et” PHYTOLOGIA _ An international journal to expedite botanical and phytoecological publication | Vol. 67 July 1989 No. 1 CONTENTS \ R.B. SHAW, S.L. ANDERSON, K.A. SCHULTZ and V.E. DIERSING, Floral inventory for the U.S. Army Pifion Canyon Maneuver Site, Colorado «+1 / R.C.H.M. OUDEJANS, New names and new combinations in the genus , Euphorbia [oy (Euphorbiaceae) PIeT II Iiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiitt) 43 d M. KORD and T. HATHOUT, Effects of Metolachlor and Alachlor on permeability and lipid synthesis in some plants eccccccccccccccccccccccccoocs 3() : _G.L. NESOM, The separation of Trimorpha (Compositae: Astereae) from Erigeron SbubGaeehadanmebnesudecaccccenesccresebcecsenudscbnobesececocsascncacswedtc 61 2 G.L. NESOM, Infrageneric taxonomy of New World Erigeron (Compositae: Astereae) SSSOSSSSHSSSSSSSSSSSSHSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSESSSSSSOSSESS 67 * J. ORTEGA O., Dos nuevos registros de Aristolochia (Aristolochiaceae) para Veracruz, México SOSSSOSSSSOSSSSSHSSSOSSSSSHSSSSSSSSSSSSSHSSSSOSSHHSSHSSOSSOSOOOEE 94 4 M. LOPEZ F. and A. MORALES M., Contribution to the lichen flora of Venezuela, Vi | SSCHHSSHSSSSSSSSSHSSSSSSSSOSSSSSOSSSSSSSSOHSHSHSSOSSSSOSHSHHOSOHSSOSSOSSOOS 1 00 /G.L. NESOM, New combinations in Ericameria (Compositae: Astereae) SOOSSTOHOSSSSSSHSSOSSHSOSSSHSSSHSHSSSSOHSOSHSOSSHOSSSOSSSOSSHHOSOOOSSOSOSSOOSEOOOS 1 04 , J. OLSEN, A new species of Verbesina section Verbesinaria from the Dominican Republic SSCHOOHSSSSSSSSSSSSHSOSSSHSSHSHSSSSSHSSSSSSSSSSHSSHHOSSOSSSSOSSSSSHSOSSESSES 107 S) s Published by Michael J. Warnock 185 Westridge Drive Huntsville, Texas 77340 U.S.A. ' Price of this number $3.00; for this volume $16.00 in advance or $17.00 after close of this volume; $5.00 extra to all foreign addresses; 512 pages constitute a complete volume; claims for numbers lost in the mail must be made immediately after receipt of the next following number for free replacement; back volume "prices apply if payment is received after a volume is closed. Phytologia (July 1989) 67(1):1-12. FLORAL INVENTORY FOR THE U.S. ARMY PINON CANYON MANEUVER SITE, COLORADO R.B. Shaw!', S.L. Anderson!, K.A. Schultz’ & V.E. Diersing? ' Department of Range Science, Colorado State University, Fort Collins, Colorado 80523 USA 2 Environmental Resources Team, Environmental Division, U.S. Army Corps of Engineers, Construction Engineering Research Laboratory, Champaign, Illinois 61820 USA ; _ ABSTRACT The me, (ebiGiline tn the U.S. Army’s Land Condition/Trend Analysis Program is to complete a floristic inventory of their training lands. :Plant. specimens were collected from May 1985 through’ ‘August’ 1988 ‘at the Piftion Canyon Maneuver Site in southeastern Colorado. A total of 359 species of flowering plants were colledtédsat; the site. One taxon, Haplopappus fremontii Gray var. monocephalus (A. Nels.) Hall (Asteraceae), is proposed for the Federal Threatened and Endangered Species List. Three species [Portulaca parvula Gray, Sapindus saponaria L. var. drum- mondii (Hook. & Arn.) L. Benson and Amorpha nana Nutt.| are on the Colorado Natural Area List of Special Concern. KEY WORDS: Floristics, Colorado, endangered species. INTRODUCTION One of the initial steps in implementing the U.S. Army’s Land Condi- tion/Trend Analysis Program (LCTA) is to do a complete floristic and veg- etational inventory of military installations. In the spring of 1985 a study was begun at the Pinon Canyon Maneuver Site (PCMS) to describe the veg- etation, produce a species and ecological checklist, and establish permanent field plots to determine the influence of tracked vehicular traffic on the native vegetation. The following species list is based on collections made from May 1985 through August 1988. Thus far, we have collected 359 species from 65 families and 220 genera. Voucher specimens for these collections are in the Range Science Herbarium, Colorado State University. The 244,000 acre PCMS is located in southeastern Colorado (Figure 1). Average elevation at the site is approximately 5000 ft above sea level. The 2 BHY TOL O.GwrA volume 67(1):1-42 July 1989 region receives about 12 inches of precipitation each year. The climate is semiarid and temperate continental where maximum precipitation coincides with maximum temperature. The vegetation at PCMS is a complex mosaic of grasslands, shrublands and woodlands. The mosaic of vegetation is due in part to the diverse soils (31 soil series or complexes reported) that have developed from sandstone, limestone, basalt and shale parent material. The dominant vegetation types are grasslands that cover extensive areas. These grasslands are typical of the shortgrass steppe or desert grassland and are dominated by blue grama (Bouteloua gracilis), black grama (B. eriopoda), western wheatgrass (Agropy- ron smithit) and galleta (Hilaria jamesiz). Most common interstitial species are ring muhly (Muhlenbergia torreyt), tumblegrass (Schedonnardus panic- ulatus), squirreltail (Sttanion hystriz) and Fendler threeawn (Aristida fend- leriana). The shrublands at PCMS are, for the most part, composed of typical grassland understory vegetation with an overstory of shrubs or succulents. The most abundant erect succulent is tree cholla (Opuntia imbricata). Some of the most common shrubs are small soapweed ( Yucca glauca, pale wolfberry (Lycium pallidum), common winterfat (Ceratoides lanata) and Bigelow sage- brush (Artemisia bigelovii). When soils are extremely sandy, sand sagebrush (A. filifolia) dominates the community. Alkali soils are typically dominated by dense stands of fourwing saltbush (Atriplex canescens) with an