelphia. The Joseph R. Mason, 1808-1842" in George C. Groce and David H.Wal- lace, Dictionary of artists in America, 1564-1860 (New Haven, 1957) 428, singularly makes no reference to Mason's association with Audubon! . Marshall B. Davidson, Introduction, Original water-color paintings of John James Au- dubon for the Birds of America (New York, 1966) plate 211, but Mason's drawing of Hydrangea quercifolia, plate 83, as usual makes no record of his participation. . Aloysius Enslen, Austrian gardener whom Frederick Pursh had known in Europe. He collected for Prince Lichtenstein as far south as New Orleans, where he first found the copper-colored Iris “on the banks of the Mississippi River near New Orleans,” but the records in Vienna evidently lost.Enslen's discoveries, including the iris, were cir- culated in the Philadelphia coterie. John Lyon listed the “sp. nova. copper coloured flowers,"then undescribed,on his broadside of garden novelties from America (Trans. Amer. Philos. Soc. 53 (pt. 2) (1963) 57).The latest account on Enslen is J. Ewan, editor's introduction, reprint, F.Pursh, Flora Americae Septentrionalis (Cramer, Vaduz,Germany, 1979) 15-16.See also J.Ewan,"From Calcutta and New Orleans, or, tales from Barton’s greenhouse,” Proc. Amer. Philos. Soc. 127 (1983) 125-134. The type specimen of Iris fulva Ker. was destroyed in the bombing of World War Il. See also Margaret Stones, Flora of Louisiana ... with botanical descriptions by Lowell Urbatsch (Louisiana State Univ. Press, Baton Rouge, 1991). . William Paul Crillon Barton, 1786-1856, M.D. and Prof. of Botany, University of Penn- sylvania, drew 49 native species, colored the plates himself, for his Vegetable materia medica of the United States (Philadelphia, 1817-18) but it would have been Barton's ambitious three-volume Flora of North America (Philadelphia, 1821-23) that possibly involved Joseph Mason. Barton's wife, Esther Sergeant, assisted with the drawings of both works. See J. Ewan, “History of Philadelphia horticulture: chronology, dramatis personae,” From seed to flower, Philadelphia 1681-1876 (Penn. Hort. Soc., 1976) 59,65. . See note 20. Davidson introduction, xi-xxix. Color reproductions from the Audubon collection at the New York Historical Society. . Louis Francois Tainturier, fl. 1825-1840, market-gardener (?), correspondent of W.J. Hooker, whose eight letters, 1824-1836, are preserved at Kew, lived at 47 Burgundy St.,acc.Gibson’s guide and directory, 1838, and Michel's New Orleans directory, 1840.The scattered references to Tainturier in Torrey and Gray, Flora 1:15, the collections in the Philadelphia Academy, and in T. Nuttall, Sylva 1:194, represent correspondence with Hooker and not the unlikely correspondence of Tainturier with American botanists. Ewan, Old South. W.J. Hooker, Nov. 21, 1825, to John Richardson, in part,“| am even keeping in view our projected Flora of British N. America and am extending my cor- respondence to all parts of that Continent, in order that our portion of it may in due NO <2 No NO NO NO Ww ie) f EWAN, NOTES ON LOUISIANA BOTANY AND BOTANISTS, 1718-1975 2293 NO ial NO Ke) LoS) (=) WwW (8) WW WwW W (oon time, be the better illustrated. | have an excellent correspondent even at New Or- leans, who sent me large collections from that neighbourhood & some way up the banks of the Mississippi.” Hooker-Richardson letters, Kew. . For College of Orleans see Joseph A. Breaux, Publ. Louisiana Hist. Soc. 7 (1915) 136- 142. Ewan, Old South. Ewan, French naturalists 173. Could there be a connection be- tween the “herbarium of 8000 plants well dried and well preserved” and Tainturier? Joseph Lakanal announced at the College of Orleans in 1822. See John Charles Dawson, Lakanal, the regicide (University, Ala., 1948) 133. . S.H. Scudder, Psyche 8 (1899) 306-308. Harry B. Weiss and Grace M. Ziegler, Thomas Say, early American naturalist (Springfield, Ill, 1931) cites six Barabino references. . E.T.Hamy, Travels of the naturalist Charles A. Lesueur in North America, 1815-1837, Mil- ton Haber, transl. (Kent, Ohio, 1968) 66. . Ewan, Bibliog. 38,52,56.For more on Lesueur and Bory see M.Ly-Tio-Fane, Le géographe et le naturaliste a I'lle-de-France, 1801, 1803 (Port Louis, Mauritius, 2003) . $.G.Morton, Synopsis of the organic remains of the Cretaceous group of the United States (Philadelphia, 1834) 63. Though Morton did not botanize, Asa Gray memorialized him in the celastraceous genus Mortonia, in Plantae Wrightianae 1 (1852) 34,“to that most eminent American naturalist, the late Samuel George Morton, author of Crania americana."For sidelights on the“American Golgotha,”see Frank Spencer, Trans. Stud. Coll. Physicians Philadelphia ser. 5,5 (1983) 321-338, and W.J. Bell, Jr.in C.C. Gilespie, ed., Dictionary of scientific biography 9 (1974) 540—541. eal . Daubeny,son of an Anglican cleric, is identified by Arnold Thackray as a chemist and geologist, not as a botanist (in C.C. Gilespie, ed., Dictionary of scientific biography 3 (1971) 585-586. At the Oxford meeting of the British Association for the Advance- ment of Science when the Bishop of Oxford hoped to conclude the sessions by “smashing Darwin,” Daubeny supported the Origin. See Leonard Huxley, Life and let- ters of Joseph Dalton Hooker (London, 1918) 1:521.The Catalogue of the archives of the Oxford Botany Department, now deposited in the Bodleian Library, lists 135 Daubeny titles. Ewan, Old South. . Popular geography of plants; or a botanical excursion round the world (L. Reeve, Lon- don, 1855) 113. The work was edited by Daubeny, with the author given as“E.M.C"A ms record equated this with Emily M.Cox, but the National Union Catalog ascribes the authorship to Maria E.Catlow, author of popular works on insects and shells. . C.G.B. Daubeny, Oxford, 4 March 1843, to WJ. Hooker, Hooker correspondence, Kew. . Thomas Drummond's Louisiana and Texas herbarium records were the first widely distributed among the world’s botanical centers. Some of the best narrative in the field of botanical exploration, Samuel W. Geiser, Naturalists of the frontier (Dallas, 1948) ed. 2.55-78, provided a background. . T. Drummond, New Orleans, 20 May 1832, to WJ. Hooker, Hooker correspondence, Kew . Bot.Mag., plate 3441.1835. Geiser provides a map of Drummond's collecting localities. . Ibid. 1948, 78. 2294 BRIT.ORG/SIDA 21(4) 37. The collections of Joseph C.Frank, 1782-1835,M.D.,in the Missouri Botanical Garden herbarium were acquired with the Bernhardi Herbarium.George Engelmann, St. Louis, 17 Feb. 1842, to Asa Gray, comments on Frank's botanizing in western Pennsylvania (Asa Gray correspondence, Harvard). Ronald L. Stuckey, Castanea 39 (1974) 263-272, on Frank's Ohio collections. Agnes Chase, Contrib. U.S. Natl. Herb. 28 (1929) 32, reported the type specimen of Paspalum frankii Steud., labeled “New Orleans, 1837," is in the Drake Herbarium, Paris. . See R.L. Stuckey, Scientific publications of Charles Wilkins Short (New York, 1978) i-v Ewan, Problems 200. Ewan, Bibliog. 20. 39. J.Ewan,"Josiah Hale, M.D., Louisiana botanist, Rafinesque’s pupil,” . Soc. Bibliogr. Nat. Hist. 8 (1977) 235-243. 40. Ibid. 237. Ewan, Bibliog. 25. Sargent-Cocks letters, numbers 140, 192, 246. 41. Specimen simply annotated by Asa Gray “Louisiana, Hale” Gray Herbarium, Harvard. 42. Ibid. 238. J. Hale, New Orleans, 6 June 1838, to John Torrey, Torrey correspondence, N.Y. Botanical Garden. 43. J.Hale,Canton, Mississippi, 7 Nov. 1855, to G.Engelmann.Engelmann correspondence, Missouri Botanical Garden. For Samuel Barnum Mead, 1799-1880, M.D., see the over- looked privately printed Alice L.Kibbe, Afield with plant lovers and collectors (Carthage, lll, 1953) 5-42. 44, New Orleans Academy of Sciences archives now preserved at Howard-Tilton Me- morial Library, Tulane. 45. J. Ewan. “Only ten feet less.” In James E. Guncke, ed., Current topics in plant science (New York, 1969) 155-166. . R.S.Cocks,"William Marbury Carpenter, a pioneer scientist ol Louisiana,” Tulane Gradu- ates’Mag. 3 (1914) 122-127. Also John Duffy, ed., Rudolph | history of medicine in Louisiana (Baton Rouge, 1962) passim. . W.M. Carpenter, Jackson, La, 15 June 1839, to John Torrey, Torrey correspondence, N.Y. Botanical Garden, courtesy of Susan Fraser, archivist. 48. Charles Lyell, Second visit to the United States of North America (New York, 1850) 2:111. Carpenter accompanied Lyell in March, 1846, about New Orleans, 2:106-107, and his observations around Port Hudson, 2:138-139, and the hydrography of the Missis- sippi River, 2:188-189, were reported by Lyell. 49. J.Karlem Riess,"John Leonard Riddell,” Tulane Stud. Geol. Paleontol. 13 (1977) 1-110. John Duffy, ed., Rudolph Matas history of medicine in Louisiana (Baton Rouge, 1962) 2:85-86, passim. Ralph W. Dexter,"Early career of John L. Riddell as a science lecturer in the nineteenth century,” Ohio . J. Sci. 88 (1988) 184-188. Ewan, Problems 200. DLB. 50. Otto Juettner, Daniel Drake and his followers (Cincinnati, 1909) 202—203.J.L. Riddell, to Amos Eaton, quoted by Wm. M. Smallwood, New York History 18 (1937) 183. “Prior to Agassiz, no other individual contributed nearly so much to American cul- ture through the actual study of natural history as did Amos Eaton"Wm.M.and Ma- bel S.C. Smallwood, Natural history and the American mind (New York, 1941) 283, ina Ww co pes Ov aw ™ MN EWAN, NOTES ON LOUISIANA BOTANY AND BOTANISTS, 1718-1975 2295 notable chapter, 249-284.When publication of Eaton, Manual of botany (Albany, 1817) was rejected, 63 Williams College students underwrote its printing (262). 52. Hubert C. Skinner,“The remarkable Dr. John Riddell,” Linn’s Weekly Stamp News (Sid- ney, Ohio) 40 (1967) 23. 53. Agnes Chase,“Durand Herbarium," Bartonia 17 (1935) 40-45. 54. Laurence J. Dorr,“Identity of Riddellia Raf,” Taxon 41 (1992) 80-83, see 81. 55. Hermia N. Clokie, Account of the herbaria of the Department of Botany in the University of Oxford (Oxford, 1964) 154 and 232. 56. Ewan, Problems 199-200, for figures of Riddell herbarium labels. 57. J.Ewan,"Riddell’s place in the phytography of Louisiana.” Amer. J. Bot. 50 (1963) 631, abstract. 58. J Hist. Med. Allied Sci. 28 (1973) 102. 59. J.Ewan, Rocky Mountain naturalists (Denver, 1950). 60. John H. Barnhart, Bi phical pon botanists (Boston, 1965) 3 vols.Ewan, Bibliog 29. 61. A.Gray,"Report on the botanical report of A. Featherman,”Amer. J. Sci. ser. 3. 2 (1871) 374-375. Ewan, Bibliog. 29-30. R.L. Wilbur, Rhodora 57 (1955) 101. 62. J.Ewan,“Roots of the California Botanical Society,” Madrofio 24 (1987) 1-17. Photo- graph of the Lemmons’ exhibit at the New Orleans Cotton Exposition, fig. 2, refs. 16 63. J.B.S. Norton,“Joseph F. Joor,” Bot. Gaz. 23 (1898) 271-274. portr. Copy of obituary in SouthW. Presbyterian for August 25, 1892, in Tulane University Scrap-book, vol. 2, Jan. 1, 1889—July 31, 1893, pp. 110-111, Howard-Tilton Memorial Library. . Engelmann letters, Missouri Botanical Garden archives. . Joor papers, 1868-1893, Tulane Univ., Howard-Tilton Memorial Library, Special Colls. . Verified by Anne S. Bradburn, Tulane Univ. Herbarium, October, 1978. 67 For the Langlois herbarium (said to amount to 20,000 specimens) see A.O. Tucker, M.E. Poston, and H.H. IItis, taxon, ee Gee) 196- el 68 A.B.Langlois, Catalog g cryptog dela Ba Louisiane (St. Etienne, open 1887). 69. A.B.Langlois, St. Martin's Church, St. Martinville, La.,6 Oct. 1896, to E.L. Greene, Notre Dame Univ. archives. 70. J.M.Langlois, Breaux Bridge, La.,4 Dec. 1900, to E.L. Greene, Notre Dame Univ. archives. 71. Shirley C.Tucker,"Langlois’s collection sites of Louisiana lichens,” Bryologist 73 (1970) 137-142. See also Eric Sundell, Tulane Stud. Zool. Bot. 21 (1979) 43. 72. New York Times for Feb. 7, 1934. 73. Ernest J. Palmer,"Benjamin Franklin Bush,” Amer. Midl. Naturalist 18 (1937) ii-vi. portr., bibliog. of his writings. For the field collecting of Ernest Jesse Palmer, 1875-1962, in Louisiana on behalf of the Arnold Arboretum and the Missouri Botanical Garden, see Sargent-Cocks letters, passim. 74. B.F.Bush,“Notes on the botany of some southern swamps,” Gard. & Forest 10 (1897) 514-516. Leitneria is known today from Brazoria and Chambers counties, Texas, where OV K OV OV Nn 2296 BRIT.ORG/SIDA 21(4) Drummond first discovered it. For comprehensive report on Leitneria see R.B.Chan- nell and C.E. Wood, Jr. J. Arnold Arbor. 43 (1962) 435-438. Useful early account by William Trelease, Missouri Bot. Gard. Annual Rep. 6 (1895) 65-90. frontis., plates 30-44. . Sargent-Cocks letters 2-6, portr. Ewan, Bibliog. 35-43. . S.M.Walters, Shaping of Cambridge botany (Cambridge, 1981) 75-82. . R.S.Cocks, New Orleans, 20 May 1906, to E.L.Greene, Notre Dame Univ. archives. . For Ida Ann Slocum Richardson, d. 1910, second wife of Tobias Gibson Richardson, 1827-1892,M.D.,see Mary Gehman, Women and New Orleans,a history (New Orleans, 1988) 115,127; and Charlotte Seidenberg, New Orleans garden (New Orleans, 1990) 457-458. — . R.S.Cocks,"Grasses of Louisiana,” Proc. Louisiana Soc. Naturalists [1900] 125-131. CS. Sargent, Bot. Gaz. 66 (1918) 437. . Sargent-Cocks letters 94. . Ibid. 98. . Ibid. 7. . Harold A. Dundee and Douglas A.Rossman, Amphibians and reptiles of Louisiana (Ba- ton Rouge, 1989) 4. . Ecology 9 (1928) 216-229. . Sci. Monthly 26 (1928) 19-27. . Louisiana out-of-doors (New Orleans, 1933). 190 pp., 110 illus. One dollar. . Edgar Anderson, Introgressive hybridization (New York, 1949).P.Viosca,“Irises of south- eastern Louisiana,” Bull. Amer. Iris Soc. 57 (1935) 3-55. . New Orleans States-Item for 28 Aug. 1961, portr. Memorial number, Tulane Stud. Zool. 9 (1962) no.5. . Clair Alan Brown, Ph. D. thesis:“Odontia, an epixylous fungus of family Hydnaceae.” See Bot. Gaz. 96 (1935) 640-675. . Louisiana trees and shrubs (Baton Rouge, 1945). Ewan, Bibliog. 62 . CA. Brown and Donovan S. Correll, Ferns and fern allies of Louisiana (Baton Rouge, 1942). Ewan, Bibliog. 59 . CA.Brown, Wild fh 5 of Louisiana (Baton Rouge, 1972). See Eric Sundell, Tulane Stud. Zool. Bot. 21 (1979) 46. . For Ada Nesta Dunn Ewan (1908-2000) see L.J. Dorr and Alan T.Whittemore, Taxon 49 (2000) 817-818. ADDITIONS AND EMENDATIONS TO THE WILD ORCHIDS OF NORTH AMERICA, NORTH OF MEXICO Paul Martin Brown! Research Associate University of Florida Herbarium (FLAS) lori useum of Natural History 9 Dickinson Hall, PO Box 110575 Gainesville Florida 32611-0575, U.S.A. ABSTRACT The Wild Orchids sie America, North of Mone eae Marin Brown ang aan aon Univer- me Press of Florida, 2003 ISBN 0-8130-2572-9 checkli of the orchids of an region. In the few years since Cee was ee, for that work many new taxa have been described, other pertinent facts have been published, and six corrections of fact need to be added to the original manuscript. Those additions and corrections, including literature refer- ences and photographs, are presented here to enable the reader to update this volume to the close of 2005 RESUMEN The Wild Orchids of North America, North of Mexico (Paul Martin Brown y Stan Folsom, University Press of Florida, 2003 ISBN 0-8130-2572-9) ha servido como catdlogo anotado e ilustrado de las orquideas de esa region. En los pocos anos desde que se finalizo la investigacion del trabajo se han descrito muchos taxa nuevos, se han publicado otros hechos importantes, y se han hecho seis correcciones que necesitan anadirse al manuscrito original. Estas adiciones y correcciones, que incluyen ene bibliograficas y fotografias, se presentan aqui para posibilitar al lector poner al dia en 2005 este volumen Since the compilation and publication of The Wild Orchids of North America, North of Mexico in April 2003 (research completed in May 2002), six errors have been noted, several older forms revived, num taxa described, new com- binations published, range extensions noted, ‘lost’ species rediscovered, and sig- nificant publications presented. Whether one embraces all of the subspecific taxa is not the point of this publication. These taxa have been described and are present in living material, and therefore they deserve some recognition. Recent reassessments of older generic concepts (ie, Gymnadeniopsis) and ongoing molecular work (ie. Piperia/Platanthera) have resulted in several transfers and new combinations. Whereas this is not intended in any way to be a revised or second edition of The Wild Orchids of North America, North of Mexico, those combinations and transfers (although in some cases a more accurate assess- ment of the individual taxon) are usually listed as synonyms, with literature | Address for correspondence: 10896 SW 90" Terrace, Ocala, Florida 34481, U.S.A.naorchid@aol.com SIDA 21(4): 2297 — 2319. 2005 2298 BRIT.ORG/SIDA 21(4) references, under the appropriate species. Additions and corrections are ar- ranged in original page order of The Wild Orchids of North America, North of Mexico. Page 2 Amerorchis rotundifolia (Banks ex Pursh) Hultén small round-leaf orchis emend: forma angustifolia (Rousseau) P.M. Brown—narrow-leafed form Brown, PM.and S.N. Folsom. 2006. Wild Orchids of the Canadian Maritimes and Northern Great Lakes Region, p. 284. add: forma rosea PM. Brown—pink-flowered form Brown, PM. 2004 North American Native Orchid Journal 10: 34. forma wardii PM. Brown—bicolored form Brown, PM. 2004. North American Native Orchid Journal 10: 34. 7 Calopogon barbatus (Walter) Ames bearded grass-pink add: forma lilacinus PM. Brown-—lilac-flowered form forma albiflorus PM. Brown—white-flowered form Brown, PM. 2003. North American Native Orchid Journal 9: 33. Brown, PM. and S.N. Folsom. 2004. Wild Orchids of the Southeastern United states, pp: 26-27, Brown, PM. and S.N. Folsom. 2005. Wild Orchids of Florida: updated and ex- panded edition, pp. 40-41. 8 Calopogon multiflorus Lindley many-flowered grass-pink dd: forma albiflorus PM. Brown—white-flowered form Brown, P.M. 2004. North American Native Orchid Journal 10: 21. Brown, PM. and S.N. Folsom. 2005. Wild Orchids of Florida: updated and ex- panded edition, pp. 42-43. Calopogon oklahomensis D.H. Goldman Oklahoma grass-pink add: forma albiflorus RM. Brown—white-flowered form BROWN, 2299 Brown, 2003. North America Native Orchid Journal 9: 33-34. Brown, PM. and SN. Folsom. 2004. Wild Orchids of the Southeastern United States, pp. 30731. 10 Calypso bulbosa (Linnaeus) Oakes var. americana (R. Brown) Luer eastern fairy-slipper add: forma biflora PM. Brown—two-flowered form Brown, 2004. North American Native Orchid Journal 10: 35. Brown, PM.and S.N. Folsom. 2006. Wild Orchids of the Canadian Maritimes and Northern Great Lakes Region, pp. 32-33. 13 add: Cleistes Xochlockoneensis RM. Brown Ochlockonee hybrid rosebud orchid (C. divaricata x C. bifaria) Brown, PM. 2004. North American Native Orchid Journal 10: 22. Brown, PM. and SN. Folsom. 2005. Wild Orchids of Florida: updated and ex- panded edition, p. 53. add: Corallorhiza maculata (Rafinesque) Rafinesque var. ozettensis Tisch Ozette coralroot Brown, PM. and S.N. Folsom. 2006. Wild Orchids of the Pacific Northwest and Canadian Rockies, pp. 40-41. Tisch, E. 2001. Madrono 48: 40-42. 18 Corallorhiza striata Lindley var. vreelandii (Rydberg) L.O. Williams Vreeland’s striped coralroot correct spelling to: Todsen 19 Corallorhiza trifida Chatelain early coralroot forma verna (Nuttall) PM. Brown—yellow-stemmed/white-lipped form Brown, PM.and S.N. Folsom. 2006. Wild Orchids of the Canadian Maritimes and Northern Great Lakes Region, p. 284. Corallorhiza wisteriana Conrad 2300 BRIT.ORG/SIDA 21(4) Wister’s coralroot a forma cooperi PM. Brown—cranberry-pink colored form Brown, P.M. 2004. North American Native Orchid Journal 10: 22. Brown, PM. and S.N. Folsom. 2005. Wild Orchids of Florida: updated and ex- panded edition, pp. 62-63. 20 Cranichis muscosa Swartz moss-loving cranichis emend: rediscovered in Collier County, Florida in 2004 Clusman, R. 2004. Native Orchid Conference Journal 1(2): 26-28. 21 Cypripedium acaule Aiton pink lady’s-slipper, moccasin flower aaa forma lancifolium House—narrow-leaved form House, H. 1924. New York State Museum Bulletin 254: 236. 23 Cypripedium fasciculatum Kellogg ex S. Watson clustered lady’s-slipper add: forma purpureum PM. Brown—mahogany-flowered form forma viride PM. Brown—green-flowered form Brown, PM. 2004. North American Native Orchid Journal 10: 36. Brown, PM. and S.N. Folsom. 2006. Wild Orchids of the Pacific Northwest and Canadian Rockies, pp. 56-57. 24 Cypripedium guttatum Swartz spotted lady’s-slipper add: forma albiflorum Lee—white-flowered form Lee, Y.N. 1996. Flora of Korea: 1164. Cypripedium kentuckiense C.F Reed ivory-lipped lady’s-slipper add: forma summersii PM. Brown—concolorous yellow-flowered form Brown, PM. 2002. North American Native Orchid Journal 8; 30-31. BROWN, 2301 Calypso bulbosa var. americana forma biflora p. 10 Calopagon multiflorus forma albiflorus p.8 Corallorhiza maculata var. ozettensis p. 16 Corallorhiza wisteriana forma cooperi p. 19 2302 BRIT.ORG/SIDA 21(4) af Cypripedium passerinum Richmond sparrow’s-egg lady’s-slipper, Franklin’s lady’s-slipper forma minganense (Victorin) PM. Brown—dwarf form Brown, P.M. and S.N. Folsom. 2006. Wild Orchids of the Canadian Maritimes and Northern Great Lakes Region, p. 284. 28 add: Cypripedium x herae Ewacha & Sheviak Queen Hera’s hybrid lady’s-slipper (C. parviflorum var. pubescens X C. reginae) Sheviak, C.S. 2004. Orchids 73(4): 296-299. 29 Cyrtopodium punctatum (Linnaeus) Lindley cowhorn orchid photograph is incorrect (is C. macrobulbon), see new photograph add: Cyrtopodium macrobulbon (La Llave & Lexara) G.A. Romero-Gonzalez & G. Carnevali Fernandez-Concha é~ giant cowhorn orchid southwestern Florida: Mexico photographed (1999) in Monroe County, Florida; introduced? Brown, PM. and S.N. Folsom. 2005. Wild Orchids of Florida: updated and ex- panded edition, p. 73. 30 Dactylorhiza aristata (Fischer ex Lindley) S06 var. arista Fischer’s orchid add forma albomaculata PM. Brown—white/pink spotted lip form Brown, P.M. 2004. North American Native Orchid Journal 10: 37. jae d Dactylorhiza aristata (Fischer ex Lindley) S06 var. kodiakensis Luer & Luer f Kodiak orchid a forma alba PM. Brown—white-flowered form Brown, PM. 2004. North American Native Orchid Journal 10: 37. BROWN, 2303 32 Deiregyne confusa Garay Hildago ladies’-tresses Funkiella confusa (Garay) Szlachetko, Rutkowski, & Mytnik emend: discovered at Big Bend National Park in 2004 photo in Luer (and on p. 32) taken in Durango, Mexico and does not appear to exhibit pubescent sepals of D. confusa and may be D. durangensis, see new photograph Coleman, R. 2006. Native Orchid Conference Journal 3: in press. Szlachetko, D.L., P Rutkowski, and J. Mytnik. 2005. Contributions to the taxo- nomic revision of the subtribes Spiranthinae, Stenorrhynchidinae and Cyclopogoninae (Orchidaceae) in Mesoamerica and the Antilles. Polish Botani- cal Studies 20; 229. 45 Insert the following and remove from the indicated pages in Platanthera. Gymnadeniopsis clavellata (Michaux) Rydberg var. clavellata Platanthera clavellata (Michaux) Luer var. clavellata (details see p. 78) little club-spur orchis Gymnadeniopsis clavellata (Michaux) Rydberg var. ophioglossoides (Fernald) WJ. Schrenk Hake +] : 7] 17 KALA] 5 ©) J} |e ] Hae | iS ey JA\DKA R -] | oOo f oO p. 79) northern club-spur orchis Gymnadeniopsis integra (Nuttall) Rydberg Platanthera integra (Nuttall) Lindley (details see p. 84 yellow fringeless orchis Gymnadeniopsis nived (Nuttall) Rydberg Platanthera nivea (Nuttall) Lindley (details see p. 87) snowy orchis Rydberg, PA. 1901. in Britton, Manual of the Flora of the Northeastern United Slates, pi 9> Brown, PM. 2002. North American Native Orchid Journal 8: 32-40. el Gymnadeniopsis clavellata (Michaux) Rydberg var. clavellata little club-spur orchis add: forma wrightii (Olive) PM. Brown—spurless form Brown, PM.and S.N. Folsom. 2006. Wild Orchids of the Canadian Maritimes and Northern Great Lakes Region, p. 284. Olive, L. 1951. Bulletin of the Torrey Botanical Club 78(4): 289-291. 1951. 2304 BRIT.ORG/SIDA 21(4) (yrtopodium puncatum p. 29 Cyrtopodium macrobulbon p.29 Deiregyne confusa p. 32 BROWN, 2305 49 emend: Hexalectris revoluta Correll var. revoluta recurved crested coralroot Texas; Mexico Catling, PM. 2004. Native Orchid Conference Journal 1(2): 14-16. rare and local in the mountains of southern Texas [photograph and drawing are H. revoluta var. colemanii| add: Hexalectris revoluta var. colemanii PM. Catling Coleman’s crested coralroot se Arizona Coleman, R.A. 1999. North American Native Orchid Journal 5(): 312-15. Catling, PM. 2004. Native Orchid Conference Journal 1(2): 14-16. this new variety was recently described from southeastern Arizona 50 Hexalectris spicata (Walter) Barnhardt var. spicata crested coralroot correct: forma albolabia to white-lipped form add: forma wilderi PM. Brown—albino form forma lutea PM. Brown—yellow-flowered form PM. Brown. 2004. North American Native Orchid Journal 10: 23. Brown, PM. and S.N. Folsom. 2005. Wild Orchids of Florida: updated and ex- panded edition, pp. 134-135. 51 Hexalectris warnockii Ames & Correll Texas purple-spike add: forma lutea PM. Catling—yellow-flowered form Catling, PM. 2004. Native Orchid Conference Jou rnal 12): 24. 57 insert correct name: Listera banksiana Lindley Listera caurina Piper northwestern twayblade Brown, PM. 2004. North American Native Orchid Journal 10: 2-12. Brown, PM. and SN. Folsom. 2006. Wild Orchids of the Pacific Northwest and Canadian Rockies, pp. 260. Lindley, J. 1840. Genera and Species of Orchidaceous Plants p. 455. 2306 BRIT.ORG/SIDA 21(4) the rule of priority dictates that Listera banksiana (1840) must be used over L. cdaurind (1898) 58 Listera cordata (Linnaeus) R. Brown var. cordata heart-leaved twayblade add: forma tetraphylla Lavoie—four-leaved form Lavoie, G. 1984 Provancheria 7: 92. 63 emend: Malaxis paludosa (Linnaeus) Swartz bog adder’s-mouth incorrect photograph (is Platanthera sparsiflora!); replace with new photograph 64 Malaxis spicata Swartz Florida adder’s-mouth add: forma trifoliata PM. Brown—three-leaved form Brown, P.M. 2003. North American Native Orchid Journal 9: 34. Brown, P.M. and S.N. Folsom. 2004. Wild Orchids of the Southeastern United States, pp. 126-27. Brown, PM. and S.N. Folsom. 2005. Wild Orchids of Florida: updated and ex- panded edition, pp. 154-55. 68 Pelexia adnata (Swartz) Sprengel glandular ladies’-tresses emend: discovered in Collier County, Florida in 2004 Brown, PM. and S. Folsom. 2005. Wild Orchids of Florida: updated and expanded edition, pp. 172-73. 69 Piperia candida Morgan & Ackerman (Platanthera candida (Morgan & Ackerman) R.M. Bateman: homonym) slender white piperia Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(1): 21. 70 Piperia colemanti Morgan & Glicenstein Platanthera colemanii (Morgan & Glicenstein) R.M. Bateman BROWN, 2307 Hexalectris revoluta var. revoluta p. 49 Hexalectris spicata var. spicata forma lutea p. 50 Hexalectris revoluta var. colemanii p.49 Malaxis paludosa p. 63 2308 BRIT.ORG/SIDA 21(4) Coleman’s piperia Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(1): 21 es id pcoanee TLS a watson ‘i (S. Watson) R.M. Bateman ae stoutspire orchid Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(): 21. 71 Piperia elegans Lindley Platanthera elegans Lindley Platanthera elegans Lindley subsp. maritima (Rydberg) R.M. Bateman elegant piperia Lindley, J. 1835. Genera and Species of Orchidaccous Plants, p. 285. Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(1): 21. ge rid ie ns Hasear subsp. decurtata Morgan & Glicenstein in ‘Lindley subsp. decurtata (Morgan & Glicenstein) R.M. Bateman ohat ee piperia Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(1): 21. 72 Piperia elongata Rydberg Platanthera elongata (Rydberg) R.M. Bateman long-spurred piperia Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(1): 21. Piperia leptopetala Rydber Platanthera leptopetala (Rydberg) R.M. Bateman lace orchid Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(1): 21. 73 Piperia michaelii Greene Platanthera michaelii (Greene) R.M. Bateman Michael’s piperia Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(): 21. Piperia transversa Suksdorf Platanthera transversa (Suksdorf) R.M. Bateman flat-spurred piperia Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(1): 21. 74 Piperia unalascensis (Sprengel) Rydberg Alaskan piperia BROWN, 2309 forma olympica PM. Brown—dwarf montane form Brown, PM. 2004. North American Native Orchid Journal 10: 37. Piperia yadonii Morgan & Ackerman Platanthera yadonii (Morgan & Ackerman) R.M. Bateman Yadon’s piperia Bateman, R.M. 2003. Botanical Journal of the Linnaean Society 142(): 21. 75 Platanthera blephariglottis Lindley [delete var. blephariglottis] northern white fringed orchis 76 emend and insert with text on p. 79: Platanthera conspicua (Nash) PM. Brown Platanthera blephariglottis Lindley var. conspicua (Nash) Luer southern white fringed orchis Brown, PM. 2002. North American Native Orchid Journal: 8: 3-14. 79/94 add: Platanthera convallariifolia (Fischer) Lindley lily-leaved rein orchis rare on Alaskan islands; eastern Asia Sheviak, CJ. 2002. in Flora of North America, vol. 26 pp. 559-60. Brown, PM. and SN. Folsom. 2006. Wild Orchids of the Pacific Northwest and Canadian Rockies, pp. 132-33. [note corrected spelling to convallariifolia] 80 Platanthera dilatata (Pursh) Lindley var. albiflora (Chamisso) Ledebour bog candles range extension: east to southwestern South Dakota 81 Platanthera flava (Linnaeus) Lindley var. flava southern tubercled orchis range extension: east to southern New Jersey 83 Platanthera hookeri (Torrey) Lindley Hooker’s orchis add: forma oblongifolia J.A. Paine) PM. Brown—narrow-leaved form Brown, PM. and S.N. Folsom. 2006. Wild Orchids ofthe Canadian Maritimes and Northern Great Lakes Region, p. 284. 2310 emend: Platanthera huronensis (Nuttall) Lind green bog orchis incorrect line art; replace with new —_— ey ~~ igure 88 correct spelling to: blunt-leafed rein orchis few-flowered blunt-leafed rein orchis 89 Platanthera orbiculata (Pursh) Lindley pad-leaved orchis add: forma longifolia (Clute) PM. Brown—narrow- leaved forn m forma pauciflora Jennings) PM. Brown—few BRIT.ORG/SIDA 21(4) Brown, PM.and S.N. Folsom. 2006. Wild Orchic Isoft ve Canadian Maritimes and Northern Great Lakes Region, p. 284. 90 Platanthera peramoena (A. Gray) A. Gray purple fringeless orchis add: forma doddsiae PM. Brown—white-flowered form Brown, PM. 2002. North American Native Orchid Journal 8: 30-31. Brown, P.M. and S.N. Folsom. 2004. Wild Orchids of the Southeastern United States, pp. 26-27 Platanthera praeclara Sheviak & Bowles (c} dion Fimbriella } western prairie fringed orchis viak & Bowles) Szlachetko & Rutkowsk Szlachetko, D. and P. Rutkowski 2000. Acta Botanica Fennica 169: 380. 91 Platanthera psycodes (Linnaeus) Lindley small purple fringed orchis add —_ orma fernaldii (Rousseau & Rouleau) PM. Brown—dwarf form Victorin, M. 1957. Bulletin de Jardin Botanique de l’Etat 27: 370. Brown, PM.and S.N. Folsom. 2006. Wild Orchids of the Canadian Maritimesand Northern Great Lakes Region, p. 284. BROWN, 2311 92 emend: Platanthera sparsiflora (S. Watson) Schlecter [delete var. sparsiflora] few-flowered rein orchis Platanthera stricta Lindley slender bog orchis range extension: east to southwestern South Dakota 94 add the following hybrids: Platanthera Xapalachicola PM. Brown & S. Stewart Apalachicola hybrid fringed orchis (P. chapmanii X P. cristata Brown, PM. 2003. North American Native Orchid Journal 9: 35. Brown, PM. and SN. Folsom. 2004. Wild Orchids of the Southeastern United States:p 63: Platanthera x beckneri PM. Brown Beckner’s hybrid fringed orchis (P conspicua x P.cristata) Brown, PM. 2002. North American Native Orchid Journal: 8: 3-14. Brown, PM. and S.N. Folsom. 2004. Wild Orchids of the Southeastern United States, p.107. Platanthera Xlueri PM. Brown Luer’s hybrid fringed orchis (P. conspicua X P. ciliaris) Brown, PM. 2002. North American Native Orchid Journal: 8: 3-14. Brown, PM. and S.N. Folsom. 2004. Wild Orchids of the Southeastern United States, p. 166. Platanthera Xosceola PM. Brown & S. Stewart Osceola hybrid fringed orchis (P. chapmanii X P ciliaris) Brown, PM. 2003. North American Native Orchid Journal 9: 35. Brown, P.M. and S.N. Folsom. 2004. Wild Orchids of the Southeastern United States, p. 167. Platanthera X vossii Voss’ hybrid rein orchis New nothogenus and combination: emend to: x Platanthopsis vossii (Case) PM. Brown Brown, PM. 2002. North American Native Orchid Journal 8: 32-40. BRIT.ORG/SIDA 21(4) 2312 Platanthera xlueri p.94 Piperia unalascensis forma olympica p. 74 SS SS Platanthera huronensis p.83 — =~ Cochlidium linearifolium (Desv.) Maxon ex C. Chr. (Grammitidaceae) PERU. Amazonas: Prov. Bagua: Comunidad de Yamayakat, Bosque primario, 05°03'24'S, 78°20'17"W, 600 m, Vasquez 23793, et al. (MO). Cyathea amazonica R.C. Moran (Cyatheaceae) PERU. Loreto: Prov. Maynas: Quebrada Tamshiyacu, Caseria Alianza, 4°05'S, 72°58'W, 130 m, Gentry et al. 29231 (MO); San Miguel, 2 km below Indiana, 3°30'S, 73°02'W, 130 m, Gentry 65689, et al. (MO); ca. 5 km NW from ack of Gen Gen at Rio Mormon, 3°37'S, 73°17' W, Tuomisto 3034 (AMAZ, TUR) (three additional ll ie 1 by Moran 1995, all from Prov. May nas). Cyathea boliviana R. M. Tryon (Cyatheaceae) PERU. Cuzco: Proy. Paucartambo: carretera a Pilcopata, 2650 m, Leon 2204 (USM). Cyathea brevistipes R.C. Moran (Cyatheaceae) PERU. Pasco: Prov. Oxapampa: Foster et al. 10532 (F not seen, USM) (Moran 1991). SMITH ET AL., NEW RECORDS OF PTERIDOPHYTES FROM PERU 2329 Cyathea caroli-henrici M. Lehnert (Cyatheaceae) PERU. Cuzco: Prov. Urubamba: 2050 m, Benino 1127 (CUZ) (Lehnert 2003:180). ee concordia B. Leon & R.C. Moran (Cyatheaceae) PER azonas: Coos del condoy aig 2050 m, 3°52'48'S 78°26'00"W, Beltran & Foster 1183 (F cae type and only n & Moran 1996). ohne herzogii Rosenst. ae PERU. Cuzco: Prov. La Convencion: Cordillera Vilcabamba, 12°38'S, 73°38'W, 1760 m, Dudley 10588 (NA not seen, USM). Junin: Proy. Satipo: Northern Cordillera Vilcabamba, E slope, upper Rio Puyeni watershed, 11°33'35"S, 73°38'W, 2090 m, Boyle et al. 4749 (USM). Distinct at species rank from Cyathea caracasana (Klotzsch) Domin, where it was included by Tryon (1976: 77 Cyathea palaciosii R.C. Moran (Cyatheaceae) ERU. Amazonas: Prov. San Martin: 5°41'S, 77°48'W, 1950 m, van der Werff 16721, et al. (MO not seen, UC) (Moran 1995). Cyathea thelypteroides A.R. Sm. (Cyatheaceae) PERU. Amazonas: Bagua Distr., upper slopes and summit of Cerro Tayu, 05°15'56'S, 78°22'07" W, 1030 m, van der Werff 16323, et al. (HoLotypE: USM; isotypes: MO, UC) (Smith 2005b). Cyathea tortuosa R.C. Moran (Cyatheaceae) PERU. Amazonas: Prov. Bagua: Dist. lmaza, Yamayakat, 5° oe ee S; iS 20'23"W, 400 m, Vasquez 21790, etal.(MO not seen, UC). Madre de Dios: Prov. Manu: C lla, Rio Palotoa 10-15 km NNW of ane 12°35'S, 71°18'W, 650-700 m, Foster 10703 (F UC); San Martin: along Yorongos-La Florida road near Rioja, 1000 m, van der Werff 16542, et al. (MO not seen, UC, USM). Also Depts. Pasco and Loreto (Moran 1991:100) Cyathea werffii R.C. Moran (Cyatheaceae) PERU. Amazonas: Puerto Nazareth, 540 m, Ellenberg 3489 (UC), a paratype (Moran 1991:94). Cyathea windischiana A.R. Sm. (Cyatheaceae) PERU. Amazonas: Bagua Distr, Cerro Tayu, ca. 1 hour from Chiriaco, 05°15'56"S, 78°22'07"W, 800 m, van der Werff 16207, et al. (HoLotypE: UC; isotype: MO) (Smith 2005b). Cyathea xenoxyla M. Lehnert (Cyatheaceae) PERU. Amazonas. Proy. Condorcanqui: Cordillera del Condor, Puerto de la Vigilancia Alfonso Ugarte (PV 3), cabeceras del Rio ae tributaria al oeste del Rio Cenepa, 3°55.0'S, 78°25.4'W, 1000-1300 m, Beltran & Foster 1083 (USM). Proy. Chachapoyas: road Chachapoyas-Mendoza, 52 km from yee ca. 10 km aa Molinopampa, 6°14.2'S, 77°35.9'W, 2400 m, Lehnert 229 (GOET, UC, USM). Cuzco. Prov. La Convencion: Dist. Echaraté, Llactahuaman, N del Rio Apurimac, NE del Pueblo Libre, S de la Cordillera de Vilcabamba, 12°51'55.5"S, 73°30'40"W, 1650 m, Baldeon et al. 3077 (USM). Pasco. Prov. Oxapampa: trail to summit of Cordillera Yanachaga via Rio San Daniel, 10°23'S, 75°27' W, 2500 m, Smith & ae 7846 (USM). San Martin. Prov. Rioja: road Moyobamba-Pedro Ruiz, Km 395, Lehnert 216 (GOET, UC, USM). Ucayali. Prov. Coronel Portell: Dobson (?), Ferreyra s.n. (USM). (Lehnert 2003:177). Danaea acuminata Tuomisto @ R.C. Moran (Marattiaceae) PERU. Specimens from Depts. Loreto and Madre de Dios cited by Tuomisto and Moran (2001:28), e.g., Madre de Dios: Prov. Manu: 4 km S of village of Boca Colorado, 12°38'S, 70°25'W, 300 m, Tuomisto 13589, et al. (CUZ, TUR, U, UC, USM) Danaea bipinnata Tuomisto (Marattiaceae) PERU. Specimens from Dept. Loreto cited by Tuomisto and Moran (2001:33), e.g, Loreto: Mariscal 2330 BRIT.ORG/SIDA 21(4) Ramon Castilla, 20.5-26 km NW from village of ree Rio Yaguasyacu, 3°12'S, 72°10'W, 100- 150 m, Tuomisto 11387, et al. (AMAZ, TUR, UC, L Danaea leprieurii Kunze (Marattiaceae) PERU. as from Depts. Loreto and Madre de Dios cited by Tuomisto and Moran (2001;50), e.g s, Upper Rio Itaya, ca. 5 km SSE of village Carbajal, 4°19'S, 73°35'W, 100-200 m, Tuomisto 10116, et al. (AMAZ, TUR, UC, US). Madre de Dios: Prov. Manu: 4 km S from village of Boca Colorado, 12°38'S, 70°25'W, 300 m, Tuomisto 13592, et al. (CUZ, TUR, UC, USM) Dennstaedtia arcuata Maxon (Dennstaedtiaceae) All Peruvian specie cated as D. ware ce Seobe e M. aS by Tryon and Stolze (1989b) are referred to D.arcuata, by} vas not mentioned by Tryon and Stolze. Dennstaedtia auriculata H. Navarrete & B. Ollg. (Dennstaedtiaceae) FER) Cuzco: Prov. Convencion: Amaibamba, 1700 m, Vargas 9805 (UC). This specimen cited as D. s (Willd.) Maxon, by Tryon and Stolze (1989b:100), a species restricted to the Antilles by Kiavaerdie se Ollgaard (2000). Dennstaedtia cornuta (Kaulf.) Mett. (Dennstaedtiaceae) PERU. Amazonas: |2 km E of La Peca, 1700 m, Barbour 2498 (MO not seen, UC). Pasco: Prov. Oxapampa: Paleazu, 10 km N of Oxapampa, 10°32'S, 75°23'W, 2100 m, D. Smith 8479 (MO not seen UC); along road Chatarra-Cacazu, 10°32'S, 75°04'W, 890 m, van der Werff 18292, et al. (MO not seen, UC). This name not mentioned by Tryon and Stolze (1989b), but probably some of the specimens they cited as D. dissecta are D. cornuta; the former is considered to be restricted to the Antilles, s Mexico, and nw 1MA)) i; N | Ollg 2000) South America. See Navarret Dennstaedtia coronata (Sodiro) C. Chr. (Dennstaedtiaceae) RU. San Martin: Proy. Huallaga: abajo de La Morada, cerca al Rio Guabayacu, 6°57'S, 77°32'W, 1900- 2000 m, Quipuscoa & Bardales 962 (UC). Name not mentioned by Tryon and Stolze (1989b). See Na- varrete and Ollgaard (2000). Dennstaedtia mathewsii (Hook.) C. Chr (Dennstaedtiaceae) PERU. Cajamarca: Proy. San Ignacio: La Coipa, Vista Florida (camino a la Laguna), 5° 26'10"S, 78° 56'00"W, 1900-2000 m, ll ee 4048, with Garcia (MO not seen, UC); Huarango, El. Progresso, 5° 19'15"S, 78° 40'00"W, 1300-1450 m, Campos 6276, et al. (MO not seen, UC). Pasco: Prov. Oxapampa Parque Nacional ee El Huampal, 10°11'S, 75°34'W, 1200 m, van der Werff 17908, et al. (MO not seen, UC). Name treated as synonymous with D. arborescens (Willd) Maxon by Tryon and Stolze (1989b). See Navarrete and Ollgaard (2000), who considered D. arborescens to be restricted to the Antilles. Dennstaedtia obtusifolia pees : 1, none epoonotacotaceas) Specimens cited as D. dissecta (Sw.) T. Moore by 198 Dz obtusifol ia. According to Navarrete ee Oll a (2000), D. dissecta is estricted to the Antilles, Mesoamerica, and nw S. America, and is not known from Per } me ata grandifolium (Sw.) Sw. roe U. Amazonas: Proy. Bongara: Shillac, N by trail from Pedro Ruiz, 5°49'S, 78°O1'W, 2300 m, D. Smith a (an S. 4908 (MO not seen, eek Les =i paparnes) Suu a, ca. 1900 m, Young & Eisenberg 355 (MO not seen, UC). Th specimens of this species from Peru, but all as var. andicola Stolze We believe that this variety is a cooed species and that the correct name at species rank is Diplazium balliviani Rosenst., type from Bolivia. The true D. grandifolium, type from SMITH ET AL., NEW RECORDS OF PTERIDOPHYTES FROM PERU 2331 the West Indies, does indeed occur in Peru, but apparently no specimens were seen by Tryon and Stolze. We note that the two Peruvian SESSIens of D.grand aes eee are from 1900 and 2300 m while 16 of 18 specimens seen of D. I the souther from low elevation, 100- , both from Bolivia, extend to 1100 m. Diplazium immensum Stolze (Athyriaceae) PERU. San Martin: Knapp & Alcorn 7749 (F, MO, neither seen). Cited by Tryon and Stolze (1991:77), under D. macrophyllum Desv., and as D. immensum by Stolze et al. 1994:41). wines ene oe R. M. Tryon & A. F Tryon (Athyriaceae) RU. a: Dist. Ollantaytambo, Huaytampo, 13°09'02"S, 72°30'28"W, 2300 m, Ear 5 ae aD Rie not seen, UC) Diplazium longisorum (Baker) C. Chr. (Athyriaceae) RU. San Martin: Prov. Rioj Rioja-Pedro Ruiz, 1450 m, van der Werff 15558 (MO not seen, UC). ace alae ial Stolze (Athyriaceae) PERU tin: along Rioja-Pedro Ruiz, about bridge Serranoyacu, 1170 m, van der Werff 16777, et ao (Stolze et 1994:72). sl teesdals eaion Hieron. (Athyriaceae) RU. Amazonas: Prov. Bagua: Rio Urubamba ae on Cerro Tapur above Hda. Misqui, ca. 7 km S of Bagua eae 1200 m, Hutchison 1483 (UC, USM). Loreto: [Prov. Alto Amazonas]: above ponee de Meenas mouth of Kile santiago, 200 m, ee 6129 (UC). Madre de Dios: Prov. Manu ocha Cashu B 11°53'S, 71°924'W, 400 m, Tuomisto 13177, et al. (CUZ, TUR, UC, USM); Parque Nacional dai Manu, Cocha Cashu Biological Station, Foster P-84-57, P-84-86 (UC). Several of the specimens cited by eS ane pes (1991:79) as D. striatum ae ) iS Presl are D. tabalosense, which differs from D. striatum by costules, and veins abaxially, less incised pi (blades never 2- ane even at the base), and narrower indusia ca. 0.1- 0.2(0.3) mm wide. Known from Venezuela, Ecuador, Peru, and Bolivia. Diplazium wolfii Hieron. eee PERU. Junin: Prov. Satipo: Gran Pajonal, Mapari, ca. 12 km SW of Chequitavo, 74°23'W, 10°45'S, 1300 m, D. Smith 6789 (MO not seen, UC, USM) (see oe et al. 1994:22). ee hamnpacbe (Mett. ex H. Christ) Alston (Dryopteridaceae) PERU. Cajam v. San Ignacio: Chirinos, localidad de Pacasmayo, 5°15'00"S, 78°55'00"W, 1750 m, Campos & ae "4504 (MO not seen, UC, USM Elaphoglossum exsertipes Mickel ee eS PERU. Amazonas: near border with Dept. San Martin, 5°41'S, 77°48'W, 2000 m, van der Werff 16650, et al. (MO not seen, UC). San Martin: along road Rioja-Pedro Ruiz, El Mirador, 5°40'29"S, 77°46'25"W, 1850 m, van der Werff 15659 (MO not seen, UC Elaphoglossum palmarum M. Kessler & Mickel lanes PERU. San Martin: Prov. Rioja: Pedro Ruiz-Moyobamba road, Km 390, Venceremos, 5 1800-1900 m, D. Smith 4503 (MO not seen, UC). Previously cited by cnr (in Tryon & aon ior ns as E. litanum (Sodiro) C. Chr, which is now known only from Ecuador. Elaphoglossum tovarense (Mett. ex Kuhn) T. Moore ex C. Chr., vel aff. (Dryopteridaceae) PERU. Amazonas: Prov. Bagua: Yamayakat, 04°55'S, 78°19'W, 320 m, Jaramillo 1189, et al. (MO). 2332 BRIT.ORG/SIDA 21(4) Enterosora trichosora (Hook.) L-E. Bishop (Grammitidaceae) PERU. Amazonas: near border with Dept. San Martin, 5°41'S, 77°48'W, 2000 m, van der Werff 16667, et al. (MO not seen, UC). Equisetum myriochaetum Schltdl. @ Cham. (Equisetaceae) PERU. Amazonas: Prov. Bagua: 25 km E of La Peca, 5900 ft, Barbour 2880 (MO). Cajamarca: Huarango, Nuevo Mundo, 1600-1700 m, Campos & Nunez 4623 (MO). Synonymized under E. giganteum by Tryon and Stolze (1994:15), but well characterized by Hauke (1963). Eriosorus hirtus (Kunth) Copel. (Pteridaceae) PERU. Piura: Prov. Ayabaca: Cerro de Aypate, Comunidad Campesina de Tacalpo, anexo Yanchala, 4°42) 82'S, 79°34'150'W, 2800-2880 m, Quipuscoa S. 643, et al. (F not seen, UC). Eriosorus novogranatensis A. F Tryon (Pteridaceae) PERU. Amazonas: San Martin, 5°41'S, 79°48'W, 1940 m, van der Werff 16729 (MO not seen, UC, USM), 16747, etal. (MO not seen, UC). Huperzia dichotoma (Jacq.) Trevis. (Lycopodiaceae) ERU. Pasco: Proy. Oxapampa: along road Chatarra-Pto. Bermudez, 10°30'S, 75°03'W, 700 m, van der Werff 18161, et al. (MO not seen, UC). a firma (Mett.) Holub, vel aff. (Lycopodiaceae) RU. Amazonas: Prov. Leymebamba: Dist. Leymebamba, ruta Laguna de Los Condores, La Atalyaya, slvededare de La Fila, 6°49.056'S, 77°44.134'W, 3000-3500 m, Quipuscoa S. 1206, et al. (F not seen, ee cen’ Se (Hymenophyllaceae) Je Maria, 800 m, Aguilar 304 (UC). Perhaps most (1989a O) = a specimens ened . Tryon and Stolze (1 559) as H.apiculatum Mett. ex Kuhn from Peru are better referred to H.dendritis,a low-elevation, snailer species (fronds mostly 3-5 cm long) with ovate, rather than obovate involucres, included, shorter receptacles (generally <1 mm long vs. often slightly exserted receptacles ca.2mm long in H.apiculatum), and smaller sporangia (ca. 0.3 mm long vs. 0.5 mm long) Meinl haat platylobum Bosch ey lees ERU. Puno: [Proy. Carabaya]: San Gaban, Lechler 2489 (Ho JS; soTyPE: F). Treated by ee (1947) and by Tryon and Stolze (1989a:68) as a synonym ol HL vaivatun Hook. & Grev,, but we believe the two are specifically distinct. Jamesonia verticalis Kunze (Pteridaceae) RU. San Martin: Dto. Huallaga, Saposoa, entre El Tambo y Jalca de El Rayo, caminoa Leymebamba, 2800-3200 m, Quipuscod S. 2484, et al. (F not seen, UC). Lellingeria aff. suspensa (L.) A.R.Sm.@ R.C. Moran (Grammitidaceae) PERU. Amazonas: near border with Dept. San Martin, 5°41'S, 77°48'W, 2000 m, van der Werff 16686, et al. (MO not seen, UC). Lindsaea bolivarensis V. Marcano (Lindsaeaceae) PERU. Loreto: Prov. Maynas: ca. 7 km E of oil palm plantation at Rio Maniti, 3°38'S, 72°56'W, 100-150 m, Tuomisto 4721, et al. (TUR, UC, USM); trail from the village of Panguana towards Rio Maniti, about m from Rio Amazonas, 3°53, 73°5'W, 100-200 m, Tuomisto 6124, et al. (TUR, UC, USM); Explorama / EER reserve, 5-15 km E of ie mouth of Quebrada Sucusari at lower Napo, 3°15'S, 72°50'W, 100- i Tuomisto 7755, etal. (AMAZ, KSP TUR); 2 km SE of the village Santa Ana at lower oe 4°6'S, 73°7'W, 100-200 m, Tuomisto 8863, with Oré (AMAZ, TUR, USM) (Marcano 1989:254). SMITH ET AL., NEW RECORDS OF PTERIDOPHYTES FROM PERU 2333 Lindsaea coarctata K.U. Kramer (Lindsaeaceae) RU. Loreto: Prov. Maynas: close to the village Nina Rumi at lower Rio Nanay, 20 km SW of Iquitos, 3°51'S, 73°23'W, 100-200 m, Tuomisto 5863 (AMAZ, TUR, USM). Prov. Mariscal Ramon Castilla: 3-4 km W from village of Puerto Izango, Rio Yaguasyacu, 3°18'S, 72°O1'W, 100-150 m, Tuomisto 11227, et al. (AMAZ, TUR, UC, USM). Prov. Requena: 3 km E from Requena, 5°04'S, 73°48'W, 150-200 m, Tuomisto 12764, et al. (AMAZ, TUR, USM); 4 km SE from Requena, 5°05'S, 73°49'W, 150-200 m, Tuomisto 12778, etal. (AMAZ, TUR, USM). a>) fea Lindsaea hemiptera K.U. Kramer (Lindsaeaceae) PERU. Loreto: Prov. Maynas: 14 km NW from the carretera to Nauta at km 40, 3°56'S, 73°30'W, 100- ney Tuomisto 3557, et al (AAU, AMAZ, TUR, USM, Z); Experimental station ‘El Dorado’ of INIA, Km 25 along road Iquitos-Nauta, 3°57'S, 73°25'W, 100-200 m, Tuomisto 13049, with Ruokolainen re TUR, UC, USM). Lindsaea javitensis Humb. & Bonpl. ex Willd. (Lindsaeaceae) PERU. Loreto: Prov. Maynas: Experimental station ‘El Dorado’ of INIA, Km 25 along road Iquitos- Nauta, 3°57'S, 73°25'W, 100-200 m, Tuomisto 13051, with Ruokolainen (AMAZ, TUR, UC, USM). Lindsaea tetraptera K.U. Kramer (Lindsaeaceae) PERU. Loreto: Es Maynas: Experimental station of UNAP at Puerto Almendras along Rio Nanay, 20km Iquitos, 3°51'S, 73°22'W, 100-200 m, Tuomisto 6439, with Ruokolainen (AMAZ, TUR ore close to the experimental station of UNAP at San Gerardo, km 13 of the road Iquitos- Nauta, 3°55'S, 73°22'W, 100-200 m, Tuomisto 7217, with Ruokolainen (AMAZ, TUR, USM); Experi- mental station ‘El fiigrade of INIA, Km 25 along road Iquitos-Nauta, 3°57'S, 73°25'W, 100-200 m, Tuomisto 13048, with Ruokolainen (AMAZ, TUR, U M). Lomariopsis prieuriana Fée tie aiccen®) PERU. Loreto: Prov. Maynas: C. Nueva Paleta, Rio Napo, 3°01'S, 73°21'W, Flores st 353, 363 oo San Martin: Prov. Rioja: road Rioja-Pedro Ruiz, 1450 m, van der Werff 15554 (UC). Additional colle tions from Loreto cited by Moran (2000:93). Megalastrum mollis A.R. om (Diyoptendaceae) PERU. Amazonas: Bagua Distr. Is B 5°16'57"S, 78°23'10'W, 750 m, van der Werff 16300, et al. (HOLOTYPE: nue: isoTyPE: MO) (Smith. 2005b), Melpomene assurgens (Maxon) A.R. Sm. & R.C. Moran (Grammitidaceae) RU. Cajamarca: Prov. San Ignacio: Tabaconas, Santuario Nacional Tabaconas-Namballe, 5°15'00'S, 79°19'00"W, 2300-2400 m, Campos 5691, et al. (MO not seen, UC, USM). Discussed under Grammitis moniliformis (Lag. ex Sw.) Proctor by Tryon and Stolze (1993:99) Melpomene peruviana (Desv.) A.R. Sm. & R.C. Moran (Grammitidaceae) PERU. Ancash: Prov. Huaras: N of Laguna Llanganuco, 9°03.1'S, 77°36.2' W, 4400, Lehnert 273 (UC). Ayacucho: Proy. Huamanga: Ayacucho- ercanuay aS above Denes, kaa Sh Oe 3'W, 4200 m, Lehnert 339 (UC, USM); Ayacucho-Andahuaylas, bel Vilcas Hi 1, 13°22'S 73°58'W, 4200 m, Lehnert 333 (UC, USM). Synonymized under Cravnonesiabe onAts (Poir) C. V. Morton by Tryon and Stolze (1993:102). Metaxya lanosa A.R.Sm. & Tuomisto (Metaxyaceae) Cited by Smith et al. (2001) from Loreto ladditional collections are now known: PERU Prov. Maynas: Distr. Iquitos, El Dorado-INIA, Varillal Alto Humedo, Mc Daniel & rene not seen, USM): Moran 3630 (USM); Rimachi 7526 (USM Microgramma acatallela Alston (Polypodiaceae) 2334 BRIT.ORG/SIDA 21(4) PERU. [Loreto]: Prov. Maynas: Iquitos, Estacion Biologica, Allpahuayo-IIA PBosque primario, 03°53'S, 73°25'W, 130 m, Jaramillo & Marcos 703 (MO not seen, UC). Pasco: Dist. Pozuzo, Parque Nacional Yanachaga Chemillén, 10°11'S 75°34'W, 1225 m, nein se etal. See not seen, i ); El Huampal, van der Werff 17930A, etal. (MO not seen, UC). M piloselloides, as defined by Tryon and Stolze (1993:152), are M. peat the former is erly concider to be restricted to the Antilles, s Mexico, and G (e.g., Mickel & Smith 2004:3 aie Aare fosteri B. Leon & H. Beltran (Polypodiaceae) -PER ov. Padre Abad: Cordillera Azul del Biabo, cabaceras del Rio Pisqui, 8°28'45.6'S, 73° eer i! a m, Beltran 3643, et al. Giocotyre: USM) (Leén & Beltran 2002). saan al caucanum (Hieron.) A.R.Sm.,, vel aff. (Grammitidaceae) PERU Prov. Oxapampa: road to Chacos, near top of ridge, 10°35'S, 75°06'W, 2400-2700 m, van der a nesae 18573, etal. (MO not seen, UC); Chacos, 10°37'S, 75°17'W, 2500 m, van der Werff 17697, et al. (MO); Parque Nacional Yanachaga-Chemillen, Abra Yanachaga, 10°22'S, 75°27'W, 2900 m, Vdsquez 28455, etal. (MO not seen, UC). Pecluma robusta (Fée) M. Kessler & A.R. Sm. (Polypodiaceae) PERU. San Martin: Prov. Huallaga: Dist. Saposoa, Arriba de Zarumilla, al NO del pueblo, 6°34'55'S, 77°23'06'W, 1350 m, Quipuscoa S. 2096, et al. (F not seen, UC) (Kessler & Smith 2005). Pleopeltis disjuncta M. Kessler & A.R. Sin. (Polypodiaceae) PERU. Cuzco: Cook & Gilbert 1510 (US), as cited by Kessler and Smith (2005). See also Tryon and Stolze (1993:137), under Polypodium furfuraceum Schltdl & Cham Pleopeltis stolzei A.R. Sm. (Polypodiaceae) Tr TE ] 1993-143) il yon an d Stolze (1993:143), Kessler and Smith (2005). A synonym is Pleopeltis PERU. Amazonas: see macrocarpa (Bory ex Willd.) Kaulf. var. laciniata Stolze. Polybotrya sessilisora R.C. Moran (Dryopteridaceae) PERU. Loreto: Prov. Loreto: 1.5 km E of road to Iquitos at Km 10 from Nauta, 4°28'S, 73°34'W, 100-150 m, vomit 4435, et al. (AAU, AMAZ, TUR, U, USM); Rio Tigre, 2 km E of the village Paraiso, 3°57'S, 7'W, Tuomisto 13977, et al. (AMAZ, USM, TUR); Rio Tigre, 7 km NW of the village era a, ue 75°22'W, Tuomisto 14560, et al. (AMAZ, TUR, UC, USM). Prov. Maynas: Explorama/ACE Reserve, 15-23 km E of mouth of Quebrada Bea at lower Napo, 3°15'S, 72°45'W, LOO- i m, yee 7853, etal. (AMAZ, KSP, TUR, UC); Upper Rio Itaya, about 5 km SSE of the village Carbajal, O'S, 73°35'W, LOO-200 m, Tuomisto 10120, et ia (AMAZ, TUR, US). + Polypodium appressum Copel. (Poly podiaceae) PERU. Amazonas: |zuchaca, 6°19'40'S, 77°31'05' W, van der Werff 16936, etal. (MO not seen, UC). Junin: Prov. Tarma: Agua Dulce, 1900 m, Woytkowski 35479 (UC). The species was subsumed under the much more common P.fraxinifolium Jacq. by Tryon and Stolze (1993:132) Polypodium attenuatum Humb. & Bonpl. ex Wil 2 (Polypodiaceae) PERU. Tumbes Hwy to“El Caucho,” 400 m, Coronado 230(UC). Misd by Tryon, in 1957. Polypodium (Pleopeltis) fayorum R.C. Moran & B. Ollg. (Polypodiaceae) PERU. Cajamarca: Prov. Santa Cruz: 1800 m, Sagastegui & Leiva 14092 (UC) (Moran & Ollgaard 1998:437) Polypodium funckii Mett. (Poly podiaceae) PERU. Amazonas: P care ae Chachapoyas-Mendoza, Km 52, Lehnert 232 (UC). Huanuco: Carpish, 2800 m, C tere 70 (UC SMITH ET AL., NEW RECORDS OF PTERIDOPHYTES FROM PERU 2335 Polypodium giganteum Desv. (Polypodiaceae) PERU. Pasco: Prov. Oxapampa: along road Chatarra-Cacazu, 10°32S, 75°04'W, van der Werff 18272, et al. (MO not seen, UC). Polypodium gilliesii C. Chr. (Polypodiaceae) Stork et al. 9283 (UC), Stork & Vargas 9338 (UC), Tryon & Tryon 5418 (BM, UC), from Dept. Lima, ie 12956, et al. (UC), from Dept. La Libertad, as well as other specimens seen from Dept. Lima an pt. Lambayeque from low elevations are this species, and not P. lasiopus Klotzsch, as cited Tryon Stolze (1993:130). Polypodium gilliesii citer from P. laste pus: where it was eynonyanized by bo th Hensen (1990) and Tryon and Stolze (1993 3), b ly glabrous Or nearly glabrous blades (lacking long, septate hairs) and generally longer, narrower, somewhat diverging, and darker rhizome scales arnost prominently se seen in i. Veneztielan specimens, the type locality); in addition, P. Peru, while P gillliesii is recorded from 400-800(-1300) m, nanan on lomas. weed acs intricatum M. ea & A.R. Sm. (Polypodiaceae) RU. Amazonas: Prov. Luya: Dist. Camporredondo, Tullanya, Pascana, La Palma, 6°04'35'S, are W, 2710 m, Vasquez sn et a Mo not seen, UC, USM). Cajamarca: Campos & Nunez 4647 (UC). San Martin: Prov. Rioja: Pedro -Moyobamba road, Km 390, 77°45'W, 05°50'S, 1800 m, Smith 4414 (MO not seen, UC, USM). a also i: yon and Stolze (1993:130), most specimens cited un- der P loriceum L. Polypodium kunzeanum C. Chr. (Polypodiaceae) PERU. Amazonas: Prov. Bongara: Shillac, north by trail from Pedro Ruiz, 5°49'S, 78°O1'W, 2300 m, D. Smith 4945, with Vasquez S. (MO not seen, UC). Huanuco: between Chinchao and Puente Durand, 2000 m, Coronado 85 (UC). Cited by yon and silze euan as a synonym of P. sessilifolium Desv., g ee | i re gul ar venation, some- which we believe is not closely related. P e times with two included and themselves ang astomcsing veins within an areole, cordate-based, non- adnate pinnae (the basalmost with | | pping the rachis), and non-clathrate rhizome scales. Both collections seen from Peru, neither cited by Tryon and Stolze (although Smith 4945 was anno- tated as P triseriale vel aff.), are indicated as growing terrestrially. Polypodium loriciforme Rosenst. (Polypodiaceae) PERU. Pasco: Prov. Oxapampa: Dist. Chontabamba, carretera a la Suiza, 10°39'S, 75°27'W, 2130-2210 m, Monteagudo 4539, et al. (MO not seen, UC). Polypodium maritimum Hieron. (Polypodiaceae) PERU. Cajamarca: Prov. San Ignacio: San José de Lourdes, 4°59'22"S, 78°53'03'W, 2020 m, Vdsquez 26336, et al. (MO not seen, UC); San José de Lourdes, camino al Cerro Picorana, 5°O1'40"S, 78°54'30"W, 2100-2200 m, Campos 5507, et al. MO not seen, UC). Polypodium polystichum Link (Polypodiaceae) PERU. Huanuco: Prov. Huanuco: Dist. cee ba, Hacienda Mercedes, Cotirarda, 1560 m, Mexia 8216a (UC); Dist. Churubamba, Hacienda Exito, bank of Rio Ysabel, 1100 m, Mexia 8163 (UC). Pasco: Proy. Oxapampa: Gran Pajonal, vicinity of ee 10°45'S, 74°23'W, 1250 m, D. Smith 5161 (MO not seen, UC). Related to P.fraxinifolium Jacq. but with spreading, long-acuminate scales. Polypodium (Pleopeltis) tweedianum Hook. (Polypodiaceae) PERU. Cajamarca: Prov. Contumaza, entrada al Bosque Cachil, 2500 m, Sagdstegui 15107, et al. (F not seen, UC). Related to P. pycnocarpum C. Chr, but with sharply bicolored rhizome scales with a scle- rotic mid-stripe Polystichum cochleatum (Klotzsch) Hieron. (Dryopteridaceae) 2336 BRIT.ORG/SIDA 21(4) PERU. See Tryon and Stolze (1991:54), under P. pycnolepis (Klotzsch) T. Moore; the name accepted b Tryon and Stolze for this species is, in our opinion, a synonym of their P. orbiculatum (Desv.) J. Rémy Polystichum rufum M. Kessler & A.R. Sm. (Dryopteridaceae) PERU. San Martin: Prov. Lamas: Dist. panies below pean Eyanceucas Mission, Lamas, Belshaw 3428(UC; GH, US not seen). $ Ro and Stolze (1991:52) Gg 7 osenst. by Tryon f f Polystichum stuebelii Hieron. (Dryopteridaceae) E co: Prov. Urubamba: Dist. Ollantaytambo, Huaytampo, 13°09'02"S, 72°30' 28"W, 2400 m, Calatayud 109, et al. (MO not seen, UC). Pasco: Prov. Oxapampa: Dist. Oxapampa, Parque Nacional Yanachaga Chemillén, cercanias del Refugio el Cedro, 10°32'S, 75°21'W, 2420 m, Monteagudo 3814, et al. (MO not seen, UC). Probably included within P montevidense (Spreng.) Hieron. by Tryon and Stolze (1991), but the name P stuebelii not mentioned by them Polytaenium brasilianum (Desv.) Benedict (Vittariaceae) PERU. Cuzco: Prov. Convencion: Tupitari, 2000 m, Vargas C. 3440 (UC). This species was recognized as distinct by Tryon (1964), and the Vargas specimen cited as representative, but Antrophyum brasilianum (Desv.) C. Chr. was synonymized by Tryon and Stolze (1989b:87) under A. cajenense (Desv.) Spreng. [= a tea i Hattee Desv) Benedict hes atter differs in nav ng oblenbeglate (vs. ellip- tic) blades and darkened (vs iall towards the base of the blades. Pteris consanguinea Mett. ex Kuhn (Pteridaceae) PERU. Amazonas: Prov. Bagua: |2 km E of La Peca, 1700 m, Barbour 2497 (MO not seen, UC). Cajamarca: Prov. San Ignacio: San José de Lourdes, 5°01'00"S, 78°57'00"W, 1500-1600 m, ol 3890, et al. (MO not seen, UC); San José de Lourdes, 4°59'22"S, 78°53'03"W, 2020 m, Vasquez 26293, et al. (MO not seen, UC). San Martin: Proy. Huallaga: Dist. Saposoa, al sur de Anazco Pueblo, 2000 m, Quipuscoa S. 2328 et al. (F not seen, UC). Specimens placed here would key to P decurrens C. Presl in Tryon and Stolze (1989b:77), but that species differs in a number of respects (see Prado & Windisch 2000). Pteris muricatopedata Arbelaez (Pteridaceae) PERU. Huanuco: Prov. Leoncio Prado: Dtto. Emileo Baldizan, Tingo Maria-Pucallpa La Divisora road, 1600 m, Rimbach 4993 (NY not seen). Pasco: Prov. Oxapampa: 4-5 km N of Mallampampa, 2400 m, Smith & Canne 5786 (NY not seen). Additional collection from Pasco cited by Arbelaez (1995:177). Saccoloma membranaceum Mickel (Dennstaedtiaceae, temporarily) PERU. Amazonas: Dist. Bagua, Imaza, Quebrada Almendro, 5°18'S, 78°20'W, 400 m, Vasquez 26106, et al. (MO not seen, UC); Dist. Bagua, along road Imaza-Chiriaco, 5°03'24'S, 78°20'17"W, 400 m, van der Werff 16182, et al. (MO not seen, UC); Dist. Imaza, Tayu Mujaji, Comunidad de Wawas, 5°15'56"S 78°22'07'W, 900 m, Vasquez 24644, et al. (MO not seen, UC). Pasco: Prov. Oxapampa: along road Chatarra-Cacazu, 10°32'S, 75°04'W, 890 m, van der Werff 18249 (MO not seen, UC) Salpichlaena hookeriana (Kuntze) Alston (Blechnaceae) PERU. Loreto: Proy. Maynas: ca. 10 km SW of Iquitos at zoological park, Moran 3672 (UC); 7 km E of the oil palm plantation at Rio Maniti, 3°38'S, 72°56'W, 100-150 m, Tuomisto 4771, et al. (AAU, AMAZ, TUR, U, USM); Explorama/ACEER reserve, 23-31 km E of the mouth of Quebrada Sucusari at lower Napo, 3°14'S, 72°39'W, 100-200 m, Tuomisto 7880, et al. (AMAZ, KSP. TUR, US); Upper Rio Itaya, about 6 km SSE of the village Carbajal, i YS, 73°35'W, 100-200 m, Tuomisto 10083, et al. (AMAZ, NY, TUR). rov. Mariscal Ramon Castilla: 2-5 km SW from village of Puerto Izango, Rio Yaguasyacu, 3°18'S, 72°OI'W, 100-150 m, Tuomisto 11252, et al. (AMAZ, TUR, UC, USM). Prov. Requena: 2 km N from the biological station of Jenaro Herrera, 4°52'S, 73°39'W, 150-200 m, Tuomisto 12785, et al. (AMAZ, TUR, SMITH ET AL., NEW RECORDS OF PTERIDOPHYTES FROM PERU 2337 USM). Prov. Loreto: 5 km upriver from mouth of Rio Pucacuro, 3°17'S, 74°59'W, Tuomisto 14123, et al. (AMAZ, TUR, USM). Madre de Dios: Prov. Manu: 4 km S from the village of Boca Colorado, 12°38'S, 70°25'W, ao m, tose poe et al. Ue. TUR, ees subsumed by yen and Stolze ae 70) J ly jy oe Ie A oe dis inguished b strongly under isti dimeeohic amet a See ca. 1-2 mm wide and the buds in the ae of Seale pinnae. Tryon and Stolze also cited two specimens from Loreto (not seen) tk kely this species. Schizaea fluminensis Miers ex J. Sturm (Schizaeaceae) PERU. Loreto: Maas 6336, et al. (AMAZ, USM), Mejia s.n. (USM). Cited by Vasquez (1997) for Loreto; see also Leon et al. (in press). Selaginella arthritica Alston (Selaginellaceae) PERU. Huanuco: Prov. Leoncio Prado: along road from Tingo Maria to Pucalpa, less than | km N of Sortilegio, 9°13'16"S, 75°50'15' W, 1310 m, Croat 81744, & Sizemore (MO not seen, UC, USM) Selaginella fragilis A. Braun (Selaginellaceae) PERU. Loreto: Prov. Requena: 140 m, van der Werff 1010, et al.