if PHYTOLOGIA An international journal to expedite plant systematic, phyltogeographical fn dnd ecological publication eek LIBRARY CONTENTS NESOM, G.L., Two new species of Penstemon subg. Habroanthus Scroohularianede front MeKicO~...>sv. Senos) fopanen na pede leone lode wea ld ecco. 333 ~NESOM, G.L., A new species of Leucophyllum (Scrophulariaceae) from ET te OS eM R ce A.C peta Moen Seep Rete ca a nha ta lad Ee nN 3437 . TURNER, B.L., An overview of the North American species of Menodora OMG islet heehee Stace aii ret eS noc dectns ecto teins sThia oes koe vd gat Coeovtied 340 _NESOM, G.L., A new species of Nama (Hydrophyllaceae) from north- Cy ay THLE beta ert ep eR RRBAON.9' gD RL REE gta NRE 357 _ JORDAN, R., Putative natural hybrid of Eupatorium capillifolium x E. glaucescens from Hardin County, Texas .............ccececececeeeeeeeeeeees 360 _ - LEMKE, D.E., The genus Solanum (Solanaceae) in Texas.............00006. 362 . _BROWN, L.E. & J. SCHULTZ, Arthraxon hispidus (Poaceae), new to MOL el ua tt AE Rela a ody ath metabo te Mat wishinn suion Rea pcbiwe esi vaaicasigoawtee 379 “LUTHER, H.E., Two new species of Aechmea (Bromeliaceae)............. 382 _JONES, S.D., G.D. JONES, & J.K. WIPFF, Kosteletzkya depressa, section KRasteleizkya(MalwaGede), THO WetO-LeXAS 1 iota. paviln psiscvee eho Uren ae 387 _SORENG, RJ., Notes on new infraspecific taxa and hybrids in North Fe wel ia Be FGM aro Mg Loy. Val ei” Cech AIMDIENC aA OND a A a ho NR ieee 390 Contents continued on the inside cover. Published by Michael j. ‘Warnock 185 Westridge Drive Huntsville, Texas 77340 U.S.A. PHYTOLOGIA is printed on acid free paper. (Contents continued) y _.NESOM, G.L. & M.A. BAKER, First report of #rigeron velutipes — (Asteraceae) fromthe United States «0.00.2. 5. ae 414 — _ NESOM, G.L., A new species of Erigeron (Asteraceae: Astereae) from — northwestern New MeXiCO:...0....0.:..30 1k ees 416 > ERTTER, B., New combinations in Potentilla and Horkelia (Rosaceae) in California ey ie A ets en ee 420 Books received. eC eee 423 Corrections: and additions 0.0 oye oe eee 428 2 . Ae Se PHYTOLOGIA (ISSN 00319430) is published monthly with two volumes per year by Michael J. Warnock, 185 Westridge Drive, Huntsville, TX 77340. Second Class postage at Huntsville, TX. Copyright ©1991 by PHYTOLOGIA. Annual domestic individual subscription (12 issues): $36.00. Annual domestic institutional subscription (12 issues): $40.00. Foreign and/or airmail postage extra. Single copy (previous to volume 67), $3.00 (add $.50 per copy postage and handlin [$1.00 per copy foreign]). Back issue sales by volume: $17.00 per volume 4 (not all available as complete volumes); $21.00 per volume 67-present; add | per volume postage US ($4.00 per volume foreign). POSTMASTER: Send ad changes to Phytologia, 185 Westridge Drive, Huntsville, TX 77340. Phytologia (November 1991) 71(5):333-336. TWO NEW SPECIES OF PENSTEMON SUBG. HABROANTHUS (SCROPHULARIACEAE) FROM MEXICO Guy L. Nesom Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT Two new species of Penstemon subg. Habroanthus are described from México: P. luteus (yellow flowered) from Coahuila and P. gal- loensis (blue flowered) from Nuevo Leon. KEY WORDS: Penstemon subg. Habroanthus, Scrophulariaceae, México Study of specimens recently collected in northeastern México by the Hinton family has revealed the existence of two previously undescribed species of Pen- stemon. The morphology of the anther sacs (dehiscing from the apices inward, usually leaving the central portion indehiscent) in both of these species places them in Penstemon subg. Habroanthus Crosswh. (Crosswhite 1967; Holmgren 1984). The taxonomic position of each species within the subgenus is discussed following its description. Penstemon luteus Nesom, sp. nov. TYPE: MEXICO. Coahuila. Mpio. Arteaga, Sierra E] Coahuilon, scattered in pine woods, 3100 m, 17 Jun 1991, Hinton et al. 21009 (HOLOTYPE: TEX!). Ex affinitate Penstemoni subg. Habroanthi Crosswh., imprimis proprius floribus tubularibus luteisque et inflorescentia floribus nu- merosis in pedunculis ac pedicellis longis. Herbs 0.6 m tall; stems and leaves completely glabrous, eglandular, slightly glaucous. Leaves opposite, linear-lanceolate, entire, epetiolate, the uppermost (below the inflorescence) 6-8 cm long, 5-10 mm wide. Inflorescence a thyrse of ca. 20 verticillasters, not secund, the cymes 3-7 flowered on peduncles 4-6 cm long and pedicels 10-15 mm long; sepals green, lightly striate, 6-7 mm long, ovate-acuminate with entire margins; corollas yellow (as noted on the collector’s label), drying yellow, apparently without other coloration, glabrous 333 334 PHY TOLOGLIA volume 71(5):333-336 November 1991 outside, the palate sparsely villous with long, yellowish hairs, tubular, 25-35 mm long, gradually expanded, slightly if at all ventricose, ca. 8-10 mm wide at the throat, apparently without ventral grooves, strongly bilabiate, the lower lobes 7-8 mm long and strongly reflexed, the upper lobes 4-5 mm long and projecting past the lower; fertile stamens nearly reaching the throat but not exserted, the anther sacs glabrous, widely divergent, 3 mm long from tip to tip, dehiscent from the distal ends, leaving the proximal portion indehiscent and often slightly gibbous, the suture margins minutely sharp papillate; staminode glabrous, ca. half the length of the fertile stamens. Mature fruits not seen. Known only from the type collection, represented by a specimen with the inflorescence and upper portion (45 cm long) of a stem of a single plant. All previously described species of subg. Habroanthus produce blue to red flowers. There is no other yellow flowered species of Penstemon in México, and such are rare within the genus over its entire range of geography and morphology. Even if the flowers of P. luteus were some other color, however, it still could not be identified as any previously known species. Besides P. luteus, P. barbatus (Cav.) Roth is the only other species of subg. Habroanthus that occurs in the high sierra of southeastern Coahuila. Plants of P. barbatus are generally smaller and produce flowers with red corollas in few flowered inflorescences on thinner stems, exserted stamens, and staminodes as long as the fertile stamens. Penstemon subg. Habroanthus has been divided into two sections, based on flower color and morphology: sect. Habroanthus (= sect. Glabri [Rydb.| Pennell) produces ventricose, blue to violet flowers; sect. Elmigera (Reichb.) Benth. produces tubular, gradually ampliate, red flowers. These differences apparently are correlated with hummingbird pollination in sect. Elmigera vs. wasp pollination in sect. Habroanthus (Crosswhite 1967). Penstemon luteus represents a third mode of variation within subg. Habroanthus, and although its yellow, tubular flowers are probably hummingbird pollinated, it justifiably could be accorded sectional status on a phenetic basis, since flower color is the primary distinction between the two previously recognized sections. It is not clear, however, that the relatively few species of sect. Elmigera represent a monophyletic group (Crosswhite 1965, 1976), and P. luteus may itself have been derived from a red flowered ancestor. Penstemon barbatus (the type of sect. Elmigera) hybridizes with the blue flowered species P. laevis Pennell, P. speciosus Dougl. ez Lindl., and P. letophyllus Pennell of sect. Habroanthus as well as with P. palmeri A. Gray of subg. Penstemon (Holmgren 1984). Penstemon sect. Elmigera, which has not been the subject of a study or even a recent taxonomic summary (not even by Crosswhite 1965 or 1967), comprises eight species: P. barbatus, P. wislizeni (A. Gray) Straw, P. imberbis ( Willd.) Poir., P. labrosus (A. Gray) Hook., P. eatonit A. Gray, P. cardinalis Woot. & Standl., P. regalis A. Nels. (or P. cardinalis subsp. regalis [A. Nels.] Nisbet & Jackson), and P. henricksoni Straw. Penstemon barbatus occurs from central Nesom: Two new Penstemon from México 335 México northward to Coahuila and Texas and to Durango, Chihuahua, and the southwestern United States; P. wislizeni, P. imberbis, and P. henricksonu are endemic to México (Straw 1959, 1976); P. labrosus occurs in California and Baja California Norte, and the last three species are restricted to the southwestern United States. Apart from the two species described in the present paper, all the species of sect. Habroanthus are restricted to the western United States. Penstemon galloensis Nesom, sp. nov. TYPE: MEXICO. Nuevo Leon. Mpio. Galeana, Cerro El Gallo, oak woods, 2100 m, 16 Aug 1987, Hinton et al. 19166 (HOLOTYPE: TEX!). Penstemoni barbato (Cav.) Roth similis sed differt floribus azure- is, sepalis ovatis marginibus denticulatis, antherarum sacculis pagi- nis dense minute papillatis, et staminodio apice papillato longitu- dine ca. 2/3 staminum partes aequanti. Perennial herbs ca. 0.6 m tall; stems and leaves completely glabrous, eg- landular. Leaves opposite, the cauline linear-lanceolate, 7-10 cm long 2.5-4.0 mm wide, the basal oblanceolate to spatulate, 10-12 cm long, the blades 10-22 mm wide. Inflorescence a thyrse of ca. 8 verticillasters, apparently not secund, the cymes 2-3 flowered on peduncles 2-4 cm long and pedicels 1-2 cm long; sepals greenish, glabrous, ovate-deltate-acuminate with denticulate-serrulate margins, lightly striate, the lobes 6-7 mm long; corollas purple, ventrally whitish, with dark purple stripes running onto the lower lip, glabrous out- side, lightly bearded on the palate and proximal portions of the lower lobes, 27-30 mm long, tubular, slightly ventricose, apparently without prominent ventral grooves, ca. 5-6 mm wide at the throat, strongly bilabiate, the upper lobes 7-8 mm long, the lower lobes 6-7 mm long and strongly reflexed; fertile stamens definitely exserted from the corolla tube, the anther sacs densely and minutely papillate, widely divergent, 4 mm long from tip to tip, dehiscent from the distal ends, leaving the proximal third of each sac indehiscent and often slightly gibbous, the suture margins smooth; staminode distinctly papillate at the apex, ca. 2/3 the length of the fertile stamens. Mature fruits not seen. Known only from the type collection. Penstemon galloensis is similar in its vegetative morphology and strongly bilabiate corollas to P. barbatus, but the sepals of the latter are triangular, the corollas red, the anther sacs smooth, and the staminode equaling the length of the fertile stamens, without a papillate apex. In Crosswhite’s taxonomic arrangement of subg. Habroanthus (1967), P. galloensis would be placed with other blue flowered species in sect. Habroanthus series Habroanthus. The other species of this group are restricted to the western United States, and the evo- lutionary relationship of P. galloensis to these is not clear. The new species is compared here in the Latin diagnosis to P. barbatus because the latter is 336 PHY FOL GIA volume 71(5):333-336 November 1991 the only putatively related species, besides P. luteus, that grows in northeast- ern México. In a key that includes the majority of the related taxa of sect. Habroanthus in the western United States (Holmgren 1984), P. galloensis runs with difficulty to the vicinity of P. speciosa Dougl. ez Lindl., the most widely distributed species of the section, and P. laevis Pennell. These are perhaps closely related to the new species, but both differ from it in their more com- pact inflorescences, flowers with broader throats, and smooth anther sacs that are sigmoidally twisted. ACKNOWLEDGMENTS I thank Dr. B.L. Turner, Dr. Andrew McDonald, and Dr. T.P. Ramamoor- thy for their review and comments on the manuscript. LITERATURE CITED Crosswhite, F.S. 1965. Hybridization of Penstemon barbatus (Section Elmi- gera) with species in section Habroanthus. Southw. Naturalist 10:234- 237. 1967. Revision of Penstemon section Habroanthus (Scrophular- iaceae). Amer. Midl. Naturalist 77:1-41 (I: Conspectus, pp. 1-11; II: Series Speciosi, pp. 12-27; III: Series Virgati, pp. 28-41). . 1970. Revision of Penstemon (Section Leptostemon) Series Race- most. Amer. Midl. Naturalist 84:237-243. - Holmgren, N. 1984. Penstemon. Intermountain Fl. 4:370-455. Straw, R.M. 1959. Los Penstemon de México. I. Sobre la confusion entre Penstemon lanceolatus y Penstemon imberbis. Bol. Soc. Bot. Méx. 24:39- 52. . 1976. A new species of Penstemon (Scrophulariaceae) from Mexico. Madrono 23:263-265. Phytologia (November 1991) 71(5):337-339. A NEW SPECIES OF LEUCOPHYLLUM (SCROPHULARIACEAE) FROM NUEVO LEON, MEXICO Guy L. Nesom Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT Leucophyllum hintoniorum sp. nov. is described from eastern Nuevo Leén, México, near Aramberri. It is distinctive in its obovate- oblanceolate, concolorous leaves densely invested with nearly sessile dendritic trichomes, uniseriate glandular trichomes extending above the remainder of the vestiture, and purple flowers with yellow spots and eciliate corolla lobes. KEY WORDS: Leucophyllum, Scrophulariaceae, México Study of specimens recently collected in Nuevo Leon by the Hinton fam- ily has revealed the existence of a remarkably distinctive and previously un- described species of Leucophyllum. The genus is known to harbor narrow endemics (Henrickson & Flyr 1985), and it is not surprising to discover yet another. Leucophyllum hintoniorum Nesom, sp. nov. TYPE: MEXICO. Nuevo Leon. Mpio. Aramberri, E of Aramberri, gypsum hillside, 1300 m, 14 May 1991, Hinton et al. 20929 (HOLOTYPE: TEX!; Isotype: MEXU!). Leucophyllo fiyru B. Turner similis foliis concoloris obovati- oblanceolatisque, calycis lobis stipitati-glandulosis, et corollis glan- dulosis purpureisque sed differt foliis longioribus, vestimento tri- chomatibus brevi-stipitatis dendriticisque, pedicellis ac calycum vestimento pariter confertis, et corollis sparsim minute glandulosis flavi-maculatis in tubo inferno. Alternately branched shrubs ca. 1 m tall, apparently not thorny. Silvery vestiture completely obscuring the stem, leaf, and calyx surfaces, of a dense layer of relatively uniform dendritic trichomes on short stipes (less than 0.1 337 338 PHY POO GIA volume 71(5):337-339 November 1991 mm high), with fewer, conspicuous, barely stipitate dendritic trichomes with much longer and thicker radii projecting above the basal layer; young stems, leaves, and calyces densely invested with prominent stipitate glands on unis- eriate stipes 0.3-0.5 mm long and extending well above the remainder of the vestiture, these apparently persistent only on the calyces. Leaves alternate, entire, obovate-oblanceolate, broadest in the distal one-fourth, 20-35 mm long, 5-11 mm wide at the broadest point, obtuse at the apex, gradually tapered to the base, without a clear petiole. Flowers with tomentose pedicels 2-3 mm long; calyces 6.0-6.5 mm long, lobes lanceolate, 5 mm long, 1.2 mm wide, densely invested with dendritic hairs and stipitate-glandular trichomes, sparsely glan- dular inside but without dendritic hairs; corollas purple, with yellow spots on the lower tube, 16-20 mm long, tube broadly ampliate, to 7 mm wide at the throat (pressed), sparsely pilose on inside floor with crinkled hairs 1-2 mm long, sparsely stipitate-glandular on the outside, the lobes obovate, subequal, 6-7 mm long and wide, emarginate, eciliate, nearly glabrous inside; stamens 4, anthers and filaments glabrous; styles 10-12 mm long, sparsely pilose; ovaries glandular at the apex, otherwise glabrate. Mature capsules not seen. Known only from the type collection. Leucophyllum hintoniorum is similar to L. flyri B. Turner (to which it will key in Henrickson & Flyr 1985) in its concolorous, obovate-oblanceolate leaves without a clear petiole, persistently stipitate-glandular calyx lobes, the glands extending well above the other vestiture, and purple corollas glandular on the outside. The new species differs from L. flyra in its longer leaves (20-35 mm long vs. 10-27 mm long), distinctive vestiture of barely stipitate dendritic hairs (vs. tall dendritic hairs), pedicels and calyx with equally dense vestiture (vs. more dense on the pedicels), and corollas sparsely and minutely glandular (vs. relatively densely and long stipitate glandular) on the outside, with yellow (vs. purple) spots on the lower tube. As noted by Henrickson & Flyr (1985), the relationships of L. flyrii are obscure, and although it is compared here with L. hintoniorum because of their distinctive stipitate glandularity that does not occur elsewhere in the genus, the two species do not appear to be closely related. Three other species occur in the same area of southern Nuevo Leén as Leu- cophyllum hintoniorum: L. frutescens (Berland.) 1.M. Johnston, L. pruinosum I.M. Johnston, L. revolutum Rzed., and L. zygophyllum 1.M. Johnston. All of these are immediately different from L. hintoniorum in leaf morphology as well as other features. The new species is clearly different from the widespread and variable L. frutescens, which has lavender to pink corollas with ciliate lobes and distinctly bicolored leaves with a vestiture of dendritic trichomes with relatively longer axes, lacking stipitate glandular trichomes. Nesom: New Leucophyllum from Nuevo Leon 339 ACKNOWLEDGMENTS ‘I thank Dr. B.L. Turner and Dr. T.P. Ramamoorthy for their review and comments on the manuscript. LITERATURE CITED Henrickson, J. & L.D. Flyr. 1985. Systematics of Leucophyllum and Ere- mogeton (Scrophulariaceae). Sida 11:107-172. Phytologia (November 1991) 71(5):340-356. AN OVERVIEW OF THE NORTH AMERICAN SPECIES OF MENODORA (OLEACEAE) B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A systematic overview of the North American species of Menodora is presented, including a key to the fourteen species recognized. Distri- bution maps are provided for all species, including varieties. Four new species are proposed, all of which are restricted to México: M. hinto- niorum B. Turner, a gypsophile from Nuevo Leon; M. jaliscana B. Turner, from Jalisco and Michoacan; M. potosiensis Henrickson ez B. Turner, from San Luis Potosi; and M. tehuacana B. Turner, from Puebla and Oaxaca. One new combination is proposed: M. magni- flora (Steyerm.) B. Turner. KEY WORDS: Menodora, Oleaceae, México Steyermark (1932) provided a revision of the genus Menodora as part of the requirements for a Masters Degree program at Washington University, St. Louis. In this, he recognized seventeen species: ten occurring in North America; five in South America; and two in Africa. Steyermark’s presentation was an excellent one for its time, especially since there were relatively few collections from North and South America in the fifteen herbaria from which he obtained loans. Unfortunately, his study was not based upon knowledge of populational variation in the field; additionally, he failed to provide carefully constructed distributional maps for the taxa - concerned. The present overview of the North American species has been prepared so as to provide a backdrop for the description of the several new taxa proposed here. My personal experience with populational variation in the field has been largely with the Texas species, namely Menodora heterophylla Moric. er A. DC., M. longiflora Engelm. ez A. Gray, and M. scabra Engelm. ez A. Gray. Limited though this was, it proved critical in my interpretation of the 340 Turner: Overview of North American Menodora 341 intraspecific categories proposed by Steyermark and, by extrapolation, my evaluation of specific categories in general. The distributional maps included here are based upon approximately 1000 specimens from the following herbaria (numbers shown in parenthesis): F (64); GH (239); LL (200); MEXU (101); MO (60); TEX (386). KEY TO NORTH AMERICAN SPECIES OF MENODORA 1. Fruiting pedicels to some extent recurved or arcuate. ................ (2) be eEriiting, pedicelsastitily exact. ct ae: sek cesses ee, tn. a oe eee amet, o\= (9) 2.(1) Leaves deeply lobed, mostly 1-3 cm wide. ...... M. heterophylla 2. lueaves' mostly entire, 01-10 crm Wide. Woes oc ale «iene Delain ero ale (3) 3.(2) Stems deeply sulcate, at least some of the lower leaves 3 parted; central Qaeda Sad noentedim ition N sake cess eee ete eee aaa M. mezicana 3. Stems rounded, not deeply sulcate, the leaves entire; Oaxaca to Nuevo LeOt.” die web ecicme Cos x nee Maanin nae de stee tes ae aieiee Deleciideieisoutesme (4) 4.(3) Leaves linear lanceolate to oblong lanceolate, mostly 1-3 mm wide, (4-)5-7 times as lorig asiwide. c.c5 a.com tae stewie nase sv aiiweae obits (5) 4. Leaves mostly ovate to elliptical oblong; 4-12 mm wide; 3-4(-5) times AG HONG ASUWAGE nc snc da data a Ave amar Ger Maale pra cama abana niet (6) 5.(4) Stems stiffly erect, 15-40 cm high, intricately branched below; leaves opposite throughout; Nuevo Leon, gypseous soils. ..... M. hintoniorum 5. Stems spreading from the base, 3-15 cm high, the stems not intricately branched, leaves mostly alternate (rarely not); widespread. M. coulteri 6.(4) Stems hirtellous-scabrous or scabrous-puberulent, the hairs mostly somewhat reflexed and only 0.10-0.15 mm long. .............. (7) 6. Stems hirsute-pilose with mostly spreading hairs 0.2-0.5 mm lOnperatire: Pas eet re ey Se eee eee ne acres ee centre (8) 7.(6) Suffruticose spreading herbs 5-20 cm high; leaf blades minutely but markedly glandular punctate above and below (under magnification); Oaxaca tol\Guanajuaton. ®. qeaqet. is he ster veut. secs 3. S. sisymbrifolium 6. Leaves more or less entire to merely lobed. .....4. S. campechiense 7. Lower anther 6 mm or more long; corolla more than 1.5 cm across; stigma entire or only. very shehtlycapitate=s seein coon: ac ankle. peas (8) 7. Lower anther 5 mm or less long; corolla less than 1.5 cm across; stigma (CEN EES at 3 ABER OTe ee ae ere AP Rie PAN on AES 6. S. heterodorum 8. Perennials from a woody or corky base; seeds plump. 9. S. tenuzpes 8: Taprooted) annuals; seeds flattened. ....:...7<-4 lease ose (9) 9. Lower anther 6-8 mm long; corolla to 1.7 cm across. .....8. S. davisense 364 PHYET OL :O'G: LA volume 71(5):362-378 November 1991 9. Lower anther 10 mm or more long; corolla more than 2 cm across. ..... 2: sated Maiaicnereausths cus, Scaiteay? sietaraia-s 2 amet tne 7. S. citrullifolium 10. Pubescence of lower leaf surface and stems all or chiefly composed of stellate or dendritic hairs. .. 2. -cm101).4 oe eee (11) 10. Pubescence of the lower leaf surface and stems composed entirely of simple hairs or occasionally lacking. .-..5=..5-s9eeeeeeens (15) 1). Herbagée unarmed... 0i:s0.s20- 00 Sep oo 2 (12) It. Herbage prickly. 20... secs: neis oss ees gereaels oe pe eer (13) 12. Leaves ovate to ovate elliptic, densely tomentose with stellate hairs; inflorescence terminal”... ....:..'. 2.0+oe (eee eae 10. S. erranthum 12. Leaves narrowly elliptic to oblanceolate, moderately pubescent with dendritic hairs; inflorescence lateral. .......... 11. S. capsicastrum 13. Stems and lower leaf surface silvery canescent due to the dense pubescence. nb Sepia ato aa ema tus ne eters ae pn nt 2. S. elaeagnifolium 13. Stems and lower leaf surface not as above, the typically tawny pubescence NOt SO GENSE. so a. ne seca » daisies wis eiba ore noose le (14) 14. The 4 to 8 rayed stellate hairs sessile on lower leaf surface; fruit 1-2 EM 1 GlaMEtOK 6). ca.36 00.0 Gop een cae 12. S. carolinense 14. At least some of the usually 8 or more rayed stellate hairs stipitate on lower leaf surface; fruit 2.5-3.0 cmin diameter. 13. S. dimizdiatum 15. Leaves.odd pinnate. oi... ds0- smae.2n asin sacle eee (16) U5. Leaves simples 05)... 5 biys 2 acaja afee, avers cia: vm oleuaceua Shenae See (18) 16. Pubescence of young stems glandular; corolla stellate, white; leaflets linear oblong to oblong lanceolate; pseudostipular leaves similar to the regular leaves. 2.5.0.2 060.0 > o0cee eee 14. S. jamesu 16. Pubescence not glandular; corolla rotate to rotate stellate, light purple or rarely white; leaflets ovate to ovate elliptic or obovate; pseudostipular- leaves semiovate), ......:.).25..62. sees (17) 17. Plant essentially glabrous; interstitial leaflets minute or lacking. ....... DOE Sy bay Hear sat a APOE EERE jE a OR 13a 15. S. leptosepalum 17. Plant more or less strigose; interstitial leaflets usually prominent. ..... Pe eee eo nies Cures se eT ee 16. S. fendleri 18> Leaves deeply. pinnatafid. ., «..255.55ecune oop ee 17. S. triflorum Lemke: Solanum in Texas 365 18. Leaves entire or merely sinuate, sometimes hastately lobed. ..(19) 19. Plants woody perennials, erect or scandent shrubs or subshrubs; leaves typicallyobloug or hastately ovate! .. 2.00.5. iia roe eee ees (20) 19. Plants annual or perennial herbs; leaves typically ovate. ........... (21) 20. Leaves broadest at or above the middle, tapered at base. ........ Bis 7's a nicer ndnalage SO Mis ccc ee 2 eee 18. S. pseudocapsicum 20. Leaves broadest below the middle, not tapered at base. ......... SeCaieRs wii apap kas 6a sid jul snows soos in sts RE ete See 19. S. triquetrum 21. Anthers 2.6-4.9 mm long; style usually exserted at least 1 mm beyond the anthers; strong petennials. .4. 25) -2aaveuensnu eee s oe 20. S. douglasi 21. Anthers 1.2-2.6 mm long; style barely, if at all, exserted beyond the anthers; annuals or short lived perennial herbs. .................. (22) 22. Calyx greatly enlarged in fruit, enveloping at least half of the berry; fruit brownish green at maturity. ..,.......... 21. S. sarrachoides 22. Calyx not enlarged and enveloping the berry; fruit purplish black Bt FRAUOTIEY., 2 oon. es cat snote aeiege e ob ace ee eee meee (23) 23. Seeds mostly > 1.8 mm long; calyx lobes subequal in fruit. ........... Raa Bees ks ee peste seem iias Sean y ob op aae ie eu wees 22. S. interius 23. Seeds mostly < 1.8 mm long; calyx lobes markedly unequal in fruit. MPS cio Bh Sia: RGN psa Sa wai 4e Me Vath ee ao Kees 23. S. ptycanthum 1. Solanum capsicoides All. Perennial herbs, occasionally woody at the base, to 1.2 m tall, the stems pubescent with minute glandular capitate hairs and very sparsely villous with long jointed hairs, provided throughout with straight, broad based stramineous prickles to 10 mm long. Leaves ovate in outline, the blades repand to pinnately lobed, obtuse at the apex, slightly cordate at the base, 5-15 cm long, 3-12 cm wide; petiole to 5 cm long. Inflorescence short, few flowered. Calyx 5-10 mm long, the triangular-ovate lobes 3-4 mm long, prickly. Corolla white, 10-15 mm long, 12-25 mm in diameter. Stamens subequal, the anthers 3-5 mm long. Fruit scarlet, 20-50 mm in diameter. Seeds orbicular-ovoid, flat, 4.0-5.5 mm long, reticulate. S. aculeatissemum auct., non Jacq. Solanum capsicoides is a member of Solanum sect. Acanthophora, a group of somewhat weedy species with a center of distribution in southeastern South 366 PHYS OL OG TA volume 71(5):362-378 November 1991 America. The species is apparently native to coastal areas of eastern Brazil (Nee 1979), but has become successfully established in open, lowland habitats in many tropical and subtropical areas of the world, including the Atlantic and Gulf coastal plains of the U.S. The only Texas specimens seen are early 20th century collections from Chambers County. 2. Solanum elaeagnifolium Cav. Rhizomatous perennial herbs to 1 m tall, often woody at the base, the stems and branches densely covered with many rayed stellate hairs and sparsely to copiously armed with small acicular prickles to 3 mm long. Leaves narrowly lanceolate to oblong, entire to sinuate or undulate, obtuse at the apex, at the base rounded to tapering, 3-10(-15) cm long, 0.5-2 cm wide, with a few scattered acicular prickles on the midvein below; petioles to 2 cm long. Inflo- rescence 3 to 7 flowered. Calyx 5 angled, the triangular-ovate, attenuate lobes 6-8 mm long. Corolla blue to violet or rarely white, 20-35 mm in diameter. Anthers equal, 6-9 mm long. Fruit yellowish but turning black with age, 10-15 mm in diameter. Seeds brown, shining, ovoid or oblong, 3-5 mm long, nearly smooth. Chromosome number, n = 12. S. terense Engelm. & A. Gray, S. roemerianum Scheele. Solanum elaeagnifolium is widespread throughout much of the southcentral U.S., from Kansas and Missouri south to Louisiana, Texas, Arizona, and ad- jacent México. It is our most abundant nightshade, occurring in waste places, disturbed soils, and open areas throughout most of the state. The uncommon white flowered plants are referred to f. albsflorum Cockll. 3. Solanum sisymbrifolium Lam. Annual herbs to about 0.8 m tall, the stems villous-pubescent with a mix- ture of glandular capitate and stellate hairs, armed with straight prickles to 6 mm long. Leaves ovate, the blades to 16 cm long, deeply pinnatifid, the lobes sinuate or again pinnatifid, pubescent above and below with a mixture of glan- dular capitate and stellate hairs; petioles to 5 cm long. Inflorescence terminal, soon becoming lateral, 3 to 5 flowered. Calyx 10-15 mm long, divided into 5 lanceolate to ovate lanceolate lobes 7-10 mm long. Corolla and anthers not seen. Fruit red, very loosely and completely or incompletely enclosed by the prickly calyx, 15-20 mm in diameter. Seeds yellowish brown, ovoid, flattened, 1.5-2.0 mm long, minutely reticulate. Solanum sisymbriifolium is a native of Brazil that is occasionally adventive or escaped from cultivation in much of the eastern U.S. In Texas it is known from a 1927 collection from Wood County. Lemke: Solanum in Texas 367 4. Solanum campechiense L. Annual herbs to 0.6 m tall, the stems spreading, greenish, with large stipi- tate and much smaller sessile stellate hairs, armed with thin walled, yellowish prickles to 20 mm long. Leaves oblong to ovate, the blades more or less en- tire to somewhat 5 or rarely 7 lobed, the lobes coarsely dentate, obtuse at the apex, cordate basally, 5-12 cm long, 6-7 cm wide, densely pubescent. with sessile stellate hairs and prickly along the veins above and below; petioles to 5 cm long. Inflorescence few-flowered. Calyx 6-8 mm long, the lobes ovate- lanceolate, acute, 3-5 mm long, accrescent to the fruit at maturity. Corolla pale violet, 15-20 mm in diameter. Stamens subequal, the anthers 4-5 mm long. Fruit red, 10-15 mm in diameter. Seeds yellowish brown, ovoid, plump, 2.0-2.3 mm long, dorsally tuberculate. Solanum campechiense is a native of México and the West Indies that has been occasionally collected from fields and disturbed ground in brushlands in the Rio Grande Valley (Cameron, Hidalgo, and Zapata counties) with a disjunct collection from Nueces County. 5. Solanum rostratum Dun. Spreading, taprooted annual herbs to 0.7 m tall, the stems stellate pubescent and densely beset with straight, sometimes broad based, prickles to 8 mm long. Leaves ovate to broadly ovate in outline, the blades 7-16 cm long, once or twice pinnatifid, the ultimate lobes rounded or obtuse, stellate pubescent above and below, and prickly along the main veins below; petioles to 10 cm long. In- florescence 7 to 12 flowered. Calyx 6-13 mm long with linear lobes 6-10 mm long, nearly hidden by the dense covering of stellate hairs and prickles. Corolla yellow, 25-35 mm in diameter. Anthers unequal, the four uppermost yellow, 6-8 mm long, the lowermost suffused with purple, arcuate, 10-14 mm long. Fruit 9-12 mm in diameter, tightly invested by the prickly, accrescent calyx tube. Seeds dark brown, ovoid, flattened, 2.0-2.6 mm long, minutely foveolate. Chromosome number, n = 12. Solanum rostratum is an aggressive, weedy species native to the south- central U.S. from Nebraska to Texas and south into México, but it is widely adventive elsewhere. It is frequently found in disturbed soils and waste places throughout the state. 6. Solanum heterodorum Dun. Spreading annual herbs from a slender taproot, to about 0.7 m tall, the stems with scattered glandular capitate hairs and yellow prickles to 8 mm long. Leaves broadly ovate to deltoid in outline, 4-11 cm long, twice pinnatifid, the 368 PHY DTOLOG ELA volume 71(5):362-378 November 1991 ultimate lobes obtuse or rounded, above with a mixture of glandular and eglandular hairs, below with scattered stellate and short stipitate glandular hairs, prickly along the main veins below; petioles to 5 cm long. Inflorescence 5 to 9 flowered. Calyx 10 mm long with lanceolate lobes 7-8 mm long. Corolla violet or blue, 10-17 mm in diameter. Anthers unequal, the four uppermost yellow, 2.5-4.0 mm long, the lowermost purple tinged and about 1 mm longer than the upper anthers. Stigma distinctly capitate. Fruit 9-12 mm in diameter, tightly invested by the prickly, accrescent calyx tube. Seeds dark brown, ovoid, flattened, 2.5-2.9 mm long, minutely foveolate. Chromosome number, n = 12. Whalen (1979) recognized three varieties in Solanum heterodorum: vat. heterodozum, var. novomezicanum Bartl., and var. setigeroides M.D. Whalen. Only var. setzgeroides is known from Texas, having been collected from El Paso, Hudspeth, and Culberson counties. Correll & Johnston (1970) included Solanum heterodozum in S. citrulli- folium A. Braun. The two species are quite distinct, however, being easily distinguished by corolla size (<1.5 cm diam. in S. heterodorum versus >2 cm diam. in S. cttrullifolium) and stigma morphology (capitate in S. heterodorum versus entire in S. citrullifolium). 7. Solanum citrullifolium A. Braun Much branched and spreading taprooted annual herbs to about 0.7 m tall, the stems and branches pubescent with a mixture of simple glandular capitate hairs and a few longer uniseriate hairs, armed with straight yellow prickles 2-7 mm long. Leaves broadly ovate in outline, irregularly bipinnatifid and usually pinnatisect near the base, the lobes obtuse or rounded, 4-10 cm long, glandular pubescent, occasionally with a few long uniseriate hairs above and scattered, stellate hairs below; petioles to 5 cm long. Inflorescence 6 to 10 flowered. Calyx 4-5 mm long with linear-lanceolate lobes 2-3 mm long. Corolla violet or blue, 25-35 mm in diameter. Stamens unequal, the 4 uppermost with yellow anthers 6-10 mm long, the lowermost anther arcuate, violet-tinged terminally, 11-16 mm long. Stigma entire. Fruit 8.5-11.5 mm in diameter, closely invested by the prickly, accrescent calyx tube. Seeds ovoid, flattened, dark brown, 2.3-2.9 mm long. Chromosome number, n = 12. Whalen (1979) recognized three varieties in Solanum citrullifoluum, two of which occur in Texas. Var. setigerum Bartlett is found on various substrates in the Trans-Pecos, and in eastern Chihuahua and western Coahuila; it is characterized by stems that are densely covered with spreading or retrorse acicular prickles. Var. cztrullifolium, characterized by stems with scattered subulate prickles, is found primarily on igneous soils in the Trans-Pecos and adjacent México, with disjunct populations in the Llano Uplift area. Although Whalen reported var. citrullifolium to be very scarce in central Texas, having Lemke: Solanum in Texas 369 been collected only twice since 1930, the last time in 1949, I found the plants to be locally common in disturbed roadside habitats in Burnet County in 1989. 8. Solanum davisense M.D. Whalen Taprooted annual herbs 0.4-0.7 m tall, the stems and branches viscid pubescent with a mixture of simple glandular capitate hairs and fine spread- ing uniseriate hairs, armed with straight yellow prickles 3-7 mm long. Leaves broadly ovate in outline, 2 to 3 times pinnatifid, the ultimate lobes narrowly deltoid to lanceolate, 5-10 cm long, prickly along the main veins, weakly strigu- lose above, below with few-rayed stellate and scattered short stipitate glandu- lar hairs; petioles to 3 cm long. Inflorescence 5 to 9 flowered. Calyx 4-5 mm long with linear-lanceolate lobes 2-3 mm long. Corolla violet or blue, 13-20 mm in diameter. Anthers unequal, the 4 uppermost yellow, 4.0-5.5 mm long, the lowermost arcuate, violet tinged at the apex, 5.5-8.5 mm long. Stigma weakly capitate. Fruit 8-10 mm in diameter, closely invested by the prickly, accrescent calyx tube. Seeds ovoid, flattened, dark brown, 2.6-3.0 mm long. Chromosome number, n = 12. Solanum davisense occurs sporadically on igneous soils and in sandy or gravelly streambeds, mostly above 1500 m, in the Davis, Chisos, and Chinati mountains of Trans-Pecos Texas, and in the Sierra del Carmen of adjacent Coahuila, México. 9. Solanum tenuipes Bartlett Spreading perennial herbs from woody or corky roots and rhizomes, 20- 70 cm tall, the stems viscid pubescent with a mixture of glandular capitate and spreading uniseriate hairs or occasionally glabrate, armed with widely spaced pale yellow prickles 2-8 mm long. Leaves broadly ovate, to 15 cm long, 2 to 3 times pinnatifid, the ultimate lobes usually rounded or obtuse at the apex, glandular capitate and often with scattered uniseriate eglandular hairs above, below with abundant short stipitate glandular capitate and few to numerous stellate hairs, prickly along the main veins below; petioles to 6 cm long. Inflorescence 6 to 9 flowered. Calyx 5-8 mm long with lanceolate to linear lobes 3-5 mm long. Corolla blue or violet, 25-35 mm in diameter. Anthers unequal, the 4 uppermost yellow, 7-10 mm long, the lowermost tinged with purple apically, arcuate, 12-20 mm long. Fruit wholly enclosed by the prickly accrescent calyx tube or nearly so, 7-10 mm in diameter. Seeds dark brown, reniform, 2.7-3.6 mm long, minutely foveolate. Chromosome number, ripest Be 370 PHYTOLOGIA volume 71(5):362-378 November 1991 Solanum tenuzpes is similar in aspect to S. citrullifolum, but can be easily distinguished by its perennial habit, smaller fruits (7-10 versus 8.5-11.5 mm in diameter), and larger seeds (2.7-3.6 versus 2.3-2.9 mm long). The species is found on calcareous or gypseous soils in open desert or semidesert in west Texas and northern Mexico. Whalen (1979) recognized two varieties in Solanum tenuipes, both of which occur in Texas. In var. tenuzpes the largest leaves are thrice pinnatifid and the seeds 3.1-3.6 mm in length; it is distributed from eastern Coahuila north into Brewster, Terrell, Val Verde, and Maverick counties. Var. latisectum M.D. Whalen has leaves only twice pinnatifid and seeds 2.7-3.1 mm long; it is distributed primarily in northern México (Chihuahua, Coahuila, Durango) and adjacent Texas (Presidio, Brewster, Terrell, Crockett, and Maverick counties). 