under- story of alkali sacaton (Sporobolus airoides). Black greasewood (Sarcobatus vermiculatus) frequently is found in dense stands along seasonal streams and arroyos. The woodlands are composed almost entirely of one seeded juniper (Ju- niperus monosperma) and pinyon pine (Pinus edulis). Understory shrubby vegetation tends to vary with parent material. On limestone outcrops, grease- bush (Glossopetalon meionandra), Bigelow sagebrush and James frankenia (Frankenia jamesit) are the most common shrubs. On sandstone parent ma- terial, the most common understory shrubs are wax currant (Ribes cereum), skunkbush sumac (Rhus trilobata), true mountain mahogany (Cercocarpus montanus), common hoptree (Ptelea trifoliata) and litteleaf mockorange ( Pht- ladelphus microphyllus). Riparian woodlands are dominated by plains cot- tonwood (Populus deltoides), willow (Saliz sp.) and five-stamen tamarix ( Tamariz pentandra). The species list is in alphabetical order by family and species. Author citation, common synonyms, common name and a four letter symbol com- posed of the first two letters of the genus and specific epithet are given for each taxon. The first column after the common name refers to functional group (F=forb, G=grass, T=tree, S=shrub, $=succulent and V=vine) The second column refers to whether the plant is native (N) or introduced (I). The Shaw et al.: Floral inventory of Pinon Canyon Site ‘OPEIOIOD ‘aS JeAnaueyy UOAUeD UOUIg “| aunBl4 SaINE SLIWIT 430- STE ECL $1ajawolly S 319s oOposojo7y 4 PHY TO I-0OG ra volume 67(1):1-42 July 1989 third column refers to the longevity of the plant (A=annual, B=biennial and P=perennial). The fourth column depicts whether the plant flowers during the cool season (C=March to June), warm season (W=July to September) or evergreen (E). PLANT SPECIES LIST - PINON CANYON MANEUVER SITE Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season AGAVACEAE Yucca glauca Nutt. SMALL SOAPWEED (YUGL) S N P Cc ALISMATACEAE Alisma triviale Pursh (syn. A. plantago- aquatica L. var. americanumR. & S. AMERICAN WATERPLANTAIN (ALTR) F N 1B WwW cf Alisma sp. WATERPLANTAIN (AL-X) F N P Cc Sagittaria cf latifolia Willd. COMMON ARROWHEAD (SALA) F N 1p Ww AMARANTHACEAE Amaranthus albus L. TUMBLEWEED (AMAL]) F N A Ww Amaranthus graecizans L. (syn. A. blitoides S. Wats.) PROSTRATE PIGWEED AMARANTH (AMGR) F N A WwW Shaw et al.: Floral inventory of Pinon Canyon Site 5 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season AMARANTHACEAE (continued) Amaranthus retroflezus L. ROUGH PIGWEED (AMRE) F N A Ww ANACARDIACEAE Rhus trilobata Nutt. SKUNKBUSH SUMAC (RHTR) S N P Cc Tozicodendron rydbergii (Small) Greene POISON IVY (TORY) Wi N ie C APOCYNACEAE Apocynum cannabinum L. INDIAN HEMP (APCA) r N P Cc ASCLEPIADACEAE Asclepias arenaria Torr. SAND MILKWEED (ASAR) F N P w Asclepias asperula (Dcne.) Woodson (syn. A. capricorn Woodson ssp. occidentalis Woodson) ANTELOPE HORNS (ASAS) F N |e: C Asclepias engelmanniana Woodson ENGELMANN MILKWEED (ASEN) E I P Ww Asclepias incarnata L. SWAMP MILKWEED (ASIN) B N |e Ww Asclepias macrotis Torr. LONGHORN MILKWEED (ASMA) io N P Ww Asclepias speciosa Torr. SHOWY MILKWEED (ASSP) ¥ N P Cc 6 PHYTOLOGIA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season ASCLEPIADACEAE (continued) Asclepias subverticillata (Gray) Vail POISON MILKWEED (ASSU) F N P WwW Asclepias verticillata L. WHORLED MILKWEED (ASVE) F N P W Asclepias viridiflora Raf. GREEN MILKWEED (ASVI) F N 1B W BORAGINACEAE Cryptantha cinerea (Greene) Cronq. var. jamesti Cronq. [syn. C. jamesii (Torr.) Pays.| JAMES CRYPTANTHA (CRCI) F N P C Cryptantha minima Rydb. CRYPTANTHA (CRMI) F N P C Cryptantha thyrsiflora (Greene) Payson CLUSTER CRYPTANTHA (CRTH) F N Proas Cc Lappula diploma (F. & M.) Guerke [syn. L. redowskii (Hornem.) Greene var. cupulata (Gray) M.E. Jones; L. terana (Scheele) Britt.] CUPSEED STICKSEED (LADI) F N A C Lappula redowskti (Hornem.) Greene BLUEBUR STICKSEED (LARE) 12 N A C Lithospermum incisum Lehm. NARROWLEAF GROMWELL (LIIN) ry 2 ~ Q Onosmodium molle Michx. var. occidentale (Mack.) I. Johnst. WESTERN MARBLESEED (ONMO) F N P .@; Shaw et al.: Floral inventory of Pinon Canyon Site ¢ Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season CACTACEAE Coryphantha vivipara (Nutt.) Britt. & Rose PINCUSHION CACTUS (COVI) $ N P C Echinocereus viridiflorus Engelm. HEDGEHOG CACTUS (ECVI) $ N B C Opuntia imbricata (Haw.) DC. TREE CHOLLA (OPIM) $ N i C Opuntia phaeacantha Engelm. PRICKLY PEAR (OPPH) $ N 13g C Opuntia polyacantha Haw. var. polyacantha PLAINS PRICKLY PEAR (OPPO1) $ N P C Opuntia polyacantha Haw. var. trichophora (Engelm. & Bigel.) Coult. HOARY PRICKLY PEAR (OPPO2) $ N P C CAMPANULACEAE Lobelia cardinalis L. CARDINAL FLOWER (LOCA) F N P Ww CAPPARIDACEAE Cleome serrulata Pursh ROCKY MOUNTAIN BEEPLANT (CLSE) F N A W Polansia dodecandra (I..) DC. ssp. trachysperma (T. & G.) Iltis (syn. P. trachysperma T. & G.) CLAMMYWEED (PODO) E N A C 8 Poa YY PeOVrO GA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season CAPRIFOLIACEAE Sambucus canadensis L. AMERICAN ELDER (SACA) S N BE Ww Symphoricarpos occidentalis Hook. WESTERN SNOWBERRY (SYOC) S N 1B Cc Symphoricarpos oreophilus Gray (syn. S. rotundifolius var. oreophilus M.E. Jones MOUNTAIN SNOWBERRY (SYOC) S N P Cc CARYOPHYLLACEAE Arenaria hookeri Nutt. var. hookeri HOOKER SANDWORT (ARHO) F N P C Paronychia cf sessiliflora Nutt. CREEPING NAILWORT (PASE) F N je C CELASTRACEAE Glossopetalon meionandra Koehne. {syn. Forsellesia meionandra (Koehne) Heller] GREASEBUSH (GLME) S N Ee C CHENOPODIACEAE Atripler argentea Nutt. TUMBLING SALTBUSH (ATAR) F N A W Atripler canescens (Pursh) Nutt. FOURWING SALTBUSH (ATCA) S N 2 C Shaw et al.: Floral inventory of Pinon Canyon Site 9 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season CHENOPODIACEAE (continued) Ceratoides lanata (Pursh) Howell {syn. Eurotia lanata (Pursh) Moq.