(MO not seen, UC). Subsumed by Tryon and Stolze (1994:84) under S. parkeri (Hook. & Grev.) Spring, but this specimen, at least, seems spe- cifically distinct, with much narrower penultimate divisions and long-flagelliform branch apices. oe leucoloma Alston ex Crabbe & Jermy (Selaginellaceae) PERU. Pu ail from Aricoma Pass to Santo Domingo, 5800 ft, McCarroll 126, pro parte (MICH). ee by eae (1995:366). Selaginella macilenta Baker (Selaginellaceae) PERU. Junin: Chanchamayo, 750 m, Kunkel 347 (S). Cited by Valdespino (1995:322). Selaginella moritziana Spring (Selaginellaceae) PERU. Pasco: Prov. Oxapampa: San Alberto, 10°32'S, 75°21'W, 2400 m, van der Werff 18597, et al. (MO not seen, UC). Selaginella pearcei Baker, type from Peru, was subsumed under S. novae-hollandiae (Sw.) Spring by Tryon and Stolze (1994:77), but it is referred to S. moritziana var. pearcei (Baker) Valdespino, ined., by Valdespino (1995). Selaginella palmiformis Alston ex Crabbe & Jermy (Selaginellaceae) PERU. Loreto: Prov. Mariscal Ramon Castilla: ca. 3 km S of Huanta, 3°17'S, 72°51'W, 100-150 m, Tuomisto 5231, et al. (AAU, AMAZ, TUR, UC, USM). Prov. Maynas: surroundings of tourist lodge of Explorama Tours at Rio Sucusari, 3°10'S, 72°52'W, 100-150 m, Tuomisto 5818, et al. (AAU, AMAZ, TUR, UC, USM); Maynas, trail from village of Panguana towards Rio Maniti, ca. 8 km from Rio Amazonas, 3°53'S, 73°05'W, 100-200 m, Tuomisto 6101, et al. (AMAZ, TUR, UC, US); Explorama/ACEER Reserve, 1.6-3.3 km E of mouth of Quebrada Sucusari at lower Napo, 3°15'S, 72°53'W, 100-200 m, Tuomisto 7712, et al. (AMAZ, KSP. TUR, UC, US). Prov. Mariscal Ramon Castilla: 3-4 km W from village of Puerto Izango, Rio Yaguasyacu, 3°18'S, 72°01'W, 100-150 m, Tuomisto 11232, et al. (AMAZ, TUR, UC, USM) sea aa tomentosa Spring (Selaginellaceae) RU. Amazonas: Rio Cenepa, second ridge E of Huampami, 900-1000’, Berlin 633 (MO, UC). This species was ee under S. Semi cU ate Cc pie ee by Tryon and Stolze (1994), but S. tomentosa is easily distinguished by the decidedly short-hairy stems. It has been recognized by nearly all other pteridologists, e.g., by Alston et al. (1981:306). - Sticherus aurantiacus Ostergaard & B. Ollg. (Gleicheniaceae) PERU. San Martin: [Proy. San Martin]: Tarapoto, 750 m, Williams 5972 (US); between Moyobamba and Huallaga, Sttibel 1103 (B). 2338 BRIT.ORG/SIDA 21(4) Sticherus boliviensis (Maxon & C. V. Morton) J. Gonzales, comb. ined. (Gleicheniaceae) PERU. Puno: Prov. Sandia: Limbani, 3200-3450 m, Metcalf 30539 (GH, MO, US). Sticherus = Snes nee comb. ined. (Gleicheniaceae) PERU. Amazonas: Prov. Bagua: between Aramango and Montenegro, 275 m, Lépez 4163(GH). Cuzco: Asuncion, 1200 m, sia 7908 (GH), ae between Inambari and Quincemil, 500-650 m, Vargas C. 16487 (H). Huanuco: Proy. Leoncio Prado: Tingo Maria, 710 m, Tryon & Tryon 5263 (GH, USM). vee Prov. Chanchamayo: La Merced, Kunkel 652 (GH). Loreto: [Prov. Alto Amazonas]: Pumayacu, betwee n Balsapuerto and Moyobamba, 600-1200 m, Krug 3249 (MO, NY). San Martin: road Reena ne imaguas, Km 12-15, 2250 m, Hickok 646 (GH). See Gonzales (2003). Sticherus ferrugineus (Raddi) J. Gonzales, comb. ined (Gleicheniaceae) PERU. Cuzco: Prov. Convencion: Rio Apurimac, above Hacienda Luisiana, Wade 1333 (GH). Loreto: Rio Maranon Valley, between mts. of Rio Pastaya and Rio Huallaga, San Lorenzo, 150 m, Killip 29219 (NY, US). See Gonzales (2003). Sticherus lanosus (H. Christ) J. Gonzales, comb. ined. (Gleicheniaceae) PERU. Cuzco: 5 km N of Aguas Calientes, 2000 m, Solomon 3172 (MO); Urubamba, Machu Pichu, 2000 m, Saunders 1232 (GH). Huanuco: Cerros del Sira, 9°25'S, 74°44'W, 1560 m, Dudley 13193 (GH). Puno: Carabaya, Ollachea-San Gaban road, Chacaneque, Boeke 3139 (MO, NY); Valle de Marcapata, 2000 m, Herrera 1592 (US). See Gonzales (2003 pe Sticherus lanuginosus (Fée) Nakai (Gleicheniaceae) Previously often called S. penniger (Mart.) Copel. [= Gleichenia pennigera (Mart.) T. Moore, in Tryon & Stolze 1989a], which is considered a synonym of S. pruinosus (Mart.) Ching, by Gonzales (2003). Sticherus melanoblastus Ostergaard & B. Ollg. (Gleicheniaceae) PERU. Pasco: Proy. Oxapampa: La Suiza Nueva, 10°38'S, 75°27'W, 2240 m, van der We Be om (MO not seen, UC). Mentioned as possibly in Peru by Ostergaard Anderson and Ollgaard (200 Sticherus velatus (Kunze) Copel. (Gleicheniaceae) PERU. Huanuco: Pampayacu, Jul 1829, Poeppig s.n. (HoLoTyre: W, the fragment of Poeppig s.n. at US annotated as isotype of S. velatus is in fact S. lanosus). Cuzeo: La Convencion, Valle Santa Ana, 1000- 1500 m, Herrera nae (US); omnes | HoepPig 1117 (W). See Gonzales ee) ums ppecies synony- mized under Glei ld tomentosa ( rex Sw.) Spreng by Trvon and § Tectaria microsora ™ R. Sm. (Dryopteridaceae) PERU. Amazonas: Pro Distr. Imaza, Comunidad ake de ek 4° 55'S, 78° 19'W, 680 m, Rodriguez R. 967, et a “oto HUT, tsorypes: MO not seen, UC, USM), Distr. Imaza, region del Maranon, comunidad de Yamayakat, Sicnede cae Rio one 04° 55'S, 78° 19'W, 550 m, Vasquez 19644 (MO, UC); same locality, 600 m, Rodriguez R. 288 (MO, UC, USM); Dist. Bagua, along road from Chiriaco towards Bagua, 05° 16'57"S, 78° 23), LO"W, 800 m, van der Werff 16260 (MO, UC). Prov. Condorcanqui: Distr. Fl Cenepa, region N iental del Maranon, Rio Cenepa, comunidad Tutino, 04° 33'S, 78° LO'W, 350 m, Vdsquez 18404, et al. (MO, UC); see Smith (2005b). Tectaria pilosa (Fée) R-C. Moran (Dryopteridaceae) PERU. Amazonas: Prov. Bagua: Dtto. oi Comunidad de Yamayakat, 05°03'24'S, 78°20'17 W, 450 m, Rojas 567, et al. (MO not seen, UC). Prov, Condorcanqui: Dtto. El Cenepa, Comunidad Aguaruna Pagki-Suwa, Rio Cenepa, quebrada Tayo, o4e 31'35"S, 78°10'34"W, 289 m, ae 22151, et al.(MO not seen, UC, USM). elena Prov. La Mar: between Santa Rosa and Hacienda Luisiana, 640 m, a. Wasshausen & E 630 (US not seen, USM). Loreto: [Prov. Alto haces: above Pongo de SMITH ET AL., NEW RECORDS OF PTERIDOPHYTES FROM PERU 2339 Manseriche, bank of Rio Santiago, 200 m, Mexia 6354 (UC). Prov. Maynas: Explor Napo Camp at Rio Sucusari, 03°20'S, 72°55'W, 120 m, van der Werff 12926, et al. (MO not seen, UC). Treated by Tryon and Stolze (1991:25) as a variant of T. incisa, but we think T. pilosa is adequately distinct, at species rank. They also cited specimens from Depts. Cajamarca, Huanuco, Madre de Dios, and Pasco that may be this species. Tectaria pubens R.C. Moran (Dryopteridaceae) PERU. Loreto: Prov. Maynas: ca. 50 km an nriver igs quires Explorama Lodge, ca. 120 m, Moran 3647 (HoLoTYPE: MO; IsoTyPE: UC); addi llocality cited by Moran (1992:138). Terpsichore chrysleri (Copel.) A.R. Sm. (Grammitidaceae) PERU. Amazonas: [Prov. Bongara]: Laguna de Pomacochas, 2550 m, van der Werff 15809, et al. MO not seen, UC). Cajamarca: Prov. San Ignacio: Tabaconas, El Pajonal, camino al Paramo y al Cerro Coyona, 5°17'30"S, 79°16'02"W, 2250 m, Ca bie 5728, et al. (MO not seen, UC); San José de Lourdes, 5°00'S, 78°54'W, 1800 m, Vdsquez 26157, et al. (MO not seen, ve Supe ume under ‘Sramimnns cae (L ss taosto ey Tryon and Stolze os 104), but easily most) G. asplenifolia are, in fact, Terpuchigrs chrysleri, specifically van oe Werff 8608 (UC) and Woytkowski 35485 (UC), but Terpsichore asplenifolia (L.) AR. Sm.s. str. also occurs in ia eae mollissima (Fée) A.R. Sm. (Grammitidaceae) PER : Prov. Oxapampa: Parque Nacional Yanachaga, El Huampal, 10°H'S, 75°34'W, 1200 m, van ee Werff 17846. et al. (MO not seen, UC). San Martin: Prov. Rioja: Moyobamba-Pedro Ruiz, Km 383, 5°50'S, 77°30'W, ca. 2000 m, Lehnert 221 (UC, USM). Terpsichore subtilis (Kunze ex Klotzsch) A.R. Sm. (Grammitidaceae) PERU. Amazonas: near border with San Martin, 5°41'S 77°48'W, 2000 m, van der Werff 16675, et al. O) ee youngii B. Leon & A.R. Sm. (Grammitidaceae) U. Cuzco: near San Lorenzo, 2300-2500 m, Leon & Young 4487 (HOLoTyPE: USM; Isotype: UC) (Leon . aoe 2003:84). Thelypteris cinerea (Sodiro) A.R. Sm. (Thelypteridaceae) PERU. Amazonas: road Chachapoyas-Mendoza,a little past Molinopampa, 6°14'11"S, 77°35:49"W, 2400 m, van der Werff 15095, et al. (MO not seen, UC). Thelypteris steyermarkii A.R. Sm. (Thelypteridaceae) ERU. Cuzco: Prov. Paucartambo: Kosiipata Valley, Km 150, San Pedro, Rio Union and Rio Kosnipata junction, 1800 m, in sandy beach-river, Nunez 11958 (MO). Trichomanes accedens Hook. (Hymenophyllaceae) PERU. Amazonas: Quebrado El Almendro, 5°14'40'S, 78°21'24"W, 430 m, van der Werff 14560, et al. (MO not seen, UC). Loreto: Prov. Requena: 140 m, van der Werff 10112, et al. (MO not seen, UC). Prov. Maynas: Mishana, along Rio Nanay, 140 m, van der Werff 10193 (MO not seen, UC); Experimental station of UNAP at Puerto Almendras along Rio Nanay, 20 km air distance from Iquitos, 3°51'S, 73°22'W, 100-200 m, Tuomisto 6429, with Ruokolainen (TUR, US, USM). Discussed and considered probably synonymous with T. cristatum Kaulf. by Tryon and Stolze (1989a:98). Trichomanes dactylites Sodiro (Hymenophyllaceae) PERU. Cajamarca: Prov. San Ignacio: San José de Lourdes, laderas del Cerro Picorana, 4° 58'00'S, 78°53'01'W, 2500-2540 m, Campos 5933, et al. (MO not seen, UC). 2340 BRIT.ORG/SIDA 21(4) Trichomanes pilosum Raddi (Hymenophyllaceae) PERU. Amazonas: Prov. Bagua: Dist. Imaza, Quebrada Almendra, 5°14'40"S, 78°21'34"W, 400 m, van der Werff 16124, et al. (MO not seen, UC) Trichomanes spruceanum Hook. (Hymenophyllaceae) PERU. Loreto: Prov. Maynas: Dist. Iquitos, Puerto Almendras, 130 m, on white sand, van der Werff 9845, et al. (MO not seen, UC). DELETIONS FROM THE FLORA Elaphoglossum peruvianum (L.D. Gomez) Mickel (Dryopteridaceae) Treated by Mickel (in Tryon & Stolze 1991:170, under Peltapteris), but now regarded asa heterotypic synonym of Elaphoglossum moorei (E. Britton) H. Christ, according to Moran and Mickel (unpub- lished ms.); we also subsume Peltapteris in Elaphoglossum. Megalastrum yungense (H. Christ & Rosenst.) A.R. Sm. & R.C. Moran es * x 505 ss XS SS 825 es BS SoS Jo Or oS $50 S558 SESS JOOS SO OI Freee Cee SS2OO eetatetate esececeeat Richland C Setseces ~ seecen s oS KRY RSS a ROOD Legend ——— Streams CI Giles County Physiographic Provinces CES) Eastern FEg Outer uter Nashville Basin im ie exe) Western Highland Rim Fic. 1. Map of Giles County, Tennessee showing location of the Western Highland Rim, Eastern Highland Rim, Outer C | (Nashville) Basin, major streams, and th (ad 1 fi Griffith et al. 1998) 7 1) t | 2346 BRIT.ORG/SIDA 21(4) of the county’s soils. On high winding ridges in east-central and northeastern Giles County, on outliers of the WHR, cherty acidic soils of the Bodine-Fuller- ton-Dellrose (BFD) association occur, occupying 10% of the county. The Dellrose- Bodine-Mimosa association (DBM) is a cherty and rocky soil found on steep slopes, ridgetops, low-lying knobs, and deep hollows. These soils vary from slightly to strongly acid and make up 27% of the county’s soils. In the OCB in valleys of the Elk River, Richland Creek, and their larger tributaries, in bot- tomlands, on terraces, and on adjacent uplands, soils of the Staser-Armour- Maury (SAM) association occur. These soils are often phosphatic and are neu- tral to acidic; 32% of the county’s soils are of this association (True et al. 1968). Giles County lies within the Tennessee River drainage system. One of the Tennessee River's major tributaries, the Elk River, crosses the southeastern quar- ter of the county. The main tributary of the Elk River is Richland Creek. This large creek enters Giles County from the northeast, flows south through the central portion of the county, and empties into the Elk River in the south-cen- tral section. Both the Elk River and Richland Creek are medium-sized streams lined mostly by agricultural fields and wooded slopes. Limestone bluffs are fre- quent along the Elk River and the lower section of Richland Creek. Both streams are characterized by having a shallow channel, a slow to moderate flow,a rocky or gravelly substrate, and numerous gravel bars and shoals. Other important streams in the county include Big, Bradshaw, Buchanan, Indian, Shoal, Sugar, and Weakley creeks. These streams occur in the smaller valleys and are lined mostly by small fields and wooded slopes. They, like the Elk River and Richland Creek, have a rocky or gravelly substrate and frequent gravel bars, shoals, and riffles. While most of the streams in the county are like those described above, Piney Creek on the EHR of extreme southeastern Giles County is a slow-flow- ing stream that resembles streams of the southeastern Coastal Plain. It has a muddy substrate and is bordered mostly by wet flatwoods. The study area is located in Képpen’s Cfa climatic type and is character- ized by a mild rainy climate with hot summers and lacks a distinct dry season (Ackermann 1941). The average growing season is 190 days and extends from April 13 to October 20. The mean annual temperature is 15°C. July is typically the hottest month with an average temperature of 32°C, while January is the coldest with an average temperature of -0.4°C. Annual precipitation totals 139.04 cm with snow accounting for about 12.19 cm. February and March are the wettest months and September and October the driest (True et al. 1968). Giles County is within the eastern portion of Braun’s (1950) Western Me- sophytic Forest Region approximately 80-90 km west of the Mixed Mesophytic Forest Region. The Western Mesophytic Forest is transitional to surrounding forest regions and lacks a combination of characterizing dominants. Local cli- mate, topography, and soil conditions influence vegetational characteristics of a particular area (Braun 1950, Chester 1995). Most of the county’s forests clearly — ~— ESTES, FLORA UF , LENNESSEE 2347 fit into Braun’s Western Mesophytic Forest Region but in the eastern portion of the county, on sheltered north slopes and in ravines, forests with qualities of Braun’s (1950) Mixed Mesophytic Forest Region occur. METHODS Specimens were collected between July 1998 and September 2004. Thirty plant communities, including all types known to occur within the county, were sampled during the study. Collecting sites were located by driving throughout the county, and by consulting county road, topographic, and county soil sur- vey maps. Specimens were identified using standard field manuals: Small (1933), Fernald (1950), Radford et al. (1968), Cronquist (1980), Isely (1990), Gleason and Cronquist (1991), and Yatskievych (1999). In addition to native taxa, many non- native species were collected or observed during the survey. Non-native taxa listed in the annotated checklist include only those taxa that appeared to be naturalized or persistent. Most voucher specimens have been deposited in the herbarium at the University of Tennessee (TENN). Some specimens have been deposited in the herbaria of Austin Peay State University (APSC), Middle Ten- nessee State University (MTSU), or the Vanderbilt (VDB) collection at the Bo- tanical Research Institute of Texas. Furthermore, the herbaria at MTSU, TENN, and VDB were consulted for species not collected during this study. All such specimens encountered were checked for accuracy and each was annotated. The status of federal and state listed rare species was taken from the Tennessee Natu- ral Heritage Program (2003). All information concerning the location, habitat, and population status of rare species discovered during the inventory was pro- vided to the Tennessee Natural Heritage Program to aid in their protection. Lastly, the plant communities of the county were qualitatively described ac- cording to their physiographic location, soil association, topographic position, and species composition. RESULTS AND DISCUSSION Floristic Summary Of ca. 2000 voucher specimens, 1186 species and infraspecific taxa were col- lected, resulting in 912 county records. An additional 22 taxa were observed during the study for which vouchers were not collected. Therefore, the total flora of Giles County included 1208 species and infraspecific taxa representing 138 families and 553 genera distributed among 36 pteridophytes, 5 gymnosperms, 28 monocots, and 883 dicots. Asteraceae was the largest family with 15] taxa, followed by Poaceae (109), Cyperaceae (75), Fabaceae (63), Rosaceae (53), Lamiaceae (34), Liliaceae (33), Brassicaceae (31), Ranunculaceae (30), and Scrophulariaceae (30). The woody flora of the county was quite large with 254 taxa. The largest woody genera were Quercus (18 taxa, including one hybrid), Prunus (10), Rubus (9), Carya (8), Crataegus (7), Acer (6), and Salix (6). Cornus, 2348 BRIT.ORG/SIDA 21(4) Euonymus, Hypericum, Ulmus, and Vaccinium each had five taxa. Aesculus (in- uding one hybrid), Ilex, Lonicera, Pinus, Rhus, Smilax, and Vitis each had four taxa. Woody genera represented by three taxa included Celtis, Clematis, Fraxinus, Hydrangea, Magnolia, Rhododendron, Rosa,and Viburnum. The larg- est herbaceous genera were Carex (47), Juncus (14), Solidago (14), Polygonum (13), Eupatorium (12), Symphyotrichum (12), Viola (12), Desmodium (1), Dichanthelium (1D, Asclepias (10), Cyperus (10), Helianthus (10), and Ranun- culus (10). Seventy-nine percent of the flora (955 taxa) consisted of native spe- cies while introduced taxa accounted for 237 taxa (20%) (Wofford and Kral 1993). An additional eight species (<1%) were native to portions of Tennessee but did not appear to be native to Giles County. A complete summary of the flora of Giles County is provided in Table 1. ae Rare/Protected Plants In Giles County, 17 taxa are considered rare and are tracked by the Tennessee Division of Natural Heritage (Table 2). Of these, two are federally threatened (LT), two are state endangered (E), five species are listed as state threatened (T), one is listed as special concern-proposed threatened (S-PT), three species are listed as special concern (S), and the remaining four taxa are threatened or of special concern due to commercial exploitation (FCE, S-CE) (Tennessee Natu- ral Heritage Program 2003). Prior to this survey, seven rare species had been reported from the county. However, one previously reported rare taxon, Galium asprellum Michx.(R. Kral 64888, VDB), was based ona misidentified specimen of Galium mollugo,a non- native species. Another, Allium tricoccum, was reported from the county based on a Kral collection (R. Kral 54907, VDB). Upon closer inspection, it was deter- mined that the specimen best corresponds with A. tricoccum var. burdickii in- stead of the more eastern A. tricoccum var. tricoccum. Two additional taxa, Leavenworthia exigua var. exigua and Schoenolirion croceum, were reported from the county based on collections made in the 1950s; unfortunately the popu- lations were not relocated during the study and these taxa are presumed extir- pated from the county. A fifth species, Arenaria lanuginosa, was collected for the first time in Tennessee from Giles County in 1948 along bluffs of the Elk River (AJ. Sharp, S. Fairchild, & E. Clebsch 9840, TENN). As of 1999, the species was listed as endangered and possibly extirpated from the state (Tennessee Natural Heritage Program 1999). In the summer of 2000, A. lanuginosa was re- discovered along two bluffs of the Elk River after not having been observed for over 50 years (Estes & Chester 2001). A third Giles County population was found in 2001 (Estes 2004). Asa result of this study, I] rare taxa were discovered that represent new county records. The most significant were Apios priceanaand Helianthus eggertii (United States Fish and Wildlife Service 1993, 1999), both listed as federally ESTES, FLORA VP OILED LUUNT I, TENNESSEE 2349 Taste 1. Summary of the vascular flora known from Giles County, Tennessee. Species and Infraspecific Taxa Group Families Genera Native Non-native Total Pteridophytes 14 25 36 ) 36 Gymnosperms 2 2 2 3 5 Angiosperms 19 101 231 53 284 Dicots 103 425 686 197 883 Total 138 553 955 253 1208 Taste 2. The rare vascular plant species known from Giles County, Tennessee. Scientific Name Federal Status State Status Apios priceana ne E? Helianthus eggertii LT i baa anemia eee E E icifuga rubifolia a Jug ans cinerea alk a f | i eer ji is SN rr llaria fontinalis T Xyris is ace var. | ee S-PT* aa stanea asm S? .exigua S he metanehile taleai cus S * Allium tricoccum var. burdickii T=CE® Lilium michiganense T-CE Hyd tic canqdenci S-CE’ Panax quinquefolius S-CE * Rare taxon known from Giles County prior to this study ' Taxon listed as threatened in the United States ’ Taxon listed as endangered in Tennessee Taxon listed as threatened in Tennessee “Taxon currently listed as special concerned-proposed threatened in Tennessee ° Taxon listed as special concern in Tennessee ® Taxon listed as threatened in Tennessee due to commercial exploitation ‘Taxon listed as special concern in Tennessee due to commercial exploitation threatened by the United States Fish and Wildlife Service. Estes (2004) pro- vided a more detailed discussion of the significance of these and other notable rare or uncommon plants of Giles County. 2350 BRIT.ORG/SIDA 21(4) Additional Noteworthy Collections A large established population of the Asiatic species Achyranthes japonica var. hachijoensis was discovered in extreme southern Giles County on a wooded floodplain of the Elk River at the base of a limestone bluff ca. 3 river miles up- stream from Limestone County, Alabama. Medley et al. (1985) first reported this species as new to North America based on material collected from northeast- ern Kentucky and adjacent West Virginia. This species also has been reported from Ohio (Flora of North America Editorial Committee 2003). The discovery of A. japonica var. hachijoensis in Giles County represents a state record, a range extension of ca. 500-600 km to the southwest, and the first report of the species from the Tennessee River watershed. Euonymus kiautschovicus, an introduced shrub native to China, was col- lected from two localities in Giles County, one along a roadside embankment at the edge of disturbed woods and the other from a weedy bank between a stream and roadside. This species has not previously been reported from Ten- nessee. At both sites, only one or two plants were found and did not seem to exhibit strong invasive potential. Eupatorium X pinnatifidum, a hybrid species originating from a cross be- tween E. capillifoliumand E. perfoliatum, was collected for the first time in Ten- nessee from flat uplands on the EHR at the edge of a clearing adjacent to wet flatwoods dominated by Acer rubrum, Pinus taeda, Quercus nigra, and Q.phellos. Both E. capillifolium and E. perfoliatum were growing within 10 m of the E. x pinnatifidum plants. Atthe site where E. X pinnatifidum occurred, several individuals of Baccharis halimifolia,a species formerly restricted to the southeastern Coastal Plain were observed. The shrubs were approximately 1-2 m tall. One month prior to the discovery of B. halimifolia in Giles County, one individual was discovered ca. 87 km to the NNE in Rutherford County, Tennessee (Estes 2004). This is the second report for B. halimifolia from the state. Baccharis halimifolia should be considered an exotic species in Tennessee and is to be expected in disturbed areas across the southern half of the state. third species found in Giles County that deserves special mention is Amaranthus powellii. This species was discovered in August 2003 ona roadside next toa cornfield in association with Amaranthus palmeri, A. spinosus, Eragrostis pectinacea, and Senna obtusifolia. At that time, it had not been previously docu- mented from the state; but shortly thereafter, during the preparation of this manuscript, the Flora of North America Editorial Committee (2003) reported A. powellii from Tennessee. This is apparently the second report for the species from the state. Plant Communities Thirty qualitatively defined plant communities were sampled in the county. ESTES, FLORA OF TENNESSEE 2351 These were divided into four sections, those that occurred on the HR, those of the OCB, aquatic or riverine communities, and anthropogenically altered or maintained habitats. The 20 communities included for the HR and OCB and the three habitats listed for the Streams and Riverine Communities section were generally natural communities, although most have sustained some type of human-related disturbance. The seven types of Disturbed Sites sampled in- cluded areas that generally lacked natural qualities, appeared recently affected by anthropogenic disturbances, or were maintained as unnatural communities by human activities. HIGHLAND RIM 1. Oak-Hickory Forests (OH).—The oak-hickory community was the dominant upland forest type on dry to submesic sites on the HR, occupying broad upland flats, ridge-tops, and upper slopes of south and west-facing hillsides. This com- munity was mostly associated with the Mountview, Bodine, and Dickson series of the MFP BME and BFD soil associations. The canopy contained several spe- cies of oak including Quercus alba, Q. coccinea, Q. montana, Q. stellata, and Q. velutina as well asa few hickory species, including Carya alba, C. glabra, and C. ovata var. ovata. Less dominant but common species were Acer rubrum, Nyssa sylvatica, Oxydendrum arboreum,and Sassafras albidum. In the sul py and understory Amelanchier arborea, Cornus florida, Kalmia latifolia, Rhododen- dron canescens, Styrax grandifolia, Vaccinium arboreum, and V.stamineum were occasional to common constituents. Woody vines common to this community included Smilax glauca, S. rotundifolia, Toxicodendron radicans, Vitis aestivalis var. aestivalis,and V. rotundifolia. Subshrubs and low shrubs were Chimaphila maculata, Hypericum hypericoidessubsp. multicaule,and Vaccinium pallidum. The herbaceous layer was sparsely vegetated when compared to more mesic forests, but included Carex picta, Coreopsis major, Cunila origanoides, Danthonia spicata, Desmodium rotundifolium, Dichanthelium dichotomum var. dichotomum, Eurybia hemispherica, Solidago erecta, S. hispida, S. ulmifolia,and Viola hirsutula. 2. Rich Forests (RF).—Rich mesophytic forests were primarily found on lower to middle north and east facing slopes, in ravines, and in narrow stream valleys. Such sites were associated mostly with soils of the Bodine and Dellrose series of the BMF soil association. The canopy often contained Acer nigrum, A. saccharum, Aesculus flava (mostly EHR), Carya cordiformis, Fagus grandifolia, Fraxinus americana, Quercus alba, Q. rubra, Tilia americana, T. heterophylla, and Ulmus rubra. Woody vines of this community included Menispermum canadense, Parthenocissus quinquefolia, and Smilax tamnoides. Common un- derstory shrubs or small trees were Asimina triloba, Carpinus caroliniana, Dirca palustris, Hydrangea cinerea, Lindera benzoin, and Staphylea trifolia. In the her- baceous layer Actaea pachypoda, Adiantum pedatum, Anemone acutiloba, 2352 BRIT.ORG/SIDA 21(4) Botrychium virginianum, Cardamine concatenata, Carex cumberlandensis, C. albursina,C. kraliana, Diplazium pycnocarpon, Erythroniumamericanumsubsp. harperi, Hydrophyllum appendiculatum, Jeffersonia diphylla, Oxalis illinoensis, Pachysandra procumbens, Phacelia bipinnatifida, Phlox divaricata, Polemonium reptans, Trillium cuneatum, T.flexipes, T. stamineum, Valeriana pauciflora, and Viola pubescens var. scabriuscula were present, in addition to many others. 3. Flatwoods (FW).—This community occurred only in extreme southeast- ern Giles County on the EHR along Piney Creek in association with soils of the Guthrie and Taft series of the MFP soil association. The site was level and the soil saturated, often for much of the year. Acer rubrum, Nyssa biflora, N. sylvatica, Quercus nigra, and Q. phellos dominated the canopy while Q. lyrata and Pinus taeda were occasional associates. Common small trees and shrubs were Arundinaria gigantea, Cornus foemina, Crataegus marshallii, Hypericum crux- andreae, H. hypericoides subsp. hypericoides, Ilex verticillata, Photinia pyrifolia, Rhododendron canescens, Styrax americana, Vaccinium fuscatum, Viburnum dentatum var. lucidum, and V. nudum var. nudum. Noteworthy herbaceous spe- cies of this community included Arisaema triphyllum subsp. pusillum, Carex flaccosperma, C. gigantea, C. intumescens, C. joorii, Chasmanthium laxum, Galium obtusum, Gratiola neglecta, G. virginiana, Isotria verticillata,Osmunda cinnamomea, O. regalis, Platanthera flava var. flava, and Triadenum virginicum. Clearings and wet meadows within the flatwoods community supported a number of herbaceous species that were not found elsewhere in the county. Some of these species are more common on the southeastern Coastal Plain. Examples include Agalinis purpurea, Eryngium prostratum, Gratiola pilosa, Ludwigia linearis, Mikania scandens, Mitreola petiolata, Rhexia mariana var. interior, R. mariana var. mariana, R. virginica,and Spiranthes vernalis. 4. Bluffs and Rock Outcrops (RO).—Outcroppings of limestone and less fre- quently shale generally were located on the slopes of hills or often in narrow ravines or along watercourses, mostly in the strongly dissected portion of the HR. Such outcrops were mostly associated with Rockland of the BMF soil asso- ciation. The rock outcrops were almost exclusively located in or closely sur- rounded by forested areas. In some places, sizeable bluffs also occurred. Two types of bluff or rock outcrop communities of the HR were identified: those that were relatively moist and those that were dry to xeric. The more mesic bluffs and rock outcrops were located mostly on north or east facing exposures while the drier ones were typically associated with south or west facing aspects. Shrubs or woody vines associated with moist sites included Decumaria barbara, Hy- drangea cinerea, and Philadelphus hirsutus. Herbaceous species common to moist bluffs were Aquilegia canadensis, Arabis laevigata, Asplenium rhizophyllum, Cystopteris bulbifera, C. tennesseensis, Galium triflorum, Heuchera villosa, Parietaria pensylvanica, Saxifraga virginiensis, Sedum ternatum, Solidago caesia, and Thalictrum dioicum. On drier sites, some of the above species also were — ESTES, FLORA OF TENNESSEE 2353 present but notable additions included And i, Asclepias verticillata, Cheilanthes lanosa, Oxalis violacea, Packera obovata, Pellaea atropurpurea, Pleopeltis polypodioides subsp. michauxiana, Polymnia canadensis, Sedum Solidago sphacelata, and Woodsia obtusa 5. Barrens (RB).—Portions of western Giles County lie within the Barrens of the southwestern Highland Rim described by Shanks (1958). Barrens were mostly encountered on soils of the Mountview and Dickson series of the MFP and BMF soil associations. The county’s highest quality barrens were found on the undissected portion of the HR in the southwestern quarter of the county but other good examples occurred on thin ridges and even in some valleys throughout the western edge of the county. They occurred mostly at the mar- gins of oak-hickory forest, often along roadsides, and were apparently princi- pally maintained as open areas by periodic mowing. Without periodic distur- bance the barrens likely would be succeeded by oak-hickory forest. Although the barrens themselves lacked trees for the most part, species that occasionally intruded from surrounding oak-hickory forest included Oxydendrum aor, Pinus taeda, Quercus coccinea, Q. marilandica, Q. stellata, and Sassa- Small trees and shrubs of the barrens were Ceanothus americanus, alas angustifolia, Rhus copallinum, Rubus flagellaris, Salix humilis var. humilis, Vaccinium arboreum, and V. stamineum. Perennial grasses and mem- bers of the Asteraceae and Fabaceae dominated the barrens. The most impor- tant grasses were Andropogon gerardii, A.gyrans, A. virginicus, Dichanthelium spp., Panicum anceps, Saccharum alopecuroidum, Schizachyrium scoparium var. divergens, S.scoparium var. scoparium, and Tridens flavus var. flavus. Other her- baceous species were Ambrosia bidentata, Angelica venenosa, Asclepias amplexicaulis, Aureolaria pectinata, Chamaecrista fasciculata, C. nictitans, Conyza densis var. pusilla, Coreopsis tripteris, Desmodium spp., Eupatorium spp. Euphorbia corollata, Eurybia hemispherica, Galium pilosum, Helianthus spp. (including H. eggertii), Hypericum drummondii, H. gentianoides, Lespedeza spp., Liatris spicata, L. squarrulosa, Parthenium integrifolium, Pteridium aquilinum var. latiusculum, Silphium mohrii, Solidago spp., Spiranthes lacera var. gracilis, Symphyotrichum spp, Veronicastrum virginicum, and Viola sagittata. 6. Marshes (RM).—Marshes were more or less open wetland areas with standing water and were dominated by herbaceous vegetation and shrubs. Those surveyed occurred at the headwaters of small streams and in natural upland depressions. This community was restricted to soils of the Guthrie, Taft, and Lee series of the MFP soil association. Some were created by beaver activ- ity; others were created by human actions such as stream damming and road construction. Trees and large shrubs were mostly restricted to their margins. Important large tree species were Acer rubrum, Liquidambar styraciflua, Nyssa sylvatica, Quercus phellos, Salix nigra,and rarely Betula nigra. Shrubs included Cephalanthus occidentalis, Itea virginica, Photinia pyrifolia, Rhododendron pulchellum, 2354 BRIT.ORG/SIDA 21(4) canescens, Vaccinium corymbosum, V.fuscatum, and Viburnum nudum vat. nu- dum. Noteworthy emergent or marginal herbaceous species were Carex crinita var. brevicrinis, C joorii,C. lupuliformis, Hibiscus moscheutossubsp. moscheutos, Juncus effusus, J. repens, Osmunda cinnamomea, O. regalis, Panicum rigidulum var. pubescens, Platanthera ciliaris, Proserpinaca palustris, Rhexia mariana var. interior, Rhynchospora glomerata, Saccharum baldwinii, Scirpus cyperinus, Sparganium americanum, Typha latifolia, and Viola x primulifolia. Few float- ing or submerged aquatics were observed; the most abundant were Lemna mi- nor, Potomogeton diversifolius, and Spirodela polyrrhiza 7. Wet Meadows (WM).—Wet meadows were frequently encountered in valleys along small streams or in upland swales. They occurred on soils of the Lobelville, Lee, and Lynnville series associated with the MFP, BME and BED soil associations. These sites appeared to be maintained as open areas by periodic grazing or mowing thereby preventing the establishment of large shrubs and trees. The most common shrubs found in the wet meadows included Alnus serrulata, Amorpha fruticosa, Cephalanthus occidentalis, Cornus amomum, and Salix sericea. Common herbaceous taxa were Acorus calamus, Alisma sub- cordatum, Apios americana, Asclepias incarnata subsp. incarnata, Bidens aristosa, B. cernua, Boehmeria cylindrica, Carex frankii, C. vulpinoidea, Commelina virginica, Dichanthelium dichotomum var. ramulosum, D. scoparium, Eupato- rium fistulosum, E. perfoliatum, Hibiscus moscheutos subsp. moscheutos, Hymenocallis occidentalis, Impatiens capensis, Juncus coriaceus, Lobelia cardinalis, L. siphilitica, Ludwigia alternifolia, Mentha x piperita, Mimulus alatus, Panicum rigidulum var. rigidulum, Polygonum hydropiperoides, P sagittatum, Rhexia virginica, and Solidago gigantea. At one site on an outlier of the HR in northeastern Giles County, a wet meadow was found that contained three species that are uncommon in Middle Tennessee: Panicum verrucosum, Paspalum fluitans, and Trachelospermum difforme. 8. Acidic Seeps (AS).—This community type was restricted to forested ra- vine bottoms in extreme northwestern Giles County in the upper sections of small HR streams. The soils of acidic seeps are part of the Lee series, a member of the BMF soil association. The surrounding forest type was predominantly oak-hickory. Some species common to the oak-hickory community were present among the seeps but were mostly restricted to dry hummocks. Such taxa in- cluded Acer rubrum, Liquidambar styraciflua, Nyssa sylvatica, Oxydendrum arboreum, and Quercus alba. Shrubs common to this community were Alnus serrulata, Cephalanthus occidentalis, Euonymus americana, Itea virginica, Photinia pyrifolia, Rhododendron alabamense, R. canescens, and Vaccinium fuscatum. Important herbaceous taxa were Bartonia virginica, Carex atlantica subsp. capillacea, C. intumescens, Osmunda cinnamomea, O regalis, Phlox glaber- rima, Platanthera clavellata, and Thelypteris novaboracensis. An unidentified ESTES, FLORA OF TENNESSEE 2355 species of Sphagnum, a non-vascular plant, was one of the dominant ground cover species in this community. 9. Calcareous Seeps (CS).—This type of wetland community occurred along the western edge of Giles County on lower slopes of ravines and hollows along small streams, by waterfalls, or above wet bluffs. The seeps o urred over lime- stone, had a thin, gravelly, continuously waterlogged substrate, and were level to slightly sloping. This community was restricted to soils of the BMF associa- tion, specifically the Rockland type or Lynnville and Lobelville soil series. Most were found among forests but the seeps themselves usually lacked large trees. Shrubs commonly associated with the seeps were Alnus serrulataand Lindera benzoin. The woody vine Decumaria barbara was often associated with the seeps in the southwestern corner of the county. Herbaceous species found were Cardamine pensylvanica, Carex leptalea, Chelone glabra, Cuscuta compacta, Dichanthelium dichotomum var. ramulosum, Dryopteris celsa, Equisetum hyemale var. affine, Glyceria striata, Impatiens capensis, Juncus coriaceus, Lobe- lia cardinalis, Oxypolis rigidior, Samolus valerandi subsp. parviflorus, and Sol- idago patula. 10. Riparian Areas (RIP).—T his community included those areas that were located along riparian zones in the HR and was found in conjunction with a variety of series belonging to the BMF soil association. Many species were found in this broad community type although few species were restricted to it. Trees included a few species of Quercus and Acer as well as Carpinus caroliniana, Juglans cinerea (rarely), J. nigra, Liquidambar styraciflua, Morus rubra, an Platanus occidentalis. Smaller trees and large shrubs were Alnus serrulata, Arundinaria gigantea, Cornus amomum, Corylus americana, Crataegus calpo- dendron, Hamamelis virginiana, Lindera benzoin, Salix caroliniana, and S. sericea. Many species of wildflowers inhabited this community, including most of the species listed for the wet meadow community above. Additional note- worthy herbaceous taxa were Elymus macgregorii, Equisetum hyemale var. af- fine, and Mertensia virginica. OUTER CENTRAL BASIN 1. Rich Forests (BRF).—In the OCB, rich woodlands occurred most often on middle to upper, north and east facing slopes, along river bluffs, or in sheltered hollows associated with soils of the Bodine, Dellrose, and Culleoka series of the MEP. BME and BED soil associations. Unfortunately, much of this forest com- munity was heavily logged or grazed in the past. The dominant canopy species of the Basin’s rich woodlands included Acer nigrum, A. saccharum, Aesculus flava, Carya cordiformis, Carya ovata var. ovata, Fagus grandifolia, Fraxinus americana, Juglans nigra, Liriodendron tulipifera, Quercus muehlenbergii, Q. rubra, Prunus serotina, Tilia americana var. americana, and I. americana vat. heterophylla. A rare component of some of the forests was Juglans cinerea. 2356 BRIT.ORG/SIDA 21(4) Smaller woody species that were sometimes present included Aesculus pavia, Asimina triloba, Carpi nus caroliniana, Gleditsea triacanthos, Morus rubra,and Ulmus rubra. Euonymus atropurpurea, E. americana, Hydrangea cinerea, Lindera benzoin,and Staphylea trifolia were frequent shrubs. Notable herbaceous taxa were Actaea pachypoda, Arisaema triphyllumsubsp. triphyllum, Botrychium virginianum, Carex blanda, C. kraliana, Claytonia virginica, Cynoglossum virginianum, Delphinium tricorne, Impatiens pallida, Jeffersonia diphylla, Osmorhiza claytonii, Panax quinquefolius, Phacelia bipinnatifida, Podophyllum peltatum, Sanguinaria canadensis, Thalictrum thalictroides, and Trillium cuneatum. 2. Dry Forests (BDF).—Dry forests occurred along ridge-tops and on middle to upper, south or west facing and occasionally east facing slopes. This commu- nity was most frequently associated with Rockland soils of the DBM and SAM soil associations. These forests, like the remainder of forested land in the county, have been heavily logged and grazed. They contained Acer saccharum, Aesculus glabra, Carya glabra, C. ovata var. ovata, Celtis laevigata, Fraxinus americana, F quadrangulata, Juniperus virginiana, Quercus alba, Q. muehlenbergii, Q. shumardii, Robinia pseudoacacia, Ulmus alata, and U. serotina. Common small trees and shrubs were Cornus florida, Crataegus crus-galli, C. intricata, Forestiera ligustrina, Frangula caroliniana, Ostrya virginiana, Prunus americana, Rhus aromatica, Rosa setigera, Symphoricarposorbiculatus,and Viburnum rufidulum. Frequent woody vines included Bignonia capreolata, Cocculus carolinus, and Smilax bona-nox. Agrimonia pubescens, Anemone virginiana, ania platyneuron, Camassia scilloides, Carex albicans var. albicans, C. cep| hal; Chasmanthium sessiliflorum, Elymus villosus, Heliotropium tuberosum, a inflata, Melica mutica, Nothoscordum bivalve, Scutellaria ovata, and Viola palmata were the common herbaceous species. 3. Limestone Karst Woods (LKW).—This ity type was restricted to asmall area in the central portion of the county. These forests occurred on small rocky knobs over Ordovician-aged Lebanon or Ridley limestone where soil was sufficiently deep to support forests. Limestone karst woods were associated with Rockland soils of the SAM soil association. The rock outcrops in these commu- nities were often massive (1-4 m tall) and covered several ha. Common tree spe- cies of the karst woods included Celtis laevigata, Diospyros virginiana, Fraxinus americana, F. quadrangulata, Juniperus virginiana, Quercus muehlenbergii, Q. shumardii, Tilia heterophylla, Ulmus alata, and U. serotina. In the subcanopy and shrub layer, Callicarpa americana, Cercis canadensis, Forestiera ligustrina, Frangula caroliniensis, Rhamnus lanceolata, Rhus aromatica, Sideroxylon lycioides, Staphylea trifolia, Symphoricarpos orbiculatus, and Viburnum rufidulum were found. Frequent herbs were Cynoglossum virginianum, Del- phinium tricorne, Erythronium albidum, Frasera caroliniensis, Nemophila aphylla, Osmorhiza claytonii, Packera obovata, Polygonatum biflorum, Trillium ESTES, FLORA OF | TENNESSEE 2357 cuneatum, T. sessile, and Vicia minutiflora. On the rock outcrops in the forest, species such as Aquilegia canadensis, Arabis laevigata, Asplenium resiliens, A. rhizophyllum, A. ruta-muraria, Cystopteris bulbifera, Heuchera villosa, Pellaea atropurpurea, Perideridia americana, Pleopeltis polypodioides subsp. michauxiana, Polymnia canadensis, Ranunculus micranthus, Saxifraga virginiensis,and Woodsia obtusa were present. 4. Cedar Barrens (CB).—This community type was represented by only a few degraded examples that were restricted to the slopes of small knobs over Ordovician limestone in central Giles County. Cedar barrens were found on Rockland soils of the SAM association. They occurred on open slopes sur- rounded by rocky limestone karst woods within close proximity to cedar glades. Juniperus virginiana was abundant near the margins of the barrens and as iso- lated individuals among them. Most of the woody species were the same as those listed for the glades with a few exceptions such as Quercus stellata and Vaccinium arboreum. They were dominated by a variety of perennial grasses and herbs. Noteworthy herbaceous taxa were Allium canadense var. canadense, Andropogon virginicus, Aristida oligantha, Asclepias viridiflora, Carex cherokeensis, Chasmanthium sessiliflorum, Croton capitatus, Cuphea viscosissima, Daucus carota, Dichanthelium malacophyllum, Eupatorium altissimum, E. hyssopifolium, Euphorbia corollata, Festuca arundinacea, Geranium carolini- anum, Lespedeza procumbens, Lobelia spicata, Onosmodium molle subsp. molle, Panicum flexile, Potentilla recta, P. simplex (plants in this community may be referable to the variety argyrisma Fern.), Rudbechia triloba, Sabatia angularis, Silphium trifoliatum var. latifolium, Sporobolus clandestinus, and Tridens flavus var. flavus. 5. Cedar Glades (CG).—The limestone glades of Giles County were dis- tributed mostly along a 16 km stretch of a 32 km section of a fault line that runs diagonally from NW to SE across the southern half of the county. Along this line, they were mostly along the western edge and to the west of Richland Creek. Like most other glades of Middle Tennessee, these were associated with Ordovician-aged Lebanon and Ridley limestone of the Stones River Group (Miller et al. 1966). They occurred on level to slightly sloping sites with shallow gravelly soil or exposed limestone. Cedar glades were mostly associated with soils of the Rockland type, part of the SAM association. Such sites often were wet in the winter and spring and were dry during summer and autumn. The woody species commonly associated with glade margins included Celtis laevigata, Carya ovata var. australis, Forestiera ligustrina, Frangula caroliniana Juniperus virginiana, Quercus muehlenbergii, Rhus aromatica, Rosa carolina, R. setigera, and Sideroxylon lycioides. In the open portion of the glades, common herbaceous species included Amphiachyris dracunculoides, Astranthium integrifolium, Carex cherokeensis, Cyperus squarrosus, Dichanthelium malaco- phyllum, Erigeron strigosus var. calcicola, Geranium molle, Glandularia 2358 BRIT.ORG/SIDA 21(4) canadensis, Grindelia lanceolata, Heliotropium tenellum, Houstonia purpurea var. calycosa, Hypericum sphaerocarpum, Hypoxis hirsuta, Isanthus brachiatus, Leavenworthia torulosa, Malvastrum hispidum, Manfreda virginica, Minuartia patula, Nothoscordum bivalve, Oenothera triloba,O lium molle subsp. molle, Oxalis priceae subsp. priceae, Opuntia humifusa, Packera anonyma, Panicum flexile, Penstemon tenuiflorus, Phemeranthuscalcaricus, Ranunculus fascicularis, Rudbeckia triloba, Ruellia humilus, Salvia lyrata, Scutellaria parvula, Sedum pulchellum, Sisyrinchium albidum, Symphyotrichum priceae, Verbena simplex, and Viola egglestonii. 6. Bluffs and Outcrops (BO).—Other than cedar glades and man-made road- cut bluffs, rock outcrops and bluffs were generally uncommon in the OCB. Some small bluffs were located near caves or springs. Larger bluffs were mostly con- fined to watercourses, particularly the larger streams suchas the Elk River and lower section of Richland Creek. The soils of this community are part of the Rockland type of the DBM and SAM soil associations. Woody species associ- ated with bluffs included Fraxinus quadrangulata, Hypericum frondosum, Juniperus virginiana, Quercus muehlenbergii, and Philadelphus pubescens var. pubescens. Common herbaceous taxa encountered were Asplenium resiliens, Cheilanthes alabamensis, C. lanosa, Cystopteris bulbifera, Heuchera villosa, Mateled gonocarpos, Pellaea atropurpurea,and Sedum pulchellum. 7. Marshes (BM).—Marshes were rare in the OCB, mostly restricted to the flood plains of large streams. This community was restricted to soils of the Lynnville and Newark series of the SAM soil association. All of those sampled were formed either by man or by beaver activity. Common trees included Fraxinus pennsylvanica, Platanus occidentalis, and Salix nigra with Cephalanthus occidentalis the common shrub. Emergent aquatics were repre- sented by Leersia oryzoides, Scirpus cyperinus,and Typha latifolia. Floating and submerged aquatics included Heteranthera reniformis, Lemna minor, Potomogeton diversifolius, P. foliosus, Spirodela polyrrhiza, and rarely Azolla caroliniana. 8. Wet Meadows (BWM).— Wet meadows occurred mostly in the flood plains of larger streams and were associated with the Newark, Lynnville, and Armour series of the DBM and SAM soil associations. This community type was domi- nated by various herbaceous wetland species. Representative taxa of this com- munity included Alopecurus carolinianus, Ammannia coccinea, Asclepias incarnata subsp. incarnata, Bidens cernua, B. frondosa, Carex frankii, C. vulpinoidea, Cyperus strigosus, Echinochloa muricata, Gratiola neglecta, Iva annua, Juncus acuminatus, J. effusus, Leersia oryzoides, Lobelia siphilitica, Ludwigia glandulosa, L. palustris, Lycopus rubellus, L. virginicus, Panicum dichotomiflorum, Polygonum hydropiperoides, Rorippa sessiliflora, Rotala ramosior, Rumex altissimus, and Sagittaria australis. 9. Limestone Wet Meadows (LWM).—This community type was found in ea ESTES, FLORA OF GILES COUNTY, TENNESSEE 2359 low areas within or near limestone cedar glades or along roadside ditches through gladey areas where the ground was thoroughly saturated for lengthy periods in spring. The soils of this ity are part of the Talbott and Roellen series of the SAM soil association. Some are also associated with the Rockland type. The major difference between the Limestone Wet Meadow Community and other wet meadows of the OCB is that the limestone wet meadows are thin- soiled depressions over limestone bedrock. Characteristic species of this plant community included Carex festucacea, C. granularis, Cuphea viscosissima, Eleocharis bifida, E. obtusa, Hypericum sphaerocarpum, Isoetes butleri, Juncus filipendulus, Leavenworthia torulosa, Mecardonia acuminata, Nothoscordum bivalve, Scirpus atrovirens, and S. lineatus. 10. Riparian Areas (BRIP).—This ity type occurred along the flood plains of medium to large streams in association with soil of the Staser, Lynnville, and Armour series of the SAM soil association. Tree species common tothisco y were Acer negundo, A. saccharinum, Celtis laevigata, Praxinus pennsylvanica, Gleditsea triacanthos, Populus deltoide i, Sali nigra, Ulmus americana, and rarely Penile nigra. Asimina triloba, Cornus drummondii, and Staphylea trifolia were occasional to common understory spe- cies. At several sites, Arundinaria gigantea was the dominant understory spe- cies. Aristolochia tomentosa,a high-climbing woody vine, was restricted to this community type. Herbaceous species found in this community included Ageratina altissima, Alliaria petiolata, Chasmanthium latifolium, Elymus macgregorii, E. virginicus, Laportea canadensis, Microstegium vimineum, Phlox paniculata, Silphium perfoliatum, and Verbesina alternifolia. Quere STREAMS AND RIVERINE COMMUNITIES 1. Small Streams and Springs (ST).—Small streams and springs were common throughout Giles County. Species commonly found in this community included Lemna minor, Mentha x piperita, Rorippa nasturtium-aquaticum, Spirodela polyrrhiza, and Veronica anagallis-aquatica. 2. Large Streams and Rivers (RIV).—The two largest streams in Giles County are the Elk River and Richland Creek. These two streams are approximately 25-75 m wide in most places and are characterized by having a slow to moder- ate current with more or less shallow waters. The substrate is mostly composed of large rocks, gravel, and silt. Aquatic species were generally rare and were con- fined to the shallow and often swift-flowing riffles or slow-moving backwater areas. Representative taxa found in this community included Heteranthera dubia, Justicia americana, Podostemum ceratophyllum, and Potomogeton nodosus. 3. Gravel Bars (GB).—Large gravel bars were frequent along the Elk River and Richland Creek. Large woody species generally were not abundant due to perioidic flooding. Woody species that were common in this community in- cluded saplings of Betula nigra, Platanus occidentalis, Populus deltoides, Salix caroliniana,and S. nigra. The common herbaceous taxa were Cuscuta sp., Diodia 2360 BRIT.ORG/SIDA 21 = 4) virginiana, Dysphania ambrosioides, Eclipta prostrata, Justicia americana, Polygonum spp., and Samolus valerandi subsp. parviflorus. DISTURBED HABITATS A number of anthropogenically modified or maintained communities were sampled during the survey. These areas were found in all three physiographic sections and often were surrounded by some of the more natural habitats dis- cussed above. The most common of the disturbed sites sampled included farm ponds and man-made lakes (FP); lawns, old homesites, cemeteries, etc. (LAW): pastures and fields (PAS); crop fields (CRP); roadsides and railroad tracks (RD); abandoned lots and bare ground (DIS), and urban thickets (UT). These sites appeared to receive higher amounts of disturbance than other communities and contained higher numbers of non-native and weedy species compared to more natural communities ANNOTATED CHECKLIST Nomenclature follows the PLANTS database (USDA NRCS 2000) except for the genus Crataegus, which follows Wolford and Chester (2002); newly described species (Campbell 2000, Allison & Stevens 2001, Smith 2001); and those plants covered in published volumes of Flora of North America (1993, 1997, 2000, 2002a, 2002b, 2003), with the exception of Carya glabra and C. ovalis and in- fraspecific taxa within Lindera. Juncus biflorusand J. marginatus are maintained here as distinct species following Radford et al. (1968). The arrangement of the checklist follows that of Wofford and Kral (1993), whereby the taxa are divided into three major groups: PTERIDOPHYTES, GYMNOSPERMS, and AN- GIOSPERMS (MONOCOTS, DICOTS). Families and genera are arranged alpha- betically within the three major divisions. An asterisk (*) precedes the scientific name of each taxon not native to Tennesseee (sensu Wofford & Kral 1993). Two asterisks (**) signify taxa that are native to portions of Tennessee but not to Giles County. Entries denoted by the symbol “@” represent recently described taxa (for recently described taxa, a lit- erature citation is provided in brackets following all other information). A cross (+) indicates taxa collected from Giles County prior to this study (mostly based on Chester et al. 1993, 1997): those without a cross represent county records. Taxa preceded by “A” were observed but not collected. Rare taxa are listed in bold type and are enumerated in Table 2. Following the scientific name and authority, an abbreviation is listed for the relative abundance of each taxon within the county. An abbreviation for the physiographic area(s) where the taxon was collected or observed and an abbreviation for the habitat are listed next. The abbreviations used in the check- list to represent relative abundance, physiographic province, and habitat types are listed in Table 3. ESTES, FLORA OF , TENNESSEE 2361 Taste 3. Abbreviations used in the checklist of vascular plants known from Giles County, Tennessee. Symbols Used in the Checklist * = Non-native species @ = Recently described taxon A = Taxon observed but not collected Relative Abundance V =Very rare O = Occasional R = Rare F = Frequent S = Scarce C = Common | = Infrequent X = Extirpated Physi phic Area OCB = Central Basin HR = Highland Rim (both EHR & WHR) EHR = Eastern Highland Rim WHR = Western Highland Rim Plant Communities AS = Acidic Seep BDF = Central Basin Dry Forest BRF = Central Basin Rich Forest BM = Central Basin Mars BO = Central Basin Bluff/Outcrop BRIP = Central Basin Riparian Area BWM = Central Basin Wet Meadow FP = Farm Pond/Lake GB = Riverine Gravel Bar LAW = Old Homesite/Lawn LKW = Limestone Karst Wood LWM = Limestone Wet Meadow OH = Oak-Hickory Forest PAS = Pasture/Field RB = Highland Rim Barren RD = Roadside/Railroad RO = Highland Rim Bluff/Outcrop RIP = Highland Rim Riparian Area RF = Highland Rim Rich Forest WM = Highland Rim Wet Meadow ST = Small coe UT = Urban Thick The classification used to categorize the relative abundance of each taxon follows that of Murrell and Wofford (1987) and is as follows: very rare, found in a single locale with few individuals; rare, one or two localities, generally small populations; scarce, several localities, or scattered small populations; infrequent, scattered localities throughout; occasional, well distributed but not abundant anywhere; frequent, generally encountered; and common, characteristic and dominant. Taxa not observed over the last 20 years are listed as extirpated. When taxa were found in all three physiographic areas (Outer Central Ba- sin, Eastern Highland Rim, Western Highland Rim) the word ALL is listed. For taxa collected from both the Eastern and Western Highland Rims the designa- tion HR is given, otherwise each physiographic area is listed separately. Addi- tional habitat information is listed where necessary. For each entry, the physi- ographic and habitat designations are listed in their order of abundance, starting with the most abundant. 2362 BRIT.ORG/SIDA 21(4) After the information concerning relative abundance, physiographic prov- ince, and habitat, the author's collection number is given in parenthesis. For specimens collected by the author housed at TENN only the author's collection number is given (e.g. 01077); however, for specimens housed in herbaria other than TENN, both the author's collection number and herbarium acronym where the specimen is deposited are given (e.g.01349, VDB). For taxa listed in the check- list that were collected by others, the collector's name, collection number, and the herbarium acronym where the specimen is housed are included (e.g. R. Kral 39577, VDB) PTERIDOPHYTES ASPLENIACEA tAsplenium playreon ) - 7 P—F; ALL; BDF, BRF, FW, LKW, OH, RF; (02 ieclentom a lens oe OCB, WHR; LKW, BO, RO; (00339). tAsplenium rhizophyllum L—O;ALL;BO, RO, LKW, RF, BRF; (02825) tAsplenium ruta-muraria L.—R; OCB; LKW, BO; (04452). AZOLLACEAE Azolla caroliniana Willd.—R; OCB; BM; (03521). BLECHNACEAE Woodwardia areolata (L.) T. Moore—R; HR; FW,AS; (00031). DENNSTAEDTIACEAE Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underwood ex Heller—O; HR; RB, OH; (02142) DRYOPTERIDACEAE tAthyrium filix-femina (L.) Mertens. var. asplenioides (Michx.) Farw.—O; HR; RF; (00914). tCystopteris bulbifera (L.) Bernh.—O; ALL; LKW, BO, RO; (06784 tCystopteris protrusa ea Blasdell—l; OCB, WHR; BRF, RF; (0219 tCystopteris tennesseensis se WHR, OCB; O; (J. Shaver 10220, TENN). A Beparia acrostichoides (Swartz) M. Kato—V; HR; RF. Diplazium pycnocarpon (Spreng.) M. Broun—t; WHR, OCB; RF, BRF; (01537). ee aie celsa (W. Palmer) Knowlt., Palmer, & Pollard—V; WHR; CS, WM; (02127). Dryopteris marginalis (L.) Gray—V; WHR; RF; (00984) +Onoclea sensibilis L.—S; HR; CS, FW; (03763). tPolystichum acrostichoides (Michx.) Schott—C; ALL; RF, BRF, FW; (00970). +Woodsia esr (Spreng.) Torr. subsp. obtusa— OQ} R: LKW, BDF, CG, BO, RO; (00666) EQUISETACEAE Equisetum hyemale L. var.affine (Engelm.) Calder & R.L. Tayor—R; WHR; CS, RIP; (00129). ISOETACEAE /soetes butler’ Engelm.—V; OCB; LWM; (03043). LYCOPODIACEAE Diphasiastrum digitatum (Dillenius ex A. Braun) Holub—R; WHR; OH, RD; (01607) OPHIOGLOSSACEAE Botrychium biternatum (Sav.) Underwood—S; HR; FW, RB; (05030). Botrychium dissectum Spreng.—R; WHR; RF; tBotrychium virginianum (L.) Sw.—F; ALL; RF, BRF, KW; (01936). tOphioglossum engelmannii PrantlL—R; OCB; CG; (01954). ma re vulgatum L.—R; EHR, OCB; FW, W; (00605). OSMUNDACEAE ap ie cinnamomea L.—|; HR; RM, FW,AS, ST; 05). ae regalis L. var. soe Gray—I; HR; RM, FW, AS, ST: (00588). POLYPODIACEAE tPleopeltis polypodioides (L.) Andrews & Windham subsp. michauxiana (Weatherby) Andrews & Windham— HR; LKW, BDF, BO, RO, on trees and rocks; (01783). (Willd.) PTERIDACEAE tAdiantum pedatum L.—1; ALL; RF, BRF; (02187). ESTES, FLUKA UF , TENNESSEE +Cheilanthes alabamensis (Buckl.) Kunze—S; OCB; BO, LKW; (00652). re lanosa (Michx.) D. C. Eat.—R; OCB, HR; BO, RO; (02757). ug aea sat a Link—O; OCB, WHR; LKW, BO, RO, CG; (00663). SELAGINELLACEAE + Selaginella apoda (L.) Spring—V;WHR; RM; (A. J. Sharp, E. Clebsch, & A.Clebsch 9814, TENN). THELYPTERIDACEAE tPhegopteris hexagonaptera (Michx.) Fee—|; ALL; RF, BRF; (04931) tThelypteris noveboracensis (L.) Nieuwl.—S; HR; AS, F W; (00916) a= GYMNOSPERMS CUPRESSACEAE Juniperus virginiana L.var. virginiana—F; ALL; CG W, BDF, RDE,RB, RD, PAS, BO, RO; (01616). PINACEAE ** Pinus echinata P. Mill —R; WHR,OCB ipiohably planted); OH, RB, RD, few if ar ly native stands; (03150) **Pinus strobus L_—V; WHR, tree plantation with seedlings; (02140) Pinus taeda L.—S; ALL; FW, OH, RB, native only in extreme southern part of county; (0036/7). ** Pinus virginiana P. Mill.—R; WHR; RD, no native Vy fant 21) stands observed; (01631). ANGIOSPERMS: MONOCOTS ACORACEAE *Acorus calamus L.—R; WHR, OCB; WM, BWM; (03772). AGAVACEAE Manfreda virginica (L.) Salisb. ex Rose—S; OCB; CG, CB, BO; (00821). ** Yucca filamentosa L.—R; OCB; RD, LAW, doubt- fully native to county; (04954). ALISMATACEAE Alisma subcordatum Raf.—S; HR; WM, FW; (00831). Sagittaria australis (J. G. Smith) Small—R; WHR, OCB; AS, BWM; (05426). Sagittaria montevidensis Chamisso & Schlechtendal subsp. calycina (Engelm.) Bogin—S; WHR; FP, WM; (01333). —_ 2363 ARACEAE Arisaema dracontium (L.) Schott—R; OCB; LKW; (02115 ). Arisaema triphyllum (L.) Schott subsp. pusillum (Peck) Huttleston—R; EHR; FW; (00553). tArisaema triphyllum (L.) hott subsp. triphyllum—F; ALL; RF, BRF, LKW; (01743). COMMELINACEAE *Commelina communis L.—O; ALL; BRIP, BWM, CRP. GB, RD, RIP; (03745). *Commelina diffusa Burm. f—R; OCB; LWM, RD; (02715) Commelina erecta L. var. erecta—R; OCB; CG, BO; (06427). tCommelina virginica L.—S; ALL; WM, BWM; (03864) tTradescantia subaspera Ker Gawler—S; WHR, OCB; RF, BRF, BRIP, BO; (00980) Tradescantia virginiana L.—S; WHR; RF, OH; (01920) CYPERACEAE Bulbostylis eae illaris (L.) C. B. Clarke ex J.D. Hoo R; RB, wet ruts; (05289). +Carex iibicans Willd. ex armen var. albicans— O; ALL; RF, BRF, LKW, OH; (04399). Ibol Schwein.—R;WHR; RM, WM; Carex (03276). Carex albursina E. Sheldon—R; HR; RF; (04401). +Carex amphibola Steud.—|; OCB, WHR; BRF, RF; (05914) = Carex atlantica Bailey subsp. capillacea (Bailey Rezn aly AS; (00424 VDB). Car udel.—V; OCB; BM; (03522). ae baileyi Britt. ay WHR; RM; (00036 VDB). Carex basiantha Steudel—R; OCB; LKW; (05899). Carex blanda Dewey—S; OCB; BDF, BRF, PAS, CG; (04a 6). Carex cephalophora Willd. ex Willd. —S; WHR; RF; (00614). ae eee Schwein.—S; OCB; CG, CB, W; (02039). ae eee Torr. & Hook.—R; WHR; RB; (04905). Carex crinita =e var. brevicrinis Fern.—R; WHR, ' 53); Carex aes Naczi, Kral, & Bryson.—l; ;RF; (05910). = Carex oe is Michx. var. debilis—S; HR; AS, FW; (00428 VDB). 2364 Carex digitalis Willd.—R; WHR; AS; (02265). Care nine Schkuhr ex Willd —l; OCB, WHR, WM, RM; (03243). gee Dewey—R; EHR; FW; We0ee): +Carex frankii Kunth—O; ALL; BWM, RM, BM, RM (00668). tCarex gigantea ge Carex mranilan Muhl. ex Willd.—R; a on < Ay Fa sa | oe V, CAIN, Tt ' : 03249). tCarex intumescens Rudge—S; HR; AS, WM, FW; — (00967). Carex jamesii Schweinitz—R; OCB; BRF; (05834). Carex joorli Bailey—S; HR; FW, WM, RM; (01322 VDB). Carex kraliana Naczi & Bryson—R; OCB; BDF; (0065 4). tCarex laevivaginata (Kukenth.) Mackenzie—R; WHR; CS; (02 Carex leavenworthii Dewey—S; OCB; BDF; (00649) Carex leptalea Wahlenb —R:WH R; CS: (02261 ). Carex lupuliformis Sartwell ex Dewey—R; WHR M; (00786). Carex lupulina Willd.—S; HR; WM, FW; (04206). Carex lurida Wahlenb.—S; HR; AS, FW; (00953). Carex muehlenbergii Schkuhr ex a var.enervis Boott—R; OCB; PAS, CG; (05 Carex normalis eee. Carex picta Steud.