10. Solanum erianthum D. Don Shrubs or small trees to 3 m tall, the young stems velvety tomentose throughout with a dense stellate indumentum. Leaves ovate to ovate elliptic or elliptic, entire, acute to acuminate at the apex, rounded at the base, 5-20 cm long, 2-12 cm wide, densely tomentose with stellate hairs below, less densely vestitured above; petioles to 8.5 cm long. Inflorescence usually terminal, many flowered. Calyx 6-8 mm long, tomentose, with ovate lobes 2.5-3.5 mm long. Corolla white, 15-18 mm in diameter, the lobes ciliate. Anthers equal, 2-3 mm long. Fruit yellow, 10-20 mm in diameter. Seeds yellowish brown, ovoid, flattened, 1.5-2.0 mm long, minutely reticulate. S. verbascifolium auct. Solanum erianthumis distinctive among the Texas species of Solanum by its habit and densely tomentose, almost felty leaves. The species is a pantropical weed, widely distributed in the Old and New World. In Texas, it is known from several collections in thickets along the margin of the Resaca del Rancho Viejo in Cameron County and from a single collection in loose sand at the edge of live oak mottes in Kenedy County. 11. Solanum capsicastrum Link Small shrubs to about 0.7 m tall, the stems densely pubescent with small dendritic hairs. Leaves narrowly elliptic to oblanceolate, entire to sinuate or undulate, revolute, obtuse or rounded at the apex, tapering at the base, 1-7 cm long, 0.5-1.5 cm wide, pubescent above and below with short dendritic hairs; petioles to 1 cm long. Flowers solitary or 2-3 in lateral inflorescences. Calyx 6-7 mm long, the lanceolate lobes 3-4 mm long. Corolla white, to 15 mm in diameter. Fruit orange-red or scarlet, 8-10 mm in diameter. Seeds orange-brown, flattened, 2.5-3.5 mm long, minutely reticulate. a Lemke: Solanum in Texas 371 Solanum capsicastrum, the false Jerusalem cherry, is a native of Brazil that occasionally escapes from cultivation. In Texas the species has been collected from several localities along the Gulf coastal plain (Harris, Colorado, and Victoria counties). 12. Solanum carolinense L. Rhizomatous perennial herbs to approximately 1 m tall, pubescent through- out with 4 to 8 rayed sessile stellate hairs and armed with stout yellow prickles to4 mm long. Leaves ovate elliptic to ovate, shallowly lobed or toothed, obtuse to acute at the apex, rounded or cuneate at the base, 3-12 cm long, 1.5-5.0 cm wide, prickly along the veins below; petioles to 1.5 cm long. Inflorescence 3 to 8 flowered. Calyx 5-7 mm long, the lanceolate-acuminate lobes 2.0-3.5 mm long. Corolla pale violet to occasionally white, 20-30 mm in diameter. Anthers equal, 6-8 mm long. Fruit yellow, 10-20 mm in diameter. Seeds yellowish-brown, orbicular-ovate, flattened, 2.2-2.6 mm long, minutely rugose. Solanum carolinense occurs throughout the eastern U.S. as far west as Minnesota and Texas. In Texas, the species is found primarily on sandy soils of the Piney Woods region, as far west as Grayson, Anderson, and Brazoria counties. The white flowered plants are known as f. albiflorum Benke. 13. Solanum dimidiatum Raf. Rhizomatous perennial herbs to about 1 m tall, the stems and branches densely vestitured with 8 to 12 rayed, often stipitate, stellate hairs and spar- ingly armed with small stout prickles to 5 mm long. Leaf blades ovate, sinu- ately 5 to 7 lobed, entire to undulate, rounded at the apex, at the base rounded to truncate or cordate, 6-15 cm long, 5-10 cm wide, both surfaces densely beset with 8 to 12 rayed sessile and stipitate stellate hairs, the midrib below pro- vided with short subulate prickles or these sometimes wanting; petioles stout, to 3 cm long. Inflorescence 4 to 20 flowered. Calyx 7-10 mm long, the 5 or 6 triangular-ovate, acuminate lobes 3-5 mm long. Corolla bluish purple to vio- let, rarely white, 30-50 mm in diameter. Anthers equal, 8-12 mm long. Fruit pale yellow, 25-30 mm in diameter. Seeds light brown, oval, 3.8-4.3 mm long, minutely rugose. Chromosome number, n = 12. S. torrey: A. Gray. Solanum dimidiatumn is a prairie species of the southcentral U.S. (Kansas and Missouri south to Texas). It is similar in general aspect to S. carolinense, but differs in the presence of stipitate, as well as sessile, steliate hairs on the leaf blades, larger anthers (8-12 mm versus 6-8 mm long), and a larger fruit (2.5-3.0 cm versus 1-2 cm in diameter). Although their ranges in Texas overlap 372 PHY TOLO GAA volume 71(5):362-378 November 1991 slightly, S. dimzdiatum is primarily a species of prairie soils in the central] part of the state, while S. carolinense is restricted to the sandier soils of east Texas. The white flowered individuals have been treated as f. album (Waterfall) Correll. 14. Solanum jamesi Torr. Stoloniferous and tuber bearing perennial herbs, occasionally becoming suffrutescent, usually to about 0.3 m tall, rarely to 0.5 m, the stems with a mixture of coarse flattened uniseriate hairs and much smaller glandular capi- tate hairs. Tubers often numerous, globose to ellipsoid, white or light brown, 5-20 mm in diameter. Leaves odd pinnate, to 15 cm long, only rarely with small interstitial leaflets, the rachis somewhat winged; leaflets 7-11, linear oblong to lanceolate, to 6 cm long and 2 cm wide but usually much smaller, strigose with numerous flattened uniseriate hairs and scattered glandular hairs above and below. Inflorescence few flowered, the pedicels distinctly articulate at or above the middle. Calyx 3.5-8.0 mm long, irregularly lobed for about half its length. Corolla white, 15-25 mm in diameter. Anthers equal, 3-4 mm long. Style well exserted beyond the anthers. Fruit purplish black, 8-9 mm in diameter. Seeds not seen. Solanum jamesii is found at elevations of 1400 m and above in the south- western U.S. (Arizona, Utah, Colorado, New Mexico, Texas) and adjacent Mexico. In Texas the species is relatively rare in the mountains of the Trans- Pecos (El Paso, Hudspeth, Culberson, and Jeff Davis counties). 15. Solanum leptosepalum Correll Stoloniferous perennial herbs, occasionally becoming suffrutescent, to 0.5 m tall, the stems sparsely appressed strigose throughout or glabrous. Leavés odd pinnate, to 20 cm long, sparsely pubescent at the margins, the interstitial leaflets absent or minute; leaflets 5(7), ovate to ovate-elliptic, the lowermost pair much reduced, to 7 cm long and 3.5 cm wide. Inflorescence few flowered, the pedicels articulate at or well above the middle. Calyx 6-8 mm long, the ovate-lanceolate long acuminate lobes 3-5 mm long. Corolla purplish to white, 20-25 mm in diameter. Anthers equal, 6-8 mm long. Style well exserted beyond the anthers. Fruit not seen, reported by Correll (1952) to be about 15 mm in diameter. Seeds not seen. Solanum leptosepalum was described by Correll (1952) from a single col- lection and has since been sporadically collected from the mountains of Trans- Pecos Texas and adjacent Coahuila, México. I am not convinced that the taxon is specifically distinct from the widespread and variable Mexican species S. verrucosum Schlecht. Lemke: Solanum in Texas 373 16. Solanum fendler: A. Gray Stoloniferous and tuber bearing perennial herbs, occasionally becoming suffrutescent, to about 0.5 m tall, the stems more or less strigose. Tubers globose to ellipsoid, white or suffused with purple, to 3 cm long. Leaves odd pinnate, to 10 cm long, with or without small interstitial leaflets; principal leaflets 5-9, ovate to oblong elliptic or obovate, at the apex obtuse to acute, to 8 cm long and 3.5 cm wide. Inflorescence 3 to many flowered, the pedicels distinctly articulate well above the middle. Calyx 4-6 mm long, the ovate- lanceolate acuminate lobes 3-4 mm long. Corolla blue or purplish, rarely white, usually less than 30 mm in diameter. Anthers equal, 4-6 mm long. Fruit purplish black, 6-8 mm in diameter. Seeds not seen. Solanum fendleri is distributed in mountains from Colorado south through Arizona, New Mexico, and Texas to Zacatecas, México. In Texas the species has been collected from Brewster, Culberson, and Jeff Davis counties. Var. tezense Correll, restricted to the Davis Mts., is a taller and more open plant than var. fendlerz. The dense pubescence of the calyx imparts a grayish color to the inflorescence. The white to whitish lavender corolla is more deeply lobed, with narrower and more acute sinuses, and the calyx is irregularly and more deeply lobed than typical. 17. Solanum triflorum Nutt. Taprooted annual herbs, erect or ascending, to 0.4 m tall or long, the stems more or less strigose. Leaves ovate to elliptic, deeply pinnatifid, the lobes linear or nearly so, 1-4 cm long, 0.5-2.0 cm wide, strigose above and below; petioles to 2.cm long. Inflorescence 1 to 3 flowered. Calyx 4-5 mm long with lanceolate lobes 3-4 mm long. Corolla white, 8-10 mm in diameter. Anthers equal, 2.0- 2.5 mm long. Fruit green at maturity, 10-15 mm in diameter, containing 4-8 sclerotic granules. Seeds yellowish-brown, ovoid, flattened, 2.0-2.5 mm long, minutely reticulate. Solanum triflorum is very distinctive by its unarmed, deeply pinnatifid leaves. The species is widely distributed across the western half of the U.S. and is occasionally adventive eastward to the Atlantic states. In Texas it has been collected in Culberson and Hemphill counties. 18. Solanum pseudocapsicum L. Small shrubs to about 1.2 m tall, the stems puberulent with small dendritic hairs or glabrous. Leaves narrowly lanceolate to oblong or oblanceolate, entire or somewhat sinuate, revolute, obtuse or acuminate at the apex, tapering at the base, 2-10 cm long, 0.5-1.5 cm wide, sparsely to densely pubescent with 374 PHYTOL O CHA volume 71(5):362-378 November 1991 small dendritic hairs or rarely glabrous; petioles to 1 cm long or the leaves subsessile. Flowers solitary or 2-3 in lateral inflorescences. Calyx 4-6 mm long, the deltoid to lanceolate lobes 1.5-3.0 mm long. Corolla white, 10-15 mm in diameter. Anthers equal, 2.5-3.0 mm long. Fruit orange-red or rarely yellow, 10-20 mm in diameter. Seeds yellowish brown, flattened, 2-4 mm long, minutely reticulate. Solanum pseudocapsicum is a native of the Old World that has become naturalized throughout much of the New World tropics and subtropics. In Texas the species is known from several collections in Jasper County, the most recent dating from 1954. 19. Solanum triquetrum Cav. Erect suffruticose perennials to 0.5 m tall or more commonly scandent and reaching 2 m in length, the young stems somewhat striate, glabrous. Leaves deltoid to cordate, usually hastate or hastately 3 or 5 lobed, the central lobe ovate to lanceolate or linear lanceolate, 0.5-5 cm long, glabrous or with a few short marginal hairs. Inflorescence 2 to 5 flowered, the pedicels articulate at the base. Calyx 1.5-3.0 mm long, the deltoid to linear lanceolate lobes 0.5-1.5 mm long. Corolla white or violet tinged, 10-15 mm in diameter. Anthers equal, 2-4 mm long. Fruit red, 10-15 mm in diameter. Seeds yellowish brown, ovoid, flattened, 2.5-3.5 mm long, rough textured. S. lindhetmerianum Scheele. Solanum triquetrum is common on slopes, disturbed ground, and in thick- ets in central, west, and south Texas and adjacent México, often flowering throughout the year. The species is highly variable, especially in regard to its habit and leaf shape. 20. Solanum douglasi Dun. Perennial herbs or subshrubs, usually to about 1 m tall but occasionally to as much as 3 m, the stems sparsely to densely puberulent with simple ap- pressed or subappressed hairs or less often villous with spreading hairs, small glandular capitate hairs often present as well. Leaves ovate to narrowly ovate elliptic, the blades entire to sinuate or irregularly dentate, acute to obtuse at the apex, basally cuneate to subtruncate, 3-8 cm long, 1.5-4.0 cm wide, sparsely to densely strigose above and below; petioles slightly winged, to 3 cm long. Inflorescence 4 to 6 flowered, the flowers usually disposed in umbelliform cymes or occasionally solitary. Calyx 1-2 mm long, with ovate to lanceolate or rounded lobes less than 1 mm long. Corolla white, sometimes tinged with purple, 10-18 mm in diameter. Anthers equal, 2.5-4.9 mm long. Style exserted beyond the anthers by at least 1 mm; stigma only slightly expanded. Fruit purplish black, 5-10 mm in diameter, containing 4-10 sclerotic granules. Seeds Lemke: Solanum in Texas B75 yellowish brown, ovoid, plump, 1.2-1.5 mm long, minutely reticulate. Chro- mosome number, n = 12. Solanum douglasi is widespread, but infrequent, from Louisiana and Texas to California, south through México to Central America. In Texas the species has been collected from all regions of the state but is poorly represented from the High Plains and Rolling Plains. Solanum douglasi, S. sarrachoides Sendt., S. intertus Rydb., and S. pty- canthum Dun. er DC. constitute the Texas representatives of the Solanum ni- grum L. complex (Solanum sect. Solanum), a taxonomically difficult group of weedy species that are often adventive far from their original habitats (Steb- bins & Paddock 1949). Recent studies by Heiser et al. (1979), Schilling & Heiser (1979), and Schilling (1981) have helped to clarify the delimitation of the North American members of the complex. Reports of Solanum villosum Mill. from Texas (Correll & Johnston 1970) were based in part on misidentified densely puberulent specimens of S. dou- glaszi. 21. Solanum sarrachoides Sendt. Taprooted annual herbs, erect or decumbent, to 0.8 m tall or long, the stems viscid pubescent with uniseriate glandular capitate hairs. Leaves triangular ovate to ovate, entire to undulate or dentate, apically acute, at the base cuneate to truncate, 2-5 cm long, 1-3 cm wide, sparsely glandular pubescent marginally and on the main veins above and below; petioles slightly winged, to 2 cm long. Inflorescence umbellate, 2 to 6 flowered. Calyx 1.5-2.0 mm long, the lanceolate lobes 0.5-1 mm long. Corolla white, 5-7 mm in diameter. Anthers equal, 1.5- 2.0 mm long. Fruit brownish green to nearly black, the lower one third to one half loosely invested by the enlarged calyx, 6-10 mm in diameter, containing 5-7 sclerotic granules. Seeds yellowish brown, ovoid, 1.7-2.4 mm long, minutely reticulate. Chromosome number, n = 12. Solanum sarrachoides is a native of South America that is adventive and widespread in fields and disturbed areas throughout much of North America. In Texas the species is known from a single collection in the Chisos Mts. of Brewster County (Sperry 475 [SRSC]). This specimen was misidentified as S. villosum Mill., a tetraploid Eurasian member of sect. Solanum that, like S. sarrachoides, is glandular pubescent, but lacks the enlarged, accrescent calyx. No specimens referable to S. villosum have been seen from Texas. The specific epithet is spelled “sarachoides” by some authors. 376 PHY TOLOGIA volume 71(5):362-378 November 1991 22. Solanum interius Rydb. Deeply rooted annual or short lived perennial herbs to 0.8 m tall, the stems sparsely to densely strigose. Leaves ovate to triangular ovate or rhombic, entire, undulate or sinuate, apically obtuse, acute or short acuminate, at the base cuneate, rounded, or subtruncate, 3-7(-10) cm long, 1-4 cm wide, sparsely to densely strigose, especially below; petioles slightly winged, to 3 cm long. Inflorescence umbellate, 2 to 6 flowered. Calyx 1.5-2.0 mm long with ovate lobes 1.0-1.5 mm long. Corolla white or blue-violet, occasionally with purple stripes, 15-20 mm in diameter. Anthers equal, 1.6-2.0 mm long. Fruit dull purple-black, 5-10 mm in diameter, containing 2 or rarely 4 sclerotic granules. Seeds yellow to brown, ovoid, flattened, 1.9-2.3 mm long, minutely reticulate. Chromosome number, n = 12. S. nigrum L. var. interius (Rydb.) F.C. Gates. Solanum interius is found in sandy soils throughout much of the Great Plains, from North Dakota to Idaho, south to Texas and New Mexico. Our col- lections are primarily from sandy sites in the High Plains (Hartley, Hemphill, Ochiltree, and Wheeler counties) with disjunct collections from Taylor and Jeff Davis counties. 23. Solanum ptycanthum Dun. ez DC. Taprooted annual herbs 0.3-0.6 m tall, rarely to 1 m or more in height, the stems glabrous to moderately strigose, unarmed. Leaves very variable, ovate to ovate lanceolate, entire to sinuate dentate, acute, acuminate, or obtuse at the apex, at the base cuneate or rounded to subcordate, 5-15 cm long, 2-5 cm wide, glabrous to moderately strigose; petioles usually winged, to 4 cm long. Inflorescence umbellate, 2 to 4 flowered. Calyx 1.5-2.0 mm long, unequally 5 lobed, the triangular to lanceolate or rounded lobes 1.0-1.5 mm long, unequal and partially fused in fruit, not reflexed. Corolla white or tinged with purple, 10-15 mm in diameter. Anthers equal, 1.4-1.9 mm long. Fruit purplish black, 5-9 mm in diameter, usually containing 6 or more sclerotic granules. Seeds yellowish brown, ovoid, plump, 1.5-1.9 mm long, minutely reticulate. Chromosome number, n = 12. Solanum ptycanthum is widespread throughout eastern North America, west to North Dakota and Texas. The species is found in most regions of the state in open woodlands, along streams and roadsides, and in waste places. These plants were treated as Solanum americanum Mill. by Correll & John- ston (1970). Heiser et al. (1979) have argued, however, that this name should be applied to those plants previously recognized as S$. nodiflorum Jacq. and that the name S. ptycanthum should be used for plants previously referred to as S. americanum. Although S. americanum (as S. nodiflorum) was reported from coastal Texas by Jones et al. (1961), I have seen no specimens from the state and do not consider the species to be a member of our flora. if Lemke: Solanum in Texas Sih Cultivated species: Solanum diphyllum L. is occasionally planted as an ornamental shrub and has been collected from Harris County (Houston) and Travis County (Austin). The species is easily recognized by the 1-2 small, rounded pseudostipular leaves produced at the base of each simple elliptic- ovate leaf and by the subumbellate inflorescences borne on lignified lateral peduncles, these bearing prominent pedicel scars in fruit. ACKNOWLEDGMENTS Appreciation is extended to Guy Nesom and Mark Bierner for their critical reviews of the manuscript. LITERATURE CITED Correll, D.S. 1952. Section Tuberartum of the genus Solanum of North Amer- ica and Central America. U.S.D.A. Agric. Monogr. 11:1-243. Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of Teras. Texas Research Foundation, Renner, Texas. 1881 pp. D’Arcy, W.G. 1973. Flora of Panama. Family 170. Solanaceae. Ann. Missouri Bot. Gard. 60:573-780. Edmonds, J.M. 1972. A synopsis of Solanum sect. Solanum (Maurella) in South America. Kew Bull. 27:95-114. Heiser, C.B., Jr., D.L. Burton, & E.E. Schilling. 1979. Biosystematic and taxometric studies of the Solanum nigrum complex in eastern North America. Pp. 513-527 in J.G. Hawkes, R.N. Lester, & A.D. Skelding, eds., The Biology and Tazonomy of the Solanaceae, Academic Press, London, U.K. 738 pp. Jones, F.B., C.M. Rowell, Jr., & M.C. Johnston. 1961. Flowering plants and ferns of the Texas coastal bend counties. Welder Wildlife Foundation Publ. B-1, Sinton, Texas. 