| COMMON WINTERFAT (CELA) S N P C Chenopodium cf album L. LAMBSQUARTERS GOOSEFOOT (CHAL) F I A W Chenopodium incanum (S. Wats.) Heller GOOSEFOOT (CHIN) F N A Ww Chenopodium cf leptophyllum Nutt. SLIMLEAF GOOSEFOOT (CHLE) F N A Ww Kochia scoparia (L.) Schrad. KOCHIA (KOSC) F I A Ww Salsola iberica Sennen & Pau (syn. S. kali L.) RUSSIAN THISTLE (SAIB) F I A W Sarcobatus vermiculatus (Hook.) Torr. BLACK GREASEWOOD (SAVE) Ss N 12 C COMMELINACEAE Tradescantia occidentalis (Britt.) B. Smyth PRAIRIE SPIDERWORT (TROC) ef N P Cc 10 PHY? O.eCGTA Alphabetic by Family Then by Species COMPOSITAE Ambrosia cf acanthicarpa Hook. (syn. Franseria acanthicarpa (Hook.) Cov.] ANNUAL BURSAGE (AMAC) Ambrosia psilostachya DC. var. coronopifolia (T. & G.) Farw. (syn. A. coronopifolia T. & G. WESTERN RAGWEED (AMPS) Antennaria obovata E. Nels. PUSSYTOES (ANOB) Antennaria parvifolia Nutt. LITTLELEAF PUSSYTOES (ANPA) Arctium minus Bernh. COMMON BURDOCK (ARMI) Artemisia bigelovit Gray BIGELOW SAGEBRUSH (ARBI) Artemisia dracunculus L. WILD TARRAGON (ARDR) Artemisia filifolia Torr. SAND SAGEBRUSH (ARFI) Artemisia frigida Willd. FRINGED SAGEBRUSH (ARFR) Artemisia ludoviciana Nutt. LOUISIANA SAGEBRUSH (ARLU) Aster cf ericoides L. WHITE ASTER (ASER) volume 67(1):1-42 July 1989 Life Native/ Cool/Warm Form Introduced Longevity Season 1 N A W in N | Ww F N P Cc 18 N Pp Cc F I B WwW S N P Mi! 1g N iB Ww S N P Ww ip N Pp Ww r N P Ww F N P Ww Shaw et al.: Floral inventory of Pinon Canyon Site 11 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season COMPOSITAE (continued) Aster cf falcatus Lindl. WHITEPRAIRIE ASTER (ASFA) F N Ie Baccharis wrightii Gray WRIGHT BACCHARIS (BAWR) F N P Brickellia cf brachyphylla (Gray) Gray BRICKELLBUSH (BRBR) F N EB Brickellia cf californica (T. & G.) Gray CALIFORNIA BRICKELLBUSH (BRCA) F N P Centaurea repens L. (syn. C. picris Pall.) KNAPWEED (CERE1) F I P Chrysothamnus nauseosus (Pall.) Britt. RABBITBRUSH (CHNA) S N P Cirsium cf undulatum (Nutt.) Spreng. WAVYLEAF THISTLE (CIUN) F N B Conyza canadensis (L.) Cronq. CANADIAN HORSEWEED (COCA) F I A Coreopsis tinctoria Nutt. PLAINS COREOPSIS (COTI) 1p I A Dyssodia aurea (Gray) A. Nels. DOGWEED (DYAU) F N A Erigeron divergens T. & G. var. cinereus Gray SPREADING FLEABANE (ERDI) Ei N B W W 12 PHY TOLOCIA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season COMPOSITAE (continued) Erigeron pumilus Nutt. var. pumilus LOW FLEABANE (ERPU) F N P C cf Evaz prolifera Nutt. FLUFFWEED (EVPR) F N A C Gaillardia pinnatifida Torr. BLANKET FLOWER (GAPI) F N Ie Cc Grindelia squarrosa(Pursh) Dun. CURLYCUP GUMWEED (GRSQ) 19 N P Af Gutierrezia sarothrae (Pursh) Britt. & Rusby [syn. Xanthocephalum sarothrae (Pursh) Shinners] BROOM SNAKEWEED (GUSA) F N P W Haplopappus fremontti Gray var. fremontii [syn. Oonopsis foliosa (Gray) Greene] FREMONT GOLDENWEED (HAFR1) F N i Ww Haplopappus fremontii Gray var. monocephalus (Nels.) Hall (syn. Oonopsis monocephala A. Nels. RAYLESS GOLDENWEED (HAFR2) F N iB W Haplopappus spinulosus (Pursh) DC. [syn. Machaeranthera pinnatifida (Hook.) Shinners] SPINY GOLDENWEED (HASP) Fr N iP Cc Helianthus petiolaris Nutt. PRAIRIE SUNFLOWER (HEPE) F N A W Helianthus sp. SUNFLOWER (HE-X) F A W Shaw et al.: Alphabetic by Family Then by Species Floral inventory of Pinon Canyon Site Life Native/ 13 Cool/Warm Form Introduced Longevity Season COMPOSITAE (continued) Heterotheca horrida (Rydb.) V. Harms [syn. Chrysopsis hispida (Hook.) DC.; Chrysopsis horrida Rydb.| GOLDASTER (HEHO) Heterotheca villosa (Pursh) Shinners var. angustifolia (Rydb.) V. Harms [syn. Chrysopsis villosa (Pursh) Nutt.; var. angustifolia (Rydb.) Cronq.] HAIRY GOLDASTER (HEVI) EF N 1 Hymenopappus filifolius Hook. FINE LEAF HYMENOPAPPUS (HYFI)_ F N le Hymenopappus tenutfolius Pursh HYMENOPAPPUS (HYTE) F N B Hymenozys acaulis (Pursh) Parker STEMLESS HYMENOXYS (HYAC) F N 12 Iva azillaris Pursh POVERTYWEED (IVAX) ig N ie Lactuca serriola L. (syn. L. scariola L.) PRICKLY LETTUCE (LASE) E I A Lactuca tatarica(L.) C.A. Meyer ssp. pulchella (Pursh) Stebbins [syn. L. pulchella (Pursh) Stebbins; L. oblongifolia Nutt.] CHICORY LETTUCE (LATA) F N P Leucelene ericoides (Torr.) Greene [syn. Aster arenosus (Heller) Blake] HEATH ASTER (LEER) EF N P 14 PHYTOLTOGTA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season COMPOSITAE (continued) Liatris punctata Hook. DOTTED GAYFEATHER (LIPU) F N es Ww Lygodesmia juncea (Pursh) Hook. RUSH SKELETONPLANT (LYJU) F N P Ww Melampodium cinereum DC. [syn. M. leucanthum T. & G. PLAINS BLACKFOOT (MECI) E N 1B Cc Nothocalais cuspidata (Pursh) Greene [syn. Microseris cuspidata (Pursh) Sch.] FALSE DANDELION (NOCU) E N P Cc Palafozia rosea (Bush) Cory var. macrolepis Rydb. [syn. P. macrolepis (Rydb.) Cory] PALAFOXIA (PARO) F N A Mf Pectis angustifolia Torr. FETID-MARIGOLD (PEAN1) F N A W Picradeniopsis oppositifolia (Nutt.) Rydb. [syn. Bahia oppositifolia (Nutt.) DC.] PLAINS BAHIA (PIOP) F N P WwW Ratibida columnifera (Nutt.) Wooton & Standley UPRIGHT PRAIRIE CONEFLOWER (RACO) F N P Ww Ratibida tagetes (James) Barnh. SHORTRAY PRAIRIE CONEFLOWER (RATA) F N P Ww Shaw et al.: Floral inventory of Pinon Canyon Site 15 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season COMPOSITAE (continued) Senecio cf pseudaureus Rydb. var.:flavulus (Greene) Greenm. (syn. S. flavulus Greene) GOLDEN GROUNDSEL (SEPS) F N P C Senecio riddellii T. & G. RIDDELL GROUNDSEL (SERI) F N P W Senecio tridenticulatus Rydb. GROUNDSEL (SETR) F N [2 C