—l; WHR; OH; (01613). Carex pianispicata neces WHR; RF; (05972). Carex prasina Wahlenb,—R; WHR; AS, RF; (00614). Carex radiata (Wahlenb) Small.—R; WHR; RF; 971). Carex rosea Schkuhr ex Willd.—!; WHR, OCB; RF BRF, AS; (03680) tCarex scoparia Schkuhr ex Willd.—X;WHR, OCB; WM; (K. Rogers 34914, TENN). tCarex stipata Muhl.ex Willd.—R; WHR, OCB;WM, WM; (K. Rogers 34913, TENN). Carex stricta Lam.—R; WHR; AS; (02260). Carex styloflexa Buckl.—R; WHR; AS; (00427 VDB). Carex swanii (Fern.) Mackenzie—V; WHR; WM; Carex texensis (Torr.ex L.H. Bailey) L.H. Bailey—R; > RF, OH; (04400). Carex tribuloides Wahlenb.—V; WHR; BF; (04913). Carex virescens Muhl. ex Willd.—V; WHR; RF, OH?: (H.R. DeSelm s.n., TENN). tCarex vulpinoidea Michx.—l; OCB, WHR; BWM, WM, LWM; (03242). BRIT.ORG/SIDA 21(4) Torr. & Hook.—V; WHR; DIS; ee ial (06790). tCyperus echinatus (L Ae me OCB, WHR; PAS, BWM, LWM, WM, RB; (01257). Cyperus flavescens L.—R; rie RD, wet ditch; 02836). Cyperus lancastriensis Porter ex Gray—S; OCB; Cyperus odoratus L.—S; OCB; PAS; (01205) tCyperus pseudovegetus Steud.—R; EHR; WM; (05028) Cyperus refractus Engelm. ex Boeckl.—R; OCB; LWM; (03644) *Cyperus rotundus L.—V; OCB; CG; (04100). Cyperus squarrosus L.—S;OCB; CG, LWM; (04105), tCyperus strigosus L—O; ALL; WM, BWM; (04177). reid bifida S.G. Smith—R; OCB; CG, CB, M; (03267); [Smith 2001]. een engelmannii Steud.—R; EHR; WM; (02352). Eleocharis microcarpa Torr.var.filiculmis Torr—v; EHR; FW; (05390) Eleocharis obtusa (Willd.) J. A. Schultes—O; ALL; : M, LWM: RM, BM; (06288) Eleocharis cf. tenuis (Willd) Schultes—V; EHR;WM; (06297). Fimbristylis autumnalis (L.) Roemer & Schultes— R; WHR; WM, FP; (01331). Isolepis carinata Hook. & Arn. ex Torr.—V; EHR; WM; (02347) tRhynchospora capitellata (Michx.) Vah|—S; HR; Rhynchospora corniculata (Lam.) Gray—R; HR; Rhynchospora glomerata (L.) Vahl—S; HR; WM, RM, FW; (03780 tRhynchospora recognita (Gale) Kral—X; OCB; BO; (A. J. Sharp, S. Fairchild, & E. Clebsch 9892, TENN). t Scirpus atrovirens Willd—l; ALL: WM, BWM, LWM; (00998) tScirpus cyperinus (L.) Kunth—S; WHR: RM; (01323), Scirpus pendulus Muhl.—V; OCB; LWM; (04903). Scleria pauciflora Muhl. ex Willd. var. pauciflora— >; WHR; RB; (05288). Scleria oligantha Michx.—S; WHR; OH, RO; (05154). Scleria triglomerata Michx.—S; HR; FW, OH; 5026). ESTES, FLORA OF TENNESSEE DIOSCOREACEAE *Dioscorea polystachya Turczaninow—O; OCB, HR; BRIP. RIP; (05311) Dioscorea villosa L_—O; ALL; BRF, RF; (00608). IRIDACEAE *Belamcanda chinensis (L.) DC—S;OCB, WHR; CG, PAS, LAW; (01474). Iris cristata Sol.ex Ait.—I; HR; RF; (01849). *Iris germanica L—R; EHR; RD; (04376). ae tae eae oe io WHR; CG,CB, F, BO, RO; (0 tS ee pein in es ALL; BWM, WM, PAS, BRIP, RIP; (02200). JUNCACEAE ~ acuminatus Michx.—S; EHR, WHR; FW, M, BWM; (01012 VDB). jee ea Ell_—R; EHR; WM; (06296). tJuncus coriaceus Mackenzie—l; ALL; WM, BWM, CS, AS; 64). HR; WM; (02339). ‘sunaie eae ae —R; WHR, OCB; RM, BWM; (05291). Juncus effusus L.—l; ALL; RM, WM, BM, BWM; (02844). Juncus a ae Buckl.—R; OCB; LWM, CG; (03246). Juncus | ee ARE —R; WHER; RB, DIS; (05304). Juncus marginatus Rostk.—R; EHR; WM; (02356). Juncus repens Michx.—R; WHR; RM; (01324). Juncus scirpoides Lam.—R; EHR; WM; (02777). Juncus secundus Beauv. ex Poir.—R; WHR, OCB; RB, CG; (05287). Juncus tenuis Willd —O; ALL; PAS, BWM, WM, BM; (03862). Juncus torreyi Coville—V; OCB; BWM; (03690). Luzula acuminata Raf. var. carolinae (S. Wats.) ern.—R; WHR; RF; (02139). tLuzula echinata (Small) Hermann—l; ALL; RF, RF; (02136). LEMNACEAE Lemna minor L.—O; ALL; FP ST; (01473). Spirodela polyrrhiza (L.) Schleid.—S; WHR; FP; (03682). LILIACEAE Allium canadense L. var. Saal ALL; RB, PAS, CB, CG, RD; (02321) Alli rnuum Roth—V; OCB; BO; (00835). *tAllium sativum L.—xX; OCB; RD; (A. J. Sharp, A. Clebsch, E. Clebsch 40, TENN). 2365 tAllium tricoccum Sol. var. burdickii Hanes—R; WHR, OCB; RF, BRF; (00406). *Allium vineale L—l; OCB, WHR; RD, PAS, LAW; (02317) Amianthium (caliaa (Walt.) Gray—R;WHR; RF, OH; (02117). Te ne L.—S; OCB, WHR; RD, PAS; (04919). Pree scilloides (Raf.) Cory—S; OCB; BRF, BDF; (01793). Chamaelirium luteum (L.) Gray—R; WHR; OH; (03297). Erythronium albidum Nutt.—S; OCB; BRF, LKW; (01662). Erythronium americanum Ker-Gawl subsp. americanum—R; WHR; RF; (05908). Erythronium americanum Ker-Gawl subsp. harperi ae Parks and Hardin—O; ALL; RF, BRF; 98). Pe A en (L.) L—I;ALL;RD,WM, ST, LAW; (02357). *Hyacin soe nonscripta (L.) Chouard—V;WHR; RD; (01 ‘ymenoca occidentalis (J.Le Conte) Kunth— |; WHR; WM, RF; (01334). tHypoxis hirsuta (L.) Coville—S; OCB, WHR; CG, H; (01950 A Lilium michiganense Farw.—V, WHR; RF. Maianthemum racemosum (L.) Link subsp. racemosum—O; ALL; RF, BRF, LKW; (02029) *Muscari ped Guss. ex Ten.—l; OCB; C PAS; (003 38). *Narcissus poeticus L—R; WHR; LAW; (00421). *A Narcissus pseudonarcissus L—O;ALL;LAW,RD, PAS. tNothoscordum bivalve (L.) Britt —S; OCB;CG,CB, M, LKW, BO; (01653). *Orni ee umbellatum L.—S;OCB;LAW, PAS, CG; (019 Po signatumb a orum (Walt.) Ell —O; ALL; RF, BRF, LKW Bionartes laugnos (Michx.) D. Don—V; WHR; RF; (02034). tSchoenolirion croceum (Michx.) Wood.—x; OCB; CG, LWM; (E. Quarterman 5218, speci- men formerly at VDB, now apparently lost). tTrillium cuneatum Raf.—C; ALL; RF, BRF, LKW; (01934). Trillium flexipes Raf—R; WHR; RF; (01733). Trillium sessile L_—l; OCB; LKW, BDF, BRF; (01916). 2366 t Trillium stamineum Harbison—l; ALL; RF, BRF; (01805). Uvularia grandiflora Sm.—l; WHR; RF; (01666). Uvularia perfoliata L.—R; WHR; RF; (H.R. DeSelm Uvularia sessilifolia L—R; EHR; FW (06293). ORCHIDACEAE Cypripedium parviflorum Salisb. var. pubescens Willd.) Knight—V; WHR; RF; (02030). Goodyera pubescens Willd. R. Br.ex Ait.f—R; WHR; 85). Isotria verticillata Raf —V; EHR; FW; (02800). Platanthera ciliaris (L.) Lindl. WHR; RM and associated wet woods; (02745). Mee clavellata (Michx.) ae WHR; AS, RM; (0095 7). Platanthera flava (L.) Lindl. var. flava—V; EHR; FW; (06283 Platanthera peramoena (Gray) Gray—V; OCB; BWM; (03868). Spiranthes cernua (L.) Rich.—R; WHR; PAS, RD; (06880). Spiranthes lacera (Raf.) Raf. var. Hate (Bigelow) Luer—S; OCB, WHR; CB, RB; (02880). Spiranthes vernalis Engelm.& Gray—V; EHR;WM; ( ). 04943 Tipularia discolor (Pursh) Nutt.—l; ALL; RF, BRF; 801). POACEAE “Agrostis gigantea Roth—R; OCB; WM; (06276). Agrostis hyemalis (Walt.) B. S. P—R; EHR; WM; (02348). Agrostis perennens (Walt.) Tuckerman—l;ALL;OH, FW, BDF; (03669) Alopecurus carolinianus Walt.—R; OCB; BWM; (02206) Andropogon gerardii Vitman—S; WHR; RB, RO; (03684), Andropogon gyrans Ashe—R; WHR; RB, RM; (05677) Andropogon ternarius Michx.—R; WHR; RB; Andropogon virginicus L.—C; ALL; PAS, RB, RD; (06884). Aristida longespica Poir—R; WHR; DIS; (06799). — oligantha Michx.—R; OCB, WHR; CB, RB; 90). oti Gales (Thunb.) Makino—R; EHR; WM, FW; (04195). BRIT.ORG/SIDA 21(4) tArundinaria gigantea (Walt.) Chapm.—O; ALL; BRIP, RIP. BRF, RF, FW; (00250). oe erectum (Schreb. ex Spreng.) Beauv.—l|; WHR; RF; (02848). enue Ne Schrad.—|;OCB, WHR; PAS, Bromus os Muhl. ex Willd.—l; WHR; RF; (06413 R romiie racemocrs<¢ L.—V; OCB, EHR; (01943). *Bromus sterilis L.—V; OCB; CG, PAS; (02044). *Bromus tectorum L.—|;OCB;RD, PAS, CG; (04457). Chasmanthium latifolium (Michx.) Yates—|;WHR, RIP BRIP. LKW; (01482). Chasmanthium laxum (L.) Yates—R;EHR;FW,WM; (02776). Chasmanthium sessiliflorum (Poir.) Yates—l:OCB, WHR; LKW, 0787) inna arundinacea L 01532). *Cynodon dactylon (L.) Pers. —O; ALL; PAS, LAW, RD, DIS; (05397). *Dactylis glomerata L_—O; ALL; PAS, RD; (02096). Danthonia spicata (L.) Beauv. ex Roemer & J. A. Schultes—l; HR; OH, (02257). Dichanthelium acuminatum (Sw.,) Gould & CA, Clark var. acuminatum—R; OCB; BWM; (02210). Dichanthelium acuminatum (Sw.) Gould & C. A. Clark var. lindheimeri (Nash) Beetle—S;WHR; RB: (03663). Dichanthelium sal oe Gould & C.A.Clark— - HR; OH, RF; (03759). tDichanthelium commutatum (J. A. Schultes) Gould—l; ALL; BRF, RF; (H.R. DeSelm 1654, TENN). Dichanthelium depauperatum (Muh. 18). Bichantheltt ium dichotoniaite (L.) Gould—O; HR; OH, FW; (05276). Dichanthelium laxiflorum (Lam.) Gould—O; ALL; DF, LKW, BRF, RF; (00762 VDB) OCB; CG, CB, BDF; (00646 VDB),. Dichanthelium microcarpon (Muhl. ex Ell.) ohlenbrock—l; ALL; WM, RM, BWM, FW; (00788 VDB). Dichanthelium sphaerocarpon (Ell.) Gould var. ana oe Gould & C. A. Clark—S; << Gould—S; Fea scoparium (Lam.) Gould—S; HR; WM W; (04183). ESTES, FLORA OF UNTY, TENNESSEE *Digitaria ciliaris (Retz.) Koel.—l; OCB, WHR; DIS, CRP. LAW, RD, GB, CG; (02729 VDB). * Digitaria sanguinalis (L.) Scop.—|; OCB; PAS, RD; (05264) Digitaria violascens Link—V; WHR; DIS; (06798). *Echinochloa colona (L.) Link —l;OCB;BWM,LWM,; (04081) ac ee — (L.) Beauv.—S; WHR; GB, DIS; (02 rehinocl oa muricata (Beauv.) Fern. va muri —|; ALL; WM, BWM, BM, RM; ae *Fleusine ie (L.) Gaertn.—l; OCB; DIS,CRP RD; 176). + Elymus glabriflorus (Vasey ex LH.Dewey) Scribn. & Ball —R; WHR; RD; (05313). Elymus hystrix L—O; OCB, WHR; BDF, BRF, BRIP, LKW, RF, RIP; (00947). @Elymus macgregorii R. Brooks & J.J.N. Camp- bell—l; WHR, OCB; RIP. BRIP; (02320); [Camp- bell 2000). tElymus riparius Wieg.—l; WHR, OCB; RF, BRIP; (00946). be villosus Muhl. ex Willd.—O; OCB, WHR; BO, ake RO; an i; Elymus 7 WHR, OCB; RIP, BRIP, RF,RD, PAS: (05298), ¥C liq fake EI All.) Vign ex Janchen—S; “OCB: DIS, CRE.RD; Gn *Eragrostis curvula (Schrad.) Nees—S; OCB; RD; (06271). Eragrostis frankiiC.A. Mey ex Steud.—R; OCB; CG; Eragrostis pectinacea es Nees ex Steud.—I; OCB; PAS, CRP, RD; (052 Eragrostis spectabilis ean Sees —S; HR; RB; Festuca di a Schreb.—C; ALL; PAS, DIS, LAW, RD; (02211). +Festuca subverticillata (Pers.) Alexeev.—|; WHR; RF; (02185) Glyceria striata (Lam.) A.S.Hitchcock—|;WHR,CS, AS, ST, WM; (02024) *Hordeum pusillum Nutt.—l; ALL; DIS, RD, PAS; (021 43). ¥4tLJ mT { | Ti VDB). Leersia oryzoides (L.) Sw.—l; ALL; BM, BWM, RM, WM, FW (0 Bet): tLeersia virginica Willd.—O; ALL; BRIP, RIP, GB, ST; BWM, WM; (02978). L.—V;OCB;RD;(R.Kral 64891, 2367 *Lolium perenne L. subsp. ce (Lam.) Husnot.—l; ALL; DIS, RD, PAS; (02197). Melica mutica i eee RE BO, RO; (00610 *Microstegium vimineum (Trin.) A. Camus—F; ALL; RF, BRF, BDF, DIS, RD, ST, BM, BWM, FW; (05468). *Miscanthus sinensis Anderss.—R; WHR; PAS; 6327). Muhlenbergia schreberi J.F.Gmel.—l; WHR, OCB; OH, RO, CG, PAS; (05646). ae sobolifera (Muhl.) Trin—R; WHR; RF; 47). Hate sylvatica Torr.ex Gray—S;WHR; RF; (05148). aoa anceeay pine —0O;ALL;RB, PAS; (03655). —S;0C AP CRP. Panicum dichotom ee on —|; ALL; BWM, WM M, GB, DIS; (04 Panicum see ae —l; OCB; GG, CB, BO, PAS; (0410 Pani philadelphi Bernh.subsp.gattinger! (Nash) Freckmann & Lelong—!; OCB; CG, CB, BO, RD; (05652). Panicum rigidulum Bosc. ex Nees var. pubescens (Vasey) Lelong—V; WHR; RM; (02748). Panicum rigidulum Bosc.ex Nees var. rigidulum— |; OCB, WHR; BWM, WM, (03856) Panicum verrucosum Muhl.—R; HR; WM; (04073). Panicum virgatum L.—R; OCB; RD, DIS; (06272). *Paspalum dilatatum Poir—O;ALL; DIS, PAS, LAW; tPaspalum distichum L—x; OCB; river bottom, S. Treanor Jr.s.n., TENN). Paspalum fluitans (Ell) Kunth.—V; WHR; WM, RM; 04086) Paspalum laeve Michx. var. pilosum Scribn.—R; EHR, OCB; WM, LWM; (05021) *Paspalum urvillei Steud.—V; WHR; RD; (06774). *Phleum pratense L.—S; OCB; PAS, RD; (05411). *Poa annua L.—O; ALL; DIS, PAS, LAW, RD; (04403). Poa autumnalis Munhl. ex Ell—tl; HR; RF; (02085 VD *Poa compressa L.—R; OCB; PAS; (04953). *Poa pratensis L.—O; ALL; DIS, PAS, LAW, RD; (04450) oa sylvestris Gray—l; HR; RF, RO; (01930). *Poa trivialis L.—\V; WHR; CS; (02095 VDB). Saccharum alopecuroidum (L.) Nutt.—O; ALL; PAS, RB, RD; (04109). 2368 Saccharum baldwinii Spreng.—V; WHR; RM; Saccharum aiganteum (Walt.) Pers.—R; EHR; WM, RB; (04197). echiedenyauyr seopanina (Michx.) Nash var. R.\A/UIP- PR. (NDE Schiccehy sini seonartinn (Michx,) Nash var. scoparium—l; HR; RB, OH, RO; (03649). *t Secale cereale L.—S; OCB; CG, RD; (02209). *Setaria faberi Herrm.—O; OCB; CRP DIS, PAS, RD; (05413). acne (Poir.) Kerguelen—O; ALL; BWM, WM D, RB; (02733 VDB). *tSetaria ils L.) Beauv. var. major (Gaudin) ospichal—R; OCB; CRP; (H.R. DeSelm 1652, TENN). *Setaria viridis (L.) Beauv. var. viridis—l|; OCB; CRP IS, PAS, RD; (05266). Sorghastrum nutans (L.) Nash—R; WHR; RB; (02916). “Sorghum halapense (L.) Pers.—F: ALL; DIS, PAS, CRP, RD; (00776). a aca (Rydb.) Rydb.—R; EHR; W; (02 Pate Pesce Scribn.—l; ALL; BRF, L 51). Sporobol land — TA ary Pe tinus (Biehler) A. S. Hitche. — S$; OCB; CG, CB; (05649), — compositus (Poir.) Merr. var. compositus—R; OCB; CG; (05487). Sporobolus ona (Torr. ex Gray) Wood—|; OCB; CG, CB; (04103). Tridens flavus (L.) A.S. fete var. flavus—O; ALL; B, PAS, LWM, RD; (04169). pia myuros (L.) K. ie Gmel.—R; WHR; RB; 20). es eanoh Hea ie var. glauca (Nutt.) B; (00587). *Vu ‘EB WHR; R PONTEDERIACEAE *A Eichhornia crassipes (Mart.) Solms—R; OCB, WHR; FP, observed in two separate ponds over me course of two or three years but een in either pond in 2004, bean ee dubia i MacM.—R; OCB; RIV, swift shallows; (03517). Heteranthera reniformis Ruiz & Pavon—V: OCB; BM; (03528). POTOMOGETONACEAE Potomogeton diversifolius Raf.—R; WHR; RM; (02752 BRIT.ORG/SIDA 21(4) ae oe Raf. subsp. foliosus—R; OCB; B ee ar Poir—V; OCB; RIV, swift shallows; (03520). SMILACACEAE t Smilax bona-nox L.—F;OCB, WHR; BDF, LKW,CG, BRIP. RIP. BO, RO; (00208), Smilax glauca Walt.—|; ALL; OH, RB, FW, BDF; 00022). Smilax hugeri (Small) J.B. Norton ex Pennell—S; WHR, OCB; RF, LKW; (00604). Smilax eal Hook.—S; WHR, OCB; OH, RF, BDF, PAS; (01472). Smilax der L.—O; ALL; OH, FW, BDF; (00200) tSmilax tamnoides L.—l; WHR, OCB; RF. BRF, RIP. BRIP; (00162 SPARGANIACEAE Sparganium americanum Nutt.—R; WHR; RM; (03782) TYPHACEAE Typha oo ia L.—V;WHR;RD, roadside ditch; (00 Typha i 1a L.—1; ALL;RM, BM, BWM, FP; (03693). XYRIDACEAE Xyris laxifolia Mart. var. iridifolia (Chapman) Kral.—V; EHR; WM; (06298). ANGIOSPERMS: DICOTS ACANTHACEAE Justicia americana (L.) VahI—l; OCB; RIV, GB; (03519). Ruellia pluie UJ. F. Gmel.) Steud.—I; WHR; RB, 992). Ruellia finns Nutt.—S; OCB; CG, CB; (00784). Ruellia strepens L_—S; OCB; LKW, BDF, CG; (04091). sania \A/URD. OL (0 S; Wt WA, AP, negundo Lk ALL: BRIP, RIP. FW. (00142 TSU). Nc 77 hae st Michx. f.—l; WHR, OCB; RF, BRF; (01922). Acer aes L. var. trilobum Torr. & Gray ex K. Koch—O; ALL; OH, FW, RM, RD; (00826). Acer saccharinum L.—O; OCB; BRIP. BM; (05319). tAcer saccharum Marsh.—C; ALL; RF, BRF; (04499); [including both var. saccharum and var. schneckii Rehd.]. ESTES, FLORA OF , TENNESSEE AMARANTHACEAE *Achyranthes japonica (Miq.) Nakai var. hachijoensis Honda—R; OCB; BRIP; (06419). Amaranthus hybridus L.—l; OCB; CRP, DIS, RD; 46). (037 *Amaranthus palmeri S.Wats.—S; OCB; CRP, DIS, RD; (04175) *Amaranthus powellii S. Wats.—R; OCB; CRP; (05273). *Amaranthus spinosus L.—O; ALL; PAS, DIS; (02714). ANACARDIACEAE +Rhus aromatica Ait.—l; OCB; LKW, CG, CB, BO, BDF; (01970). Rhus fel L.var. ee Engl.—O; ALL; RB, OH, BDF, PAS, RD; (03664). tRhus glabra ee ALL; ae RD; (03672). Rhus hirta (L.) Sudworth—S; OCB, WHR; BRF, PAS, RD; (03983 Toxicodendron radican BDF, RF, BRF, FW, BRIP. RIP. (03675). ANNONACEAE Asimina triloba (L.) Dunal—F; ALL; RF, BRF, BRIP RIP; (00401) APIACEAE Angelica venenosa 00976). —= Greenway) Fern.—|; WHR; RB; *tCarum carvi L.—V;OCB;RD; (R. Kral 64893, VDB). tChaerophyllum procumbens (L.) Crantz.—1;OCB; BRF, RF; (05839). tChaerophyllum tainturieri Hook—O; ALL; PAS, RD, DIS; (02316). Cicuta maculata L.—R; OCB; BWM; (02835). *Conium maculatum L.—l; OCB; DIS, PAS, CRP. BWM, BRIP; (04501). Cryptotaenia canadensis (L.) DC.—; ALL; RF, BRF; (00943) *Daucus carota L.—F; ALL; PAS, DIS, RD, RB; (03656). Erigenia bulbosa (Michx.) Nutt.—S; WHR; RF; (01634 Fry 791). tEryngium yuccifolium Michx.—V; WHR; RB; (R. Carter & T. Smith 3133,V et (Michx.) C. B Clarke—lI; ALL; RF, BRF, LKW, 6). Osmorhiza ee: (Torr.) DC.—S; WHR, OCB; RF, LKW; (04911 Kuntze—C; ALL; OH, tratum Nutt. ex DC.—V; EHR; WM; 2369 Oxypolis rigidior (L.) Raf —S; WHR, OCB; AS, CS, ST, WM, BWM; (01184). *+Pastinaca sativa L—R; OCB; PAS, RD; (04942). Perideridia americana (Nutt. ex DC.) Reichenb.—V; OCB; LKW; (021 12), tPtilimnium capillaceum (Michx.) Raf—S; EHR, OCB; WM, BM; (02774) Sanicula canadensis L.var.canadensis—O;ALL;RF, BRF, BDF; (00945). Sanicula odorata (Raf.) K. M. Pryer & L. R. Phi e—|; WHR; RF; (00934). an smallii Bickn.—R; WHR; OH; (02849). Sanicula trifoliata Bickn.—S; WHR, OCB; RF, BRF; (02728). os da trifoliatum (L.) ea var. qureum —O;WHER; RF, RO; (02684). ee pment ) ae OCB; DIS, PAS, CG; (03704). +Zizia aurea (L.) W. D. J. Koch—S; WHR; RIP; (02130). APOCYNACEAE msonia_ tabernaemontana Walt. var. tabernaemontana—V; WHR; RF; (04912). Apocynum cannabinum L.—|;ALL; WM, BWM, RD; (00983). ale difforme (Walt.) Gray—V;WHR; M; (04000). *Vinca major L.—R; OCB; RD, BDF; (01788). *Vinca minor L.—S;OCB, WHR; LAW, LAW; (01632). AQUIFOLIACEAE tllex decidua Walt.—S; OCB; BDF, RD; (06793). llex longipes Chapman ex Trel —R; WHR, OCB; OH, LKW; (02871). Ilex opaca Ait.—S; WHR, OCB; OH, BDF; (00282). llex verticillata (L.) Gray—R; EHR; FW; (00555). ARALIACEAE Aralia racemosa L.—V;WHR; RF; (01269). Aralia spinosa L—l; ALL; RF, ie . (00961). *A Hedera helix L—R; OCB; L Panax aca L.—S; ae Fe BRF, LKW; (00933). ARISTOLOCHIACEAE Aristolochia serpentaria L_—|; WHR; RF; (02824). +Aristolochia tomentosa Sims—l|; OCB; BRIP; ) adense L.—O; ALL; RF, BRF; (01736). ASCLEPIADACEAE Asclepias amplexicaulis Sm.—R;WHR; RB; (03661). 2370 Asclepias exaltata L.—V; WHR; RF; (02268). Asclepias incarnata L. subsp. incarnata—l; ALL; M, RM, BM; (03859), eee quadrifolia Jacq.—R; WHR; RF; (02129). ASC fs) 3671). st tuberosa L. eee interior Woods.—O; PAS, RD, RB; (04 ace ias gee irs ae RB, OH; (02249). cillata L—V; WHR; RO; (05158). asclepies vitidiflora Raf_—R; OCB; CG, CB; (03715). Asclepias viridis Walt.—R; OCB; CG, PAS; (02108). ynanchum laeve (Michx.) Pers.—l; OCB, WHR; DIS, BRIP, RD; (03514). Matelea caroliniensis Jacq.) Woods.—|; WHR; RF, RO; (02323). tMatelea gonocarpos (Walt.) Shinners—l; OCB, HR; LKW, BDF, BO, RO; (00657). ASTERACEAE Achillea millefolium L.—O,; ALL; PAS, DIS, RB, RD; (02138). tAgeratina altissima (L.) King & H. E. Robins.—I; R; RF, BRF, BDF, DIS, RD; (04066). Ambrosi anenisiOl ia L.—F;ALL; DIS, RB,CG, PAS, CRP. RD; (04068). Ambrosia bidentata Michx.—R; WHR;RB; (02739), Ambrosia trifida L.—O; OCB, WHR; BRIP. RIP. DIS, R aS Amphiachyris dracunculoides (DC.) Nutt—S;OCB; CG, CB, PAS; (04099) tAntennaria plantaginifolia (L.) Richards.—O; ALL; OH, BDF, LKW; (01715 APSC). Antennaria solitaria Rydb.—S; WHR; OH; (01874). *Anthemis cotula l.—R,; OCB; PAS; (04952). “Arctium minus Bernh.—S; OCB; DIS, PAS; (06432). Arnoglossum atriplicifolium (L.) H. £. Robins.—| RF, RIP; (03872). tArnoglossum muehlenbergii (Schultz-Bip.) H obins.—R; WHR; RF; (00950), “Artemisia annua L._—O; OCB; DIS, RD, PAS, CG; (04093). ieee integrifolium (Michx.) Nutt. subsp. rifolium—S; OCB; CG, CB, LKW; (01773). me halimifolia L—V;EHR;FW, DIS; (04202), Bidens aristosa (Michx.) Britt —l; WHR: WM, RM; (01517). Bidens bipinnata L.—l; OCB; CG, LKW, BDF, DIS; (02887). tBidens cernua L.—S; OCB; BWM; (04088). Bidens frondosa L.—S; OCB; BWM, DIS; (05467). eptas syriaca L.—O; OCB; PAS, BRIP, RD; BRIT.ORG/SIDA 21(4) Boltonia asteroides (L.) L'Hér—R; EHR; WM; ce Brickellia eupatorioides (L.) Shinners—V; OCB; CG, (04097). *Carduus nutans L—F;ALL;DIS, ie RD,CG; (02041). *Centaurea biebersteinii DC.—R; OCB: RD; (D. Sunbrink CENMA-I-65MMS5-5/21/98, TENN). *Centaurea cyanus L.—R; OCB; CRP, PAS; (02047). Chrysopsis mariana (L.) Ell—S;WHR;OH; (02898). Chrysopsis pilosa Nutt.—R; WHR; RB; (02842). *Cichorium intybus L.—l; OCB; RD, PAS, DIS; (06438). tCirsium altissimum (L.) Hill—l; WHR; RIP. RF; 50). Cirsium discolor (Muhl. ex Willd.) Spreng.—O; OCB, WHR; PAS, RB, RD; (05402). Cirsium muticum Michx.—V; WHR; RB; (01301). *+Cnicus benedictus L—X; OCB; alfalfa field: (T.T. Jackson 20432, TENN). Conoclinium coelestinum (L.) DC.—O; ALL; WM, BWM, PAS, RD; (03857). Conyza canadensis (L.) Cronq. var.canadensis—l|; ALL; DIS, PAS, RD; (01191). Conyza canadensis (L.) ae var. pusilla (Nutt.) Crong.—R; WHR: RB; (02858). Coreopsis ienccolaka a OCB; highway roadcut; (02208 EKU). Coreopsis major Walt.—O; WHR; OH, RB; (00033). Coreopsis tinctoria Nutt.—R; WHR, OCB; RD, PAS; Coreopsis tripteris L.—l; WHR; RB; (01183). *Crepis pulchra L.—R; OCB; CG; (03240). seca oe (Michx.) Greene—R; WHR; OH; (02905). Doell ie umbellata (P. Mill.) Nees var. umbellata—R; WHR; RM, WM; (02747). *Eclipta ae (L.) L—t; ALL; BWM, WM, GB; (02761). Elephantopus carolinianus Raeusch.—|; ALL; BDF, O RD; (01258 VDB) Elephantopus tomentosus L.—S; WHR: OH; (02746) Erechtites hieraciifolia (L.) Raf.ex DC.—1; ALL; RB, OH, DIS, RD; yee Erig (L.) Pers.—l; OCB, WHR; PAS, RD; ” (02453), Erigeron philadelphicus L—O;WHR, OCB; PAS, RD; 811). tErigeron pulchellus Michx.—S;WHR; OH, RF, RO; (00119) ESTES, FLURA UT , TENNESSEE a igeron sure Muhl. ex Willd. var. calcicola Allison—S; OCB; CG; (03647) [Allison and ee Dei Erigeron strigosus Muhl. ex Willd. var. strigosus— O; ALL; PAS, RB, RD; (00783). Eupatorium album L. var. album— (02828) Eupatorium altissimum L—R;OCB;CG,CB; (04098). Eupatorium capillifolium (Lam.) Small—l; ALL; PAS, RD, DIS; (04200). rapa oe Barratt—!; WHR, OCB; M, BWM R; WHR; RB; on): Pp lum Mera L—l;WHR, OCB; RB, CB; (02891). Eupatorium perfoliatum L.—l|; ALL; WM, BWM, RM, BM; (03853) Fry patorit aa) pilosum W. alt. —R; EHR; FW; (05029). Eupatort ium x pinnatifidum Ell—V; EHR; WM, FW; (04199). Eupatorium purpureum L.—S;WHR; RF; (00948). Eupatorium rotundifolium L. var. ovatum (Big- elow) Torr.—O; HR; RB, OH, WM; (03737). Eupatorium serotinum Michx.—|;OCB, WHR; PAS, DIS; (01316). Eupatorium sessilifolium L.—S; WHR; OH, RF; (01094). Eurybia hemispherica (Alexander) Nesom—l; WwW OH; (01137) Fleischmannia incarnata (Walt.) King & H.E.Rob- ins.—l; OCB; BDF, LKW; (0417 *Galinsoga quadriradiata Cut OCB; RD; (00647 Gamochaeta purpurea (L.) Cabrera—t; ALL; PAS, RB, OH, BDF, RD; (02066) Grindelia lanceolata Nutt. var. lanceolata—R;OCB; CG, CB; (02687). Helenium amarum (Raf.) H.Rock—O;ALL;PAS,CG, RD; (03706). tHelenium autumnale L—S;OCB, EHR; LWM,WM; 93). Helenium flexuosum Raf.—S; HR; WM; (00001). elaninus pa ius L.—S; EHR;WM,; (04198). a V; OCB; RD; (05420). Helianthus airoibehs L—l;WHR; RB, OH; (01389). Helianthus decapetalus L.—R; WHR; RF; (01093a,b) Helianthus eggertii Small —R;WHR;RB; (02851). Helianthus hirsutus Raf—l; WHR; RB, OH; (01139). Helianthus microcephalus Torr. & Gray—l; WHR; RB, OH; (01305). 2371 Helianthus silphioides Nutt —R; WHR; RB; (03755) Helianthus strumosus L.—V;WHR; RF, RD; (03871). +Helianthus tuberosus L—O;OCB, WHR; BRIP, RIP; (01034) Heliopsis helianthoides (L.) Sweet—l; WHR; RF; — = } Hieracium gronovii L—|; WHR; OH, RB; (02903). ee radicata L.—R; WHR; PAS; (03756). *lvaann OCB; BWM; (05462). tKrigia ene (Walt.) Blake—O; WHR; OH; (00289). tKrigia caespitosa (Raf.) ali ALL; CRP, DIS, BWM, GB, WM, CG; (02214). Krigia dandelion (L.) Nutt.—R; ii (02056). tLactuca canadensis L._—O; ALL; PAS, RB, RD; (03665) Lactuca floridana (L.) Gaertn.—l; WHR; RF, RIP RD; *Lactuca saligna L.—R; OCB; RD; (06280). *L actuca serriola L.—l; OCB; RD, PAS, DIS; (03674). *leucanthemum vulgare Lam.—O; ALL; PAS, RD, CG, CB, DIS; (0177 Liatris spicata (L.) Willd. var. spicata—S; WHR; RB; (02841). Liatris squarrulosa Michx. S; WHR; RB; (O01 522). *Matricaria discoidea DC.—V; OCB; CRP; (01670 VDB). Mikania scandens (L.) Willd.—R; EHR, FW, WM; (04187 Packera oe (Wood) W. A. Weber & A. L6 ALL; RB, CG, CB, PAS; (02065). +Packera g aul (Poir) C. Jeffrey—l; ALL; BWM, M,CS, CRP, RD; (02097). Packera obovata (Muhl. ex Willd.) W. A. Weber & A. Love—l; WHR, OCB; OH, RO, LKW, BO; (00456). Parthenium integrifolium L. var. integrifolium—l; Pluchea camphorata (L.) DC.—S; ALL; WM, BWM; (04082). tPolymnia canadensis L.—|; OCB, WHR; LKW, BDF, BO, a O; (02713). altissima L.—l|; WHR; RF; (01479). Beudgnaot ium setae ium (L.) Hilliard & —S;WHR; RB; (01 Pooapo aa ae cere DC.—|; ALL; BWM _ PAS, RB; (05163). Ratibida a fanata (Vent.) Barnh.—R; OCB; rocky PAS; (04898) 2372 ee fulgida Ait. var. fulgida—R; WHR; RB; (01048). ruth fulgida Ait. var. umbrosa (C.L.Boynton eadle) Crong.—R; WHR; CS, ST; (06787). iti hirta L. var. hirta—O; ALL; RB, OH, RD, PAS; (00982 VDB) Rudbeckia hirta L.var. pulcherrima Farw.—l|; OCB; PAS, RD; (00210). Rudbeckia laciniata L. var. laciniata—|; WHR; RF, CS; (01053). eae triloba L—O; OCB; CG, CB, BDF, PAS, 00779 a linifolius (L (L.) B.S. P—S;WHR; RB, OH; (01390). Silphium asteriscus L. var. asteriscus—l|; WHR; OH, RB, RF; (03752) tSilphium mohrii Small—S; WHR; RB; (00922). Silphium perfoliatum L.—S; OCB; BRIP,. PAS, RD; (01342) Silphium trifoliatum L. var. latifolium Gray—R; OG CB; (01046). Smallanthus uvedalius (L.) Mackenzie ex Small— |; HR; RF, RD; (05423) Solidago arguta Ait.var.caroliniana Gray—|; WHR; Solidago caesia L.—|; ALL; RF, BRF, BO, RO; (06794). ae canadensis L var. i Torr. & Gray— F; ALL; RB, PAS, RD; (04111). Pursh—I; ee OH; (02900). Soidage flexicaulis L_—R; WHR; RF; (02726). Solidago gigantea Ait —|; ALL; WM, BWM; (01129). Solidago hispida Muhl. ex Willd. var. hispida—l; : (02902). Solidag Solidago missouriensis Nutt. var. fasciculata Holz.—R; WHR; RB; (01047). Solidago nemoralis Ait —O;WHR; RB, PAS; (01521). Solidago odora Ait. var. odora—|; WHR; RB; —_ (02869). Solidago patula Muhl. ex Willd.—R; WHR; CS; Solidago rugosa P. Mill. ssp. aspera (Ait.) Crong.— |; HR; OH, RB, WM, FW; (06781). Solidago sphacelata Raf —R; WHR; RO; (02846). ae ulmifolia Muhl. ex Willd.—l; WHR; OH; (05274). Osos asper (L.) Hill—l; EHR, PAS; (02340), Symphyotrichum cordifolium (L.) Nesom—|l;WHR; RF; (01481). Symphyotrichum drummondii (Lindl.) Nesom— 5; OCB, WHR; BDF, RO; (05638). BRIT.ORG/SIDA 21(4) Symphyotrichum dumosum (L.) Nesom—|; WHR; RB; (02868). parece lanceolatum (Willd.) Nesom— - (05465) a Smpyrichan set ile A. & D. L6ve— O; ALL; RF, BRF, PAS, RD; (01349 VDB). Pr eects ontarione (Wieg.) Nesom—R; OCB; M; (05645). Symphyotrichum sees (Ait.) Nesom—l; WHR; RB, OH; (02972). pl ichum pil (Willd.) N FALL; PAS, RD, DIS; (01471). Symphyotrichum priceae (Britt.) Nesom—R;OCB; 2. Symphyotrichum shortii(Lindl.) Nesom—O; ALL; BDF, BO, RO, OH; (01382). Symphyotrichum undulatum (L.) Nesom—R; Ww OH; (02974), a aed urophyllum (Lindl.) Nesom—O; F BRF, RO; (01536 VDB). Pile ae .) OC.—R; OCB; RD; (01974). *Taraxacum officinale G.H. Weber ex Wiggers— PAS, RD, DIS, CRP; (06441). Verbesina alternifolia (L.) Britt. ex Kearney—O; OCB; RIP, RF, BRIP. WM, BWM; (01261) Verbesina virgir es L.—O;WHR, OCB; RIP BRIP.CG, CB, PAS; (01131). tVernonia sia (Walt.) Trel.ex Bran.& Cov.— RB, WM, BWM; (03860). ae staumatiimn Lt ALL; PAS, DIS; (04095) BALSAMINACEAE Impatiens capensis Meerb.—O; ALL; RF, BRE. WM, WM, CS, AS; (01032). eee pallida Nutt.—R; WHR, OCB; RF, BRF; (0112 BERBERIDACEAE *Berberis bealei Fortune— nee oe (05915). “A Berberis thunbergii DC.—V; OCB; PAS. tCaulophyllum thalictroides (L.) rene —R; WHR; mal Jeffersonia diphylla (L.) Pers.—S; WHR, OCB; RF, BRF; (00385). *A Nandina domestica Thunb.—V; OCB; highway roadcut. tPodophy flum peltate 1m LC ALL; BRF, RF, BDF: (01746). ESTES, FLORA OF GILES COUNT ¥, FENNESSEE BETULACEAE Alnus serrulata (Ait.) Willd. —O;HR; RIP.CS, AS, FW; ) (00316 Betula nigra L.—S; ALL; RM, FW, BRIP; (01142). Carpinus caroliniana Walt.—F; ALL; RF, BRF, RIP, BRIP; (02857). Corylus americana Walt.—|; WHR; OH, RF; (00962). tOstrya virginiana (Mill) K. Koch—O; ALL; BDF, W, OH, RF, BRF, BO, RO; (05278 BIGNONIACEAE iigneiva oe L.—O; ALL; RF, BRF, BDF, LKW, tCampsis ae re S RF, BRF, BDF, RD; (02 B O; ALL; g). *Catal . econ Walt.—R; OCB; BRIP; (02 89). ban ae laaue pen Warder ex Engelm.—S; OCB; BORAGINACEAE *t Buglossoides arvensis (L.) |. M. Johnston—S; OCB; CG, RD, DIS; (01660) Cynoglossum ee L.—l; ALL; BRF, BDF, LKW, RF; (01 953). ie sin ; dic (ee EHR; FW. R; He eletopium tenellurn (Nutt.) Torr—R; OCB; CG, CB; (02690). bees canescens (Michx.) Lehm.—V; B; VDB). Quarterman 526 Laem Saas um Rugel ex —|;WHR, OCB; OH, RF, BDF, ar RB; (02882). Mertensia Gane (L.) Pers.ex Link—l; WHR, OCB; RIP, BRIP; (00127 +Myosotis macrosperma Engelm.—O; ALL; RF, BRF, BDF, LKW; (01958). t+Onosmodium molle Michx. subsp. molle—S; OCB; CG, CB, PAS; (04849). BRASSICACEAE *Alliaria petiolata (Bieb.) Cavara & Grande—S; 82) Bea thaliana (L.) Heynh.—R; EHR; CRP; (04394). wee ae (Muhl. ex Willd.) Poir. var. laevigata—O; ALL; RF, RO, BRF, BO, LKW ci *tBarbarea verna (P. Mill.) Aschers.—S; WHR; RD, RIP (02134). Baek: vulgaris Ait. f—O; ALL; PAS, RD; (00 435). 2373 *+Brassica rapa L.—O; ALL; CRP, PAS, RD, DIS; ) 00328). *+Capsella bursa-pastoris (L.) Medik.—l; OCB; CRP ies DIS; (01748). Ci a tata O. E. Schulz—l; WHR; PF; J (03039). tCardamine bulbosa (Schreb.ex Muhl.) B.S. P—l; WHR; CS; (01727). tCardamine concatenata (Michx.) Sw.—F; ALL; BRF, RF; (01701). Cardamine diphylla (Michx.) Wood.—R; OCB; BRF; (05832). *Cardamine hirsuta L.—C; ALL; CRP, DIS, BRF, BDF, RF, LKW; (01774) tCardamine parviflora L_—R;OCB, WHR; LKW, RO; (AJ.Sharp,C. J.Felix,&W.Adams 10930,TENN). Cardamine pensylvanica Muhl.ex Willd.—S;WHR, OCB; CS, BWM; (04453). tDescurainia pinnata (Walt.) Britt. subsp. brachycarpa (Richards) Detling—xX; OCB; CG; We Quarterman, J. Baskin, & S. Oakland 64-52, DB). “der inia sophia (L.) Webb ex Prantl—V; OCB; RD; (R. Kral 64884, VDB), Draba brachycarpa Nutt.ex Torr. & Gray—R; EHR, OCB; CRP, CG; (04395). *Draba verna L._—l; OCB; CG, RD, DIS; (01661). *tErysimum repandum L.—R; OCB; CRP, RD; (03077), arte le (Michx.) Steud.—S; WHR; RF; (02186). ‘eavenrti exigua Rollins var. exigua—x; OCB; CG; (E. Quarterman, J. Baskin & S. Oak- land 64-61,VDB). oo torulosa Gray—S;OCB; CG, LWM; (01603). *tLepidium campestre (L.) Ait. f—R; OCB; CG; (01964). K+] ij al ifl — Schrad.—V; OCB; RD; (R. Kral 64894, 2 May 1980, VDB). ie ee L.—I|; OCB; CG, CB, PAS, RD, een eee Rollins—R; OCB; CRP; (01672). *Rorippa nasturtium-aquaticum (L.) Hayek.—O; ALL; ST; (01768) Rorippa sessiliflora (Nutt.) A.S. Hitchc.—R; OCB; WM; (04396). Sibara virginica (L.) Rollins—S; OCB, WHR; CRP; (01671). 2374 *tSisymbrium officinale (L.) Scop.—S; OCB; CG, RD; (01969) *Th Sai asp! arvense L.—R; OCB: RD; (01790). UXACEAE Pachysandra procumbens Michx.—l; WHR; RF; 10) CACTACEAE Opuntia humifusa (Raf.) Raf. var. humifusa—t; OCB; CG, CB, BO, PAS; (01626). CALLITRICHACEAE Callitriche heterophylla Pursh — subsp. heterophylla—S; WHR; RM; (02203). CAMPANULACEAE tCampanulastrum americanum (L.) Small—t; R, OCB; RF, BRF; (00655). Lobelia ag L.—l; WHR; WM, RM, RIP. ST: (01134). Lobelia rae L.—l; WHR; RD; (02702). Lobelia puberula Michx.var.puberula—|;WHR; RB; (01300). tLobelia siphilitica L—O; ALL; BWM, WM; (05418). A Lobelia spicata Lam.— B; CB. tTriodani nee fata (L.) Nieuivl var.perfoliata— |; WHR, OCB; DIS, RD; PAS; (02313). CAPPARACEAE *Cleome hassleriana Chod.—V;WHR; GB; (02732). CAPRIFOLIACEAE *Lonicera fragrantissima Lindl.& Paxton—R;WHR, OCB; RD, UT; (03038 *Lonicera japonica Thunb.—C; ALL; BRF, BDF, WM, BRIP, RIP PAS, RD; (02256). *Lonicera maackii (Rupr.) Herder—R; OCB; RD, UT; 70) (05670). tlonicera sempervirens L.—|; WHR; OH, RO; 081). Sambucus nigra L.subsp.canadensis (L.) R.Bolli— O; ALL; BWM, BM, WM, RM, FW, BRIP. RIP. CS; (02826). bac at ee se iculatus Moench—C; ALL; BDF, RF, BO, RD; (05644). L. var. lucidum Ait.—R; EHR; \VAih rt. FW; (02790 EKU). Viburnum nudum L. var. nudum—R; HR; FW, RM; (05013) tViburnum rufidulum Raf—O; ALL; BDF, LKW, BRE, F:(01973) BRIT.ORG/SIDA 21(4) CARYOPHYLLACEAE tArenaria lanuginosa (Michx.) Rohrb.—R; OCB, WHR; 2 *+Arenaria serpyllifolia L.—R; OCB; RD; (02194). *tCerastium Hee tees Desportes ex Pers.—S; OCB; CG, RD; (01784). Cerastium aaa renee Ege ex Gray) B.L. R R; WHR; OH; (05906). eae fontanum see . subsp. vulgare artman) Greuter & Burdet—S; OCB; RD;(A J. Sharp, C. J. Felix, & B. Adams 10957, TENN). *Cerastium glomeratum Thuill—S; WHR; LAW; (02092 VDB) tCerastium nutans Raf—S; WHR; RO, RF; (01924 VDB) *Dianthus armeria L.—l; OCB; RD, PAS; (02196). *Holosteum umbellatum L.—S; OCB; CRP, CG; (01674). tMinuartia patula (Michx.) Mattf—l; OCB;CG, CB, BO; (00363), *Myosoton aquaticum (L.) Moench—R;WHR;CS, ST; (02090 VDB). *Saponaria officinalis L.—l; OCB, WHR; RD, PAS; 77). *Silene latifolia Poir. subsp. alba (P. Mill.) Greuter rdet—R; WHR; RD; (02723). Silene stellata (L.) Ait. f—S; WHR; RF; (01045), t Silene virginica L—O; ALL; RF, OH, RO, BRF, BDF; — — 00595), Stellaria fontinalis (Short & Peter) B.L.Robins. — *Stellaria media (L.) Vill. subsp. media—C; ALL; RF, BRF, LKW, LAW, PAS, RD; (01867 Stellaria pubera Michx.—I; HR; RF; (01810). CELASTRACEAE Celastrus scandens L.—V; WHR; RIP; (01529). *Euonymus alata (Thunb.) Sieb.—R; EHR; FW, OH; (02049). tEuonymus americana L.—F; ALL; RF, BRF, LKW; a 22). S; OCB; LKW BDF BRF; (02889). *Euonymus jf (Turcz.) Hand.-Maz.—R;OCB; UT, LKW *Euonymus aera Loesener.—R; OCB; BDF, RD, BRIP. UT: (05674). CHENOPODIACEAE *Chenopodium album L.—O; ALL; DIS,RD, PAS, GB, CRP; (04172) ESTES, FLORA OF TENNESSEE Chenopodium standleyanum Aellen—V;WRR RF; *Dysphania ambrosioides (L.) Mosyakin & Clemants—l; OCB; PAS, GB, DIS; (01198). CISTACEAE Lechea minor L.—V;WHR; RB (05309). +Lechea mucronata Raf.—S; WHR; RB; (02843). hcl l tifolium Ell—R; WHR; RB; (06776). Biypen icum crux-andreae (L.) Crantz—R; EHR; FW, = Hypericum drummondii (Grey. & Hook.) Torr. & ray—|; WHR; PAS, RB; (01189). Hypericum frondosum Michx.—S; OCB; BO, BDF; (00985). Hypericum gentianoides (L.) B. S. P—R; WHR; RB; (02854). Hypericum hypericoides (L.) Crantz subsp. hypericoides—S; HR; FW,OH, AS, WM; (02703). +Hypericum hypericoides (L.) Crantz subsp. multicaule (Michx. ex Willd.) Robson—|; ALL; BDF; (000 tHypericum muniim L.—l; ALL; WM, RM, BWM, * Hypericum perforatum L.—R; OCB; RD; (04917). Hypericum prolificum L.—R; OCB; BDF; (00774). Hypericum aa Lam.—l; ALL; RB, PAS, OH, BDF; (01 Hypericum oe Michx.—l; OCB; CG, LWM, CB; (02688). Triadenum virginicum (L.) Raf.—R; EHR; FW; (04189) BS oa + I+ a | iadenum walteri J.G.Gmel.) Gleason—VR;WHR; CS; (06380). CONVOLVULACEA Cal ae aah Pursh—R; WHR; RB; (2078 DB). Ca = ee silvatica (Kit.) Griseb. subsp. fraterni- florus (Mackenzie & Bush) Brummitt—F; OC WHR; BRIP, RIP, BWM, WM, RD; (03754). *Dichondra caroliniensis Michx. —R; OCB; LAW, (06440). in R; OCB; RD; (01340). "pomoea poe eee ian —S;OCB; CRP DIS, RD; 054 Bae acunosa L.—R; OCB; CRP; (01336). tlpomoea pandurata (L.) G. F. W. Mey.—l; WHR; OH, RB; (00991). 2375 *Ipomoea purpurea (L.) Roth—S; OCB; RD, CRP; (06881) CORNACEAE Cornus alternifolia L. f—\V; WHR; RF; (01757). +Cornus amomum P. Mill.—l; ALL: WM, RM; ST, AS, CS, BWM; (01528). +Cornus ee. A.Mey.—|;OCB; BDF, LKW, BRIP, BO, CG, CB; (04108). tCornus oe L.—F; ALL; RF, BRF, BDF, OH; Cornus foemina P. Mill.—R; EHR; FW; (02341). Nyssa biflora Walt.—R; HR; RM, FW; (02793). tNyssa sylvatica Marsh.—O;ALL;OH, BDF, BRF, FW, RM; (02925). CRASSULACEAE +Penthorum sedoides L.—l; ALL; WM, BWM, RM, BM; (02725). Ba pa um Michx.—l;OCB; CG, BO, LKW; (00 ee als Bunge—R; WHR; RO; (02088). A Sedum ternatum Michx.—S; WHR; RF, RO. CUCURBITACEAE *Citrullus vulgaris (Thunb.) Matsumura & Nakai— V; WHR; DIS; (05302). *Cucurbita pepo L.—R; OCB; PAS, RD; (01194). +Melothria pendula L.—R; EHR; FW; (04185). Sicyos angulatus L.—|; OCB; BRIP; (04090). CUSCUTACEAE Cuscuta compacta Juss. ex Choisy —S; WHR; CS, ST, RIP; (01299) Cuscuta gronovii Willd. ex J.A. Schultes—R; EHR; 03) Cuscuta pentagona Engelm.—|; OCB, WHR; PAS, RD; (05429). DIPSACACEAE *Dipsacus fullonum L.—R; OCB; PAS, RD; (05173). EBENACEAE Diospyros virginiana L.—F; ALL; PAS, BDF; (01908 ELAEAGNACEAE *Elaeagnus umbellata Thunb.—R;WHR, OCB; OH, RD, PAS; (03148). ERICACEAE Gaylussacia baccata ( EHR; FW; (02803). Wangenh.) K. Koch—V; 2376 tKalmia latifolia L.—l; HR; OH, RO; (02894), tOxydendrum arboreum (L.) DC.—O; ALL; OH, BDF; (03658). Rhododendron alabamense Rehd.—R; WHR: AS, Rhododendron canescens oo ) Sweet—I; HR; H, FW, RM, AS, RB; (01872). ee ake periclymenoides Shinners—R; HR; OH, AS; (01802). tVaccinium arboreum Marsh.—|; ALL;OH, RB, CB; (02888). Vaccinium corymbosum L.—R; EHR; FW; Saas Vaccinium fuscatum Ait.—l; HR; OH, AS, FW, R (03026). (Michx. —_ tVaccinium pallidum Ait.—|l; WHR; OH, RB; (00423), Vaccinium stamineum L.—|; HR; OH, RB; (01758). piesaincaae \A/UR. RR-(ND RV VITA, IAD, \U 861). Acalypha stevia Riddell OCB; CRP, PAS, RD; (04 tAcalypha enmbolden Raf.—X; OCB; BO; (A. J. Sharp, S. Fairchild, & E. Clebsch 9859, TENN). Acalypha es L.—O; ALL; RD, DIS, CRP. LWM, CG, GB; (05 *Chamaesyce maculata (L.) Small—l; OCB; PAS, RD, DIS; (05392), Chamaesyce nutans (Lag.) Small—l; ALL; PAS, RB R *Chamaesyce prostrata (Ait.) Small—R; OCB;CRP a Croton capitatus Michx.—S;OCB; CG, CB; (05484). tCroton monanthogynus Michx.—l;OCB;CG, CB BO, RD; (05617). iii corollata L.—l; WHR; RB, OH, RD; “porbi cyparissias L.—R; OCB; RD, LAW; (016 tpt deniers Michx.—l; OCB; BO, CG, CB; (00648). Ephab mercurialina Michx.—R; OCB; BRF; (03185). Phyl ons caroliniensis Walt.—|; ALL; WM, BWM; (04118). Tragia cordata Michx.—V; WHR; RO; (05159). BAC “Albizia julibrissin Durazz.—|; ALL; DIS, BRF. RF. RD; ). tAmorpha fruticosa L.—S; WHR, OCB; WM, CG; (01130 APSC). BRIT.ORG/SIDA 21(4) Amorpha nitens Boynt.—R; OCB; CG; (00780). tAmphicarpaea bracteata (L.) Fern.—l; ALL; RF, (01599), tApios americana Medik.—|; ALL; WM, BWM, ST: (02855). Apios priceana B. L. Robins.—R; WHR; RF, RO; (02710). Astragalus canadensis L.—R; OCB; BO; (06275). tCercis canadensis L.—F; BDF, LKW, OH, RF, PAS; (01718) as see Gel Se Greene—I; PAS, RB, RD; (01 eae nictitans i 5 eens R;RB; 02823) Clitoria mariana L.—|; WHR; RB; (00233). *Coronilla varia L—l; ALL; RD; (02181). Crotalaria sagittalis L_—R; WHR; RB; (02852). esmanthus illinoensis (Michx.) MacM. ex B. L. Robins, & Fern.—R; OCB; BWM, PAS; (04126). Desmodium ciliare (Muhl. ex Willd.) DC. var Rae 5; ven die (02740), dium um (Muhl. ex Willd.) DC. ex Loud.—R; WHR: RE (01 256) Desmodium glutinosum (Muhl.ex Willd.) Wood— S; WHR; RF; (00994) Desmodium laevigatum (Nutt.) DC_—R;WHR;RD: (06786). Desmodium nudiflorum (L.) DC.—1; WHR; OH, RF; (03750). Desmodium nuttallii (Schindl.) Schub.—l; OCB, WHR; PAS, RD, RB; (01319) Desmodium paniculatum (L.) DC.—1; ALL; PAS, OH, FW; (04120) tDesmodium pauciflorum (Nutt.) DC.—I; HR; RF; Desmodium perplexum Schub,—R; WHR; RF; (01530). Desmodium rotundifolium DC.—l; WHR; OH, RB; (02901). Desmodium viridiflorum (L.) DC.—S; WHR; RB; 2856). Galactia volubilis (L.) Britt —l; WHR, OCB; RB, OH, CG, CB; (03705). Gleditsia triacanthos L.—O; OCB; PAS, BRF. BRIP: (05400). *Kummerowia stipulacea (Maxim.) Makino—R; WHR; RD; (02706). “Kummerowia striata (Thunb.) Schind|.—F; ALL: PAS, sh RD; ” 399), *Lath tus L.—R; EHR; WM, PAS; (02358). ESTES, FLORA OF GILES COUNTY, TENNESSEE *Lathyrus latifolius L—R; WHR; RD; (03740). *lespedeza bicolor Turcz.—S; WHR; RD, PAS, RB; (03668). *lespedeza cuneata (Dum.-Cours.) G. Don—C; ALL; PAS, RD, RB; (04071). Lespedeza frutescens (L.) Hornem.—S; WHR; RB; (02878). Lespedeza hirta (L.) Hornem. subsp. hirta—|; WHR; Lespedeza procumbens Michx.—|; WHR, OCB; RB, OH, CB; (02862) Lespedeza repens (L.) W. Bart.—S; WHR; RB; 39). Lespedeza violacea (L.) Pers.—R;OCB; CG; (05488). Lespedeza virginica (L.) Britt. —l; WHR; RB; (01141). *Medicago lupulina L—F; ALL; PAS, RD; (01786). *Melilotus albus Medik.—F; ALL; PAS, RD, DIS; (00755). *Melilotus officinalis (L.) Lam.—l; WHR; RD; e018 Mil ricrophylla Dry.—S; WHR; RB; (02314). Orbex fie pedunculatum (P. Mill.) Rydb. var. pedunculatum—R; WHR; RB, OH; (02077). Pediomelum tee (Torr. & Gray) Rydb.—V, see X;OCB; CG, moist soil; (D. McGavock SU). mek us eae: (L.) B. S. PR—R; WHR; RB; (01091). *tPueraria montana (Lour.) Merr. var. nee (Willd.) Maesen & S. Almeida—S; OCB, W DIS, RD; (03670 **Robinia hispida hy var. hispida—R; WHR; LAW, RD; (01185). Robinia pseudoacacia L.—O; ALL; BDF, BRF, RF, PAS; (05286). Senna marilandica (L.) Link—R; OCB, WHR; CG, PAS, RF; (06433) *Senna obtusifolia (L.) Irwin & Barneby—I; OCB, WHR; CRP; (05269) te oo (Muhl. ex Willd.) Britt — sso nae ao S.P—I;WHR; RB; (02700). ephrosia virginiana (L.) Pers.—l; WHR; RB; (02182). *t Trifolium campestre Schreb.—O; ALL; PAS, RD; (00600). *Trifolium incarnatum L.—R; OCB; CB; (01962). *Trifolium pratense L.—F; ALL; PAS, RD; (00770). *Trifolium repens L.—C; ALL; LAW, PAS, RD, CG; (03271) 2377 tVicia caroliniana Walt.—|; WHR; RF, OH; (01722). *Vicia grandiflora Scop.—R;OCB; PAS, RD; (01770). tVicia minutiflora F.G. Dietr—l; OCB, WHR; LKW, F, RF; (00448) *Vicia sativa L.subsp.nigra (L.) Enrh.—l;OCB; PAS, (01769) par *Vicia villosa Roth subsp. varia (Host) Corb.—S; WHER; RD; (01005). FAGACEAE Castanea dentata (Marsh.) Borkh.—S; WHR, OCB; OH, BDF; (00973). *Castanea mollissima Blume—R; OCB; RD, LAW; (00141). tFagus grandifolia Ehrh.—F; ALL; BRF, RF, FW; (00135). Quercus alba L.—F; ALL:OH, BDF, RF, BRF; (02924). Quercus bicolor le —V; OCB; BWM; (00988). pale Mun hh.—l; HR; OH; (05314). cata Michx.—O; HR; OH; (01311). a X filialis Little—V;WHR;RM, WM; (02909) tQuercus imbricaria Michx.—S; OCB, WHR; BDF, OH; (04107 Quercus lyrata Walt.—R;HR; RM,WM, FW; (02907). Quercus marilandica Munchh.—S; HR; OH, RB; (02926). Quercus michauxii Nutt.—R; OCB; BRIP; (02922). Quercus montana Willd.—O; ALL; OH, BDF; (05904). tQuercus muehlenbergii Engelm.—C; ALL; LKW, F RF, RO; (04094) Quercus nigra L_—S; EHR; FW; (04113). t+Quercus pagoda Raf—S; OCB; BDF, BRIP; (00007). tQuercus phellos L.—l; HR; FW, RM, WM, OH; (02910) Quercus rubra L.—O; ALL; RF, BRF, LKW; (01476). tQuercus shumardii Buckl—C; ALL; BDF, BRF, LKW, BO, RF; (05422) Quercus stellata Wangenh.—O; ALL; OH, BDF; (04112) Quercus velutina Lam.—O; ALL; OH, BDF; (00960). FUMARIACEAE tCorydalis flavula (Raf) DC.—l; OCB, WHR; BRF, BDF, RF, LKW; (03139). Dicentra cucullaria (L.) Bernh.—R; WHR; RF; (01698 GENTIANACEAE Bartonia paniculata (Michx.) Muhl.—V; EHR; FW; =o inica (L.) B.S. P—V;WHR; AS; (01310). i . OGICOLHG VITGIT 2378 Frasera caroliniensis Walt.—l; WHR, OCB; RF, OH, BDF, LKW; (00963 Obolaria virginica L.—R; WHR; OH; (03140). Sabatia angularis (L.) Pursh—l; WHR, OCB; RB, CG; (03712). Sabatia brachiata Ell_—R; WHR; RB; (03651). GERANIACEAE Geranium carolinianum L. var. carolinianum—t: ALL; PAS, RD, CG; (01951). *Geranium dissectum L.—S; OCB; CRP. LAW, PAS: (01946). Geranium maculatum L.—|; HR; RF; (01762). *Geranium molle L.—S; OCB; LKW,CG,CB; (01775). GROSSULARIACEAE [tea virginica L.—S; HR; AS, RM, WM, FW; (00549), HALORAGACEAE *Myriophyllum aquaticum (Vell.) Verdc.—R; OCB; FP, BWM; (05280) Proserpinaca palustris L.—R; HR; RM, FW, ST; (00561). HAMAMELIDACEAE Hamamelis virginiana L.—I; HR; RF; (01050). Liquidambar styraciflua L.—O; ALL; FW, OH, BDF, RM; (00790) HIPPOCASTANACEAE Aesculus flava Ait.—|; ALL; BRF, RF; (05838). tAesculus glabra Willd. —|; OCB; BDF, PAS; (00818 haat x hybrida DC.—V;WHR; RF; (00408 VDB). Aesculus pavia L.—O;, ALL; BRF, BDF, RF; (01798). HYDRANGEACEAE t+Decumaria barbara L.—R; WHR; RF, RIP: CS: ce Hydrangea arborescens L.—R; OCB; BRF; (03515). tHydrangea cinerea Small—F; ALL; RF, BRF, RO, BO; 2452). Hydrangea quercifolia Bartr.—R; WHR; OH; (02100). Philadelphus hirsutus Nutt.—S; WHR; RF, RO; (02188). tPhiladelphus pubescens Loisel. var, pubescens— S; OCB; BO, BDF; (A. J. Sharp, S. Fairchild, & E. Clebsch 9864, TENN HYDROPHYLLACEAE Hydrophyllum late WHR; BRF, RF; (01932). Mycrophyllum ee Nutt.—R; WHR; RF; (02145). ulatum Michx.—l; OCB, BRIT.ORG/SIDA 21(4) tNemophi ss a(L.) Brummitt—O; OCB;LKW, BDF, BRF: (01717). tPhacelia bipinnatfida Michx.—O; ALL; RF, BRF; (00402). Phacelia dubia (L.) Trel.var.interior Fern.—V; OCB; CG; (01782). tPhacelia purshii Buckl—O; OCB; PAS, CG, RD; (03264). JUGLANDACEAE a cordiformis (Wangenh.) K.Koch—O; ALL; F, BRF; (05421). a glabra (Mill.) Sweet—O; ALL; OH, RF, BDF, BRF; (00028). Carya ovalis (Wang.) Sarg.—O; OCB, WHR; RF, BDF, RF; (00057). **Carya illinoinensis (Wangenh.) K.Koch—R;OCB; RD; (02923). Carya laciniosa (F. Michx.) Loudon—R; OCB; BRF, BDF; (05387) ba ovata (P.Mill.) K. Koch var. australis (Ashe) —I; OCB; BDF, LKW, BO, CG, CB; (00611). eae ovata (P. Mill.) K. Koch var. ovata—F; ALL; BDF, OH, BRF, RF, LKW; (0540 Carya tomentosa (Poir.) Nutt —O: ALL; OH, BDF; Juglans cinerea |.—S; ALL; RF, BRF, RIP; (00249). Juglans nigra L.—F; ALL; BRIP RIP BRF, RF; (05285). LAMIACEAE tAgastache nepetoides (L.) Kuntze—S; WHR; RF, RIP; (R. Kral 36930, TENN), Ajuga reptans L.—R; OCB; LAW hi ciliata (L.) Benth.—l; OCB, WHR: CG,LKW, F; (03273). sei He (Pursh.) Benth.—R;WHR, OCB RF, P; (03741). ae ae nepeta (L.) Savi—V; OCB; BDF, BO; (01003 VDB) Colli j 4| L.—I; WHR; RF; (01533). Cunila origanoides (L.) Britt. —R; WHR; OH; (02973). *Glechoma hederacea L.—O; ALL; PAS, LAW, RD, RF, BRF; (01702). Hedeoma hispida Pursh—R; OCB; PAS; See Isanthus brachiatus (L.) B. S. P.—l; OCB; (05650). “tLamium amplexicaule L.—F; ALL; PAS, LAW, RD; (01619) *Lamium purpureum L.—F; ALL; PAS, LAW, RD; (00360) ESTES, FLUKA UF TENNESSEE Lycopus americanus Muhl. ex W. Bart.—S; OCB, EHR; LWM, FW; (06788). Lycopus rubellus Moench—|;WHR, OCB; CS, BWM; Lycopus virginicus L—l: OCB; BWM, BM; (05250) *+Marrubium vulgare L—X;OCB; thin limestone soil; (A. J. Seto C.J. Felix,& B. Adams 10968, TENN). *Mentha X piperita L—O;WHR,OCB; WM, BWM, BM; (01038). +Monarda bradburiana Beck—S;WHR; OH, RB, RF; Monarda fistulosa L.—|; WHR; RB; (00964). *+Perilla frutescens (L.) Britt —F; ALL; PAS, DIS, RD, GB; (04069). *Prunella vulgaris L.—O; ALL; PAS, BWM, RD; (00940). eae loomisii Nutt —O; WHR; RB, RD, PAS, OH; (02845). Pycnanthemum Cee (Michx.) Pers.—R; EHR; 0630 Ms Pycnanthemum tenuifolium Schrad.—l; WHR; RB; (03650) t Salvia lyrata L—O; ALL; PAS, RD, OH, BDF, LKW, CG, CB; (00609). Salvia urticifolia L—R; WHR; OH, RB; (02252). Scutellaria elliptica Muhl. ex Spreng. var. hirsuta (Short & Peter) Fern.—l; WHR; RF; (05151). Scutellaria incana Biehler var. incana—R; WHR; Cry itellaria intearifolia ER: EHR; WM; (02338). Scutellaria lateriflora L.—|; WHR, OCB; WM, BWM; (01265) ta Hill—R; OCB; BDF; (00656). +Scutelt aria parvula Michx. var. parvula—S; OCB; CG, CB; (01785). +Teucrium canadense L.—l|; OCB; BWM, BM, PAS; 91). Trichostema dichotomum L.—V; WHR; RB; (02860). LAURACEAE +Lindera benzoin (L.) Blume var. benzoin—O;ALL; RF, BRF, RIP. BRIP; (02977) Lindera benzoin (L.) Blume var.pubescens (Palmer & Steyermark) Rehd. —R; OCB; BM; (03886). +Sassafras albidum (Nutt.) Nees—O; ALL; OH, BDF, PAS, RB; (0365/7). 2379 LINACEAE Linum medium (Planch.) Britt. var. texanum (Planch.) Fern.—R; WHR; RB; (02741). Linum striatum Walt.—|; HR; WM; (02795). tLinum virginianum L.—R; WHR; RD, RB; (03686) LOGANIACEAE Mitreola petiolata (J.F.Gmel.) Torr.& Gray—R; EHR; Spigelia marilandica (L.) L—O; ALL; RF, BRF, BDF, BO, RO; (00993). LYTHRACEAE Ammannia coccinea Rottb.—R; OCB; BWM; (05414) uphea viscosissima Jacq.—S; OCB; LWM,CG, CB, BO; (02875). Rotala ramosior (L.) Koehne—I; ALL; WM, BWM, RM, FP; (04079 MAGNOLIACEA shot tulipi ee ALL;RF, BRF; (001 36). Magnolia acuminata (L.) L—R; WHR; RF; (01755 VDB). *Magnolia grandiflora L_—R; OCB; LAW, RD; ). tMagnolia macrophylla Michx.—R; HR; RF; (02121). MALVACEAE aoe pee moscheutos L. subsp. moscheutos—|; M, RM, FW, BWM, BM; (01133). sc oe L.—S; WHR, OCB; RF, RD, LAW; 44), mee trionum L.—V; OCB; CRP; (01339). see ei . ispidum (Pursh.) Hochr.—R; OCB; CG; (05 *tSida spinosa jieec, ALL; PAS, DIS, RD, CG; (05482). MELASTOMATACEAE Rhexia mariana L. var. interior (Pennell) Kral & Bostick—R; HR; RM, FW; (02796 VDB) Rhexia mariana L. var. mariana—S; EHR; WM; (02769). Rhexia virginica L_—S; HR; WM, FW; (02704). MENISPERMIACEAE Calycocarpum lyonii (Pursh) Gray—S; OCB, WHR; BRIP. RF; (03734). Cocculus see (L.) DC.—O; OCB, WHR; BDF, BRF, RO, LKW, CG, RD; (01470). tMenispermum canadense L.—l; WHR, OCB; FF, ) ES =_ =m 2380 MOLLUGINACEAE Mollugo verticillata L.—l; OCB; CRP; (01177). MONOTROPACE Monotropa nee —V, WHR; RF; (02830), “Broussonetia papyritera (L.) ah ex Vent.—S; OCB; UT, BDF, RD, LAW; (05 *Maclura pomifera (Raf) see —0; OCB; PAS, B “Morus alba L.—R; EHR, OCB; UT; (05016). Morus rubra L.—O; ALL; BRF, BDF, BRIP. RIP, RF; (03875). OLEACEAE Chionanthus virginicus L.—S; WHR; OH, FW; (02794) tForestiera ligustrina (Michx.) Poir.—l; OCB; LKW, G *Forsythia viridissima Lind!.—R; OCB; LAW: (03033 Fraxinus americana L.—F; ALL; RF, BRF, BDF, (05279). tFraxinus pennsylvanica Marsh.—O; ALL; BRIP. RIP. WM, WM, FW; (00154). aii cuadrangulata Michx.—l; OCB; LKW, CG, BDF, BO, BRF; (00653). “Jasminum pease Lindl.—R; OCB; RD; (01604) *Ligustrum sinense Lour.—C; ALL; BRIP RIP. BREF. RF, F, LAW, LKW; (02198), ONAGRACEAE is lutetiana L. subsp. canadensis (L. schers & Magnus—l; HR; RF; (00949), pom coloratum Biehler—S; OCB, WHR; M, WM; (R. Kral 36940, VDB). ee alternifolia L.—l;ALL; WM, RM, BWM, BM, AS; (02705). Ludwigia decurrens Walt.—S; WHR, OCB; WM, BWM, FP, GB; (01330) Ludwigia glandulosa Walt.—R; OCB; BWM; YS ae eae A —V; EHR; WM; (02778). is (L.) Ell—O; ALL; RM, BM, WM, BWM, er FP; (01146). *Ludwigia uruguayensis (Camb.) H. Hara—V;WHR; ST; (06266). nothera biennis L.—O; ALL; PAS, RD; (02919). recon laciniata Hill—S; OCB; CG, DIS, RD, CRP: (01966). BRIT.ORG/SIDA 21(4) *Oenothera speciosa Nutt.—S; EHR, OCB; RD; (00558). Oenothera triloba Nutt.—S; OCB; CG; (01779). OROBANCHACEAE Conopholis americana (L.) Wallr.f —S; WHR, OCB; RF, BRF, LKW; (01787). tEpifagus virginiana (L.) W. Bart—S; WHR; RF; (01534) OXALIDACEAE Oxalis illinoensis Schwegm.—R;WHR; RF; (01929). tOxalis priceae Small subsp. priceae—S; OCB; CG; (01949), Oxalis stricta L. — PAS, DIS,RD, LAW, RB, OH, BDF, GB; (02 tOxalis woten ‘ = OCB, WHR; CG,LKW, BO, RO; (01721). PAPAVERACEAE *A Papaver dubium L.—R; OCB; ee ie canadensis ms ALL: RF, BRF; (01664). ee diphyllum (Michx.) Nutt.—R; HR; RF; (01803) PASSIFLORACEAE tPassiflora incarnata L.—F; PAS, RD; (03747). Passiflora lutea L.—S; WHR; RF; (01251). PHY TOLACCACEAE Phytolacca americana L. var.americana—F; ALL; PAS, RD; (00763) PLANTAGINACEAE Plantago aristata Michx.—R; WHR; PAS, RB: (01192). aries lanceolata L.—F; ALL; PAS, LAW, RD; (009 Peat rugelii Dcne.—F; ALL; PAS, LAW, RD; (00951). ia virginica L.—l; WHR, OCB; RD, CG; 01). PLATANACEA Platanus oe L.—C; ALL; BRIP, RIP BM, WM, RM, WM, FW; (04930) PODOSTEMACEAE Podostemum ceratophyllum Michx.—R; OCB; RIV, swift shallows; (03488). POLEMONIACEAE +Phlox amoena Sims—R; WHR; RB; (00351). t Phlox divaricata L. subsp. divaricata—O; HR; RF, RIP; (01723). ESTES, FLORA OF TENNESSEE tPhlox glaberrima L_—R; WHR; AS; (00956). Phlox paniculata L_—R; WHR, OCB; RIP RF, BRIP, RD; 32). Polemonium reptans L.—O; ALL; RF, BRF, RIP; (01734). POLVGALACEAE bigua Nutt.—R; WHR; RB; (02742). | iata V; EHR; WM; (06301). Shoals incarnata ee WHER; RB; (02853). Polygala sanguinea L.—S; HR; WM, RM; (01148). POLYGONACEAE *Polygonum aviculare L—S;ALL; RD, GB; (02354). *+ Polygonum caespitosum Blume var.longisetum (de Bruyn) A.N. a O; ALL;GB, ST, BRIP RIP. RF, BREF, DIS, ( *Polygonum Derreies an & Zucc.—R; OCB; UT; (06436). *+ Polygonum hydropiper L.—X; WHR; RM; (A. J. Sharp, E. Clebsch, & A. Clepsch 9822, TENN). ae ee glee Michx.—l; ALL; WM, M, BM, FW; (02760). eka lapathifolium L.—S; ALL; WM, BWM; (05162). tPol eee pensylvanicum L.—O; WHR, OCB; M, BWM; (05463). *D lyaon| m persicaria L._—l; OCB; BWM, RD, DIS: b (051 68). tPolygonum punctatum Ell.—l; EHR, OCB; WM, BWM; (03858). Polygonum sagittatum L.—S; HR; WM; (01381). tPolygonum scandens L.—S; WHR; RO; (06328) Polygonum setaceum Baldw.—R; WHR; GB, ST; (01253). t Polygonum virginianum L.—O; ALL; RF, BRF, BDF, LKW; (01127) *Rumex acetosella L—l; ALL; PAS, RD; (01854). Rumex altissimus Wood—l; ALL; BWM, BM, WM, RM; (02062) *Rumex conglomeratus Murr.—V; OCB; BWM; (064 42). *Rumex crispus L.subsp.crispus—F; ALL; PAS, DIS, RD; (02204) *Rumex obtusifolius L_—R; WHR; RD; (02829). PORTULACACE AE tClaytonia virginica L.—O; ALL; RF, BRF, BDF; 99). 016 +Phemeranthus calcaricus (S. Ware) Kiger—R; OCB; CG; (03703) 2381 *Portulaca oleracea L.—|; OCB; CG, CRP, RD; (04102). PRIMULACEAE Dodecatheon meadia L.—R; WHR; RF; (03134 MTSU). q hi 4 iliata | L.—R; WHR; RIP; (05294). “bumech nummularia L.—S; OCB; BWM, BRIP; (02201). Lysimachia quadrifolia L—R; WHR; RF; (03679). tSamolus valerandi L. subsp. parviflorus (Raf) Hultén—S; WHR, OCB; CS, ST, GB; (03761). PYROLACEAE Chimaphila maculata (L.) Pursh—F; ALL; OH, BDF; (00231). RANUNCULACEAE tActaea pachypoda Elliott—l; ALL; RF, BRF; (01807 +Anemone acutiloba (DC.) G. Lawson—O; HR; RF; (00317) Anemone americana (DC.) H. Hara—R; WHR; OH; (01633). Anemone quinquefolia L. var. quinquefolia—R; WHR; RF; (03137). tAnemone virginiana L.—|;WHR, OCB; RB, CG, CB, BDF; (01049). tAquilegia canadensis L.—|; ALL; BO, RO, LKW, CG; (00342). Cimicifuga rubifolia Kearney—R; WHR; RF; 2912). +Clematis catesbyana Pursh—R; OCB; CG, CB, BDF; 03252). “Clematis enue DC.—R; OCB; RD; (01341). Smalle Rydb.— —R; a BO, Clematis versicolor BOF: (064 23). Clematis viorna L—R; WHR; RF; (04914). Clematis virginiana L—O; OCB, WHR; BRIP RIP, RD; 60). *Consolida ajacis ( (03645). tDelphinium tricorne Michx.—l; OCB, WHR; BRF, RF, LKW; (01794). Enemion biternatum Raf—R; WHR; RF; (01696). Hydrastis canadensis |.—S; WHR, OCB; RF, BRF; (01266). tMy curtis minim | L.) Schur—V; OCB; LKW; us L.—R; OCB; PAS, CRP. RD; (E. ‘Ousnennen 5261,VD +Ranunculus abortivus L. = ALL; BRIP RIP. LAW, RD, PAS, FW, BRF, RF; (01669). *Ranunculus bulbosus L.—S; OCB; RD; (05837). 2382 tRanunculus fascicularis Muhl. ex Bigelow—S; ;CG, CB, LKW; (00343) Ranunculus hispidus Michx. var. hispidus—l; WHR; RF, OH; (01865). Ranunculus hispidus Michx. var. nitidus (Chap- man) 7. Duncan—S; EHR, OCB; WM, FW, BWM; (01668) tRanunculus micranthus Nutt.—S;OCB; LKW, BO; (01778) —, *tRanunculus parviflorus L.—V; OCB; RD; (R. Kral DB). Ranunculus pusillus Poir—S; ALL; WM, BWM; (02054). t+Ranunculus recurvatus Poir.—l; HR; RF; (00602). *Ranunculus sardous Crantz—C; ALL; PAS, RD, LWM, CG; (00939 VDB). tThalictrum dioicum L.—R; WHR; RF, RO; (01868). Thalictrum revolutum DC.—S; WHR, OCB; WM, BWM; (03865). peace tea ) oo & B. Boivin— ALL; RF, BRF, LKW; (016 <- RHAMNACEAE Berchemia ae (Hill) K. Koch—S; OCB, WHR; RE RF; (02128). Ceanothus americanus L.—S; WHR; RB; (00232). Fengu caroliniana (Walt.) Gray—O; ALL; BRF, F, LKW,R _R Rhamnus lanceolata Pursh.—R; OCB: LKW, CG; (02885), ROSACEAE sti os Ait.—S; WHR, OCB; WM, M; (01132). eae en Wallr.—S; OCB; BDF; (01178). TAGE: boat lata Malas |; WHR; RF; (01262). x. f.) Fern. —O; HR; OH, RB, FW. (00128). Ar nelar rchier canader 1515 (L.) Medik.—R; WHR; RB, OH: (02141). *Chaenomales speciosa (Sweet) Nakai—V; OCB; BDF (06273) Crataegus ca anne (Ehrh.) Medik.—S; WHR OCB; RF, BRF; (01915). Crataegus collina Chapm.—R; OCB, WHR; PAS; rer AIG rea(h atdesusc crus-galli L—S; OCB, WHR; PAS, BDF, OH, RB; (03854). Crataegus intricata Lange—O; ALL; PAS, RF, BRF, DF; (02137) BRIT.ORG/SIDA 21(4) Crataegus marshallii Egglest.—R; EHR; FW; (00552). Crataegus pruinosa (Wend. f.) K. Koch—R; OCB, WHR; DF; (00226). Crataegus spathulata Michx.—V; OCB; BDF; (s.n.). *Duchesnea indica (Andr.) Focke—F; ALL; PAS, RD, W:; (05967) Fragaria virginiana Duchesne—R; OCB; CG; (04458). tGeum canadense Jacq.—l|; WHR, OCB; RF, BRF, LKW; (00944). tGeum vernum (Raf.) Torr. & Gray—R; OCB; BRF; tGeum virginianum L.—X; OCB, WHR; densely ooded slopes; (H. K. Svenson 8811, TENN). Malus angustifolia (Ait.) Michx.—S; HR; RB, OH, PAS; (01754) *Malus pumila P.Mill—R; OCB; PAS, LAW; (00252). Photinia melanacarpa (Michx.) Robertson & Phipps—R; HR; FW, RB; (03186). Photinia pyrifolia (Lam.) Robertson & Phipps—l; HR; AS, RM, WM, FW; (01847) Physocarpus opulifolius (L.) Maxim.—V; OCB; BO; (00836 APSC). Porteranthus stipulatus a ex Willd.) Britt — S; WHR; B; (0225 *Potentilla nee L.—l; ae RD, PAS, CG, CB; (02099), tPotentilla simplex Michx.—|; WHR;OH, RB; (0185 7); [Plants of cedar glades and cedar barrens in the county may belong to the variety argyri- sma ee while plants from other habitats to belong to the oe ee ae americana Lae L; BDF, BRF, RF, PAS, RD, CG; (01 tPrunus angustifolia ieee —O;OCB;PAS,RD,CG; *Prunus caroliniana Ait.—V;OCB; UT, RD; (00352), *Prunus cerasifera Enrh.—R; OCB; PAS; (05409). Prunus hortulana Bailey—R; OCB; RD; (01745). *A Prunus mahaleb L.—R; OCB; BDF, LAW. tPrunus mexicana S. Wats.—S; OCB; BDF, PAS; (00260 VDB). Prunus munsoniana W.Wight & Hedrick—R; WHR; RD; (00248). *Prunus persica (L.) Batsch—l; ALL: RD, PAS, LAW; (06446). Prunus serotina Ehrh.—F; ALL; BRF, RF, BDF, OH, PAS; (01791). ESTES, FLORA OF TENNESSEE *Pyracantha fortuneana (Maxim.) Li—V;OCB; PAS; (05632). *Pyrus calleryana Decne.—R; EHR; RD; (05424). Pyrus communis L.—S; OCB; PAS, LAW; (05968). Rosa carolina L.—S;OCB, WHR;CG, CB, RB; (02109 *Rosa cae he ex eae ALL;PAS, BDF, BRF, RF, BRIP. RIP RD; (0529 Rosa setigera Michx.—O; cate BDF, PAS, RD, BWM, (03890). Rubus alumnus Bailey—R; WHR; OH; (00032). Rubus argutus Link—C; ALL; PAS, RD; (05410). *A Rubus bifrons Vest ex Tratt—R; OCB; PAS, RD. *Rubus discolor Weihe & Nees—R; EHR; RD; (05425). Rubus flagellaris Willd. —l; HR; RB, OH, PAS; (00398), Rubus occidentalis L—l; ALL: BRF, RF, BDF;(06445). a Rubus pensilvanicus Poir.—R; EHR; RF; (05405). A Rubus phoenicolasius Maxim.—V; OCB; BRF. Rubus trivialis Michx.—R; OCB; RD; (01789). *Spiraea prunifolia Seib. & Zucc.—S; WHR, OCB; RD, LAW; (01700 *Spiraea thunbergii Seib.—R; OCB; CG, LAW: (01615 RUBIACEAE tCephalanthus occidentalis L_—l; ALL; WM, RM, BWM, BM, FW, FP; (00764). *Cruciata pedemontana (Bellardi) Ehrend.—R; OCB; RD; (021 95). Diodia teres Walt.—O; ALL; PAS, RB, CB, CG, RD; tDiodia virginiana L.—l; OCB, WHR; BWM, WM, PAS; (01143) +Galium aparine L—F; ALL; BRF, RF, LKW, PAS, RD, BRIP, RIP; (01961). Galium circaezans Michx.—l; ALL; RF, BRF, LKW; (05147). *+Galium mollugo L_—V; OCB; PAS; (R. Kral 64888, VD Calne l Bigelow—R; EHR; FW; (00554). *Galium pafisiohve iat OCB; RD; (06439). tGalium pilosum Ait.—l!; WHR; RB, PAS; (00761). Galium tinctorium L.—l; ALL; WM, BWM; (02833). Galium He Michx.—O; ALL; RF, BRF, LKW, RO, BO; (02451). Houstonia ae L.—l; HR; OH; (00429). Houston! iene L.var.calycosa Gray—S; OCB; CGC 274). B; (03 Houstonia purpurea L. var. purpurea—l; ALL; RF, OH, BRF, BDF; (02126) aS 2383 Houstonia pusilla Schoepf—S; OCB, WHR; PAS, CRP; (01635) tMitchella repens L.—S; HR; FW, OH; (05394). *+ Sherardia arvensis L.—|; OCB; PAS, LAW; (01939). RUTACEAE *Poncirus trifoliata (L.) RafimS; OCB; PAS, BDF; (05835) Ptelea trifoliata L.—R; OCB; BDF, BRIP; (03742). SALICACEA *Populus a L.—S; WHR; RD, LAW; (03873). *A Populus X canescens (Ait.) Sm.—V; OCB; PAS. Bi Raves Bartr. ex Marsh.—l; OCB; BRIP DIS “A pat us hee L.—V; WHR; PAS. Salix caroliniana Michx.—l; WHR, OCB; RIP, GB; (03749). Salix humilis Marsh. var. humilis—R; WHR; RB; (01910). Salix interior Rowlee.—|; OCB; BWM, BM, BRIP, GB, RD; (01948). Salix nigra eer —F;ALL;BRIP RIP BM,RM, BWM, WM, FP; (00160). *Salix X aE Simonkai—R; OCB; RD, BRIP; (05969) Salix sericea Marsh.—R; WHR; WM, ST; (00066). SAPINDACEAE *Cardiospermum halicacabum L.—S;OCB; BWM, PAS; (01201), cae on ie oides L—|;OCB; BDF, LKW, CG, CB; (05633). SAURURACEAE Saururus cernuus L.—R; OCB, EHR; BM, BWM, FW; 87). SAXIFRAGACEAE +Heuchera americana L.var.americana—S; WHR; RF; (02133). tHeuchera villosa Michx.—O; ALL; RO, BO, LKW; (06281); [Giles County material is referable var. macrorhiza (Small) Rosl, But. & Lak}. Mitella diphylla L—R; WHR; CS, RF; (01766). +Saxifraga virginiensis Michx.—O; ALL; RO, BO, LKW, RF, BRF; (0162 Tiarella cordifolia L. var.collina Wherry—O; HR; PF; (03135). SCROPHULARIACEAE Agalinis gattingeri (Small) Small—S;WHR; RB, OH; (02914). 2384 Agalinis purpurea (L.) Pennell—R; EHR; WM, FW; (04184) Aureolaria flava (L.) Farw.—l; WHR; OH; (02897). ra pectinata (Nutt.) Pennell—S;WHR; RB, OH; (03683). tAureolaria virginica (L.) Pennell—R; EHR; FW, OH; (02798). tChelone glabra L.—|; WHR, OCB; WM, BWM, CS, AS, ST; (01 Dasistoma macrophylla (Nutt.) Raf—S; WHR OCB; RF, RD, BDF; (00935). tGratiola neglecta Torr—S; EHR, OCB; WM, FW, BWM; (02059). Gratiola pilosa Michx.—R; EHR; WM; (05020). Gratiola virginiana L.—S; HR; FW, AS; (03190). Leucospora multifida (Michx.) Nutt.—R;OCB; CRP; (05318) Lindernia dubia (L.) Pennell var. anagallidea hx.) Cooperrider—V; WHR; RB, DIS; (06791). Lindernia ire (L.) Pennell var. dubia—S; EHR, M, BWM; (05014). Wee acuminata (Walt.) Small var. acuminata—R; OCB; LWM, CG; (02893). ea us alatus Ait.—l; HR; WM; (05022). ia tomentosa (Thunb.) Sieb. & Zucc. ex ud.—O; ALL; BRF, RF, RD; (05404). ata aris eanaeenee L.—S; WHR; RF; (02131). +Penstemon nc Small—l; OCB, WHR; PAS, RD, BRIP, BO; (02098). tPenstemon tenuiflorus Pennell—R;OCB; CG, CB; (03269). Scrophularia marilandica L.—R; OCB, WHR; BRF. RF; (05155). *Verbascum blattaria L.—l; OCB, PAS, CG, RD; (03639). *Verbascum thapsus L.—F; ALL; PAS, RD; (00965). “t+ Veronica agrestis L._—R; OCB; CRP. RD; (00321). A Veronica anagallis-aquatica L.—R; OCB; ST. *tVeronica arvensis L.—l; ALL; RD, DIS, BDF; (01759) *Veronica hederifolia L.—R; WHR; RO; (01720). “Veronica peregrina L. subsp. peregrina—l; OCB, WHR; CRP, RD, DIS; (02093 APSC). *Veronica persica Poir.—R;OCB; RD, LAW; (04351 weronicg polita Fries. —R; WHR;RD; (00122 VDB). Veronicastrum virginicum (L.) Farw.—R; WHR; RB; (02707). SYS BRIT.ORG/SIDA 21(4) SIMAROUBACEAE *Ailanthus altissima (P. Mill.) Swingle—O; ALL; BRF, BDF, RF, RD; (05312) SOLANACEAE “Datura stramonium L.—l; ALL: PAS, RD, DIS; (04128) *A Nicandra physalodes (L.) Gaertn.—R; OCB; CRP. Physalis ot L.—I; OCB, WHR; CRP, DIS, RD; (01335). ee heterophylla Nees—S; OCB; oe (04921). tPhysalis longifolia Nutt. var. subglabrata (Mack enzie & Bush) Crong.—l; OCB; RD, PAS; (01206). Physalis virginiana Mill. var. virginiana—R; WHR; OH, RB; (02250). Solanum carolinense : var. carolinense—O; ALL; PAS, RD, DIS; (0231 Solanum aa Dunal—S; WHR; RF, RB; (05290). STAPHYLEACEAE tStaphylea trifolia L—F; ALL;RF, BRF, BRIP RIPLKW; (01526) STYRACACEAE Styrax americanus Lam.—R; EHR; FW; (02071). Styrax grandifolius Ait.—S;WHR; OH; (02082). THYMELAEACEAE tDirca palustris L.—l;WHR, OCB; RF, BRF; (02976). TILIACEAE t Tilia americana L.var.americana—l|; ALL: RF, BRF: (03874). tTilia americana L. var. heterophylla Loud.—F; ALL; RF, BRF; (04900). ULMACEAE tCeltis Gaus a —F; ALL; BDF, LKW, PAS, BRIP, RIP; (03709). Celtis ee L.—R; WHR; RIP; (02920). Celtis tenuifolia Nutt.—R;OCB, WHR; LKW, CG, RB; (01388 VDB). Ulmus a He Michx.