165 pp. Nee, M. 1979. Patterns in biogeography in Solanum, section Acanthophora. Pp. 569-580 in J.G. Hawkes, R.N. Lester, & A.D. Skelding, eds., The Biology and Tazonomy of the Solanaceae, Academic Press, London, U.K. 738 pp. Schilling, E.E. 1981. Systematics of Solanum sect. Solanum (Solanaceae) in North America. Syst. Bot. 6:172-185. 378 PHYTOLOGIA volume 71(5):362-378 November 1991 & C.B. Heiser. 1979. Crossing relationships among diploid species of the Solanum nigrum complex in North America. Amer. J. Bot. 66:709- 716. Stebbins, G.L., Jr. & E.F. Paddock. 1949. The Solanum nigrum complex in Pacific North America. Madrono 10:70-81. Whalen, M.D. 1979. Taxonomy of Solanum sect. Androceras. Gent. Herb. 11:359-426. Phytologia (November 1991) 71(5):379-381. ARTHRAXON HISPIDUS (POACEAE), NEW TO TEXAS Larry E. Brown & Jeff Schultz Spring Branch Science Center Herbarium, 8856 Westview Drive, Houston, Texas 77055 U.S.A. ABSTRACT Arthrazon hispidus (Thunb.) Makino is reported new to Texas from Cass County. This collection is a slight range extension from adjacent localities in Arkansas and Louisiana. KEY WORDS: Arthrazon, Poaceae, Texas Arthrazon hispidus (Thunb.) Makino, an introduced Asian grass, was found on a roadbank along U.S. highway 59 in Cass County, northeast Texas. The nearest sites to this locality are Claiborne Parish, Louisiana (Allen 1980), and Pike and Clark counties in Arkansas (Smith 1988). Neither Johnston (1989, 1990) nor Hatch et al. (1990) listed this species for Texas. This grass has spread to the following states since Kiger’s (1971) distribu- tional summary: West Virginia, Ohio, Kentucky, Indiana (Cusick 1986) and now Texas. Van Welzen (Blumea 27:255-300, 1981) has lately revised the genus. It is of interest to note that Microstegium vimineum (Trin.) A. Camus, another introduced Asian grass and reported as a frequent associate in other states, was found recently in nearby Bowie Co., Texas (Nixon et al. 1987). Specimen collected: UNITED STATES. Texas: Cass Co., along highway 59, 0.25 mi N of Linden, 22 Sep 1991, Jeff Schultz 0308A (SBSC,TAES). The plants were growing in wet to moist soils in the full sun of an open area. Some associated plants were Panicum spp., Onoclea sensibilis L., Vernonia missurica Raf., and Helianthus angustifolius L. Figure 1 is a drawing of this Texas collection. 379 380 PHY TOLOGIA volume 71(5):379-381 November 1991 Figure 1. Arthrazon hispidus. A. portion of plant with: 1. adventitious roots, 2. foliage leaf, 3. bractlike leaf subtending the inflorescence, 4. inflorescence. B. two spikelets on rachis: 1. closed spikelet, 2. open spikelet. The pedicelled spikelet is reduced to the small pedicel at the base of the spikelet. Brown & Schultz: Arthrazon hispidus, new to Texas 381 ACKNOWLEDGMENTS We thank Charles Peterson (SBSC) and Kancheepuram N. Gandhi (NCU) for their helpful comments. We also thank Ellis Grenier of Houston Commu- nity College for the illustration and Marcia Braun of the Houston Community College Central Library for some technical support. LITERATURE CITED Allen, C.M. 1980. Grasses of Louisiana. The University of Southwestern Louisiana, Lafayette, Louisiana. Cusick, A.W. 1986. Distributional and taxonomic notes on the vascular flora of West Virginia. Castanea 51:56-65. Hatch, S.L., K.N. Gandhi, & L.E. Brown. 1990. Checklist of the Vascular Plants of Texas. MP-1655. Texas Agric. Expmt. Sta., College Station, Texas. Johnston, M.C. 1989, 1990. A List, Up-dating the Manual of the Vascular Plants of Teras. Published by the author, Austin, Texas. Kiger, R.W. 1971. Arthrazon hispidus (Gramineae) in the United States: Taxonomic and floristic status. Rhodora 73:39-46. Nixon, E.S.,S$.C. Damuth, & M. McCrary. 1987. Five additions to the Texas flora. Sida 12:421-422. Smith, E.B. 1988. An Atlas and Annotated List of the Vascular Plants of Arkansas, 2nd edition. Published by the Author, Fayetteville, Arkansas. Phytologia (November 1991) 71(3):382-386. TWO NEW SPECIES OF AECHMEA (BROMELIACEAE) Harry E. Luther Marie Selby Botanical Gardens, 811 South Palm Avenue, Sarasota, Florida 34236 U.S.A. ABSTRACT Two new species of Aechmea are described: A. haltonii (subgen. Podaechmea) from Panama and A. flemingii (subgen. Aechmea) from Guadeloupe. KEY WORDS: Aechmea, Bromeliaceae, Guadeloupe, Panama Aechmea flemingii Luther, sp. nov. (Figure 1). TYPE: GUADELOUPE. Grande Terre: Pterocarpus woods N of La Raizet airport, Conrad Flem- ing s.n. legit, 14 May 1985, (SEL 85-241), flowered in cultivation, 10 July 1991, H.E. Luther s.n. (HOLOTYPE: SEL; Isotypes: K,US). A Aechmea strobilina (Buerling) L.B. Smith & R.W. Read, cui affinis, spinis foliorum perminoribus, bracteis florigeris majoribus castaneisque differt. Plant flowering 0.6 m tall, very densely clustering. Leaves densely rosu- late, spreading, 0.8-1.3 m long, bright green. Leaf sheaths elliptic, 13-15 x 8-10 cm, entire, subdensely dark punctate lepidote, somewhat castaneous to- ward the base abaxially. Leaf blades ligulate, acute to rounded and apiculate, pungent, 3-5 cm wide, laxly and inconspicuously antrorse serrate with dark 0.5 mm long spines, inconspicuously punctate lepidote. Scape erect, mostly concealed within the leaf sheaths. Scape bracts erect, tubular-imbricate, nar- rowly lanceolate, attenuate, entire, the sheaths castaneous abaxially, the blades green. Inflorescence densely digitate with 4 to 10 branches, 20 x 15 cm, mostly concealed within the rosette. Primary branches lustrous. castaneous, orangish green to bronzish green, with a pale green margin. Branches subsessile, slightly spreading, each with 1-3 sterile bracts at the base, 8-12 x 3 cm, 10 to 25 flow- ered. Floral bracts erect, densely imbricate, concealing the rachis and most of 382 Luther: Two new species of Aechmea 383 Figure 1. Aechmea flemingii. A. leaf. B. inflorescence. C. floral bract. D. flower. E. sepal. F. flower, longitudinal section. G. petal. 384 PHY TO L°O'G TA volume 71(5):382-386 November 1991 the flowers, ovate, acute, mucronate, 32-37 mm long, carinate, coriaceous, pale punctate lepidote especially near the keel, lustrous, orangish green or bronzish green, drying castaneous with a pale margin. Flowers sessile, erect. Sepals asymmetrical, oblong to obovate, mucronate, 15-17 mm long, the adaxial pair carinate, green. Corolla erect, tubular, barely opening at anthesis. Petals ligu- late, obtuse, 20-23 mm long, each with a pair of 7 mm long, basal appendages, bright yellow. Fruit an ellipsoid dark blue berry. Paratype: GUADELOUPE. type locality, clone of the holotype, 14 May 1985, Conrad Fleming s.n. (SEL). Aechmea flemingii closely resembles A. strobilina (Buerling) L.B. Smith & R.W. Read from the Caribbean coast of Panama, but is clearly distinct due to its small and lax leaf serrations and larger floral bracts. In addition, the primary and floral bracts of A. strobilina are brightly colored, (rose red or orange), and the petals are pale (white or cream). The name honors the collector, Conrad Fleming of St. Croix, Virgin Islands, a well known plantsman. Aechmea flemingw is evidently a narrow island endemic probably over- looked due to its inconspicuous flowering habit with the inflorescence mostly concealed within its large leafy rosette. Aechmea haltonii Luther, sp. nov. (Figure 2). TYPE: PANAMA. Prov. Coclé: El Copé, lithophyte, Aug 1985, J. Halton s.n. (HOLOTYPE: SEL). A Aechmea mezicana Baker, cui affinis, laminis foliorum tri- angularibus, acutis, pungentibus; scapis bracteis primariisque ser- ratis; sepalis petalisque brevioribus differt. Plant flowering 0.75-1.50 m tall, clustering. Leaves densely rosulate, erect to spreading, 30 to 45 in number, 35-90 cm long, coriaceous. Leaf sheaths elliptic, 15-25 x 6-15 cm, entire, densely brown punctate lepidote, dark cas- taneous toward the base abaxially. Leaf blades triangular, acute, pungent, 2-6 cm wide, subdensely serrate with dark 3-8 mm long straight or antrorse spines, subdensely pale punctate lepidote, bright green. Scape erect, 25-65 cm x 8-12 mm, fugaciously white lanate, bright red or pink. Scape bracts elliptic, acute, pungent, serrate, the lowest subfoliaceous, the upper narrowly elliptic, erect and much exceeding the internodes, bright red or pink with green tips. Inflorescence laxly tripinnate, 40-60 x 20-40 cm, many flowered. Primary bracts elliptic, acute, pungent, serrate, exceeding the naked, sterile bases of the branches, often reflexed, gradually diminishing in size toward the apex of the inflorescence, thin coriaceous, nerved, sparsely pale lanuginous lepidote, bright red or pink. Primary branches with a flattened 5-90 mm long, sterile, Luther: Two new species of Aechmea 385 ed 5mm 5cm Figure 2. Aechmea haltonu. A. leaf, adaxial surface. B. branch of inflorescence. C. primary bract. D. flower. E. sepal. F. petal and stamen. 386 PHY TL ODO GIA volume 71(5):382-386 November 1991 ebracteate, sparsely pale lepidote base, spreading at 45-90° from the main axis, 2-25 cm long. Secondary branches with a 1-5 mm long base, 1 to 7 flowered. Floral bracts narrowly triangular to filiform, attenuate, 1.00-1.25 mm long, red. Flowers with a slender 2-3 mm long, green, glabrous pedicel, polystichously spreading at ca. 45° from the axis at anthesis. Sepals very asymmetrical, trun- cate, 4 x 4 mm, the broad lateral wing exceeding the 1 mm long dark terminal mucro, glabrous, green with red margins. Corolla erect, tubular, barely open- ing at anthesis. Petals oblanceolate, obtuse, or emarginate, slightly cucullate, 10-11 mm long, each with a pair of basal appendages, pink. Ovary ellipsoid, glabrous, pale green, epigynous tube ca. 1 mm deep, placentae apical. Fruit a globose white berry 6-8 mm long. Paratypes: PANAMA. Type locality, clone of the holotype (SEL 85-1447), flowered in cultivation, 12 Dec 1990, H. Luther s.n. (SEL); March 1991, J. Anderson s.n. (SEL); 28 Jul 1991, H. Luther s.n. (PMA,US); Prov. Veraguas, 2 km above Santa Fe on road to Cerro Negro, 2200 feet, epiphyte, 23 Sep 1991, C. Skotak s.n. (SEL). Aechmea haltonii seems to be most closely related to A. mezicana Baker, a species known from México to western Ecuador. It can be immediately dis- tinguished by its triangular and conspicuously dark serrate leaf blades, serrate scape and primary bracts, broader inflorescence, and smaller flowers. The name honors the original collector, the late Joseph S. Halton, first Display Greenhouse Manager at the Marie Selby Botanical Gardens, who in- troduced many ornamental and interesting tropical plants to horticulture. ACKNOWLEDGMENTS I thank Dr. John T. Atwood and Dr. Gregory K. Brown for their comments on the original draft of the manuscript. Phytologia (November 1991) 71(5):387-389. KOSTELETZKYA DEPRESSA, SECTION KOSTELETZKYA (MALVACEAE), NEW TO TEXAS 'Stanley D. Jones, 7Gretchen D. Jones, & 'J.K. Wipff 1S.M. Tracy Herbarium (TAES), Department of Rangeland Ecology and Management, Texas A&M University, College Station, Texas 77843-2126 WS “Department of Biology, Texas A&M University, College Station, Texas 77843-3258 U.S.A. ABSTRACT Kosteletzkya depressa (L.) O. Blanchard, Fryxell, & Bates, sec- tion Kosteletzkya (Malvaceae) previously unreported in Texas has been found in Cameron County. KEY WORDS: Kosteletzkya, Kosteletzkya depressa, section Kostel- etzkya, Malvaceae, Texas Kosteletzkya K. Presl, a genus of 17 species, occurs in the New World in North, Central, and South America, and in the Old World from Africa and Madagascar to Europe and Malesia (Fryxell 1988). Kosteletzkya depressa (L.) O. Blanchard, Fryxell, & Bates, section Kosteletzkya, was first described by Linnaeus (1753) as Melochia depressa L. Pres] (1835) was the first to move it into the genus Kosteletzkya, but failed to use the oldest available specific epithet. Seven others followed in renaming this taxon, maintaining it in the genus Kosteletzkya, but it was not until 1978 that the oldest correct epithet was recognized by O. Blanchard, Fryxell, & Bates. For a detailed nomenclatural history, see Fryxell (1988). Kosteletzkya depressa is reported from southern Florida, the Caribbean Islands, is widely distributed in México including Baja California, and also occurs in South America at least to Ecuador. Cory & Parks (1937), Correll & Johnston (1970), Gould (1975), Hatch et al. (1990), nor Richardson (1990) listed K. depressa as occurring in Texas, strongly suggesting that this is a re- cent introduction. Based on previously mapped distributions (Fryxell 1988), 387 388 P4IEVT OL OGIA volume 71(5):387-389 November 1991 the Cameron County station is approximately 200 air mi. (324 km) from the closest previously known station in southern Tamaulipas, México. There are now two species of Kosteletzkya represented in Texas. The following key dif- ferentiates them. KEY TO TEXAS KOSTELETZKYA 1.. Petals 8 to: 12 mm long; calyx 4 to’8:mm long:) 5. jcc¢h. eee SS ESOS e Er ane Oe K. depressa (L.) O. Blanchard, Fryxell, & Bates 1. Petals 30 to 45 mm long: calyx 8 to 13 mm long. 2202 e eee ae Jarek uid aoa ee ne re Wem fo ey Bier Apel hace Dy, K. virginica (L.) A. Gray Our specimen is a robust tap rooted annual about 2 m tall with the lower stem having a conspicuous narrowly rhombic pattern of vasculature (at least when dry). It is much branched above with many diffuse leafy panicles spread- ing to about 1 m. Flowers are solitary in the leaf axils, rotate, whitish fading to pink. The staminal column is bright yellow. The specimen has stinging trichomes (especially along the lower portion of the plant). The trichomes are of two types, stellate and simple. The simple trichomes are appressed to ascending. The capsules have some scattered stellate trichomes with uncinate trichomes on the margins of the wings. The leaves are lanceolate to hastate with stellate pubescence on both surfaces. The leaves are reduced upwards. Specimen collected: UNITED STATES. Texas: Cameron Co., 20 October 1991, S. & G. Jones 7787 & J.K. Wipff (BRIT,ob[O.J. Blanchard’s personal herbarium],PAUH,pf[Paul A. Fryxell’s personal herbarium],TEX). The pop- ulation consists of about ten plants. The collection site is 3.5 mi. (5.7 km) south on Farm Road 1847 from its junction with Farm Road 106, SE of La Tina. The longitude is 97° 28’ 00” W and the latitude is 26° 09’ 50” N, E of Cross Lake referencing the Laguna Atascosa 7.5’ Quad. sheet. The habitat is a low lying drainage area with permanent to semipermanent water with the elevation being about 10 feet (3 m) above sea level. The soils are in the Lomalta series of the Laredo-Olmito association and are specifically Lomalta clay (LM). Permeability is very slow, with the available water capacity low. The soil is dark gray to dark grayish brown, calcareous; slightly to moder- ately saline. The geology of the site is Alluvium formation (Qas) (Recent). Associated species include Mimosa pigra L., Prosopis glandulosa Torr., Cype- rus ochraceus Vahl, Marsilea macropoda A. Br., Borrichia frutescens (L.) DC., Polygonum sp., Paspalum sp., and Eleocharis sp. Jones et al.: Kosteletzkya depressa, new to Texas 389 ACKNOWLEDGMENTS We thank O.J. Blanchard, Jr. (Long Island State University) for verifying our collection and sharing his expertise. We thank Paul A. Fryxell (USDA) for technical directions and reviewing this manuscript, and Robert Lonard (PAUH) for reviewing this manuscript. LITERATURE CITED Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of Teras. Texas Research Foundation, Renner, Texas. Cory, V.L. & H.B. Parks. 1937. Catalogue of the Flora of Tezas. Texas Agric. Exp. Sta. Bull. 550. Gould, F.W. 1975. Texas Plants-A Checkhst and Ecological Summary. Texas Agric. Exp. Sta. Bull. MP-585. Fryxell, P.A. 1988. Malvaceae of México. Syst. Bot. Monogr. 25:1-522. Hatch, D.L., K.N. Gandhi, & L.E. Brown. 1991. Checklist of the Vascular Plants of Texas. Texas Agric. Exp. Sta. Bull. MP-1655. Linnaeus, C. 1753. Species Plantarum. 674. Presl, K.B. 1835. Reltg: Haenk. 2:131. t. 70. Richardson, A. 1990. Plants of Southernmost Teras. Gorgas Science Foun- dation, Inc., Brownsville, Texas. Phytologia (November 1991) 71(5):390-413. NOTES ON NEW INFRASPECIFIC TAXA AND HYBRIDS IN NORTH AMERICAN POA (POACEAE) R.J. Soreng L.H. Bailey Hortorium, Cornell University, Ithaca, New York 14853 U.S.A. ABSTRACT Five new infraspecific combinations and three new subspecies are proposed. Poa abbreviata subsp. marshii, P. arctica subsp. lanata, P. cusickii subsp. purpurascens, P. hartzii subsp. alaskana, P. hartzii subsp. ammophila, P. laxa subsp. banffiana, P. pratensis (subsp. alpigena] var. colpodea, and P. secunda subsp. juncifolia. Poa evagens is assigned to Deschampsia caespitosa subsp. brevifolia, and the putative parentage of P. x fibrata (P. pratensis x P. secunda subsp. guncifolia) is discussed. Comments on the new taxa, new combinations, relationships of these taxa, additional chromosome counts, and keys are provided. KEY WORDS: Poa, Deschampsia, grasses, Poaceae, taxonomy, apomixis, hybrids, polyploidy, North America, arctic, alpine - 1. Poa abbreviata R. Br. subsp. marshii R.J. Soreng, subsp. nov. TYPE: U.S.A. Idaho: Blaine Co., Sawtooth Mts., head of Boulder Creek Canyon, 10,000’, granite talus, 2 Aug 1937, J.W. Thompson 14083 (HOLOTYPE: US 1649210!; Isotypes: CAS,CU!,F,GH,NY!,RSA!,WTU) (Fig. 1). A P. abbreviata R. Br. subsp. abbreviata et subsp. pattersoni (Vasey) A. Love, D. Love, & Kapoor lemmatibus glabris et callo arachnideo differt. Perennial. Culms 5-15 cm tall, slender, from small dense tufts with narrow bases, shoots intravaginal. Upper culm leaf sheath margins fused 1/10-1/4 the length; ligules 1-3 mm long, smooth; blades 1.0-1.5 mm wide, folded and inrolled, lacking papillae, abaxially smooth, adaxially scabrous on and between 390 Soreng: New taxa and hybrids in North American Poa 391 Figure 1. Poa abbreviata R. Br. subspp. a. subsp. marshii R.J. Soreng, habit (Thompson 14083; HOLOTYPE: US 1649210!). b. subsp. abbreviata, habit. c. subsp. abbreviata, spikelet. d. floret (Holmen 6626). e. subsp. marshii, floret (Hitchcock & Muhlick 11146). f. subsp. pattersonii, (Vasey) A. Love, D. Love & Kapoor (jordalii form), floret (Cantlon & Gillis 57-797). g. subsp. pattersonzi, floret (Soreng & Spellenberg 1165). 