Solidago mollis Bartl. VELVETY GOLDENROD (SOMO) F N ie Ww Solidago multiradiata Ait. (syn. S. ciliosa Greene) GOLDENROD (SOMU) F N P Ww Solidago petiolaris Ait. DOWNY GOLDENROD (SOPE) EF N Pp W Solidago sparsiflora Fray THREE-NERVED GOLDENROD (SOSP) F N P W Stephanomeria pauciflora (Torr.) A. Nels. DESERT WIRELETTUCE (STPA) F N P Ww Tarazracum officinale Weber COMMON DANDELION (TAOF) F I P Cc Thelesperma megapotamicum (Spreng.) O. Ktze. GREENTHREAD (THME) EF N P C Thelesperma subnudum Gray NAVAJO-TEA GREENTHREAD (THSU) F N P Cc 16 PHY TOLOGITIA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season COMPOSITAE (continued) Townsendia hookeri Beaman EASTER DAISY (TOHO) F N P Cc Tragopogon dubius Scop. WESTERN SALSIFY (TRDU) F I B C Verbesina encelioides (Cav.) Benth. & Hook. GOLDEN CROWNBEARD (VEEN) F N A WwW Xanthium strumarium L. COCKLEBUR (XAST) F I A Ww Zinnia grandiflora Nutt. ROCKY MOUNTAIN ZINNIA (ZIGR) F N Ee C CONVOLVULACEAE . . Convolvulus arvensis L. FIELD BINDWEED (COAR) V I P Cc Convolvulus equitans Benth. BINDWEED (COEQ) Vv N P W Evolvulus nuttallianus R. & S. ARIZONA EVOLVULUS (EVNU) F N Pp Cc Ipomoea leptophylla Torr. BUSH MORNING-GLORY (IPLE) Vv N le Cc CRUCIFERAE Camelina microcarpa Andiz. LITTLEPOD FALSEFLAX (CAMI) E I A C Descurainia pinnata (Walt.) Britt. PINNATE TANSYMUSTARD (DEPI) EF N A Cc Shaw et al.: Floral inventory of Pinon Canyon Site Li Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season CRUCIFERAE (continued) Descurainia sophia (L.) Webb. FLIXWEED TANSYMUSTARD (DESO) F I A C Draba reptans (Lam.) Fern. CAROLINA DRABA (DRRE) F N A C Erysimum asperum (Nutt.) DC. WESTERN WALLFLOWER (ERAS) F N EB C Lesquerella fendleri (Gray) Wats. FENDLERS BLADDERPOD (LEFE) F N Pp C Stanleya pinnata (Pursh) Britt. PRINCES PLUME (STPI) F N 4 C Thelypodium wrightit Gray WRIGHT THELYPODY (THWR) F N B Ww CUCURBITACEAE Cucurbita foetidissima H.B.K. BUFFALO GOURD (CUFO) V I P WwW CUPRESSACEAE Juniperus monosperma (Engelm.) Sarg. ONESEEDED JUNIPER (JUMO) AL N P Cc Juniperus scopulorum Sarg. ROCKY MOUNTAIN JUNIPER (JUSC) T N P C 18 PHY TOLOGIA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season CYPERACEAE. Carez brevoir (Dew.) Mack. SEDGE (CABR) F N ib Cc Carez foena Willd. SEDGE (CAFO) F N | Cc Carer gravida Bailey var. lunelliana (Mack.) F.J. Herm. HEAVY SEDGE (CAGR) F N P Cc Carez cf lanuginosa Michx. WOOLLY SEDGE (CALA) \3 N P C Carex rerantica Bailey SEDGE (CAXE) F N P S Cyperus filiculmis Vahl. [syn. C. lupulinus (Spreng.) Marcks ssp. lupulinus] FERN FLATSEDGE (CYFI) F N P Ww Cyperus schweinitzti Torr. SCHWEINITZ FLATSEDGE (CYSC) B N Salil cS Ww Eleocharis palustris (L.) R. & S. (syn. E. macrostachya Britt.) COMMON SPIKESEDGE (ELPA) F N P Cc Eleocharis sp. SPIKESEDGE (EL-X) F |e Cc Scirpus acutus Muhl. TULE BULRUSH (SCAC) F N E Cc Scirpus americanus Pers. AMERICAN BULRUSH (SCAM) F N Pp Ww Scirpus pallidus (Britt.) Fern. BULRUSH (SCPA1) F N P C Shaw et al.: Alphabetic by Family Then by Species EQUISETACEAE Equisetum laevigatum A. Braun SMOOTH HORSETAIL (EQLA) Equisetum variegatum Schleich. VARIEGATED HORSETAIL (EQVA) EUPHORBIACEAE Croton terensis (K1.) Muell. Arg. TEXAS CROTON (CRTE) Euphorbia dentata Michx. TOOTHED EUPHORBIA (EUDE) Euphorbia fendleri T. & G. FENDLER EUPHORBIA (EUFE) Euphorbia cf glyptosperma Engelm. RIDGESEED EUPHORBIA (EUGL) Euphorbia lata Engelm. HOARY EUPHORBIA (EULA) Euphorbia marginata Pursh SNOW-ON-THE-MOUNTAIN- EUPHORBIA (EUMA) Euphorbia missurica Raf. [syn. Chamaesyce missurica (Raf.) Shinners] MISSOURI EUPHORBIA (EUMI) Euphorbia cf serpens H.B.K. ROUND LEAVED SPURGE (EUSE) Euphorbia spathulata Lam. SPURGE (EUSP) Floral inventory of Pinon Canyon Site 19 Life Native/ Cool/Warm Form Introduced Longevity Season 1h N P C E N Pp Ww lp N A W F N A Cc E N P C EF N A C ie N le C F N A Ww 18 N A W Ie N A Ww EF N A C 20 PANO OF Gia Alphabetic by Family Then by Species EUPHORBIACEAE (continued ) Euphorbia stictospora Engelm. MAT SPURGE (EUST) volume 67(1):1-42 July 1989 Life Native/ Cool/Warm Form Introduced Longevity Season Tragia nepetaefolia Cav. (syn. T. urticifolia Michx.; T. betonicifolia Nutt. NOSEBURN (TRNE) FRANKENIACEAE Frankenia jamesti Torr. JAMES FRANKENIA (FRJA) FUMARIACEAE Corydalis aurea Willd. GOLDEN SMOKE (COAU) GERANIACEAE Erodium cicutarium (L.) L’Her. FILAREE (ERCI1) GRAMINEAE Agropyron cristatum (L.) Gaertn. CRESTED WHEATGRASS (AGCR) Agropyron dasystachyum (Hook.) Scribn. THICKSPIKE WHEATGRASS (AGDA) Agropyron smithii Rydb. WESTERN WHEATGRASS (AGSM) E N A WwW F N P Cc S N 127 Cc In N A C F I A C G I P C Q Zi ~ Q Shaw et al.: Floral inventory of Pinon Canyon Site Alphabetic by Family Then by Species GRAMINEAE (continued) Agrostis stolonifera L. REDTOP BENTGRASS (AGST) Andropogon gerardi Vitman BIG BLUESTEM (ANGE) Aristida adscensionis L. SIXWEEKS THREEAWN (ARAD) Aristida divaricata Humb. & Bonpl. POVERTY THREEAWN (ARDI) Aristida fendleriana Steud. [syn. A. purpurea Nutt. var. longiseta (Steud.) Vasey] FENDLER THREEAWN (ARFE) Aristida longiseta Steud. [syn. A. purpurea Nutt. var. robusta (Merrill) A. Holmgren & N. Holmgren] RED THREEAWN (ARLO) Aristida purpurea Nutt. PURPLE THREEAWN (ARPU) Avena fatua L. WILD OAT (AVFA) Bothriochloa saccharoides (Sw.) Rydb. (syn. Andropogon saccharoides Sw.) SILVER BLUESTEM (BOSA) Bouteloua curtipendula (Michx.) Torr. SIDEOATS GRAMA (BOCU) Bouteloua eriopoda (Torr.) Torr. BLACK GRAMA (BOER) Zyl Cool/Warm Form Introduced Longevity Season 22 PHY TOLOG IA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season GRAMINEAE (continued) Bouteloua gracilis (H.B.K.) Lag. BLUE GRAMA (BOGR) G N P W Bouteloua hirsuta Lag. HAIRY GRAMA (BOHI) G N 1B Ww Bromus japonicus Thunb. JAPANESE