—O; ALL; LKW, BDF, PAS, OH; (0161 tUlmus americana L.—l; ALL; BRIP, RIP, BDF; — Vent.) (00082). *A Ulmus pumila L.—R; OCB; PAS, RD, LAW. Ulmus rubra Muhl.—O; ALL; RF, BRF; (03765). Ulmus serotina Sarg.—l; OCB, WHR; BRF, BDF, RF, PAS; (02921) ESTES, FLORA OF TENNESSEE URTICACEAE Boehmeria cylindrica (L.) Sw.—l; ALL: WM, RM, BWM, BM; (009 42). Laportea canadensis (L.) Weddell—l; ALL; RF, BRF, RIP, BRIP; (00941) + Parietaria pensylvanica Muhl.ex Willd.—S;WHR; RF, RO; (04915 +t Pilea pumila (L.) Gray—l; ALL; RF, BRF, ST; (05167). tUrtica chamaedryoides Pursh.—l; ALL; RF, BRF; (01799) eam VALERIANACEAE Valeriana pauciflora Michx.—R;WHR; RF; (01913 *+ Valerianella (04456 ). tValerianella radiata (L.) Dufr—O;WHR, OCB; PAS, RD, RF; (02125 APSC) tValerianella umbilicata (Sullivant) Wood—l: CB; CRP, RD, PAS, LWM; (04455). VERBENACEAE tCallicarpa americana L.—|; OCB; BDF, LKW; 8). (0364 (01 eee leptostachya L.—l; ALL; RF, BRF, BDF; (00 - (02832), *Verbena brasiliensis Vell—R; OCB; PAS, BRIP; (03489). Verbena hastata L—R;WHR;WM, RM; (03753). Verbena simplex Lehm.—O; OCB, WHR; GG, CB, BO, RD, RB; (02043). Verbena urticifolia L.—l; OCB, WHR; PAS, RD; (00778 VIOLACEAE Hybanthus concolor (7. F.Forst) Spreng.—|!; WHR; RF; (01763). ~ar locusta (L.) Lat.—l; OCB; PAS, DIS; Glandularia canadensis (L.) Nutt.—R; OCB; CG; 704). 667). tPhyla lanceolata (Michx.) Greene—|; ALL; WM, 2385 tViola bicolor Pursh—O; ALL; RD, PAS, CRP; (00362) Viola cucullata Ait.—l; WHR, OCB; CS, ST, WM, BWM; (00378) Viola egglestonii Brainerd—R; OCB; CG; (01624). Viola hirsutula Brainerd—S; WHR; OH; (03142). tViola palmata L.—l; ALL; BDF, RF, BRF, LKW; 02132). Viola * primulifolia L—R;WHR; Dt Seah tViola pubescens Ait. var. scabriuscula Schw ex Torr.& Gray—O; ALL; BRF, RF; (00404 ie Viola sagittata Ait—R; WHR; RB; (03147 tViola sororia Willd.—O; RF, BRF, PAS, LAW, BRIP, RIP; (01695). tViola striata Ait—|; ALL; RF, BRF, RIP BRIP; (01797). Viola edie Ell —S;WHR; RF, OH; (02031); [Giles unty material is referable to variety ce mma (DC.) Harper]. Viola walteri House—R; OCB, WHR; BRF, RF; (03061) VISCACEAE Phoradendron leucarpum (Raf.) Reveal & M.C. Johnston—l; OCB; epiphytic on trees; (05675). VITACEAE ** Ampelopsis arborea (L. (05317). mae cordata Michx.—l; OCB; BRIP; (05316). Q wa — Koehne—R; OCB; BWM; Pras Cee : Be ee —F: ALL; F BRE, RIP P LKW; ( Vitis ee we Var. ae ALL; OH, BDF; (02267). Vitis cinerea (Engelm.) Millard var. baileyana on) Comeaux—S;WHER; RF RIP; (02266). Vitis ae a ia Michx.—O; ALL; OH, BDF; 0191). tVitis vulpina L.—F; OCB, WHR; BRIP, RIP, BWM; ACKNOWLEDGMENTS I would like to thank the following individuals for their assistance during this project. Eugene Wofford, curator of the University of Tennessee Herbarium, and Beware Chester, professor of biology at Austin Peay State University, served as offered helpful advice throughout the project. Claude Bailey, Roger McCoy, and David Lincicome of the Tennessee Natural Heritage Program de- serve thanks for providing information concerning several rare species. Robert 2386 BRIT.ORG/SIDA 21(4) Kral, curator of the Vanderbilt Herbarium at the Botanical Research Institute of Texas kindly verified several problematic specimens and loaned material for study, and Ed Schilling, professor of botany at the University of Tennessee, as- sisted with the identification of some specimens of Helianthus. Chris Fleming prepared the figure. Lastly, Eugene Wotford, Hal DeSelm, Edward Chester, Rob- ert Kral, Joey Shaw, and twoa us reviewers provided many helpful com- ments that improved the eu REFERENCES AckERMAN, E.A. 1941. The K6ppen classification of climates in North America. Geogr. Rev. 31:105-111 Atwison, J.R.and T.E.Stevens. 2001.Vascular flora of Ketona dolomite outcrops in Bibb County, Alabama. Castanea 66:154-205. Braun, E.L. 1950. Deciduous forests of eastern North America. The Blakiston Company, Philadelphia, Pennsylvania. Campsett, J.J.N. 2000. Notes on North American Elymus species with paired spikelets |: E. macgregorii sp. nov. and E. glaucus ssp. mackenzii comb. nov. J. Kentucky Acad. Sci. 61: 8 Crester, E.W.1995. An overview of forest diversity in the Interior Low Plateaus: Physiographic Province. Pp. 109-115 in: Landis, T.D.; Cregg, B., tech. coords. National Proceedings, For- est and Conservation Nursery Associations. Gen. Tech. Rep. PNW-GTR-365. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. Cuester, E.W., B.E. Worroro, R. Krat, H.R. DeSeim, and A.M. Evans. 1993. Atlas of Tennessee vas- cular plants. Volume 1. Pteridophytes, gymnosperms, angiosperms: monocots. Misc. Publ. No. 9. The Center for Field Biology, Austin Peay State University, Clarksville, Tennessee. Cuester, E.W., B.E. Worrorp, and R. Krat. 1997. Atlas of Tennessee vascular plants. Volume 2. Angiosperms: dicots. Misc. Publ. No. 13. The Center for Field Biology, Austin Peay State University, Clarksville, Tennessee. Cronauist, A. 1980. Vascular flora of the southeastern United States. Volume 1. Asteraceae. The University of North Carolina Press, Chapel Hill, North Carolina. DeSeim, H.R. 1959. A new map of the Central Basin of Tennessee. J. Tennessee Acad. Sci. 34:66-—72. Estes, D. 2004. Noteworthy collections: Middle Tennessee. Castanea 69:69-74, Estes, D. and E.W.Cuester. 2001. Arenaria lanuginosa rediscovered and a new county record for Apios priceana in Tennessee. J. Tennessee Acad. Sci. 76:16. (Abstract). FENNEMAN, N.M. 1938. Physiography of eastern United States. McGraw Hill Book Company, New York, New York FerNnacd, M.L. 1950.Gray’s manual of botany. Eighth edition. American Book Company, New York, New York. Fiora OF NortH America EpiroriAL Committee (eds.). 1993. Flora of North America north of ESTES, FLORA OF GILES COUNTY, TENNESSEE 2387 Mexico.Volume 2.Pteridophytes and | Oxford University Press, New York, New York. Frora of NortH America EpitoriaL Committee (eds.). 1997. Flora of North America north of Mexico. Volume 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford University Press, New York, New York. Fiora oF NortH America EprroriAL Committee (eds.). 2000. Flora of North America north of Mexico.Volume 22.Magnoliophyta:Alismatidae, Arecidae, Commelinidae (in part),and Zingiberidae. Oxford University Press, New York, New York. Flora of NortH America Eprtoria. Commirtee (eds.). 2002a. Flora of North America north of Mexico. Volume 23. Magnoliophyta: Commelinidae (in part): Cyperaceae. Oxford Uni- versity Press, New York, New York. Fora of NortH America Eprroriat Committee (eds.). 2002b. Flora of North America north of Mexico. Volume 26. Magnoliophyta: Liliidae: Liliales and Orchidales. Oxford University Press, New York, New York. Fiora of NortH America EprroriAL Committee (eds.). 2003. Flora of North America north of Mexico. Volume 4. Caryophyllidae, Part 1.Oxford University Press, New York, New York. Gteason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. Second edition. The New York Botanical Garden, Bronx, New York. GrirritH, G.E., JM. Omernik, and S. Azeveoo. 1998. Ecoregions of Tennessee. 1:940 000. Albers equal area projection. U.S. Geological Survey, Reston, Virginia. Isecy,D.1990.Vascular flora of the southeastern United States. Volume 3, Part 2.Leguminosae (Fabaceae). The University of North Carolina Press, Chapel Hill, North Carolina. Meotey, M.E., H. Bryan, J. MacGrecor, and J.W. Tuierer. 1985.Achyranthes japonica (Miq.) Nakai (Amaranthaceae) in Kentucky and West Virginia: New to North America. Sida 11: 92-95 Miter, R.A., W.D. Harveman, D.S. FuLterton, C.R. Sykes, and R.K. Garman (eds.). 1966. Geologic map of Tennessee. West-Central Sheet. Tennessee Department of Conservation, Divi- sion of Geology, Nashville, Tennessee. Murrett, Z.E.and B.E.Worrorp. 1987. Floristics and phytogeography of Big Frog Mountain, Polk County, Tennessee. Castanea 52:262-290. QuarterMaN, E.and R.L. Powe. 1978. Potential ecological/geological natural landmarks on the Interior Low Plateaus. United States Department of the Interior, Washington, Dis- trict of Columbia. Ravrorb, A.E., H.E. Antes, and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas. The University of North Carolina Press, Chapel Hill, North Carolina. SHANKS, R.E. 1958. Floristic regions of Tennessee. J. Tennessee Acad. Sci. 33:195-210. Smatt, J.K. 1933. Manual of the southeastern flora. The University of North Carolina Press, Chapel Hill, North Carolina. SMITH, S.G. 2001. Taxonomic innovations in North American Eleocharis (Cyperaceae). Novon 11:241-257. 2388 BRIT.ORG/SIDA 21(4) Tennessee NATURAL HeriTace Procram. 1999, Tennessee rare plant list. Division of Natural Heri- tage, Tennessee Department of Environment and Conservation, Nashville, Tennessee. Tennessee NATURAL HERITAGE PROGRAM. 2003. Tennessee rare plant list. Division of Natural Heri- tage, Tennessee Department of Environment and Conservation, Nashville, Tennessee. True, J.C., JF. Campsett, and E.P. Davis. 1968. Soil survey of Giles County, Tennessee. United States Department of Agriculture Soil Conservation Service, Washington, District of Columbia. United States Fish AND Witouire Service. 1993. Recovery plan for Price’s Potato-Bean (Apios priceana). Jackson, Mississippi. Unite States FisH AND WILDLIFE Service. 1999. Recovery plan for Helianthus eggertii (Eggert’s Sunflower). Atlanta, Georgia. USDA (Uniteb States DeparTMeNT oF Acricutture). 2000. Natural Resources Conservation Ser- vice. The PLANTS patasase (http://plants.usda.gov/plants/). National Plant Data Center, Baton Rouge, Louisiana [as accessed December 29, 2003]. Worrorb, B.E. and E.W. CHester. 2002. Guide to the trees, shrubs, and woody vines of Ten- nessee. The University of Tennessee Press, Knoxville, Tennessee. Worroro, B.E.and R. Krat. 1993. Checklist of the Tennessee vascular plants. Sida, Bot. Misc 10. Botanical Research Institute of Texas, Fort Worth, Texas. YATSKIEVYCH, G. 1999. Steyermark’s flora of Missouri. Volume 1. Revised edition. The Missouri Botanical Garden Press, St. Louis, Missouri THE VASCULAR FLORA OF MONTGOMERY COUNTY, ARKANSAS Travis D. Marsico! University f Arkansas Herbarium (UARK) Bi arch Center 141 le Arkansas 72701, U.S.A. S) 4 = Fayettevi ABSTRACT A floristic inventory in Montgomery County, Arkansas, documented 1, i. vascular plant taxa. _— nine taxa (5% of county total) in Montgomery County are considered to be of conser cern and are tracked by the Arkansas Natural Heritage Commission. One eee and Sou taxa (12% of county total) were introduced, 21 (2% of county total) of which are considered invasive. The families with the most taxa represented in the flora are Asteraceae (144), Poaceae (120), Cyperaceae (82), Fabaceae (75), and Rosaceae (38). Elatine triandra Schkuhr represents a species never before collected in Arkansas. A single federally ee species, Ptilimnium nodosum (Rose) Mathias, was noted. Hydrilla verticillata (L-£.) Royle, a federally listed noxious weed was collected. Montgom- ery County has one of the highest oe of sensitive species in Arkansas, but has fewer than 20% of the State’s known alien species, probably due to its eke diversity of rare habitats and a rela- tively intact landscape. With 1,110 taxa, Montgomery County is the sixth best-collected county in Arkansas. However, since most Arkansas counties have fewer than 800 documented taxa, there is a A 4 clear need for increased floristic work if the species that grow in Arkansas are to be known. RESUMEN © evé a cabo un inventario floristico en el condado de Montgomery, Arkansas, que proporciono una lista de L110 taxa ge Benes vasculares. Cmeuenla y mueve taxa (5% del total) del condado de r la Comision Herencia Natural Mont de ieee ue Natural Medige Cc OT ree treinta ay cuatro taxa ee del total it le condado) son introducidos y 21 (2% del total de familias con un mayor numero de taxa oe re eon en la flora son aes ( 144), Poaceae a Cyperaceae (82), Fabaceae (75), y Rosaceae (38). Eohite anare Sara es una sap ates que fue colectada por primera vez en Arkansas. F peligro, Ptilimnium nodosum (Rose) Mathias. Se colecto Hydrilla verticillata (L. f. ) Royle, una especie nociva federal mencionada. El condado de Montgomery tiene una de las mas elevadas Be ana de especies sensibles de Arkansas, pero tiene menos del 20% de las especies introducidas en el estado, probablemente debido al gran numero de habitats raros y un paisaje ee intacto. Con L110 taxa, el condado de Montgomery es el sexto condado m eee de Arkansas. Sin embargo, puesto que la mayoria de los condados de Arkansas tienen menos de 800 taxa documentados, hay una necesidad de mas estudios floristicos con el fin de conocer las especies que crecen en Arkansas. ‘Present Address: Department of Biological Sciences, Galvin Life Science 107, University of Notre Dame, Notre Dame, Indiana 46556, U.S.A, tmarsico@nd.edu. SIDA 21(4): 2389 — 2423. 2005 2390 BRIT.ORG/SIDA 21(4 INTRODUCTION Floristic studies have long provided botanists with baseline knowledge of plant species distribution and habitat preference, so proportionately more is known from locations that have had extensive inventories (Duncan 1953). Descriptive studies are needed before subsequent ecological endeavors can be conducted. No systematic inventory of the Ouachita Mountains’ flora has been conducted to date with the exception of Hot Springs National Park (Palmer 1926; Scully 1937, 1941, 1942). In addition, certain areas of botanical interest in the Quachitas (e.g., Rich Mountain, Mount Magazine, and Albert Pike) have received much attention by botanists, but very frequently for their charismatic plant species, and a comprehensive flora accompanied by vouchered specimens is lacking, This study is the first one of its scale in the physiographic region of the Oua- chita Mountains of Arkansas. Geography, Geology, and Soils Montgomery County is located in west central Arkansas in the Central Oua- chita Mountains (Fig. 1). The county covers 2,023 km? (781 mi2), and is charac- terized by east-west trending ranges of the Ouachita Mountains, including the Fourche, Caddo, and Cossatot mountains. The Fourche Mountains are located in the northern part of the county. The central portion of the county consists of a large basin that contains the Ouachita and Caddo rivers. The Caddo Moun- tains are south of the basin, and southwest of the Caddo Mountains are the Cossatot Mountains. Elevation in Montgomery County ranges from a low of 49 m in the very southeast corner of the county along Sugarloaf Creek to a high of 673 mat Slatington Mountain in the southwest (490 to 2209 ft). Geologically the area is old and the exposed rock and soils are varied. The oldest rock in the state, which dates from the Cambrian and lower Ordovician periods, is Collier Shale, which is partially exposed in Montgomery County (Braden 1999), The east-west geology of the Ouachita Mountains displays the rock in decreasing age both north and south of the center due to anticline fold- ing during the Pennsylvanian Period. All the sedimentary rocks in the Ouachita Mountains were deposited be- lore the tectonic events of the late Pennsylvanian. Each rock type was deposited ina horizontal stratum, but was subsequently twisted and folded into its present shape (Palmer 1926). Thrust faulting and solange due to conbne aka) oon gave the Ouachita Mountains their compressed anticli (Snidet 1982). The Ouachita fold belt extends from southwest Alabama north igus Arkansas, and southwest through Oklahoma, Texas, and into Mexico. The 2,100 km course of the Ouachitas is only exposed for 500 km (Flawn 1959). The rocks are shale, sandstone, quartzite, conglomerate, and novaculite (Palmer 1926). No- vaculite, the slightly metamorphosed product of chert (Guccione 1993), is mined for high-quality whetstone in the area, and is interesting both geologically and 2391 MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS dIHSU3NMO CNV (ees ajeus pue suospueg Fe : Seutay ee ; SunseAOn AE eer Co = 2a ajeug ee auospues “ff, A901039 J savou , puebe7] | y, Arkansas. fa Fic. 1. Geol 2392 BRIT.ORG/SIDA 21(4) botanically. Additionally, novaculite glades are habitats unique to the Ouachita Mountains. Substantially due to its geology, Montgomery County is rich in re- gional endemics of both plants and animals. The area has a great diversity of habitats in close proximity to one another (Braun 1950) due to its rough and varied topography. Soil orders in the region are Entisols and Ultisols. Entisols are young soils with little or no evidence of developed soil horizons. In the Ouachita Moun- tains they form in the vicinity of stream courses where erosion is extreme and equals or exceeds soil formation. Ultisols are old soils that form in warm, hu- mid climates with a seasonal dry period under forest vegetation. Ultisols are the dominant soils in Montgomery County (Soil Survey Staff 1998, 1999). Soils in the area are commonly thin and in many places, including steep slopes and glades, the parent geologic material has far greater influence than that of the soil. Climate The climate of Montgomery County and the central Ouachita Mountains is broadly described as humid sub-tropical (Bailey 1995). The climate of the state of Arkansas is influenced by its latitude between 33 and 37 degrees north, the prevailing westerlies, polar fronts from the continent, and Gulf of Mexico mois- ture (Baldwin 1984). The following climate data were compiled from the Na- tional Oceanic and Atmospheric Administration (NOAA) observations col- lected in Mount Ida, Arkansas, in central Montgomery County, from 1931 through 2000, but represent 66 years of observation due to missing values from 1948, 1949, 1950, and 1954. The average annual high temperature is 23°C (73°F), the average annual low temperature is 8°C (47°F), and the overall average an- nual temperature is 15°C (60°F). Temperatures have ranged from an all-time high of 47°C (116°F) on August 10, 1936, toa low of -29°C (-21°F) on February 2, 1951. Even though extremes in precipitation or temperature occur, they are rare. The average hottest day for a given year is 39°C (103°F), but the most common value in the 66 years of data collection was 38°C (100°F). The average coldest temperature for a year is -15°C (5°F), and the most common value was -13°C (8°F). The average period without a freeze (growing-season) is typically between 200 and 240 days in the Ouachita Mountains province (USDA 1981). The area receives some of the highest rainfall in Arkansas (Reinhold 1969) with an annual average of 141 cm (55 in.). The range of annual rainfall, how- ever, is incredibly varied, with a maximum of 213 cm (84 in.) recorded in 1945 anda minimum of 83cm (33 in.) recorded in 1936. The Ouachita Mountains are high enough to draw more rainfall from passing storm systems than other natu- ral regions of Arkansas. Precipitation is seasonal with the spring months of March, April, and May commonly being wettest and typical dry times in July through September. Snow is limited in the region, and one out of every five years lacked snow entirely. The average annual snowfall for the area is 12 cm (5 in). MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS 2393 The greatest yearly snowfall recorded was 47 cm (19 in) in 1988. Ice storms are uncommon, but may be severe. In December 2000, an ice storm struck the re- gion and dramatically opened the forest canopy. There were long-term changes in vegetation structure due to the severity of that storm. Anthropogenic Influence Montgomery County and the greater Ouachita Mountain region were occupied by early Americans in the Dalton cultural period in the early Holocene for hunt- ing grounds and short-term settlements. There is evidence that even in rugged areas of the Cossatot Mountains by the late Caddo period (15 Century), hunt- ing and gathering was supplemented by small plots of maize, beans, and squash (Early 2000). Although First Nations peoples mined and farmed small plots, their impact to the landscape in the area appears small compared to that after European settlement. European descendants typically from Mississippi, Alabama, and Georgia settled in the Ouachita Mountains in the 19'" Century. They were predominately of English or Scotch-Irish heritage, and lived subsistence lives with only small farms, often surrounded by virgin woodland (Smith 1986). When mechanized logging arrived in the Ouachitas around 1900, the vegetation began to change rapidly as large tracts of shortleaf pine timber were logged. Virtually no virgin timber remained by 1950. The Arkansas National Forest was set aside in 1907 (changed to the Ouachita National Forest in 1926). Sustained yield forestry be- gan to replace “cut out and get out” practices in the 1920s, but as a valuable timber resource the areas within the Ouachita National Forest experienced heavy logging. In the 1970s Weyerhaeuser Company began intensively manag- ing its stands as monocultures of loblolly pine. The Ouachita National Forest began to use even-age management as well (Smith 1986). Such management regimes are now unacceptable on public lands. It is important to note that al- though none of the land is in its pre-settlement state, and all of it is managed, the majority of Montgomery County retains the landscape’s original character, and allows for a great number of native species to persist or thrive. The Oua- chita National Forest and Montgomery County will likely keep their semi-natu- ral character long into the future. Ever since the warming and drying after the most recent ice age, plant com- munities in the region have experienced human induced fires. Though Euro- American settlers in the South were casual in their feelings toward fire, national policy prevented fire in an effort to prevent damage to property. However, U5. Forest Service policy in the South had always contained provisions for allow- ing fire, and interest in using fire as a management tool increased (Pyne 1982). Currently, the Ouachita National Forest prescribes burns to limit potential for catastrophic fire and improve wildlife habitat (http://www.fs.fed.us/r8/ ouachita/fire/fire_management%20.shtml). 2394 BRIT.ORG/SIDA 21(4) Although ecosystem degradation by land management practices is always a concern, most of Montgomery County is in federal land holding, and ecosys- tem destruction from development is of little concern. A potential problem, now, and the second most important cause for the decline of imperiled species is the introduction of non-native, invasive species (Stein et al. 2000). Common exotic invasive species in Montgomery County include Elaeagnus umbellata, Lespe- deza cuneata, Ligustrum sinense, Lonicera japonica, and Microstegium vimineum. Ecological Systems The Ecological Systems Database (NatureServe 2003) was used as a framework for classifying ten plant communities in Montgomery County. The communi- ties are defined on a meso-scale, allowing for patterns of ecological variability while remaining recognizable to guide conservation and land managers’ needs (Comer et al. 2003). NatureServe lacked anthropogenically managed or created communities that did not fall under the categories of “natural” or “semi-natu- ral.” These include easily recognizable areas such as pastures, roadsides, and abandoned pits or quarries. A distinct community dominated by Pinus taeda was not included for the Ouachita Mountains. The only natural community that was lacking from the published list was the Shale Glade Ecological Sys- tem. Shale glades are important botanically in Montgomery County because they are areas where unique plant assemblages form and include species found nowhere else in the county. (PIN) Ozark-Ouachita Shortleaf Pine-Oak Forest and Woodland.—The Ozark-Ouachita Shortleaf Pine-Oak Forest and Woodland covers the largest land area in Montgomery County. It is classified as a natural or semi-natural, vegetated, and upland matrix. The thread that ties this variable system together is the presence and often dominance of Pinus echinata. The hardwood compo- nents, dominated by various Quercus species, vary with slope, aspect, and mois- ture conditions (Dale & Ware 1999). There are other hardwood canopy species in the system such as Carya spp. and Prunus serotina, but they are less abun- dant than oaks. The canopy ranges from completely closed to more commonly open with as little as 40% canopy cover. The system covers a wide range of to- pography from level to steep slopes, most aspects, and is not tied to a specific topographic feature (e.g., streams) or geology. Understory species include Vaccinium spp., Solidago spp., Monarda spp., and Schizachyrium scoparium as dominants. (LOB) Ouachita Mountain Planted Loblolly Pine Forest.—This ecological system is not listed by NatureServe (2003) for Montgomery County probably because Pinus taeda is thought to be exotic to the Ouachita Mountains. The loblolly pine system is similar to the shortleaf pine system in the Ouachitas except that it is dominated by Pinus taeda rather than Pinus echinata. The com- MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS 2395 munity is located only in the southeastern portion of the county where the topography is less rugged and slopes are less steep than in other areas. Com- mon herbaceous species include Asclepias variegata, Lactuca canadensis, Pseudognaphalium helleri, Clitoria mariana, and Rhexia mariana. Except for its location in the Ouachita Mountains, the community matches closely the Pinus taeda forest alliance of the West Gulf Coastal Plain Pine-Hardwood For- est (NatureServe 2003). Sites in Montgomery County range from obvious plan- tations with trees in straight rows to a more natural system managed by fire. Although all sites may represent plantings in this area, the system (at least non- plantation sites) can follow the semi-natural classification of the Ozark-Oua- chita Shortleaf Pine-Oak Forest and Woodland with a different dominant canopy species, or the West Gulf Coastal Plain Pine-Hardwood Forest in a more northern location. (MES) Ozark-Ouachita Mesic Hardwood Forest.—The Ozark-Ouachita Mesic Hardwood Forest is classified as a natural or semi-natural vegetated up- land small patch system. This community may be found on low, north-facing slopes and along river terraces in areas that are not distinctly riparian. Quercus alba and Q. rubra are common oak dominants, but the classic mesic species are Fagus grandifolia and Acer barbatum. Tilia americana is another canopy spe- cies, and Asimina triloba and Magnolia tripetala may be found in the under- story. In the Crystal Campground area, the mesic forest includes a population of Pinus strobus, planted for timber in 1910, but subsequently naturalized, and it isthe only reproducing population known in Arkansas. The herbaceous layer of this community contains a wide range of spring ephemeral species such as Trillium recurvatum, Cypripedium kentuckiensis, and Podophyllum peltatum. Mesic forest habitat provides greater moisture to vegetation thus supporting different species than the shortleaf pine-oak forest and woodland system. RIP) Ozark-Ouachita Riparian —The Ozark-Ouachita Riparian commu- nity is variable in vegetation, but has one main topographic feature that ties the system together—streams. The system is classified as natural or semi-natu- ral, vegetated, and upland. The spatial pattern is linear. Canopy species may vary, but typically include Liquidambar styraciflua and Platanus occidentalis as canopy dominants. Acer spp. and various Quercus spp. are also canopy spe- cies in the riparian ecosystem. Betula nigra occurs infrequently. The understory and shrub layers often consist of Lindera benzoin, Alnus serrulata, Hamamelis vernalis, Carpinus caroliniana, and Ostrya virginiana. The herbaceous layer is diverse and ly consists of Festuca subverticillata, Osmorhiza longistylis, Galium aparine, Viola pubescens, and Elymus virginicus. Certain riparian sites also include the Ouachita Mountain endemic, Hydrophyllum brownei. The ri- parian system does not often include Fagus grandifolia in its canopy. The ri- parian zone in Montgomery County is typically found from the immediate riverbank through a system of periodically flash flooded terraces. The size of ey — 2396 BRIT.ORG/SIDA 21(4) the stream dictates the distance away from the stream that the riparian com- munity is encountered. The substrate consists of soils that are rich and well- drained and often with abundant gravel. (SEE) Ouachita Mountain Forested Seep.—The Ouachita Mountain Forested Seep community is characterized as a natural, small patch, vegetated wetland. All seepage areas have water coming from below the ground surface. Seeps may occur at the headwaters of streams or along riparian areas. They are saturated or very moist throughout the year. The canopy may be dominated by Liquidam- bar styraciflua, Quercus alba, Acer rubrum, and Magnolia tripetala, which is also common in the understory. The coverage of the canopy is variable from fully covered to quite open. However, due to the soft substrate, which allowed for easy uprooting, much of the canopy in many seeps in Montgomery County was dramatically opened by an ice storm in December 2000. Subcanopy spe- cies commonly encountered are Magnolia tripetala, Ilex opaca, Carpinus caroliniana, and Corylus americana. Aside from the saturated soil, the herba- ceous layer provides a distinctive sign of a forested seep. Ferns are abundant and diverse and include Osmunda cinnamomea, Osmunda regalis, Athyrium filix-femina, Onoclea sensibilis, and others. (NOV) Ouachita Novaculite Glade and Woodland.—The Ouachita Novacu- lite Glade and Woodland system is defined as a small patch of natural occur- rence that is vegetated and upland. The diagnostic feature for the community is novaculite geology. The system is found from 450-640 m (1476-2100 ft.) in el- evation and is a mosaic of open glades, outcrops, and woodlands. Dominant species include Quercus stellata, Quercus marilandica, Quercus rubra,and Carya texana. The endemic Quercus acerifolia is found only in this and the Ouachita Montane Oak Forest system. A common and often distinctive member in the subcanopy is Ptelea trifoliata. The herbaceous layer is dense with grass species including Bromus spp., Danthonia spicata, Dichanthelium spp., and Schizachyrium scoparium. Ambrosia arte misiifolia, Helianthus divaricatus,and Helianthus hirsutus are other common associates. Trees are often stunted and gnarly due to drought, fire, wind, and ice, all of which are thought to play im- portant roles in the maintenance of this system. The Ouachita Montane Oak Forest is a similar community to the Novacu- lite Glade system except that it lacks novaculite substrate. The inclusion of this system under the Novaculite heading is appropriate here because the montane oak forest has a limited extent in Montgomery County. It is only found in high- elevation areas in the northwest part of the county. Although the geology dif- fers, vegetation is remarkably similar in both high-elevation communities, and does not warrant a separate community designation in Montgomery County. (SHA) Ouachita Mountain Shale Glade.—The Ouachita Mountain Shale Glade community Is characterized as a small to large patch, natural or semi- MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS 2397 natural vegetated, upland system. The soil is very thin and shale (the dominant substrate) is often exposed at the surface. This ecosystem is found on level or slightly sloped topography in the basin regions of the Ouachita Mountains. There is often no tree canopy associated with this system, but Juniperus virginiana and Carya texana have become common canopy species. There is little shrub layer, as the system is dominated by a diverse array of herbs and grasses. The shale substrate acts as a fragipan, creating very wet surface condi- tions in the early spring that dry throughout the summer, when the system appears nearly barren. This hydro-xeric phenomenon characterizes the shale glade. Herbaceous species include Ialinum calycinum and Dodecatheon meadia, as well as many members of the Asteraceae including Coreopsis grandiflora, Silphium laciniatum, and Solidago spp. (ROA) Ouachita Mountain Upland Herbaceous with Regular Interval Hu- man-Induced Disturbance.—This community includes those habitats that are managed by,at minimum, yearly grazing or mowing: roadsides, roadside ditches, pastures, and cemeteries. These are linear, small patch, or large patch, anthro- pogenic upland vegetated systems. They are typically without a tree or shrub layer and consist mostly of grasses and other herbaceous species such as Daucus carota, Bidens spp., Trifolium campestre, Paspalum spp., Loliumspp.,and Tridens flavus. The species growing in these habitats are adapted to full sun and a range of moisture conditions. Although not considered by NatureServe, this system is important to the categorization of the plant species in the Ouachita Moun- tains. Pastures and roadsides favor grass species, which tolerate mowing/graz- ing, serve as entry points and corridors for invasive species, represent novel habitat for natives, and make up a large land area. (WEE) Ouachita Mountain Upland Herbaceous-Shrubby with Single Ma- jor Human-Induced Disturbance.—This community is common along aban- doned Forest Service roadways, but also is used for any land not actively man- aged, suchas abandoned lots, pits, or mines. It is a linear or small patch vegetated, upland anthropogenic system. Though much less important in terms of land area covered than Regular Interval Disturbance system, it is distinct from it. The main difference between the ecosystems is type of disturbance. The Regu- lar Interval Disturbance system has regular, at least annual mechanical distur- bance, whereas this weedy community begins with a single major disturbance only. After this usually vegetation-voiding initial event, colonization and suc- cession occur undisturbed, unless aided by plantings to reduce erosion. This system is characterized by weedy or early successional herbaceous and shrub species and represents an ever-changing continuum from unvegetated bare ground to late successional stages. These habitats are abundant with non-na- tive species such as Ligustrum sinense and Lespedeza cuneata. Other abundant early colonizers include Ambrosia spp. and Acalypha virginica. 2398 BRIT.ORG/SIDA 21(4) (WET) Ouachita Mountain Human Created/Maintained Still Water Wet- land and Gravel Bar.—This system is a variable system that includes anthropo- genic lakes and ponds (none of which are natural in the Ouachita Mountain landscape), a beaver created upland marsh, and natural gravel bars of rivers and streams. These are wetland systems that can often be described as early successional. Regulated lakes and ponds in the region have a fluctuating shore- line allowing for wetland plant growth and colonization that follows the water level. Stream gravel bars, though natural, mimic this pattern due to rapid water level changes in streams and rivers. Gravel bars also experience vegetation-void- ing disturbance during flash floods. These systems are dominated by a mix of wetland herbaceous and shrub species and weedy early successional plants. True aquatic species include Nymphaea odorata, Potamogeton spp., and Utricularia gibba. Examples of emergent or terrestrial species in this system are Justicia americana, Xanthium strumarium, and Cleome hassleriana. Richardson Bot- toms, a beaver created upland marsh, though a unique community in the Oua- chita Mountains, fits under this category for ecological system description. METHODS Voucher collections were made from August 2001 through October 2003. Sev- eral primary collecting sites were established by conducting pilot searches early in the study, reviewing topographic maps, and consulting Ouachita National Forest and Arkansas Natural Heritage Commission unpublished document accounts (Orzell 1985; Bates 1993; Robison & Marsh undated). The primary sites are representative of the diversity of habitats found within the political bound- ary of Montgomery County. At primary sites, collections were made in each phase (spring, summer, fall) of the growing season. Auxiliary sites were visited only once or twice throughout the study period. Vouchers collected according to standard collecting methodology, and material was compared to UARK specimens and keyed with pertinent floras (e.g, Radford et al. 1968; Smith 1994a; Diggs et al. 1999; Yatskievych 1999) for the majority of the identifications. A ive voucher specimen for each taxon was deposited at the University of Arkansas Herbarium (UARK). Specimens were assigned a community type from which they were collected in order to providea high resolution of species distribution within Montgomery County. After all collections were identified, Smith (1988) by way of the Texas AWM Bioinformatics Working Group website thttp://wwwesdl tamu.edu/FLORA / cgi/kartesz_ar_ page_click?county=Montgomery) was consulted for species collected from Montgomery County by previous investigators. Taxa that were found on Smith’s list that were not collected in the current field study (2001- 2003) were noted. Subsequently, a search was conducted for the listed specimens Identifications of specimens found at UARK were verified, and those at other MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS 2399 herbaria were accepted as correctly identified without review. Collections made by EL. Hardcastle, CS. Reid, D.X Williams, and CT. Witsell in Montgomery County since 1988 were also reviewed. RESULTS Plant collections resulted in 1,013 taxa at and below the species level, +74 gen- era, and 139 families. Further investigation led to the discovery of additional species collected by others for Montgomery County. These additions bring the total known taxa for Montgomery County to 1,110 species and subspecific as- signations, 506 genera, and 143 families. The plant families with the greatest number of taxa are as follows: Asteraceae (144), Poaceae (120), Cyperaceae (82), Fabaceae (75), and Rosaceae (38). Forty-three taxa that had previously been re- ported from Montgomery County could not be supported by voucher speci- mens, and have subsequently been excluded from the flora. The specimens ei- ther never existed (a verbal report only), could not be found, or have been annotated as some other taxon. Fieldwork resulted in the addition of one species not previously known to occur in Arkansas: Elatine triandra Schkuhr. Elatine triandra is native to the United States and the collection represents a range expansion from its known distribution. This wetland species was collected on the muddy shores of a re- cently constructed lake southwest of Mount Ida. Fifty-nine of the 1,110 taxa in Montgomery County are tracked by the Ar- kansas Natural Heritage Commission. Species of special concern, therefore, make up 5% of the flora in Montgomery County. One population of the listed, federally endangered species Ptilimnium nodosum (Rose) Mathias was located. The voucher is a photograph (TD. Marsico 3247, VARK), since a permit to col- lect endangered species was not obtained. The location of the population had been studied by Hardcastle and Williams (2000), and as stated in their report, thousands of individuals were observed. Non-native species have recently gained the attention of land managers. One hundred and thirty-four (12%) taxa of the 1,110 of the Montgomery County flora are represented by species categorized as non-native. Of those, 21 are con- sidered invasive. Montgomery County also tains a native invasive, Baccharis halimifolia L. Invasive status follows the working list from the Rare and Inva- sive Plants of Arkansas Conference (RIPAC) in October 2003 (Arkansas Native Plant Society 2003). One of the taxa collected is a federally listed noxious weed, Hydrilla verticillata (L£.) Royle. Collections made during the course of the study revealed that the riparian and roadside communities are home to the greatest number of species, whereas the planted loblolly pine forest, forested seeps, and shale glades had the fewest species. Also of note is the three communities with the greatest percentage of introduced taxa are those that are human created and/or dominated (Table 1). 2400 BRIT.ORG/SIDA 21(4) Taste 1. Summary of plant collections by community type in Montgomery County. Because only Marsico collections were assigned to a community, 1013 taxa are used in “total” calculations. Com- munity codes are lob—Planted loblolly pine forest, mes—Mesic hardwood forest, nov—Novacu- lite glade and woodland and montane oak forest, pin—Shortleaf pine- iis ee and woodland, rip—Riparian, ic ae roadside ditch, ee and cemetery, see ted seep, sha—Shale glade, wee—Weed h as abandoned ONF roadways, abandoned lots, pits, or mines, wet— Wetlands nelading iskes. sends mudflats, gravel bars, and a marsh. Number of Number of Total Taxa Percent of County — Percentin Native Taxa Introduced Taxa (N+I)=T Total Marsico abitat (N) I) Collections Introduced (T/1013)*100 (I/T)*100 LOB 9] 4 95 9% 4% MES 283 14 297 29% 5% NOV 196 15 211 21% 8% PIN 289 14 303 30% 5% RIP 418 36 454 45% 9% ROA 355 68 423 42% 19% SEE 167 4 171 17% 2% SHA 145 13 158 16% 9% WEE 149 55: 202 20% 36% WET 330 46 376 37% 14% DISCUSSION The 1,110 taxa documented for Montgomery County places it sixth among the best collected counties of the state. While it is important to remember that bo- tanical diversity relates to land area, topographic diversity, and land use prac- tice, many places in the state of Arkansas are still not well known botanically. The best collected county is Washington with 1,355 taxa, and the worst is Woo- druff with 347 (Smith 1994b). The top five best collected counties, except for Pulaski, are in the Ozark Plateaus Natural Division. All counties in Arkansas that have had systematic botanical inventories conducted have over 900 known taxa; therefore, it is probable that all counties in the state have this potential. Still, 65% of Arkansas counties have fewer than 800 documented taxa. This underscores the need for increased floristic work if the flora of Arkansas is to be fully understood. With 59 species of special concern, Montgomery County has one of the highest proportions of sensitive plant species in Arkansas. When combined with animals, the Ouachita Mountains have a high level of endemism (Robison & Allen 1995). Contributing to the diversity, Montgomery County contains dis- junct species from the Ozark Plateaus and the nearby Gulf Coastal Plain. Also, shale and novaculite glades, forested seeps, and mesic and riparian communi- ties are hot-spots for rare species. The high number of rare habitats provides adequate conditions for the plants that are specific for those habitats. MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS 2401 After habitat destruction, alien or introduced species are the second-great- est threat to imperiled species (Stein et al. 2000). Montgomery County has fewer than 20% of the known alien species in Arkansas, but over one-third of the to- tal native species known to occur in the state. While the flora of Arkansas is comprised of about 24% non-native taxa (Arkansas Vascular Flora Committee, pers. comm.), Montgomery County has only half that (12%). One possible rea- son for this difference is the relatively intact landscape of Montgomery County. Even though anthropogenic influence has been pronounced, human activities have not prevented the landscape from retaining many of its original qualities. For example, where there was once pine-dominated forest, this system remains today. Human population remains low, and agriculture is not a prominent force. In contrast, the state of Missouri has seen its numbers of introduced taxa in- crease by 35% from the early 1960s through the late 1990s. This is at least par- tially attributable to unprecedented environmental changes including a highly and continually disturbed landscape, favoring the predominate annual and biennial habit of introduced taxa (Yatskievych 1999). Therefore, it is not sur- prising that the vast majority of introduced species in Montgomery County are found in the three main human created or dominated landscapes of roadsides, wet areas, and abandoned roadways or lots. Over one-third of the introduced taxa collected in this study were gathered from the community classified as “weedy areas,” which by definition would include a high number of weeds. Overall, roadside and riparian habitats include the highest diversity of spe- cies in Montgomery County. This is due primarily to the widespread nature of each of these ities ina linear system. While roadsides and streamsides do not amount to a majority of the Montgomery County landscape, they cut through all other communities, allowing for high numbers of species exclusive to those communities al ide specie at oe out from other communities The planted loblolly pine forest, glades, and forested seeps have the lowest overall diversity oe due to the very low amount of land area each covers in the county. ANNOTATED CHECKLIST OF VASCULAR PLANT TAXA Taxa below all represent vouchers from Montgomery County, Arkansas. They are listed alphabetically by family, then alphabetically by genus, spectic epi- thet, and subspecific designation where appropriate. For all vouchers collected by LD. Marsico, following the Latin name is a collection number correspond- ing toa specimen deposited at UARK, the habitat codes for where the taxa were collected, whether the taxa are native (N) or introduced (D, and any other spe- cial designation such as a species of special concern (SC) or invasive (I*). Taxa listed that were not collected by the author have the Latin name followed by the collector of the voucher, his/her collection number, the herbarium in which the voucher is located, and its status as native or introduced. Habitat codes are 2402 BRIT.ORG/SIDA 21(4) as follows: lob—Planted loblolly pine forest, mes—Mesic hardwood forest, nov— Novaculite glade and woodland and montane oak forest, pin—Shortleaf pine- oak forest and woodland, rip—Riparian, roa—Roadside, roadside ditch, pasture, and cemetery, see—Forested seep, sha—Shale glade, wee—Weedy areas such as abandoned ONF roadways, abandoned lots, pits, or mines, wet—Wetlands in- cluding lakes, ponds, mudflats, gravel bars, and a marsh. All nomenclature fol- lows The PLANTS Database (USDA, NRCS 2004). ACANTH Dicliptera Dae (Pursh) Spreng., 4626, rip, N Justicia americana (L.) Vahl, 3138, pin, rip, roa, wet, N Ruellia humilis Nutt., 2642, nov, roa, sha, N p Ruellia pee: L.,4250, rip, see, N ERACEAE Acer barbatum Michx., 834, mes, N Acer leucoderme Small, 2754, nov, N, SC Acer negundo L., 3261, rip, r ee, N Acer rubrum L. vat. Pe (Hook. & Arn. ex Nutt.) Sarg., 4955, pin, rip, wet, N Acer rubrum L.var. rubrum, 5053, lob, mes, pin, rip, a, see, wet, N ane ae L., ae. rip, wet, N \ce var. saccharum, 5692, nov, rip, N AGAVACEAE Manfreda virginica (L.) Salisb. ex Rose, 3742, nov, roa, sha, N Yucca arkansana Trel.,4994, pin, roa, N Alismataceae Alisma subcordatum Raf, 5539, roa, wet, N Echinodorus cordifolius (L.) Griseb., 3295, wet, N — oe Michx. var. graminea, 5485, Coicne ei lia Willd., 4027, roa, N Sagittaria platyphylla ean J.G.Sm., 5771, wet, N Amaranthaceae Amaranthus retroflexus L., R. DB. Thomas, 131846, LU,| Amaranthus spinosus L.,4414, wet, N lresine rhizomatosa Standl., 3877, rip, wee, N Anacardiaceae Rhus aromatica Ait. var. aromatica, 3947, nov, pin, , Rhus copallinum L., 5997, nov, pin, roa, wee, N Rhus glabra L., 5534, pin, roa, wee, Toxicodendron pubescens P. Mill., 5061, pin, N Toxicodendron radicans (L.) Kuntze, 2651, lob, nov, pin, rip, roa, see, N Annon pense ak .) Dunal, 5447, mes, nov, rip, N Apiaceae Chaerophyllum procumbens (L.) Crantz var. procumbens,4727,mes, nov, rip, roa, wee, wet, N Chaerophyllum tainturieri Hook. var. tainturieri, Cicuta pan L., 2308, rip, roa, N inadensis (L.) DC., 5456, mes, pin, ans Daucus caret ot 5105, roa, wee, wet, | ia bulbosa (Michx.) Nutt. 540, rip, N ee jum prostratum Nutt. ex DC., 5473, lob, rip, t Eryngium yuccifolium Michx., 3580, rip, roa, N Hydrocotyle prolifera Kellogg, 4068, rip, wet, N Hydrocotyle verticillata Thunb. var. verticillata, 3890, rip, Osmorhiza longistylis (Torr.) DC., 1626, mes, nov, rip, see, N Oxypolis rigidior (L.) Raf, 3572, mes, pin, rip, N Ptilimnium capillaceum (Michx.) Raf., 5478, wet, N Ptilimnium nodosum (Rose) Mathias, 3247, wet, N, SC, Federally Endangered Ptilimnium nuttallii (DC) Britt. 2562, pin, sha, wee, wet, panicuta pea L., 5503, lob, mes, nov, pin, e, sha, wet, a odorata (Raf.) K.M.Pryer & LR. Phillippe, Sanicula smallii Bickn., 1603, me Spermolepis inermis (Nutt.ex DC. ) Mathias & Con- stance, 2601, wee, N MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS Taenidia integerrima (L.) Drude, 1744, mes, rip, N Thaspium barbinode (Michx.) Nutt. 3436, mes, rip, N Thaspium trifoliatum (L.) Gray, 2514, mes, rip, see, By rev [al if VETiofs (TU ds.) Link, 2590, pin, wee, | Tepoeaine aethusae Nutt.ex DC., 5452, pin, rip, N , Zizia aurea (L.) W.D.J.Koch, 4984, pin, rip, roa, wet, N Apocynaceae Amsonia ciliata Walt. var. ciliata, E.Sundell, 11139, U Amsonia hubrichtii Woods., 3153, rip, sha, wet, N, SC Amsonia tabernaemontana Walt.,4861,mes, Nov, bi Ls eee roa,N Trace! spend It.) Gray, 5521, lob, pin, rip, roa, sha, wee, wet, N Vinca major L., 4762, pin, roa, * Aquifoliaceae llex ambigua (Michx.) Torr, ae mes, N llex decidua Walt., 3041, rip, roa, llex longipes Chapman ex Trel., os mes, rip, N, SC [lex opaca Ait. var. opaca, 5467.5, mes, rip, see, N llex vomnitoria Ait., 5733, lob, mes, nov, pin, N Araceae Arisaema dracontium (L.) Schott, 5506, rip, wet, N Arisaema triphyllum (L.) Schott, 1831,mes, rip, see, Araliaceae Aralia spinosa L.,4197, mes, wee, N Aristolochiaceae Aristolochia serpentaria L., 3350, mes, pin, N MUSOOONG a, Sims, 5597, rip,N Asclepiadac Asclepias ea ia Michx., 2392, sha, N Asclepias quadrifolia Jacq., 1602, mes, pin, see, N Asclepias tuberosa L. ssp. interior Woods., 2248, nov, roa, sha, N Asclepias variegata L.,5492,lob,mes, pin, roa, see sclepias verticillata L., 3188, nov, roa, N Matelea baldwyniana (Sweet) Woods., 2748, nov, rip, roa, sha, N 2403 Matelea decipiens (Alexander) Woods., 334, mes, N Matelea gonocarpos (Walt.) Shinners, 2360, rip, N Aspleniaceae Asplenium platyneuron ee B.S.P, 5509, lob, mes nov, pin, rip, roa, sha, w Asplenium bradleyi D.C.Eat., i. eee 100549, U ARK,N Asplenium trichomanes L.,J.L. Roberts, 254, UARK, N Asteraceae Achillea millefolium L., 2394, roa, sha, N Ageratina altissima (L.) King & H.E. Robins. var. altissima, 4534, mes, nov, pin, rip, N Ambrosia artemisiifolia L.4109, nov, pin, roa, wee N Ambrosia bidentata Michx., 4287, roa, wee, N Ambrosia trifida L., 4161, rip, roa, wee, N Antennaria parlinii Fern.ssp. fallax (Greene) Bayer & Stebbins, 984, pin, N Antennaria parlinii Fern. ssp. parlinii, 717, mes, N Antennaria plantaginifolia (L.) Richards., 4670, mes, nov, pin, rip, sha, N ossum plantagineum Raf., 2395, pin, roa, a,N pe integrifolium (Michx.) Nutt., 1544, rip, roa, wee, N Baccharis halimifolia L., 5130, mes, roa, wet, N* Bidens aristosa (Michx.) Britt. 4081, nov, pin, rip, et,N Bidens bipinnata L., J.E. Moore, 3347, UCAC, N Bidens discoidea (Torr. & Gray) Britt., 6044, roa, N Bidens frondosa L., 4604, roa, wet, N Boltonia diffusa Ell., 3868, see, rip, w Brickellia eupatorioides (L.) Shinners, 4510, pin, N Carduus nutans L., 5089, roa, I* Centaurea cyanus L., J. Hauser, 255, APCR, | Chrysopsis pilosa Nutt., 4028, roa, wee, wet, N > Cirsium altissimum (L.) Hill,3970,mes, pin, rip, roa, wet, Cirsium eat ie Fern.& Schub., 1856, mes, pin, rip, r Cirsium discolor been ex Willd.) Spreng., 6002, lob, roa, Cirsium horridulum Michx., 4943, see, N Conoclinium coelestinum (L.) DC., 4001, mes, rip, , wet, N Conyza canadensis (L.) Cronq. var. canadensis, 4091,mes, nov, pin, N 2404 Conyza canadensis (L.) Cronq. var. pusilla (Nutt.) , 4286, roa, wee, N Coreopsis grandiflora Hogg ex Sweet var.grandi- flora, 2101 pin, sha, N Coreopsis grandiflora Hogg ex Sweet var. harveyana (Gray) Sherff, 2286, lob, nov, pin, e,N ‘ psis lance latal . 1742, pin, rip, roa, wee, N Coreopsis palmata re 2003, pin, N Coreopsis tinctoria Nutt. var. tinctoria, 2199, sha, wee, N Coreopsis tripteris L., 3708, lob, rip, roa, see, N Croptilon divaricatum (Nutt.) Raf., D. M. Moore, 410229, UARK,N Doellingeria sericocarpoides Small,4011, see, wee, N Echinacea pallida (Nutt.) Nutt. 2380, pin, roa, sha, N Echinacea pupurea (L.) Moench, 2414, mes, nov, in, rip, roa, wet, N Eclipta prostrata (L.) L., 5737, wet, N saslpei ded carolinianus Raeusch., 3883, mes, rip, s oe tomentosus L., 3373, lob, mes, see, wet, N Erechtites hieraciifolia (L.) Raf. ex DC. var. hieraciifolia, 3509, pin, roa, sha, wet, N sail annuus (L.) Pers., 2489, mes, pin, rip, roa, Figron Di abaladlges L.,791,rip,N Erig hx., 880, mes, N Erigeron strigosus Muhl.ex Willd., 1562, mes, nov, rip, roa, sha, wee, wet, N 2 oe torium al 0,nov,N Eupatorium fistulosum Barratt, 3824, see, N Eupatorium perfoliatum L. var. perfoliatum, 4365 ee a abt spe mes, pin, N 25, lob, pin, N Fupatori lum serotinum aan ae lob, pin, rip, roa, W iN Eurybia hem ishperica (Alexander) Nesom, 3800, lob, nov, pin, roa, sha, w ene ee agai (L.) Cabrera, 2118, nov, roa, N Grindelia lanceolata Nutt, 3895, rip, N Helenium amarum (Raf.) H. Rock, 3778, rip, roa, wet, N Helenium flexuosum Raf. 5176, pin, rip, roa, wet, N BRIT.ORG/SIDA 21(4) Helianthus angustifolius L., 4488, rip, wet, N Helianthus divaricatus L.,5524,lob,mes, nov, pin, rip, roa, sha, wee, w ae hirsutus Raf, 3051 nov, pin, rip, roa, N Helianthus mollis Lam., R. D. Thomas, 128892A, NLU, N 1 DIAAG I! Helianth li plantagineus (Torr. & Gray) Shinners 3886, rip, N, SC Helianthus silphioides Nutt., 4274, mes, roa, N elianthus simulans E.E.Wats., 477, roa, N Heliopsis helianthoides (L.) Sweet, 1720, mes, rip, a Hieracium gronovii L., 2004, nov, pin, rip, roa, see, lonactis linariifolius (L.) Greene, 4589, pin, N Krigia biflora (Walt.) Blake var. biflora, 5041, mes, rip, see, wet, N ae piace oe pee 4696, rip, N 1(L.) Nutt., 4807, rip, wet, N ig) id Visinicall ari 4899, roa, N Lactuca canadensis L., 3175, lob, mes, nov, pin, roa sha, N Lactuca floridana (L.) Gaertn.,6001, mes, roa, wee Sl L., 3208, rip, wee, | L., Pias, 959, NLU, | nacanilie naan vulgare Lam., 2283, roa, sha, wee wet, N aa serrio Liatris aspera Michx., 3627, nov, N Liatris elegans (Walt.) Michx., yee roa, N Liatris pycnostachya Michx., 3713, lob, rip, roa, N Liatris squarrosa (L.) Michx. var. compacta Torr. & Gray, 2648, nov, N, SC Liatris squarrosa (L.) Michx. var. squarrosa, 3400, roa, N Liatris squarrulosa Michx., 3944, nov, pin, roa, N Marshallia caespitosa Nutt.ex DC. var. caespitosa, Ifo2, Packera aurea (L.) A.& D.Love, 4766,mes, pin, rip, Packera glabella (Poir.) C. Jeffrey, Hawkins, 18, APCR, N Packera obovata (Muhl. ex Willd.) W.A. Weber & 075, mes, rip, wet, N Packera tomentosa (Michx.) C. Jeffrey, 4698, roa, N Parthenium integrifolium L. var. integrifolium, 7,t0a,N Pityopsis graminifolia (Michx.) Nutt., 4587, nov, pin, N MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS Pluchea camphorata (L.) DC.,4225, pin, see, wee, N Polymnia cossatotensis A.B. Pittman & V. Bates, 675,mes,N, Prenanthes altissima L., 4351, pin, rip, see, N Pseudognaphalium hell eri (Britt.) A. Anderb. ssp. eller’, 4620, lob, p aN ce ee um ie (L.) Hilliard & sp. obtusifolium, 4238, mes, rip, wee, N ae carolinianus (Walt.) DC., 2345, roa, N wee, wet, eee fulgida Ait. var. tg (C.L.Boynt. & eadle) Cronq., 4044, w oe ia grandiflora (D. oe J.F.Gmel. ex DC. grandiflora, 5633, pin, roa, sha, N Rudbeckia hirtaL.,5507,lob,mes, pin, rip, roa, sha, e, wet, N rue laciniata L.var.laciniata,3341,mes, rip, Rudbeci subtomentos Pursh, 3577, rip, N aL. var. triloba, 3422, pin, rip, roa, NUUVECKIG C wet, N Silphium asteriscus L.var.asteriscus, 3570,mes, rip, roa, N Silphium integrifolium Michx., 3582, mes, pin, rip roa, N Silphiu 3232, roa, sha,N Siphium perfoliatum L. var. perfoliatum, 3872,rip, Mm laciniatiym ie roa, N Silphium radula Nutt., Mrs. J. Miller, 176, UARK, N Smallanthus uvedalius (L.) Mackenzie ex Small, 64, rip, wee Solidago arguta Ait. var. bootii (Hook.) Palmer & maree, 56954, BRIT, N Solidago auriculata Shuttlw. ex Blake, 335, mes, Steyermark, D N, SC olidago caesia L. var. caesia, 4428, mes, pin, rip, wee, N Solidago canadensis L., 6024, lob, mes, nov, roa, wet,N Solidago hi pier Muhl. ex Willd, 4586, mes, pin, rip, ro Solidago reno Ait.,4022,lob, nov, pin, rip, roa a, W Solidago se Ait.,4332, nov, pin, rip, sha, N olidago ouachitensis C.& J. Taylor, 4214, mes, rip, Solidago petiolaris Ait., 4406, nov, pin, rip, sha, wet, N Solidago radula Nutt., 4393, nov, pin, N 2405 Solidago rugosa P. Mill., 6006, lob, N ee ie ia Muhl. ex Willd. var. microphylla Gray, 4171, mes, pin, rip, roa, wet, N, SC a ie Muhl. ex Willd. var. palmeri a 4336, mes, NOV, pin, rip, roa, sha, wee, s ae ae Muhl. ex Willd. var. u/mifo 3630, nov, N Sonchus oleraceus L., 3200, wee, | Symphyotrichum anomalum (Engelm.) Nesom, 0, mes, Nov, pin, roa, N Symphyotrichum drummondit(Lindl.) Nesom var. texanum (Burgess) Nesom, 4492, nov, rip, sha, N Symphyotrichum dumosum (L.) Nesom, 4478, wet, N ee lateriflorum (L.) A. & D. Love, see, N smote oblongifolium (Nutt.) Nesom, J. Logan CAC, smotyrchan ontarione (Wieg.) Nesom, 4623, rip, Pe eae oolentangiense (Riddell) Nesom var. oolentangiense, 4622, nov, pin, sha, ae patens ae ) Nesom var.patens, smo pilosum anil Nesom, 6023, n, rip, roa, wee, wet, Simptyotichur racemosum (Ell.) Nesom, 4594, pin Spycam turbinellum (Lindl.) Nesom, 4593, Cea eine lanceolatum (Willd. s, rip, see, sha, N Taraxacum officinale G.H. Weber ex Wiggers, roa, wee, wet, N & | Thelesperma filifolium (Hook.) Gray, D. M. Moore, 54-112, UARK, N Verbesina alternifolia (L.) Britt. ex Kearney, 4163, Nesom, rip, N Verbesina helianthoides Michx.,5470,mes, pin, rip, roa, sha, Verbesina virginica L. var. virginica, 4222, rip, roa, Vernonia nee oe ssp. baldwinii, 3549, nov, pin, rip, roa, sha, w Vernonia baldwinii ss 0 interior (Small) Faust, , pin, roa / KAvAl ssp fasciculata, 3611, rip, N, SC 2406 Vernonia lettermannii Engelm. ex Gray, D. M. Moore -290, UARK,N, Vernonia missurica Raf., 3711, lob, pin, see, wet, N Xanthium strumarium L., 4421, rip, wet, N Azollaceae zolla mexicana Schlecht. & Cham. ex K. Presl, J. Peck, 9451 Balsaminaceae Impatiens capensis Meerb., 5458, rip, see, wet, N di daceae andina domestica Thunb.,4797, mes, l* ee ane oe L.,5515,mes, nov, rip, see, wet, N Betulaceae Alnus serrulata (Ait.) Willd., 2326, mes, rip, see, wet, N Betula nigra L., 6038, rip, wet, N Carpinus caroliniana Walt., 5437, rip, see, wet, N Corylus americana Walt. 5445, mes, rip, see, N Ostrya virginiana (P. Mill.) K.Koch, 5467, mes, nov, pin, rip, roa pidhenlacene ni lata L., 1011, rip,N Campsis vacieoel Seem. ex aan 6000, lob, a, wee, wet Catalpa bignonioides Walt., Blechnaceae Woodwardia areolata (L.) T.Moore, 3550, pin, see, wet, N 1680, wet, N Boraginaceae Cynoglossum ae rile & Drummond, Mrs. Miller, 179, U Cyl 10glossum virgi iNnianum L , 1694 ,mes, see, N Hackelia virginiana (L.) |.M.Johnston,D.M.Moore 430153, aN N im indi eliotropit L., 4070, rip, wet, | bithormennum avon Michx,, 4942, rip, N Myosotis verna Nutt mes, rip, see, wet, N Onosmodium a Michx. ssp. subsetosum (Mackenzie & Bush) Cochrane, 4401, nov, N Brassicaceae Arabis canadensis L., 2361, sha, N Arabis laevigata (Muhl. ex Willd.) Poir. var. laevigata, 1812, ri Arabis missouriensis Great 1657, mes, nov, pin, rip, roa, wet, N L DUrUUled Aill.) Aschers., 1295, pin, rip, roa, | BRIT.ORG/SIDA 21(4) Brassica rapa L., 769, roa, | Cardamine angustata O.E. Schultz, 576, rip, see, N, Cardamine . bosa (Schreb. ex Muhl.) B.S.P, W. Butler, 28, UCAC, N aes eaneaenati (Michx.) Sw., 619, mes, pin, rip, see, sha,N Cardamine hirsuta L., 688, mes, rip, roa, see, sha, wet, | codon sane oI var. arenicola (Britt.) O.E. Schultz, M.C. Black, 44, UARK, N ne ae Muhl. ex Willd., 4713, Draba aprica Beadle, E. Sundell, 11143, UAM, N, SC Lepidium virginicum L. var. medium (Greene) C.L. Hitche., 2179, roa, w Lepidium virginicum L. var. virginicum, 2570, rip, Oa, wee, wet, N Rorippa nasturtium-aquaticum (L.) Hayek, C. Reid, 1961, UARK,N Rorippa palustris (L.) Bess. ssp. fernaldiana (But- ters & Abbe) Jonsell, 4819, roa, wet, Selenia aurea Nutt., 696, sha, wet, N Streptanthus maculatus Nutt ssp. obtusifolius (Hook.) Rollins, 5699, nov, N, SC la Buddlejaceae Polypremum procumbens L., 3297, wet, N Cabombaceae Brasenia schreberi J.F.Gmel., 5021, wet, N Cactaceae Opuntia humifusa (Raf.) Raf. var. humifusa, 2362, nov, roa, sha, N Callitrichaceae ee bec dees Pursh ae heterophylla, es, pin, roa, sha, we Campanulaceae Campanulastrum americanum (L.) Small, 3643, Lobelia appendiculata A. DC., 5069, lob, pin, roa, N Lobelia cardinalis ., 4264, rip, roa, see, wet, N Lobelia inflata L., 3454, mes, rip, wee, N obelia puberula Michx., 6028, lob, mes, pin, roa, wee, wet, N Lobelia siphilitica L., 4467, roa, N Lobelia spicata Lam., 2374, lob, pin, rip, sha, N Triodanis lamprosperma McVaugh, 1903, nov, pin, roa, sha, wee, wet, N MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS Triodanis leptocarpa (Nutt.) Nieuwl., 1760, sha, N — ei (L.) Nieuwl. var. biflora (Ruiz on) Bradley, 2220, roa a aoe iata (L.) Nieuwl. var. perfoliata, 11, nov, pin, rip, roa, wet, 1 Capparaceae Cleome hassleriana Chod., 3866, wet, Polanisia dodecandra (L.) DC., G. E. ae 5297, R,N Caprifolia eee ae Mrs. J. Miller,63, UARK, N Lonicera japonica Thunb., 1673,mes, rip, see, wee wet, |* Lonicera sempervirens L., 993, lob, mes, noy, rip, roa, sha, N Sambucus nigra L. ssp. canadensis (L.) R. Bolli, 5,mes, rip, roa, wee, wet, N Symphoricarpos orbiculatus Moench, 3637, nov, pin, rip, roa, N Viburnum nudum L., 2446, see, N Viburnum prunifolium L., 1134, pin, N Viburnum rafinesquianum J.A. Schultes, 1026, Viburnum rufidulum Raf. 2010, mes, nov, pin, roa, Caryophyllaceae Arenaria serpyllifolia L.,4781, wee, | 2407 Chenopodium standleyanum Aellen, 4404, nov,N Cistaceae Lechea mucronata Raf., 3677, lob, N Lechea tenuifolia Michx,, 1296, roa, sha, wee, wet, N Clusiaceae st drummondii (Grev. & Hook.) Torr. & 4, pin, roa, sha, wee, wet, N nei gentianoides (L.) B.S.P, 3617, nov, pin, a,N Hypericum gymnanthum Engelm. & Gray, 3242, roa, Hypericum hypericoides (L.) Crantz, 3748, nov, pin, : Ip, TOd, , ' Hypericum lobocarpum 3285,rip,N Hypericum mutilum L., 5736, rip, roa, see, wet, N Hypericum prolificum L., 3185, mes, pin, rip, roa, sha, wet, N Hypericum pseudomaculatum Bush, 2267, nov, sha, wee, wet, N Hypericum punctatum Lam., 3284, mes, Nov, rip, ; t,N Triadenum walteri J.G.Gmel.) Gleason, 4305, wet, N Commelinaceae Commelina communis L. var. communis, 2587, mes, rip, wee, wet, pelle ina diffusa Burm. f.,4611, wet, N ta. 3559, rip, wee, wet, N stium glomeratum Thuill., 759, mes, rip, roa, wet, | Dianthus armeria L., 5490, nov, roa, sha, wee, | Minuartia muscorum (Fassett) Rabeler, 2331, rip, roa, sha, N Paronychia fastigiata (Raf.) Fern. var. fastigiata, J. E. Moore, 3088, U aponaria officinalis L., 3440, wet, Silene stellata (L.) Ait.f,, 3259, nov, pin, rip, roa, see N 1282, mes, nov, pin, rip, roa, N Silene virginica L., Stellaria media (L.) Vill. ssp. media, 881, mes, nov, pin, rip, wee, wet, | Stellaria media (L.) Vill. ssp. pallida (Dumort.) Aschers. & Graebn., 686, roa, wet, | Celastraceae uonymus americana L., 1792, mes, rip, N Ceratophyllaceae + hy} ef. mL., 5486, wet, N Chenopodiaceae C podium ambrosioides L., 4416, rip, wet, | omme}lina virginica rip, see, wet, N Murdannia keisak (Hassk.) Hand.