392 PRY POLO GTA volume 71(5):390-413 November 1991 the veins. Panicles 2-5 cm long, lanceolate, the branches scabrous angled. Spikelets with 2-4 florets, 5-6 mm long; glumes subequal, 1-3 veined, the first slightly shorter, second frequently slightly exceeding the lower lemma; calluses of the lower lemmas webbed (sometimes minutely); lemmas 3.5-4.0 mm long, glabrous, smooth; palea keels scabrous; rachillas smooth. Flowers perfect; anthers 0.6-1 mm long. Habitat. High alpine slopes rocky slopes. Distribution and Specimens. U.S.A. California (Mono Co., White Mts., Morefield & Ross 4695.2 RSA!, with subsp. pattersonu M. & R. 4695.1 RSA!). Idaho (Blaine Co. [TYPE]; Butte Co., Lemhi Range, Diamond Pk., Moseley 507 NY!; Custer Co., Lost River Mts., Leatherman Pass, Hitchcock & Muhlick 11146 CU!,GH,NY!,US,WTU). Nevada (White Pine Co., Shell Creek Range, Shell Pk., N. Holmgren, Reveal, & Lafrance 2245 NY!). Subspecies marshii is distinguished from all other Poa abbreviata only in having entirely smooth, glabrous lemmas (Fig. 1). It was independently de- scribed as a subspecies by D.D. Keck in an unpublished manuscript, and by V.L. Marsh, for whom the subspecies is named, in his unpublished disser- tation (1950). Both authors cited two collections from Idaho (Hitchcock & Muhlick 11146, and J.W. Thompson 14083). Three additional specimens of subsp. marshii have been discovered in Idaho, Nevada and California. With one exception, subsp. marshizis allopatric to other subspecies of P. abbreviata, suggesting that the proposed taxon is more than a local population variant or a hybrid. Recognition of the new subspecies involved restricting the circumscription and reevaluating the geographic range of subsp. pattersoni (Vasey) A. Love, D. Love, & Kapoor. The latter taxon is frequently confused (as it is in the Intermountain Flora) with Poa glauca Vahl sensu lato (s. lat.) and P. secunda J.S. Presl s. lat. Poa abbreviatas. lat. and P. glauca can be distinguished from one another with relative ease by observing the base of the plant. The new shoots of P. abbreviata develop intravaginally, the bases of both flowering and nonflowering shoots are clothed in sheaths with blades, and the prophylls are generally more than 1 cm long. Most new shoots of P. glauca develop extrav- aginally, breaking laterally out of old sheaths, the basal leaves are bladeless (cataphylls), nonflowering shoots of a season are few in number, and prophylls are generally less than 0.6 cm long. Poa abbreviata also has more slender culms and lustrous spikelets. These differences in ramification and aspect are correlated with differences in anther length: 0.2-1.2 mm long in P. abbreviata, and 1.2-2.5 mm long in P. glauca. The anthers of P. secunda are even longer (1.5-3.0 mm), the lemmas are only weakly keeled and never have an isolated single tuft of hair (web) on the dorsal surface of the callus, and the longest panicle branches exceed 1.8 cm in length. (Many specimens previously filed under P. pattersonii Vasey have been annotated as other species. Those I have verified as this subspecies are listed below. See also comments under P. laza.) Soreng: New taxa and hybrids in North American Poa 393 Under this interpretation P. abbreviata subsp. pattersoni and subsp. marshu are only known to be sympatric in the White Mountains of California. KEY TO THE SUBSPECIES OF POA ABBREVIATA 1. Lemmas glabrous; callus of basal florets webbed; U.S.A. California, Idaho, Wewadan Meio, owen itte DA naiof Pinnde Sala ats, SAO Te subsp. marshi 1. Lemmas pubescent; callus of basal florets webbed or not. ............. 2 2. Lemma intervein regions abundantly pubescent, callus glabrous (rarely distinctly webbed); arctic tundras; circumboreal, north of 60° N (rare in western North America and eastern U.S.S.R.) (map p: 147.5 Hulten; 1974); 2n = 28, 42) 70,76. -..... subsp. abbreviata 2. Lemma intervein regions glabrous or sparsely puberulent, callus frequently webbed; alpine to arctic tundras; Far eastern U.S.S.R. CANADA. Alberta, British Columbia, sw Northwest Territories, Yukon. U.S.A. Alaska, California, Colorado, Montana, Utah, Wyo- pT Gg CRE: bilan eo re OTe a RYT subsp. pattersont 2. Poa abbreviata R. Br. subsp. pattersonii (Vasey) A. Love, D. Love, & Kapoor, Arctic & Alpine Res. 3:142. 1971. BASIONYM: Poa pat- tersoni Vasey, Contr. U.S. Natl. Herb. 1:275. 1893. TYPE: U.S.A. Colorado: mts. about the head waters of Clear Creek, 11-14,000’, top of Mt. McClellan near Grays Peak, 19 Aug 1885, Patterson 154 (Isotypes: NY!,US 556757!,US s.n.!). Poa jordalii A. Pors., Canad. Field-Naturalist 79:82, fig. 1. 1965. Poa abbreviata R. Br. subsp. jordaliz(A. Pors.) Hultén, Bot. Not. 126:468. 1973. TYPE: U.S.A. Alaska: south slope of Brooks Range, Bet- tle’s River, in alpine tundra on limestone, elev. 2000’, L.H. Jordal 2284 (HOLOTYPE: CAN!; Isotype: US 1980583!). Chromosome number. 2n = 42, U.S.A. Montana: Anaconda-Pintlar Wilderness, Mt. Tiny, Soreng & Spellenberg 1165; Colorado: Hoosier Ridge, Soreng et al. 2548, Mt. Evans, Soreng et al. 2555. These three new counts agree with the two previous counts for subsp. pattersonii from Colorado (Love et al. 1971), and a report for “subsp. jordalix” (A. Pors.) Hultén from the U.S.S.R., as well as the most frequent number reported for subsp. abbreviata (Love & Love 1975). Distribution and Verified Specimens. CANADA. Alberta (Waterton Lakes N.P., Avion Ridge, Brettung 17288 US!). British Columbia (Antimony Mt., 394 PHYEOL OCIA volume 71(5):390-413 November 1991 50° N x 122° W, Tisdale 1938 UBC!; Summit Pass, 58° N x 124° W, Raup & Correll 10651 CAN!, 10705 CAN!,UBC!, 10704 CAN!). Northwest Territories: District of Mackenzie (Mackenzie Mts., 63° N x 128° W, Cody 16690 CAN!; Nahanni N.P., 62° N x 127° W, Talbot T 6145 CAN!). Yukon (Sheep Mt., 61° N x 139° W, Krajina & Hoefs 1970 UBC!). U.S.A. Alaska (Kanayut Lake, 68° N x 151° W, Spetzman 1958 s.n. US!,US!; Ambresvajun Lake, 68° N x 144° W, A. & C. Batten 75-482 CAN!, DAO!; Bettle’s River, 67° N x 150° W,Jordal 2284 CAN!,US!; Ivishak River, 68° N x 147° W, Hettinger 657 CAN!; Jag Mt., 69° N x 144° W, Cantlon & Gillis 57-797 CAN!. California (Mono Co., White Mts., Morefield & Ross 4695.1 (RSA!). Colorado (Chaffee Co., Mt. Harvard, Neely & Carpenter 2349 NY!; Clear Creek Co., Mt. McClellan, 19 Aug 1885, Patterson 154, NY!,US!, 23 Aug 1892, Patterson 154, US 748851! [Hitchcock 1935, fig. 235], Grays Peak, Shear 690 1/2 US!, Swallen 1407 US!, Soreng et al. 2555 NMC!; Gilpin Co., James Peak, C.F. Coz 491 US!; Gunnison Co., Conundrum Pass, J. Bar- rell 70b-55 US; Lake Co., Mt. Elbert, L. & E. Kelso 5055 & 5059 DAO!; Larimer Co., Longs Peak, Hitchcock 16278 US!, Longs Peak trail, Ulke in 1918 [labeled British Columbia] CAN!; Park Co., Mt. Bross, Weber et al. 2097 DAO!, Mt. Bross, E. Hartman & Rottman 2313; Summit Co., Hoosier Ridge, Soreng et al. 2548). Montana (Gallatin Co., Mt. Hyalite, 1 Aug 1902, Blank- inship s.n. US!, Lava Peak, Blankinship s.n. US!; Deer Lodge Co., Mt. Tiny, Soreng & Spellenberg 1165 NMC!). Utah (Duchesne Co., Uinta Mts., Kings Peak, Harrison et al. 10070 US!; Grand Co., La Sal Mts., Mt. Waas, Maguire et al. 16383 CAN!,CU!; San Juan Co., La Sal Mts., Mt. Peal, Maguzre et al. 16384 CU!). Wyoming (Park Co., Abasroka Mts., Evert 6260; Teton Co., Two Ocean Mt., Hitchcock 23172 US!; Yellowstone N.P., Soda Butte, Tweedy 634 NY!,US!). U.S.S.R. (Wrangel Island, Petrousky & Polozova 6005 DAO!). Poa jordali A. Pors. was based on plants from the Brooks Range of Alaska. The new species was said to lack a web on the callus, and to be pubescent only on the keel and marginal veins of lemmas. However, most of the material from that region has at least a vestige of a web on calluses of basal lemmas (Fig. 1). The web and intervein puberulence are also variable in occurrence in P. abbreviata subsp. pattersonii of the U.S. Rocky Mountains (Fig. 1). The only characters that I have found to be correlated with geography are: 1) the hairs of the lemma veins are often shorter (0.10-0.15 mm long), and broadened toward the blunt apex, in Alaskan, Yukon, and British Columbia material, as opposed to always longer (0.3-0.5 mm long), not broadened toward the apex, the apex acute in more southerly material; 2) the panicles of the northern plants are more exerted, average 1 cm shorter, and are more sparsely flowered. However, these differences in hairs and panicles are not consistent enough to warrant subspecific distinctions. Hultén reduced P. jordali to a subspecies, commenting that, “It is the Rocky Mountain counterpart to P. abbreviata, and Soreng: New taxa and hybrids in North American Poa 395 here regarded as a major race of that species.” However, P. abbreviata subsp. pattersonz, established two years earlier, has priority. Poa abbreviata grades through the P. jordalu form toward the closely re- lated P. lettermani Vasey, a more diminutive, yet more commonly occur- ring species, of similar habitat and range (ranging north to 60° N, British Columbia). Poa lettermanii is distinguished from P. abbreviata by its lemmas being glabrous or infrequently sparsely puberulent on the base of the keel, shorter (2.5-3.0 mm long), the first and second exceeded in length by both glumes, and anthers being 0.2-0.6 mm long. Its only reported chromosome number, 2n = 14 (A. Love, pers. comm.), differs from those reported for P. abbreviata. Where the one species stops and the other begins requires closer attention. 3. Poa arctica R. Br. subsp. lanata (Scribner & Merr.) R.J. Soreng, comb. et stat. nov. BASIONYM: Poa lanata Scribner & Merr., Contr. U.S. Natl. Herb. 13:72, fig. 16. 1910. TYPE: U.S.A. Alaska: Aleutian Islands, 17 Jul 1899, Coville & Kearney 2191 (HOLOTYPE: US 376421!). Poa malacantha V. Komarov, Bot. Mat. (Leningrad) 5(10):148. 1924. TYPE: U.S.S.R. Kamchatka, Komarov 2832 (LE). Poa komarovu Rosch., Izv. Glavn. Bot. Sada SSSR, 26(3):286. 1927. TYPE: U.S.S.R. Kamchatka, Komarov 1080 (LE). Distribution. CANADA. Alberta, British Columbia, Northwest Territo- ries, Yukon. Outside North America. Far eastern arctic U.S.S.R. U.S.A. Alaska. (map and figs., Hultén 1974). Applying species rank to Poa lanata is unworkable. The few characters (e.g., spikelet size, coloration, and rachilla pubescence) said to distinguish P. lanata from P. arctica are not correlated or grade continuously between the taxa. Poa malacantha, another taxon in sect. Poa, has also been recognized in North America (Hultén 1974; Tzvelev 1983). Tzvelev (1983, English trans- lation) separates P. malacantha from P. lanata as having “Rachilla almost always somewhat pilose; plants of the Far East [and Alaska], usually form- ing turf, but often with short creeping underground shoots.” versus “Rachilla glabrous; plants [of the northern Far East, including Alaska] with fairly long creeping underground shoots, usually not forming turf.” However, these vari- ations and all combinations of them occur in my collections from Alaska and those of UBC, V, and CAN, with no evidence of underlying geographical or ecological pattern. These taxa have almost identical geographic distributions (maps, Hultén 1974). As such, I believe P. malacantha to be synonymous with subsp. lanata. 396 PEO¥ PO LO Gd A volume 71(5):390-413 November 1991 KEY TO POA ARCTICA SUBSP. ARCTICA AND SUBSP. LANATA 1. Spikelets mostly greater than 6 mm long; lower lemmas 4.5-6.0 mm long; culm leaf blades often flat and 2-6 mm broad; Far eastern U.S.S.R. CANADA. Alberta, British Columbia, Northwest Territories, Yukon. W.S.A.. Alaska. se iotth.2nagith. acs oo nee aee ee subsp. lanata 1. Spikelets less than 6 mm long; lower lemmas 3.0-4.5 mm long; culm leaf blades often folded and less than 2(2.5) mm broad; circumboreal, south to New. Mexicoin U.S:A. ...... coca... es bee eee subsp. arctica (including: P. brintnelli Raup, P. arctica subsp. longiculmis Hultén, P. longipila Nash, and P. williamsu Hultén) 4. Poa cusickii Vasey subsp. purpurascens (Vasey) R.J. Soreng, comb. et stat. nov. BASIONYM: Poa alpina L. var. purpurascens Vasey, De- scr. Cat. Grasses U.S. 79. 1885. Poa cusickit Vasey var. purpurascens (Beal) [error for Vasey] C.L. Hitchc., Vasc. Pl. Pacific Northwest 1:659. 1969 [(Vasey) C.L. Hitchc. Emend. C.L. Hitchce., Fl. Pacific Northwest 659. 1973]. TYPE: U.S.A. Oregon: Mt. Hood, 4000-6000’, Aug 1981, T. Howell (HOLOTYPE: US 556826; Isotypes: GH!,ORE!,US 133409!. (map, Soreng 1991.). Soreng (1991) discussed the nomenclatural problems, morphological vari- ability, and breeding system associated with this taxon and recognized it as Poa cusickit Vasey subsp. epilzs (Scribner) W.A. Weber var. purpurascens (Vasey) C.L. Hitchc. The new combination is proposed to be consistent with several floras in progress that restrict the use of more than one infraspecific rank. 5. Poa laxa Haenke subsp. banffiana R.J. Soreng, subsp. nov. TYPE: CANADA. Alberta: Rocky Mountains, vicinity of Sunshine Ski Lodge, south of Healy Creek: Wa-wha Ridge, Standish Hump, alpine slopes and ridges, 7800’, 11 Aug 1945, Porsild & Breitung 14092 (HOLOTYPE: CAN!) (Fig. 2). Poa larva Haenke var. occidentalis Vasey er Rydb. & Shear?, U.S.D.A. Div. Agrostol. Bull. 5:32. 1897, nom. nud. SYNTYPES: U.S.A. Colorado: Grays Peak, Shear 690 and Rydberg 2440. [Not having seen the types of this recently, I can not be certain of its place- ment here, but several collections from alpine Colorado with narrow, smooth branched panicles, anthers ca. 0.8 mm long, and extravagi- nal branching closely approach P. lara subsp. banffianal. Soreng: New taxa and hybrids in North American Poa 397 Figure 2. Poa laza Haenke subsp. banffiana R.J. Soreng. a. habit (Porsild & Breitung 14092, HOLOTYPE: CAN!). b. spikelet (Porsild & Breitung 13960). 398 PHY TOG lA volume 71(5):390-413 November 1991 A P. laza Haenke subsp. laza, subsp. fleruosa (Smith) Hylan- der, subsp. fernaldiana (Nannf.) Hylander, et P. paucispicula Scrib- ner & Merrill callo glabro, lemmatis nervis intermediis puberulis differt. A P. abbreviata s. lat. et P. lara subsp. fernaldiana inno- vationibus frequenter extravaginalibus differt. A P. paucispicula paniculis contractis, ramis 2-4 (nec 1-2) brevioribus differt. Perennial. Culms 8-25 cm tall, from small tufts, shoots mainly extrav- aginal, vegetative shoots common and concurrent with the flowering shoots. Upper culm leaf sheath margins fused 1/5-1/3 the length; ligules 2.0-3.5 mm long, acute or lacerate, smooth; blades 1-2(-3) mm wide, flat or folded, prow tipped. Panicle 2.5-8.0 cm long, narrowly open, secund, the branches fairly strict (not flexuous), smooth or sparsely scabrous, 2-3(-5) per node, steeply ascending with 2-6 spikelets, the longest ones 1.2-3.0 cm long. Spikelets with 2-5 florets, 4.0-5.5 mm long; glumes 3 veined, broadly lanceolate, the second glume nearly equal to or longer than the first lemma (ratio 0.92/1-1.13/1); cal- lus glabrous; lemmas 3.0-4.6 mm long, lanceolate, the apex obtuse to acute, distinctly villous on the keel (for at least half the length) and marginal veins, usually sparsely puberulent on the base of the intermediate vein of at least one side of the lemmas, the surface smooth or sparsely finely muriculate; palea keels sparsely scaberulous; rachillas smooth. Flowers perfect; anthers 0.8-1.1 mm long. Chromosome number. 2n = 84, meiosis normal (U.S.A. Montana: Glacier N. P., Pigan Pass, Soreng & Spellenberg 1187 NMC). Habitat. High alpine, moist ground, slopes, ridges, and ledges. Distribution and Specimens. CANADA. Alberta (Lake Agnes, Malte in 1917; Porsild & Brettung 13029, Healy Creek, 13508 & 18513, Quartz Ridge, 13882, Citadel Peak, 13960, Wa-wha Ridge, 14092, Bow River Pass, 14869 & 14878, Upper N. Saskatchewan River, 16052). British Columbia (east sidé Simpson Pass, Lid & Porsild 717; Yoho National Park, Valley of Ten Peaks, Ulke in 1922; all the above at CAN!). U.S.A. Montana (Glacier National Park, Soreng & Spellenberg 1197, NMC!). The identity of my 1980 collection/cytological voucher, 1137, had plagued me for years until I found a series of matching material at CAN from the northern Rocky Mountains that A.E. Porsild had tentatively set aside as a new species. This series of collections is clearly allied to the Poa lara complex of sect. Oretnos (Asch. & Graebner) V. Jir., where I believe it is best placed as a new subspecies in P. laza. The presence of subspecies of Poa laza s. lat. in North America has long been debated. Nannfeldt (1935) excluded subsp. fleruosa (Smith) Hylander, and subsp. laza s. str. as strictly European (the former reaching Iceland; the latter strictly continental). He described P. fernaldiana Nannf. (P. laza subsp. fernaldiana) as the eastern North American counterpart. Soreng: New taxa and hybrids in North American Poa 399 Polunin (1959) equivocated on the geographic range, including Poa pseu- doabbreviata Rosch. within his broad delimitation of the species. Scoggan (1978), citing several maps, recognized subsp. fleruosa from the coastal re- gions of northcentral Baffin Isl., Greenland, and Ungava south to Labrador. Porsild & Cody (map 164, 1980) recognize P. fleruosa Smith (with the older P. laza in synonymy) from scattered locations in eastern Canada south to the Gaspe Peninsula. Having studied the material called Poa lara from North America and Eu- rope at CAN, CU, DAO, NY, and US, I have come to the conclusion that all of their vouchers from the high Nearctic including Greenland called P. lara subsp. fleruosa (except a few problematical immature specimens from seepage areas on the Ungavian Peninsula, Wolsenhome s.n., from Port Burwell and Wakeham Bay [CAN!!]) are referable to P. glauca s. lat. (see also Nannfeldt’s rejection of records in the historical literature for P. glaucas. lat.; p. 60, 1935). Specimens from the Atlantic Provinces, Labrador and Newfoundland, south to the Gaspe Peninsula, and northern New England are referable to P. laza subsp. fernaldiana. Distribution and Representative specimens of P. laza subsp. fernaldiana: CANADA. Newfoundland (Gros Morne, Bouchard et al. 84159 CAN!; High- lands of St. John, Deer Pond Brook, Fernald & Long 27405 CU!). Labrador (Lanse au Clair, Rev. Waghome in 1894). Quebec (Matane Co., Mont-Blanc, Gallo 1125 DAO!, 1128 DAO!, Mont-Blanc, Swallen 3093 CU!; Mt. Logan, Fernald & Pease 248738 CAN!, Mt. Fortin, Fernald & Pease 24873 CAN!; Gaspe Co., Mt. Blanc, Swallen 3471 CAN!, Mt. Jacques-Cartier, Raymond et al. 1877 CAN!, Rolland-Germain 1178 DAO!, Mt. Le Vieillard, Fernald et al. 25445 CAN!, Tabletop Mt., Scoggan 1257 CAN!, 1622 CAN!, Fernald & Collins 162 CAN!). U.S.A. Maine (Mt. Katahdin, Allard 5212!). New Hampshire (Mt. Wash- ington, numerous collections including the type, Williams & Robinson Plantae Ezsiccatae Grayanae 123 CU!, and cp-DNA voucher, Soreng 3401 CU!; Mt. Lafayette, 31 Jul 1863, W. Boott CU!). New York (Mt. Marcy, Erskins in 1954, 2n = 42 DAO!). Vermont (Mt. Mansfield, Woodward in 1911). Poa glauca is best distinguished from North American P. laza by its up- permost culm sheaths being open 1/10-1/5 their length; pruinose glumes, the second glume being 0.78-0.97 x as long as the first lemma; more firm, densely, finely muriculate lemma surfaces; short, stout, more or less scabrous angled (occasionally smooth) panicle branches; and longest anthers over 1.2 mm long. The other three subspecies of Poa lara exhibit a habit and panicle very similar to those of subsp. banffiana, but differ by uniformly smooth panicle branches, and presence of a web on the callus, and (in subsp. fernaldiana) by predominantly pseudo-intravaginal innovations (7.e., innovations intravaginal in origin, but with a short prophyll and one or more tubular (intravaginal prophylls are not tubular) bladeless leaves below the first leaf with a blade, 400 PHY POLO: GA volume 71(5):390-413 November 1991 as opposed to a long prophyll and no bladeless leaves), and thinner, often filiform, leaves. Although A.S. Hitchcock reports P. laza subsp. fernaldiana (P. fernaldiana) as lacking a web at the base of the lemmas, I have found a web to be present (though often sparse and short) on at least the lower florets within spikelets of most material. The new subspecies differs from Poa pseudoabbreviata in that the sheaths of the upper culm leaves are open 1/5-1/3 their length; panicle branches are more stout, smooth or only sparsely scabrous, more numerous per node, with more spikelets per branch; lemma pubescence is longer; rachillas are smooth; and anthers are longer. Poa pseudoabbreviata has upper culm leaf sheaths open 1/6-1/5 their length; open panicles with distinctly scabrous, elongate, capillary branches bearing 1-2 spikelets; short, 0.05-0.15 mm long lemma pubescence; scabrous rachillas; and anthers 0.2-0.6 mm long (recently revised, and mapped, Cody et al. 1990). Subspecies banffiana differs from Poa paucispicula in having shorter, more erect, less flexuous panicle branches that are frequently very sparsely scabrous, and have more numerous spikelets, and lemmas sometimes with puberulent intermediate veins (on at least one side) and a glabrous callus. It differs from P. abbreviata s. lat. in having panicles more open, with longer, more smooth branches, and high proportion of extravaginal branching. Subspecies banffiana occurs at the southern terminus of the range of Poa paucispicula. Porsild & Breitung collected both taxa at Bow River Pass (14875 and 14874 both P. paucispicula, 14873 a mixture of the two taxa) and Quartz Ridge (13832). Although subsp. banffiana agrees so closely with P. laza that it can not be reasonably distinguished, its characteristics are intermediate be- tween those of P. paucispicula and P. abbreviata, and further investigation may reveal it to be a stable hybrid between the latter species. One strongly webbed specimen from Oregon, and several webless specimens from Colorado, remain indistinguishable from P. lara except by their more narrow erect pan- icles, and these could be placed in P. abbreviata subsp. pattersoni except for their obvious extravaginal innovations and smooth or nearly smooth panicle branches! (Colorado [Clear Creek Co., Grays Peak, 13000’, 15 Aug 1885, Let- terman s.n. CAN!, Jul 1886, Letterman 6 US!, Mt. Evans, Summit Lake, 8 Sept 1956, Weber s.n. RSA!; Kingston Peak, Coz 487 US!]