BROME (BRJA) G I A Cc Buchloe dactyloides (Nutt.) Engelm. BUFFALOGRASS (BUDA) G N |e WwW Calamagrostis neglecta (Ehrh.) Gaertn. [syn. C. stricta (Timm.) Koel.] SLIMSTEM REEDGRASS (CANE) Q Zz ae) Q Cenchrus longispinus (Hack.) Fern. SANDBUR (CELO) G N Pp WwW Dactylis glomerata L. ORCHARDGRASS (DAGL) G I Ie , Cc Dichanthelium oligosanthes (Schult.) Gould var. scribnerianum (Nash) Gould (syn. Panicum scribnerianum Nash) SCRIBNER PANICUM (DIOL) G N P Cc Distichlis spicata (L.) Greene var. stricta (Torr.) Beetle INLAND SALTGRASS (DISP) G N P Ww Echinochloa crusgalli(L.) Beauv. BARNYARD GRASS (ECCR) G I A WwW Shaw et al.: Floral inventory of Pinon Canyon Site 23 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season GRAMINEAE (continued) Elymus canadensis L. CANADA WILDRYE (ELCA) G N P C Eragrostis cilianensis (All.) Mosher STINK GRASS (ERCI2) G N A Ww Eragrostis spectabilis (Pursh) Steud. PURPLE LOVEGRASS (ERSP) G N r Ww Erioneuron pilosum (Buckl.) Nash [syn. Tridens pilosus (Buckl.) Hitchc.] HAIRY FALSE TRIDENS (ERPI) G N |e W Glyceria stricta (Lam.) Hitchc. FOWL MANNAGRASS (GLST) G N P C Hilaria jamesii (Torr.) Benth. GALLETA (HIJA) G N 1e Ww Hordeum jubatum L. FOXTAIL BARLEY (HOJU) G N P C Hordeum pusillum Nutt. LITTLE BARLEY (HOPU) G N A C Koeleria pyramidata (Lam.) Beauv. [syn. K. cristata (L.) Pers.] PRAIRIE JUNEGRASS (KOPY) G N P C Lycurus phleoides H.B.K. COMMON WOLFTAIL (LYPH) G N 1B W Muhlenbergia arenacea (Buckl.) Hitchc. EAR MUHLY (MUAR1) G N P W 24 PHYTOLOGIA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season GRAMINEAE (continued) Muhlenbergia arenicola Buckl. SAND MUHLY (MUAR2) G N EB W Muhlenbergia asperifolia (Nees. & Mey.) Parodi ALKALI MUHLY (MUAS) G N ie WwW Muhlenbergia racemosa (Michx.) B.S.P. GREEN MUHLY (MURA) G N P W Muhlenbergia torreyi (Kunth) Hitchc. RING MUHLY (MUTO) G N P Ww Munroa squarrosa (Nutt.) Torr. FALSE BUFFALOGRASS (MUSQ) G N A Ww Oryzopsis hymenoides (R. & S.) Ricker INDIAN RICEGRASS (ORHY) G N E C Oryzopsis micrantha (Trin. & Rupr.) Thurb. LITTLESEED RICEGRASS (ORMI) G N P C Panicum capillare L. COMMON WITCHGRASS (PACA) G N A W Panicum obtusum H.B.K. VINE MESQUITE (PAOB) G N P Ww Panicum virgatum L. SWITCHGRASS (PAVI2) G N P Ww Phleum pratense L. TIMOTHY (PHPR) G I 12 Cc Phragmites communis Trin. [syn. P. australis (Cav.) Trin.] COMMON RED REED (PHCO) G N le Cc Shaw et al.: Alphabetic by Family Then by Species Floral inventory of Pinon Canyon Site Life Native/ 25 Cool/Warm Form Introduced Longevity Season GRAMINEAE (continued) Poa bigelovii Vasey & Scribn. BIGELOW BLUEGRASS (POBI) Poa pratensis L. KENTUCKY BLUEGRASS (POPR) Poa sandbergii Vasey SANDBERG BLUEGRASS (POSA) Polypogon monspeliensis (L.) Desf. RABBITFOOT POLYPOGON (POMO) Schedonnardus paniculatus (Nutt.) Trel. TUMBLEGRASS (SCPA2) Gekteachyreum scoparium (Michx.) Nash (syn. Andropogon scoparius Michx.) LITTLE BLUESTEM (SCSC) Scleropogon brevifolius Phil. BURROGRASS (SCBR) Sitanion hystriz Nutt. J.G. Smith SQUIRRELTAIL (SIHY) Sorghastrum nutans (L.) Nash INDIANGRASS (SONU2) Sphenopholis obtusata (Michx.) Scribn. WEDGEGRASS (SPOB) Sporobolus airoides (Torr.) Torr. ALKALI SACATON (SPAI) 26 PE OL On A volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season GRAMINEAE (continued) Sporobolus cryptandrus (Torr.) Gray SAND DROPSEED (SPCR) G N iP Ww Stipa comata Trin. & Rupr. NEEDLE AND THREAD (STCO) G N r Cc Stipa neomezicana (Thurb.) Scribn. NEW MEXICO FEATHERGRASS (STNE) G N P C Stipa robusta (Vasey) Scribn. SLEEPYGRASS (STRO) G N e Cc Stipa scribneri Vasey SCRIBNER NEEDLEGRASS (STSC) G N P Cc Stipa viridula Trin. GREEN NEEDLEGRASS (STVI) G N Pp Cc Tridens elongatus (Buckl.) Nash [syn. T. muticus (Torr.) Nash var. elongatus (Buckl. Shinners] GREEN TRIDENS (TREL) G N Pa Ww Vulpia octoflora (Walt.) Rydb. (syn. Festuca octoflora Walt.) SIXWEEKS FESCUE (VUOC) G N A C GROSSULARIACEAE Ribes aureum Pursh GOLDEN CURRANT (RIAU) S N P Cc Ribes cereum Doug}. WAX CURRANT (RICE) 5 N P C Ribes cf leptanthum Gray . TRUMPET GOOSEBERRY (RILE) S N P C Shaw et al.: Floral inventory of Pinon Canyon Site Alphabetic by Family Then by Species HYDRANGEACEAE Philadelphus microphyllus Gray var. microphyllus LITTLELEAF MOCKORANGE (PHMI) JUNCACEAE Juncus balticus Willd. BALTIC RUSH (JUBA) Juncus interior Wieg. INLAND RUSH (JUIN) Juncus torreyi Cov. TORREY RUSH (JUTO) LABIATAE Hedeoma drummondii Benth. DRUMMOND FALSE PENNYROYAL (HEDR) Marrubium vulgare L. COMMON HOARHOUND (MAVU) Monarda pectinata Nutt. PONY BEEBALM (MOPE) Salvia reflera Hornem. LANCELEAF SAGE (SARE) Teucrium laciniatum Torr. CUTLEAF GERMANDER (TELA) Zit Cool/Warm Form Introduced Longevity Season 28 RPPY ROW GT aA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season LEGUMINOSAE Amorpha fruticosa L. FALSE INDIGO (AMFR) S N P G Amorpha nana Nutt. DWARF INDIGO AMORPHA (AMNA) _ S N P Cc Astragalus crassicarpus Nutt. GROUNDPLUM MILKVETCH (ASCR)_ F N 1B Cc Astragalus gracilis Nutt. SLENDER MILKVETCH (ASGR) F N iE C Astragalus missouriensis Nutt. MISSOURI MILK VETCH (ASMI) F N ig C Astragalus puniceus Osterh. TRINIDAD MILKVETCH (ASPU) F N |e Cc Astragalus racemosus Pursh ALKALI MILKVETCH (ASRA) F N P C Astragalus shortianus Nutt. SHORTS MILKVETCH (ASSH) F N P C Caesalpinia jamesii (T. & G.) Fisher (syn. Hoffmanseggia jamesii T. & G. JAMES RUSHPEA (CAJA) F N Y C Dalea aurea Nutt. SILKTOP DALEA (DAAU) F N P Ww Shaw et al.: Floral inventory of Pinon Canyon Site Alphabetic by Family Life Native/ 20 Cool/Warm Then by Species Form Introduced Longevity Season LEGUMINOSAE (continued) Dalea candida Michx. [syn. Petalostemum candidus ( Willd.) Michx. WHITE PRAIRIE CLOVER (DACA) F N B Dalea enneandra Nutt. INDIGO BUSH (DAEN) F N P