-Mazz.,4408, wet, |* Tradescantia bracteata Small ex Britt. 2415, pin, N Tradescantia ernestiana E.S., Anderson & Woods., 8 mes, rip, roa, wet, N Tradescantia hirsuticaulis Small, 4809, nov, pin, a, wet, N Tradescantia hirsutiflora Bush, 2586, nov, wee, N Tradescantia longipes E.S. Anderson & Woods., Tradescantia occidentalis (Britt.) Smyth, 2679, lob, mes, wet di is Raf., 5052, mes, nov, rip, roa wee, wet, N bares ozarkana ES. fees & Woods., Kirkwood, K-26, UCAC, N Convolvulaceae Ipomoea hederacea Jacq., 4630, rip, | 2408 Ipomoea lacunosa L., 3855, rip, wee, wet, N Ipomoea pandurata (L.) G.EW.Mey.,3147,mes, rip, roa, wet, N Cornaceae Cornus drummondii C.A. Mey., 1575, rip, N 2230,meS, NOV, pin, rip, roa, see, Cornus florida | N Cornus foemina P. Mill. 5763, rip, wet, N Cornus obliqua Raf., 2089, mes, pin, rip, roa, N Crassulaceae Penthorum sedoides L., 5767, lob, see, wet, N Sedum nuttallianum Raf.,J.L. Roberts, 277, VARK, Sedum pulchellum Michx., 1757, sha, N Sedum ternatum Michx., 778, mes, rip, N, SC Cucurbitaceae Melothria pendula L., 3590, rip, roa, N Sicyos angulatus L.,4115, roa, wee, N Cupressaceae Juniperus virginiana L. var. virginiana, 5504, nov in, roa, sha, wet, ! Cuscutaceae Cuscuta compacta Juss. ex Choisy, 4005, rip, roa, see, wet, Cuscuta cuspidata Engelm., E. B. Smith, 3370, UARK, N Cuscuta gronovii Willd.ex J.A.Schultes, 6020, roa, Cuscuta indecora Choisy, 4226, wee, N Cuscuta pentagona Engelm., 6022, rip, sha, roa, wet, N Cuscuta polygonorum Engelm., 3642, nov, N Cyperaceae Bulbosytlis capillaris (L.) Kunth ex C.B. Clarke, 220, sha, N, SC Ibicans Willd.ex Spreng. var.albicans, 770, Carex Schwein., 2393, sha, N Carex amphibola Steud., 1588, mes, rip, see, N Carex lace Dewey, 1476, mes, rip, wet, N Carex bushii Mackenzie, 2372, roa, sha, N ae caroliniana Schwein., 5179, wet, N Carex cephalophora dine ex Willd., 2098, mes, nov pin rip roa, see Carex cherokeensis ee 4714, mes, rip, wet, N Carex complanata Torr.& Hook.,2160, lob, pin, rip, wet, N BRIT.ORG/SIDA 21(4) Carex crinita Lam., 4967, mes, rip, see, wet, N Carex davisti Schwein. & Torr., 1632, mes, N, SC Carex debilis Michx. var. debilis, 1652, mes, N Carex digitalis Willd., C. T. Bryson, 4344, UARK, N Carex festucacea Schkuhr ex Willd., 2542, rip, see, Carex flaccosperma Dewey, 5469, pin, rip, roa, wet, N Carex frankii Kunth, 5481, mes, pin, rip, roa, see, Carex glaucodea Tuckerman ex Olney, 5087, pin, Carex granularis Mubl. ex Willd. var. granu 11, roa,N Carex grayi Carey, 3603, rip, N Carex hirsutella Mackenzie, 2100, mes, nov, pin, oa, sha, wet, Carex intumescens Rudge, 4958, rip, wet, N arex jamesii Schwein., P.E. Hyatt, 6937, UARK, N Carex joorii Bailey, 3898, rip, N Carex laevivaginata (Kukenth.) Mackenzie, J. H. ttig aris, Carex latebracteata Waterfall, 5002, pin, sha, N, SC Carex laxiculmis Schwein., P.E. Hyatt, 7329, UARK, SC N, Carex laxiflora Lam., J. H. Rettig, 560, BRIT, N, SC Carex leavenworthii Dewey, 1548, rip, roa, N Carex leptalea Wahlenb., 2444, see, N Carex lupulina Muhl. ex Willd., 4959, rip, wet, N Carex lurida Wahlenb., 5460, mes, pin, rip, roa, see, Carex mile enbergii Schkuhr ex Willd, 3089, mes, v, pin, roa, sha, N Carex nigromarginata Schwein.,762, mes, pin, roa, N Carex oklahomensis Mackenzie, 2384, rip, roa, sha, N Carex oligocarpa Schkuhr ex Willd., 1435, rip, N Carex ouachitana Kral, Manhart & Bryson, 2791, Carex oxylepis Torr.& Hook. var.oxylepis, 2417,mes, pin, roa, N Carex oxylepis Torr. & Hook. var. pubescens J.K. ood, 1076, mes, see, N, SC Carex retroflexa Muhl.ex Willd. 1513, mes, rip, see, N Carex rosea Schkuhr ex Willd., 2437, mes, rip, see, N Carex squarrosa L., 5172, Carex stricta Lam.,E.B nied 3801, BRIT, N, SC MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS Carex swanii (Fern.) Mackenzie, 2739, nov, N, SC Carex texensis (Torr.) Bailey, 1499, rip, N Carex torta Boott ex Tuckerman, 1097, mes, rip, ee, N, Carex tribuloides Wahlenb., 5494, wet, N Carex vulpinoidea Michx., 5475, lob, mes, nov, pin rip, roa, see, wet, eg aristatus Rottb., E.Sundell, 1 eae echinatus (L.) Wood, 2066, ba wee, N Cyperus erythrorhizos Muhl., 4482, Cyperus esculentus L., 436, wee, wet, ae Cyperus flavescens L.,D.M Weare 32971, UARK, N Cyperus iria L., 4549, wee, wet, | Cyperus lupulinus (Spreng.) Marcks, 3725, nov, wee, N Cyperus polystachyos Rottb. var. texensis (Torr.) Fern., 4560, wet, N Cyperus pseudovegetus Steud., 5747, wet, N yperus squarros Cyperus strigosus L., 3324, roa, see, sha, wet, N Dulichium arundinaceum (L.) Britt. 3257, wet, N, SC Eleocharis acicularis (L.) Roemer & J.A. Schultes, 3300, wet, Eleocharis engelmannii Steud., 2304, rip, sha, wet, N Eleocharis lanceolata Fern., 5553, roa Eleocharis obtusa (Willd.) J.A.Schultes, 2724,mes, rip, roa, sha, wee, wet, N Eleocharis quadrangulata (Michx.) Roemer & J.A. Schultes, 5131, wet, N Fimbristylis annua (All.) Roemer & J.A. Schultes, a, N Fimbristylis autumnalis (L.) Roemer & J.A. Schultes, 4235, wee, wet, N Fimoristylis vahlii (Lam.) Link, 396, wet, N i ng Bumie nS 4419, wee, wet, N Ell., 3178, roa, N, SC lata (Lam.) Gray, 5751, rip, Dp} f int / : i c 5611, roa,N Rhynchospora glomerata (L.) Vahl, 3568, rip, N Rhynchospora recognita (Gale) Kral, 2687, lob, rip, N roa, Schoenoplectus pungens (Vahl) Palla var. pungens, Scirpus atrovirens Willd., 5578, roa, wet, N 10606, UARK, 2409 Scirpus cyperinus (L.) Kunth, 5731, lob, wet, N Scirpus georgianus Harper, 2310, mes, rip, N Scirpus pendulus Muhl., 5128, roa,N Scleria oligantha Michx., 2229, lob, nov, pin, roa sha, N Scleria triglomerata Michx., 2497, pin, N Dennstaedtiaceae Dennstaedtia punctilobula (Michx.) T. Moore, G. Oleson, 87-017, UARK, N, SC Pteridium aquilinum (L.) Kuhn var. fatiusculum Es sv.) Underwood ex Heller, 2151,mes, pin, ae se aquilinum (L.) Kuhn var. pseudo- caudatum (Clute) Heller, 5510, roa, N Deseo te nccase tifolia L., 3587, rip, | Dioscorea vi Noel 5442,mes, pin, rip, N Dryopteridac ee filix- ht (L.) Roth ssp. asplenioides ichx.) Hulten, 5491, lob, mes, rip, see, N Pee protrusa (Weatherby) Blasdell, 3413, rip, N Cystopteris tennesseensis Shaver, 2770,mes, rip, N Dryopteris celsa (Wm. Palmer) Knowlt, Palmer & ard ex Small, 2536, see, N, SC Dryopteris marginalis (L.) Gray, 2430, mes, pin, see, N Onoclea sensibilis L., 1687, see, N ee acrostichoides (Michx.) Schott, 5479, 5, pin rip see, wet Woodsia obtusa (Spreng.) Torr. ssp. obtusa, 3663 v, pin ’ Woodsia obtusa (Spreng.) Torr. ssp. occidentalis Windham, 3330, rip, sha, wee, Ebenaceae Diospyros virginiana L., 5500, nov, rip, roa, wet, N Elaeagnaceae Elaeagnus umbellata Thunb., 3592, pin, rip, roa, wet, |* Elatinaceae Elatine triandra Schkuhr, 4041, wet, N Equisetaceae Equisetum hyemale L. var. affine (Engelm.) A.A.Eat.,, 4457 ; Ericaceae Gaylussacia baccata (Wangenh.) K.Koch,V. Bates, 55, UARK, N, SC 2410 Lyonia ligustrina (L.) DC.,5529, mes, rip, see, wet, N Rhododendron prinophyllum (Small) Millais, 289, rip, see, N Rhododendron viscosum (L.) Torr., 2309, mes, pin, rip, se AN ca pees Marsh., 2657, nov, pin, rip, a, wet, Vaccinium fuscatum Ait. 2483, see, N Vaccinium pallidum Ait., 1898, mes, nov, pin, see, N Vaccinium stamineum L., 1129, mes, nov, pin, rip, N Vaccinium virgatum Ait., 5040, lob, pin, rip, see, Euphorbiaceae Acalypha gracilens Gray, 3088, nov, pin Acalypha monococca (Engelm.ex coy L.Mill. & Gandhi, 3733, nov, pin, sha, wet, N Acalypha rhomboidea Raf., 4132, roa, N Acalypha virginica L., 3136, roa, wee, wet, N Chamaesyce humistrata (Engelm.) Small, 3853, wee, wet, Chamaesyce maculata (L.) Small,4157, rip, roa, N Chamaesyce nutans (Lag.) Small, 3854, nov, rip, roa, wee, wet, N roton Michx., 4424, r Croton slandidless L. var. septentrionalis Muell.- rg., 3164, wee, wet, N Croton monanthogynus Michx., 3224, nov, roa, sha, wee, wet, N Croton willdenowii G.L. Webster, 3331, nov, sha, wet, N Euphorbia corollata L.,5519, mes, nov, pin, roa, see, sha, N bh LL; rn i 3332,mes,nov,sha, N Euphorbia dentata Michx., 6004, a oe Euphorbia spathulata Lam., 6004, Leptopus phyllanthoides (Nutt.) @ iy Webster, 5000, pin, Phyllanthus caroliniensis Walt., 3852, roa, see, wet, Tragia cordata Michx., 3340, mes, Tragia urticifolia Michx., R. D. oe 128945, Fabaceae pleas a Durazz., 3119, pin, dae wee, |* morpha fruticosa L., 4961, lob, rip, roa, BRIT.ORG/SIDA 21(4) Amorpha nitens Boynt., D. M. Moore, 55-289, Amorpha ouachitensis Wilbur, T. Huffman, sn, UARK, N Amphicarpaea bracteata (L.) Fern. var. bracteata, 49, pin, ri Amphicarpaea bracteata (L.) Fern. var.comosa (L.) Fern.,4130,r Apios americana edie 5543, pin, rip, roa, see, wet, N Astragalus canadensis L., 3548, nov, pin, N Astragalus crassicarpus Nutt. var. trichocalyx (Nutt.) Barneby, L. A. Bariola, 85, UARK, N, SC Astragalus distortus Torr. & Gray var.distortus, J.W. Gibbons, 33, UARK, N Astragalus distortus Torr.& Gray var.engelmannii (Sheldon) M.E. Jones, 4794, sha, N Baptisia alba (L.) Vent. var. macrophylla (Larisey) Isely, 4945, roa,N Baptisia bracteata Muhl. ex Ell. var. leucophaea utt.) Kartesz & Gandhi, 1981, noy, pin, rip, ,wet, N Baptisia nuttalliana Small, 2086, roa, N Baptisia sphaerocarpa Nutt., 4946 Cercis canadensis L.var. canadensis, a mes, pin, rip, roa, S Chamaecrista ane (Michx.) Greene, 3959, ! ' Chamaecrista nictitans (L.) Moench ssp. nictitans, 563,n Clitoria mariana L., 3118, lob, nov, ae rip, roa, N Crotalaria sagittalis L., 3833, w Desmodium canescens (L a0, : ° Smith, 3773, Desmodium cuspidatum (Muhl. ex Willd.) DC. ex ar. cuspidatum, 6021, roa, see, N eee a glutinosum (Muhl. ex Willd.) Wood, 3933,mes,N Nesmodittm Iqeviaqat J (Nutt.) DC., 4388, lob, N Desmodium marilandicum (L.) DC., 4087, pin, N mca nudiflorum (L.) DC., 3476, lob, mes, pin eae nuttallil (Schindl.) Schub., 3404, rip, N Desmodium ae (Muhl. ex Willd.) DC., 6009, lob, pin, Desmodium *panicutaram (L.) DC. var. paniculat 66 nov, pin, roa, see, N ee oe (Nutt.) DC., 3366, mes, pin, rip, see, N MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS Desmodium perplexum Schub.,3997,mes, pin, rip, see, wet, N esmodium rotundifolium DC.,4577,mes, N Galactia regularis (L.) B.S.P.,, 3731, lob, nov, N Britt., a Gleditsia triacanthos i 2 Kummerowia stipulacea mea Ww | ‘ fh pace Ps Al OGIaCud VOU HIS \L. 5 Makino, 237, roa, | Kummerowia striata (Thunb.) Schindl., 3312, lob, ov, roa, wet, | ae sah 1936, pin, | latifolius L., 2532, roa, | Lathyrus venosus Muhl. ex Willd., 1064, mes, nov, pin, N sages cuneata (Dum.-Cours.) G. Don, 3288, rip, roa, wee, wet, I* lspedero iol .) Hornem., 3932, lob, nov, pin a,N ee ane procumbens Michx., 4591, pin, roa, N Lespedeza repens (L.) W. Bart., 4387, lob, pin, roa, ee, wet, N Lespedeza violacea (L.) Pers., 3652, rip, wee, wet, espedeea VIRGIN EGS .) Britt. 4021, pin, roa, N al.3 ,wee, | oie peduncu atum (P. Mill.) Rydb. var. pedunculatum, 4818, mes, wet, N we Pueraria montana (Lour.) Merr.var.lobata (Willd. & S. Almeida, 4310, wee, I* Rhynchosia latifolia Nutt.ex Torr.& Gray, 5563, pin, N Robinia hispida L., 5088, roa, N Robinia pseudoacacia L., 3143, mes, Nov, pin, rip wee, wet, nen inia viscosa Vent, 1666, roa, N a (L.) Link, 3620, nov, pin, rip, N oe helvula (L.) Ell, 3148, roa, wet, N Stro, ps umbellata — ex Willd.) Britt., 5758, lob, pin, roa, wet, Stylosanthes biflora (L.) oo 4272, pin, roa, sha, N Tephrosia onobrychoides Nutt., G. Barber, 2201, ARK,N Tephrosia virginiana (L.) Pers., 2499, nov, pin, roa, N Trifolium arevense L.,5210,roa, iM Schreb., 134 coe dub um cde G ae sy UARK, | 1m L., 4854, nov, roa, wee, | Trifolium pratense L., 1150, pin, rip, roa, wee, wet, | Trifoli ling incarnat, 2411 Trifolium reflexum L., 1315,r0a, N Trifolium repens L., 1149, mes, pin, roa, wee, | rifolium vesiculosum Savi, 2604, wee, wet, | Vicia caroliniana Walt., 1119, pin, wee, wet, N Vicia minutiflora F.G. Dietr.,4723, wet, N Vicia sativa L., 4703, nov, rip, roa, wee, wet, | Vicia tetrasperma (L.) Schreb., 5108, roa, | Vicia villosa Roth ssp. varia (Host) Corb., 2069, nov, Oa, Wee, Vicia villosa Roth ssp. villosa, 2713, wet, | Wisteria floribunda (Willd.) DC., 4300, wet, I* Wisteria sinensis (Sims) DC., 3369, roa, I* Fagaceae Castanea pumila (L.) P. Mill. var. ozarkensis (Ashe) Tucker, 5630, mes, nov, pin, see, N, S Castanea pumila (L.) P. Mill. var. pumila, 195, see, N Fagus grandifolia Ehrh., 1637,mes, rip, see, N Quercus acerifolia (Palmer) Stoynoff & Hess, 3615, nov,N, Quercus alba L., 5472, mes, pin, rip, roa, see, wee, N Quercus falcata Michx., 5153, lob, mes, nov, pin, N hh., 2633, nov, pin, N fi Engelm., 4395, mes, nov, en by pin, rip, roa, sha, N Quercus nigra L., 4674, rip, wet, N Quercus pagoda Raf, 1518, rip, N Quercus palustris Muenchh., 4964, wet, N 2329, lob, pin, rip, roa, wet, N Quercus rubra L.,4391, mes, nov, N Quercus shumardii Buckl.var. shumardii, wr Quercus phellos | 1739,rip, N Quercus stellata Wangenh., 4507, nov, pin, wee, ae Quercus velutina Lam., 5620, nov, pin, N Fumariace Sige Bedi ina Engelm.,W.Butler,57,UCAC, oe is flavula (Raf) DC., 4683, rip, N Corydalis micrantha (Engelm.ex Gray) Gray ssp. australis (Chapman) G.B. Ownbey, 774, rip, N Gentiana Bartonia eneine (Michx.) Muhl. ssp., paniculata, 4007, see, Frasera eas Walt., 1750, rip, N Sabatia angularis (L.) Pursh, 3371, roa, sha, wee, N 2412 Sabatia campestris Nutt., D.M. Eggers Ware, 5483, UARK, Geraniac Geranium eee L., 1152, mes, roa, wet, N Geranium dissectum L., 1563, roa, | Geranium maculatum L., 1247, rip, N Grossulariaceae [tea virginica L., 3267, rip, see, N Ribes curvatum Small, J.L. Roberts, 617, VARK,N, SC Haloragace M pat htop ede 5549,wet,N Myriophyllum spicat Proserpinaca ie L,, . wet, N Hamamelidaceae Hamamelis vernalis Sarg., 1000, mes, pin, rip, wet, Hamamelis virginiana L., 442, mes, nov, pin, rip, e,N Liquidambar styraciflua L.,3875, pin, rip, see, wee J rashes glabra Willd., 4859, mes, nov, rip, see, N Aesculus nave L., 1058,mes, N Hydrangeaceae Hydrangea arborescens L., 5448, mes, rip, see, N ocharitaceae a verticillata (Lf) Royle, 5168, wet, | = = Hydro Hydri BySreply acess Nutt.ex Choisy, 5724, wet, N irony brownei Kral & Bates, 1712,mes, rip, see, N, SC Nemophila phacelioides Nutt., 4938, rip, roa, wet, N Phacelia hirsuta Nutt., 1148, mes, roa, wet, N Phacelia ranunculacea (Nutt.) Constance, 4690, mes, rip, N Hymenophyllaceae Trichomanes petersii Gray, J. Peck, 82457, LRU,N, SC lridaceae geri chinensis (L.) DC., J. E. Moore, sn, Iris een a” 2095, mes, Nov, pin, rip, see, N Iris pseudacorus L., 4924, wet, Iris verna L.,D.M. Moore, 51 ae UARK, N, SC BRIT.ORG/SIDA 21(4) Iris virginica L., 4965, wet, N Sisyrinchium angustifolium P. Mill, 5028, mes, rip, Sisyrinchium campestre Bickn., 4795, roa, N Sisyrinchium langloisii Greene, M. C. Black, 81, UARK, N Isoetaceae lsoetes melanopoda Gay & Durieu ex Durieu, 852 Juglandacea Carya alba (L i; Nutt. ex oi 5502, lob, mes, nov, pin, roa, wee, wet Carya cordiformis oa ) K. Koch, 5438, mes, nov, rip, N Carya texana Buckl., 3645, mes, nov, pin, sha, N Juglans nigra L., 3463, lob, mes, N Juncaceae Juncus acuminatus Michx.,4969, mes, rip, roa, wet Juncus brachycarpus Engelm., 5610, roa, N Juncus coriaceus Mackenzie, 5732, lob, mes, pin, a, see, wee, Wet, juneus dichotomus Ell., 2301, rip, N Juncus diffusissimus Buckl., 5554, rip, roa, wee, N Juncus effusus L.,5547,mes, roa, wet, N sedis inter Or IEG ne roa, N 555, lob, roa, wee, wet, Juncus secundus Beauv. ex Poir., 3384, roa, sha, wet, N Juncus tenuis Willd., 2083, pin, rip, roa, see, sha, ee, wet, N Juncus torreyi Coville, 3684, lob, wet, N Juncus validus Coville, 3535, roa, N Luzula acuminata Raf. var.acuminata, 2786, mes, N Luzula acuminata Raf. var. carolinae (S. Wats.) 613, rip, Luzula bulbosa (Wood A Smyth & Smyth, 4791, zr ila Luzula echinata ta (Small) FJ. Herm., 647, Mes, pin, rip, roa, see, N Lamiaceae Ajuga reptans L., 763,r Clinopodium be eta (Nutt.) House, 5573, roa, N Cunila origanoides (L.) Britt. 4106, mes, pin, rip, see, wee, N MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS Hedeoma hispida Pursh, 5199, sha, wet, N Hedeoma pulegicides (L.) Pers., Simpson,sn, UARK, N Isanthus brachiatus (L.) B.S.P, E. B. Smith, 3767, UARK, N | ; pl L.,676, rip, roa, N Lamium purpureum le 67, rip, roa, wet, N Lycopus rubellus Moench,4207,mes, rip, see, wet Lycopus virgil ICUS kes 4655, see, N Mentha spicata L., Simpson, sn, UABK, | Monarda_ fistulosa ssp. fistulosa var. stipitatoglandulosa (Waterfall) Scora, comb. nov.ined.,2571,nov, pin, roa, sha, wee, wet, N Monarda punctata L.,D. Demaree, 57040, APCR, N Monarda russeliana Nutt.ex Sims, 2045,mes, pin, rip, Perilla frutescens (L.) Britt.,4146, mes, pin, rip, | Physostegia angustifolia Fern., 5605, roa, N Prunella vulgaris L., 4953, lob, mes, pin, rip, roa, Pycnanthemum albescens Torr. & Gray, 3176, lob — Pycnanthemum muticum (Michx.) Pers. 3253, rip, Pycnanthemum tenuifolium Schrad., 5497, lob, Al, roa, a wet, N ichx.ex Lam., D. Demaree, 62848, NLU,N Salvia lyrata L., 904, mes, rip, roa, wet, N Scutellaria elliptica Muhl.ex Spreng. var. elliptica /mes, rip, wet, Scutel Had lateriflora L., 4049, wet, N vata Hill, 2409, mes, nov, pin, wet, N cae anlinci Nelson, 3461, mes, pin, N, SC Stachys tenuifolia Willd., 3594, mes, rip, N Teucrium canadense L. var. canadense, 3420, rip, N Trichostema dichotomum L., Simpson, sn, UARK, Lauraceae Lindera benzoin (L.) Blume, 1619, mes, rip, see, N Sassafras albidum (Nutt.) Nees, 5511, mes, nov, pin, see, wet, N Lemnaceae Lemn tialis Welw., 4030, w oes a punctata (G.F.W. Mey.) CH. a Aeon 4 ‘ 2413 Lentibulariace Urticularia Be - 4303, wet, N Liliacaeae Allium canadense L. var. canadense, 1234, pin, rip, ,wet, Allium canadense L. var. mobilense (Regel) Owenby, 2242, ae N Allium vineale L., 2191, wee, | Amianthium raUeCtONICUE (Walt.) Gray, 2459, mes, see, N Camassia scilloides (Raf) Cory, 4849, nov, N Erythronium albidum Nutt. 544, rip 7 f) Erythronium rostratum W.Wolf, 604, nov, rip, see, N Hemerocallis fulva (L.) L., 1920, nov, | Lilium michiganense Farw., Mrs. J. Miller, 166, UARK, N Maianthemum racemosum (L.) Link, 1307, mes, pin, rip, roa, see, Melanthium virginicum L., 5483, lob, pin, roa, see N Melanthium ii (J.W.Robbi n,R. Davis, 1967, APCR, N, SC Muscari botryoides (L.) P. Mill., 682, roa, | Nothoscordum bivalve (L.) Britt., 1091, mes, nov, rip, roa, sha, wet, Ornithogalum umbellatum L., 957, ri Pol am biflorum (Walt.) EIL, oo rip, see, Wood) Boa- senontim gramineum (Ker-Gawl.) Morong var. ramineum, G.E. Tucker, 15000, APCR, N, SC Trillium pusi ee Michx. var. ozarkanum (Palmer & Steye ermark) Steyermark, 609, rio, N, SC Trillium recurvatum Beck, 4941, mes, rip, N Uvularia grandiflora Sm., 2412, pin, N Uvularia perfoliata L., 701, mes, N, SC Uvularia sessilifolia L., 5044, wet, N Linaceae Linum medium (Planch.) Britt. var. texanum (Planch.) Fern., 3513, roa, wet, N Linum striatum Walt., 3280, pin, rip, roa, N Loganiaceae Spigelia marilandica (L.) L.,1439,mes, pin, rip, roa, N ycopodiaceae ee um digitatum Dill.ex A.Braun, 1117,pin, N 2414 Lythraceae Ammannia coccinea Rottb., 400, wet, N a diandra (Nutt. ex DC.) Wood, 5725, wet, N,SC Rotala ramosior (L.) Koehne, 6042, wee, wet, N Magnoliaceae Magnolia acuminata (L.) L.,4575, mes, N Mecnohaarandtions L.,2147, pin, see, N Magnolia tripetala (L.) L., 1622, mes, rip, see, N Malvaceae Abutilon theophrasti Medik., D. X Williams, ARFOO78, UARK, Callirhoe pedata (Nutt.ex Hook.) Gray, 1753, sha, N Hibiscus laevis Allioni, 3847, wet, N Malva neglecta Wallr.,, Simpson, sn, UARK, | Sida spinosa L., 3843, roa, wet, sileaceae Pilularia americana A. Braun, 5201 »sha, N, SC Melastomataceae — mariana L. var. interior (Pennell) Kral & 3282, rip N et mariana L.var. mariana, 2700, lob, wet, N Rhexia virginica L., 5723, wet, N Menisperma ee ol (Pursh) Gray, 2581, rip, see, wee _N Cocculus carolinus (L.) DC., 2732, nov, pin, rip, roa wee, wet, N NApnicnormrim conn, L., 1353, rip,N i renecunaiine o verticillata L., 402, wet, N Hee ra L., 4579, pin, N BU UE! Moraceae Maclura pomifera (Raf.) Schneid., 1723, rip, wet, N Morus rubra L., 5538, rip, roa, see, wet, N Najadaceae Najas guadalupensis (Spreng.) Magnus, 4281, wet, N Nyctaginace Mirabilis al ener ) Heimerl, 2600, roa, wee, N Nymphaeaceae Nuphar lutea (L.) Sm., 5019, wet, N Nymphaea odorata Ait., 5018, wet, N BRIT.ORG/SIDA 21(4) yssaceae Nyssa sylvatica Marsh.,4966, mes, Nov, pin, rip, roa, pedenes cana L., 5493, lob, mes, pin, rip, N saul inus pennsylvanica Marsh., T. Huffman, sn, Ligustrum sinense Lour., 4466, lob, mes, rip, roa, see, wee, |* Onagraceae Gaura demareei Raven & Gregory, 250, roa, N Gaura longiflora Spach, 3840, roa, w ae see L., 5459, lob, mes, int roa, see, wet, N Ludwigia d 15 Walt., 3885, sale wet,N Ludwigia g inpdulose Walt., 3389, En eWIgig palesltis 5 (L.) Ell., 5097, roa, wet, N enothera fruticosa L., 5010, mes, pin, N Oenothera laciniata Hill, 2580, roa, wee, N Oenothera linifolia Nutt., 4978, pin, roa, sha, N Oenothera villosa Thunb., 4140, rip, roa, N Ophioglossaceae Botrychium un (Sav.) Underwood, 4628, pin, rip, Botrychium Access Spreng., J. E. Moore, sn, UARK, N LS selec virginianum (L.) Sw., 1069, mes, rip, e,N Ophea ossum crotalophorioides Walt., C. Reid, 5, UARK, N ee ossum vulgatum L., 2363, sha, N Orchidaceae Cypripedium kentuckiense C.F. Reed, 841,mes, rip, N, SC Goodyera pubescens (Willd.) R. Br. ex Ait. f., J. E. 081, UCAC,N /sotria verticillata Raf, 5661, see, N Platanthera ciliaris (L.) Lindl., 192, rip, see, N llata (Michx.) Luer, 3787, see, N Platanthera lacera (Michx.) G. Don, 5100, roa, N Spiranthes cernua (L.) L.C. Rich., 4483, roa, wet, N Spiranthes tuberosa Raf., 208, sha, Spiranthes vernalis Engelm.& Gray, 2706, roa, wet, N Plata th aAcla Orobanchaceae Epifagus virginiana (L.) W. Bart., 3407, rip, N Orobanche uniflora ks) G Hunter, 70, UARK, N MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS Osmundaceae Osmunda cinnan , 986, rip, see, N Osmunda regalis L.,5753, mes, pin, see, wet, N Oxalidaceae Oxalis corniculata L.,675, roa, N Oxalis stricta L., 5034, lob, mes, pin, rip, roa, wee, i wet, N Oxalis violacea L., 884, mes, pin, rip, roa, see, sha N Papaveraceae fe ; iy qd ic | Sanguinaria canadensis L ,610, rip, roa, see, N Passifloraceae if| Passi PUTRI CGT lata L.,5523,rip, roa, wee, N Passiflora lutea L.,5518, nov, pin, rip, roa, wee, wet N Phytolaccaceae Phytolacca americana L., 5526, nov, pin, rip, roa, Pinaceae Pinus echinata P. Mill. 3086, pin, rip, N Pinus strobus L.,4574,mes, N Pinus taeda L., 5761, lob, nov, wet, N dieu tata Mi 578, roa, sha, wee, N Piantago ee Nutt Hardin, 606, APCR, N 2124,roa, wee, | Plantago oil jDene, 2540, rip, roa, see, N _rip, roa, wee, N P lantago virgit Platana Platanus ee 1813,rip,N Poaceae Agrostis gigantea Roth, 4544, pin, rip, | Agrostis hyemalis (Walt.) B.S.P,, 5078, rip, roa, wet, N Agrostis perennans (Walt.) Tuckerman,E.B.Smith, 3 R a Aira caryophyllea L., 1163, roa, | Aira elegans Willd.ex Kunth, D.M.Moore, 55- 291, RK, | pare carolini anus Walt., 910 lii Vit Pl sis, roa, § PasoReg remnaiitis ee 4505, rip, N virginicus,6025,roa ng itman, 3546, a nov, pin, Ar 1\dropogol | virginicus | L Anthoxanthum aristatum Boiss., 5149, roa, | Anthoxanthum odoratum L., 878, mes, | 2415 pices cen Otgiag Michx., 218, roa, sha, N Mi 4312,wee,N Arthraxon hid (Thunb.) Makino, 4254, roa, an naria gigantea (Walt.) Muhl., 5480, mes, rip, See, pee furcatus (Fluegge) A.S. Hitchc., 4613 wet, N Brachyelytrum erectum (Schreb. ex Spreng.) Beauv,, 3356, mes, nov, pin, rip, see, N Bromus catharticus Vahl, 4895, roa, | Bromus commutatus Schrad., 1682, wet, | teen hordeaceus L., 1030, wee, | Bromus japonicus Thunb. ex Murr.,2072,nov,wee, | Bromus pubescens Muhl.ex Willd., 1809, mes,nov, pin, rip, roa, sha, N Bromus racemosus L., 1569, roa, suas Bromus secalinus L., 2749, no toa, | Chasmanthium latifolium Re Yates, 2250, es, pin, rip, roa sha, N Chasmanthium laxum (L.) Yates, 3489, lob, mes, Peaemant um sessiliflorum (Poir.) Yates, 3480, pin, roa, wet Cynodon dactylon (L.) Pers., 4474, roa, wee, wet, |* P\yaeiihtea to | 5601, nov, pin, roa, sha, wee, | Danthonia spicata (L.) Beauv. ex Roemer & J.A. Se SNES: 1925, nov, pin, roa, sha, wee, N Di Beauv., Cae mes, nov, N rip, § Di aaa um acuminatum (Sw.) Gould & CA. lark var ee roa, sha, Peele jum ve (Poir.) Gould & C.A. Clark, 2022, mes, NOV, pin, rip a clandestinum (L.) Gould, 3493, eu commutatum (J.A. Schultes) ould, 1354, mes, nov, pin, rip, roa, see, N Dichanthelium depauperatum (Muhl.) Gould, 1327, pin,roa,N ee Siialead .) Gould, 2346, ee aa yreaee ae ) Gould, 2062, roa, wee, wet, Dichanthelium linearifolium (Scribn. ex Nash) Gould, 4982, mes, nov, pin, sha, N 2416 Dichanthelium oligosanthes (J.A.Schultes) Gould, 1370, wee, Dichanthelium ravenelii (Scribn. & Merr.) Gould, _pin, Dichanthelium scoparium (Lam. Ss Gould, 3240, pin, ia ar sphaerocarpon (Ell.) Gould var. sop dena eee ) Gould & CA. Clark, 5463, ob,m rip, roa, sha, wet, ce ee (Ell.) Gould var. sphaerocarpon, 2693, lob, mes, pin, sha, N Digitaria ciliaris (Retz.) Koel., 3162, roa, wee, wet, N Digitaria ischaemum eae Schreb, ex Muhl., 2,MeS, roa, wee, W Digitaria sanguinalis (L a 3775, pin, rip, wet, N Digitaria violascens Link, 213, sha, | Echinochloa colona (L.) Link, 4609, wet, | Echinochl iyi rei Aoaaed aes 5721,rip,roa, Eleusine ie (L.) Gaertn., 4409, wee, wet, | Elymus canadensis L., 2592, wee Elymus hystrix L., 5579, nov, rip, N Elymus villosus Muhl. ex Willd., 2583, nov, wee, N Elymus virginicus L. var. virginicus, 5474, pin, rip, Oa, see, wee, Wet Eragrostis capillaris (L.) Nees, 3714, nov, N Eragrostis curvula (Schrad.) Nees, 3456, mes, roa, wee, | Eragrostis hirsuta (Michx.) Nees, 4309, wee, N Eragrostis hypnoides (Lam.) B.S.P., 5994, roa, wet, N Festuca paradoxa Desv., 1961, lob, pin, N Festuca subverticillata (Pers.) Alexeev, 1621, mes, pin, rip, see, wee, N Glyceria striata tam ) A.S.Hitchc.,2325, mes, pin rip, se Holcus ee L.,5151,roa, I* Hordeum pusillum Nutt., 1558, roa, wet, N Leersia oryzoides (L.) Sw., 4377, roa, wet, N Leersia virginica Willd.,4514, lob, mes, pin, rip, see, Lolium arundinaceum eg S.J. Darbyshire, mes, roa, sha, w Lolium perenne L.ssp. a ae (Lam.) Husnot, 00, roa, sha, wee Lolium perenne L. ssp. perenne, 1794, nov, rip, roa, ,wet, | Melica mutica Walt., 5047, mes, rip, roa, wet, N No BRIT.ORG/SIDA 21(4) Melica nitens (Scribn.) Nutt. ex Piper, 2750,nov,N Micros aa vimineum (Trin.) A. Camus, 441, rip, ro nen a schreberi JF Gmel, D. M. Moore, 20407, UARK, N munerbeae sobolifera (Muhl. ex Willd.) Trin., i sylvatica Torr. ex Gray, 4219, mes, wee, N Meblenbeaia tenuiflora (Willd.) B.S.P., No collec- or listed, sn, VAM, N Oplismenus hirtellus (L.) Beauv., D. M. Moore, 410404, UARK,N ,3827,lob,nov, pin, rip, roa, me) et, N Panicum capillare L., 3736, nov, roa, N Panicum dichotomiflorum Michx.,4616, nov, pin, wet, N Panicum flexile (Gattinger) Scribn., 4308, wee, N Panicum philadelphicum Bernh.ex Trin. 217, sha, N Panicum rigidulum Bosc ex Nees, 4138, rip, roa, wet,N Panicum virgatum L., 3865, lob, nov, pin, rip, roa wee, wet, N aera dilatatum fea 3537, roa, wee, wet, | (L.) L., 378, wet, N pane nedondaaum Michx., 4267, roa, N Paspalum laeve Michx., 3202, wee, wet, N Paspalum notatum Fluegge var. saurae Parodi, 3654, lob, roa, wee, | spalum setaceum Michx., 3524, roa, wet, N Paspalum urvillei Steud., 4278, wet, | Pennisetum glaucum (L.) R.Br, 3777, pin, rip, wee, Py palum diccecrtis Phleum pratense L., 4719, sha, wet, | Piptochaetium avenaceum (L.) Parodi, in, roa, i annua L., ea he roa, wet, | x Ell., 895, mes, see, N 1330,mes, nN ay tt. Boa bulbosa L., 4910,1 roa, | Red pater L.,4912,roa,N t ay, 1114,mes, rip, N sacenailnn shooter ae (L.) Nutt.,4211,roa,N Saccharum brevibarbe (Michx.) Pers. var. contortum (Ell.) R.Webster, 3553, rip, roa, wee, N Sacciolepis striata (L.) Nash, 4497, rip, N, SC Schizachyrium scoparium (Michx.) Nash, 4355, Vv, pin, roa, sha, N MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS Secale cereale L., 1342, roa, sha, we Setaria parviflora (Poir.) Kerguelen, ae roa, sha, ’ Sorghastrum nutans (L.) Nash, 4018, roa, N Sorghum halepense (L.) Pers., 3612, rip, roa, wee, wet, I* na eee nitida (Biehler) Scribn., 2008, nov, shenooh obtusata (Michx.) Scribn., 1322, rip, roa oe clandestinus (Biehler) A.S. Hitche., R. or, 38721, UARK, N Gaeicte us compositus (Poir.) compositus, 219,sha Sporobolus vaginiflorus (Torr. ex Gray) Wood, 4320, wee, N Steinchisma hians (Ell.) Nash, 2176, rip, roa, sha, et, N Stenotaphrum secundatum (Walt.) Kuntze, 4060, roa, N Tridens flavus (L.) A.S.Hitchc. var. flavus, 3235,mes, nov, pin, roa, sha, N Tridens strictus (Nutt.) Nash, 3923, roa, wet, N Tripsacum dactyloides (L.) L., 3530, roa, N Vulpia myuros (L.) K.C.Gmel,, 2231, roa, sha, wee, Vulpia octoflora (Walt.) Rydb., 1288, pin, wet, N Podostemaceae Podostemum ceratophyllum Michx., J. E. Moore, sn, UARK, N, SC Polemoniac Phlox eae ie ssp. laphamii (Wood) Wherry, 1464, rip, wee, N Phlox paniculata L., 3588, rip, roa, N Phlox ee L. ssp. ozarkana (Wherry) Wherry, iN ' a L. ssp. pilosa, 902, mes, Nov, pin, rip, roa, N Polemonium reptans L., 1589, rip, N Polygalaceae Polygala ambigua Nutt., 2185, nov, pin, sha, wee, N Polygala polygama Walt.,D.M.Moore,4176,UARK, N Polygala sanguinea L., 5075, roa, sha, N rolygenacsae gifolium Nutt., 4651, sha, N Polygonella americana (Fisch.& C.A. Mey.) Small, 3633, nov, N Merr. var. 2417 ilare L 492, Ww oo. caespitosum var. longisetum (de Bruyn) A.N. Stewart, 3995, mes, rip, see, wee, Polygonum convolvulus L., 1921, nov, | Pol ee ee es Michx.,6041, lob, rip, = vgonum lapathotur. oe wet, N 4149, rip, roa, wet, aie Po Pol oa punctatum Ell, 3994, rip, see, wee, wet, piesa ea L., 266, rip, see, N Polygonum scanden Polygonum setaceum Baldw., 3528, roa, N Polygonum tenue Michx., 4397, nov, sha, N ,3839,mes, rip, see, N L., 325, mes, nov, rip, N Pol OGrhulil virgit vanum | Rumex acetosella L., 1553, rip, roa, ae | Rumex altissimus Wood, 14 Rumex conglomeratus Murr, i i cn 5110,UARK, | Rumex crispus L., 3206, nov, roa, wee, wet, | Rumex hastatulus Baldw., 5175, wee, wet, N Rumex obtusifolius L., 1707, rip, roa, | Rumex pulcher L., 2602, wee, | Polypodiaceae Pleopelt is polypodioides (L .) Andrews & Windham ssp. michauxianum (Weatherby) Andrews & Windham, 1111,mes, nov, pin, rip, see, sha, N Ponteder eerie ie (Sw,) Willd. 4031, wet, N Portulacaceae Claytonia virginica L., 1050, lob, mes, Nov, pin, rip roa, see, wet, N Talinum calycinum Engelm., 2644, nov, sha, N Potemogetonacea Potamogeton hese Raf.,6029, es N Potamogeton nodosus Poir., 3887, w Potamogeton p ule Tuckerman, aie wet, N Potamogeton pusillus L., 5024, we Enlereae m a L.,4782,5 lveinaehia laaceolata Walt., 2339, mes, rip, N Lysimachia quadriflora Sims, 4948, roa, Samolus valerandi L.ssp.parviflorus (Raf) Hulten, 181, mes, roa, see, wee, N Pteridaceae Art. + ilus-veneris L., Mayo, 275, HEND, N 2418 Adiantum pedatum L., 4576, mes, N Cheilanthes lanosa (Michx.) D.C. Eat., 2731, nov, n,N pl Cheilanthes tomentosa Link, W. C. Taylor, 918, U Ranunculacea Actaea Sere Ell. ye mes,N Anemone virginiana | : 8, lob rip, Sec, N Clematis reticulata ne ey wee, N Delphinium carolinianum Walt.ssp.carolinianum, 1752, mes, pin, sha, N ga tl newtonianum Moore, E. Hardcastle, R13, UARK,N, ben tricorne Michx., 1010, mes, rip, N nemion biternatum Raf., C. Hunter, 58, UARK, N 2 bats lis Schreb. var. obtusa (Pursh) Steyermark, 611,mes, rip, N Ranunculus abortivus L., 4747, rip, wet, N Ranunculus bulbosus L., 4684, rip, | Ranunculus fascicularis Muhl. ex Bigelow, 4783, sha, N Ranunculus harveyi (Gray) Britt, 4784, mes, rip, roa, Ranunculus hispidus Michx. var. nitidus (Chap- man) T.Buncan, M.C. Black, 39, UARK, N laxicaulis (Torr. & Gray) Darby, 4957, Rantincisliye wet, N Ranunculus micranthus Nutt., 4687, nov, rip, roa, wet, N Ranunculus pusillus Poir., 5508, roa, see, wet, N Ranunculus recurvatus Poir., 1505, mes, rip, see, N Ranunculus sardous Crantz, 2186, roa, wee, wet, | Thalictrum dasycarpum Fisch. & Ave-Lall., 1778, rip, alictrum revolutum DC., 3602, ri Thalictrum thalictroides Eames & Boivin, 706, mes, Ov, pin, rip, see, wet, N Rhamnaceae Berchemia scandens (Hill) K. Koch, 5742, mes, rip, panies americanus L., 5003, pin Raf, J.C. Baker 60, VAM, N aie caroliniana (Walt.) Gray, 5537,mes, nov, pin, rip, sha, wee, wet, N Rosaceae Agrimonia parviflora Ait., 4256, roa, N Agrimonia rostellata Wallr., 5740, a mes, pin, rip, BRIT.ORG/SIDA 21(4) Amelanchier arborea (Michx. f.) Fern., 5749, lob, mes pin Ip Chaenomeles speciosa (Sweet) Nakai, 744, see, | Crataegus berberifolia Torr. & Gray, 2201, rip, roa, oe wet,N Crataegus crus-galli L., 2629, nov,N Crataegus intricata Lange, D. M. Moore, 56-55, UARK, N | ee | 5532,mes, nov pin ip, wet, Crataegus spathulata Michx., 5450, pin, rip, roa, wet, N Crataeg iflora Muenchh., 4508, pin, N Geen viridis L. var. viridis, 1023, roa, N Duchesnea indi ica (Andr.) Focke, 1523, rip, | Frag chesne, 1309, rip, roa, N Geum canadense Jacq., 5454, mes, nov, rip, see, N Photinia ise ifolia (Lam.) Robertson & o 1 447, see,N pycee opulifolius (L.) Maxim., 2330, mes, rip wet, N Porteranthus stipulatus (Muhl. ex Willd.) Britt, 38, lob, nov, roa, wee, N Potentilla recta L., 2282, nov, rip, roa, sha, | Potentilla simplex Michx., 11 ae 15, NOV, pin, rip, roa, KA n / 3920, Nov, pin, rip, roa, Prunus mexicana S.Wats., 5505, nov, pin, wet, N Prunus persica (L.) Batsch, 2617, wee, | P, ] Ehrh., 5522, lob, nov, pin, roa, wee, wet,N Prunus umbellata Ell., 4856, nov, N Pyrus calleryana Dcne., 571, roa, I* Pyrus communis L., 2616, wee, | Rosa carolina L., 2040, ober mes, Noy, pin, rip, roa, N Rosa chinensis Jacq., 5139, roa, | ner enunniiere Thunb. ex Murr. 1345, roa, I* vosa setigera Michx., 5462, pin, rip, roa, see, wee, Rubus argutus Link, 5032, lob, rip, see, wet, N Rubus bushii Bailey, 1126, pin, N Rubus discolor Weihe & Nees, 5091, ro Rubus flagellaris Willd., 1908, mes, nov, pin, roa, wee,N Rubus ostryifolius Rydb., 1997, pin, N Rubus pensilvanicus Poir., 2726, mes, nov, N Rubus trivialis Michx.,2131,roa, wee, wet, N MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS Spiraea xbilliardii he (pro sp.) [douglasii x salicifolia], 2797, roa, aceae Cephalanthus occidentalis L., 3301, rip, roa, wet, Cruciata pedemontana (Bellardi) Ehrend., 5101, a, | oe teres Walt., 388, nov, roa, wee, wet, N Diodia virginiana L., 5760, rip, wee, wet, N Galit ne L., 1444, mes, nov, rip, roa, see, N Galium aikannen Gray, 1999, mes, nov, pin, see 2419 Saxifragaceae euchera americana L. var. americana, 2763,mes, nov, pin, N Heuchera americana var. hirsuticaulis (Wheelock) sake Butters & Lakela, 4835, nov, Saxifraga peas Bush, 600, rip, N Scrophulariaceae Agalin ape ata (Ell.) Raf, 4386, lob, wet, N Agalinis aula: ia rive Raf.,4501, pin, rip, N Au (Benth.) Pennell, 439, pin, Galium circaezans Michx., 3700, lob, mes, pin, rip N Galium obtusum Bigel. ssp. obtusum, 4956, sha, wet,N Galium pilosum Ait., 5528, lob, rip, wee, wet, N Galium tinctorium lhe. 271 oe roa, wet, N Hedyotis nigricans (Lam.) Fosberg, 3896, rip, sha, Houstonia caerulea L., 4804, roa, sha, wet, N Houstonia a longi ifol ig Gaertn., 4980, pin, N oustonia | a (Shinners) Terrell, 679, roa, Houstonia ouachitana (E.B.Sm.) Terrell, 1107,mes, ov, pin, rip, N,S Houstonia purpurea L.,5451, mes, pin, rip, see, N Houstonia pusilla Schoepf, 716, mes, pin, rip, roa , 1398, lob, mes, pin, rip, see, N IVETE Sherardia arvensis L., 4734, rip, roa, wee, | Rutaceae Poncirus trifoliata (L.) Raf., Ae roa, wet, I* teas ea anal ata L., 2741, nov, a lava her rcul L., 4003, fOd, N Salicac popu usa ‘ba L ae wee, | P. x Marsh. ssp deltoides,R “Dalia: sn, UCAC, N Salix caroliniana Michx., wet, N 1089, mes, rip, roa, see, Salix nigra Marsh., 4975, roa, wet, N ees Car jospermum halicacabum ke A411 19, rip, N Sapotaceae Sideroxylon lanuginosum Michx., 2751, nov, pin, rip, wet, N se ae rip, roa, wet, N Aureolaria pectinata (Nutt.) Pennell, M. Stewart, 87-529, UAM, N ratiola brevifolia Raf., 3269, rip, N Gratiola neglecta Torr., 5033, wet, N Gratiola pilosa Michx., 3382, wet, N Lindernia dubia (L.) Pennell, 5476,rip, wee, we Mecardonia acuminata (Walt.) Small, 4384, | N Mimulus alatus Ait., 3278, mes, rip, see, wet, N Nuttallanthus canadensis (L.) D.A. Sutton, 1147, roa, Pedicularis canadensis L.,4725, mes, rip, N pills aan ae Pennell, 4989, pin, rip, roa, N italis Nutt. ex Sims, 1726, mes, rip, ae) ' aN Penstemon tubiflorus Nutt., 1834, mes, N - val aria marilandi ca L., 3961, mes, rip, N 8, roa, wee, wet, | akaseum thapsus L., 3746, wee, | Veronica arvensis L.,4722, rip, roa, wet, | Veronica peregrina L.,4710, wet Veronica persica Poir., 566, roa, | Farw., 3764, pin, N ae \ ; lente (| VEFOFHCOS CUTIE VIFGIFIICUTTE XL. Selaginellaceae Selaginella apoda (L.) Spring, D.M.Moore,430154, UARK, N Smilacaceae Smilax bona-nox L., 5046, lob, mes, Nov, pin, rip roa, see, wee, wet, N Smilax glauca Walt., 5045, mes, rip, see, wee, wet, N ilax herbacea L., 3632,nov,N ilax baa Hook., 1593, mes, N oS pea wet, N Smilax tamnoides L., 3425, rip, wet, N 2420 ia aa a stramonium L., Simpson, sn, UARK, | a. angulata L., 3867, w Physalis heterophylla Nees, a rip, roa, N Physalis Rapenr ies Le Ais mes, wet, N Solanu 30, rip, roa, wee, wet, N ate Dunal,4118,mes, nov, pin, Sparganiaceae Sparganium androcladum (Engelm.) Morong, 5772 : Staphyleac Staphylea ae ae 5455, rip, N Styracaceae Halesia tetraptera Ellis var. i (Rehd. Reveal & Seldin, D.M. Moore, 69007, UARK, N Styrax grandifolius Ait. 5471, mes, rip, N Ww helypteridaceae Phegopteris hexagonoptera (Michx.) Fee, 4341, mes, pin, rip, Thelypteris noveboracensis (L.) Nieuwl., 4245, rip, Thymelaeaceae Dirca palustris L., 708, mes, N Tiliaceae Tilia americana L.var. americana, 5443, mes, nov, N pin, rip, Tiliaamericana L. var.caroliniana (P.Mill.) Castigl.,, 2105, pin, yphaceae Typha angustifolia L., 5516, wet, | Typha latifolia L.,5545, roa, wet, N Ulmaceae Celtis laevigata Willd.,6014, lob, nov, rip, sha, wet, N Celtis occidentalis L.,R.D. Thomas, 128988, NLU, N Celtis tenuifolia Nutt., 1899, nov, pin, rip, sha, N ichx., 5051, pin, rip, roa, wet, N Ulmus americana L., 6005, nov, rip, see, wet, N wills EPunE L., hank roa, | bra Muh, 1943, pin, rip, see, N Urticaceae - Boehmeria cylindrica (L.) Sw., 3250, mes, rip, roa, BRIT.ORG/SIDA 21(4) Urtica chamaedryoides Pursh, M. C. Black, 43, UARK, N Valerianaceae Valerianella longiflora (Torr. & Gray) Walp., T. tsell,01-0251, UARK, N Valerianella nuttallii (Torr.& Gray) Walp., E. Sundell Valerianella palmeri Dyal, 4678, rip, N, SC Valerianella radiata (L.) Dufr., 1320, mes, pin, rip _sha, wee, wet, N Verbenaceae Callicarpa americana L.,5513,nov, pin, rip, wet, N Glandularia canadensis (L.) Nutt. 661, mes, roa, sha, N Phryma leptostachya L.,5449, mes, nov, pin, rip, N Verbena brasiliensis Vell.,4559, rip, wet, | Verbena stricta Vent., Simpson, sn, UARK, N V urticifolia L., 3999, mes, rip, wee, N PUCTIOC Violaceae Hybanthus concolor (T.F. Forst.) Spreng., 3670, mes, N Viola affinis Le Conte, 4887, mes, pin, rip, see, N Viola bicolor Pursh, 652, nov, roa, sha, wet, N Viola lanceolata L., 4728, rip, N Viola palmata L., 4812, pin, wet, N Viola pedata L., 587, mes, nov, pin, rip, roa, sha, wee, wet, Viola pubescens Ait., 790, rip, see, N Viola sagittata Ait., 653, lob, pin, rip, roa, sha, wet, N Viola sororia Willd., 732, mes, rip, see, N Viola striata Ait., 954, rip, see, N Viola villosa Walt., 779, rip, N Vitaceae Ampelopsis arborea (L.) Koehne, 4114, rip, roa, N Ampelopsis cordata Michx., 3472, mes, roa, N sl laces guinquefolia (L.) Planch., 5446, Novy, pin, rip, ser ae ; ie saestval Michx., 3596, me erea (Er elm) Millard var. cinerea, 5517 ,fOa, wee, wet, N a palmata Valli 4048, wet, N hx. var. rotundifolia, 5050, nov, pin Nov, rip pin, rip roa, see, wet, N Scheele, V. Bates, 10447, UVARK,N Vitis see, N Laportea canadensis (L.) Weddell, 4444, rip, N Pilea pumila (L.) Gray, 3879, mes, rip, see, Vitis vuloina L.,, 5744, mes, pin, rip, wee, wet, N Xyridaceae Xyris jupicai L.C. Rich., 3889, rip, N MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS 2421 ACKNOWLEDGMENTS I gratefully acknowledge the University of Arkansas Herbarium (UARK) for its continuous support of this research effort in the form of funds and resources as well as people who proved to be an invaluable resource and support system. Funding for the project was provided by UARK, the Ouachita National Forest (ONF), and the Arkansas Native Plant Society. The Ouachita Mountains Bio- logical Station provided an excellent place to stay and work while I was in the field. Ithank Johnnie Gentry for discussions on the directions the project should take and about the manuscript. Cindy Sagers and David Krementz also pro- vided ideas and assistance. Susan Hooks (ONF) was instrumental in providing information that allowed me to choose primary collection sites. Johnnie Gen- try, George Johnson, Sarah Nunn, Jim Peck, Chris Reid, and Eric Sundell helped in identifications. I thank Rob Doster, Johnnie Gentry, Edie Hardcastle, Terry McKay (ONF), Sarah Nunn, Matt Nutt, Chris Reid, John Shadwick, and Theo Witsell (ANHC) for their company and assistance in the field. Johnnie Gentry, Cindy Sagers, David Krementz, Leesia Marshall-Rosenberger, Sarah Nunn, Krista Clements Peppers, and Staria Vanderpool, as well as three anonymous review- ers commented on the manuscript. I especially thank my wife, Katie Marsico, who not only assisted in the field and reviewed the manuscript, but supported me with her love through it all. REFERENCES Arkansas Native PLANT Society. 2003. Invasive plants in Arkansas’ natural areas. Presented at the Rare and Invasive Plants of Arkansas Conference, 23-24 October 2003, Fayetteville, AR. Baiey, R.G. 1995. Description of the ecoregions of the United States. 2"? ed. rev. and ex- panded (1° ed. 1980). Misc. Publ. No. 1391 (rev.), USDA Forest Service Washington, DC. Batowin, J.R. 1984. Arkansas tornado distribution, recurrence probability and intensity: 1950-1982.M.A. Thesis, University of Arkansas, Fayetteville. Bates, V. 1993. Quachita National Forest, Volume |: Overview of plant communities and an inventory of natural areas. Unpublished Internal Report. Braven, A. 1999.Geologic map of Arkansas.Arkansas Geologic Commission, William V. Bush, State Geologist, Vardelle Parham Geology Center, Little Rock, Arkansas. Braun, E.L. 1950. Deciduous forests of eastern North America. Hafner Publishing Com- pany, Inc., NY. Comer, P,, D. FA8ER-LANGENDOEN, R. Evans, S Gaw er, C. Josse, G. Kittet, S. MENARD, M. Pyne, M. REID, K. Scuutz, K. Snow, and J. Teacue. 2003. Ecolagical Systems of the United States: A Working Classification of U.S. Terrestrial Systems. NatureServe, Arlington, VA. Date, E.E.and S. Ware. 1999. Analysis of oak-hickory-pine forests of Hot Springs National Park in the Ouachita Mountains, Arkansas. Castanea 64:163-174. Discs, G.M., Jr., B.L. Liescome, and R.J. O’Kennon. 1999. Shinners & Mahler's illustrated flora of north central Texas. Botanical Research Institute of Texas, Ft. Worth. 2422 BRIT.ORG/SIDA 21(4) Duncan, W.H. 1953. Taxonomic collections of vascular plants in the southeastern States— their abundance and relation to production of floras. Rhodora 55:353-358. Earvy, A.M. 2000. Results of the winding stair project.|n:Forest farmsteads:a millennium of human occupation at winding stair in the Ouachita Mountains. A.M. Early, ed. Arkansas Archeological Survey, Fayetteville, AR. Fiawn, PT. 1959. The Ouachita structural belt. In: The geology of the Ouachita Mountains: A Symposium. L.M. Cline, WJ. Hilseweck, and D.E. Feray, eds. Dallas Geological Society and Ardmore Geological Society, Dallas, TX. Guccione, MJ. 1993.Geologic history of Arkansas through Time and Space.M.J.Guccione (self published), Fayetteville, AR. Harocastte, E. and D.X Wittiams. 2000. Ptilimnium nodosum: Arkansas Status Report 2000. Unpublished document. NatureServe. 2003. Ecological Systems Database, version 1.01. NatureServe, Arlington, VA. Orzett, S.L.1985. Selected acid seep forests in the Ouachita National Forest—Preliminary Botanical Assessment. Unpublished Report for the Arkansas Natural Heritage Com- mission. Pacmer, E.J. 1926. The ligneous flora of Hot Springs National Park and vicinity. Journal of the Arnold Arboretum 7:104-135, Pyne, S.J. 1982. Fire in America:A cultural history of wildland and rural fire. Princeton Uni- versity Press, Princeton, NJ. Raororb, A.E., H.E. AHLes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. The University of North Carolina Press, Chapel Hill, NC. REINHOLD, R.O. 1969. Climates of the states: Arkansas. Climatography of the United States No. 60-3. U.S. Department of Commerce, Environmental Science Services Administra- tion, Washington, D.C. Rosison, H.W. and D.L. MarsH. Undated. Biodiversity of seeps and springs on the Caddo Ranger District, Ouachita National Forest, Arkansas. Final Report of Part | to USDA For- est Service, Ouachita National Forest, Caddo Ranger District. Rosison, H.W. and R.T. Auten. 1995. Only in Arkansas: A study of the endemic plants and animals of the State. University of Arkansas Press, Fayetteville. Scutty, EJ. 1937. Ferns of Hot Springs National Park. Amer.Fern J.27:50-62, Scutty, E.J.1941. The mosses of Hot Springs National Park and vicinity. Bryologist 44:125- Scutty, EJ. 1942. Grasses of Hot Springs National Park and vicinity. Rhodora 44:70-71. Smith, E.B.1988.An atlas and annotated list of the vascular plants of Arkansas. 2" Edition. Edwin B. Smith (self published). Smith, E.B.1994a. Keys to the flora of Arkansas. The University of Arkansas Press, Fayetteville. Smith, E.B. 1994b. Plant taxonomy booklet: Materials for BOTY 4104 Lecture and Labora- tory.P.A.No.410. Printing Services Professor's Assistance Program, University of Arkan- sas, Fayetteville. Smith, K.L.1986.Sawmill:the story of cutting the last great virgin forest east of the Rockies. The University of Arkansas Press, Fayetteville. MARSICO, FLORA OF MONTGOMERY COUNTY, ARKANSAS 2423 Snioer, L.V. 1982. Integrated multidisciplinary lineament analysis of the Ouachita Moun- tains of Arkansas and Oklahoma. Master's thesis. University of Arkansas, Fayetteville. Soi Survey Stare. 1998. Dominant soil orders and suborders. Soil taxonomy. 1998. Maps and Soil Photographs, USDA, NRCS. National Soil Survey Center, Lincoln, NE. Soi Survey STAFF. 1999. Soil taxonomy:A basic system of soil classification for making and interpreting soil surveys, 2" edition. USDA, NRCS. Stein, B.A., L.S. Kutner, and J.S. Apams (eds.).2000. Precious heritage: the status of biodiversity in the United States. Oxford University Press, New York, NY. USDA, NRCS. 2004. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. USDA Sot ConservarTion Service HANDBOOK 296, 1981. Land resource regions and major land resource areas of the United States. USDA. YarskievycH, G. 1999. Steyermark’s flora of Missouri—Revised Edition. Missouri Botanical Garden, St. Louis 2424 BRIT.ORG/SIDA 21(4) Book NOTICES Timber Press Jim JERMYN (text) with photographs by WILHELM and Dieter SCHACHT. 2005. Alpine Plants of Europe: A Gardener's Guide. (ISBN 0-88192-734-1, hbk.). Timber Press Inc. 133 S.W Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $39.95, 320 pp., color photos, 8 b/w figures, 71/2" X10 — d tl l | I lated. I found it hard to = ao the beak once pee it. Ni th he history of ape alpine plants to alpine gardening, fill in the space between the rnenas photos.—Barney Lipscomb, Botanical Research In- stitute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. JouN E. Bryan. 2005. Timber Press Pocket Guide to Bulbs. (ISBN 0-88192-752-2, flexibind). Timber Press Inc. 133 S.W Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $19.95, 227 pp,, 300 color photos, 5 3/4" X 81/4" As noted in the Preface, this book “is intended to give gardeners a greater appreciation of bulbs.” The } 1h ] t ]; } RPP ey | RB, 1] i ees [ees | } ; lf. Amita’ ] a | Ov to Zantedschia elliotiana. The Guide to Bulbs gives information to more than 700 species, cultivars, and hybrids. Everyday helpful information on such things as cu and diseases is also found in the Guide.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. — tivation, maintenance, and pests SIDA 21(4): 2424. 2005 MIKANIA BUCHTIENII (ASTERACEAE: EUPATORIEAE) NEW TO ARGENTINA Maria M.Cerana Luis Ariza-Espinar Facultad de Ciencias Agropecuarias Instituto Multidi io de Biologia Vegetal (IMBIV) a Nacional - doba CONICET- Uni iversidad Nacional de Cérdoba a de Correo Casilla de Correo 495 eu acs 5000 Cordoba, Argentina macerana@agro.uncor.edu arizaesp@imbiv.unc.edu.ar ABSTRACT Mikania buchtienii B.L. Rob. (Asteraceae, Eupatorieae) is first reported as occurring in Argentina, where it has been collected in the Yungas region of the northwestern Ba et Ene county alte and Jujuy provinces). The species, which was known only from Boliv key shows the differences between M. buchtienii and M. euryanthela, a closely related species. RESUMEN Mikania buchtienii B.L. Rob. (Asteraceae, E i imera vez par ra Argentina, donde C Ee ha sido colectada en las Yungas de la region noroccidental del pais (provincias de Salta y Jujuy). La especie, que vive en Bolivia, es descripta e ilustrada, y una clave muestra las diferencias entre M. buchtienii y M. euryanthela, una especie muy afin. During an extensive taxonomic study of the genus Mikania in Argentina (Cerana 1997a, b), two specimens from the northwestern Argentina (Yungas region) came to our attention. After a literature review and comparison with similar species, we concluded that it was necessary to relocate and collect the species in its natural habitat in order to determine its identity. In July 1997, Feb- ruary and November 1998, three trips were made to Las Capillas in Jujuy Prov- ince to search for the species, but without success. In October 2000, field stud- ies were conducted in San Pedrito, Salta Province, located about 45 km from Estacién Vespucio. One flowering specimen, which also had fruits from the previous year, was collected. Other plants present were in the vegetative stage. In addition to the field studies, specimens from the following herbaria were examined: BAB, CORD, CTES, JUA, LIL, LP LPB, SL Our research has shown that the collection and herbarium specimens represent the first records of Mikania buchtienii BL. Rob. for Argentina. This is also the first known occurrence of the species from outside of Bolivia. A specimen was sent to WC. Holmes of Bay- lor University in Waco, Texas, who confirmed our determination. Mikania buchtienii B.L. Rob., Contr. Gray Herb. n. s. 64:7. 1922. (Fig. 1). Twining lianas; stems terete, striate, glabrate. Leaves simple, coriaceous, oppo- site, petiolate, inconspicuously stipulate; petioles 10-30 mm long; blades from SIDA 21(4): 2425 — 2427. 2005 2426 BRIT.ORG/SIDA 21(4) cath We i a MWe fay, AY) Ws “IN COW Hes yp! HG . Fic. 1. Mikania buchtienii B.L. Rob. A. Leaf with deltate-hastate shape. B. Leaf with deltate-ovate shape. C. Achene. D Flowering branch. E. Head. F. Stamens. G. Floret. (A, B: Cerana 1740, CORD; C-G: Schulz 5482, LIL). Scale bar = 30 mm (A,B, D), 3 mm (C, E, G), 1 mm (F). deltate-hastate to deltate-ovate or ovate, 30-90 X 20-60 mm, margins entire to slightly serrate, bases subcordate to rounded, apices acute; upper surfaces gla- brescent, lower surfaces glabrate or slightly pilose. Capitulescence paniculate to cylindrical, dense, branches terete, terminal or axillary disposed; heads ca. 10 mm long, peduncles pubescent; subinvolucral bracts lanceolate, abaxially pubescent, 2.5-3.5 x 0.7 mm, located at the base of the peduncle or beneath the head; phyllaries 5-6.7 x 1.2-1.8 mm, apex obtuse or rounded, the outer bracts oblong, lanceolate or oblanceolate, abaxially pubescent, the inner ones, lan- ceolate, glabrate, apex ciliate. Corolla white, 5-6.5 mm long, tube 2.8-3.5 mm long, limb widely campanulate to campanulate 2-3 mm long, divided into lan- ceolate corolla lobes extending almost to the base (2.3-2.6 mm long); style gla- brous. Achenes 3.5-4 mm long, more densely pubescent apically, ribs serrulate; CERANA & ESPINAR TO ARGENTINA 2427 pappus yellowish white with 45-60 barbellate bristles, 5.5-7 mm long. Phenology.—Flowering specimens were collected from August to October, but it is assumed that flowering and fruiting occur from August to March. Habitat and distribution—The original description and previ ddit reports of M. buchtienii have been based on specimens collected in upland for- ests, riverine woodlands and mattoral areas in the Yungas region of Bolivia, at 1100-3500 m (Robinson 1922a, b). The new records extend the known range to the forests and borders of woods in the Yungas region of Salta and Jujuy, Argen- tina, at l1OOO-1200 m. This biogeographical province has a wet and humid cli- mate with over 2500 mm of rain annually and temperatures between 14° and 26° C (Cabrera 1971). ional $321 QUA, SD. Salta: aia de San Manta, San : Pednito ioses de ie ee dard), 25 Aug 194¢ Schulz 5482 (LIL, BAB, CTES); General José de San Martin, San Pedrito, Empresa Panamerican a ergy,a+200 m de la Historica Escuela, 19 Oct 2000, Cerana tibeecerces BAYLU BOL az: Nor Yungas, Yolosa hacia Chuspipata Bajo y el derruml grande, ea 16'S, 67° 47° W), 19 Sep 1995, Beck 22468(L. PB); Inquisivi, “Rio C ae Ww elewioe the Rio Churu from 200 m W of Aguilani to | km above the main fork up river, 3 km SE of the ruins of Choquecamiri, 17 km N of Choquetanga (16° 42'S, 67° 20’ W), 12 Sep 1991, Lewis 40208 (LPB), Inquisivi, “Aguas Calientes de Calachaca’, in the area of the thermal springs of the Rio Calachaca Jahuira, 9 km NW of Choquetanga (16° 48'S, 67° 19’ W), 9 Mar 1991, Lewis 38244 (LI Mikania buchtienii is closely related to Mikania euryanthela (Malme) WC. Holmes, which is widespread in Paraguay, southern Brazil (State of Parana) and the provinces of Corrientes, Chaco, Formosa and Misiones in Argentina (Barroso 1959; Holmes & McDaniel 1996; Cerana 1997a, b). Both species are similar in leaf blade shape, corolla shape and very long corolla lobes. The main differ- ences between them are summarized in the following key: KEY TO DISTINGUISH MIKANIA BUCHTIENII AND M. EURYANTHELA . Leaf blades membranaceous, deltate-ovate lat te, glabrate; capitulescence paniculate, scattered; Saeer bracts slightly pubescent, a a prominent middle nerve; corolla glabrate, limb 2—-2.3 mm long divided in corolla lobes 1.5-1.7 mm long; achenes see ous Mik scuanel: Leaf blades coriaceous, deltate-hastate, deltate-ovate, or ovate, bases subcordate or rounded, upper surfaces glabrescent, lower aura glabrate or slightly pilose; capitulescence paniculate to cylindrica ucral bracts pubescent, with an inconspicuous middle net lla pubescent, limb 2-3 mm long divided into corolla lobes 2.3-2.6 mm long; achenes pubescent Mikania buchtienii ny ACKNOWLEDGMENTS We are grateful to curators of the herbaria BAB, CORD, CTES, JUA, LIL, LP, LPB, SL, and St. Beck, director of the herbaria LPB for their loan of specimens. We appreciate M. Nee (NY) for sending us digital images. We thank WC. Holmes of Baylor University, Waco, Texas, for assistance in confirming the identity of our 2428 BRIT.ORG/SIDA 21(4) plant. Guy Nesom and an anonymous reviewer offered helpful improvements. We thank managers of Panamerican Energy for providing access to the prop- erty and transportation while on-site. We thank A.P de Pereyra (FC.A., Univer- sidad Nacional de Cordoba, Argentina) for assistance with the English version of the manuscript. Financial support for the research was provided by Agencia Cordoba Ciencia and Fondo para la Investigacién Cientifica y Tecnologica (FONCyT). Thanks are also due to N. de Flury and L. Ribulgo for drawing the excellent illustration. al REFERENCES Barroso, G.M. 1959. Mikaniae do Brasil. Arq. Jard. Bot. Rio de Janeiro 16:38-333. Casrera, A.L. 1971. Fitogeografia de la Republica Argentina. Bol. Soc. Argent. Bot. 14:1—42. CerANA, M.M. 1997a. El género Mikania (Asteraceae) en la Republica Argentina. Aspectos taxonomicos y morfologicos. Tesis Doctoral. Facultad de Ciencias Exactas, Fisicas y Naturales, Universidad Nacional de Cordoba, Argentina. Pp. 1-257. Cerana, M.M. 1997b. Mikania. In: A.T. Hunziker, ed., Tribu Eupatorieae, Fl. Fanerog. Argent. 47:54—76. Hotmes, W.C.and S.McDaniet. 1996, Mikania.|n:Spichiger y Ramella, eds., Flora del Paraguay 25:208-271. Rosinson, B.L. 1922a. Records preliminary to a general treatment of the Eupatorieae, - |. Contr. Gray Herb. n.s.64:1-21. Rosinson, B.L. 1922b. The Mikanias of northern and western South America. Contr. Gray Herb. n.s.64:21-116. RANUNCULUS FICARIA (RANUNCULACEAE), NEW TO NORTH CAROLINA AND AN UPDATED KEY TO CAROLINA CONGENERS Alexander Krings Alan S.Weakley Herbarium, Department of Botany Herbarium North Carolina State University University of North Carolina Raleigh, North Carolina 27695-7612, U.S.A. North Carolina Botanical Garden Alexander_Krings@ncsu.edu Chapel Hill, a Carolina 27599-3280 kley@unc.edu Joseph C. Neal ee C. Swab Herbarium, Department of Botan North Carolina State University Raleigh, North Carolina 27695-7612, U.S.A. Department of Horticultural Science North Carolina State University Raleigh, i Carolina 27695-7609, U.S.A. e_Neal@ncsu.edu ABSTRACT Populations of the expanding exotic Ranunculus ficaria L. (Ranunculaceae) are reported escaped in North Carolina. Voucher specimens are cited and an updated key to Ranunculus L. in the Carolinas is presented. RESUMEN Poblaci le la exoti pansion R lusficaria L. (Ramunculace7®) se Sha cance Carolina del Norte. Se citan | peci y se presenta Ranunculus L.en las Carolinas. Populations of the exotic Ranunculus ficaria L. (lesser celandine) are known outside of cultivation from the northeastern United States, Oregon, and Wash- ington (Gleason & Cronquist 1991; USDA, NRCS 2005), but the species has been previously collected in the Southeast only in Kentucky, Tennessee, Virginia, and Maryland (Whittemore 1997). Ranunculus ficaria is native to much of Europe (Tutin 1976; Taylor & Markham 1978). The species was reported in New York State as early as 1890 (Hollick & Britton 1891). By the mid 1940s, it was known from several northeastern coastal states, as well as Virginia (Sargent s.n., NCSC), but had not yet reached West Virginia (Bell 1945). The taxon was not previ- ously reported for North Carolina by Radford et al. (1968), Pittillo et al. (1969), Pittillo et al. (1972), Kral (1981), Pittillo and Brown (1988), or Wibiie mere (1997). However, a population of the species was recently t n Raleigh (Wake Co., North Carolina). Twenty-nine clumps were observed, of eri seventeen were either in flower or fruit at the time of collection. Individuals were observed SIDA 21(4): 2429 — 2437. 