. Oregon [Wallowa Mts., Cusick 2493 CU!,CU!].) 6. Poa paucispicula Scribner & Merr., Contr. U.S. Natl. Herb. 13(3):69, fig. 15. 1910. TYPE: U.S.A. Alaska: Yakutat Bay, Hidden Glacier, 20 Jun 1899, Coville & Kearney 970 (HOLOTYPE: US 376352!). Poa merrilliana A. Hitchc., Amer. J. Bot. 2:309. 1915. Poa glacialis Scribner & Merr. [not Stapf. in 1906], Contr. U.S. Natl. Herb. Soreng: New taxa and hybrids in North American Poa 401 13(3):68. 1910. TYPE: U.S.A. Alaska: Hubbard Glacier, Cov- ille & Kearney 1077 (HOLOTYPE: US 376363!). Poa merrilliana was described as lacking a web on the callus, and thus could be confused with the new subspecies P. laza subsp. banffiana. The type collection of P. merrilliana is immature, but does have a distinct, if somewhat sparse, web, and is otherwise indistinguishable from P. paucispicula s. str. Most other material at US labeled or annotated by earlier workers as P. merrilliana is referable to P. pseudoabbreviata. 7. Poa secunda J.S. Pres] subsp. juncifolia (Scribner) R.J. Soreng, comb. nov. BASIONYM: Poa juncifolia Scribner, U.S.D.A. Div. Agrostol. Bull. 11:52. pl. 8. 1898. Poa juncifoha Scribner subsp. juncifolia (autonym), established by Keck in C.L. Porter, Flora Wyom., Part 3, Wyoming Agric. Exp. Sta. Bull. 418:22. 1964. TYPE: U.S.A. Wyoming: Sweet- water Co., Point of Rocks, Black Rock Springs, 13 Jul 1897, Nelson 3721 (LECTOTYPE designated in Hitchcock 1935, fig. 262: US 556860!; Isolectotypes: GH!,NY!,NY!,RM). Poa nevadensis Vasey ez Scribner, Bull. Torrey Bot. Club 10:66. 1883. TYPE: U.S.A. in 1877, Palmer 474 (Isotypes: NY!,NY!). [The geographical origin of this collection, which has been in doubt, may be Red Creek (now Paragonah), Iron Co., Utah, where Palmer col- lected 474-1/2 in June (25-) July, 1877, (P. fendleriana [Steudel] Vasey subsp. longiligula [Scribner & T. Will.) R.J. Soreng)]. Scrib- ner mistakenly believed Red Creek to be in Arizona (Scribner & Williams 1899, p. 3) (see MacVaugh 1956). In 1877, Palmer col- lected in southwestern Utah, southern Nevada, and northwest Ari- zona, his numbers are sequential-systematic, and there is no record of his collection notes. My supposition here is that 474 was one collection, subsequently split into two halves. Poa ampla Merr., Rhodora 4:145. 1902. TYPE: U.S.A. Washington: Steptoe, G.R. Vasey 3009 (Isotype: US!). The only previous application of the rank of subspecies among the approx- imately 50 taxa currently included in Poa secunda s. lat., was by D.D. Keck (1964). He proposed the name P. juncifolia subsp. porteri Keck, thereby creat- ing the autonym, subsp. juncifolia. According to ]CBN Article 57.3 (Greuter et al. 1988), the latter name has priority at this rank within any taxon in- cluding the type of P. junczfolia, unless an earlier epithet of the same rank is found. 402 PHYTOLOGIA volume 71(5):390-413 November 1991 Kellogg’s studies (1983, 1985) of Poa secundas. lat. demonstrate the futility of attempting to recognize microspecies in geographically parapatric/ecotonally sympatric, facultatively apomictic complexes. Her well considered conclusion that there is only one species agrees with interpretations of Marsh (1952) and Soreng (1985). This is also consistent with the species concept being applied in other such complexes in Poa (Tzvelev 1983; Soreng 1991). However, there is substantial variation within Poa secunda s. lat., which is indicated by the recognition of up to thirteen species and subspecies by Keck (unpublished manuscript), and by A.S. Hitchcock’s (1935) division of what he called eight species into two groups (Scabrellae and Nevadenses). The latter concept was more or less retained in the Vascular Plants of the Pacific Northwest (Hitch- cock et al. 1969) and the Intermountain Flora (Cronquist et al. 1977). An intermediate solution to what has been done by others is proposed, recognizing the two major variants as subspecies. When the recorded chromo- some numbers are graphed there are peaks at 2n = 84 and 2n = 63 (Almgard 1960; Armstrong 1937; Bowden 1961; Hiesey & Nobs 1982; Stebbins & Love 1941). [Subsp. secunda: 2n = 42, 44, 56x2, ca.61, 62, 63x5, 64, 66, 68, 70x4, 72, 74, 78x2, 80, 81x7, 82x6, 83, 84x23, 85x3, 86x5, 87, 88, 90x2, 91, 93, 94x2*, 98, 99, 104, ca.106 [82% > 70}. Subsp. juncifolia: 2n = 42, 44, 60, 62x7*, 63x29, 64x11, 65x2, 66, 68x2, 70x3, 78, 84, 96, 97, 100 [87% < 68. Two of these counts are new (*, 2n = 63, Nevada, Soreng 821, 2n ca. 94; Montana, Soreng & Spellenberg 1135). The hexaploid juncifolia count (2n = 42) was added after reassessment of the identity of the voucher spec- imen (Soreng 1991)|. The nanaploid and duodecaploid peaks correspond to consistent differences in ecology, anatomy, and gross morphology. Subspecies secunda (including Poa canby: [Scribner] Howell, P. gracillema Vasey, P. incurva Scribner, P. sandbergiit Vasey, P. scabrella [Thurb.| Benth.) has chromosome numbers centered around 2n = 84. It usually occurs in well drained soils of low salinity or alkalinity. The basal leaf blades are often thin and wither early, a correlate of having long cells that are fusiform in outline with thin, smooth walls. The lemmas are almost invariably softly to crisply puberulent, though in certain individuals or geographic regions the hairs may be extremely sparse, and easily overlooked. Ligules of leaves on sterile shoots are usually acute or acuminate, greater than 2 mm long, and may be glabrous or scabrous. Subspecies juncifolia has chromosome numbers centered around 2n = 63. It usually occurs in deep, frequently poorly drained, alkaline or saline, soils. The leaf blades are thickened and persistent, a corollary of having some proportion of the long cells rectangular in outline with more or less: thickened sinuous walls. The lemmas are glabrous or scabrous, or rarely sparsely and minutely crisp puberulent across the base (the latter puberulent phase is particularly evident on the high plains east of the Rocky Mountains where the taxon may intergrade with Poa arida Vasey). Ligules of leaves on sterile shoots are usually Soreng: New taxa and hybrids in North American Poa 403 truncate or obtuse, mostly less than 2 mm long, and scabrous (acute, longer, and sometimes glabrous in P. nevadensis Vasey forms). The substantial discontinuity in morphology is evident in that most col- lectors have distinguished individuals of these two extremes with reasonable success. They do breed true, and the differences are stable in transplant stud- ies. It is the consistent parapatric to ecotonally sympatric occurrence and high frequency of intermediates between these races (I estimate 10-20%) that makes consistent application of species rank impossible. 8. Poa secunda J.S. Pres] subsp. juncifolia (Scribner) R.J. Soreng x P. praten- sis L. Poa x lumosa Scribner & T. Will. (pro spec.)., U.S.D.A. Div. Agrostol. Circ. 9:5. 1899. TYPE: U.S.A. California: Mono Lake, 1866, Bolander |a rhizomatous plant] (HOLOTYPE: US 748920!). Poa x fibrata Swallen (pro spec.), J. Wash. Acad. Sci. 30:210. 1940. TYPE: U.S.A. California: Siskiyou Co., Shasta Valley, 3 mi. south of Grenada, 2600’, Wheeler 3629 (HOLOTYPE: US 1646953!; Iso- types: CAS! ,NY!). Apparent hybrids between Poa pratensis and P. secunda subsp. junczfolia are usually sterile anthered intermediates, and cccur over much of the geo- graphic range of overlap of the parents. J.T. Howell first suggested that this may be the origin of P. fibrata Swallen (pers. comm.). I have encountered several sites in California and Oregon, including those investigated in a study of in vivo P. fibrata for the California Fish & Game Department in 1986, at which both parents and intermediates occur. In addition, I have seen sporadic examples of the nothotaxon from British Columbia and Oregon in herbaria (CAS, OSC, UBC, US, V). Hiesey & Nobs (1982) present ample evidence of the crossing ability of these taxa, and at least temporary stabilization by gametophytic apomixis or vegetative reproduction in F, hybrids. Occurrence of counterfeit hybridization should not be ruled out (DeWet et al. 1984). The hybrids evidently have had multiple origins, and, as in the case of the type and some other localities of P. fibrata, they are sometimes ephemeral. An older name for Poa fibrata is P. limosa (there may be older names yet, but finding them would require exhaustive search of some 50 types). Rhi- zomatous plants like P. limosa still grow on the western shores of Mono Lake. Rather than apply a nothotaxon epithet, I suggest that these plants be referred to by their hybrid combination. 9. Poa pratensis L. [subsp. alpigena (Blytt) Hiit.] var. colpodea (Th. Fries) R.J. Soreng, comb. nov. BASIONYM: Poa stricta Lindeb. subsp. 404 PAY ToOlh OCA volume 71(5):390-413 November 1991 colpodea Th. Fries, Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 26:138. 1869. Poa alpigena (Blytt.) Lindman var. colpodea(Th. Fries) Scholand., Skr. Svalbard Ishavet 62:89. 1934. Poa rigens R. Br. subsp. colpodea(Th. Fries) D. Love, Taxon 17(1):89. 1968. Poa pratensis L. subsp. colpodea (Th. Fries) Tzvelev, Novosti Sist. Rast. 9:47. 1972. TYPE: Spitsbergen (“Liefdebay”). Poa alpigena (Blytt.) Lindman f. vivipara Rosch. in Komarov [nom. nud.|, Fl. U.S.S.R. 2:390. 1934. Tzvelev (1983) recognized the major ecological forms of the Poa praten- sts complex as subspecies. Poa pratensis s. lat. is a compilospecies, including many facultatively apomictic and reticulating lineages. The character combi- nations defining each of the major forms overlap and are too subtile and poorly documented to maintain species recognition. Variety colpodea is a viviparous form of the high arctic islands and coasts. It occurs at the northern edge of the geographic range of subsp. alpzgena and extends north of that (map. 7, Hultén 1964). This is well north of any other subspecies of P. pratensis in the arctic. The possibility that it is a hybrid between P. arctica R. Br. and P. praten- sts subsp. alpigena, is being studied (D. Goldman, S. Aiken, J.I. Davis, R.J. Soreng, unpublished data), as is the possibility that vivipary in var. colpodea may be a fixed or plastic response of subsp. alpigena to the more extreme northern climate. The morphology of the plants, especially of the sometimes normally developed spikelets within panicles, is typical of subsp. alpigena, and is here considered, as it has been by most recent authors, a subset of variation within the latter subspecies. 10. Poa hartzii R. Br. subsp. ammophila (A. Pors.) R.J. Soreng, comb. et stat. nov. BASIONYM: Poa ammophila A. Pors., Sargentia 4:12. 1943. TYPE: CANADA. Northwest Territories: District of Mackenzie, Cape Dalhousie, 70° 20’ N x 125° 55’ W, forming colonies on sandy hills back of coast, 7-14 Aug 1927, A.E. Porsild 2704 (HOLOTYPE: CAN!; Isotype: C!) (fig. 143 and map 158, Porsild & Cody 1980.). Habitat. Arctic, coastal hills, in sandy to clayey soils and stabilized dunes. Distribution and Specimens. Continental arctic Canada. Northwest Ter- ritories, District of Mackenzie, from Darnley Bay and Cape Parry Peninsula west to the Mackenzie River Delta (69° 20’-70° 20’ N x 124°-135° W) (A.E. & R.T Porsild 2154*, 2311-, 2422*, 2704-, 2827 (all CAN!), Cody & Ferguson 10363- DAO!, Scotter & Zoltai 25592a* DAO! (papillae present = *, absent = -). Porsild & Cody (1980) report it from one collection farther west, in Alaska, but this is probably Tieszen 854, which belongs to Poa hartzii subsp. alaskana R.J. Soreng). Soreng: New taxa and hybrids in North American Poa 405 Polunin (1959) and Scoggan (1978) submerged Poa ammophila A. Pors. into P. hartztz. Poa hartzii is a principally Nearctic, high latitude, psam- mophilic species, occurring on arctic islands from Wrangel Island eastward to Svalbard. Although it is traditionally placed in sect. Abbreviatae, its affinities to other species of that section are not strong, and hybrid origins from sev- eral different parents have been postulated (Edmondson 1980; Polunin 1959; Tzvelev 1983). Because P. hartziz has extravaginal branching, long anthers, somewhat weakly keeled lemmas, and diffuse callus hairs, I suggest it be in- cluded in sect. Secundae Marsh ez R.J. Soreng. (That section may have had a hybrid origin, but it is distinct from sect. Abbreviatae s. str.). Unlike most Poa (except sections Andinae Nicora, Secundae, and Arctopoa [Griseb.] Tzvelev), P. hartzii has villous hairs distributed around the callus and a smooth transi- tion of the callus into the base of the lemma (fig. 3), the plesiomorphic state in the tribe Poeae (Soreng 1990). Most Poa have the derived state of hairs arising from the dorsal side of the callus in a single tuft and a shght sulcus on either side of the web origin of the somewhat laterally compressed callus. (The sulcus and compression are also present in derived species that have lost the web.). The almost invariably sterile, early abortive anthers (lacking any pollen), high, dysploid chromosome numbers (2n = 63-70), and lack of obvious morphological variation, in plants from the Canadian Archipelago, arctic coast of Quebec (Cayouette 1984, the first report for the mainland), and Greenland, confirm that P. hartziiis predominantly obligately and autonomously apomic- tic. It has chloroplast-DNA restriction sites diagnostic for Poa (R. Soreng, J-I. Davis, & S. Aiken, unpublished data). Porsild & Cody (1980) maintain that plants from the western continen- tal Canadian arctic are not Poa hartzii, but a distinct species, P. ammophila. These plants have more normally developed, sometimes pollen bearing anthers (1.5-1.8, vs. aborted and 0.8-1.5 mm long). In addition, they often have papil- lae on the long cells of the adaxial surface of the leaf blades. They also have shorter ligules (1.5-3.0 mm long), shorter lemmas (3.0-4.6 mm long), elongate trachilla internodes, consistently smooth panicle branches (rarely with a few scabers), a greater tendency to branch intravaginally, and more drab spikelets as compared to P. hartzii s. str. (ligules 2-7 mm long, lemmas 3.9-5.3 mm long, panicle branches often scabrous, mixed or predominantly extravaginal branching, and lustrous spikelets). The most consistent characteristics distin- guishing subsp. ammophila are the lack of hairs on the callus, and the shorter, fine, crisply puberulent hairs of the lemmas (< 0.3 mm long), versus callus hairs present and at least some callus or basal hairs of the lemmas being (0.5-) 0.8-2.0 mm long, and villous. Although the pubescence characteristics are con- stant elsewhere in the ranges of the two taxa, there is one collection (Porsild’s “luxuriant form,” 2706 CAN!) from near the type locality of subsp. ammophzla, which has callus hairs, long lemma hairs, ligules 3 mm long, and lacks papillae. This (and Parmellee 3214 from 100 km east of the range of subsp. ammophila; 406 PHYPYT OLOGIA volume 71(5):390-413 November 1991 Cc Figure 3. Spikelets of Poa hartzii R. Br. subspp. and P. secunda J.S.Presl subsp. secunda. a. Poa secunda subsp. secunda (Corcoran 9). b. subsp. hartziz (Porsild 18636). c. subsp. ammophila(A. Pors.) R.J. Soreng (Porsild & Porsild 2154). d. P. hartzit subsp. alaskana R.J. Soreng (Murray & Johnson 7159; HOLOTYPE: C!). Soreng: New taxa and hybrids in North American Poa 407 69° 35’ N x 120° 44’ W), is clearly indistinguishable from typical P. hartzii and has caused confusion about recognition of subsp. ammophila. With the exception of this one plant, from a possibly mixed or polymorphic population, as P. hartzi, there is no problem distinguishing these taxa. However, an additional problem occurs with regard to inland plants col- lected from around Great Bear Lake, identified as Poa ammophila by Porsild & Cody (1980) (Porsild 17007- CAN!, and Corcoran 9- DAO!). The Corcoran specimen has quite scabrous branches and a habit typical of P. secunda and is best placed in P. secunda subsp. secunda (also identified by D.D. Keck as P. canbyi [= P. secunda]). In the absence of papillae on the leaf blades (which are variable in occurrence among plants of subsp. ammophila from the coast, but also occur in taxa of Poa sect. Secunda subsect. Halophytae V.L. Marsh ez R.J. Soreng), there is no combination of characters by which to distinguish 17007 from subsp. ammophila or P. secunda. Poa secunda s. lat. is usually distinguished from all other Poa by having weakly keeled lemmas. Although the Great Bear Lake specimens have weakly keeled lemmas, so do some spec- imens of coastal subsp. ammophila (also noted by Porsild); keels of lemmas of P. hartzi s. str. are not always well defined either. The habits of the plants from the arctic coast and Great Bear Lake regions vary considerably, from spreading and tufted as in P. hartzws. str., to erect and tightly tufted as in P. secunda. Poa secunda is native as far north as 63-64° N in eastern Alaska, the Yukon and western District of Mackenzie, directly south of the range of subsp. ammophila, and often reaches into the alpine. Whether P. secunda is native at Great Bear Lake (Sawmill Bay, 65° 43’ N, Corcoran 9) is not known, but it is entirely possible that its range extends farther north than currently known, into the Mackenzie River delta. Eventually subsp. ammophila may be proven to be a hybrid between P. secunda and P. hartzit. 