Dalea jamesii (Torr.) T. & G. JAMES DALEA (DAJA) F N im Dalea purpurea Vent. [syn. Petalostemum purpureum (Vent.) Rydb. PURPLE PRAIRIE CLOVER (DAPU) F N 12 Glycyrrhiza lepidota Pursh AMERICAN LICORICE (GLLE) F N P Hedysarum boreale Nutt. NORTHERN SWEETVETCH (HEBO) F N le Hoffmanseggia drepanocarpa Gray SICKLEPOD RUSHPEA (HODR) F N P Lathyrus eucosmus Butters & St. John {syn. L. brachycalyz Rydb. ssp. eucosmus (Butters & St. John) Welsh] BUSH PEAVINE (LAEU) F N P Lupinus pusillus Pursh RUSTY LUPINE (LUPU) F N A Melilotus alba Medic. WHITE SWEETCLOVER (MEAL1) EF I A 30 PHYTOLOGIA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season Melilotus officinalis (L.) Pallas YELLOW SWEETCLOVER (MEOF) F I A Cc Ozytropis lambertii Pursh LAMBERT CRAZYWEED (OXLA) F N iE Cc Psoralea tenuiflora Pursh SLIMFLOWER SCURFPEA (PSTE) iD N P C Sophora nuttalliana B.L. Turner (syn. S. sericea Nutt.) SILKY SOPHORA (SONU1) F N iB Cc Victa americana Muhl. AMERICAN VETCH (VIAM) Vv N 1B Cc LILIACEAE Allium teztile A. Nels. & Macbr. TEXTILE ONION (ALTE) F N iP Cc Calochortus cf gunntsonii S. Wats. GUNNISON MARIPOSALILY (CAGU) F N P. C Leucocrinum montanum Nutt. SAND LILY (LEMO) F N P C LINACEAE Linum lewisii Pursh [syn. L. perenne L. var. lewisii (Pursh) Eat. & Wright] LEWIS FLAX (LILE) F N P Cc Linum cf rigidum Pursh YELLOW FLAX (LIRI) F N A C Shaw et al.: Floral inventory of Pinon Canyon Site 31 Alphabetic by Family Cool/Warm Then by Species Form Introduced Longevity Season LOASACEAE Mentzelia albicaulis (Hook.) T. & G. WHITESTEM MENTZELIA (MEAL2) Mentzelia nuda (Pursh) T. & G. {syn. Nuttallia nuda (Pursh) Greene] BRACTLESS MENTZELIA (MENU) Mentzelia oligosperma Nutt. STICKLEAF (MEOL) MALVACEAE Sphaeralcea angustifolia (Cav.) D. Don [syn. S. angustifolia (Cav.) D. Don var. cuspidata| NARROWLEAF GLOBEMALLOW (SPAN) Sphaeralcea coccinea (Pursh) Rydb. SCARLET GLOBEMALLOW (SPCO) MARTYNIACEAE Martynia louisianica P. Mill. [syn. Proboscidea louisianica(P. Mill.) Thell.] DEVIL’S CLAW (MALO) NYCTAGINACEAE Abronia fragrans Nutt. SNOWBALL SANDVERBENA (ABFR) Mirabilis multiflora (Torr.) Gray COLORADO FOUR-O’CLOCK (MIMU) 32 PHY POLO ETA volume 67(1):1-42 Alphabetic by Family Life Native/ July 1989 Cool/Warm Then by Species Form Introduced Longevity Season NYCTAGINACEAE (continued) Mirabilis hirsuta (Pursh) MacM. (syn. Orybaphus hirsutus Sweet) HAIRY UMBRELLAWORT (MIHI) F N P Mirabilis linearis (Pursh) Heimerl. [syn. Ozybaphus linearis (Pursh) Robinson] NARROWLEAVED UMBRELLAWORT (MILI) F N P Tripterocalyz micranthus (Torr.) Hook. SANDPUFF (TRMI) 1D N P ONAGRACEAE Calylophus hartwegit (Benth.) Raven ssp. lavandulifolius (T. & G.) Towner & Raven [syn. C. lavandulifolius (T. & G.) Raven; Oenothera lavandulaefolia T. & G.] LAVENDERLEAF EVENING PRIMROSE (CAHA) F N iP Gaura coccinea Pursh SCARLET GAURA (GACO) ig N P Gaura parviflora Doug}. SMALLFLOWER GAURA (GAPA) F N B Oenothera caespitosa Nutt. TUFTED EVENING PRIMROSE (OECA) fF N P Oenothera harringtonii Wagner, Stockhouse & Klein EVENING PRIMROSE (OEHA) ig N P Shaw et al.: Alphabetic by Family Then by Species OROBANCHACEAE Orobanche multiflora Nutt. BROOMRAPE (ORMU) PAPAVERACEAE Argemone cf hispida Gray [syn. A. Floral inventory of Pinon Canyon Site 33 Life Native/ Cool/Warm Form Introduced Longevity Season platyceras Link. & Otto var. hispida (Gray) Prain|] HEDGEHOG PRICKLYPOPPY (ARHI) PINACEAE Pinus edulis Engelm. PINYON PINE (PIED) Pinus ponderosa Dougl. PONDEROSA PINE (PIPO) PLANTAGINACEAE Plantago patagonica Jacq. (syn. P. purshi R. & S.) WOOLLY PLANTAIN (PLPA) POLEMONIACEAE Gilia acerosa (Gray) Britt. (syn. G. rigidula Benth. GILIA (GIAC) Ipomopsis laziflora (Coult.) V. Grant (syn. Gilia laziflora (Coult.) Osterh. GILIA (IPLA) F N 12 Cc T N P E a N le E 1p N A Cc 1 N EB Cc a N A Cc 34 PHY TOLO GTA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season POLEMONIACEAE (continued) Ipomopsis pumila (Nutt.) V. Grant (syn. Gilia pumila Nutt.) DWARF GILIA (IPPU) F N A C Ipomopsis spicata (Nutt.) V. Grant (syn. Gilia spicata Nutt. SPIKE GILIA (IPSP) F N 184 Cc Phlox longifolia Nutt. LONGLEAF PHLOX (PHLO) F N If Cc POLYGONACEAE Eriogonum annuum Nutt. ANNUAL ERIOGONUM (ERAN) 1p N A Ww Eriogonum effusum Nutt. BUSHY ERIOGONUM (EREF) F N E Ww Eriogonum jamesii Benth. var. jamesit JAMES ERIOGONUM (ERJA) F N PA WwW Eriogonum lachnogynum Torr. ERIOGONUM (ERLA) Hy N B W Eriogonum tenellum Torr. MATTED WILD BUCKWHEAT (ERTE) F N 1b Ww Eriogonum cf umbellatum Torr. SULPHUR ERIOGONUM (ERUM) E N EP W Shaw et al.: Alphabetic by Family Then by Species POLYGONACEAE (continued) Rumez crispus L. CURLY DOCK (RUCR) Rumerz stenophyllus Ledeb. NARROWLEAF DOCK (RUST) POLYPODIACEAE Woodsia oregana D.C. Eaton OREGON WOODSIA (WOOR) PORTULACACEAE Portulaca oleracea L. COMMON PURSLANE (POOL) Portulaca parvula Gray PURSLANE (POPA) Talinum parviflorum Nutt. PRAIRIE FAMEFLOWER (TAPA) RANUNCULACEAE Clematis ligustictfolia Nutt. WESTERN VIRGINSBOWER (CLLI) Delphinium cf virescens Nutt. ssp. penardii (Huth) Ewan PLAINS LARKSPUR (DEVI1) Floral inventory of Pinon Canyon Site 5 Life Native/ Cool/Warm Form Introduced Longevity Season F I Pp C E I | 2s C E N Pp C F N A Cc EF N A C F N P W V N Pp W F N Pp C 36 PHY TOLDOGIA Alphabetic by Family Then by Species RANUNCULACEAE (continued) Delphinium virescens Nutt. ssp. wootonti (Rydb.) Ewan (syn. D. wootonit Rydb.) WOOTON LARKSPUR (DEVI2) ROSACEAE Amelanchier alnifolia Nutt. SASKATOON SERVICEBERRY (AMAL) Cercocarpus montanus Raf. TRUE MOUNTAIN MAHOGANY (CEMO) Physocarpus monogynus (Torr.) Coult. MOUNTAIN. NINEBARK (PHMO) Potentilla cf arguta Pursh var. arguta WHITE CINQUEFOIL (POAR) Prunus americana Marsh. AMERICAN PLUM (PRAM) Prunus pensylvanica