2005 2430 BRIT.ORG/SIDA 21(4) only in areas receiving partial shade for a significant portion of the day. They were absent from portions of the lawn receiving full sunlight. Voucher collection: U.S.A. North Carolina. Wake Co.: Raleigh, backyard lawn of residential home on Van Dyke Avenue, in partial shade, absent from portions of lawn receiving full sun, 11 Apr 2005, Krings 1271(AUA, F FLAS, GA, LSU, MISS, NCSC, NCU, TEX, UNA, US, USE USCH, VDB) The species has also been observed on a private property in Chatham Co., where it had apparently been planted in the past, but is now escaping short distances into nearby natural areas. Due to summer senescence, vouchers of this popula- tion have not been taken this year. Several attempts have been made to morphologically distinguish diploid from polyploid plants of R. ficaria (Marsden-Jones 1935; Marsden-Jones & Turrill 1952; Lawalrée 1955). Unfortunately, most of these have proven unreliable (Heywood & Walker 1961; Jones 1966; Taylor & Markham 1978). Arguing that too great a reliance had been placed on chromosome counts, Sell (1994) recog- nized five subspecies (Table 1), with the caution that these taxa could be recog- nized with ease only if cultivated or examined at intervals through their flow- ering and fruiting periods. If only seen once in the field or from a single specimen, certain identification would remain difficult. The following key was provided by Sell (1994) to facilitate identification: 1. Leaf blades to 8 X 9 cm; petioles to 28 cm; flowers to 60 mm diam; achenes to 5.0 xX 3.5mm 2. Stems rather robust, but straggling; bulbils present in leaf axils after flowering subsp. a 2. Stems robust and erect; without bulbils in leaf axils after flowering recente 1. Leaf blades to 4 X 4 cm;petioles to 15 cm; flowers to 40 mm diam; achenes to 3.5 X 2.2mm eaves crowded at base with few on short ste ae calthifolius 3. Leaves less crowded at base and more numerous on the elongate st 4, Bulbils not present in leaf axils after flowering; achenes well- ios subsp. ficaria subs oe) 4. Bulbils present in leaf axils after flowering; achenes poorly developed Sp. bulbilifer USDA, NRCS (2005) reported only R. ficaria var. bulbifera Marsden-Jones for the United States. Sell (1994) pointed out that this name is illegitimate, being a later homonym of R. ficaria var. bulbifer Albert, which Sell treated in synonymy under subsp. bulbilifer Lambinon. Based on Whittemore’s (1997) description of the species, the flower sizes of individuals of subsp. bulbilifer in the United States would be on the upper end compared with European individuals as rec- ognized by Sell (1994) (Table 1). Following Sell (1994), the individuals of the Wake Co., North Carolina popu- lation are tentatively referable to subsp. ficariiformis (EW. Schwartz) Rouy & Fouc, previously not reported for the United States. Though there are some more KRINGS ET AL TABLE 1. The five tioles; ptl 2431 subspecies of Ranunculus ficaria L. recognized by Sell (1994). Fls=flowers; ree ls=petals. subsp. bulbilifer Lambinon subsp. calthifolius subsp. chrysocephalus subsp. ficariiformis subsp. ficaria (Reichenb.) Arcangeli _P.D. Sell (F.W. Schultz) Rouy & Fouc. Tetraploid Diploid Tetraploid Tetraploid Diploid Ptio< 15cm Ptio<7cmlong PtioS21cmiong Ptio S 28cm Ptio< 15cm long long long Axillary bulbils Axillary bulbils Axillary bulbils Axillary bulbils fe bulbils globular absent ovoid or bse globular Fils = 25mm Fls = 30mm Fils = 60 mm Fis =50mm Fls 20-40 mm diam Ptls 6-11 diam Ptls 10-15 diam Ptls 18-25 diam Ptls 17-26 diam Ptls 10-20 2-5 mm, not 5- ; < 9-15(-18)mm, 4-12 mm, 4-9 mm, often contiguous not contiguous contiguous or contiguous or contiguous overlapping overlapping Occurs through- Restricted to east- Occurs in the Occurs in the Restricted to out mostrange — centraland south-eastern central and western Europe of the species, eastern Europe Mediterranean western although rare in region Mediterranean Mediterranean Region region diminutive plants in the population, a number of individuals bear leaves > 4 cm diam and petals = 17 X 6 mm, that are contiguous to overlapping (see Table 1 for a comparison of characters among subspecies). On some plants, ellipsoid axillary bulbils are evident. However, subspecific taxa of R.ficaria are not uni- formly accepted. Citing extensive intergradation in form, Whittemore (1997) did not recognize any subspecific taxa in his treatment of Ranunculus L. for the Flora of North America. To help address the continuing disparity in taxo- nomic treatments, detailed studies are needed to examine the distribution of haplotypes and potential morphological correlations. Such an approach has been useful for other expanding exotic plants (e.g., Saltonstall 2002, 2003a-c; Saltonstall et al. 2004). Further investigation is also needed to determine more precisely the timing and mode of introduction, as well as the rate of spread of the species. Survey of herbarium collections may shed more light on this mat- ter. Pertinent studies regarding the life history, pollination biology, and ecology have been published by Marsden-Jones (1935, 1936) and Taylor and Markham (1978). To improve collections, Sell (1994) suggested that specimens should be taken late when fruit and bulbils are developed. However, at this stage any flow- ers remaining open are typically late ones, which are generally smaller than 2432 BRIT.ORG/SIDA 21(4) Fic. 1.2 lus ficaria L. A. Habit: B. Flowers: C. Tuberous roots. B d Kri 1271 i ~s KRINGS ET Al 2433 those when the plant first came into flower. To improve our understaning of the distribution and ecology of subspecies, botanists should note flower sizes on an initial visit to populations and then check the bulbils and fruits at a later date (Sell 1994). Ranunculus ficaria (Fig. 1) can be distinguished from its Carolina conge- ners by the combination of unlobed, reniform to suborbicular leaves, tuberous roots, typically three sepals, yellow petals > 10 mm long (sometimes partially fading to white with age), and pubescent, beakless achenes. An updated key to Carolina congeners, largely adapted from Whittemore (1997), is provided be- low. Following arguments presented by Nesom (1993), we diverge from Whittemore (1997) in treating R. carolinianus DC. as a species, rather than asa variety of R. hispidus Michx. Distribution, habitat, and frequency information follows Weakley (2005) (Mt = Mountains; Pd = Piedmont; Cp = Coastal Plain). Unless otherwise indicated, provincial distributions and comments apply equally to North Carolina (NC) and South Carolina (SC). Asterisks indicate ex- otic species. KEY TO RANUNCULUS IN THE CAROLINAS 1. All leaves unlobed 2. Leaf blades reniform to suborbicular or orbicular, bases shallowly to deeply cordate. 3. Roots tuberous; petals 2 10 mm long; achenes pubescent beaks abse . ficaria L. Hs (NC); Deen rich forests and bottomlands, mesic suburban forests, law aturalize locally from ees ae rare] } | | an | urved 3. Roots filiform; petals < 3.5 mm long; a¢ g Bi apOrey us: 7 [Mt, Pd, Cp; au fields, disturbed areas, bottomlands, lawns 2. Leaf blades ovate to lanceolate, bases truncate, rounded-obtuse to cuneate ee cordate in R. laxicaulis, then petals 2-6 mm lon Ae 4, Petals 1-3, 1.5-2 mm long R. pusillus Poir. (Mt (NC), Pd, Cp; marshes, ditches, other wet habitats; common (uncommon in Mt)] 4. Petals 4-6,5-8 mm lon 5. Proximal cauline leaf blades 5.9-12.2 cm long; petals 5; achenes to 1.8mm long R. ambigens S. Wats. [Pd (NC), Cp (NC); marshes; rare] 5. Proximal cauline leaf blades to 5.7 cm long; petals 4-6; achenes 0.8-1 mm long R. laxicaulis (Torr. & A. Gray) arby [Cp; marshes; rare] _ All or some leaves lobed or compound. 6. Leafy stems creeping and rooting at the nodes, or floating in water (then rootless). ie ae 3-foliolate. 8. Achene margins 0.4-1.2 mm wide R. carolinianus DC. [Mt, Pd, Cp; swamp forests, wet woodlands, open marshy wetlands; uncommon] 2434 BRIT.ORG/SIDA 21(4) 8. Achene margins 0.1—0.2 mm wide *R. repens L. [Mt (NC), Pd (NC), Cp; low meadows, disturbed areas; uncommon] 7. Leaves simple, lobed, parted, or dissected. = 1 cm long; floral receptacles glabrous; petals white; achenes < 1.6 mm long R. hederaceus L. [Cp; oastal brackish marshes, other circumneutral soils; rare] 9. Leaves = 1.2 cm long; floral ee. sparsely hispid; petals yellow achenes = 1.8mm long R. flabellaris Raf. [Cp (NC); pools in ecu of small stream swamps, other stagnant or slow moving waters; rare] 6. Leafy stems erect or if pani nee rooting only at the base (rarely cone at the nodes in R. sceleratus), never floating. 10. Style absent; achene margins atk and corky; emergent ee or on wet soil R. sceleratus L. [Pd (NC), Cp; marshes, ditches, and stream margins; uncommon] 10. Style present; achene el not corky; various habitats, but not aquatic. 11. Basal leaves variously unlobed to deeply divided; achenes thick-lenticu- lar or asymmetrically thick-lenticular to ae 1.2-2 times as wide as thic 12. Stems villous R. micranthus Nutt. [Pd (NC); rich forests; rare} 12. Stems glabrous. 13. Sepals glabrous; achene beaks 0.1-0. ai mm long R.abortivus L. Pd, Cp; low Nei: disturbed areas, potomiad lawns ] 13. Sepals hispid;achene beaks 0.6-1 mm long______ R. alleghaniensis Britton Mt (NC, SC?); cove forests, rich forested slopes; uncommon] . Basal leaves always deeply lobed or celia achenes strongly com- pressed, at least 3-15 times as wide a 14. Achenes spinose or papillose (sometimes smooth in R, sardous). 15. Petals 1-2 mm long; receptacles glabrous; achenes finely papil- late,each with a hooked bristle. 6. Flowers pedicellate; sepals 5 *R. parviflorus |. [Mt, Pd, Cp; disturbed areas; common (rare in Mt)] 16. Flowers sessile; sepals 3 *R, sila Spreng. [Pd (NC); lawns and ditches; rare] 15. Petals 4-10 mm long; receptacles pilose or hispid; ee coarsely papillate (but not terminating in hooked bristles), spi- nose, or tuberculate. 17. Sepals 5} ling;achenes 5—9, borne in a single whorl, lon spinose *R. arvensis a [Mt (NC), Pd: fields, disturbed areas; rare] 17. Sepals reflexed; achenes 13-60, borne in ovoid or globose heads, papillose to spinose, 18. Basal leaves simple; achene beaks 2—2.5 mm es icatus L. [Pd (SC), Cp (SC); ditches sh ae rare] 18. Basal leaves compo ind; achene beaks to 0.7 mm long. KRINGS ET AL., 2435 19. Petals 7-10 mm long; achenes sparsely papillate or sometimes smooth *R. sardous Crantz [Pd, Cp; low fields, disturbed areas; uncommon] 19. Petals 4-5 mm long; achenes kat tuberculate .trilobus Desf. [Cp (SC); sere roadsides, ditches; rare] 14. Achenes smooth, glabrous or pubesce 20. Petals 3-5 mm long; achene ae markedly recurved R. recurvatus Poir. (Mt, Pd, Cp; bottomland forests, cove forests, swamps, mesic slope forests; common] 20. Petals = 7 mm long; achene beaks more or less straight, not markedly recurved. . Sepals ae along a defined fold 1-3 mm above base. 21 22. Stem bases bulbous, corm-like; petals 9-13 mm X 8-11 mm *R. bulbosus L. [Mt, Pd, Cp; fields, roadsides, disturbed areas; common re in South Carolina)] 22. Stem bases not bulbous; petals 7-10 * 4-8 mm *R. sardous Crantz [Pd, Cp; low fields, disturbed areas; uncommon] 21. Sepals spreading (sometimes reflexed from base with age 23. Basal leaf blades 3-5-parted, pentagonal in outline *R. acris L.[Mt (NC), Pd (NC), Cp; pastures, fields, roadsides, disturbed areas; common (uncommon in Pd, rare in Cp)] 23. Basal leaf blades 3-5- nee ovate to ene in outline. 4. Tuberous roots abse R. hispidus Michx. [Mt, Pd; rich moist forests, creek banks, mesic to dry woodlands and forests, bottomlands; common] 24, Tuberous roots present R. fascicularis Mubhl. ex Bigelow [Mt (NC), Pd; wet flats with prairie affinities, rocky rens and glades over mafic rocks, ultramafic outcrop barrens, limestone barrens; rare] ACKNOWLEDGMENTS We are grateful to Charles W. Averre (Prof. Emeritus, Dept. Plant Pathology, NCSU) for bringing the plant to our attention and the thoughtful reviews of a previous version of the manuscript by Michael Denslow, Carl Keener, and an anonymous reviewer. REFERENCES Bett, FH. 1945. The genus Ranunculus in West Virginia. Amer. Midland Nat. 34:735-743. Gteason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York Botanical Garden, New York. 2436 BRIT.ORG/SIDA 21(4) Heywooo,V.H.and S.Watker. 1961.Morphological separation of cytological races in Ranun- culus ficaria L. Nature (London) 189:604. Houtick, A.and N.L. Britton. 1891. Flora of Richmond Co.,N.Y—Additions and new localities, 1890. Bull. Torrey Bot. Club 18:213-214. Jones, B.M.G. 1966. Variation in Ranunculus ficaria. Proc. Bot. Soc. British Isles 6:275. Kral, R.1981.Some distributional reports of weedy or naturalized foreign species of vas- cular plants for the southern states, particularly Alabama and middle Tennessee. Cas- tanea 46:334-339. Lawatrte, A. 1955. Flore Générale du Belgique. Vol. 2. Bruxelles. Marsbden-Jones, E.M. 1935. Ranunculus ficaria Linn.: Life-history and pollination. J. Linn. Soc London (Bot.) 50:39-55. Marspen-Jones, E.M. 1937. Pollination of Ranunculus ficaria L. by insects. J. Bot. (London) 75:133-141. Marsben-Jones, EM. and W.B. Turritt. 1952. Studies on Ranunculus ficaria. J. Genetics 50: 522-534. Nesom, G.L. 1993. Ranunculus (Ranunculaceae) in Nuevo Ledn, with comments on the RP. petiolaris group. Phytologia 75:391-398. Pirtito, J.D., J.H. Horton, and K.W. Greencee. 1969. Additions to the vascular flora of the Caro- linas.|.J. Elisha Mitchell Sci. Soc. 85:18-22. Pirtito, J.D., J.H. Horton, and K.E. Herman. 1972. Additions to the vascular flora of the Caroli- nas. Il. J. Elisha Mitchell Sci. Soc. 88:144-152 Pirmitto, J.D.and A.E. Brown. 1988. Additions to the vascular flora of the Carolinas. Ill. J. Elisha Mitchell Sci. Soc. 104:1-18. Pocan, E. and H. Wcissto. 1986. Studies in Ranunculus ficaria L. Vil. Additions to chromo- some numbers. Acta Biologica Cracoviensia, Series Botanica 28:87-92. Raproro, A.E., H.E. AHies, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. The University of North Carolina Press, Chapel Hill. SALTONSTALL, K. 2002. Cryptic invasion of a non-native genotype of the common reed, Phragmites australis, into North America. Proc. Natl. Acad. Sci. USA 99:2445-2449, SALTONSTALL, K.2003a. Microsatellite variation within and among North American lineages of Phragmites australis. Molec. Ecology 12:1689-1702. SALTONSTALL, K. 2003b. Genetic variation among North American populations of Phragmites australis: implications for management. Estuaries 26(2B):445-452, SALTONSTALL, K. 2003c. A rapid method for identifying the origin of North American Phragmites populations using RFLP analysis. Wetlands 23:1043-1047. SALTONSTALL, K., PM. Peterson, and R. Sorene. 2004. Recognition of Phragmites australis subsp. americanus (Poaceae: Arundinoideae) in North America: evidence from morphologi- cal and genetic analyses. Sida 21:683-692. Set, PD. 1994. Ranunculus ficaria L.sensu lato.Watsonia 20:41-50. Taytor, K.and B.MarkHam. 1978. Biological flora of the British Isles Ranunculus ficaria L.(Ficaria verna Huds.; F ranunculoides Moench). J. Ecology 66:1011-1031. a KRINGS ET Al 2437 Tutin, T.G. 1964. Ranunculus L.|n:T.G. Tutin et al., eds. Flora Europaea, Vol.1 Cambridge Uni- versity Press, Cambridge. Pp. 223-237. USDA, NRCS. 2005. The PLANTS Database, Version 3.5 (http://plants.usda.gov). Data com- piled from various sources by Mark W. Skinner.National Plant Data Center, Baton Rouge. Weaktey, A.S. 2005. Flora of the Carolinas, Virginia,and Georgia:Working draft, 10 June 2005. University of North Carolina Herbarium, Chapel Hill. Wuittemort, A.T. 1997. Ranunculus.In: Flora North America Editorial Committee (eds.), Flora North America, Vol. 3. Oxford University Press, New York. Pp. 88-135. 2438 BRIT.ORG/SIDA 21(4) BOOK NOTICE Cuares Borwe (ed.). A C.S. Rafinesque Anthology. 2005. (ISBN 0-7864-2147-9, pbk.). McFarland & Company, Inc., Box 611, Jefferson, NC 28640, U.S.A. (Orders: wwwincfarlandpub.com, 800-253-2187). $45.00, 271 pp., 17 b/w figures, Fie wo From the back ee ee American naturalists, C.S. Rafinesque (1783-1840) is second only to Audubon in the popular interest he eas This interests is due in part to his colorful life and pro- vocative personality, but he is al Qo. — or devising Latin scientific names for more plants than any other naturalists who ever lived ~ ae a erent number in the animal kingdom, as well. This passion for nome nclature has ke pt would Say g natural- ists. Yet his taxonomic writings made up only a part of his ext 4 There are at least six previous books on Scie also edited by Charles Boewe. Rafinesque’s essays covered in this Saline are 1) Antiquities, 2) Linguistics, 3) Society, 4) Education, 5) Public Lectures, 6) Popular Science, 7) Phytogeography, 8) Natural Science, and 9) Metaphysics A great fan of Rafinesque was the Canadian born botanist, Lloyd H. Shinners (SMU). Ruth Gins- burg, in her 2002 biography, Lloyd Herbert Shinners: By Himself, discusses Shinners’ admiration for Ratinesque. The following excerpt gives some insight into the life of Rafinesque and his influence on at least one botanist: “Constantin Samuel Rafinesque was a naturalist, traveler, and writer. sh in Constantinople, he spent much of his youth in Italy and came to the United States in 1815. He was not trained in Botany) but he read widelya in ibaa sanguages and became deeply et in natural history, well as botany. Shinners admired his wide knowl- edge, his voracious appetite for dicsonere aac publication and scpacielly his courage in defending unpopular positions. Lack of eaplarnacy, organization, and understanding of their limitations were other characteristics that they shared. If you enjoy reading about great naturalists then you will definitely enjoy Boewe's A CS. Rafinesque Anthology.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A SIDA 21(4): 2438. 2005 THE OCCURRENCE OF CERASTIUM PUMILUM (CARYOPHYLLACEAE) INOKLAHOMA Bruce W. Hoagland Amy Buthod Oklahoma Biological Survey Oklahoma Biological Survey and Department of Geography University of Oklahoma BY of Oklahoma Norman, Oklahoma 73019, U.S.A. Norman, Oklahoma 73019, U.S.A. ee bhoagland@ou.edu Gloria Caddell Department of Biology University of Central Oklahoma Edmond, Oklahoma 73034, U.S.A ABSTRACT Cerastium pumilum (Caryophyllaceae) was recently reported for Oklahoma without documenta- tion. This paper documents the occurrence of Cerastium pumilum W. Curtis in Oklahoma. The first collection from Oklahoma was made in 1981 from Hughes County but no additional collections were made until 2003. Since that time, C. pumilum has been documented in eight additional counties. Thus, C. pumilum is currently known from a total of 11 Oklahoma counties RESUMEN ie 12 | C } 1] rer : de O\vlah : ] ae Fste L an lod | iadeC ti ilum W.Curtisen Oklahoma. La pri ] de Oklahoma se Rese en 1981 en Bughe an poe no se plelenn epecelOnes adicionales hasta 2003. Desde entonces, C. pumilum se h Asi pues, C. pumilum — se conoce actualmente de un total de 1] condados de Oklahoma. Cerastium pumilum W. Curtis (Caryophyllaceae) is an annual plant species native to Europe and southwest Asia (Diggs et al. 1999). It has been reported from the adjacent states of Arkansas, Kansas, Missouri, and Texas, but not Colo- rado (USDA-NRCS 2005). Morton (2005) mapped C. pumilum in Oklahoma but did not provide voucher documentation. The Oklahoma distribution record in Morton (2005) is based on a 1981 collection by Steve Stephens. Voucher collection: OKLAHOMA. Hughes Co.: 3.5 mi W of Calvin, roadside shoulder, red sandy soils, few plants, 7 Apr 1981, Steve Stephens 90325 (KANU). However, C. pumilum is not listed in Taylor and Taylor (1994) and a search of Hoagland et al. (2005) revealed that no collections had been deposited in Okla- homa herbaria prior to 2003, when the authors collected it from three counties. Since then, it was collected from five additional counties in 2004 and three in 2005. All specimens reported below are deposited in the Robert Bebb Herbarium at the University of Oklahoma (OKL), with the exception of Caddell and Rice SIDA 21(4): 2439 — 2440. 2005 2440 BRIT.ORG/SIDA 21(4) (GMC 1039) which is deposited at the University of Central Oklahoma (CSU). Since these specimens represent locales covering a broad geographic extent, additional county records are no doubt forthcoming. Bac oneL van -ace specimens ORT AY OMA. Beckham Co.: 1.1 mi E of 1-40 from jet. with Hwy 34, TIN, R22W, sec 35, disturbed area, 3 Apr 2003, Buthod and Hoagland, AB3649. Cleveland Co.: Nor- man, TON RIW sec 32, disturbed area, 7 May 2005, Amy Buthod, AB6762. Kay Co.: Blackwell city a T27N, RLW, sec 23, disturbed area, 29 May 2003, Buthod, Hoagland, and Fagin, AB3822. Lincoln er Turnpike (I-44) at exit 171, TI5N, R5E, sec 32, disturbed grassland, 6 May 2004, Buthod and ae AB6155. Major Co.: ca. 3 mi W of Bouse Junction on Hwy 412, T22N, R1OW, sec 6, grazed pasture, 22 Apr 2004, Buthod and Shannon, AB4563. Oklahoma Co.: Forest Park in Oklahoma City, T12N, R3W, sec 13, disturbed grassland, 27 Apr 2004, Buthod, Hoagland, and Fagin, AB4609. Oklahoma Co.: Lake Arcadia, TI14N, R2W, sec 34, disturbed area, 3 Mar 2005, Amy Buthod and Lacy Brookshire, AB6251. Ottawa Co.: Approximately 7.2 mi NE of Grove, T26N, R23E, sec 33, disturbed area, 22 Apr 2004, Buthod, ree a Brookshire, AB4544. Pawnee Co.: 5 mi W of Cleveland on Hwy 64, T2IN, R7E, sec 15, disturbed area, 1 Apr 2004, Buthod and Hoagland, AB4521 Stephens Co.: 1.6 mi SW of Bray at Clear Lake, T2N, ROW sec 29, disturbed area, 13 Mar 2003, Hoagland and Buthod, AB3421, Woodward Co.: Alabaster Caverns State Park, T26N, RI8W, sec 28, disturbed area along road- side near Canyon Campground, 7 May 2005, Caddell and Rice, GMC 1039. ACKNOWLEDGMENTS We thank Richard Rabeler for reviewing the paper and for bringing the Stephens specimen to our attention and Craig Freeman for providing the label data from that specimen. REFERENCE Diccs, G.M, B.L. Liescome, and RJ. O'KeNNoN. 1999. Shinners & Mahler's illustrated flora of north central Texas. Botanical Research Institute of Texas (Fort Worth) and Austin Col- lege (Sherman, TX). HOAGLAND, B., A. BUTHOD, |. BUTLER, P. CALLAHAN-CRAWFORD, W. EN A. UDASI, ane R. Tyat. 2005. Oklahoma vascular plants database. [online]. Available: http://www.biosurvey.ou.edu. (Accessed on 1 March 2005). Morton, J.K.2005.Cerastium Linnaeus. In:Flora of North America Editorial Committee (eds), Flora of North America north of Mexico, vol.5 Magnoliophyta: Caryophyllidae, part 2. New York (NY): Oxford Press. Tayior R. J.and CS. Taytor. 1994. An annotated list of the ferns, fern allies, gymnosperms, and flowering plants of Oklahoma. Southeastern Oklahoma State University, Durant. USDA-NRCS 2005. The PLANTS database [online]. Available: http://plants.usda.gov/plants. National Plant Data Center, Baton Rouge, LA. (Accessed on 1 March 2005). FIRST RECORD OF NYMPHOIDES INDICA (MENYANTHACEAE) IN TEXAS Ken Saunders Freshwater Resources cl Inland Fisheries Divisi Texas Parks and Wildlife ee San Marcos, Texas 78666, U.S.A. ABSTRACT The first Texas record of N d new for Texas. RESUMEN Se cita por primera vez para Texas Nymphoides indica (L.) Kuntze (Menyanthaceae). Nymphoides specimens were collected from Soldier Springs, Uvalde County, Texas in April 2002 and subsequently planted in small water gardens upon return to the laboratory. Specimens from this locale had previously been curso- rily identified as N. aquatica (Bill Carr, The Nature Conservancy of Texas, pers. comm.). However, upon inspection of flowers from reared specimens, it was determined that the species was instead N. indica (independent species verifi- cation by David Lemke, Texas State University Biology Department), which rep- resents a new species record in Texas. Voucher specimen: U.S.A. Texas. Uvalde Co.: Soldier Springs, ca 2 km S. of the U.S. Hwy 90 bridge and a short distance upstream of County Boat ze Som Nunn crossing), floating in water with Trichocoronis rivularis and other species, f 1 pool, 18 Apr 2002, Saunders & Lemke s.n. (SWT). It has since been observed in at least one other spring system upstream of Sol- dier Springs, also within the Nueces River basin (Chad Norris, Texas Parks and Wildlife Department, pers. comm.). Two native Nymphoides species are known to occur in the southeastern United States (Wood 1983; Jacono 2002), namely N. cordata ( Ell.)Fern and N. aquatica) (J.E Gmel.) Kuntze. Nymphoides indica (L.) Kuntze (water snowflake) is native to both New and Old World tropical regions, but like many other in- troduced aquatic macrophytes, has been brought to the United States for orna- mental uses. Ornduff (1969) did not find consistent morphological difference between New World plants known as N. humboldtiana (H.B.K.) Kuntze, and Old World plants known as N. indica (L.) Kuntze. Ornduff thus concluded the two must be conspecific and since the name N. indica has priority both Old and New World specimens should be referred to by this name. Occurrence of wild populations of N. indica, until now has been documented only in two SIDA 21(4): 2441 — 2443. 2005 2442 BRIT.ORG/SIDA 21(4) southwestern counties in Florida (Wunderlin & Hansen 2004) where it is spreading rapidly Jacono 2002). Nymphoides indica isa floating perennial with tuberous and adventitious, spur-like roots; it can propagate through rhizomes. Within its native range, it occurs in shallow ponds and stream pools at elevations below 1500 m (Ornduff 1969; Sivarajan & Joseph 1993). Flowers are produced in cymose, umbel-like clusters, which occur on the petiole on the same node as the spur-like roots. Flowers have four to eight petals covered with copious marginal hairs produc- ing afurry appearance, hence the c name water snow! lake (Wood 1983). The flowers are fairly small and slender (1.7-2.7 cm wide), white with a center area of light yellow (Ornduff 1969; Jacono 2002). Anthers are often black and seeds range from 1.0-1.7 mm in length (Ornduff 1969). The springs and pool at Soldier Springs lie entirely within the flood plain of the Nueces River but during normal flows are separated from the main course by a point of land. The pool flows into the river at the downstream end of the point along the eastern edge of the channel. Spring flows emanate from gravel and cobble substrates within a short spring run at the head of the pool. The pool has depths in excess of 1.3 m (Bill Carr, op cit), is up to 30 m wide, and measures a few hundred meters in length. Brune (1981) classified the springs as moderately large, meaning they have a mean discharge between 0.79 and 7.93 m?/s. At the time of the Nymphoides indica collection, the spring run and pool were densely vegetated with various aquatic macrophyte species. Among the most prolific was Trichocoronis rivularis, a species with restricted global dis- tribution and found only in a few southwestern Texas and northern Mexico spring systems and cienegas (Bill Carr, op cit). Trichocoronis rivularis together with N. indica formed areas of very dense vegetation extending from the banks out into mid pool. Both species were distributed throughout the pool but were most dense from the mouth of the spring run downstream to about the middle of the pool. Other common macrophytes included Justicia americana, Ludwigia sp., Myriophyllum sp., Nasturtium officinale, and Potamogeton illinoensis. ACKNOWLEDGMENTS Thanks to Bill Carr, The Nature Conservancy of Texas, for providing background information, and to David Lemke, Texas State University, Biology Department for verification of specimen identification. Nicholas Tippery and Guy L. Nesom provided helpful review comments. REFERENCES Brune G. 1981.Springs of Texas Vol. 1. Branch-Smith, Inc., Fort Worth, Texas. Jacono, C, 2002. Florida's floating-hearts — know Nymphoides. Aquaphyte 22(1):insert. SAUNDERS, NYMPHOIDES INDICA IN TEXAS 2443 Ornburr, R. 1969. Neotropical Nymphoides (Menyanthaceae): Meso-American and West Indian species. Brittonia 21:346-352. SivaraJaAN, VV, and K.T. JosepH. 1993. The genus Nymphoides Séguier (Menyanthaceae) in India. Aquatic Bot.45:145-170. Woop, C.E. 1983. The genera of Menyanthaceae in the southeastern United States. J. Ar- nold Arbor. 64:431-445. WUNDERLIN, R.P. and B.F. Hansen. 2004. Atlas of Florida vascular plants [S.M.Landry and KN. Campbell (application development), Florida Center for Community Design and Re- search.]. Institute for Systematic Botany, University of South Florida, Tampa. 2444 BRIT.ORG/SIDA 21(4) Book NOTICES Miros_av M. GRANDTNER (compiler). 2005. Elsevier's Dictionary of Trees. Volume 1: North America with Names in Latin, English, French, Spanish and other Languages. (ISBN 0-444-51784-7, hbk.). Elsevier B.V. Radarweg 29, PO. Box 211, 1000 AE Amsterdam, THE NETHERLANDS. (Orders: wwwelseviercom). $199.00, 1489 pp., 61/2" x 91/2" The general compiler, Miroslav M. Grandtner, had numerous scientific collaborators to help with this incredible index. Each entry (genus and species) is numbered and the names bolded for easy eevee and sia anne of the name, family (with distribution), synonym(s), and common pd y noted). The list tallies up some 8,778 entries which includes genera. “Indexes” include DL ist of Synonyms of Latin Names; 2) English Names; 3) Noms Frangais; 4) Nombres Espanoles; 4) Other Names [languages]; and concludes with 5) Trade Names. Texas ASM University Press JOHN WATSON, CAROLE PETERSON, and DEANNA Payne. 2005. In Our Backyards: Public and Private Gardens of the Texas Coastal Bend. (ISBN 0-9766235-0-1, hbk.). Botanical and Nature Institute of South Texas, Inc. d.b.a. the Corpus Christi Botanical Gardens & Nature Center. (Orders: Texas ASM University Press, 4354 TAMU, College Station, TX 77843-4354, U.S.A. (979-458-3982, 979- 847-8752 fax). $39.95, 120 pp., color photos, 11" x 81/4". Publisher comments.—This wonderful new book showcases efforts of area gardeners to beautify our region, discusses env poner: issues peaciie ii - oastal coe and Hoks these topics by cui how g aan ane land issues which concern us all.’ . And to help showcase these gardens are beautiful and crisp color photographs. Timotuy Brusu. 2005. Nesting Birds of a Tropical Frontier: The Lower Rio Grande Valley of Texas. (ISBN 0-58544-490-1, pbk.). Texas AGM University Press, 4354 TAMU, College Station, TX 77843-4354, U.S.A. (Orders: 979-458-3982, 979-847-8752 fax). $24.95, 245 pp., color photos, 4 maps, 61/4" x 91/4" Contents.—Preface, Introduction, The Lower Rio Grande Valley of Texas, Ecological Diversity and his- tory, Habitats and Birds of the Valley, The Spectacular, Annual Cycle, Breeding Birds of the Valley, Species, Accounts and Summaries, Concluding Remarks, References, and Index. SIDA 21(4): 2444, 2005 MUSCARICOMOSUM (LILIACEAE) NEW TO TEXAS Lee M.Luckeydoo Botanical betes Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. LLuckeydoo@BRIT.org ABSTRACT M i (L.) Mill. (Lili cultivated persisting population in Parker County, Weatherford, Texas. This eee non-cultivated, is a new record for the state of Texas. RESUMEN Muscaric sum (L.) Mill. (Lili ) i una poblacid on persi sultivada en Parker Coasts We: athe ford, Texas. Esta especie, no cultivada, es una nueva cita para el estado de Texas. Muscari comosum (L.) Mill. (tassel grape-hyacinth, feather hyacinth) has not been reported for Texas by Correll and Johnson (1970), Hatch et al. 1990), Jones et al. (1997), Diggs et al. (1999), or Flora of North America (2002). This species has been reported for several other states in the U.S.: South Carolina (Haldeman 2001), Oregon, Missouri, Illinois, Ohio, Pennsylvania, Tennessee, Virginia, North Carolina, Georgia (Kartesz 1999), and Kentucky (Kartesz 1999; Jones 2005). A Texas collection of Muscari comosum is in the Tracy (TAES) Herbarium, Texas ASM University (Brazos Co. Ness s.n., 10 Apr 1899). The label information states that this was a “single specimen found on campus” (Kruse pers. comm. 2005). No further information was available as to this specimen having been cultivated or existing as a persisting population. The Texas collection reported here was submitted to the Botanical Research Institute of Texas by a homeowner in Weatherford, Texas (Parker County). The species had not been planted in the yard since the owner’s family moved into the home in 1948. The plants were first noted in the yard approximately “20 years ago.” This isa non-cultivated persisting population that is currently grow- ing in two groups. The homeowner has observed a gradual increase in the num- ber of plants of M. comosum over time. The plants thus appear to be spreading slowly. Muscari comosum (Fig. 1) isa perennial that regenerates from a bulb and is native to the Mediterranean region (Lopez Alonso & Pascual Reguera 1989). This species hasa distinctive flowering scape (raceme), which has fertile, dark purple, urn-shaped flowers that are spaced along the top half of the axis and a tassel of mene pure flowers at the tip of the axis. Synonyms for this plant include Hyacinth 1s comosus L. SIDA 21(4): 2445 — 2447, 2005 2446 BRIT.ORG/SIDA 21(4) Fic. 1. M ; Wl ion by Pi ] h Red 6 (1759 1840) (M | y R, Mall y 1986). J t LUCKEYDOO, MUSCARI COMOSUM NEW TO TEXAS 2447 Voucher specimen: TEXAS: r Co.: Weatherford, owner's family have lived in home since 1948, these plants were not seen a 1980/1990- occurring in 2 clumps (15 and 9 scapes); flowers ea purple with tassel at top of Salle fragrant, apeat SN. 15 Apr 2005. A photograph, dated 1992, of comosum in homeow ner Sy ACKNOWLEDGMENT Thanks to Barney Lipscomb of BRIT for providing assistance with identifica- tion, Monique Reed for her valuable comments, and Dale Kruse of the Tracy Herbarium for providing collection data. REFERENCES Corrett, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner. Diccs, G.M., B.L. Liescome, and R.J. O’Kennon. 1999. Shinners & Mahler's illustrated flora of North Central Texas. Sida, Bot. Misc. 16. Fora of NortH America Eoitorial Committee (eds.) 2002. Flora of North America north of Mexico. New York and Oxford. 26:317. Hatoeman, J.H. 2001. Noteworthy collection, South Carolina. Castanea 66:308. Hatcn, S.L., KN. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Bull. MP-1655. Jones, S.D., J.K. Wiere, and PM. Montcomery. 1997. Vascular plants of Texas: a comprehensive checklist including synonymy, bibliography, and index. Univ. of Texas Press, Austin. Jones, R.L.2005.Plant life of Kentucky: An illustrated guide to the vascular flora. Univ. Press of Kentucky, Lexington. Kartesz, J.T. 1999.A synonymized checklist and atlas with biological attributes for the vas- cular flora of the United States, Canada, and Greenland. First Edition.|In:Kartesz, J.T.and C.A. Meacham. Synthesis of the North America flora, version 1.0. North Carolina Bo- tanical Garden, Chapel Hill, NC. Kruse, D. 3 May 2005. Personal communication. Tracy Herbarium. Texas A&M University, College Station. Lopez Atonso, D. and L. Pascuat Recuera. 1989. Population structure and pattern of geo- graphic variation in Muscari comosum along its range of distribution. Genetica 78: 39-49. Mattary, P and F. Mattary. 1986. A Redouté treasury, 468 watercolors from Les Liliacees of Pierre-Joseph Redoute. Plate 153. Vendome Press, New York. 2448 BRIT.ORG/SIDA 21(4) ANNOUNCEMENT Digital Atlas of the Virginia Flora ls Online The Digital Atlas the Virginia Flora,a significantly augmented and somewhat revised version of edition III of the Atlas of the Virginia Flora (hardcopy 1992), is now available on the Virginia Tech website at http://www .biol.vt.edu/ digital xatlas. The Atlas project for the Virginia flora was initiated by Alton M. Harvill, Jr, of Longwood College, Farmville, Virginia, in the mid 1970s. With assistance from a group of other Virginia botanists who were also collecting extensively in the state, Harvill produced three hardcopy editions of the Atlas (1977/1981, 1986, and 1992). Following Dr. Harvill’s retire thi [which he had named and incorporated as the Virginia Botanical Associates (VBA)| continued his work by accumulating new records from their own and others’ more recent field work (e.g. records from the Virginia Division of Natural Heri- tage). The group also targeted numerous taxonomic issues needing resolution. Ultimately, the VBA set the goal of making available on-line a modified and updated version of Atlas II] and began developing a checklist. Thomas F Wieboldt compiled and maintained the developing checklist at the Massey Her- barium at Virginia Tech, and he also modified family circumscriptions to fol- low (largely) the Angiosperm Phylogeny Group (APG2). The initial database was populated with records from the 1992 hardcopy edition; then VBA mem- bers added new records to the database. The Digital Atlas of the Virginia Flora is very mucha work in progress. This is especially true in regard to mapping of infraspecific taxa that are currently “buried” in species maps because the specimens on which these maps were based had been identified only to species level. Users of the web-site are invited to correspond with the VBA regarding the Digital Atlasofthe Virginia Flora. Please direct questions, comments, and/or any new data to Thomas F. Wieboldt, Massey Herbarium, Virginia Tech, Blacksburg, VA 24061; phone (540) 231-5746; e-mail: wieboldt@vt.edu. SIDA 21(4): 2448. 2005 HYDROCOTYLE SIBTHORPIOIDES (APIACEAE) NEW FOR TEXAS AND NOTES ON INTRODUCED SPECIES Barney L. Lipscomb George M. Diggs, Jr. Botanical Research Institute of Texas Department of Biology 509 Pecan Street in College Fort Worth, Texas 76102, U.S.A. Sherman, Texas 75090, U.S.A. barney@brit.org Botanical ey Institute of Texas gdiggs@austincollege.edu ABSTRACT 1.,J asl ‘heh eae Ain 5 =| bP I 1] a ] pa fe : 1 i to the yee Flor a. Introduced pec in the East Texas flora, as g four pa rticularly problem- atic ne (Cuscuta japonica, pane ramosa, Solanum viarum, aaa Triadica sebifera) are discussed. RESUMEN Ty ] ] og Ms } ee ee | A ‘ \ la flora de’ oes Se discuten especies a iaeesewea: enla ‘lore del ae de Texas y uatro especies particularmente problemans as Ciseuta japonica, Orobanche ramosa, Solanum ana y Iriadica sebifera). Hydrocotyle is a cosmopolitan genus of approximately 130 species of creeping perennial herbs, including a number grown as ornamental ground covers (Mabberley 1997) or cultivated in water gardens or other aquatic habitats. Hydrocotyle sibthorpioides Lam., lawn marsh-pennywort or lawn water-pen- nywort, isa native of Asia (Mabberley 1987, 1997) but is widely cultivated. The species, usually described as a lawn weed, is naturalized in a number of locali- ties in the eastern United States (Arkansas, Florida, Delaware, Georgia, Indiana, Louisiana, Kentucky, Maryland, New Jersey, North Carolina, Ohio, Pennsylva- nia, South Carolina, Tennessee, Virginia, and West Virginia) and in California and Hawaii (Gleason & Cronquist 1963; Strausbaugh & Core 1978; Constance 1993; Kartesz 1999; USDA PLANTS Database 2005). It was not reported from Oklahoma (Taylor & Taylor 1994); however, it is reported from both Arkansas and Louisiana (USDA PLANTS Database 2005). Despite this proximity, the spe- cies has not been previously reported from Texas (Correll & Johnston 1970; Stan- ford 1976; Hatch et al. 1990; Jones et al. 1997; Diggs et al. 1999) A collection made in 2001 in Dallas is apparently the first documented occurrence of this species for Texas. Voucher specimen: TEXAS. Dallas Co.: spreading in landscape, 3511 Overbrook, Dallas, 5 Jun 2001, B. Lipscomb 3502 (BRIT). SIDA 21(4): 2449 — 2456. 2005 2450 BRIT.ORG/SIDA 21(4) Since originally observed at the collection locality, the species has persisted and spread in the lawn. Because this plant is low-growing and inconspicuous, par- ticularly when growing with taller grasses, we suspect that it is more wide- spread in Texas yards and has simply escaped notice. It has likely been inten- tionally planted into other localities as a ground cover or accidentally introduced with soil, compost, or cultivated plants. Hydrocotyle sibthorpioides can be recognized by the following description (modified from Gleason & Cronquist 1963, 1991; Radford et al. 1968; Strausbaugh & Core 1978; and Constance 1993). Delicate glabrous, perennial, terrestrial herb; stems creeping; leaves petiolate, the petioles 0.5-2+ cm long; leaf blades reni- form to suborbicular, shallowly 5-7-lobed, 3-12+ mm wide, minutely crenate; umbels simple, capitate, with 3-10 sessile flowers, peduncled, the peduncles 5- 15 mm long; flowers sessile, whitish; fruits round, 1-1.5 mm wide. Mar-Sep. Kartesz (1999) listed H. rotundifolia Roxb. as a nomenclatural synonym. An il- lustration (Fig. L), reprinted from Hiroe and Constance (1958), and a photograph (Fig. 2) are provided. The five species of Hydrocotyle known to occur in Texas can be distin- guished using the following key modified from those in Gleason and Cronquist (1963), Radford et al. (1968), Constance (1993), and Diggs et al. (1999): 1. Leaves peltate (= petiole attached to middle of lower surface of leaf blade). 2. Flowers in a simple umbel (= all flowers in inflorescence attached at the same point) H. umbellata L. 2. Flowers in a branched umbel or in on along an inflorescen is (definitely not all attached at the same poin 3. Flowers in a branched pai seiete more than 2 branches H. bonariensis Lam. 3. Flowers in whorls along an unbranched inflorescence axis, forming an inter- rupted spike or spike-like raceme or the axis with only 2 branches ____ H.. verticillata Thunb. . Leaves not peltate, the petiole attached at base of a notch. 2. Plants aquatic; stems and petioles fleshy; individual flowers and fruits with short H. but distinct pedicels ranunculoides Lf. 2. Plants terrestrial; stems and petioles thread-like; individual flowers and fruits essile H. sibthorpioides Lam. Hydrocotyle sibthorpioides is yet another introduced species (considered here as those originating outside the United States) added in recent years to the flora of Texas. Such non-native species are variously referred to as alien, exotic, or foreign. Their effects on the Texas flora have been commented upon a number of times in recent years (e.g, Diggs et al. 1999; O’Kennon et al. 1999: see Diggs et al. 2006, in press, for a detailed discussion), and there is growing concern in the state about their potential impacts. Recently (2003), a bill authorizing the Texas Department of Agriculture to publish a list of noxious plants was passed (Hibbs 2003; Texas Parks & Wildlife 2003), and that list is now available on-line (Texas Administrative Code 2005). However, Texas still has no single authority in G. Nakai, del. Fic. 1. Habit of Hyd Pr ae ee : Ws us 2, Constance 1958, fig. 4). gd ? f be charge of addressing invasive species issues, and detailed policies for effective and coordinated control efforts are still lacking. This deficiency could prove costly to the state in the future, both economically and ecologically. Fortunately, significant attention is now focused on the problem, and a major collaborative conference addressing the issue, the state-wide Texas Invasive Plant Conference (The Pulling Together Initiative), was held in November of this year (TexasInvasives.org 2005). Data recently obtained for volume one of the Illustrated Flora of East Texas (Diggs et al. 2006, in press) indicate that of the 3,402 total species known for East Texas, 619 species or 18% of the East Texas flora are introduced. Unfortu- nately, very little is known regarding the percentage of Texas’ land area cov- ered primarily by such exotics or the percent biomass represented by alien spe- cies in a variety of Texas habitats. Certainly vast areas of grazing land and roadsides are vegetated almost exclusively by exotics such as King Ranch 2452 BRIT.ORG/SIDA 21(4) F ~_ tf Fic. 2. Phot h af Uv, J “bah. rey Inhatn b BIL Linscomh) svt Pv }s bluestem (Bothriochloa ischaemum (L.) Keng var. songarica (Rupr. ex Fisch. & C.A. Mey.) Celerier & Harlan), Johnson grass (Sorghum halepense (L.) Pers.), Ber- muda grass (Cynodon dactylon (L.) Pers.), and tall fescue (Festuca arundinacea Schreb.). Casual observation and anecdotal evidence would suggest that many other habitats are seriously affected as well. A particularly troubling example is the eastern Asian Triadica sebifera (L.) Small [Sapium sebiferum (L.) Roxb], usually known as Chinese tallow tree or as popcorn tree, which has been widely used in landscaping in East Texas, in part because of its brilliant fall color. How- ever, this species is now widely recognized as one of the most serious invasive exotics in East Texas and in the adjacent Gulf Prairies and Marshes (e.g., Barrilleaux & Grace 2000; Keay et al. 2000; Loos 2002). It is particularly prob- lematic in invading and destroying native Coastal Prairie habitats and is show- ing a rapid increase in sapling populations in some floodplain forests of the Big Thicket National Preserve (Harcombe et al. 1998; Keay et al. 2000). In fact, in areas that had previously been primarily native forest, one can now find large numbers of young individuals of Triadica sebifera, sometimes being swarmed over by yet another invader, Japanese climbing fern (Lygodium japonicum (Thunb. ex Murray) Sw.). A table in the Illustrated Flora of East Texas (Diggs et al. 2006, in press) lists 41 species recently (since 1997) introduced into East Texas (and new to the state). This listing is almost certainly incomplete, in part because of species discovered 2453 since the book went to press. While many of these exotics, such as Hydrocotyle sibthorpioides, may be innocuous and present little danger of becoming invasive or otherwise of concern, a number pose serious conservation or economic threats. A weed with the potential to be economically devastating, Orobanche ramosa L. (hemp broom-rape or branched broom-rape), is now spreading in the west cen- tral part of East Texas and has been reported from at least 22 counties (Texas Cooperative Extension 2003). Recently (2004), it was discovered as far north as Dallas County (J. Quayle, pers. comm.). It is apparently being spread widely by highway mowing equipment. This chlorophyll-less native of southern and cen- tral Europe is a well known root parasite of agricultural crops, including bean, cabbage, celery, eggplant, pepper, potato, and sunflower and has the potential to have a significant economic impact in Texas. In heavily infested areas in other parts of the world, hemp broom-rape has been known to cause total crop failure (USDA undated). It is classified as a federal noxious weed (USDA Natural Re- sources Conservation Service 2002). Cuscuta japonica Choisy, Japanese dodder, is another federal noxious weed. Though currently reported in Texas only from Houston (Harris County), there is concern about its possible spread (Camilli 2002; Huber 2002). It is an aggressive parasitic vine which attacks a variety of woody plants and has the potential to have ser logical and ec if not eradicated. Solanum viarum Dunal, tropical soda apple, is yet another fed- eral noxious weed recently reported from Texas (Reed et al. 2004). This peren- nial, prickly shrub native to Brazil and Argentina is an alternate host to several viral diseases and pathogens that attack other members of the Solanaceae. It is thus a threat toa number of vegetable crops including tomatoes, potatoes, egg- plants, and peppers (Byrd et al. 2004). A final well known example is Salvinia molesta Mitchell, giant salvinia, a federal noxious weed that in recent years has been found in abundance at Toledo Bend Reservoir on the Texas-Louisiana bor- der Jacono 1999b) and at numerous other localities in the state. This South Ameri- can native (Forno & Harley; Forno 1983; US. Geologic Survey 2000) is a serious threat to aquatic habitats in Texas (Diggs et al. 2006, in press). Considered “one of the world’s worst weeds” Jacono 1999b), it has been introduced by humans to fresh waters of Africa, Asia, Australia, s Europe, New Zealand, North America, and the South Pacific and has resulted in severe economic and environmental problems (JJacono 1999a, 1999b; Garbari et al. 2000). The plants can grow rapidly, cover the surface of lakes and streams, and form floating mats that shade and crowd out native plants. Additionally, the mats (sometimes to a meter thick) re- duce oxygen content, degrade water quality, and can cause physical problems including hindering boats, clogging irrigation and drainage canals, and block- ing water intakes (Thomas & Room 1986; Jacono 1999a, 1999b, 1999c; Wood et al. 2001; Moran 2004). The four species just discussed are among the 29 species currently on the Texas Noxious Plant List (Texas Administrative Code 2005). Unfortunately, 2454 BRIT.ORG/SIDA 21(4) among the 619 introduced species documented for East Texas, there are numer- ous other detr imental e exotics that could certainly join them on the list. Further, without major on prevention, there will almost certainly be additional problematic introduced species added to the Texas flora in the years to come. ACKNOWLEDGMENTS We would like to thank Robert George for locating and scanning the illustration and for obtaining a number of references and Guy Nesom and Monique Reed for reviewing the manuscript. We also thank Lorine Gibson for her assistance REFERENCES BarriLteaux, T.C. and J.B. Grace. 2000. Growth and invasive potential of Sapium sebiferum (Euphorbiaceae) within the coastal prairie region: The effects of soil and moisture re- gime. Amer. J. Bot. 87:1099-1106. Byro, J.D, Jr., C.T. Bryson, and R.G. Westsrooks. 2004. Tropical soda apple (Solanum viarum Dunal): Identification and control. http://www.ceris.purdue.edu/napis/pests/tsa/ tsa2004-fs.pdf. Accessed Oct 2005. CAMILLI, K. 2002 oan Asian Dodder: A new invasive plant detected in Texas. http:// /other/texas.html. Accessed Oct. 2005. Corrett, D.S. and MC. eee 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner Constance, L. 1993. Apiaceae. In: J.C. Hickman, ed. The Jepson manual: Higher plants of California. Pp. 136-166. Univ. of California Press, Berkeley. Diccs, G.M. Jr, B.L. Liescome, and R.J. O’Kennon. 1999, Shinners & Mahler's illustrated flora of North Central Texas. Sida, Bot. Misc. 16. Diccs, G.M. Jr., B.L. Liescome, M.D. Reeo, and R.J. O’Kennon. 2006 (in press). Illustrated flora of East Texas, Vol. 1. Sida, Bot. Misc. 26. Forno, |W. 1983. Native distribution of the Salvinia auriculata complex and keys to spe- cies identification. Aquatic Bot. 17:71-83. Forno, |W. and K.L.S. Hartey. 1979. The occurrence of Salvinia molesta in Brazil. Aquatic Bot. 6:185-187 Gareari, F., A. Giovannini, and D. Marcuetti. 2000. Salvinia molesta D.S. Mitchell (Salviniaceae nuova per la flora d'Italia. Arch. Geobot. 6:73-78. Gieason, H.A. and A. Cronauist. 1963. Manual of the vascular plants of northeastern United States and adjacent Canada. Van Nostrand Reinhold Company, New York. Gteason, H.A.and A. Cronauist. 1991. Manual of the vascular plants of northeastern United States and adjacent Canada, 2nd ed.Van Nostrand Reinhold Company, New York. Harcn, S.L., K.N. Ganoul, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Misc. Publ. No. 1655. Harcomee, PA,, R.B.W. Hatt, J.S. Gurtzenstein, and D.R. Strenc. 1998. Sensitivity of Gulf Coast forests to climate change. In: G.R. Guntenspergen and B.A. Vairin, eds. Vulnerability of coastal wetlands in the southeastern United States: Climate change research results, 7 2455 1992-97. Biological Science Rep. USGS/BR/BSR-1998-002. U.S. Dept. of the Interior, U.S. Geol. Survey, National Wetlands Research Center, Lafayette, LA. Hisss, D.A. 2003. Title 4. Agriculture. http://www.sos.state.tx.us/texreg/archive/Octo- ber172003/PROPOSED/4.AGRICULTURE.html#18. Accessed Feb 2004. Hiroe, M.and L. Constance. 1958, Umbelliferae of Japan. Univ. California Publ. Bot. 30:1-144. Huser, K. 2002. Stranglehold: Giant dodder threatens urban landscape. Houston Chronicle 18 May 2002, page 8D. Jones, S.D., J.K. Wiper, and P.M. Montcomery. 1997.Vascular plants of Texas:A comprehensive checklist including synonymy, bibliography, and index. Univ. of Texas Press, Austin. Jacono, C. 1999a. Have you seen this plant? Texas Parks and Wildlife information sheet. Texas Parks and Wildlife, Austin. Jacono, C. 1999b. Salvinia molesta (Salviniaceae), new to Texas and Louisiana. Sida 18:927- Jacono, C. 1999c. Salvinia molesta (D.S. Mitchell) invades the United States! Aquatics 21:4-9. Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland, 1st ed. In: Kartesz, J.T.,and CA. Meacham. Synthesis of the North American Flora, CD format, Version 1.0. North Carolina Botanical Garden, Chapel Hill. Keay, J., WE. Rocers, R. Lankau, and E. Siemann. 2000. The role of allelopathy in the invasion of the Chinese tallow tree (Sapium sebiferum). Texas J. Sci.52(4) Suppl.:57-64. Loos, PM. 2002. An alien among us. Newsletter Native Plant Soc. Texas 20:6. Massertey, DJ. 1987.The plant book, a portable dictionary of the higher plants. Reprinted with corrections in 1989. Cambridge Univ. Press, Cambridge, England, U.K. Maseertey, DJ. 1997. The plant book, a portable dictionary of the higher plants, 2nd ed. Cambridge Univ. Press, Cambridge, England, U.K. Moran, R.C. 1992. The story of the molesting Salvinia. Fiddlehead Forum 19:26-28. O'KeNNON, R J., T.M. Barktey, G. M. Dicos, Jr, and B.L. Liescoms. 1999. Lapsana communis (Asteraceae) new for Texas and notes on invasive exotics. Sida 18:1277-1283. Quavte, J. Jerr). Personal communication, 2004. Plant collector, discoverer of the recently described S io quaylii (Asteraceae), naturalist,and botanical consultant of Fort Worth, TX. Raproro, A.E., H.E. AHtes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. The Univ. of North Carolina Press, Chapel Hill. Reeo, M.D., M. KetcHersio, and R.L.THompson. 2004. Solanum viarum (Solanaceae) —tropical soda-apple—confirmed from Texas. Sida 21:1171-1174. Stanroro, J.W.1976.Keys to the vascular plants of the Texas Edwards Plateau and adjacent areas. Published by the author, Brownwood, IX. STRAUSBAUGH, P.D. and E.L. Core. 1978. Flora of West Virginia, 2nd ed. Seneca Books, Inc., Grantsville, West Virginia. Tayior, R.J.and C.E.S. Taytor. 1994. An annotated list of the ferns, fern allies, gymnosperms and flowering plants of Oklahoma, 3rd ed. Southeastern Oklahoma State Univ., Durant. Texas Apministrative Cove. 2005. Quarantines and Noxious Plants, Chapter 19. State of Texas. 2456 BRIT.ORG/SIDA 21(4) http://info.sos.state.tx.us/pls/pubreadtac$ext.TacPage?sl=R&app=9&p_dir=&p_ rloc=&p_tloc=&p_ploc=&pg=1&p_tac=&ti=4&pt=1&ch=19&rl=300. Accessed Oct 2005. Texas Cooperative EXTENSION. 2003. Keep watch for branched broomrape (Orobanche ramosa) A major threat to U.S. crops. http://www-aes.tamu.edu/mary/brmrape/brmrape.htm 2002. Accessed Jan 2003. TexAsINvasives.orG. 2005. http://www.texasinvasives.org/. Accessed Oct 2005. Texas Parks AND WicouiFe. 2003c. Recent legislation affects Texas fish and wildlife. http:// www.tpwd.state.tx.us/news/news/030626a.phtml. Accessed Feb 2004 THomas, PA, and PM. Room. 1986. Taxonomy and control of Salvinia molesta, Nature 320: 581-584. USDA (U.S. Department of Agriculture). Undated. Keep watch for branched broomrape (Orobanche ramosa). http://www-aes.tamu.edu/mary/brmrape/brmrape.htm. Accessed Oct 2005. USDA Naturat Resources CONSERVATION Service. 2002. Federal noxious weeds. http:// plants.usda.gov/cgi_bin/topics.cgi?earl=noxious.cgi. Accessed Dec 2002. USDA PLANTS Datasase. 2005. PLANTS profile for Hydrocotyle sibthorpioides Lam. http:// plants.usda.gov/cgi_bin/topics.cgi?earl=plant_profile.cgi&symbol=HYSI. Accessed Oct 2005. U.S. GEoLoaic a 2000. Salvinia molesta, giant salvinia. http://nas.er.usgs.gov/ferns/. Accessed 2 Woop, M., B. rie and J.Garcia. 2001.Attack on giant salvinia. U.S.D.A. Agric. Res. 49:4-6; also on-line at: http:// usda.gov/is/AR/archive/nov01/giant1101.htm.Accessed Aug 2004. ANNOUNCEMENT 2457 ANNOUNCEMENT 2005 Delzie Demaree Travel Award Recipients The 17th Annual Delzie Demaree Travel Award was presented at the 52nd An- nual Systematics Symposium (7-11 Oct 2005) at the Missouri Botanical Gar- den. Three students were presented the Travel Award: Mario Blanco, University of Florida; Pedro Fiaschi, Virginia Commonwealth University; and Lina S. Juswara, Ohio State University. The 2005 Travel Awards were underwritten by 1) Delzie Demaree Travel Award Endowment, 2) Members of the Delzie Demaree Travel Award Commit- tee, and 3) John Clayton Chapter of the Virginia Native Plant Society. Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research Institute of Texas, to Barney Lipscomb, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 1-817- 332-7432; Email: barney@brit.org. Thank you. 2006 Applications for the Delzie Demaree Travel Award Applications for the 2006 Delzie Demaree Travel Award should include a letter from the applicant telling how symposium attendance will benefit his/her graduate work and letter of recommendation sent by the major professor. Please send letters of application to: Dr. Donna MLE. Ware, PO. Box 8795, Herbarium, Biology Department, The College of William and Mary, Williamsburg, VA 23185- 8795, U.S.A. 1-757-221-2799; Email: ddmware@wm.edu. The period for receiv- ing applications will end three weeks prior to the date of the symposium if a sufficient number of applications are in hand at that time. Anyone wishing to apply after that date should inquire whether applications are still being ac- cepted before applying. The Systematics Symposium dates for 2006 are 13-14 October. The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended 35 out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer, discoverer, and teacher. His teaching career as a botanist began in Arkansas at Hendrix College in 1922. He also taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas A&M and Arkansas State University at Jonesboro where in retired as professor emeritus in 1953. On of the things he enjoyed most as a botanist was assisting students with their field botany research. SIDA 21(4): 2457. 2005 2458 BRIT.ORG/SIDA 21(4) Book REVIEWS Mary Pratt. 2005. Practical Sci for Gardeners. (ISBN 88192-718-X, hbk.). Timber Press Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $24.95, 175 pp., b/w drawings, conversion chart, appendices, glossary, index, 6" x 9" In Practical Science for Gardeners, Mary Pratt gives the reader some of the science behind common and successful gardening practices. This book will be especially interesting for gardening enthusi- asts that may not have any botanical background. The book is written in a very conversational a with clever chapter titles such as “Nature arms race and how to negotiate a peace deal,” for the chap- ter discussing weeds, pests ane met Hoes of ans Ne areas pa science book can be foc of as horticulture “lite,” off why tices are effective and ee are not. Pratt also includes quite a bit of personal charm and alates some pocone pareeang Shiialie gies. basic cellular building blocks, plant anatomy, plant hormonal chet edsiey soil science, pest control, biodiversity, basic genetics, genetically modified plants, plant classification and pent species naming. After reading “ information “ue Les sia read: ers will und y important g do what ina plant, the benefits of different fentilizer eerieney ratios, why light and placement are so important to plants, what happens when plants are water stressed, processes that occur in a seed during germination, ways to determine plant nutrient deficiencies, the importance of biodiversity in the garden, what soil pH is, and how to correct improper soil pH, what are some aia aaa pest -_ a biocontrol strategies, and what does Benencany modified me Ip ] x ters are presented in a way to h apply the scientific information discussed to practical garden uses. Each chapter includes a anus of boxes that provide gardening — ns 1. These suggestions include some practical information for solving or preventing dorden problems For instance, antes nealing ae information oS on hormones and their functions in the plant g suge how to ripen tomatoes, and the effects of pruning plants. Pratt al I for further reading on topics presented in each chapter. Important words within each sha apter are bolded, and many of these bolded words are defined ina Ss eaae found at the end et the text, This book will be of particular important for British gar he inf sented in the chapter on biodiversity i is sees to many environments, the examples and sug. gested species given are specific for the UK. One theory of succession the development of woodland, grasslands and wetlands gardening habitats are Pree | A variety of gardening strategies for success and suggested species are included in the discussion on each of these gardening habitats. Readers outside of the UK be cautious in using the species suggested, what does well in one area, may be an aggressive invasive in another (Eurasian Milfoil, Myriophyllum a for example). Pre tical Science for d i eq Aeterna] 4 ] ,and can be a useful book for gar- ractices. Practical Science for Gardeners presents a ae overview of plant and Rorniculearals science behind gardening that shou courage readers to learn more of the science on topics of interest to them; the author has Gan lly included suggestions tO more advance rate by SHARC topic. The information is presented i InN a Way ! g think about why g or not be valid. / +} : aT ick ] 1 a | SIDA 21(4): 2458. 2005, BOOK REVIEWS 2459 Pick upa copy of Mary Pratt’s eae ann Gardeners; reading it will be like enjoying a con- versation over the fence with well-informed gardening neighbor—Lee Luckeydoo, Herbarium, Bo- tanical Research Institute of Texas, ee ee Street, Fort Worth, TX 76102-4060, U.S.A. BeN-ERIK VAN Wyk. 2005. Food Plants of the World. (ISBN 0-88192-743-0, hbk.). Timber Press Inc. 133 S.W. Second Ave, a 450, Portland, OR 97204-3527, US.A. (Orders: wwwtimberpress.com, berpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $39.95, 480 pp., many color photos, 6 1/2) x S1/2- Have you ever looked famili pinauce, beans, pies or Spices in he le and asked your- self—What is that, where does it at parts can you eat and what etal: is needed? Author Ben-Erik van a has ae hea answers to such questions in his book, Food Plants of the World. van Wyk, has constructed a food plant encyclopedia, listing information on 354 food plants and their close telat: complete wat iat ent photographs. Food Bae ca Wong would bea g he book lists primary crops by regions: ie. central Asia, Africa, etc. This is ee : major food categories, such as cereals, pulses, fruits, vegetables, herb, sugars, beverage plants, spices and flavors. Descriptions of the major food categories include information on general anatomy, basics on processing and cooking, and other interesting facts. Within each food category, are lists of the “main” plants by common names, followed by the scientific names. This list of pri- mary foods) in ae Aes is important as a abe plone to determine the scientific name of d f in the index. The author has included some intriguing color ane of foods within a category with common names in the captions, for instance for cereals, the photograph shows lines of different grains. Note that the author admittedly uses the labels of fruit and vegetable in the “grocery,” not in the botanical sense ulk of the text is the food plant “encyclopedia” in alphabetical order by specific epithet. Each food entry contains vibra color i amages of ene ee as wel as the edible portions. Following the food photographs I origin and | sumed, culageacea and harvesting, uses, ener Sale: assorted notes, common names in vari- ous languages and the family to which the species belongs. The descriptions are very thorough and the common names in a languages can be very helpful when shopping in different groceries or ac for foods and spic uthor has included handy f fter the food dia. The first is a section dis- cussing tl i ds. The chemiel structures of various sugars, es fats, pres vitamins and minerals are imeluded in the discussion. The descriptions of the aa t small portion of the sae of how the body Digeccnes such mo aa d ds, and often, the author has provideda of the various compounds are derived from. The second reference included is a quick a to Re ee food ae ais common name, cable Dorsons, country of ae and base nutri- tional values. The rema d index Food Plants othe or ¢ by Ben- Erik v van a Wyle is mehly recommenced for ee in toed sci- The text is thorough an sharp, and on color shotosraphs are fantastic in helping to Ste the plant in the field and the edible portions. This could be a wonderful reference book or gift for cooks, horticultural students or “foodies” in general. Definitely a book worth a look!