11. Poa hartzii R. Br. subsp. alaskana R.J. Soreng, subsp. nov. TYPE: U.S.A. Alaska: 70° 45’ N, 156° 30’ W, Mead River, forming tufts in sand dunes of point bars, 4 Aug 1980, D. Murray & Johnson 7153 (HOLO- TYPE: C! plant no. 1 [no. 2 = Deschampsia caespitosa (L.) Beauv. subsp. brevifolia (R. Br.) Tzvelev]; Isotype: ALA) (Fig. 4). A P. hartzu R. Br. subsp. hartzii et subsp. ammophila(A. Pors.) R.J. Soreng lemmatibus 5.5-7.0 mm longis, et rhachillis 1.5-2.0 mm longis, differt. Perennial; hermaphroditic; cespitose, or becoming stoloniferous in sandy soils. Culms loosely tufted, 20-45 cm tall, innovations intra and extravaginal. Upper culm leaf sheath margins fused 1/7-1/5 the length; ligules 5-7 mm long, smooth to sparsely scabrous abaxially; blades 1-3 mm wide, firm, folded and 408 PHY TOLOG FA volume 71(5):390-413 November 1991 Figure 4. Poa hartzw R. Br. subsp. alaskana R.J. Soreng habit (Murray & Johnson 7153; HOLOTYPE: C!). Soreng: New taxa and hybrids in North American Poa 409 inrolled on the margins, smooth abaxially, more or less scabrous to hispidulous adaxially on and between the veins, lacking papillae. Panicle 7-12 cm long, lanceolate, the branches smooth to moderately scabrous, sulcate. Spikelets with 3-5 florets, 5-7 mm long; glumes broadly hyaline, 3 veined, the second glume frequently exceeding the lower lemma; callus with villous hairs 1-2 mm long around the base; lemmas 5.5-7.0 mm long, thin, sparsely villous on the keel and marginal veins and between them, the margins broadly hyaline; palea keels sparsely scabrous to pilose below; rachillas glabrous or sparsely villous, the longest internodes 1.5-2.0 mm long. Flowers perfect; anthers 2.2-2.8 mm long, appearing fertile. Habitat. High arctic coastal plain, in sands along inland waterways, 100- ca. 850 m. Distribution and Specimens. U.S.A. Alaska: north slope (Mead River, vicinity of Atkasook village, 70° 39’ N, 157° 15’ W, Murray & Johnson 7158 ALA,C!, Komarkova & Duffy 595 COLO, west side of river, 10 mi. north of Atkasook village, in sand dunes, 16 Aug 1959 (1953?), Cantlon 4763 CAN!, 4782 CAN!; Mead River delta, Rothe 45 ALA; Lake Peters [2800’, 69° 20’ N, 145° 02' W], up Bear Creek, late snow area in moss bed [roots sandy], 3 Aug 1966, Treszen 854 CAN!). (I have not seen the ALA or COLO specimens, which are cited from location data sent by D. Murray.). Plants of Poa hartziz from the Mead River, northwest Alaska, and Lake Peters, northeast Alaska, are distinguished as subsp. alaskana. These are ro- bust (20-45 cm tall), stoloniferous, principally intravaginally branching, have ligules 5-7 mm long, and well developed, pollen bearing anthers 2.0-2.5 mm long. David Murray independently noted some of these distinctions (unpub- lished manuscript). Plants of subsp. hartziz have mostly abortive (empty) anthers, less than 1.5 mm long, ligules 2-5 mm long, shorter stature (15-25 cm tall), a cespitose habit, and proportionally more extravaginal branching. In addition, the Alaskan plants have abundant callus hairs about 2 mm long, and longest lemmas and rachilla internodes that are longer than in subsp. hartzu (5.5-7.0 and 1.5-2.0 mm long, versus 3.9-5.3 and 0.8-1.5 mm long, re- spectively). Like specimens of subsp. hartzii checked, the Alaskan plants lack papillae on the leaf blades. The best characters to discriminate between these geographically isolated populations and other P. hartzii are fully developed versus early abortive anthers, and the slightly larger looser habit and more vigorous growth of the spikelets. These characters quite likely will be found to overlap. The Alaskan plants may be merely sexually reproducing popula- tions within an otherwise predominantly, obligately apomictic species. The degree of morphological differentiation is consistent with recognition at the subspecific level. 410 PHY POLO Gra volume 71(5):390-413 November 1991 12. Deschampsia caespitosa (L.) Beauv. subsp. brevifolia (R. Br.) Tzvelev in Tolmachey, Fil. Sev.-Vost. Europ. Chasti SSSR 1:141. 1974. Poa evagens Simmons, Report Second Norw. Arctic Exped. in the Fram 1898-1902. Kristiana 2:165-166. 1906. TYPE: CANADA. Ellesmerlandiae meridionalis, Fram Fjord 76° 23’ N, 81° 30’ W, Simmons 4267 (LD!). The name Poa evagens has been applied to material of P. hartziw subsp. hartzii and subsp. ammophila by several botanists. An examination of the type specimen of P. evagens by myself and S. Aiken revealed that the taxon belongs to Deschampsia, not Poa; the early deciduous awns probably having contributed to the confusion. The following infrageneric taxa in Poa, proposed by Soreng (1991), are emended here. Three were incorrect according to the JCBN (Greuter et al. 1988). Subgeneric taxa other than those that include the type of the genus in which they are described, are not autonymic; each rank, even though repetition of the epithet is recommended, is independent of those in which they are included and must be separately validated (see Articles 6.8, 57 and extensions thereof). Thus taxa 14 and 17 need their types and authors repeated. Creation of new subgeneric taxa also requires that the genus be explicitly stated in the protocol (Articles 21.1 and 37). Taxa 13, 15, and 16, but not 14, 17 and 18, were validated by their usage in text and tables in Soreng (1991). 13. Poa L. subgen. Poasect. MadropoaR.J. Soreng, Syst. Bot. 16(3):512-513. 1991. TYPUS: Poa piperi A. Hitchc. 14. Poa L. subgen. Poa sect. Madropoa R.J. Soreng subsect. Madropoa R.J. Soreng, Syst. Bot. 16(3):513. 1991. TYPUS: Poa piper: A. Hitche. Emend. 15. Poa L. subgen. Poa sect. Madropoa R.J. Soreng subsect. Epiles A. Hitchc. ez R.J. Soreng, Syst. Bot. 16(3):512-513. 1991. TYPUS: Poa epilis Scribner. 16. Poa L. subgen. Poa sect. Secundae V.L. Marsh ez R.J. Soreng, Syst. Bot. 16(3):513, 523. 1991. TYPUS: Poa secunda J.S. Presl. 17. Poa L. subgen. Poa sect. Secundae V.L. Marsh ez R.J. Soreng subsect. Secundae R.J. Soreng, Syst. Bot. 16(3):523. 1991. TYPUS: Poa secunda J.S. Pres]. Emend. 18. Poa L. subgen. Poa subsect. Halophytae V.L. Marsh ez R.J. Soreng, Syst. Bot. 16(3):523. 1991. TYPUS: Poa unilateralis Scribner. Emend. Soreng: New taxa and hybrids in North American Poa 411 ACKNOWLEDGMENTS Thanks are extended to Nancy Soreng for the illustrations, Susan Aiken, Jerrold Davis, and Melissa Luckow for careful readings of the manuscript, Susan Aiken for providing live material of arctic Poa and assisting me in the annotation of most of the CAN Poa, Rupert Barneby for help with the Latin, David Murray for comments on the Alaskan taxa, John Strother for help with nomenclature, David Keck for his annotation lists of Poa, the curators of C, CAN, CU, DAO, LD, NY, RSA, UBC, US, and V for loans of specimens consulted in this study, and Vernon Leroy Marsh, wherever he may be, for his dissertation on Poa. LITERATURE CITED Almgard, G. 1960. Experiments with Poa I. Studies of Poa longifolia Trin. as a fodder grass and as a component in interspecific hybrids. Kongl. Svenska Lantbr.- Acad. Ann. 26:77-119. Armstrong, J.M. 1937. A cytological study of the genus Poa L. Canad. J. Res. 15:281-297. Bowden, W.M. 1961. Chromosome numbers and taxonomic notes on north- ern grasses IV. Tribe Festuceae: Poa and Puccinellia. Canad. J. Bot. 39:123-138. Cayouette, J. 1984. Additions et extensions d’aire dans la flore vasculaire du Nouveau-Quebec. Naturaliste Canad. 111:263-274. Cody, W.J., S.J. Darbyshire, & C.E. Kennedy. 1990. A bluegrass, Poa pseu- doabbreviata Roshev., new to the flora of Canada, and some additional records from Alaska. Canad. Field-Naturalist 104:589-591. Cronquist, A., A.H. Holmgren, N.H. Holmgren, J.L. Reveal, P.K. Holmgren. 1977. Intermountain Flora, vol. 6. Columbia University Press, New York, New York. DeWet, J.M.J., C.A. Newell, & D.E. Brink. 1984. Counterfeit hybrids be- tween Tripsacum and Zea (Gramineae). Amer. J. Bot. 71:245-251. Edmondson, J.R. 1980. Poa L. In T.G. Tutin, V.H. Heywood, N.A. Burges, D.M. Moore, D.H. Valentine, S.M. Walters, & D.A. Webb, [eds.], Flora Europaea, vol. 5, 159-167. Cambridge University Press, Cambridge, U.K. 412 Pon vel OL O-G tA volume 71(5):390-413 November 1991 Greuter, W., et al. (eds). 1988. International Code of Botanical Nomencla- ture. Regnum Veg. 118:1-328. Hiesey, W.M. & M.A. Nobs. 1982. Experimental studies on the nature of species VI. Interspecific hybrid derivatives between facultatively apomic- tic species of bluegrasses and their responses to contrasting environ- ments. Carnegie Institution of Washington Publication 636, Washington, D.C. Hitchcock, A.S. 1935. Manual of the Grasses of the United States. U.S.D.A. Misc. Pub. 200. U.S. Govt. Print. Off., Washington, D.C. Hitchcock, C.L., A. Cronquist, M. Genter & J.W. Thompson. 1969. Vas- cular Plants of the Pacific Northwest. University of Washington Publi- cations in Biology 17:648-683, Seattle, Washington. Hultén, E. 1964. The circumpolar plants. I. Kéngl. Svenska Vetenskapsakad. Handl., Fjarde Ser. 8(5):1-240. . 1974. A Flora of Alaska and Neighboring Territories. Stanford University Press, Stanford, California. Keck, D.D. Unpublished typescript, untitled [a revision of Poa of the con- tiguous western United States], 65 pages, circa 1949. Copies at US and WIS. . 1964. Poa. Pp. 17-21, in, A Flora of Wyoming, part 3, by C.L. Porter. Flora Wyom., Part 3, Wyoming Agric. Exp. Sta. Bull. 418. Kellogg, E.A. 1983. A biosystematic study of the Poa secunda complex. Ph.D. Dissertation. Harvard University, Cambridge, Massachusetts. . 1985. A biosystematic study of the Poa secunda ons J. Arnold Achar: 66:201-242. Love, A., D. Love, & B.M. Kapoor. 1971. Cytotaxonomy of a century of Rocky Mountain Orophytes. Arctic and Alpine Res. 3(2):139-163. Love, A. & D. Love. 1975. Cytotaronomical Atlas of the Arctic Flora. J. Cramer, Leutershausen, Germany. MacVaugh, R. 1956. Edward Palmer, Plant Explorer of the American West. University of Oklahoma Press, Norman, Oklahoma. Marsh, V.L. 1950. A taxonomic revision of the genus Poa of the United States and southern Canada. Ph.D. Dissertation. University of Washington, Seattle, Washington. Soreng: New taxa and hybrids in North American Poa 413 . 1952. A taxonomic revision of the genus Poa of the United States and southern Canada, part 1. Amer. Midl. Naturalist 47:202-250. Nannfeldt, J.A. 1935. Taxonomical and plant-geographical studies in the Poa laza group. Symb. Bot. Upsal. 1(5):1-316. Polunin, N. 1959. Circumpolar Arctic Flora. Clarendon Press, Oxford, U.K. Porsild, A.E. & W.J. Cody. 1980. Vascular plants of continental Northwest Territories, Canada. Natl. Mus. Nat. Sci. NM92-71/1979, Natl. Mus. Canad., Ottawa, Ontario. Scoggan, H.J. 1978. The Flora of Canada, part 2. Natl. Mus. Nat. Sci. 70(2), Natl. Mus. Canad. Scribner, F.L. & T. Williams. 1899. Poa fendleriana and its allies. U.S.D.A. Div. Agrost. Circ. 10:1-6. Soreng, R.J. 1985. Poa in New Mexico, with a key to middle and southern Rocky Mountain species (Poaceae). Great Basin Naturalist 45:395-422. —_____. 1990. Chloroplast-DNA phylogenetics and biogeography in a retic- ulating group: Study in Poa(Poaceae). Amer. J. Bot. 77(11):1383-1400. . 1991. Systematics of the “Epiles” Group of Poa (Poaceae). Syst. Bot. 16:507-528. Stebbins, G.L. & R.M. Love. 1941. A cytological study of California forage grasses. Amer. J. Bot. 28:371-382. Tzvelev, N.N. 1983. Grasses of the Soviet Union. {English translation of 1976 Russian ed.] New Delhi: for Smithsonian Institution by Amerind Publishing Company. Phytologia (November 1991) 71(5):414-415. FIRST REPORT OF ERIGERON VELUTIPES (ASTERACEAE) FROM THE UNITED STATES Guy L. Nesom and Marc A. Baker Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. and Department of Botany & Microbiology, Arizona State University, Tempe, Arizona 85287 U.S.A. ABSTRACT General collections from southern Arizona by the second author have included plants of Erigeron velutipes Hook. & Arn., a species pre- viously known only from Mexico. KEY WORDS: Asteraceae, Erigeron, range extension UNITED STATES. Arizona: Santa Cruz Co., ca. 4 km ESE of Black Peak, just N of fork in road 39, T22S R11E SE 1/4 s.31; moist soil in edge of seep on both sides of road, 27 May 1991, M.A. Baker 8431 (ASU,TEX). Erigeron velutipes Hook. & Arn. has been collected in relative abundance in México from the states of Michoacan and Jalisco northward along the Pacific coast to Sonora and inland in the northernmost part of its range to western Chihuahua. The closest known collections to the Arizona locality have been made about 500 kilometers to the south in eastcentral Sonora. The Arizona plants and those from around Basaseachic, Chihuahua, are different from most others of the species in their primarily basal stem branching and heads on long, naked, stiffly erect peduncles, but the species includes a great deal of other variability in habit. In its annual duration, nodding buds, distinctive vestiture, narrow ligules drying blue, and small achenes, Erigeron velutipes is similar to E. lobatus A. 414 Nesom & Baker: First report of Erigeron velutipes in U.S. 415 Nelson and LE. piscaticus Nesom. The last two are Arizona natives, and details of contrast among these three species are provided in the key in Nesom (1989). ACKNOWLEDGMENTS We thank Dr. B.L. Turner and Lindsay Woodruff for their review and comments. LITERATURE CITED Nesom, G.L. 1989. A new species of Erigeron (Asteraceae: Astereae) from Arizona. Phytologia 67:304-306. Phytologia (November 1991) 71(5):416-419. A NEW SPECIES OF ERIGERON (ASTERACEAE: ASTEREAE) FROM NORTHWESTERN NEW MEXICO Guy L. Nesom Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A new species of Erigeron is described from the Zuni Mountains of McKinley County, New Mexico: E. sivinskii. It is most closely related to E. consimilis and E. nematophyllus. KEY WORDS: Erigeron, Astereae, Asteraceae, New Mexico Erigeron sivinskii Nesom, sp. nov. TYPE: UNITED STATES. New Mex- ico: McKinley Co., Zuni Mts., E side of Six-Mile Canyon, T14N, R15W, Sec. 18, NW 1/4, 2200 m, several hundred plants on steep, eroded shale slopes of the Chinle Formation, with an abundance of barite nodules on the surface, associated with pinyon-juniper, Swertza radiata, Cryptan- tha fulvocanescens, and Chrysothamnus nauseosus, 3 Jun 1991, Robert Sivinski & K. Lightfoot 1687 (HOLOTYPE: UNM; Isotype: TEX!). Erigeronti consimilo Cronq. similis sed differt caulibus foliatis, caulibus foliis phyllariisque sparsim pubescentibus, capitulis mi- noribus radiis paucioribus, et acheniorum ciliis marginatis leniter evolutis. Perennial herbs arising from a thick taproot with numerous, short (1-3 cm), ascending-erect caudex branches, the upper portion of these with adherent, old leaf bases. Stems erect, 5-8 cm tall, unbranched, green, sparsely strigose with white, stiff, filiform, closely appressed trichomes evenly distributed and of even length (0.2-0.3 mm long), eglandular. Leaves green, similar in vestiture to the stems, arising in dense basal clusters from the caudex apices, erect to ascending erect, linear, mostly 12-30 mm long, 0.5-0.8 mm wide, definitely thickened, slightly flaring at the very base, the cauline strictly ascending and continuing relatively unreduced in size half way to nearly all the way up the 416 Nesom: New Erigeron from New Mexico 417 stems. Heads solitary, terminal, cupulate, 10-14 mm wide (pressed); phyllaries in 2-3 subequal series, 5-6 mm long, linear-lanceolate with acuminate apices, relatively thin herbaceous, narrowly lanceolate with attenuate-filiform apices, minutely but prominently granular-glandular, the outer also sparsely pilose with a few (ca. 10-20 per phyllary), crisped-spreading hairs arising from along the midregion. Ray flowers 21-33, the corollas 7-9 mm long, ligules 1.0-1.5 mm wide, white with a lilac midstripe, distinctly coiling from the apices with maturity. Disc corollas 3.0-3.8 mm long, narrowly funnelform, not strongly inflated or indurated, glabrate; style branches 0.5-0.6 mm long, the collecting appendages deltate to shallowly triangular, 0.1-0.2 mm long. Achenes 2(-3) nerved, narrowly oblong, 2.8-3.1 mm long, the faces glabrous, the margins very sparsely ciliate; pappus of 21-27 barbellate bristles, with a prominent outer series of setae 0.4-0.6 mm long. Known only from the area of the type locality, named for its collector, Robert C. Sivinski, endangered species botanist for the New Mexico Forestry Division. Additional collections examined: UNITED STATES. New Mexico: McKin- ley Co., Zuni Mts., Six-Mile Canyon (type locality): 25 May 1990, Robert Swinski & A. Cully 1425(UNM); T14N, R15W, Sec. 7, SW 1/4, several dozen plants on hard, sodic shale outcrop of Chinle Formation, in pinyon-juniper with Sarcobatus vermiculatus and Hilaria jamesii, 3 Jun 1991, R. Sivinski & K. Lightfoot 1686 (TEX, duplicate at UNM not seen). Plants of Erigeron sivinskit Nesom are very similar and clearly closely re- lated to those of E. conszmilis Cronq., which occurs in western Colorado, east- ern Utah, and northeastern Arizona. Plants of both taxa produce a taproot with thick caudex branches, linear, erect, and densely massed basal leaves, solitary heads, ray flowers with coiling ligules, achenes with glabrous faces and ciliate margins, and a pappus with a prominent outer series of setae. The vestiture of minute, white, closely appressed trichomes is particularly distinc- tive and, with the coiling ligules, is a critical factor in the placement of these species in sect. Wyomingia (A. Nels.) Cronq. (Cronquist 1947; Nesom 1989). The nature of the relationship between the typical members of this section, however, and the “FE. compactus group,” of which E. szvinskii and E. consimilis are members, remains unclear. On the basis of their strong similarity and relatively less marked disparity, Blake (1950) considered Erigeron consimilis to be only varietally distinct from E. compactus §.F. Blake, which occurs from eastcentral California through Nevada to western Utah. These two taxa also were recently treated as varieties of a single species by Welsh (1983), although neither he nor Blake noted the oc- currence of intermediates. According to data given by Welsh (1983) and Albee et al. (1988), the two are strongly allopatric in Utah where their ranges ap- proach each other; Welsh characterized their ranges in Utah as “Great Basin” (E. compactus) vs. “Colorado Drainage system” (E. consimilis). Each of the two taxa is geographically widely distributed, and although the populations of 418 PAY TOsO°GPA volume 71(5):416-419 November 1991 each appear to be relatively widely isolated, both taxa appear to be constant in the morphological features that distinguish them. Although E. compactus is closely similar to E. consimilis, Cronquist’s initial characterization (1947) of the latter as a separate species remains justifiable. Erigeron sivinski occurs near the southeastern corner of the range of E. consimilis and can be regarded as a third segment of the E. compactus-E. consimilis lineage. Compared to E. sivinsku, however, E. consimilis produces scapose stems, densely invested stems, leaves, and phyllaries, shorter leaves (5-20 mm long), larger heads (15-20 mm wide) with more numerous rays (30- 55 per head), and achenes with strongly developed marginal cilia. Even at the closest geographical approach of the two taxa, there is no evidence of intergradation. Further, several features of the McKinley County plants are more similar to another closely related species of the E. compactus group of sect. Wyomingia (Nesom 1989), E. nematophyllus Rydb., which ranges from southwestern Wyoming through western Colorado and into adjacent Utah. As in E. nematophyllus, the vestiture of E. sivinskit is sparse and the leaves are basal as well as disposed along at least the lower portion of the stem. Thus, while it shares distinctive and probably specialized features of leaf shape and achene vestiture with E. consimilis, E. sivinskw is more similar to E. nematophyllus in leaf disposition and vegetative vestiture. The New Mexico population occurs outside of the geographical range of both its closest relatives, and to best account for its somewhat intermediate morphological position between them, treatment as a separate species is warranted. Erigeron untermannit Welsh & Goodrich and E. carringtoniae Welsh, which were tentatively included in the E. compactus group (Nesom 1989), are differ- ent in vestiture, leaf shape, and achenal pubescence from the species discussed above and are more distantly related, if they belong with the group at all. The following key distinguishes E. sivinskii and its closest relatives. 