L. f. PIN CHERRY (PRPE) Prunus virginiana L. COMMON CHOKECHERRY (PRVI) Rosa cf woodsii Lindl. WOOD ROSE (ROWO) Rubus deliciosus Torr. BOULDER RASPBERRY (RUDE) volume 67(1):1-42 July 1989 Life Native/ Cool/Warm Form Introduced Longevity Season F N P Cc ) N Pr C ) N e Cc ) N E C F N P Ww S N P C T N je C S N ie Cc S N (2 C S N 1B C Shaw et al.: Floral inventory of Pinon Canyon Site 37 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season RUTACEAE Ptelea trifoliata L. (syn. P. baldwinii T. & G.) COMMON HOPTREE (PTTR) ) N P C SALICACEAE Populus deltoides Marsh. ssp. monilifera (Ait.) Eckenw.; syn. P. sargentti Dode) PLAINS COTTONWOOD (PODE) al N P C Populus tremuloides Michx. QUAKING ASPEN (POTR) ih N P C Saliz cf amygdaloides Anderss. PEACHLEAF WILLOW (SAAM) T N iB Cc Salix exigua Nutt. ssp. interior (Rowlee) Cronq. (syn. S. interior Rowlee) SANDBAR WILLOW (SAEX) S N Ie C Saliz sp. WILLOW (SA-X) S P C SANTALACEAE Comandra umbellata (L.) Nutt. BASTARD TOADFLAX (COUM) F N P C SAPINDACEAE Sapindus saponaria L. var. drummondii (Hook. & Arn.) L. Benson (syn. S. drummondii Hook. & Arn.) SOUTHERN SOAPBERRY (SASA) ah N iz C 38 PHYTOLOGTsA Alphabetic by Family Then by Species SAXIFRAGACEAE Heuchera parvifolia Nutt. LITTLELEAF ALUMROOT (HEPA) SCROPHULARIACEAE Castilleja integra Gray ORANGE PAINTBRUSH (CAIN) Castilleja sessiliflora Pursh DOWNY PAINTBRUSH (CASE) Penstemon angustifolius Nutt. var. caudatus (Heller) Rydb. NARROWLEAF PENSTEMON (PEAN2) Penstemon auriberbis Pennell PENSTEMON (PEAU) Penstemon barbatus (Cav.) Roth. var. torreyi (Benth.) Keck TORREY PENSTEMON (PEBA) Verbascum thapsus L. FLANNEL MULLEIN (VETH) SOLANACEAE Chamaesaracha coronopus (Dun.) Gray GREEN FALSE NIGHTSHADE (CHCO) Lycium pallidum Miers PALE WOLFBERRY (LYPA) volume 67(1):1-42 July 1989 Life Native/ Cool/Warm Form Introduced Longevity Season ig N iP C F N Pp C F N Ie C F N iP C if N |e Cc 1p N P Ww 12 I B Cc E N P Cc S N P C Shaw et al.: Floral inventory of Pinon Canyon Site Alphabetic by Family Then by Species Physalis hederifolia Gray var. cordifolia (Gray) Waterfall (syn. Physalis fendleri Gray) CLAMMY GROUNDCHERRY (PHHE) Physalis lobata Torr. [syn. Quincula lobata (Torr.) Raf.] PURPLEFLOWER GROUNDCHERRY (PHLO1) Physalis longifolia Nutt. var. longifolia [syn. P. virginiana P. Mill. var. sonorae (Torr.) Waterfall LONGLEAF GROUNDCHERRY (PHLO2) Physalis longifolia Nutt. var. subglabrata (Mack. & Bush) Cronq. (syn. P. subglabrata Mack & Bush) TAPERLEAF GROUNDCHERRY (PHLO3) Solanum elaeagnifolium Cav. SILVERLEAF NIGHTSHADE (SOEL) Solanum nigrum L. (syn. S. ptycanthum Dun.) BLACK NIGHTSHADE (SONI) Solanum rostratum Dun. BUFFALOBUR NIGHTSHADE (SORO) Solanum triflorum Nutt. CUTLEAF NIGHTSHADE (SOTR) Form Introduced Longevity Season 40 PHAT OE O GPA Alphabetic by Family Then by Species TAMARICACEAE Tamariz pentandra Pallas (syn. T gallica L.) FIVE-STAMEN TAMARIX (TAPE) TYPHACEAE Typha angustifolia L. NARROWLEAF CATTAIL (TYAN) Typha latifolia L. COMMON CATTAIL (TYLA) ULMACEAE Celtis reticulata Torr. NETLEAF HACKBERRY (CERE2) UMBELLIFERAE Conium maculatum L. POISON HEMLOCK (COMA) Cymopterus montanus T. & G. MOUNTAIN SPRING PARSLEY (CYMO) Lomatium orientale Coult. & Rose ORIENTAL LOMATIUM (LOOR) Musineon divaricatum (Pursh) Nutt. LEAFY MUSINEON (MUDI) volume 67(1):1-42 July 1989 Life Native/ Cool/Warm Form Introduced Longevity Season S I P Cc F N r Cc F N P Cc Jb N P C ip I B Cc F N P C F N bi Cc 15 N P Cc Shaw et al.: Alphabetic by Family Then by Species URTICACEAE Parietaria pensylvanica Muhl. PENNSYLVANIA PELLITORY (PAPE) VERBENACEAE Phyla cuneifolia (Torr.) Greene [syn. Lippia cuneifolia (Torr.) Steud.] WEDGELEAF FOGFRUIT (PHCU) Verbena bipinnatifida Nutt. (syn. V. ambrosifolia Ry db.) DAKOTA VERVAIN (VEBI) Verbena bracteata Lag. & Rodr. PROSTRATE VERVAIN (VEBR) VIOLACEAE Viola nuttallii Pursh NUTTALL VIOLET (VINU) VITACEAE Parthenocissus vitacea (Knerr) Hitchc. THICKET CREEPER (PAVI1) Vitis longi Prince (syn. V. acerifolia Raf. LONGS GRAPE (VILO) Floral inventory of Pinon Canyon Site 41 Life Native/ Cool/Warm Form Introduced Longevity Season F N A C 8 N I? W E N le C F N A Cc F N P Cc Vv N P C V N le Cc 42 PHYTOL O.GTA volume 67(1):1-42 July 1989 Alphabetic by Family Life Native/ Cool/Warm Then by Species Form Introduced Longevity Season ZYGOPHYLLACEAE Tribulus terrestris L. PUNCTURE VINE (TRTE) la I A W ACKNOWLEDGMENTS This work was funded by the U.S. Army Corps of Engineers, Engineering & Housing Support Center (CEHSC-FN), Mr. Donald M. Bandel, Chief of Cultural and Natural Resources. We thank Chris Bern, Kathy Cushman, Jay Wipff, Meg Van Ness, Jon Alstad and Carrie Maenius-Mosley for their assistance. We also thank Richard D. Laven and Dana Quinney for reviewing and commenting on the manuscript. Phytologia (July 1989) G7(1):413-19. Rob C.H.M. Oudejans van Konijnenburglaan 12, 3925 XB Scherpenzeel, The Netherlands ABSTRACT New names and combinations are published for Euphorbia in anticipation of future publication of a listing of all names applied to the tribe Euphorbieae (Euphorbiaceae). KEY WORDS: Euphorbiaceae, Euphorbia, nomenclature. During the preparation of a world list of all species names published in the tribe Euphorbieae (Euphorbiaceae) with their distribution (Oudejans, in preparation), several new names and new combinations appeared to be necessary. The list will contain over 10,000 names, including synonyms, mis- spellings, etc. Prior to the publication of the list, for better accessibility, all name changes will be published in this paper. Currently, the tribe contains the following genera: Euphorbia L., Mon- adenium Pax, Synadentum Boissier, Endadenium Leach, Neogutllauminia