—Lee Luckeydoo, Her- barium, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 21(4): 2459. 2005 2460 BRIT.ORG/SIDA 21(4) JAN Wrepr. 2005. Trees, Shrubs, and Vines of the Texas Hill Country. (ISBN |- 58544-426-X, pbk.). Louise Lindsey Merick Natural Environment series; no. 39. Texas AGM University Press, John H. Lindsey Bldg., Lewis St. 4354 TAMU, College Station, TX 77843-4354, U.S.A. (Orders: wlawrence@tamuedu, http://www.tamu.edu/upress, 800-826-8911). $23.00 hbk., 246 pp., color photos, index, 53/4" x 81/2". Each of 126 species is illustrated by at least one color photograph. A short is provided for each species with a brief commentary on aspects of its biology, pre isaac tics, and uses (for landscape ee or wi ue ors ie habitat). Within gymnosperms, dicots, and lly by family, ae and species, one species per page. The first one- if ‘th of the book ere nicely written sections on red cedar management, healthy streams, benefits of native species, landscaping with woody ae and invasive species. At the end hal ] ld are appendices with information on native plant nurseries and demonstration gardens and other resources for native plants. A multi-page chart gives “color or aroma,” “season,” “potential height,” “site preference,” and “wildlife uses” for 117 native species. The book surely will quickly be in the hands of many residents of central Texas and will be much used and valued. It potentially provides a aa service toward Hill Country conservation. r identification of woody plants, Wrede’s book is the successor of Daniel Lynch’s Native & een Woody Plants of Austin & the Hill Country (1981) and could have Supe ented it. Surpris- ingly, though, the newer poet oes not ee a eee set of Ns uy secs treated and : us- ] trated in the earlier one aaa nyllantoes, Berchemia Sean NS, Bernardia sree ieapolia, Buddleja racemosa, Castela texana,C ] , Ficuscarica, Fraxinus pennsylvanica, Galphimia hryallis) angustifolia, sin nigra, Ligusieum sinense, Mimosa biuncifera, Nicotiana glauca, Pavonia lasiopetala, ulus alba, Prunus caroliniana, Pauls mexicana, Quercus macrocarpa, uae sinuatd, nase eee ata, Sesbania drummondii, Symp! ‘pos orbiculatus, Thamnosma oy T]] tex eee further Hill Contes omissions, i dj ing | Amorpha roemeriana, Convolvulus arvensis, Condalia viridis, Croton texensis var. alabamensis, Da ionaie: Diospyros virginiana, Forestiera reticulata, Juglans major, Ligustrum lucidum, one rum quihoui, Matelea edwardsensis, Mimosa texana, oa nen pe ee enn Rubus bifrons, Salix exigua, Salvia ballotiflora. Al he most common Hill ountry species of prickly pear, the Texas state plant), O. imbric wee O. davisii, O. macrorhiza, and O. phaeacantha, Hawthorns are treated as “Crataegus sp.,” acceding to the myth that “The taxonomy of the Hill Country Crataegus is uncertain,” and continuing “and there may be several species that are very hard to separate, so all can be conveniently treated as one.” Crataegus crus-galli, C. greggiana, C. reverchonii, C. tracyi, C. turnerorum, C. uvaldensis, and C. viridis are Hill Cou ountry species sand are no more difficult to identify than the Prunus species. The smaller photo (p. 148) shows C. crus-galli, the larger C. tracyi. Finally, in view of the lengthy and useful discussion of cedar management (all pre- sumably assumed to be Juniperus ashei), at least a mention of J. pinchotii and J. virginiana would have been appropriate, although neither is abundant in the Hill Country.—Guy Nesom and Bob O’Kennon, Botanical Research Institute of Texas, Fort Worth, TX, 76012-4060, U.S.A. SIDA 21(4): 2460. 2005 BOOK REVIEWS 2461 Dariusz L. SZLACHETKO, PloTR RUTKOWSKI, and JOANNA eas es Contubuaons f th Quhtiwh Qn to the Taxonomic Revision 0 and Cyclopogoninae (Orchidaceae) in Mesoumeries aad the Antilles. (ISBN 83-89648-18-0, pbk; ISSN 0867-0730). Polish Botanical Studies 20. 1B Publisher, Polish Academy of Sciences, W. Szafer Institute of Botany, Lubicz 46, PL- 31-512 Krakow, POLAND. (Orders: ed-office@ib-pan.krakow.pl). Euro 90,00; ca. USD $105.46, 387 pp. 646 figures (including maps, types, line drawings, and color plates), 17 x 24 cm. This long- aia fe Gausemts 4 Szlachetko and his associates is a major cou poaon to the tax- my and of these three Spiranthoid subtribes. Treating 170 species in 35 a, it is ee illustrated with not on maps for each species but diagnostic morphological ee seas y- mee photographs of type specimens, and a sprinkling of color plates and is an excellent reference r those interested in this particular group of orchids. Covering Mexico, Central America, and th ae eee es are also found in saa United oe in once Soule tee Hens: southern Dn N + or Arizona genera a new combinations are used ten ae of cross-referenced synonyms are oe are two nese oe of ane volume. The lack of a general index, although the afore- ti lin g current treatments, requires going back and forth from the Table of Contents to the ee the es of genera and species is by subtribes and not alphabetical. Following what unfortunately appears to be a trend in publications on this region, when species are also found in the adjacent United States the information on them is either lacking or erroneous, primarily because it is based up work done more than 25 years ago. For some reason distributional work presented in several recent major North American publications was not consulted. Significant corrections that should be made concerning US distribution would include: Pages 14-17: Mesadenus polyanthus (Reichenbach f.) Schlecter listed for USA and is not found here, whereas M. lucayanus (Britton) Schlecter is not listed for the USA and is present in Florida. Pages 146-153: The treatment of Sacoil l rank or synonym is curiously miss- eee Szlachetko has treated in other peuliction: as S. lanceolata var. squamulosa. It is present throughout much of the range of the work as well as in central Florid a 209-210: ee aos ona a ree Burns-Balogh is ree eunoncoualy for USA and figure 351 is $ wn Lae aes by C. Luer in Arizona and known from an ees in Soutnweste Arizona and we Page 229: Funk Rutko oe be ee Deiregyne confusa ae, omits USA (Texas) from the range wee 1 g (Ames & C. Schweinfurth) Szlachetko on page 232 is listed for USA but there. Page 299: ee aus Svar) saison omits USA (Florida) from the range. Page 302: Cyclopogon omits USA (Florida) from the range. Page 313: Blend adnata (Swartz) Poiteau ex Richard omits USA (Florida) from the range. Supporting documentation for all of the above may be found in the appropriate generic treatments within the Orchidaceae, Flora of North America, volume 26.—Paul Martin Brown, Ocala, FL 34481 U.S.A, naorchid@aol.com SIDA 21(4): 2461. 2005 2462 BRIT.ORG/SIDA 21(4) Marcaret Mee. 2004. Margaret Mee’s Amazon: Diaries of an Artist Explorer. (ISBN 1-85149-454-5, hbk.). Antique Collectors’ Club in association with The Royal Botanic Gardens, Kew, Sandy Lane, Old Martlesham, Woodbridge, Suffolk, IP12 45D, UK. (Orders: wwwantiquecollectorsclub.com, 01394-389950, 01394- 389999 fax, email info@antique-acc.com). $59.50, 319 pp, water color paintings, O1s/2* <1 17/2". Some books are a slow read, this was one. Not ae it was is eee -hardly! Encounters with snakes, armed poachers, malaria, electric eels, and storms to say the least. What made the book a slow read was that the book is al t book, { filled with full-page botanical paintings of Margaret Mee, the Ge ae adine who journeyed to the Amazon to paint the rare and un- known flowers indigenous to the region. The reader is faced with the choice, at every turn of the page: do I read the wild story or do I gaze at the magnificent art? It is impossible to do both. The complex, emia: Worley peat oi the Amazon are executed in such exacting detail, you must pause and stare ere reading. Yet the adventure the artist recounts of hunting plants via dugout canoe, is so oripellings you dare not slow down to study the life-like paintings. Margaret Mee’s Amazon Diaries eae Artist oe. isa ne ke book bursting with big repro- kly ‘in asi ane pe eucons of ne anishes waterc ples | Lt places visited Sl | 1 Ls swam in the thee water e the river, fearful of the currents, and went eollectings ina occu canoe ith two young Indian boys. Gilberto climbed high in a tree, and from a rotting branch, which | feared would fall upon him, threw down a strange bromeliad which | had seen from afar the plant was not in flower, but I had no doubt that this was a new species.” Mee did not begin her travels deep into the fertile rainforests until the age of forty-seven. She continued until seventy-six. So focused was she on her mission to document the flora of the vanish- ing Amazon, that she would often return to a distant outpost, where a rare plant had been spotted i months earlier, just for the opportunity to record it in Hees Ever focused on her mission to paint rare and u species, she brushed off the misery of mosquitoes, hunger, humidity and drenching rains. The artist writes in her diary of finding Acacallis cyanea—the blue orchid. “I walked until I was exhausted, wading through streams and then, with soaking canvas shoes, ploughed through black, Swamy ground. But | was delighted with the results of my journey, for I had materia for many ed argaret Mee wi p land painted, she al inted | in the bottom of a rocking canoe, and wh hi I de either method of panuine diffi- cult, plants were piled into the bottom of ae fa and painted l ater. She cer the most complex of plants witha deft han plai he of-factly, “the flowers fell in golden showers as | ae ie cendiile I painted the ane seated in the boat as we moved upstream, for vine flowers are delicate and ephemeral.” Her work combines the rare ability to commit to paper, oe strictest ' botanical — ofa pene i] ] while at the same time, imbuing it with artistr piel of the Moont lower, which blooms briefly and only at night. She set up an wall night vigil to nin { the night bloomer. As it unfurled she painting furiously, by torchlight..until es at which: time the eat {lower withered and was no more. The resulting body of work y known images of the nocturnal beauty. Place das book at eit ear Read the wild adventure of this tireless explorer or gaze at the magnificent paintings of a brilliant botanical artist ..or do both. You will be lured back to it nightly, like a moth to a Moonf lower. SIDA 21(4): 2462, 2005 BOOK REVIEWS 2463 Cynthia Padilla ye ee ey ‘a } + { hat 4 7 rs J 1: oe * . Ee Pan [o} ts reviews i I pl trait siemens d lleaderon sketching s rT ¢ B | | +1 ‘ + Os http: Vina ee oomes com/ Join other | F fod FE s ft botanicalar t Davip R. Foster and Joun D. Aser. 2004. Forests in Time: The Environmental Consequences of 1,000 years of change in New England. (ISBN 0-300-09235- O, hbk.). Yale University Press, 302 Temple Street, PO. Box 209040, New Haven CT 06520-9040, U.S.A. (Orders: www.yale.edu/yup, 203-432-0960, 203- 432-0948 fax). $45.00, 447 pp., b/w figures, tables, index, bibliography, contributors, bibliographic essay, 6" x 91/4" The book, Forests in Time, is composed of essays by multiple authors that discusses the history of a New England forest. Harvard University acquired nearly 3,000 acres to establish the Harvard Forest, a study site located i in Petes, Massacausetts as an area to conduct ecological Seige The au- thors stated that 1 lissues 1 the his- tory of a ern area. This book was easy to read, and provided graphs and tables to help the reader understand the ecolonica ee of a forest, although some of these graphs were harder to nterpret than others. Th ailed pictures throughout the book helps the reader de- ee a better perspective of how this a changed through time. The book is divided into five main sections: background to ecological studies, regional be modern forest landscape, under- sae lores Sosy os, and pesone Paes oon ie tudies, conducted in the f ane fa reasons for Long pom Ecological Research (ITER). The text helps readers understand a response to environmental, anthropogenic, and a factors. The landscane ae are studied at four spatial scales: site, landscape, sub-region, and re Each of these spatial scales are oa by hydrology, humans, elias and Ss oer of the cosystems. B g-term studies must be conducted in order to understand any changes that occur. The second section describes oe changes of the forest. The Harare pOrese landscape has seen a variety of cl i eing a tundra, boreal forest point in its life. These mee occurred cheotigl natural (wind, pests, and fire), as well as anthropo- genic enh American) disturb Ff, . aoe | et peda section expel i nen al land both oe) and indirectly. The influence of several historical a ] ¢. 4 1 RA dp ] g rrospec lt Hill including pri lter the soi 1 f the soil. The an used several graphs to support the change i in abundance of gpecific plants at each site a the effects of land-use history. These graphs showed a strong correlation between the carbon to nitrogen ratio and nitrification in ones soils an eau” ee -use ee Studies on the ie term eis uence pon the et have a more ecousidereble inf a than urban areas or dof smaller vegetation. All of es enanee ae worked i in creatine eine piesene forest ecosystem. | h long- Oo Oo term Lae d d ] I i lesi 1 tosimulate | i ffect on forest ecosystems, ehe process of nitrogen satucation soil ene and litter and root influences on soil This research al for long term study of these various effects. There were several controls SIDA 21(4): 2463. 2005 2464 BRIT.ORG/SIDA 21(4) used throughout the experiments: nutrient fluctuation, “ecophysiological performance, population ynamics, vegetation structure, and ecosystem productivity.” The fifth section and conclusion reviewed lessons learned from research done in the forest. This research is leading to a better understanding of forest systems and ideas to improve land con- servation. a land and forest are constantly changing, and people that manage this land need to — “find t landscape change into long term planning”. This constant c aange pias that in sarctal systems, such as ane forest, long term studies must be conducted because they help to develop t Detter conser bj aL This book was very interesting, aa will help readers understand the importance of long term ecological research. This book was written in a way that it is easy for those new to science and ecology to understand. It is recommended to persons interested in New England terrestrial ecology, distur- bance effects on forest structure, and long-term research locations.—Keri McNew, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. Doua_as E. Sortis, PAMELA S. Soitis, PETER K. ENpRESS, and MARK W. Cuase. 2005. Phylogeny and Evolution of Angiosperms. (ISBN 0-87893-817-6, pbk.) Sinauer Associates, 23 Plumtree Rd, Sunderland, MA 01375-0407, U.S.A., (Orders: 413-549-1118 fax; orders@sinauer.com; wwwsinauer.com). $59.95, 370 pp., numerous b&w higures, G 1/2" 11", Rare is the book that has you cursing its basi ots and thesis, yet draws you to explore it often. This is such a book. In many ways, it is ed, a classic. So you might as well buy a copy and place it within easy reach next to Cronquist’s Integrated System. In essence, Phylogeny and Evolution aah a satis is the magnum opus of the Angiosperm Phylogeny Group (APG). Although the authors’ style is somewhat detached, as if to explain the ac- tions of the APG asa third or. they are nee movers and shakers of the APG. They know its workings and ora from the g pond up. Although leaders in the field of angiosperm molecu- — ar sees s, they are all classically trained, mostly in the 1970s and are well versed in plant mor- pholo ¥ This fac ctis ee in the text, and that is the reason for my oxymoronic relationship with this boo Gasman of the chapters is such that t Sad rey fall into three natural sections. Chapters | and 2 provide general background and concepts. Chapters 3 to 9 are detailed accounts of molecular sup- portand ee aoaae nan ee of the major ede of e sae classification (sequentially, the b ocots, basal eudicots car yophyllids rosids. and asterids. C naples 10-13 examine the application ai ee APG classification t t t [ of evolutionary diversification he angiosperms. Why I love this book: It is a readable and elaborated explanation of the current APG classification. Besides incorporat- ing new publications to update APGII, the Se present new analyses to answer questions raised during tl { the book its Not only do the authors a the APG ae a into ‘umes but they also characterize the mor- phology of the taxa (especially orders) recognized in the APG classification. In particular, they report any synapomor phies that corroborate the molecularly defined clades. This is es- pecially helpful in cases with morphologically divergent taxa that are united on molecular rounds. The authors are to be commended for a writing style that keeps this material from becoming tedious. SIDA 21(4): 2464, 2005 BOOK REVIEWS 2465 The book is a gold mine of data and references, citing both recent molecular and morphological studies. Combined with Cronquist’s Integrated System, it provides a commanding window into the comparative literature on angiosperms It summarizes many new insights into genomics, morphogenetics, and the regulation of gene t expression. In short, it is a great resource for keeping up-to- Why | hate loving Phylogeny and Evolution of Angiosperms: I thoroughly disagree with the philosophical basis of the authors’ conclusions. This philosophy is: “DNA sequence may is ne ace that trumps all.” The last section of each character- ized taxonomic g ted to a discussion of “Character ila ora Likewise, the last four chapters Seams general trends across angiosperms. In these discussions, every mor- phological pve that is discussed is evaluated for omdod By its estabutien on a the molecular cladogram—no exceptions. Am | on this latest bandwagon? Is one to ana that a a of three genes (two of which are plastid metabolic genes) adequately sample the genome of angiosperms? Even for the subclades based on as many as seven genes, is that an adequate sample? Given how little is known about gene regulation, expression of phenotype, and interaction of the genome with } é sal } + J} } } 11) 4 bes 7 ] 4 c L jo) oo Jc oO premature, at best, and potentially disruptive and destabilizing to systematics. Indeed, the DNA is information that contains messages in much the same way that language composed of symbols does. Is not the basing of our classification on the comparisons of nucleotide matches much the same as saying we have translated a message by simply analyzing the occurrence frequency of letters in the message? } L ld So lassically trained botanist do with this book? Obtain a copy; use it to keep up-to- date with this bandwagon and perhaps launch your own research Biobee oe to it often as you read molecular or other systematic papers. Use it to memorize the APG n to be conver- sant with molecular systematists. magine that others may hate to love this book for other reasons. What if you are a mo- lecular oo with limited backerounes in mompneleeys Obtain a COPY, smugly aeteene wi ue APG ae Then, force yourself so skillfull ould it be that the authors, too, will come to hates to lave hee own book? On page 19 they say, “A new eee i is needed to promote further progress in seed a relationships in general and angiosperm origins in particular. Given the direction that the molecular and genetic data are taking us, this new pateien ied be es more ewe pan even mene author suppose. Why will ?—Roger W. Sanders, Research Associate, Ravana Research facture of Texas, 509 Pecan ees Fort Worth, TX 76102-4060, U.S.A., rsanders@britorg. SIDA 21(4): 2465. 2005 BIENNIAL REPORT AND UppaTE 2004-2005 SIDA, CONTRIBUTIONS TO BOTANY Source of current research in classical and modern systematic botany for readers throughout the world. ...bringing out the best in botanical science for plant conservation and education ae all authors, we thank you for choosing Sida, Contributions to Botany as your forum to disseminate information and knowledge gained by scien- tific inquiry. Volume 21 is the largest volume of Sipa ever published. Sipa is pub- lished twice a year. Volume 21 has 2511 pages, 290 authors, 224 published con- tributions, and includes 237 new names and new combinations. Published papers are available online in Adobe Acrobat format (PDF files); the PDF files are true representations of the hard copy of Sipa . A word of thanks to the 298 individuals acknowledged on pages 2469- 2470, who generously supported Sipa ee their time and expertise in re- viewing 227 manuscripts submitted for volume 21. All manuscript sul are peer-reviewed by distinguished reviewers, your support is deeply appreci- ated. If by chance you reviewed a manuscript and your name was left out, the error rests solely on the shoulders of this editor and I truly apologize. Please bring any omissions to my attention. Below are the dates of publication for each of the four issues of volume 21, subscriptions for calendar year 2006, and distribution of Sipa. We also bring to your attention (see page 2468) the Mary M. Hennen Scientific Publications En- dowment, established to provide support to the future of BRIT’s scientific publi- cation program (Sida, Contributions to Botany and Sida, Botanical Miscellany). The index to volume 21 (2004-2005 follows and the issue concludes with a printed copy of the current guidelines to contributors. The online version is at http://www.brit.org/sida/SubmitPaper.htm. We thank all authors, reviewers, subscribers (individuals, institutions, or- ganizations), and readers for your continued interest and support. It is our plan to continue bringing you the best sources of current research in classical and modern systematics with your continued support. Wishing you the best in 2006 and beyond.—Barney Lipscomb (BRIT), Editor; John W. Thieret (NKU), Associ- ate Editor; and Félix Llamas (LEB), Contributing Spanish Editor. Dates of publication 21(1), pp. 1-510: 20 August 2004 21(2), pp. 511-1248: 28 December 2004 21@), pp. 1249-1954: 5 August 2005 21(4), pp. 1955-2511: 21 December 2005 SIDA 21(4): 2467. 2005 2468 BRIT.ORG/SIDA 21(4) Subscriptions and distribution.—Sipa is distributed in over 90 countries. There are 860 subscriptions, 345 domestic and 515 foreign. Brazil has the most sub- scribers (38) outside of the U.S.A. Sina is available through paid subscription or scholarly exchange. Online information is available at http://www britorg/sida/ subscriptions.htm. Annual subscription rates for Sida, Contributions to Botany for 2006. Domestic (USA only) |) Subscribing institutions (universities, libraries, and other institutes including agents) $75.00 2) Individual personal subscriptions $39.00 Outside the USA 1) Subscribing institutions (universities, libraries, and other institutes including agents) $85.00 2) Individual personal subscriptions $39.00 Mary M. Hennen Scientific Publications Endowment Established.—In honor of his wife, Mary, BRIT research associate, professor emeritus of botany at Purdue University, the world’s pre-eminent authority on the systematics and biogeog- raphy of the rust fungi, Joe Hennen has established the Mary M. Hennen Scien- tific Publications Endowment to provide support to the future of BRIT’s scien- tific publication program (Sida, Contributions to Botany and Sida, Botanical Miscellany). Joe and Mary (life-science librarian at Purdue University) have long recognized that publication of scientific botanical information is an integral part of conserving our natural heritage. Every gift to the endowment will make a difference for the future of Sida and Sida, Bot. Misc. There are many different ways to make a gift; use a credit card, give a check, stock, pledge, bequest, trust, memorials, employer/employee matching contributions, installments, etc. The BRIT Press seeks your support to ensure innovation and excellence in preparation, manufacture, and distri- bution of botanical research and scientific discoveries for the twenty-first cen- tury. For more information about the BRIT Press or anyone interested in making a contribution to the Mary M. Hennen Scientific Publications Endowment— which supports Sida and Sida, Bot. Misc.—contact Barney Lipscomb at the Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 76102-4060, US.A.; 817-332-7432; barney@brit.org: sida@brit.org. SIDA 21(4): 2468. 2005 REVIEWERS 2469 REVIEWERS: VOLUME 21 (2004-2005) 298 reviewers: several individuals reviewed more than one manuscript Thank you for supporting Sida, Contributions to Botany. Acedo, Carmen Bailey, Jr., Claude J. Baird, Baldwin, Bruce G. Ball, Peter W. Ballard, ia Harvey E. Ballenger, Julie Barber, Susan C. Barkworth, Mary E. Barlow-lrick, Patricia Barrie, Fred R Baskin, Carol C. Baskin, Jerry M. Baum, Bernard R. Brunell, Mark S. Buck, William R. Burnham, Robyn Burns, Jean Campbell, Julian Cardellina, John Carlquist, Sherwin SIDA 21(4): 2469. 2005 Carlsward, Barbara S. Carranza, Eleazar Carstens, Kenneth C. ies Richard mbers, Kenton L. a appa, C.C. Christensen, Knud Ib Church, cons A Clark, Lynn Columbus, i — Consaul, Laurie L. Cornejo iets Cragg, Gordon Craven, Lyn A. Cruden, Robert W. Das, AP. Davidse, Gerrit Davila, Patricia D. Delprete, Piero G. Denslow, Michael W. Dorr, L Dowlin RM. Duvall, Tae Eggli, Urs Elisens, Wayne J. Endress, Peter K. Essig, Frederick B. Evans, Dan K Ezcurra, Cecilia Fantz, Paul R. Farmer, Susan B Ferguson, Carolyn J. Fernandez-Concha, rman Carnevali Fishbein, Mark Ford, Bruce A. Freeman, Craig C. Freire, Susana Edith Fryxell, Paul A. Galloway, ee Gandhi, Kanchi N Garland, vm A. George, Robert - man ae ane ah Gillespie, Lynn J. oldblatt, Peter Gomez-Sanchez, Maricela Gonzalez-Elizondo, M. Socorro Graff, Pamela Grant, Verne Hall, David W. Hartman, Ronald L. oe Stephan L. Haynes, Robert R. Hennen, Joe Henrickson, James Herrera Arrieta, Yolanda Hess, Bill Hill, Steven R. aoe Cc Holst, Bruce K. Horne, Francis R. Howcroft, N.H.S. Humber, Richard A. Hyatt, Philip Janovec, John Jarvis, Charlie Johnson, George P. Jones, Thomas H. Judd, Walter S. Judziewicz, Emmet]. Kaul, Robert B. Kawasaki, Lucia Keeley, ee iC. Keener, Carl S Kennedy, Robin C. Kephart, Susan R. Khan, Zaheer Kirkbride, Jr, Joseph H. Knapp, Sé Kral, Robert Krapovickas, Antonio Kumar, Muktesh Lammers, Thomas G. Lamont, Fric Lance, Ron Landrum, Leslie R. Lee, Linda Lemke, David E. wis, David Livshultz, Tatyana amas, Félix Lohmann, Lucia G. Lonard, Robert Lowry II, Porter P. MacDougal, John acklin, James A. Magill, Robert E. Magrath, Lawrence kK. Malmquist, Ann Marcondes-Ferreira, Washington Martin, David L Martinez Diaz, Mahinda 2470 McDonald, J.A. McDowell, Tim McKenzie, Paul McMullen, Conley K. McNeill, John Mejia, Teresa Mellicham Lawr neneas a Mill, Robert Molpus, Jane Moore, Gerry Morefield, James D. Moreno, Nancy P. Mueller, Jochen Murray, David F Murrell, Zack Naczi, Robert FC. Nasir, Syed Mahmood Nee, Michael Neill, Amanda Nelson, Allan D. Nelson, John Nepokroeff, Molly Nesom, Guy Nevling, Lorin I. Nicolson, Dan H. Nordman, Carl Norris, Dan Noyes, Richard D. Oberlies, Nicholas H. Olmstead, Richard G. Orzell, Steve L. SIDA 21(4): 2470. 2005 Palmer, Michael W. Paratley, Robert Peck, James H. Peet, Robert Pennington, T.D. Pérez Garcia, Blanca Perkins, Ken Peterson, ae M. Pfister, Donald H. Phillips, Sylvia Plunkett, Gregory M. Poll, Elfriede Poole, Jackie Porter-Utley, Kristen E. Powell, A. Michael Pyne, Milo Rabeler, Richard Reed, Monique D. Rettig, Jeff H Revesl. Ties L. Roberts, Roland P. Robertson, Kenneth R. Rock, Janet H. Rohrer, Joseph R. Rossman, Amy Rumely, John H. Rutishauser, R. Ryburn, Adam K. Saar, Dayle E. Salazar, Jackeline Sanchez del Pino, Ivonne Sanchez, Carlos Sanders, Roger Schilling, eas E; Schultz, Joann Schuyler, Alfred — Semple, John C Shaw, Joey Sheviak, Charles J. Shulkina, Tatyana Siddiqi, Akmal Simpson, Beryl B. Singhurst, Jason Skinner, Mark Smith, Don Smith, Latimore Smith, S. Galen Soltis, Douglas E. Stewart, Scott Stoynoff, Nick A. rother, John L. Stuckey, Ronald L. Su ic Sutter, Robert D. Syring, John Taylor. Charlotte M. Taylor, David D. Tejero, Daniel BRIT.ORG/SIDA 21(4) Telfair Il, Ray C. Thiede, Joachim Thieret, John W. ) Dale Tippery, Nicholas Tucker, Gary Tye, Alan Umber, Ray Van Horn, Gene S. Vanderpool, Staria S. Wagner, Warren L. Wallnofer, Bruno , Grady Weigend, Maximilian Wilken, Dieter H. Williams, Justin Williams, Norris H. Wilson, Barbara L. Wunderlin, Richard P. Yang, LY. Aleck Yatskievych, George A. Yin-Long Qiu Yokota, Masatsugu Zamudio, Sergio Zarucchi, James L. Zmarzty, S Zona, Scott A. INDEX 2471 INDEX TO 224 TITLES WITH 293 AuTHoRs: VOLUME 21 (2004-2005) We know you have a choice. Thank you for choosing and supporting Sida, Con- tributions to Botany. A case of disputed orthography: is it Echinochloa colona; or is_ it Echinochloa colonum (Gramineae)? by Danie. B. Warp —2.1(4):2171 A floristic survey of Fort Matanzas National Monument, St. Johns County, Florida by Wenpy B. ZomLEFER, David E. GIANNASI, WALTER S. Jupp, Lisa M. Kruse, and Ketty A. BetTinGeR—21(2):1081 A lectotype for Stachys floridana (Lamiaceae) by JOHN B. Nevson and DanieL B.Warp—21(4):2119 A multivariate morphometric study of the aster genus Sericocarpus (Asteraceae: Astereae) by MicHette R. LEONARD, RACHEL E. Cook, and JOHN C. SempLe—21(3):1471 A new Citharexylum (Verbenaceae) from Isla Socorro, Revillagigedo Archipelago, Mexico by José Luis Leon be LA Luz and FERNANDO CHIANG —21(2):547 A new combination in Antennaria (Asteraceae: Gnaphalieae) from North America by RANDALL JAMES BayeR— 21(2):767 A new combination in Dendrocalamus (Poaceae: Bambusoideae) by MuktesH Kumar, RemesH M., and N. UNNiKRISHNAN— 21(1):93 A new combination in Matelea (Apocynaceae: Asclepiadoideae) for an endemic Jamaican vine by ALEXANDER KrinGs—21(3):1515 A new combination in Persicaria (Polygonaceae) by Craig C. FReEMAN— 21(1):291 A new combination in Stenotus SIDA 21(4): 2471. 2005 (Asteraceae) by Cates A. Morse— 21(4):2093 A new combination in the genus Packera (Asteraceae: Senecioneae) by Desra K. Trock—21(3):1643 A new name and a new combination in Solidago subsect. Glomeruliflorae (Asteraceae: Astereae) by RacHet E. Cook and JoHn C. Semple —21(1):221 A new record of the devil's cigar, Chorioactis geaster (Pezizales: Ascomycota), from Collin County, Texas by JOHN E. UBeLAKER and JANE K. STARKS— 21(3):1939 A new. species of Calycadenia (Asteraceae) from north central Califor- nia by Rosert L. Carr and Geraco D.CaRR— 21(1):259 A new. species of Centaurium (Gentianaceae) from Trans-Pecos Texas by B.L. TurNeR—21(1):87 A new species of Erigeron (Asteraceae: Astereae) from northwestern California by Guy L. Nesom and THomas W. Netson— 21(2):673 A new species of Leucophyllum (Scrophulariaceae) and comments on relationships of the genus by James HENRICKSON—2.1(1):1 A new species of Matelea (Apocynaceae: Asclepiadoideae) from Hispaniola by ALEXANDER Krincs—21(3):1519 A new species of Ptilimnium (Apiaceae) from the Atlantic coast by ALAN S. WEAKLEY and Guy L. Nessom—2.1(2):743 A new species of Sabatia (Gentianaceae) 2472 from Saline County, Arkansas by James S. PRINGLE aNd THEO WitseLL—2 1(3):1 249 A new species of Scutellaria (Lamiaceae from Guerrero, Mexico by Bitue L. Turner and James L. Revea.—21(2):679 A new species of Symphyotrichum (Asteraceae: Astereae) from a serpentine barren in western North Carolina by Gary L. KAUFFMAN, Guy L. Nesom, ALAN S. WEAKLEY, THOMAS E. Govus, and Laura M. CoTTERMAN— 21(2):827 A preliminary analysis of Clematis (Ranunculaceae) in sub-Saharan Africa by Freperick B. Essic—21(2):861 putative hybrid of Schoenoplectus saximontanus and S. hallii (Cyper- aceae) from Oklahoma by Marian Smith, Paut McKenzie, Paice Mettter-CHerry, and GALen SMiTH—21(1):475 A quantitative study of the vegetation sur- rounding yellow lady-slipper orchid (Cypripedium kentuckiense, Orchi- daceae) populations at Fort Polk in west central Louisiana by CHartes M. ALLEN, SARA THAMES, SPENCER TRICHELL, aNd JEREMY WHITE — 21(1):409 review of Crataegus series Rotundifoliae (Rosaceae) in western Canada by J.B. PHipps and R.J. O’Kennon— 21(1):65 revision of Agoseris apargioides (Asteraceae: Lactuceae) by Gary |. Bairro — 21(2):7 15 revision of the Psidium grandifolium complex (Myrtaceae) by Leste R. LANDRUM—2.1(3):1335 revision of eset ane Graphe- phorum (Poaceae: Pooideae: Aveninae in North America north of México by Victor L. Finot, Paut M. Peterson, Ropert J. Ss > > > > > <7 SIDA 21(4): 2472. 2005 BRIT.ORG/SIDA 21(4) SORENG, and FERNANDO QO. ZULOAGA— 21(3):1419 A second population of Agalinis navasotensis (Scrophulariaceae) con- firmed from Tyler County, Texas by Mo- Nique Dusruce Reed, JOHN F. Hays, JuorrH M. CANNE-HILLIKER, DANA Price, JASON SINGHURST, and Eric KeitH—21(3):1927 A synopsis of the genus Otostylis (Orchidaceae: Maxillarieae subtribe Zygopetalinae) with a new record from southern Peru by Micuet CHocce, JOHN P. JaAnovec, and Eric A. CHRISTENSON— 21(2):841 A taxonomic review of the Eriophorum russeolum—E. scheuchzeri complex (Cyperaceae) in North America by Jacques CayoueTtE—21(2):791 A two new species of Calyptranthes (Myrtaceae) from Ecuador by Maria LucIA KAWASAKI and Bruce K. Horst—21(4):1955 A vascular flora survey of emergent creek bed microhabitats of Kisatchie bayou tributaries in Natchitoches Parish, Loui- siana by Ray NeyLano, Metissa HeNNIGAN, and Lowett E. UrgatscH—21(2):1141 About 450 miles up the Red River in a large prairie: Peter Custis’ surviving botanical information from the Red River Expedi- tion of 1806 by MicHact H. MacRoserts and Barpara R. MacRoserts—21(2):963 Acerca de la identidad de Baccharis subsculpta (Asteraceae: Astereae) by ARTURO GRANDA Paucar y DANiet A, GIULIANO—21(2):705 Additions and emendations to The Wild Orchids of North America, North of Mexico by Paut Martin BRowN—21(4):2297 Additions to and noteworthy records for the vascular flora of West Virginia by Cyn- INDEX THIA M. Morton, JOHN Kartesz, BONNIE ISAAC, and Rosert Coxe—21(1):481 Allotoonia, a new neotropical genus of Apocynaceae based on a subgeneric segregate of Echites by J. Francisco Mo- RALES and Justin K. WittiaMS—21(1):133 Allozyme diversity in Eleutherococcus senticosus and E. brachypus (Araliaceae) from China and its implica- tion for conservation by SHI-LIANG ZHOU, Jun Wen, and De-Yuan Honc—21(2):993 Alternanthera paronichyoides (A t| and Rumex maritimus (Polygonaceae) new to Okla- homa by Bruce W. Hoactand and Amy BuTHOD—21(2):1199 Amaranthus acanthobracteatus (Amaranthaceae) by James HENRICKSON— 21(1):11 An annotated, preliminary checklist of the vascular flora of Camp Butner, North Carolina by ALEXANDER Krincs and Carlyle FRANKLIN—21(2):1131 An evaluation of Anthenantia (Poaceae) by Rogert KrAL—21(1):293 Annotated checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin coun- ties, Texas by Larry E. Brown, Barsara R. MacRoserts, MicHact H.MacRoeserts, Paut A. Harcomee, Warren W. Pruess, |. SANDRA ELSIK, and DAN JoHNSON—2 1(3):1807 Aristidae eludendae: Aristida hitch- cockiana (Poaceae)—a valid species? by Keuty W. ALLRED —21(4):2113 Asteraceae from wool mill sites in South Carolina, including new records for North America by Guy L. Nesom— 21(2):1215 Aulonemia nitida eae) (Poaceae: Bam- SIDA 21(4): 2473. 2005 2473 busoideae: Bambuseae), a new species from Guyana by Emmet J. Juoziewicz— 21(3):1263 Broadened concept of Liatris helleri (Asteraceae: Eupatorieae) by Guy L. Nesom—2.1(3):1323 Bromus hallii (Poaceae), a new combina- tion for California, U.S.A.,and taxonomic notes on Bromus orcuttianus and Bromus grandis by Jerrery M. SAareta and Paut M. Peterson—21(4):1997 Calamagrostis coahuilensis and C. divaricata (Poaceae: Pooideae: Agrostidinae), two new species from México by Paut M. Peterson, Rosert J. SorENG, and Jesus VALDES-REYNA—21(1):311 Carbon use by the endangered Texas wild rice (Zizania texana, Poaceae) by PAULA Power and Rosert D. DoyLe—21(1):389 Changes in baygall vegetation from 1986 to 2001 at Fort Polk in west central Loui- siana by CHarves M. ALLEN, JOHN Pare, SARA THAMES, SPENCER TRICHELL, aNd Lacy EZeLL— 21(1):419 Checklist of the vascular plants of Greene County, Pennsylvania by Rosert Coxe, Cyn- THIA M. Morton, Mary Joy Haywoop, Bonnie L. Isaac, and JoserH A. Isaac —21(3):1829 Chromosome numbers of Glandularia (Verbenaceae) from central and Trans- Pecos, Texas by B.L. Turner and A. MicHaet Poweltt—21(3):1657 Cliff Cudweed at specific rank in Pseudognaphalium (Asteraceae: Gnaphalieae) by Harvey E. BALLARD, JR., Daniette Sky Fetter, and Guy L. Nesom— 21(2):/77 Comments on the rediscovery and distri- bution of Cunila origanoides (Lamiaceae) in Texas by JASON R. SINGHURST 2474 and Walter C. Ho.mes—21(2):1161 Contributions to the moss flora of the Amazonian lowlands of Madre de Dios, Peru by Piers Masestyk and Joun P. JaNOovec—21(2):1231 Correct author citation for Cirsium eatonii var. eriocephalum (Asteraceae: Cardueae): erratum for Sida 21:212.2004 by Davio J. Ket—21(3):1645 Correction to the type citation of Sarrace- nia alabamensis and validation of the name Sarracenia alabamensis subsp. wherryi (Sarraceniaceae) by F.W. Case, JR.—2.1(4):2169 Croton bigbendensis (Euphorbiaceae), a new species from Trans-Pecos, Texas by B.L. TURNER—21(1):79 Cuniculotinus and Lorandersonia, two new genera of Asteraceae: Astereae and new combinations in Chrysothamnus by Lowett E. UrgatscH, ROLAND P.Roserts,and Kurt M. Neusic—21(3):1615 Cuscuta (Convolvulaceae)—the strength of weakness: a history of its name, uses and parasitism concept during ancient and medieval times by Mikal Costea and FRANCOIS J. TARDIF—21(1):369 Cuscuta indecora (Convolvulaceae) new for Canada by Minai Coste, Francois J. Taroir, and VERNON L. Harms—21(2):1229 Dahlia sublignosa (Asteraceae): a species in its own right by Daye E. Saar and Paut D. SoreNSEN—21(4):2161 Description of asexual reproduction in a Texas succulent Sedum wrightii (Crassulaceae) by Dennis A. Gravatt and JOSEPHINE TAYLOR—21(2):943 Dichanthelium scoparium — and Muhlenbergia glabrifloris: new to the flora of Ohio by RicHaro L. GARDNER, JAMES SIDA 21(4): 2474. 2005 BRIT.ORG/SIDA 21(4) S. McCormac, and Davio Minney— 21(1):465 Dichanthium (Poaceae) new to Arizona: open door for a potentially invasive spe- cies by RicHarb Fevcer, Tony L. Burcess, SAM- vet Dorsi, JOHN R. REEDER, aNd THomas R. VAN DevenDeER—21(3):1905 Diospyros torresii (Ebenaceae): a new black zapote from tropical Mexico by MircHeLt C. Provance and Anorew C. SAND- ERS—21(4):2045 Documented chromosome numbers 2005: 1. Miscellaneous counts from western Texas, mostly Trans-Pecos by A. MicHaet Powett and B.L. TuRNeR—21(3):1663 Documented chromosome numbers 2005: 2. Counts from western Texas, mostly Trans-Pecos cacti by A. MicHaet Powe. and James F. Weepin—21(3):1665 Documented chromosome numbers 2005: 3. Chromosome numbers in North American species of Silene and Stellaria (Caryophyllaceae) by J.K. Mor- TON—21(3):1669 Eleocharis mutata (Cyperaceae) new to the flora of North America north of México by Davio J. Rosen and Stantey D. Jones—21(2):1153 Emilia fosbergii (Asteraceae: Senecionae) in Texas revisited by W.P. Spencer and J.K. WiILUAMS—21(3):1937 Endemic vascular plants of the Interior Highlands, U.S.A. by Douctas ZOLLNeR, MicHaetL H. MacRoserts, BARBARA R, MacRoserts, and Douctas Lapop— 21(3):1781 Eragrostis (Poaceae: Chloridoideae: Eragrostideae: Eragrostidinae) from northeastern México by Paut M. Peterson and Jesus Vatoés-REYNA—21(3):1363 INDEX Erigeron peregrinus and Erigeron glacialis (Asteraceae: Astereae) by Guy L. Nesom—21(2):665 Estudios en las Apocynaceae neotropicales IV: notas taxondmicas en Prestonia (Apocynoideae: Echiteae) con una nueva especie de Ecuador by J.FRANcIscO Morates—21(1):159 Estudios en las Apocynaceae neotropicales V:una nueva especie, nuevos reportes y nueva sinonimia en las Apocynaceae de Bolivia by J. Francisco Moraes y ALFREDO FUENTES —21(1):165 Estudios en las Apocynaceae neotropicales X: definicidn de la verdadera identidad de Mandevilla leptophylla, con la descripcidn de dos nuevas especies y una nueva combinacién de Mandevilla (Apocynoideae: Mesechiteae) para Sur América by J. Francisco MorALeS— 21(3):1535 Estudios en las Apocynaceae neotropicales Xl: una nueva especie de Mandevilla (Apocynoideae: Mesechiteae) para Sur América, con un nuevo reporte para las Apocynaceae de Paraguay by J. Francisco Morates—2.1(3):1549 Estudios en las Apocynaceae Neotropicales XVII: una revisi6n del género Galactophora (Apocynaceae: Apocynoideae) by J. Francisco MorALES— 21(4):2053 Ethnobotanical report from mangroves of Pichavaram, Tamil Nadu State, India by K. VENKATESAN, V. BALAKRISHNAN, K.C. RAVINDRAN, and V. DevanaTHAN—2.1(4):2243 Ethnobotany of Rhodiola rosea (Crassulaceae) in Norway by TorBj@RN Atm—2.1(1):321 Eugenia inversa (Myrtaceae), a new spe- SIDA 21(4): 2475. 2005 2475 cies from Espirito Santo, Brazil by Marcos SoBRAL—2 11 (3):1465 First record of Clitoria mariana (Leguminosae) in western Texas by BriAN P.Oswatp, M BaTAINEH, and AMANDA ROUNTREE—21(1):507 First record of Nymphoides indica (Menyanthaceae) in Texas by Ken SAun- DERS—21(4):2441 Four new species of Ericaceae (Vaccinieae) from Ecuador by James L. Luteyn— 21(3):1269 Gamochaeta coarctata, the correct name for Gamochaeta spicata (Asteraceae: Gnaphalieae) by JoHn F.PRuski and Guy L. Nesom—21(2):711 Generic placement of Chaptalia hintonii (Asteraceae: Mutisieae) by Guy L.Nesom— 21(2):929 Geocarpon minimum (Caryophyllaceae), new to Texas by Eric L. KelTH, JASON R. SINGHURST, and STAN Cook—21(2):1165 Gibasis pellucida (Commelinaceae), a new and potentially weedy genus and species for Texas by Davin J. Rosen and Rosert B. FabeN—21(3):1931 Herbarium acronym correction for Ipo- moea seaania (Convolvulaceae) holo- type: Sida 21(3). 2005 by RicHaRD FELGER and Daniet F. Austin—21(4):2193 High resolution GIS mapping and current status of the ten viable populations of Short's goldenrod (Solidago shortii- Asteraceae) in Kentucky by Brent D.SMitH, Anprew 1. DENHAM, JAMES B. BECK, and Patrick J. Catie—21(2):1121 History and eponymy of the genus name Amsonia (Apocynaceae) by James S. PRIN- GLE—21(1):379 Hydrocotyle sibthorpioides (Apiaceae) 2476 new for Texas and notes on introduced species by Barney L. Liescoma and GeorGe M. Dias, JR—2.1(4):2449 ICBN clarification needed: use of ranks by Guy L. Nesom and Barney L. Liescoma— 21(4):2185 Infrageneric classification of Liatris (Asteraceae: Eupatorieae) by Guy L. Nesom—21(3):1305 Introduced species in Kansas: floristic changes and patterns of collection based on an historical herbarium by Ter- ESA M. Woops, SUZANNE C. STRAKOSH, MADHAV P. NEPAL, SEEMANTI CHAKRABARTI, NICHOLAS B. SIM- PSON, Mark H. MayrieLb, and Carotyn J. Fer- GUSON—2 1(3):1695 Ipomoea seaania, a new species of Convolvulaceae from Sonora, Mexico by RICHARD Fetcer and Daniet F. Austin— 21(3):1293 Isolation and identification of fungi asso- ciated with the rhizosphere and rhizoplane of wild and cultivated plants of Pakistan by SHAMIM AKHTAR QuRESHI, VIQAR SULTANA, SYED EHTESHAMUL-HaQue, and Mo- HAMMAD ATHAR—21(2):1019 Juniperus ashei (Cupressaceae): physiog nomy and age structure in three mature Texas stands by Caren McLemore, GLENN C. KROH, aNd JOHN E. Pinper III—21(2):1107 Kathleen M. Peterson by James L. Reveat— 21(2):1239 Las especies de Nototriche (Malvaceae) de Ecuador by Macpa CHANCO y CARMEN ULLOA ULLoA—2.1(2):693 Lectotypification and a new combination in Matelea (Apocynaceae: Asclep- iadoideae) for an endemic Hispaniolan vine by ALexander KriInGs—21(4):2081 SIDA 21(4): 2476, 2005 BRIT.ORG/SIDA 21(4) Lectotypification of Passiflora affinis (Passifloraceae) and discussion of its geographic range within the United States by Douctas H. Gotoman— 21(1):275 Mikania buchtienii (Asteraceae: Eupatorieae) new to Argentina by Maria M. Cerana and Luis Ariza-EsrinAk— 21(4):2425 Miscellaneous chromosome number re- ports for Poa (Poaceae) in North America by Rosert J. SoRENG—21(4):2195 Miscellaneous nomenclatural changes in Astereae (Asteraceae) by JOHN C. SemPLE— 21(2):759 Morfologia de tric en especies de Solanum secci6n Acanthophora (Solanaceae), presentes en Venezuela by 1 ated Fe Wittiam A. GRANADA-CHACON yY CARMEN E, Benitez DE RoJAS—21(3):1675 Muscari comosum (Liliaceae) new to Texas by Lee M. Luckeyooo—21(4):2445 Neotypification of Ceropegia palustris and Lyonia maritima (Apocynaceae: Asclepiadoideae) by ALEXANDER KRINGS— 21(3):1507 Nestotus and Toiyabea, two new genera of Asteraceae: Astereae from the west- ern United States and Canada by Rotano P. Roserts, Lowett E. UrsaTscH, and Kurt M. Neusic—21(3):1647 New combination in Conoclinium (Asteraceae: Eupatorieae) by THomas F. PATTERSON—2.1(3):1641 New combinations in Artemisia (Asteraceae: Anthemideae) by Leia M. SHULTZ—2 1(3):1637 New combinations in Eutrochium (Asteraceae: Eupatorieae), an earlier name than Eupatoriadelphus by Eric E. LaMONT—21(2):901 INDEX New combinations in Festuca californica (Poaceae) by StePHen J. DARBYSHIRE— 21(3):1455 New combinations in North America Eremogone (Caryophyllaceae) by RONALD L. HARTMAN and RicHarb K. RABELER— 21(1):237 New combinations in North American Caryophyllaceae by J.K. Morton— 21(2):887 New combinations in North American Caryophyllaceae by Ronatp L. HarTMAN and RicHarb K. RabeELER—21(2):753 New combinations in North American Symphyotrichum subgenus Astro- polium (Asteraceae: Astereae) by Scott D. SUNDBERG—21(2):903 New combinations in the genus Gundlachia and four new genera of Astereae (Asteraceae) from northern Mexico and the southern United States by Lowe tt E. UrsatscH and Rotano P. Ros- ERTS—21(1):243 New distribution records for Gamochaeta (Asteraceae: Gnaphalieae) in the United States by Guy L. Nesom—21(2):1175 New records in Pseudognaphalium (Asteraceae: Gnaphalieae) from Ne- braska and California by Steven B. RotrsmeleR and Guy L. Nesom—21(2):1205 New records of Erigeron (Asteraceae: Astereae) for Nevada by Guy L. Nesom— 21(1):473 New records of pteridophytes for the flora of Peru by ALan R. Smith, BLANCA LEON, HANNA Tuomisto, HENK VAN DER Werre, Rossin C. Moran, Marcus LeHNert, and MICHAEL KESSLER—21(4):2321 New reports of Eurybia and Aster s.str. (Asteraceae: Astereae ) from California, SIDA 21(4): 2477. 2005 2477 Idaho, and Wyoming by Luc BrouitteT— 21(1):459 New species and new combinations in Ericameria (Asteraceae: Astereae) by RoLanbD P. Roberts, Lowett E. UrsatscH, and JOHN ANDERSON—21(3):1557 New species of Gamochaeta (Asteraceae: Gnaphalieae) from the eastern United States and comments on similar species by Guy L. Nesom—21(2):717 New taxa and new combinations in North American Cirsium (Asteraceae: Cardueae) by Davio J. Ket —21(1):207 Nomenclature and typification in the ge- nus Usnea (lichenized ascomycetes)— lV. Usnea stuppea & Usnea substerilis by James C. LeNDemeR and Isaselte I. TAVARES— 21(2):637 Nomenclature and typification in the ge- nus Usnea (lichenized ascomycetes)— ill. Usnea alata & Usnea sulcata by James C. LenDemer and Isasette |. TAVARES— 21(2):643 Nomenclature of Ipomoea arborescens (Convolvulaceae) in Sonora, Mexico by Daniet F. Austin, RICHARD FetGer, and THOMAS R. VAN DeveNnbeER—21(3):1283 Nomenclature of the Virginia-bluebell, Mertensia virginica (Boraginaceae) by James S. PRINGLE—21(2):767 Notes on “coffee” from the Kentucky coffeetree (Gymnocladus dioicus, Fabaceae) by JoHN P. SpaeTH and JoHN W. THIERET—2 1(1):345 Notes on Libertia (lridaceae: Sisyrinchieae) in South America by Peter Gotosatt and Marcela Cetis—21(4):2105 Notes on Louisiana botany and botanists, 1718-1975 by JosePH 21(4):2275 A. EWAN— 2478 Notes on North American arctic and bo- real species of Erigeron (Asteraceae: Astereae) by Guy L. Nesom and Davip F. Murray—21(1):41 Notes on the distribution and nomencla- ture of North American Gentianopsis (Gentianaceae) by James S. PRINGLE— 21(2):525 Notes on the distribution of Pseudognaphalium luteoalbum (Asteraceae: Gnaphalieae) by Guy L. NesomM—21(1):463 Notes on the Matelea bayatensis- correllii-tigrina complex (Apocy- naceae: Asclepiadoideae: Gonolobinae) in the Greater Antilles and Bahamas by ALEXANDER KRINGS—21(3):1525 Notes on typification in Pluchea (Asteraceae: Plucheae) by Guy L.Nesom— 21(1):59 Noteworthy collections of Cyperus drummondii (Cyperaceae) from Texas by Davio J. Rosen—21(1):495 Osmorhiza bipatriata (Apiaceae) in Texas: taxonomic status and conservation con- siderations by Jun Wen, Porter P. Lowey Il, and James C. ZEecH—21(1):501 Palaeoanthella huangii gen.and sp.nov., an Early Cretaceous flower (Angio- spermae) in Burmese amber by GForce PoinarR JR. and Kenton L. CHAMBERS— 21(4):2087 Panphalea heterophylla (Compositae: Mutisioideae: Nassauvieae),a genus and species new for the flora of North America by JoHn F. Pruski—21(2):1225 Pappus variation in North American asters. |. Double, triple and quadruple pappus in Symphyotrichum and related aster genera (Asteraceae: Astereae) by JOHN C. SIDA 21(4): 2478. 2005 BRIT.ORG/SIDA 21(4) Sempce and Jennirer L.A. Hoop —21(4):2141 Phalaris arundinacea (Poaceae: Aveneae) a species new to Texas and a key to Phalaris in Texas by Stepan L.HatcH, DALE A. Kruse, and JENNIFER PLUHAR—21(1):487 Pharus’ primuncinatus (Poacae: Pharoideae: Phareae) from Dominican amber by Georce Poinar Jr. and Emmet J. Jupziewicz—2 1(4):2095 Pinguicula vulgaris (Lentibulariaceae) and its uses in Norway by Torsiorn ALM— 21(4):2249 Polyphyly of the genus Echites (Apocy- naceae: Apocynoideae: Echiteae): evi- dence based on a morphological cladis- tic analysis by Justin K. Wittiams— 21(1):117 Potential biological control of Lantana camara in the Galapagos using the rust Puccinia lantanae by Joarce Luis Renteria B.and Carot Ettison—21(2):1009 Pseudognaphalium canescens (Asteraceae: Gnaphalieae) and putative relatives in western North America by Guy L. Nesom—21(2):781 Pseudostellaria oxyphylla (Caryop- hyllaceae), a long overlooked species from northern Idaho by Ronato L. Hart- MAN and RicHarb K. RaBELER—21(1):175 Ptilagrostis luquensis (Poaceae: Pooideae: Stipeae: Stipinae), a new spe- cies from China by Paut M. Peterson, Ros- ert J. Sorenc, and ZHeEN-LAN Wu— 21(3):1355 Quercus montana (Fagaceae), new to Mis- souri by Marian Smith and Nancy PARKER— 21(3):1921 Ranunculus ficaria (R to North Carolina and an updated key to Carolina congeners by ALEXANDER ulaceae), new INDEX Krincs, ALAN S. Weaktey, JOSEPH C. NEAL, and Eowaro C. Swas—21(4):2429 Recognition of Phragmites australis subsp. americanus (Poaceae: Arundinoideae) in North America: evi- dence from morphological and genetic analyses by Kristin SALTONSTALL, PauL M. Peterson, and Rosert J. SoRENG—21(2):683 Rediscovery of Ponthieva brittoniae (Orchidaceae) in Everglades National Park by Jimi L.SADLE, STEVEN W. WooDMANSEE, Georce D.GANN,and THomas V. ARMENTANO— 21(3):1917 Reference conditions of the Red River floodplain and upland, Caddo Parish, Louisiana by MicHaeL H. MacRoserts and Barpara R. MacRoserts—21(3):1 793 Reflections on the taxonomy and distribu- tion of medicinal flowers of Pakistan by MoHammabd ATHAR and M. Akmat Sippiqi— 21(1):357 Reflections on William Chambers Coker, passionate botanist by Mary Coker JOSLIN—-2 1(2):977 Relationships between plant folklore and antitumor activity: an historical review by Richard W. SPiutT—21(4):2205 Response to “The Gerbera complex (Asteraceae: Mutisieae): to split or not to split” by Liliana Katinas by Guy L.Nesom— 21(2):941 Resurrection of a little-known species of Oenothera sect. Oenothera in north- eastern Mexico (tribe Onagreae: Onagraceae) by Warren L. WAGNER— 21(2):651 Revision of Fevillea (Cucurbitaceae: Zanonieae) by Georce L. Rosinson and Rr CHARD P. WuNDeERLIN—21(4):1971 Revision of Lobelia sect. Homochilus SIDA 21(4): 2479, 2005 2479 (Campanulaceae: Lobelioideae) by THo- mas G. LAMMERS—2.1(2):591 Revision of Siolmatra (Cucurbitaceae: Zanonieae) by Georce L. Rosinson and Ri- CHARD P. WUNDERLIN—2.1(4):1961 Ruppia cirrhosa (Ruppiaceae) in north central Texas by Tirrany L. Morcan and Watter C. Hommes —21(1):499 Sarracenia purpurea (Sarraceniaceae) In Louisiana by MicHaet H. MacRoserts and Bareara R. MacRoserts—21(2):1149 Sawtooth oak (Quercus acutissima, Fagaceae) in North America by Atan T. WuHiTTEMORE—21(1):447 Schoenoplectus hallii (Cyperaceae), a globally threatened species new for Texas by Rosert J. O’KENNON and CAREN McLemore—21(2):1201 Seed and capsule characters in Arcytophyllum, Bouvardia, and Manettia (Rubiaceae), with notes on A. serpyllaceum by Epwaro E. Terreit and Harotd Rosinson—21(2):911 Silene conoidea (Caryophyllaceae) new to Texas by Monique Dusrute Reep>— 21(1):493 Soil and ecological features of Hexalectris (Orchidaceae) sites by Annie B. COoLLns, James E. VaRNuM, and Marcaret BROWN- Marsoen—2.1(3):1879 Solanum viarum (Solanaceae)— Tropical Soda-Apple—confirmed from Texas by Monique DuBRute REED, Mary KeTCHERSID, and RicHarD L. THOMPSON—21(2):1171 Solidago sect. Ptarmicoidei,a new com- bination to replace a “rankless” name used by Torrey and A. Gray (Asteraceae: Astereae) by JoHN C.Sempete and Kanci N. GANDHI—2 1(2):755 Sporobolus (Poaceae: Chloridoideae: — 2480 Cynodonteae: Zoysieae: Sporobolinae) from northeastern Mexico by Paut M. PETERSON, Jesus VALDES-REYNA, aNd JUAN JAVIER Ortiz-Diaz—2.1(2):553 Sporobolus coahuilensis (Poaceae): a new record for the U.S.A from Trans- Pecos, Texas by B.L. TurNeR—21(1):455 Sporobolus heterolepis (Poaceae), new to Tennessee by Dwayne Estes and JOHN Beck —21(3):1923 State records and other noteworthy collec- tions for Kentucky by Ross C. CLark, RONALD L. Jones, TimotHy J. Weckman, RaceH L. THOMP- SON, JOHN W. THieRET, Kentucky Stare NATURE PRESERVES COMMISSION, aNd Kim FeEMAN— 21(3):1909 Staurochilus leytensis, a Philippine seg- regate of Staurochilus fasciatus (Orchidaceae: Aeridinae) by Eric A. CHRISTENSON—2.1(4):2051 Stemodia coahuilensis (Scrophulari- aceae), a new record for the United States by B.L. Turner—21(3):1935 Studies of Neotropical Compositae—I., Novelties in Calea, Clibadium, Conyza, Llerasia,and Pluchea by Joxn F. Pruski— 21(4):2023 a ymphy pyg transfer of Eurybia pygmaea from the eurybioid grade to the subtribe Symphyotrichinae (Asteraceae: Astereae) by Luc Brouittet and SUGIRTHIN| SELLIAH—21(3):1633 Systematics of Dirca (Thymelaeaceae) based on its sequences and ISSR poly- morphisms by James A. SCHRADER and WiL- LIAM R. GRaveS—21(2):11 Taxonomic notes on Krigia (Asteraceae) by KENTON L. CHAMBERS—21(1):225 Taxonomic reevaluations in North Ameri- SIDA 21(4): 2480. 2005 BRIT.ORG/SIDA 21(4) can Erigeron (Asteraceae: Astereae) by Guy L. Nesom—21(1):19 Taxonomic review of Astranthium integrifolium (Asteraceae: Aster Guy L. Nesom—21(4):2015 Taxonomic revision of the neotropical ge- nus: Erithalis (Rubiaceae: Chiococceae) by Vivian NeGRON-Ortiz—21(3):1565 Taxonomy of Hymenoxys subgenus Macdougalia (Asteraceae: Helenieae: Tetraneurinae) by Mark W. BieRNeR— 21(2):657 Taxonomy of VK de) Dy the Liatris pilosa (graminifolia) complex (Asteraceae: Eupatorieae) sy Guy L. Nesom and Jon M. Stucky —21(2):815 Taxonomy of the Polemoniaceae:Gilia and Lathrocasis by Verne Grant—21(2):531 Taxonomy of the Symphyotrichum (Aster) subulatum group and Symphyotri- chum (Aster) tenuifolium (Asteraceae: Astereae) by Guy Nesom—21(4):2125 Taxonomy, distribution, and medicinal uses of legume trees of Pakistan by Monam- MAD ATHAR and ZAHOOR AHMAD—21(2):951 The Actin | intron—a phylogenetically in- formative DNA region in Clematis (Ranunculaceae) by JonatHAn Soma, James Garey, and Freperick B. Essic— 21(2):879 The ecology of Trillium texanum (Trilliaceae) on the Angelina National Forest, Texas by MicHaet H. MacRoserts and Barsara R. MacRoperTs—2 1(3):1893 The floristic ecology of xeric limestone prai- ries in Kentucky, and a comparison to limestone cedar glades and deep-soil barrens by PJ. Law ess, JM. Baskin, and C.C. Baskin—21(2):1055 INDEX The genus Prenanthes (Asteraceae: Lactuceae) in Texas by JASon R. SINGHURST, Rosert J.O’KeNNon, and Watter C. HoLmes— 21(1):181 The Gerbera complex (Asteraceae: Mutisieae): to split or not to split by LILIANA KATINAS —2 1(2):935 The Gonolobus complex (Apocynaceae: Asclepiadoideae) in the southeastern United States by ALexanper Krincs and (Jenny) Olu-YUN XIANG—21(1):103 The occurrence of Cerastium pumilum (Caryophyllaceae) in Oklahoma by Bruce W. Hoactand, Amy BuTHob, and GLorIA CADDELL—21(4):2439 The Post Oak Savanna ecoregion:a floristic assessment of its uniqueness by MicHaeL H.MacRoserts and Bargara R.MacRoserts— 21(1):399 The rediscovery of the South American Hybanthus parviflorus (Violaceae) in North America by B. Eucene Worroro, JULIANA DE PaULA-SouzA, ALAN S. Weakley, and THomas E, Govus—21(2):1209 The vascular flora of Giles County, Tennes- see by Dwayne Estes—21(4):2343 The vascular flora of Montgomery County, Arkansas by Travis D. Marsico— 21(4):2389 The vascular flora of Rattlesnake Falls: a potential state natural area on the West- ern Highland Rim Escarpment in Tennes- see by Dwayne Estes and Jerrrey L. WALcK— 21(3):1753 The vascular plants of a forest fragment in southern Bahia, Brazil by ANoré M. Amorim, PEDRO FiascHi, JOMAR G.JARDIM, Wa.Wayt THO- MAS, BRENDA C. CLIFTON, and ANoré Mauricio V. DE CARVALHOT—21(3):1 727 SIDA 21(4): 2481. 2005 2481 Three new species of Eriocaulon (Eriocaulaceae) from peninsular India by SACHIN ANIL PUNEKAR, NiLESH VUaY MALpure, and P. LAKSHMINARASIMHAN—21(2):625 Three previously undescribed species of Vaccinium (Ericaceae) from Costa Rica and Panama by Rosert L.Witgur and James L. Luteyn—21(3):1607 Thymophylla setifolia var. greggii (Compositae) by JoHN L. StROTHER— 21(1):287 Tonestus kingii and T. aberrans are re- lated to Eurybia and the Machae- rantherinae (Asteraceae: Astereae) based on nrDNA (ITS and ETS) data: reinstate- ment of Herrickia and a new genus, Triniteurybia by Luc Brouitter, L.URBATSCH and R.P. Rogerts—21(2):889 Two new combinations in the genus Packera (Asteraceae) by Desra K. TRock— 21(1):289 Two new subspecies of Microseris laciniata (Asteraceae) from the Siskiyou Mountains by Kenton L. CHAMBERS— 21(1):193 Two new varieties of Agoseris (Asteraceae: Lactuceae) by Gary |. Bako —21(1):267 Una nueva especie de Aiphanes (Arecaceae) de la Cordillera de Mérida, Venezuela by Santos Micuet Nino, LJ. Dorr, and Frep W. StauFFER—21(3):1599 Una nueva especie de Guarea (Meliaceae) para Costa Rica by ALEXANDER RopriGuez— 21(4):2039 Una nueva especie de Struthanthus (Loranthaceae) para Costa Rica by Luis A. GonzAtez y J. Francisco MorRALES—21(1):97 Understanding Platanthera chapmanii (Orchidaceae), its origins and hybrids by 2482 BRIT.ORG/SIDA 21(4) Paut Martin BROWN—21(2):853 ButHoo, and Wayne EtistNsS—21(2):1187 Vascular flora of Hackberry Flat, Frederick Vegetation and flora of American Beech Lake, and Suttle Creek, Tillman County, Woods Nature Preserve, Clark County, II- Oklahoma by Bruce W. HOAGLAND, PRISCILLA linois by Bos Eoin, Gordon C. Tucker, and H.C. CRAWFORD, PHILLIP T. CRAWFORD, and For- JOHN E. Esincer—21(3):1861 REST JOHNSONT—21(1):429 Wilbur Howard Duncan, 1910-2005 by Vascular flora of Washita Battlefield Na- Wenoy B. ZomtererR and Davio E. GIANNASI— tional Historic Site, Roger Mills County, 21(3):1941 Oklahoma by Bruce W. Hoactanp, Amy SIDA 21(4): 2482. 2005 INDEX 2483 INDEX OF 290 AuTHorRs: VOLUME 21 (2004-2005) Thank you for choosing Sida, Contributions to Botany. Ahmad, Zahoor—21(2):951 Allen, Charles M—21(1):409, 419 Allred, Kelly W.—21(4):2113 a Alm, Torbjarn—21(1):321;21(4):2249 Amorim, André M.—21(3):1 727 Anderson, Jonn—21(3):1557 Ariza-Espinar, Luis—21(4):2425 Armentano, Thomas V.—21(3):1917 Athar, Mohammad—21(1):357;21(2):951; 1019 Austin, Daniel F.—21(3):1283, 1293; 21(4):2193 Baird, Gary |—21(1):267; 21(2):715 Balakrishnan, V—21(4):2243 Ballard, Jr, Harvey E-—21(2):777 Baskin, C.C.—21(2):1055 Baskin, JM.—21(2):1055 Bataineh, Mohammad—21(1):507 Bayer, Randall James—21(2):767 Beck, James B.—21(2):1121 Beck, Jonn—21(3):1923 Benitez de Rojas, Carmen E.—21(3):1675 Bettinger, Kelly A—21(2):1081 Bierner, Mark W.—21(2):657 Brouillet, Luc—21(1):459; 21(2):889; 21(3):1633 Brown, Larry E—21(3):1807 Brown, Paul Martin—21(2):853;21(4):2297 Brown-Marsden, Margaret—21(3):1879 Burgess, Tony L—21(3):1905 Buthod, Amy—21(2):1187, 21(4):2439 Caddell, Gloria—21(4):2439 Calie, Patrick J—21(2):1121 Canne-Hilliker, Judith M—21(3):1927 Carr, Gerald D—21(1):259 Carr, Robert L—21(1):259 1199; SIDA 21(4): 2483. 2005 Carvalhot, André Mauricio V. de— 21(3):1727 Case, Jr, FW.—21(4):2169 Cayouette, Jacques—21(2):791 Celis, Marcela—21(4):2105 Cerana, Maria M—21(4):2425 Chakrabarti, Seemanti—21(3):1695 Chambers, Kenton L.—21(1):193, 225; 21(4):2087 Chanco, Magda—21(2):693 Chiang, Fernando—21(2):547 Chocce, Miguel—21(2):841 Christenson, Eric A.—21(2):841;21(4):2051 Clark, Ross C.—21(3):1909 Clifton, Brenda C.—21(3):1727 Collins, Annie B.—21(3):1879 Cook, Rachel E—21(1):221;21(3):1471 Cook, Stan—21(2):1165 Costea, Mihai —21(1):369; 21(2):1229 Cotterman, Laura M—21(2):827 Coxe, Robert—21(1):481;21(3):1829 Crawford, Phillip T.—21(1):429 Crawford, Priscilla H.C.