1. Leaves mostly (2-)4-8 cm long, often apically broadened (1.5-2.2 mm wide), the lower margins ciliate; persistent portion of old basal leaves relatively long and slender-fibrous; achene faces pubescent, marginal cilia weakly developed or undifferentiated; outer pappus weakly developed. ........ ee ee eS eR E. nematophyllus 1. Leaves mostly 1-3 cm long, usually strictly linear (0.5-1.4 mm wide), ecil- late; persistent portion of old basal leaves short and broad; achene faces completely glabrous, marginal cilia present; outer pappus of numerous, prominent setae... 2. osc e sce coe es eae pee 1 cane 6c eee (2) 2. Stems, leaves, and phyllaries green, sparsely short strigose; stems with relatively unreduced leaves on at least the lower half; phyllaries relatively thin herbaceous; achene margins sparsely ciliate. ...... ai evra) eth olga Dita bictnh ead tw aialh eae el ald Baten cee E. sivinski Nesom: New Erigeron from New Mexico 419 2. Stems, leaves, and phyllaries gray-green, densely short strigose; stems essentially scapose, sometimes with a few, small, scattered bracts; phyllaries distinctly thickened along the margins; achene UGE CANIS, MENSELY, Cilvaten ace ciy cette ies aes hc Seelclesteiorn aster anae Pe (3) 3. Phyllaries spreading-hispidulous with thick based trichomes; heads 15-20 Wa VHC IED Soba Soe OMeOD cme ope nODot ecrscotGs E. consimuilis 3. Phyllaries strigose with thin based trichomes; heads 8-15 mm wide; rays JUSS 8 is est eA I ee a OR Sr Care Aa UT ie Pesach es E. compactus ACKNOWLEDGMENTS I appreciate the review and comments by Dr. B.L. Turner and Dr. C.R. Werth. Dr. Tim Lowrey brought the McKinley County collection to my at- tention, and Robert Sivinski sent additional material for study. LITERATURE CITED Albee, B.J., L.M. Shultz, & S. Goodrich. 1988. Atlas of the Vascular Plants of Utah. Utah Mus. Nat. Hist. Occ. Publ. No. 7. Blake, S.F. 1950. A new combination in Erigeron. Leafl. West. Bot. 6:71. 1950. Cronquist, A. 1947. Revision of the North American species of Erigeron, north of Mexico. Brittonia 6:121-302. Nesom, G.L. 1989. Infrageneric taxonomy of New World Erigeron (Com- positae: Astereae). Phytologia 67:67-93. Welsh, $.L. 1983. Utah flora: Compositae (Asteraceae). Great Basin Nat. 43:179-357. Phytologia (November 1991) 71(5):420-422. NEW COMBINATIONS IN POTENTILLA AND HORKELIA (ROSACEAE) IN CALIFORNIA Barbara Ertter University and Jepson Herbaria, University of California, Berkeley, California 94720 U.S.A. ABSTRACT Three new combinations are proposed for use in the upcoming re- vised Jepson Manual of California plants. Potentilla rimicola (Munz & I.M. Johnston) Ertter replaces P. wheeleri subsp. rimicola, while Horkelia californica subsp. dissita (Crum) Ertter and subsp. fron- dosa (E.L. Greene) Ertter replace H. elata and H. frondosa respectively. KEY WORDS: Potentilla, Horkelia, Rosaceae, California, taxon- omy As a result of preparing the treatment of herbaceous Rosaceae for the forthcoming The Jepson Manual: Higher Plants of California, several new combinations were found to be necessary. Key characters and descriptions will be found in the Manual and are therefore not included here. Potentilla rimicola (Munz & I.M. Johnston) Ertter, comb. et stat. nov. BASIONYM: Potentilla wheeler: S. Wats. var. rimicola Munz & I.M. Johnston, Bull. S. Calif. Acad. Sci. 24:18. 1925. The Potentilla wheeleri complex (Rydberg’s Subviscosae) includes a series of biogeographically interesting taxa on isolated mountain ranges in Califor- nia, Arizona, New Mexico, and northern México. Typical P. wheeler: occurs from the southern Sierra Nevada to northern Baja California. Although the distinctiveness or circumscription of possible segregates var. paupercula Jep- son (Mount San Gorgonio, San Bernardino County, California), P. luteosericea Rydb. (northern Baja California), or P. viscidula Rydb. (southern Arizona) are thus far uncertain, my initial studies of the complex indicate that at least var. rimicola is worth recognizing at the species level. A new combination is therefore needed at this time for the Jepson Manual. 420 Ertter: New combinations in Potentilla and Horkeha 421 The most distinctive difference between Potentilla rimicola and other mem- bers of the complex is that it grows in crevices of vertical rock faces; morpho- logical differences include pedicel length and characters of the fruit. Potentzlla rimicola occurs only in the San Jacinto Mountains of Riverside County, Cali- fornia, and the Sierra San Pedro Martir of northern Baja California, México. Although P. wheeleri occurs in the same mountain ranges, no intergradation between the taxa has been found. The petrophytic habit is very unusual in Potentilla s. str. but characterizes many species of Jvesia Torrey & A. Gray (Ertter 1989). These same species of [vesia differ from members of the P. wheelert complex in having pinnately rather than palmately divided leaves, but otherwise share several other intrigu- ing biogeographical and morphological similarities, including glandular hairs, tidged seeds, and + recurved pedicels. Horkelia californica Cham. & Schldl. subsp. dissita (Crum) Ertter, comb. et stat. nov. BASIONYM: Potentilla elata E.L. Greene var. dissita Crum an Jepson, Fl. Calif. 2:197. 1936. Potentilla elata E.L. Greene, Pittonia 1:100. 1887. Potentilla califor- nica (Cham. & Schldl.) E.L. Greene var. elata (E.L. Greene) E.L. Greene, Fl. Franciscana 1:66. 1891. Horkeha elata (E.L. Greene) Rydb., Bull. Torrey Bot. Club 25:54. 1898. Horkelia glandulosa Eastw., Bull. Torrey Bot. Club 32:195. 1905. Horkelia californica Cham. & Schldl. subsp. frondosa (E.L. Greene) Ert- ter, comb. et stat. nov. BASIONYM: Potentilla frondosa E.L. Greene, Pittonia 1:300. 1889. Horkelia frondosa (E.L. Greene) Rydb., Bull. Tor- rey Bot. Club 25:54. 1898. Potentilla californica (Cham. & Schldl.) E.L. Greene var. frondosa (E.L. Greene) Jepson, Man. Fl. Pl. Calif. 494. 1925. Although most treatments during the last few decades treat Horkelia (or Potentilla) californica, elata, and frondosa as distinct species, this glosses over the large number of intermediate specimens that do not fall conveniently into one taxon or another. Numerous collections from the North Coast ranges of California are particularly frustrating in their intermediacy between typical H. elata and typical H. californica (e.g., size of leaflets and degree of lobing). Intermediates in leaf morphology also blur the distinctiveness of H. frondosa, supporting the treatment of all three as infraspecific taxa. Unfortunately, the familiar epithet “elata” must be replaced, in that Po- tentilla elata E.L. Greene is a later homonym of P. elata Salisb. (Prodr. Stirp. Chap. Allerton 1796, p. 362). Britten (1916) argues convincingly that Salis- bury’s numerous names, although largely superfluous, should not be ignored. 422 PH ROW OIGHA volume 71(5):420-422 November 1991 Subspecies are used rather than varieties, both to parallel usage elsewhere in Horkelia and to avoid problems caused by the questionable identity of Poten- tilla glandulosa {| incisa Lind]. The possible synonymy with subsp. frondosa of this name and the equally problematical Horkelia grandis Hook. & Arn. is discussed in detail by Keck (1938). LITERATURE CITED Britten, J. 1916. The plants of Salisbury’s “Prodromus” (1796). J. Botany 54:57-65. Ertter, B. 1989. Revisionary studies in Jvesia (Rosaceae: Potentilleae). Syst. Bot. 14:231-244. Keck, D.D. 1938. Revision of Horkelia and Ivesia. Lloydia 1:75-142. 1938. Phytologia (November 1991) 71(5):423-427. BOOKS RECEIVED Anales del Instituto de Biologia, serie Botanica, vol. 61(1). Fernando Chiang C. (ed.). Coordinador de la Biblioteca del Instituto de Biologia, Uni- versidad Nacional Autonoma de México, Apartado Postal 70-233, 04510 Mexico, Distrito Federal, México. 1991. 63 pp. Price unknown (paper). ISSN 0374-5511. The present volume continues this important series on Mex- can botany. It contains a diversity of papers on botanical sub- jects. These range from description of a new species of Jatropha and augmentation of the description of a species of Quercus, to effects of plastic mulch on fungal growth in bean cultivation and difficulty of determining presence of mycotoxins in grain sorghum shipments. Also in this issue are papers summarizing the taxon- omy, alkaloid chemistry, and ethnobotany of Datura lanosa, and a listing of the cryptogamic and graminaceous type specimens in the National Herbarium of México. Annual Review of Phytopathology, volume 29. R. James Cook (ed.). Annual Reviews, Inc., 4139 El Camino Way, Palo Alto, California 94306. 1991. 516 pp. xii. $42.00 U.S.A. and Canada (plus G.S.T.), $47.00 elsewhere (cloth). ISBN 08243-1329-1. The current volume in this excellent series continues the tradi- tion of comprehensive review articles in the field of phytopathology. A total of 23 papers by 35 authors treat subjects ranging from a his- torical review of plant pathology as a profession and the changes in the field, to articles on various groups of plant pathogens, to molecular and ecological studies pertaining to phytopathology. 424 BAY DTOROGILA volume 71(5):423-427 November 1991 Flora Mesoamericana, Glosario Ingles-Espanol, Espanol-Inglés. Fernando Chiang C., Mario Sousa S., & Mario Sousa P. Coordinador de la Bib- lioteca del Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Apartado Postal 70-233, 04510 Mexico, Distrito Federal, Mexico. 1990. 60 pp. Price unknown (paper). ISBN 968-36-1574-0. Botanical terms are listed in each language in the two glos- saries. This book will be an invaluable resource for English speak- ing botanists working in Spanish speaking countries, and vice versa. As only 1000 copies were printed, the book may also become very difficult to obtain. Flora Mesoamericana Glosario para Spermatophyta, Espanol-Inglés. Mario Sousa S. & Sergio Zarate P. Coordinador de la Biblioteca del Instituto de Biologia, Universidad Nacional Autonoma de México, Apartado Postal 70-233, 04510 México, Distrito Federal, México. 1988 (reprinted from 1983 edition). 88 pp. Price unknown (paper). ISBN 968-36-0827-2. This book lists and defines (in Spanish) the terms to be used by authors for the Flora Mesoamericana project. In addition to the glossary itself, the “index” consists of a listing of English botanical terms with their Spanish translations. Flowering Bulbs, Indoors and Out. Theodore James, Jr. Photography by Harry Haralambou. MacMillan Publishing Co., 866 Third Avenue, New York, New York 10022. 1991. 150 pp. x. $29.95 (cloth). ISBN 0-02- 558915-6. Many (over 80) different types of cultivated bulbs are listed in this work. Information on blooming time, hardiness, planting methods, and other features of the plants are included with each entry. The book also contains advice on design and construction of gardens, combinations of bulbs that will provide blooms over extended periods of time. The photographs are of excellent quality. Books received 425 Gardens of the World. Penelope Hobhouse & Elvin McDonald (consulting editors). MacMillan Publishing Co., 866 Third Avenue, New York, New York 10022. 1991. 262 pp. x. $39.95 (hardcover). ISBN 0-02-583127-5. This wonderfully illustrated book provides a whirlwind tour of some famous and some less known gardens of the world. Gardens are grouped within the book by specialization of plant type, style of garden, or orientation of garden. Included are Japanese gardens, rose gardens, tropical gardens, gardens specializing in bulbs, formal gardens, and public gardens. While not technically oriented in a taxonomic or ecological sense, the book is a delight to browse through. Methods in Stomatal Research. Jonathan Weyers & Hans Meidner. John Wiley & Sons, Inc., Longman Scientific & Technical, 1 Wiley Drive, Somerset, New Jersey 08875-1272 1991. 233 pp. x. $84.95 (hardcover). ISBN 0-582-03483-3. This would appear to be an indispensable book for anyone (par- ticularly physiologists) studying stomata. Many methods (from microscopical to microelectrical) are described in the book, for studying stomatal structure and function. In addition to the use- fulness for physiologists, ecologists and taxonomists may find this a useful reference work when their research involves stomatal data. Passion Flowers and Passion Fruit. John Vanderplank. The MIT Press, 55 Hayward Street, Cambridge, Massachusetts 02142. 1991. 176 pp. $35.00 (hardcover). ISBN 0-262-22043-1. This well illustrated book treats over 100 of the more commonly cultivated species or Passiflora. The text describes and drawings illustrate the general morphological features of members of the genus. Methods of cultivation and propagation are described, as are the major pests and diseases of passion flowers. The bulk of the text consists of summaries of the commonly cultivated taxa. These summaries include morphological descriptions of the plants, notes about their origins, where they will grow, when they flower, and other information useful to anyone wishing to cultivate them. A brief chapter on butterflies that specialize on Passzflora is also included. A rather unusual identification key is provided for the included taxa. 426 PSHEY TO LOG TA volume 71(5):423-427 November 1991 Plant Form, An Illustrated Guide to Flowering Plant Morphology. Adrian D. Bell. With line drawings by Alan Bryan. Oxford University Press, 200 Madison avenue, New York, New York 10016. 1991. 341 pp. xiii. $95.00 (hardcover); $49.95 (paper). ISBN 0-19-854279-8 (hardcover); 0-19-854219-4 (paper). This book is abundantly illustrated, with color plates (one or more on nearly every left hand page as the book is opened) to de- pict subjects described in the text. These are accompanied by line drawings (mostly on the facing page) to clarify structures repre- sented in the photographs. The book has considerable technical merit, treating subjects of root and shoot morphology, phyllotaxy, inflorescence structure, floral morphology, and seed morphology among others. Unfortunately, while the book will be useful to many botanists, the high price will probably prevent its widespread classroom use. Book of Garden Design. John Brookes. MacMillan Publishing Company, 866 Third Avenue, New York, New York 10022. 1991. 352 pp. $40.00 (cloth). ISBN 0-02-516695-6. (paper). Th om This is a large volume with hundreds of color photographs. Concepts of garden design and style are outlined in the book. This book differs from many others in that architectural symbols and “tricks of the trade” are included here, but absent from other sim- ilar works. These are presented in such a way as to encourage in- dividuals to plan their own gardens. The principles of design and illustration discussed in the text are illustrated with photographs and drawings. A relatively limited catalogue of garden plants is in- cluded, with the idea of selecting plants to produce specific effects in the garden. The brevity of the plant catalogue is not a serious detriment to this work, since many other books contain much more complete lists of actual and potential garden plants. Books received 427 The Ecology of Mycorrhizae. Michael F. Allen. Cambridge Studies in Ecology series. Cambridge University Press, 40 West 20th Street, New York, New York 10011. 1991. 184 pp. xii. $62.50 (hardcover); $27.95 (paper). ISBN 0-521-33531-0 (hardcover); 0-521-33551-1 (paper). This book begins with a general introduction to mycorrhizae, in which basics in structure, evolution, physiology, and reproduction of mycorrhizae are treated. Examination of ecology of mycorrhizae ranges from interactions between mycorrhizae and plants, animals, and fungi. Ecological relationships ranging from species/species interactions to communities dynamics, dispersal of mycorrhizae, and mycorrhizae in relation to succession are examined. The book concludes with a chapter on future research on mycorrhizae. Phytologia (November 1991) 71(5):428. CORRECTIONS AND ADDITIONS Volume 70, issue 3, March 1991: page 220, line 7, substitute “Cistanthe calycina” for “Cistanthe calycina.” Volume 70, issue 4, April 1991, page 331, line 39, column 2, insert a comma between 234 and 336. Volume 70, issue 4, April 1991, page 348, lines 2-3, column 2, insert a new line 3: “spuria 3:467.” Volume 70, issue 4, April 1991, page 349, line 6, column 2, add “3:467” on end of line. Volume 70, issue 4, April 1991, page 349, lines 6-7, column 2, insert a new line 7: “maritima 3:467.” Volume 70, issue 4, April 1991, page 360, line 24, column 1, replace “3:277” with “3:227.” Volume 71, issue 1, July 1991, page 61, lines 36-37, remove “ssp. et var.” Volume 71, issue 1, July 1991, page 62, line 3, substitute “sampler Kunth” for glutinosa Nutt.” Volume 71, issue 3, September 1991: Contents, inside front cover, line 1, substitute “list” for “checklist.” Volume 71, issue 4, October 1991: page 330, line 32, substitute “cochisenszs” for “chihuahuaensis.” Volume 71, issue 4, October 1991: page 330, line 33, insert “ssp. sznuata” before “Berberis haematocarpa.” 428 nical Garden Libra’ CI Pees 3 5185 A «¥ % ‘ + v Po * - iit ng a x ~ hy ‘ m Pa! ang, - *% ee t ~~ hy . Information for Authors Articles from botanical systematics and ecology, including biographical sketches, critical reviews, and summaries of literature will be considered for publication in PHYTOLOGIA. 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