—21(1):429 Darbyshire, Stephen J.—21(3):1455 Denham, Andrew T.—21(2):1121 Devanathan, V—21(4):2243 Diggs, Jr, George M—21(4):2449 Dorr, L.J—21(3):1599 Dorsi, Samuel—21(3):1905 Doyle, Robert D—21(1):389 Ebinger, John E—21(3):1861 Edgin, Bob—21(3):1861 Ehteshamul-Haque, Syed—21(2):1019 Elisens, Wayne—21(2):1187 Ellison, Carol—21(2):1009 Elsik, |. Sandra—21(3):1807 Essig, Frederick B—21(2):861,879 2484 Estes, Dwayne—21(3):1753, 21(4):2343 Ewan, Joseph A.—21(4):2275 Ezell, Lacy—21(1):419 Faden, Robert B —21(3):1931 Feeman, Kim—21(3):1909 Felger, Richard—21(3):1283, 1293, 1905; 21(4):2193 Feller, Danielle Sky —21(2):777 Ferguson, Carolyn J.—21(3):1695 Fiaschi, Pedro—21(3):1727 Finot, Victor L—21(3):1419 Franklin, Carlyle—21(2):1131 Freeman, Craig C—21(1):291 Fuentes, Alfredo—21(1):165 Gandhi, Kanchi N.—21(2):755 Gann, George D.—21(3):1917 Gardner, Richard L—21(1):465 Garey, James—21(2):879 Giannasi, David E—21(2):1081;21(3):1941 Giuliano, Daniel A——21(2):705 Goldblatt, Peter—21(4):2105 Goldman, Douglas H—21(1):275 Gonzélez, Luis A—21(1):97 Govus, Thomas E.—21(2):827, 1209 Granada-Chacon, William A—21(3):1675 Grant, Verne—21(2):531 Gravatt, Dennis A.—21(2):943 Graves, William R.—21(2):11 Harcombe, Paul A.—21(3):1807 Harms, Vernon L.—21(2):1229 Hartman, Ronald L.—21(1):175, 237; 21(2):753 Hatch, Stephan L.—21(1):487 Hays, John F—21(3):1927 Haywood, Mary Joy—21(3):1829 Hennigan, Melissa—21(2):1 141 Henrickson, James—21(1):1, 11 SIDA 21(4): 2484. 2005 1923: BRIT.ORG/SIDA 21(4) Hoagland, Bruce W.—21(1):429; 21(2):1187, 1199; 21(4):2439 Holmes, Walter C.—21(1):181, 499; 21(2):1161 Hong, De-Yuan—21(2):993 Hood, Jennifer L.A.—21(4):2141 Isaac, Bonnie L—21(1):481; 21(3):1829 Isaac, Joseph A.—21(3):1829 Janovec, John P—21(2):841, 1231 Jardim, Jomar G.—21(3):1727 Johnson, Dan—21(3):1807 Johnsont, Forrest—21(1):429 Jones, Ronald L—21(3):1909 Jones, Stanley D—21(2):1153 Joslin, Mary Coker—21(2):977 Judd, Walter S—21(2):1081 Judziewicz, Emmet J.—21(3):1263; 21(4):2095 Kartesz, Jonn—21(1):481 Katinas, Liliana—21(2):935 Kauffman, Gary L.—21(2):827 Keil, David J—21(1):207;21(3):1645 Keith, Eric L—21(2):1 165; 21(3):1927 Kentucky State Nature Preserves Commis- sion,—21(3):1909 Kessler, Michael—21(4):2321 Ketchersid, Mary—21(2):1171 Kral, Robert—21(1):293 Krings, Alexander—21(1):103:21(2):1131; 21(3):1507, 1515, 1519, 1525; 21(4):2081, 2429 Kroh, Glenn C.—21(2):1107 Kruse, Dale A—21(1):487 Kruse, Lisa M.—21(2):1081 Kumar, Muktesh—21(1):93 Ladd, Douglas—21(3):1781 Lakshminarasimhan, P—21(2):625 Lammers, Thomas G.—21(2):591 Lamont, Eric E—21(2):901 INDEX Landrum, Leslie R.—21(3):1335 Lawless, PJ —21(2):1055 Lehnert, Marcus—21(4):2321 Lendemer, James C.—21(2):637,643 Leon de la Luz, José Luis—21(2):547 Leon, Blanca—21(4):2321 Leonard, Michelle R—21(3):1471 Lipscomb, Barney L.—21(4):2185, 2449 Lowry Il, Porter P—21(1):501 Luckeydoo, Lee M—21(4):2445 Luteyn, James L.—21(3):1 269, 1607 MacRoberts, Barbara R.—21(1):399; 21(2):963, 1149; 21(3):1781, 1793, 1807, 1893 MacRoberts, Michael H.—21(1):399; 21(2):963, 1149; 21(3):1781, 1793, 1807, 1893 Majestyk, Piers—21(2):1231 Malpure, Nilesh Vijay—21(2):625 Marsico, Travis D—21(4):2389 Mayfield, Mark H.—21(3):1695 McCormac, James $.—21(1):465 McKenzie, Paul—21(1):475 McLemore, Caren—21(2):1107, 1201 Mettler-Cherry, Paige —21(1):475 Minney, David—21(1):465 Morales, J. Francisco—21(1):97, 133, 159, 165; 21(3):1535, 1549; 21(4):2053 Moran, Robbin C.—21(4):2321 Morgan, Tiffany L—21(1):499 Morse, Caleb A.—21(4):2093 Morton, Cynthia M.—21(1):481; 21(3):1829 Morton, J.K.—21(2):887; 21(3):1669 Murray, David F—21(1):41 Neal, Joseph C.—21(4):2429 Negrén-Ortiz, Vivian—21(3):1565 Nelson, John B.—21(4):2119 Nelson, Thomas W.—21(2):673 SIDA 21(4): 2485. 2005 2485 Nepal, Madhav P—21(3):1695 Nesom, Guy L.—21(1):19, 41,59, 463, 473; 21(2)665,°673,.7 Vj 717, TA TE, SI, 815,027 929) 94) ed 75-2105; 12153 21(3):1305, 1323; 21(4):2015, 2125, 2185 Neubig, Kurt M.—21(3):1615, 1647 Neyland, Ray—21(2):1141 Nino, Santos Miguel—21(3):1599 O’Kennon, Robert J—21(1):65, 181, 311; 21(2):683, 1201 Ortiz-Diaz, Juan Javier-—21(2):553 Oswald, Brian P—21(1):507 Parker, Nancy—21(3):1921 Pate, John—21(1):419 Patterson, Thomas F—21(3):1641 Paucar, Arturo Granda—21(2):705 Paula-Souza, Juliana de—21(2):1 209 Peterson, Paul M—21(1):311; 21(2):553, 683; 21(3):1355, 1363, 1419; 21(4):1997 Phipps, J.B.—21(1):65 Pinder Ill, John E.—21(2):1107 Pluhar, Jennifer—21(1):487 Poinar Jr., George—21(4):2087, 2095 Powell, A. Michael —21(3):1657, 1663, 1665 Power, Paula—21(1):389 Price, Dana—21(3):1927 Pringle, James S.—21(1):379; 21(2):525, 767; 21(3):1249 Provance, Mitchell C—21(4):2045 Pruess, Warren W.—21(3):1807 Pruski, John F.—21(2):711, 21(4):2023 Punekar, Sachin Anil—21(2):625 Qiu-yun Xiang, Jenny)—21(1):103 Qureshi, Shamim Akhtar—21(2):1019 Rabeler, Richard K.—21(1):175, 237; 2112) 753 Ravindran, K.C._—21(4):2243 Reed, Monique Dubrule—21(1):493; 1223; 2486 21(2):1171;21(3):1927 Reeder, John R—21(3):1905 Remesh M.—21(1):93 Renteria B., Jorge Luis—21(2):1009 Reveal, James L.—21(2):679, 1239 Roberts, R.P—21(2):889 Roberts, Roland P—21(1):243;21(3):1557, 1615, 1647 Robinson, George L.—21(4):1961, 1971 Robinson, Harold—21(2):911 Rodriguez, Alexander—21(4):2039 Rolfsmeier, Steven B.—21(2):1205 Rosen, David J.—21(1):495; 21(2):1153; 21(3):1931 Rountree, AManda—21(1):507 Saar, Dayle E—21(4):2161 Saarela, Jeffery M—21(4):1997 Sadle, Jimi L.—21(3):1917 Saltonstall, Kristin—21(2):683 Sanders, Andrew C.—21(4):2045 Saunders, Ken—21(4):2441 Schrader, James A.—21(2):11 Selliah, Sugirthini—21(3):1633 Semple, John C.—21(1):221; 21(2):755 755,759; 21(3):1471;21(4):2141 Shultz, Leila M.—21(3):1637 Siddiqi, M.Akmal—21(1):357 Simpson, Nicholas B.—21(3):1695 Singhurst, Jason R—21(1):181;21(2):1161, 1165; 21(3):1927 Slomba, Jonathan—21(2):879 Smith, Alan R—21(4):2321 Smith, Brent D—21(2):1121 Smith, Galen—21(1):475 Smith, Marian—21(1):475;21(3):1921 Sobral, Marcos—21(3):1465 Soreng, Robert J.—21(3):1355, 1419; 21(4):2195 Sorensen, Paul D.—21(4):2161 Spaeth, John P—21(1):345 SIDA 21(4): 2486. 2005 BRIT.ORG/SIDA 21(4) Spencer, W.P—21(3):1937 Spjut, Richard W.—21(4):2205 Starks, Jane K—21(3):1939 Stauffer, Fred W.—21(3):1599 Strakosh, Suzanne C.—21(3):1695 Strother, John L.—21(1):287 Stucky, Jon M.—21(2):815 Sultana, Vigar—21(2):1019 Sundberg, Scott D.—21(2):903 Swab, Edward C.—21(4):2429 Tardif, Francois J—21(1):369; 21(2):1229 Tavares, |lsabelle |—21(2):637, 643 Taylor, Josephine—21(2):943 Terrell, Edward E.—21(2):91 1 Thames, Sara—21(1):409,419 Thieret, John W.—21(1):345;21(3):1909 Thomas, Wm.Wayt—21(3):1727 Thompson, Ralph L.—21(3):1909 Thompson, Richard L.—21(2):1171 Trichell, Soencer—21(1):409 Trock, Debra K—21(1):289; 21(3):1643 Tucker, Gordon C.—21(3):1861 Tuomisto, Hanna—21(4):2321 Turner, B.L.—21(1):455, 79, 87; 21(2):679; 21(3):1657, 1663, 1935 Ubelaker, John E.—21(3):1939 Ulloa, Carmen Ulloa—21(2):693 Unnikrishnan, N.—21(1):93 Urbatsch, Lowell E.—21(1):243-21(2):889, 1141; 21(3):1557, 1615, 1647 Valdés-Reyna, Jesus—21(1):311;21(2):553: 21(3):1363 van der Werff, Henk—21(4):2321 Van Devender, Thomas R.—21(3):1283, 1905 Varnum, James E.—21(3):1879 Venkatesan, K —21(4):2243 Wagner, Warren L.—21(2):651 Walck, Jeffrey L—21(3):1753 INDEX Ward, Daniel B—21(4):2119,2171 Weakley, Alan $.—21(2):743, 827, 1209; 21(4):2429 Weckman, Timothy J—21(3):1909 Weedin, James F—21(3):1665 Wen, Jun—21(1):501;21(2):993 White, Jeremy—21(1):409 Whittemore, Alan T.—21(1):447 Wilbur, Robert L—21(3):1607 Williams, J.K—21(3):1937 Williams, Justin K.—21(1):117, 133; 21(3):1937 SIDA 21(4): 2487. 2005 2487 Witsell, Theo—21(3):1249 Wofford, B.Eugene—21(2):1 209 Woodmansee, Steven W.—21(3):1917 Woods, Teresa M.—21(3):1695 Wunderlin, Richard P—21(4):1961,1971 Zech, James C—21(1):501 Zhen-Lan Wu,—21(3):1355 Zhou, Shi-Liang—21(2):993 Zollner, Douglas—21(3):1781 Wendy B.—21(2):1081; Zomlefer, 21(3):1941 Zuloaga, Fernando O—21(3):1419 2488 BRIT.ORG/SIDA 21(4) BOTANICAL NAMES AND SuBJECT INDEX: VOLUME 21 (2004-2005) New names (236) in bold face Acanthophora—21(3):1675 Achillea millefolium—21(2):1217 Agalinis navasotensis—21(3):1927 Agoseris—21(1):267 apargioides—21(2):715 var.maritima—21(2):716 grandiflora var. leptophylla— 21(1):267 heterophylla var. quentinii— 21(1):271 Agrostidinae—21(1):31 1 Aiphanes—21(3):1599 stergiosii—21(3):1600 Allotoonia—21(1):133, 135 agglutinata—21(1):139 caudata—21(1):145 parviflora—21(1):148 turbinata—21(1):150 tuxtlensis—21(1):153 Alternanthera paronichyoides— 21(2):1199 Amaranthaceae—21(1):1 1;21(2):1199 Amaranthus—21(1):1 1 acanthobracteatus—21(1):12 Burmese—21(4):2087 Dominican—21(4):2095 Amblyolepis setigera—21(2):1217 Ambrosia artemisiifolia—21(2):1217 American Beech Woods Nature Pre- serve—21(3):1861 Ampelaster—21(4):2141 Amphiachyris dracunculoides— Zl2hiet7 Amsonia—21(1):379 Angelina National Forest (Texas) — 21(3):1893 Antennaria—21(2):767 SIDA 21(4): 2488. 2005 pulcherrima subsp. eucosma— 21(2):768 Anthemideae—21(2):1637 Anthemis cotula—21(2):1217 Anthenantia—21(1):293 rufa—21(1):303 texana—21(1):296 villosa—21(1):299 Antitumor activity—21(4):2205 Apiaceae—21(1):501;21(2):743; 21(4):2449 Apocynaceae—21(1):103, 117,133, 159, 165, 379; 21(3):1507, 1515, 1519, 1525, 1535, 1549; 21(4):2053, 2081 Araliaceae—21(2):993 Arctium minus—21(2):1217 Arcytophyllum—21(2):911 serpyllaceum—21(2):911 Arecaceae—21(3):1599 Arenaria lanuginosa var. saxosa— 21(2):753 Argentina—21(4):2425 Aristida appressa—21(4):2117 hitchcockiana—21(4):2113 eludendae—21(4):2113 Arizona—21(3):1557, 1905 Arkansas—21(3):1 249; 21(4):2389 Artemisia—21(2):1637 annua—21(2):1217 arbuscula subsp. longiloba— 21(3):1637 biennis var. biennis—21(2):1217 globularia subsp. lutea—21(3):1638 Artemisia vulgaris var. vulgaris— 21(2):1217 Arundinoideae—21(2):683 INDEX Asclepiadoideae—21(1):103;21(3):1507, 1a S 1S 19 Fs25 Ascomycota—21(3):1939 Aster—21(1):459 alpinus subsp. vierhapperi—21(1):460 Asteraceae—21(1):19, 41,59, 181, 193, 207,221,225, 243, 259, 267, 289, 459, 463,473; 21(2):657, 665, 673, 705,711, Fallistey a We sve iso w Acy ae a cy aod eo por 827, 889, 901, 903, 929,935, 941,1121, 117561205 12 05; 21(3);1305, 1323, 1471, 1357,16 151033, 1637, 1641, 1643, 1645, 1647, 1937; 21(4):2015, 2093 Astereae—21(1):19, 41, 221, 243,459, 473: 21(2):665, 673, 705, 755, 759, 827, 889, 903; 21(3):1471, 1557, 1615, 1633, 1647 Astranthium ciliatum—21(4):2016 integrifolium—21(4):2015 Astropolium—21(2):903 Atlantic Coast—21(2):743 Aulonemia nitida—21(3):1263 Aveneae—21(1):487 Aveninae—21(3):1419 Baccharis pulchella—21(2):709 subsculpta—21(2): 705 Bahamas—21(3):1525 Bahia, Brazil—21(3):1727 Bambuseae—21(3):1263 Bambusoideae—21(1):93; 21(3):1263 Baygall—21(1):419 Bidens bipinnata—21(2):1217 frondosa—21(2):1217 pilosa—21(2):1217 polylepis—21(2):1217 Big Thicket National Preserve— 21(3):1807 Bolivia—21(1):165 SIDA 21(4): 2489. 2005 Boraginaceae—21(2):771 Boreal—21(1):41 Bouvardia—21(2):91 1 Brazil—21(3):1465, 1727 British Columbia—21(2):665 Bromus 21(4):1997 hallii—21(4):2004 orcuttianus—21(4):2006 Burmese amber—21(4):2087 Cacti—21(3):1665 Caddo Parish (Louisiana) —21(3):1 793 Calamagrostis coahuilensis—21(1):311,312 divaricata—21(1):311,315 Calea—21(4):2023 mediterranea—21(4):2024 triantha—21(4):2027 California—21(1):193, 259,459; 21(2):673, 715, 1205; 21(3):1615 Calotis cuneifolia—21(2):1217 Calycadenia—21(1):259 micrantha—21(1):261 Calyptranthes—21(4):1955 glandulosa—21(4):1955 ishoaquinicca—21(4):1957 Camp Butner (North Carolina)— 21Q)21731 Campanulaceae—21(2):591 Canada—21(1):65;21(2):1229; 21(3):1647 Canadanthus—21(4):2141 Cardueae—21(1):207; 21(3):1645 Carduus pycnocephalus—21(2):1217 Carthamus baeticus—21(2):1217 Caryophyllaceae—21(1):1 75,237,493; 21(2):753, 887, 1165; 21(3):1669; 21(4):2439 Centaurea americana—21(2):1218 2489 2490 melitensi—21(2):1218 solstitialis—21(2):1218 Centaurium—21(1):87 blumbergianum—21(1):87 Central America—21(2):591 Cerastium pumilum—21(4):2439 (record) velutinum var. villosissimum— 21(2):887 Ceratostema oyacachiensis—21(3):1271 pendens—21(3):1272 pubescens—21(3):1275 Ceropegia palustris—21(3):1507 Chaetopappa asteroids var. asteroides— 21(2):1218 Chaptalia hintonii—21(2):929 Chevreulia sarmentosa—21(2):1218 Chihuahuana—21(1):245 purpusii—21(1):246 China—21(2):993; 21(3):1355 Chiococceae—21(3):1565 Chloridoideae—21(2):553; 21(3):1363 Chorioactis geaster—21(3):1939 Chromosome numbers—21(3):1657, 1663, 1665, 1669; 21(4):2195 Chrysothamnus—21(3):1615 scopulorum—21(3):1626 var. canonis—21(3):1627 stylosus—21(3):1627 Cirsium—21(1):207 arizonicum var. bipinnatum—21(1):209 var. chellyense—21(1):209 var. rothrockii—21(1):210 var. tenuisectum—21(1):210 clavatum var.americanum—21(1):211 var. osterhoutii— 21(1):212 cymosum var. canovirens—21(1):212 SIDA 21(4): 2490. 2005 BRIT.ORG/SIDA 21(4) eatonii—21(3):1645 var. Clokeyi—21(1):212 var.eriocephalum— 21(1):212; 21(3):1645 var. hesperium—21(1):212 var. peckii—21(1):212 var. viperinum—21(1):212 edule var. macounii—21(1):213 var. wenatchense—21(1):213 horridulum var. megacanthum— 21(1):214 inamoenum—21(1):214 var. davisii—21(1):214 occidentale var. lucianum—21(1):214 ochrocentrum var. martinii— 21(1):215 pulcherrimum var. aridum—21(1):215 scarlosum var. americanum—21(1):215 var. citrinum—21(1):215 var. coloradense—21(1):215 var. congdonii—21(1):215 var. robustum—21(1):215 var. tolyabense—21(1):216 Citharexylum—21(2):547 danirae—21(2):548 Clark County (Illinois)—21(3):1861 Clematis—21(2):861,879 brachiata—21(2):865 burgensis—21(2):867 chrysocarpa—21(2):867 dolichopoda—21(2):868 grandiflora—21(2):867 hirsuta—21(2):868 longicauda—21(2):870 sigensis—21(2):871 simensis—21(2):870 uhehensis—21(2):871 villosa—21(2):87 1 INDEX viridiflora—21(2):872 welwitschii—21(2):872 Clibadium—21(4):2023 arriagadae—21(4):2029 Clitoria mariana—21(1):507 Coahuila, Mexico—21(1):1, 11 Coker, William Chambers—21(2):977 Collin County, Texas—21(3):1939 Commelinaceae—21(3):1931 Compositae—21(1):287, 1225 Conoclinium—21(2):1641 Cutis, Peter—21(2):963 Dichanthelium scoparium—21(1):465 Diospyros torresii—21(4):2046 Dirca—21(2):511 mexicana—21(2):511 occidentalis—21(2):511 palustris—21(2):51 1 Disterigma bracteatum—21(3):1279 Dittrichia graveolens—21(2):1218 Dominican amber—21(4):2095 Dracopis amplexicaulis—21(2):1218 Duncan, Wilbur Howard (1910-2005)— 21(3):1941 Dyssodia papposa—21(2):1218 Echinochloa colona—21(4):2171 colonum—21(4):2171 Echiteae—21(1):117,159 Echites—21(1):117, 133 Ecuador—21(1):159;21(2):693; 21(3):1 269; 21(4):1955 Eleocharis mutata—21(2):1153 Eleutherococcus brachypus—21(2):993 scheuchzeri—21(2):791 senticosus—21(2):993 Emilia fosbergii—21(3):1937 Eragrostideae—21(3):1363 Eragrostidinae—21(3):1363 SIDA 21(4): 2491. 2005 2491 Eragrostis—21(3):1363 barrelieri—21(3):1371 capillaries—21(3):1374 cillanensis—21(3):1374 ciliaris —21(3):1378 curtipedicellata—21(3):1378 curvula—21(3):1 38] elliottiim21(3):1383 erosa—21(3):1385 hirsuta—2 1(3):1387 hirta—21(3):1389 hypnoides—21(3):1390 intermedia—21(3):1392 lehmanniana—21(3):1396 lugens—21(3):1397 mexicana—21(3):1398 obtusiflora—21(3):1399 palmeri—21(3):1402 pectinacea—21(3):1403 pilosa—21(3):1406 reptans—21(3):1407 secundiflora—21(3):1409 sessilispica—21(3):1410 silveana—21(3):1412 spectabilis—21(3):1413 spicata—21(3):1414 superba—21(3):1415 Eremogone—21(1):237 capillaris var. americana—21(1):239 congesta var. cephaloidea—21(1):239 var. charlestonensis—21(1):239 var. crassula—21(1):239 var. glandulifera—21(1):239 var. prolifera—21(1):239 var. simulans—21(1):239 var. subcongesta—21(1):239 var. suffrutescens—21(1):239 var. wheelerensis—21(1):240 eastwoodiae var.adenophora— 2492 BRIT.ORG/SIDA 21(4) 21(1):240 peregrinus—21(2):665 ferrisiae—21(2):754 poliospermus var. disciformis— frankliniim—21(1):240 21(1):24 var. thompsonii—21(1):240 porsildii—21(1):44 kingii pupuratus—21(1):5] var. plateauensis—21(1):240 radicatus—21(1):32 var. rosea—21(1):240 robustior—21(1):21 macradenia sparsifolius—21(1):26 var. arcuifolia—21(1):240 tracyi—21(1):36 var. ferrisiae—21(1):240 uncialis—21(1):30 var. kuschei—21(1):240 var. Conjugans—21(1):30 Ericaceae—21(3):1 269, 1607 utahensis—21(1):26 Ericameria—21(3):1557 yukonensis—21(1):24, 49 arizonica—21(3):1558 Eriocaulaceae—21(2):625 linearis—21(3):1560 Eriocaulon—21(2):625 winwardii—21(3):1562 anshiense—21(2):626 Erigeron—21(1):19, 41,473; 21(2):673 kanarense—21(2):628 acris var. kamtschaticus—21(1):37 konkanense—21(2):630 breweri—21(1):20 Eriocephalum—21(3):1645 cavernensis—21(1):30 Eriophorum clokeyi var. pinzliae—21(1):28 chamissonis—21(2):795 davisii—21(1):22 xmedium subsp. album—21(2):807 decumbens—21(1):21 xmedium subsp. medium—21(2):801 denalii—21(1):51 russeolum—21(2):791 engelmannii—21(1):22 subsp. leiocarpum—21(2):804 foliosus—21(1):19-39 subsp. russeolum—21(2):800 glacialis—21(2):665 scheuchzeri var. hirsutus—21(2):671 subsp. arcticum—21(2):810 grandiflorus—21(1):43 subsp. scheuchzeri—21(2):809 greenei—21(1):28 Erithalis—21(3):1565 hultenii—21(1):41-57 angustifolia—21(3):1582 klamathensis—21(1):20 diffusa—21(3):1584 koraginensis—21(1):46 fruticosa—21(3):1584 lackschewitzii—21(1):34 harrisiim—21(3):1588 maniopotamicus—21(2):673 odorifera—21(3):1589 muirii—21(1):47 quadrangularis—21(3):1591 nanus—21(1):473 salmeoides—21(3):1592 ochroleucus—21(1):32 vacciniifolia—21(3):1593 pallens—21(1):52 Ethnobotany—21(1):321, 345, 357, 369; parryi—21(1):34 21(4):2205, 2243, 2249 INDEX Eugenia inversa—21(3):1465 Eupatoriadelohus—21(2):901 Eupatorieae—21(2):815,901;21(3):1305, 1323, 1641 Eupatorium dubium—21(2):1218 Euphorbiaceae—21(1):79 Eurybia—21(1):459; 21(2):889 merita—21(1):459 pygmaea—21(3):1633 Eurybioid—21(3):1633 Eutrochium—21(2):901 dubium—21(2):901 fistulosum—21(2):901 maculatum—21(2):902 var. bruneri—21(2):902 var. foliosum—21(2):902 purpureum—21(2):902 var. holzingeri—21(2):902 steelei—21(2):902 Evax multicaulis—21(2):1218 prolifera—21(2):1219 Everglades National Park—21(3):1917 Exploration—21(2):963 Fabaceae—21(1):345 Facelis retusa—21(2):1219 Fagaceae—21(1):447; 21(3):1921 Festuca californica—21(3):1455 subsp. hitchcockiana—21(3):1461 subsp. parishii—21(3):1461 Fevillea—21(4):1971 subgenus Anisosperma—21(4):1993 bahiensis—21(4):1977 cordifolia—21(4):1979 moorei—21(4):1984 passiflora—21(4):1993 pedatifolia—21(4):1985 pergamentacea—21(4):1988 trilobata—21(4):1990 Flaveria trinervia—21(2):1219 SIDA 21(4): 2493. 2005 2493 Florida—21(2):879, 1081; 21(3):1917 Foresteronia affinis—21(1):168 australis—21(1):168 graciloides—21(1):168 Fort Polk (Louisiana) —21(1):419 Fungi—21(2):1019 Gaillardia pulchella var. drummondii— 21(2):1219 Galactophora angustifolia—21(4):2060 calycina—21(1):169 colellana—21(4):2062 crassifolia—2 1(4):2064 pulchella—21(4):2069 pumila—21(4):2071 schomburgkiana—21(4):2073 Galapagos—21(2):1009 Gamochaeta—21(2):717,1175 americana—21(2):736 antillana—21(2):1179,1184,1219 argentina—21(2):1219 argyrinea—21(2):718, 1182 chionesthes—21(2):725, 1183 claviceps—21(2):1180, 1184, 1219 coarctata—21(2): 711,734, 1181, 1183 pensylvanica—21(2):1183, 1219 purpurea—21(2):729, 1182 simplicicaulis—21(2):1 183, 1219 sphacilata—21(2):1182 spicata—21(2):711 stachydifolia—21(2):1180, 1184 stagnalis—21(2):1177, 1184 ustulata—21(2):724, 1182 Gentianaceae—21(1):87, 525; 21(3):1249 Gentianopsis—21(2):525 detonsa subsp. nesophila—21(2):527 virgata subsp. macounii—21(2):529 Geocarpon minimum—21(2):1165 Gerbera—21(2):935, 941 2494 Gibasis pellucida—21(3):1931 Giles County (Tennessee) —21(4):2343 Gilia—21(1):531 castellanosii—21(2):539 humillima—21(2):540 latimerii—21(2):537 Glandularia—21(3):1657 Glomeruliflorae—21(1):221 Gnaphalieae—21(1):463;21(2):711, 717, 767,777,781,1175, 1205 Gnaphalium spicatum—21(2):712 Gonolobinae—21(3):1525 Gonolobus—21(1):103 Graphephorum—21(3):1419 melicoides—21(3):1423 wolfii—21(3):1425 Greater Antilles —21(3):1525 Greene County (Pennsylvania)— 21(3):1829 Grindelia lanceolata var. lanceolata— 21(2):1219 Guarea subsessilifolia—21(4):2040 Gundlachia—21(1):243, 246 diffusa—21(1):249 riskindii—21(1):249 triantha—21(1):248 truncata—21(1):250 Gutierrezia sarothrae—21(2):1219 texana var. texana—21(2):1219 Guyana—21(3):1263 Gymnocladus dioicus—21(1):345 Hardin County (Texas)—21(3):1807 Helenieae—21(2):657 Helenium amarum var. badium—21(2):1219 elegans var.elegans—21(2):1220 microcephalum var. microcephalum— 21(2):1220 Helianthus annuus—21(2):1 220 SIDA 21(4): 2494. 2005 BRIT.ORG/SIDA 21(4) Heliomeris multiflora var. multiflora— 21(2):1220 Herrickia—21(2):889, 897 horrida—21(2):897 glauca—21(2):897 var. pulchra—21(2):897 kingii—21(2):898 var. barnebyana—21(2):898 wasatchensis—21(2):897 Heterotheca subaxillaris subsp. latifolia—21(2):759 Hexalectris—21(3):1879 spicata—21(3):1879 warnockii—21(3):1879 Himatanthus sucuuba—21(1):171 Hispaniola—21(4):2081 Homochilus—21(2):591 Hybanthus parviflorus—21(2):1209 Hydrocotyle sibthorpioides—21(4):2449 (record) Hymenoxys—21(2):657 bigelovii—21(2):659 subg. Macdougalia—21(2):659 odorata—21(2):1220 Hypochaeris brasiliensis var. tweedei—21(2):1220 glabra—21(2):1220 ICBN Clarification—21(4):2185 Idaho—21(1):175, 459 Illinois —21(3):1861 India—21(1):93; 21(2):625 Interior Highlands (U.S.A.)—21(3):1781 Introduced species (Texas) —21(4):2449 Inula bifrons—21(4):2032 Ipomoea arborescens—21(3):1283 var. pachylutea—21(3):1287 seaania—21(3):1293, 1296; 21(4):2193 lridaceae—21(4):2105 INDEX lva annua—21(2):1220 axillaris —21(2):1220 xanthifolia—21(2):1220 Juniperus ashei—21(2):1107 Kansas—21(3):1695 Kathleen M. Peterson (1948-2003)— 21(2):1239 Kentucky—21(1):345;21(2):1055, 1121; 21(3):1909 Kentucky Coffeetree—21(1):345 Kisatchie Bayou (Louisiana)—21(2):1141 Krigia—21(1):225 cespitosa var. gracilis—21(1):227 x shinnersiana—21(1):230 virginica—21(2):1220 Lacmellea aculeate—21(1):169 arborescens—21(1):169 Lactuca graminifolia—21(2):1220 Lactuceae—21(1):181, 267;21(2):715 Lamiaceae—21(2):679, 1161; 21(4):2119 Lantana camara—21(2):1009 Lathrocasis—21(1):531 Legume trees—21(2):951 Leguminosae—21(1):507 Lentibulariaceae—21(4):2249 Leucophyllum—21(1):1 coahuilensis—21(1):1 Liatris—21(3):1305 cokeri—21(2):825 series Elegantes—21(3):1312 elegantula—21(2):824 series Garberae—21(3):1316 series Graminifoliae—21(3):1317 section Graminifolium—21(3):1315 helleri—21(3):1323 series Pauciflorae—21(3):1315 section Pilifilis—21(3):1314 pilosa—21(2):815 SIDA 21(4): 2495. 2005 2495 pilosa—21(2):822 section Vorago—21(3):1312 series Virgatae—21(3):1316 virgata—21(2):824 Libertia—21(4):2105 chilensis—21(4):2110 colombiana—21(4):2107 sessiliflora—21(4):2111 triccoca—21(4):2111 Lichenized ascomycetes—21(2):637, 643 Liliaceae—21(4):2445 Llerasia—21(4):2023 macrocephala—21(4):2033 Lobelia—21(2):591 aguana—21(2):610 decurrens—21(2):612 subsp. parviflora—21(2):616 ghiesbreghtii—21(2):61 1 guerrerensis—21(2):609 heteroclita—21(2):618 laxiflora—21(2):599 subsp. angustifolia—21(2):607 Lobelioideae—21(2):591 Lorandersonia—21(3):1615, 1619 baileyi—21(3):1621 linifolia—21(3):1622 microcephala—21(3):1622 peirsonii—21(3):1623 pulchella—21(3):1624 salicina—21(3):1624 spathulata—21(3):1625 Loranthaceae—21(1):97 Louisiana—21(1):419; 21(2):963, 1141, 1149; 21(3):1793;21(4):2275 botanists —21(4):2275 botany—21(4):2275 Lyonia maritima—21(3):1507 Macdougalia—21(2):657 Machaerantherinae—21(2):889 Madre de Dios (Peru) —21(2):1 231 2496 Malvaceae—21(2):693 Mandevilla—21(3):1549 amazonica—21(3):1536 colombiana—21(3):1538 gracilis—21(3):1541 hirsuta—21(3):1553 leptophylla—21(3):1535, 1544 matogrossana—21(3):1551 symphitocarpa—21(1):170 Manettia—21(2):91 | Mangroves—21(4):2243 Matanzas National Monument (Florida)— 21(2):1081 Matelea—21(3):1515, 1519 bayatensis—21(3):1525 correlliim21(3):1525 domingensis—21(4):2081 pentactina— 21(3):1519 rhamnifolia—21(3):1515 tigrina—21(3):1525, 1532 Maxillarieae subtribe Zygopetalinae— 21(2):841 Medranoa—21(1):254 parrasana—21(1):255 Menyanthaceae—21(4):2441 (record) Mertensia virginica—21(2):771 Mesechiteae—21(3):1535, 1549 Mexico—21(1):1,11, 133, 243,311,547, 553,591, 651;21(2):679, 929: 21(3):1283, 1293, 1363; 21(4):2045 Microseris laciniata—21(1):193 subsp. detlingii—21(1):200 subsp. siskiyouensis—21(1):195 Mikania buchtienii—21(4):2425 (record) euryanthela—21(4):2425 Minuartia nuttallii var. fragilis—21(2):753 var. gracilis—21(2):753 var. gregaria—21(2):754 SIDA 21(4): 2496. 2005 BRIT.ORG/SIDA 21(4) Missouri—21(3):192] Montgomery County (Arkansas) — 21(4):2389 Muhlenbergia glabrifloris—21(1):465 Muscari comosum—21(4):2445 (record) Mutisieae—21(2):929, 935,941 Mutisioideae—21(2):1225 Myrtaceae—21(3):1335, 1465 Nassauvieae—21(2):1225 National Cancer Institute—21(4):2205 Nebraska—21(2):1205 Neonesomia—21(1):252 johnstonii—21(1):254 palmeri—21(1):253 Nestotus—21(3):1647, 1650 macleanii—21(3):1651 stenophyllus—21(3):1652 Nevada—21(1):473;21(3):1615 North America—21(1):19, 41, 103, 207, 237, 243, 275,293, 447,455,511, 525; 21(2):683, 717, 753, 767, 791, 853, 887, 903, 1153, 1209, 1215, 1225; 21(3): 1419, 1471, 1669 North Carolina—21(1):225;21(2):827, 1131;21(4):2429 (record) Norway—21(1):321;21(4):2249 Nototriche—21(2):693 ecuadoriensis—21(2):694 hartwegii—21(2):696 Jamesonii—21(2):698 phyllanthos—21(2):699 Nymphoides indica—21(4):2441 (record) Odontadenia anomala—21(1):1 70 Oklahoma—21(1):429, 475;21(2):1187, 1199; 21(4):2439 (record) Frederick Lake—21(1):429 Hackberry Flat—21(1):429 INDEX Suttle Creek—21(1):429 Tillman County—21(1):429 Washita Battlefield National Historic Site —21(2):1187 Onagraceae—21(2):651 Onagreae—21(2):651 Orchidaceae—21(1):409; 21(2):841,853; 21(3):1879, 1917 Oregon—21(1):193 Osmorhiza bipatriata—21(1):501 Otostylis—21(2):841 alba—21(2):842 brachystalix—21(2):843 lepida—21(2):846 paludosa—21(2):850 Packera—21(1):289; 21(3):1643 musiniensis—21(3):1643 streptanthifolia var. borealis— 21(1):289 subnuda var.moresbiensis— 21(1):289 Pakistan—21(1):357; 21(2):951, 1019 Palaeoanthella—21(4):2088 huangii—21(4):2088 Panama—21(3):1607 Panphalea heterophylla—21(2):1221, 1225 Parasitism—21(1):369 Paronychia chartacea var. minima— 21(2):754 Parthenium hysterophorus—21(2):1 221 Passiflora affinis—21(1):275 Passifloraceae—21(1):275 Pennsylvania—21(3):1829 Peripleura arida—21(2):1221 Persicaria meisneriana var. beyrichiana— 21(1):291 Persicaria—21(1):29] Peru—21(2):841, 1231;21(4):2321 (records) SIDA 21(4): 2497. 2005 2497 Peterson, Kathleen (1948—2003)—21(2): 1239 Pezizales—21(3):1939 Phalaris—21(1):487 arundinacea—21(1):487 primuncinatus—21(4):2096 Phragmites—21(2):683 australis—21(2):683 subsp. americanus—21(2):683, 690 Pichavaram (India) —21(4):2243 Platanthera chapmanii—21(2): 853 Pluchea—21(1):59; 21(4):2023 baccharis—21(4):2035 camphorata—21(1):59 odorata—21(1):59 var, succulenta—21(1):59 Plucheae—21(1):59 Poa—21(4):2195 Poaceae—21(1):93, 293, 311,389,455, 487,553: 21(2):683; 21(3):1 263, 1355, 1363, 1419, 1455, 1905, 1923; 21(4):1977, 2095, 2113, 2171 Polemoniaceae—21(1):531 Polygonaceae—21(1):291;21(2):1199 Ponthieva brittoniae—21(3):1917 Pooideae—21(1):311;21(3):1355, 1419 Post Oak Savanna (Texas) —21(1):399 Prenanthes—21(1):181 altissima—21(1):183 barbata—21(1):185 carrii—21(1):187 Prestonia—21(1):159 amabilis—21(1):161 boliviana—21(1):166 lagoensis—21(1):170 tomentosa—21(1):171 Pseudognaphalium—21(2):777, 1205 beneolens—21(2):782 canescens—21(2):781, 782 jaliscense—21(2):1205 2498 luteoalbum—21(1):463 microcephalum—21(2):782 roseum—21(2):1 207 saxicola—21(2):777 stramineum—21(2):1221 thermale—21(2):781 Pseudostellaria oxyphylla—21(1):175, 176 Psidium australe—21(3):1338 var. argenteum—21(3):1342 var. suffruticosum—21(3):1344 grandifolium—21(3):1335, 1346 missionum—21(3):1352 Psilactis—21(4):2141 Ptarmicoidei—21(2):755 Pteridophytes—21(4):2321 Ptilagrostis luquensis—21(3):1355, 1356 Ptilimnium—21(2):743 ahlesii—21(2):744 Puccinia lantanae—21(2):1009 Quercus acutissina—21(1):447 montana—21(3):1921 Ranunculaceae—21(2):861,879; 21(4):2429 Ranunculus ficaria—21(4):2429 Ratibida columnifera—21(2):1221 Rattlesnake Falls (Tennessee)— 21(3):1753 Rauvolfia mollis—21(1):170 Red River—21(2):963 Expedition—21(2):963 Rhodiola rosea—21(1):321 (ethnobotany) Rober Mills County (Oklahoma)— 21(2):1187 Rosaceae—21(1):65 Rotundifoliae—21(1):65 Rubiaceae—21(2):911;21(3):1565 Rudbeckia hirta L. var. angustifolia— 21(2):1221 SIDA 21(4): 2498. 2005 BRIT.ORG/SIDA 21(4) Rumex maritimus—21(2):1199 Ruppia cirrhosa—21(1):499 Ruppiaceae—21(1):499 Sabatia—21(2):1249 arkansana—21(3):1250 Sarracenia alabamensis—21(4):2169 subsp. wherryi—21(4):2169 purpurea—21(2):1149 Sarraceniaceae—21(2):1149 Sawtooth oak—21(1):447 Schoenoplectus halliim21(1):475, 1201 saximontanus—21(1):475 Scrophulariaceae—21(1):1;21(3):1927, Scutellaria petersoniae—21(2):679 Sedum wrightii—21(2):943 Seed and capsule morphology— 21(2):91 1 Senecionae—21(3):1937 Senecioneae—21(3):1643 Sericocarpus—21(3):1471 asteroides—21(3):1493 linifolius—21(3):1491 oregonensis—21(3):1497 subsp. californicus—21(3):14999 rigidus—21(3):1499 tortifolius—21(3):1495 Silene—21(3):1669 conoidea—?1(1):493 drummondii subsp. striata— 21(2):887 laciniata subsp. californica— 21(2):888 ostenfeldii—21(2):888 Siolmatra—21(4):1961 brasiliensis—21(4):1962 pentaphylla—21(4):1966 Siskiyou Mountains—21(1):193 INDEX Solanaceae—21(2):1171;21(3):1675 Solanum—21(3):1675 viar'um—21(2):1171 Solidago—21(1):221; 21(2):755 caesia var. zedia—21(1):221 curtisii var. flaccidifolia—21(1):223 subsect. Maritimae—21(2):756 subsect. Multiradiatae—21(2):760 sect. Ptarmicoidei—21(2):756 shortii—21(2):1121 subsect. Triplinerviae—21(2):757 Soliva sessilis—21(2):1221 Sonchus oleraceus—21(2):1 22] Sonora, Mexico—21(3):1 283 South America—21(2):591,643, 1209; 21(4):2105 South Carolina—21(2):1215 Southeastern United States—21(1):103; 21(2):815 Sporobolinae—21(2):553 Sporobolus—21(2):553 coahuilensis—21(1):455 airoides—21(2):558 atrovirens—21(2):561 buckleyi—21(2):563 coahuilensis—21(2):566 compositus—21(2):566 contractus—21(2):568 cryptandrus—21(2):570 flexuosus—21(2):572 giganteus—21(2):574 heterolepis—21(3):1923 indicus—21(2):576 jacquemontii—21(2):578 nealleyi—21(2):579 purpurascens—21(2):580 pyramidatus—21(2):581 spiciformis—21(2):583 virginicus—21(2):584 wrightii—21(2):586 SIDA 21(4): 2499. 2005 2499 St. Jonns County (Florida) —21(2):1081 Stachys floridana—21(4):2119 Staurochilus fasciatus—21(4):205 1 leytensis—21(4):2051 Stellaria—21(3):1669 cuspidata subsp. prostrata— 21(2):888 Stemodia coahuilensis—21(3):1935 (record) Stenotus lanuginosus var. andersonii— 21(4):2093 Stipeae—21(3):1355 Stipecoma peltigera—21(1):171 Stipinae—21(3):1355 21(1):97 acostensis—21(1):98 Stuartina hamata—21(2):1221 Sub-Saharan Africa—21(2):861 Symphyotrichinae—21(3):1633 Symphyotrichum—21(2):827,903; 21(4):2141 bahamense—21(4):2131 concolor var. devestitum—21(2):762 divaricatum—21(4):2129 expansum—21(4):2133 pygmaeum—21(3):1633, 1635 rhiannon—21(2):828 squamatum—21(4):2135 subulatum—21(4):2125, 2128 var. elongatum—21(2):907 var. ligulatum—21(2):907 var. parviflorum—21(2):907 var. squamatum—21(2):908 tenufolium—21(2):904 var. aphyllum—21(2):905 Tabernaemontana vanheurckii— 21(1):172 Tagetes minuta—21(2):1221 Tamil Nadu State (India) —21(4):2243 Tennessee—21(3):1753, 1923; Cc | | YJUULITOIIti tao 2500 21(4):2343 Tetraneurinae—21(2):657 Tetraneuris linearifolia—21(2):1221 Texas Wild Rice—21(1):389 TexasS—21(1):79, 87, 181,389, 455, 487, 493,495, 499, 501, 507; 21(2):943, 1107, 1161, 1165, 1171, 1201; 21(3):1657, 1663, 1665, 1807, 1879, 1893, 1927, 1931, 1937, 1939; 21(4):2441 (record), 2445 (record), 2449 (record) Thymelaeaceae—21(2):511 Thymophylla setifolia—21(1):287 var. greggii—21(1):287 teneuiloba—21(2):1221 Toiyabea—21(3):1647, 1652 alpina—21(3):1653 Tonestus aberrans—21(2):889 kingii—21(2):889 Trans-Pecos, lexas—21(1):79,87;455, 21(3):1657, 1665 Trilliaceae—21(3):1893 Trillium texanum—21(3):1893 Triniteurybia—21(2):889, 898 aberrans—21(2):898 Trisetum—21(3):1419 aureum—21(3):1427 cernum subsp. canescens—21(3):1430 subsp. cernum—21(3):1428 flavescens—21(3):1433 montanum—21(3):1434 orthochaetum—21(3):1436 projectum—21(3):1437 sibiricum—21(3):1438 spicatum var. pilosiglume—21(3):1442 var. spicatum—21(3):1439 Tropical soda-apple—21(2):1171 SIDA 21(4): 2500. 2005 BRIT.ORG/SIDA 21(4) Tyler County (Texas) —21(3):1807, 1927 United States—21(2):1175;21(3):1507, 1647, 1781, 1935 Uropappus lindleyi—21(2):1 222 Usnea—21(2):637,643 alata—21(2):643 stuppea—21(2):637 substerilis—21(2):63 7 sulcata—21(2):643 Vaccinieae—21(3):1269 Vaccinium—21(3):1607 almedae—21(3):1607 furfuraceum—21(3):1609 luteynii—21(3):1611 Venezuela—21(3):1599, 1675 Verbenaceae—21(3):1657 Vinca major—21(1):171 Violaceae—21(2):1209 Vittadinia sulcata—21(2):1222 Washington—21(2):665 Washita Battlefield National Historic Site (Oklahoma)—21(2):1187 West Central Louisiana—21(1):409 West Gulf Coastal Plain—21(1):399 West Indies—21(3):1519 West Virginia—21(1):481 (floristic records) Western Highland Rim Escarpment (Tennessee) —21(3):1 753 Western North America—21(2):781 Wild Orchids of North America— 21(4):2297 Wyoming—21(1):459 Xanthium spinosum—21(2):1222 strumarium var. canadense— 21(2):1222 Xylovirgata—21(1):255 pseudobaccharis—21(1):256 Yellow lady-slipper orchid—21(1):409 Zizania texana—21(1):389 Zoysieae—21(2):553 INDEX 2501 236 New NAMES AND NEW COMBINATIONS: VoLuME 21 (2004-2005) Agoseris apargioides var. maritima (E. Sheld.) G.l. Baird, comb. et stat. nov.— 21(2):716 Agoseris grandiflora var. leptophylla G1. Baird, var. nov.—21(1):267 Agoseris heterophylla var. quentinii G1. Baird, var. nov.—21(1):271 Aiphanes stergiosii M. Nino, Dorr & FW. Stauffer, sp. nov.—21(3):1600 Allotoonia J.F. Morales & J.K.Williams, gen. nov.—21(1):135 Allotoonia agglutinata Jacq.) J.F. Morales & JK. Williams, comb. nov.—21(1):139 Allotoonia caudata (Woodson) J.F. Mo- rales, comb. nov.—21(1):145 Allotoonia parviflora (Sessé & Moc.) J.F. Morales & J.K. Williams, comb. nov.— 21(1):148 Allotoonia turbinata (Woodson) J.F.Mo- rales & J.K. Williams, comb. nov.— 21(1):150 Allotoonia tuxtlensis (Standl.) J.F. Morales & JK. Williams, comb. nov.—21(1):153 Amaranthus acanthobracteatus Henrickson, sp. nov.—21(1):12 Antennaria pulcherrima subsp.eucosma (Fernald & Wiegand) R.J. Bayer, comb. nov.—21(2):768 Anthenantia texana RB. Kral, sp. nov.— 21(1):296 Arenaria | g var. saxosa (A. Gray) Zarucchi, R.L. Hartman, & Rabeler, comb. et stat. nov.—21(2):753 Artemisia arbuscula subsp. longiloba (Osterhout) Shultz, comb. nov.— 21(3):1637 Artemisia globularia subsp. lutea (Hultén) Shultz, comb.nov.—21(3):1638 SIDA 21(4): 2501. 2005 Astranthium ciliatum (Raf) Nesom,comb. nov.—21(4):2016 Aulonemia nitida Judz., sp. nov.— 21(3):1263 Bromus hallii (Hitchc.) Saarela & PM. Peter- son, comb. nov.—21(4):2004 Calamagrostis coahuilensis PM. Peterson, Soreng & Valdés-Reyna, sp. nov.— 21(1):312 Calamagrostis divaricata PM. Peterson & Soreng, sp.nov.—21(1):315 Calea mediterranea (Vell.) Pruski, comb. nov.—21(4):2024 Calea triantha (Vell.) Pruski, comb.nov.— 21(4):2027 Calycadenia micrantha R.L. Carr & GD. Carr, sp. nov.—21(1):261 Calyptranthes glandulosa M.L. Kawasaki & B. Holst, sp. nov.—21(4):1955 Calyptranthes ishoaquinicca M_L. Kawasaki & B. Holst, sp. nov.—21(4):1957 Centaurium blumbergianum BL. Turner, sp. NOV.—21(1):87 Cerastium velutinum var. villosissimum (Pennell) J.K. Morton, comb. nov.— 21(2):887 Ceratostema oyacachiensis Luteyn, sp. nov.—21(3):1271 Ceratostema pendens Luteyn, sp. nov.— 21(3):1272 Ceratostema pubescens Luteyn, sp. nov.—21(3):1275 Chihuahuana Urbatsch & R.P. Roberts, gen. nov.—21(1):245 Chihuahuana purpusii (Brandegee) Urbatsch & R.P. Roberts, comb. nov.— 21(1):246 2502 Chrysothamnus scopulorum (ME. Jones) Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1626 Chrysothamnus scopulorum § var. canonis (S.L. Welch) Urbatsch, R.P. Rob- erts & Neubig, comb. nov.—21(3):1627 Chrysothamnus stylosus (Eastwood Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1627 Cirsium arizonicum var. bipinnatum (Eastw.) DJ. Keil, comb. nov.—21(1):209 Cirsium arizonicum var. chellyense (R.J. Moore & Frankton) DJ.Keil,comb.et stat. nov.—21(1):209 Cirsium arizonicum var. rothrockii (A. Gray) D.J. Keil, comb. et stat. nov.— 21(1):210 Cirsium arizonicum var.tenuisectum D.). Keil, var. nNOV.—21(1):210 Cirsium clavatum var. americanum (A. Gray) D.J. Keil, comb. nov.—21(1):211 Cirsium clavatum var. osterhoutii (Rydb.) D.J. Keil, comb. et stat. nov. 21(1):212 Cirsium cymosum var.canovirens (Rydb.) DJ. Keil, comb. et stat. nov.—21(1):212 Cirsium eatonii var. clokeyi (S.F. Blake) DJ. Keil, comb. et stat. nov.—21(1):212 Cirsium eatonii var. eriocephalum (A. Nelson) DJ. Keil, comb. nov.—21(1):212; 21(3):1645 Cirsium eatonii var. hesperium (Eastw.) DJ. Keil, comb. et stat. nov.—21(1):212 Cirsium eatonii var. peckii (L.F. Hend.) DJ. Keil, comb. et stat. nov.—21(1):212 Cirsium eatonii var. viperinum DJ. Keil, var. nov.—21(1):212 Cirsium edule var.macounii (Greene) DJ. Keil, comb. et stat. nov.—21(1):213 Cirsium edule var.wenatchense D.J. Keil, var. NOV.—21(1):213 WH BRIT.ORG/SIDA 21(4) Cirsium horridulum var. megacanthum (Nutt.) DJ. Keil, comb. et stat. nov.— 21(1):214 Cirsium inamoenum (Greene) D.J. Keil, comb. nov.—21(1):214 Cirsium inamoenum var. davisii (Cronquist) D.J.Keil,comb.et stat. nov.— 21(1):214 Cirsium occidentale var. lucianum D.J. Keil, var. NOV.—21(1):214 Cirsium ochrocentrum var. martinii (P. Barlow-lrick) D.J. Keil, comb. et stat. nov.—21(1):215 Cirsium pulcherrimum var. aridum (R.D. Dorn) DJ. Keil, comb. et stat. nov.— 21(1):215 Cirsium scariosum var. americanum (A. Gray) D.J. Keil, comb. nov.—21(1):215 Cirsium scariosum var. citrinum (Petrak) DJ. Keil, comb. nov.—21(1):215 Cirsium scariosum var. coloradense (Rydb.) DJ. Keil, comb. et stat. nov.— 21(1):215 Cirsium scariosum var. congdonii (R.J. Moore & Frankton) D.J.Keil, comb.et stat. nov.—21(1):215 Cirsium scariosum var.robustum D.J.Keil, var.nov.—21(1):215 Cirsium scariosum var. toiyabense D_). Keil, var. NOV.—21(1):216 Citharexylum danirae Leon de la Luz & Chiang, sp. nov.—21(2):548 Clibadium arriagadae Pruski, sp. nov.— 21(4):2029 Conoclinium betonicifolium var. integrifolium (A. Gray) T.F. Patterson, comb. nov.—21(3):1641 Conyza popayanensis (Hieron.) Pruski, comb. nov.—21(4):2032 Crataegus chrysocarpa var. vernonensis INDEX J.B.Phipps & O’Kennon, var. nov.— 21(1):67 Crataegus sheila-phippsiae J.B.Phipps & O’Kennon, sp. nov.—21(1):67 Croton bigbendensis B.L. Turner, sp. nov.—21(1):79 Cuniculotinus Urbatsch, R.P. Roberts & Neubig, gen. nov.—21(3):1618 Cuniculotinus gramineus (H.M. Hall Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1619 ~~ Dahlia sublignosa (P.D. Sorensen) D.E. Saar & P.D. Sorensen, comb. & stat. nov.— 21(4):2165 Dendrocalamus stocksii (Munro) M. Ku- mar, Remesh & Unnikrishnan, comb. nov.—21(1):95 Diospyros torresii M.C. Provance & A.C. Sanders, sp. noV.—21(4):2046 Disterigma bracteatum Luteyn,sp.nov.— 21(3):1279 Eremogone capillaris var. americana (Maguire) R.L. Hartman & Rabeler,comb. nov.—21(1):239 Eremogone congesta var. cephaloidea (Rydb.) R.L. Hartman & Rabeler, comb. nov.—21(1):239 Eremogone congesta var. charlestonensis (Maguire) R.L.Hartman & Rabeler, comb. nov.—21(1):239 Eremogone congesta var. crassula (Maguire) R.L. Hartman & Rabeler,comb. nov.—21(1):239 Eremogone congesta var. glandulifera (Maguire) R.L. Hartman & Rabeler, comb nov.—21(1):239 Eremogone congesta var. prolifera (Maguire) R.L. Hartman & Rabeler,comb nov.—21(1):239 SIDA 21(4): 2503. 2005 2503 Eremogone congesta var. simulans (Maguire) R.L. Hartman & Rabeler,comb. nov.—21(1):239 Eremogone congesta var. subcongesta (S.Watson) R.L.Hartman & Rabeler,comb. nov.—21(1):239 Eremogone congesta var. suffrutescens (A. Gray) R.L. Hartman & Rabeler, comb. nov.—21(1):239 Eremogone congesta var. wheelerensis (Maguire) R.L.Hartman & Rabeler, comb nov.—21(1):240 Eremogone eastwoodiae var. adenophora (Kearney & Peebles) R.L. Hartman & Rabeler, comb. nov.— 21(1):240 Eremogone ferrisiae (Abrams) R.L. Hart- man & Rabeler, comb. nov.—21(2):754 Eremogone franklinii (Doug!. ex Hook.) R.L. Hartman & Rabeler, comb. nov.— 21(1):240 Leite ct? Pat VOI, SGI eo rN A y HEV. Peck) R.L. Hartman nov.—21(1):240 Eremogone kingii var. plateauensis (Maguire) R.L.Hartman & Rabeler,comb. nov.—21(1):240 Eremogone kingii var.rosea (Maguire) R.L. Hartman & Rabeler,comb.et stat.nov.— 21(1):240 Eremogone macradenia var. arcuifolia (Maguire) R.L. Hartman & Rabeler, comb nov.—21(1):240 Eremogone macradenia var. ferrisiae (Abrams) R.L. Hartman & Rabeler, comb. et stat. nov.—21(1):240 Eremogone macradenia var. kuschei (Eastw.) R.L. Hartman & Rabeler, comb. nov.—21(1):240 Ericameria arizonica R.P. Roberts, 1pso Rabeler, comb. 2504 Urbatsch & J. Anderson sp. nov.— 21(3):1558 Ericameria linearis (Rydberg) R.P Roberts & Urbatsch, comb. nov.—21(3):1560 Ericameria winwardii (R.D. Dorn & CH. Delmatier) R.P. Roberts & Urbatsch, stat. nov.—21(3):1562 Erigeron clokeyi var. pinzliae Nesom, var. nov.—21(1):28 Erigeron davisii (Cronq.) Nesom,comb.et Stat. NOV.—21(1):22 Erigeron glacialis var. hirsutus (Cronq. Nesom, comb. nov.—21(2):671 Erigeron greenei Nesom, nom. nov.— 21(1):28 Erigeron klamathensis (Nesom) Nesom, comb. et stat. Nov.—21(1):20 Erigeron maniopotamicus G.L.Nesom & TW. Nelson, sp. nov.—21(2):673 Erigeron poliospermus var. disciformis (Crong.) Nesom, comb. et stat. nov.— 21(1):24 Erigeron porsildii Nesom & Murray, nom. nov.—21(1):44 Erigeron robustior (Cronq.) Nesom,comb. et stat. nov.—21(1):21 Eriocaulon anshiense Punekar, Malpure & Lakshmin., sp. nov.—2.1(2):626 Eriocaulon kanarense Punekar, Watve & Lakshmin., sp. nov.—21(2):628 Cc | ec Punekar, Malpure & Lakshmin., sp. nov.—21(2):630 Eriophorum xmedium subsp. album J. Cayouette, subsp. nov.— 21(2):807 Eugenia inversa Sobral, sp. nov.— 21(3):1465 Eutrochium dubium (Willd. ex Poiret) E.E. Lamont, comb. nov.—21(2):901 Eutrochium fistulosum (Barratt) E.E. Lamont, comb. nov.—21(2):901 SIDA 21(4): 2504. 2005 BRIT.ORG/SIDA 21(4) Eutrochium maculatum (L.) E.E. Lamont, comb. nov.—21(2):902 Eutrochium maculatum var. bruneri (A. Eutrochium maculatum var. foliosum (Fernald) E.E. Lamont, comb. nov.— 21(2):902 Eutrochium purpureum (\.) £.£.Lamont, comb. nov.—21(2):902 Eutrochium purpureum var. holzingeri (Rydberg) E.E. Lamont, comb. nov.— 21(2):902 Eutrochium steelei (£.£. Lamont) E.E. Lamont, comb. nov.—21(2):902 Festuca Ww, H hK hit L: (E.B. Alexeev) S.J. Darbyshire, comb. et stat. nov.—21(3):1461 Festuca californica subsp. parishii (Piper) S.J. Darbyshire, comb. nov.—21(3):1461 Fevillea subgenus Anisosperma (Silva Manso) G.Robinson & Wunderlin, comb. et stat. nov.—21(4):1 993 Fevillea bahiensis G. Robinson & Wunderlin, sp. nov.—2.1(4):1977 f.-]1 + 1 tifall nov.—21(4):2060 Gamochaeta argyrinea Nesom, sp.nov.— 21(2):718 Gamochaeta chionesthes Nesom, sp. nov.—21(2):725 Gentianopsis detonsa subsp. nesophila (Holm) J.S. Pringle, comb. nov.— 21(2):527 Gentianopsis virgata subsp. macounii (Holm) J.S. Pringle, comb. nov.— 21(2):529 Gilia castellanosii (J.M. Porter) V.E. Grant, comb. nov.—21(2):539 |. Morales, sp INDEX Gilia humillima (Brand) A.G. Day ex V.E. Grant, comb. nov.—21(2):540 Gilia latimerii (7.L. Weese & L.A. Johnson) V.E. Grant, comb. nov.—21(2):537 Guarea subsessilifolia Al.Rodr., sp nov.— 21(4):2040 Gundlachia diffusa (Benth.) Urbatsch & R.P Roberts, comb. nov.—21(1):249 Gundlachia riskindii (B.L. Turner & Lang- ford) Urbatsch & R.P. Roberts, comb. nov.—21(1):249 Gundlachia triantha (S.F Blake) Urbatsch & R.P. Roberts, comb. nov.—21(1):248 Gundlachia truncata (GL. Nesom Urbatsch & R.P. Roberts, comb. nov.— 21(1):250 Ee Herrickia glauca (Nutt.) Brouillet, comb. nov.—21(2):897 Herrickia glauca var. pulchra (S.F. Blake Brouillet, comb. nov.—21(2):897 Herrickia kingii (D.C. Eaton) Brouillet, Urbatsch & R.P. Roberts, comb. nov.— 21(2):898 Herrickia kingii var. barnebyana (Welsh & Goodrich) Brouillet, Urbatsch & R-P. Roberts, comb. nov.— 21(2):898 Herrickia wasatchensis (M.E. Jones) Brouillet, comb. nov.—21(2):897 Heterotheca subaxillaris subsp. latifolia (Buckley) Semple, comb. et stat. nov.— 21(2):759 See Ipomoea seaania Felger & Austin, sp. nov.—21(3):1296 Krigia cespitosa var. gracilis (DC.) KL. Chambers, stat. nov.—21(1):227 Krigia x shinnersiana K.L. Chambers, hy- brid nov.—21(1):230 Leucophyllum coahuilensis Henrickson, sp.nov.—21(1):1 SIDA 21(4): 2505. 2005 2505 Liatris serics El t (Al ex Nesom, comb. et stat. nov.— 21(3):1312 Liatris series Garberae Nesom, ser.nov.— 21(3):1316 Liatris series Graminifoliae Gaiser ex Nesom, ser. nov.—21(3):1317 Liatris section Graminifolium Nesom, sect. nov.—21(3):1315 Liatris series Pauciflorae (Alexander) Gaiser ex Nesom, comb. et stat. nov.— 21(3):1315 Liatris section Pilifilis Nesom, sect. nov.— 21(3):1314 Liatris series Virgatae Nesom, ser. nov.— 21(3):1316 Liatris section Vorago Nesom, sect.nov.— 21(3):1312 Llerasia macrocephala (Rusby) Pruski, comb. nov.—2.1(4):2033 Lobelia decurrens subsp. parviflora Lam- mers, subsp. Nov.—21(2):616 Lorandersonia Urbatsch, R.P. Roberts & Neubig, gen. nov.—21(3):1619 Lorandersonia baileyi (Wooton & Standley) Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1621 Lorandersonia_ linifolia (Greene) Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1622 Lorandersonia microcephala (Cronquist) Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1622 Lorandersonia peirsonii (D.D. Keck) Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1623 Lorandersonia pulchella (A. Gray) Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1624 Lorandersonia salicina (S.F. Blake) 2506 Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1624 Lorandersonia spathulata (L.C. Ander- son) Urbatsch, R.P. Roberts & Neubig, comb. nov.—21(3):1625 Mandevilla amazonica J.F. Morales, sp. nov.—21(3):1536 Mandevilla colombiana J.F. Morales, sp. nov.—21(3):1538 Mandevilla gracilis (Kunth) J.F. Morales, comb. nov.—21(3):1541 ).F. Morales, sp nov.—21(3):1551 Matelea domingensis (Alain) Krings, comb. nov.—21(4):2081 Matelea pentactina Krings, sp. nov.— 21(3):1519 Matelea rhamnifolia (Griseb.) Krings, comb. nov.—21(3):1515 Medranoa Urbatsch & Roberts, gen.nov.— 21(1):254 Medranoa parrasana (SF. Blake) Urbatsch & R.P Roberts, comb. nov.—21(1):255 Microseris laciniata subsp. detlingjii KL. Chambers, subsp. nov.—21(1):200 Microseris laciniata subsp. siskiyouensis K.L. Chambers, subsp. nov.—21(1):195 Minuartia nuttallii var. fragilis (Maquire & A.H.Holmgren) Rabeler & R.L.Hartman, comb. et stat. nov.—21(2):753 Minuartia nuttallii var. gracilis (B.L. Rob.) Rabeler & R.L. Hartman, comb. nov.— 2253 Minuartia nuttallii var. gregaria (A. Heller) Rabeler & R.L. Hartman, comb. et stat. nov.—21(2):754 Neonesomia Urbatsch & R.P. Roberts, gen nov.—21(1):252 Neonesomia johnstonii (G.L. Nesom) SIDA 21(4): 2506. 2005 BRIT.ORG/SIDA 21(4 => Urbatsch & R.P. Roberts, comb. nov.— 21(1):254 Neonesomia palmeri (A. Gray) Urbatsch & R.P. Roberts, comb. nov.—21(1):253 Nestotus R.P. Roberts, Urbatsch & Neubig, gen. nov.—21(3):1650 Nestotus macleanii (Brandegee) R.P.Rob- erts, Urbatsch & Neubig, comb. nov.— 21(3):1651 Nestotus stenophyllus (A. Gray in Torrey) R.P. Roberts, Urbatsch & Neubig, comb. nov.—21(3):1652 — Packera musiniensis (S.L. Welsh) Trock, comb. nov.—21(3):1643 Packera streptanthifolia var. borealis (Torr. & A. Gray) D.K. Trock, comb. nov.— 21(1):289 Packera subnuda var. moresbiensis (J.A. Calder & RL. Taylor) D.K. Trock, comb. nov.—21(1):289 Palaeoanthella Poinar & Chambers, gen. nov.—21(4):2088 Palaeoanthella huangii Poinar & Cham- bers, sp. nov.—21(4):2088 Paronychia chartacea var. minima (L.C. Anderson) R.L. Hartman, comb. et stat. nov.—21(2):754 Persicaria meisneriana var. beyrichiana (Cham. & Schltdl.) C.C. Freeman, comb. nov.—21(1):291 Pharus primuncinatus Judz. & Poinar, sp. nov.—21(4):2096 Phragmites australis subsp.americanus Saltonstall, PM.Peterson & Soreng, subsp nov.—21(2):690 Pluchea baccharis (Mill.) Pruski, comb. nov.—21(4):2035 Prenanthes carrii J.R. Singhurst, RJ. O’Kennon, & W.C. Holmes, sp. nov.— 21(1):187 INDEX Prestonia amabilis J.F. Morales, sp.nov. 21(1):161 Prestonia boliviana J.F. Morales & A. Fuentes, sp. nov.—21(1):166 Pseudognaphalium saxicola (Fassett H.E.Ballard & Feller, comb. nov.— 21(2):777 Pseudognaphalium thermale (E.E. Nel- son) Nesom, comb. nov.—21(2):781 Pseudostellaria oxyphylla (8.L. Rob.) RL. Hartman & Rabeler, comb. nov.— 21(1):176 Psidium australe var. argenteum (O.Berg) Landrum, comb. nov.—21(3):1342 Psidium australe var. suffruticosum (O. Berg) Landrum, comb.nov.—21(3):1344 Ptilagrostis luquensis P.M. Peterson, Soreng & Z.L.Wu, sp. nov.—21(3):1356 Ptilimnium ahlesii Weakley & Nesom, sp. nov.—21(2):744 sr Sabatia arkansana J.S. Pringle & CT. Witsell, sp. nov.—21(3):1250 Sarracenia alabamensis F.W. Case & R.B. Case, sp. Nov.—21(4):2169 Sarracenia alabamensis subsp. wherryi FW. Case & R.B. Case, subsp. nov.— 21(4):2169 Scutellaria petersoniae B.L. Turner & J.L. Reveal, sp. nov.—21(2):679 Silene drummondii subsp. striata Rydb. J.K. Morton, comb. et stat. nov.— 21(2):887 Silene laciniata subsp. californica (Durand) J.K. Morton, comb. et stat. nov.—21(2):888 Silene ostenfeldii (AE. Porsild) J.K. Morton, comb. nov.—21(2):888 Solidago caesia var. zedia R.E. Cook & Semple, var. nov.—21(1):221 Solidago curtisii var. flaccidifolia (Small) SIDA 21(4): 2507. 2005 2507 R.E.Cook & Semple, comb. et stat. nov.— 21(1):223 Solidago subsect. Multiradiatae Semple, subsect. nov.—21(2):760 Solidago sect. Ptarmicoidei (House) Semple & Gandhi, comb. nov.— 21(2):756 Staurochilus leytensis (Ames) E.A. Christenson, comb. nov.—21(4):2051 Stellaria cuspidata subsp. prostrata (Baldw. ex Ell.) J.K. Morton, comb. et stat. nov.—21(2):888 Stenotus lanuginosus var. andersonii C.A.Morse, comb. nov.—21(4):2093 Struthanthus acostensis L.A. Gonzalez & |.F. Morales, sp. nov.—2.1(1):98 Symphyotrichum — concolor devestitum (S.F. Blake) Semple, comb. nov.—21(2):762 Symphyotrichum pygmaeum (Lind!.) Brouillet & S. Selliah, comb. nov.— 21(3):1635 Symphyotrichum rhiannon Weakley & Govus, Sp. NOV.—21(2):828 Symphyotrichum subulatum svar. elongatum (Boss.) S.D. Sundb., comb. nov.—21(2):907 Symphyotrichum subulatum ligulatum (Shinners) S.D.Sundb.,comb. nov.—21(2):907 Symphyotrichum subulatum = var. parviflorum (Nees) S.D. Sundb., comb. nov.—21(2):907 Symphyotrichum subulatum squamatum (Spreng.) S.D. Sundb,, comb. nov.—21(2):908 Symphyotrichum tenuifolium var. aphyllum (R.W.Long) S.D.Sundb.,comb. nov.—21(2):905 Var. Var. Var. Thymophylla setifolia var. greggii (A. 2508 Gray) Strother, comb. nov.—21(1):287 Toiyabea R.P. Roberts, Urbatsch & Neubig, gen. nov.—21(3):1652 Toiyabea alpina (L.C. Anderson & S. Goo- drich) R.P. Roberts, Urbatsch & Neubig, comb. nov.—21(3):1653 Triniteurybia Brouillet, Urbatsch & RP. Rob- erts, gen. Nov.—21(2):898 Triniteurybia aberrans (A. Nelson) Brouillet, Urbatsch & R.P. Roberts, comb. nov.—21(2):898 <— SIDA 21(4): 2508. 2005 BRIT.ORG/SIDA 21(4) Vaccinium almedae Wilbur & Luteyn, sp. nov.—21(3):1607 Vaccinium furfuraceum Wilbur & Luteyn, sp. nov.—21(3):1609 Vaccinium luteynii Wilbur, sp. nov.— 21(3):1611 Xylovirgata Urbatsch & R.P. Roberts, gen. nov.—21(1):255 Xylovirgata pseudobaccharis (5.F. Blake) Urbatsch & R.P. Roberts, comb. nov.— 21(1):256 GUIDELINES FOR CONTRIBUTORS 2509 SIDA GUIDELINES FOR CONTRIBUTORS Sida home page: http://www brit.org/sida/ Guidelines: me //wwwbrit.org/Sida/SubmitPaper.htm Sida, Contributi IssN 0036-1488) is an Leseeal cactiny sian ie systematic botany con- taining primary h par lato, incl distics, ecology, Hosea - netics & evolution _numerical Eaxonomy, ea, and neces Sida i 1s open to alla I restricted toan COPTd hical area. ¥ Bf0bTAp All manuscripts eeubmined to Sida are jeonsidered by at ee st two revi ipts may be submitted in English or Spanish. P rp t may be waived or reduced under certain circumstances. = Please contact ae editor and male arrangements before :eeevik Manu- lime aad scripts are not f financial support. Submit juesti to oe ee Botanical Shas Institute of Texas, 509 Pecan Street, Fort Worth, TEXAS 76102-4060, U.S.A. Phones: 1-817-332-4441 voice; 1-817-332-4112 fax. Our electronic mail address is sida@brit.org. Access Sida a page at www/brit.org/sida Submissions: Please submit manuscripts in electronic format as an email attachment. If wish submissions are also accepted by USPS mail with a single hard copy and on computer disk. a may be submitted in low-resolution format (Gjpgs) as email attachments for re- view purposes only. Upon acceptance for publication, the manuscript es be submitted as an email SS you ee or returned in the mail with a single hard copy and an py. If you have fig- ures/illustrations in eecucns eet they ss uld be submutteg? in tif format, eee at the abso- lute best quality possible l lave to | isk if ee are too large for email. Typically b/w line vdeamines should be scanned at 600 opie or oe and b/w half d 1 5 tones and color photos at 300 dpi. If you need your , the editorial staff at Sida will be happy handle any electronic ajiaeation needs you have; all ign material can be returned if requested. Color digitization works best from color transparencies (slides). Consult the latest issue of Sida for format of articles and notes regarding title, author, and ad- dress. — of numbers are separated oy lala hyphens (6--8) Of use an en dash (-). Footnotes, figure legends, appendices, and dof manuscript following relerenc th d to use the APG farailice nearly all of which are on / can ie accessed through the Missouri Botanical Garden research site (Angiosperm cee Abstract Every paper should include both an English and Spanish, or another major language abstract. Specimen Citations Label information should be followed for itati tever language. Follow whatever unit if measure is provided on the label (metric or English). Use ihe collector's last name (or full name) and number or (s.n) and italicize. Abbreviate the months of the year by the first three letters of the mene paren a qa DenOd: ane Texas. Jeff Davis Co. | mi W of Fort Davis, 3 Jan 1972, Smith 118 (BRIT) da Agua de Rega, 23 km N of Seabra, road to Agua de Rega, ca. 1000 m, 24 Feb 1971 (fr), Irwin et al. 30894 (HB, MBM, MO, NY). Author Citations Use the author iati iven inthe Author Abbreviati iled at the H Royal Botanic Gardens, Kew (1992). Use et or the ampersand (&) between two authors; for more than two SIDA 21(4): 2509. 2005 2510 BRIT.ORG/SIDA 21(4) authors, restrict to the first one followed by et al. Do not italicize or underline in the manuscript the terms et, et al., ex, or in. Plant names in the text should include authorship or clearly state the botanical nomenclature being followed. Latin Descriptions or Diagnoses; English Descriptions Please provide a Latin di is for each new taxon. AL tere Latin diagnosis, an English or Spanish version is recommended ie hlighting the di ters. Write plant descriptions as incom- plete sentences with phrases separated by seaicclods Figures Please present ALL fi ( included) ified, singl f figures. A scale should be in or on the ‘llusteation or hora reduction will dutomadcally reduce the scale and subject proportionately. In halftones, sharp glossy photegray ne with good contrast are necessary for good reproduction. Please mount illustrative material (halftones, line drawings, etc.) on flexible paper so that may be mounted and ona high-speed drum scanner if need. You may also submit graphic mate- rials in electronic format. Typically b/w line drawings should be scanned at 600 dpi or higher and b/w half tones and color lee at 300 dpi. Please consult a editor for additional information if need. Color figures can be used and an additional c .Color digitization works best from color transparencies a Please submit eo prints for reviewing purposes. References This includes all of the literature cited in the text and may include other article citations the author may deem desirable. Normal text references should be cited as follows: ‘Ricketson and Pipoly (1997) stated... or ‘the latest revision (Ricketson & Pipoly 1997) when reference is used as authority for a statement. When there are three or more authors use only the name of the first author followed by et al. ‘Barrie et al, (1992a) stated’. References at the end of the article are ar ee apne betically and chronologically making use of a,b, etc. if an author 7 more than one publication in a given year. Author’s names are printed in Cap/sMALL cars, NO L CAPS; please type mae names in Cap/ SMALL CAP or in Cap/lower case except for initial capita s. Only the first letter of the initial word, es should be capitalized. Please note there is no space be- ane proper nouns, and proper adjectives of tit tween initials of authors. For journal abbreviations, use Botanico-Periodicum-Huntianum. Reference examples: Barrie, F.R., CE. Jarvis, and J.L. Reveal. 1992a. The need to change Article 8.3 of the Code. Taxon 41:508—512 Ricketson,J.and J. Pipoly. 1997.A synopsis of the genus Gentlea (Myrsinaceae) and a key to the genera of Myrsinaceae in Mesoamerica. Sida 17:697—707. [author's names in cap/lower case] Crawrorp, D.J.1983.Phylogenetic and systematic inferences from electrophoretic studies. In: S.D. Tanksley and T.J. Orton, eds. lsozymes in plant genetics and breeding, Part A. Elsevier, Amsterdam. Pp. 257—287. [author's names in cap/small cap] Sivinski, R.C. and K. LicHtroor (eds.). 1995. Inventory of rare and endangered plants of New Mexico. New Mexico Forestry and Resources Conservation Division. Energy, Minerals and Natural Resources Department. Misc. Pub. No. 4. SIDA 21(4): 2510. 2005 GUIDELINES FOR CONTRIBUTORS 2511 Abbreviations when the elolowing pene are used the period is omitted. Distance: mm, dm, cm, m, km, ft, Ap W, months: first 3 letters only, Jan, Feb, etc. Example: Potter Co.: 2 km Wo f Dot, 5 Jun 1971, ic: (SMU). Documented Plant Chromosome Numbers Refer to Sida 21(3):1663. 2005 Acknowledgments ] » eee poe | | | . ] aes: J . qe Cdata ill your ee das UNIS Recomm eneatD concie ya | . y: . - : ep might | their own aseitcen or on granting agencies. In a time where herbaria are Pere and even dy- ing, let’s not take them for granted. Floristic Papers Floristic papers are important, but they need to be more than a list of plants, they need added value, they need meat and context added. Such papers should emphasize the uniqueness of the floristic study area, physical characteristics, cultural and historical information, land use ses climate, | e records, major vegetational Lhabitat s, rare/enda ngered/special concern taxa, Tr ou disjuncts, invasives, etc. Lists in and of themselves are not that ieee ieee your ie as much as pooies The more annotations you can add to the checklist the more value it will have. A list that | | notes on rare and endangered taxa or other noteworthy taxa, State and Federal [ oO us on any rare and endangered species, and highlights and notes any endemics, disjuncts, or si nificant geograpnic est anOn TeCOIeS etc., is so nee more vena ang us use ful. : onservation 5 V al Ue to the paper. Also note which and how many species are state-listed by the Heritage Program if they exist. Summarize how many taxa occur in various classes of county numbers, which is informative for conservation of the state’s flora. Range Extensions Range extension papers will ONLY be considered for publication when the taxon being reported represents a new state record significant disjunct, doc ented exotic /invasive, rare and/or endan- gered or is a rare endemic. Questions: Email the editor (Barney Lipscomb, barney @brit.org). SIDA 21(4): 2511. 2005 - oa his photographic guide to the Wild Flowers of . a WILD FLOWERS OF MOMBACHO no is color-coded and arranged by Family NICARAGUA — Mombac within the color sections. It covers flowering plants found aks eS pe Nee TS on the whole of the Volcano; which includes both tropical cloud forest and dry, deciduous forest, at lower altitudes. The book covers 200 plants, each of which has two or three photographs and a short description in both English and Spanish giving common name(s); a short description o “oe of the plant; height; flowering period; habitat and any known use of the plant. The vast majority of plants are likely to be found in similar habitats throughout tropical My HELEN PLCKERING America and nearly one third are pan tropical, making this Ghia SO book of interest to a wide audience throughout the tropics. WILD FLOWERS OF MOMBACHO NICARAGUA Sida, Bot. Misc. No 28, 2006 issn 0823-1475 FLORES SILVESTRES DEL MOMBACHO isbn 1-889878-14-6 BY HELEN PICKERING 5° x 8.5", x + 217pp. over 500 photographs @ $15 + p&p’ “International: $7 BRIT PRESS Texas residents add $1.24 sales tax he Atlas of Texas covers about 6000 taxa. This is the result of 54 years of herbarium and fieldwork by BL. Turner, beginning in 1948 at Sul Ross State University, Alpine, Texas. In short, the senior author has examined Atlas of the scular personally, touched, or “pored over” an estimated several hundred thousand sheets in the preparation of the forth- coming Atlas volumes. Contents include an introduction, atlas of Texas ane arranged alphabetically by family, by genus, by species, and an index ah Available from Botanical Research Institute of Texas Press 9 Pecan Street Atlas of the Fort Worth, TX 76102-4060, USA Vascular Plants of Texas E-mail: sida@brit.org * Fax 1-817-332-4112 www.brit.org/Sida/Sida/24.htm Sida, Bot. Misc. No. 24, 2003, issn 0833-14 Holly Niohale Gaerey Denny, isbn (vol. 1) 1-889878-08-1 + isbn (vol. 2) 1-889878- — X= 7 1/2" < 10" dad aren Vi Vol. 1 $50 + p&h* «Vol. 2 oa + pal «Set + p&h* ) *International: $12.50 Kala) 1) *$11.50 jee $25 (set) = Texas residents add sales tax: $4.13 (vol. 1) « $3.30 (vol. 2) = $7.43 (set) pares UISIANA TEXAS GEORGIA ee AM | a a ARKANSAS FLORIDA IOWA Mis: DA KOTA "Nt YOMING JONA HAWAII neon UTA VASt oN SOUTH \KOTA | OY) = MC CUE / ? RK : XGINIA MINNESOTA CANADA ao ALBERTA BRITISH: RRITORY NO RTHWEST TERRITORIES NOVA a scom WANITOBA oe WFOUND AND “MANATOBA Q yack 1S ME RIC A PERU CHILE B ly TSINUCEN SuAANA . ENTINA ECUADOR BELIZE coud ENCH ¢ JRUGUAY SI JRINAME , ™ xy ee at eu Sa: 3 5 DE a aS a oe Eee ee ge See = SOLMIA GUAT EMALA SAN SALV, DNDURAS. GUYANA Cc CE NT CARAGUA PANAMA COSTA JBA PUERTO’ RICO. DOMIN REA ASIA ie ne DIA BURMA BANGLADESH thes ALAY JAMAICA NORTH i & Ss RACRUZ 2 MExicx QO DALGO GUADALAJARA - COAHI LA YUC CATAN “SIN ISSN 0036-1488