Croizat, Pedilanthus Poiteau, Anthostema Adr. Jussieu, Dichostemma Pierre, Calycopeplus Planchon and Cubanthus Millspaugh. All other genera will be treated in my world list as synonyms or sub- genera of the genera mentioned. This holds particularly for Chamaesyce Rafinesque and Tithymalus Gaertner, which are considered as separate gen- era by several authors (for instance, Sojak 1983; Koutnik 1987), whereas others include them in the genus Euphorbia L. (for instance, Radcliffe-Smith 1986; Carter 1988). In my opinion, for stability of nomenclature, the best solution is accepting these names as subgenera, until the tribe Euphorbieae can be studied on a world basis. Most problems of interpretation originate from the vast distribution of the genus Euphorbia L. over all continents with so many pantropic weedy species and different growth forms, resulting in publication of new species and genera on a basis too limited for “correct” concepts accepted universally. 43 44 PHY TOLOGIA volume 67(1):43-49 July 1989 Euphorbia adenoptera Bertoloni subsp. gundlachii (Urban) Oudejans, comb. nov. Basionym: FE. gundlachii |. Urban, Symb. Antill. 5:392-393. 1908. Euphorbia adenoptera Bertoloni subsp. pergamena (Small) Oudejans, comb. nov. Basionym: FE. pergamena J.K. Small, Bull. Torrey Bot. Club 25:615-616. 1898. Euphorbia alainii Oudejans, nom. nov., pro Chamaesyce montana H. Alain [=A.H. Liogier], Moscosoa 1(1):25-26. 1976. The epithet montana is not available in the genus Euphorbia. TYPE: REPUBLICA DOMINICANA. En Pinar, Alto del Toro, Sierra de Bahoruco, 2200 m, Alain & Perfa Liogier 19712. Euphorbia amandi Oudejans, nom. nov., pro E. minuta R.A. Philippi, Anales Mus. Nac. Chile (secc. 2, Bot.) 8:76. 1891. TYPE: CHILE. ha- bitat in Tarapaca, de Cacalhuay advecta. Non EF. minuta F. Loscos y Bernal & J. Pardo y Sastron, Ser. Inconf. Pl. Arag. 96. 1863. Euphorbia antonii Oudejans, nom. nov., pro £. lanceolata D.A. Larrana- ga, Escritos D.A. Larranaga 2:164. 1923. TYPE: [URUGUAY] Julio 21 de 1809. Non E. lanceolata K.[P.J.] Sprengel, Fl. Halens. Mant. 41-42. 1807. Euphorbia barnesii (Millspaugh) Oudejans, comb. nov. Basionym: Ade- nopetalum barnesii C.F. Millspaugh, Field Mus. Nat. Hist., Bot. Ser. Zest VON: Euphorbia blatteri Oudejans, nom. nov., pro E. helioscopioides E. Blat- ter, J. Bombay Nat. Hist. Soc. 36:483. 1933. TYPE: S. Waziristan, Sa- rarogha, 4000 ft, Fernandez 105. Non FE. helioscopioides F. Loscos y Bernal & J. Pardo y Sastron, Ser. Inconf. Pl. Arag. 93. 1863. Euphorbia bruntii (Proctor) Oudejans, comb. nov. Basionym: Chamae- syce bruntit G.R. Proctor, Sloanea 1:2. 1977. F Euphorbia celastroides Boissier in DC var. tomentella (Boissier in DC) Oudejans, comb. nov. Basionym: EF. multiformis Hooker & Arnott var. tomentella [P.|E. Boissier in A.P. de Candolle, Prodr. 15(2):12. 1862. Euphorbia chapmanii Oudejans, nom. nov., pro £. nudicaulis A.W. Chap- man, Fl. South. U.S. (ed. 1) 402. 1860. TYPE: UNITED STATES. near St. Joseph’s, west Florida. Non EF. nudicaulis G. Perrottet, Mem. Soc. Linn. Paris 3(1):116. 1826 (also as preprint 1824). Euphorbia chiogenes (Small) Oudejans, comb. nov. Basionym: Chamae- syce chiogenes J.K. Small, Fl. Southeast. U.S. 709,1333. 1903. Euphorbia conferta (Small) Oudejans, comb. nov. Basionym: Chamae- syce conferta J.K. Small, Fl. Southeast. U.S. 713-714. 1903. Earlier combinations made by B.E. Smith (1946) and E. Jablonski (1974) are invalid. 45 Oudejans: New names and combinations in Euphorbia Euphorbia cordatella Oudejans, nom. nov., pro £. cordata D.A. Larra- naga, Escritos D.A. Larranaga 2:164. 1923. TYPE: (URUGUAY.] Abril 5 de 1814. Non E. cordata F. von Paula von Schrank, Baier. FI. 1:747. 1789. Euphorbia cowellii (Millspaugh ex Britton) Oudejans, comb. nov. Basio- nym: Chamaesyce cowellii [C.F.] Millspaugh ex N.L. Britton, Bull. Torrey Bot. Club 43:457. 1916. Euphorbia cumulicola (Small) Oudejans, comb. nov. Basionym: Chamae- syce cumulicola J.K. Small, Man. Southeast. Fl. 794,1505. 1933. Euphorbia damasoi Oudejans, nom. nov, pro FL. ovata D.A. Larranaga, Escritos Larranaga, 2:164. 1923. TYPE: URUGUAY. Feb 25 de 1812. Non E. ovata E. Meyer ex [P.JE. Boissier in A.P. de Candolle, Prodr. 15(2):167. 1862. Euphorbia deltoidea Engelmann ex Chapman var. serpyllum (Small) Oudejans, comb. nov. Basionym: Chamaesyce serpyllum J.K. Small, Fl. Florida Keys 81. 1913. Euphorbia denisii Oudejans, nom. nov., pro £. obcordata M. Denis, Euphorb. Iles Austr. Afr. 56. 1921. TYPE: MADAGASCAR. Region de Sud-Ouest, bassin de l’Onilany, Perrier de la Bathie 9785. Non E. obcordata [I.]B. Balfour, Proc. Roy. Soc. Edinburgh 12:93. 1882-1883. Euphorbia donii Oudejans, nom. nov., pro EF. longifolia D. Don, Prodr. Fl. Nepal 62. 1825. TYPE: hab. in Nepalia, Hamilton [s.n.]. Non E. longifolia J.B.A.P.M. de Lamarck, Encycl. 2:417. 1788. Euphorbia dubovikii Oudejans, nom. nov., pro E. pinetorum Dubovik ex O.N. Dubovik & M.V. Klokov, Novosti Sist. Vyssh. Inzsh. Rast. (Kiev) 1976:105-107. 1977. TYPE: Dit. Krasnodarensis, prope Gelendzhik, p. Archipo-Ossipovka, O. Dubovik /s.n.]. Non E. pinetorum (Small) G.L. Webster, J. Arnold Arbor. 48:403. 1967. Euphorbia duckei (Croizat) Oudejans, comb. nov. Basionym: Chamae- syce duckez L. Croizat, J. Arnold Arbor. 24:185-186. 1943. Euphorbia georgei Oudejans, nom. nov., pro £. acuta G. Engelmann in W.H. Emory, Rep. U.S. Mex. Bound. 2(1):189. 1858. TYPE: UNITED STATES. Texas: stony prairies western Texas, along the San Pecos rivers, &c. Bigelow [s.n.], Schott [s.n.], Wright 1739,1749. Non E. acuta Bellard ex A. Colla, Herb. Pedem. 5:132. 1836. Euphorbia graminifolia Villars subsp. tommasiniana (Bertoloni) Oudejans, comb. nov. Basionym: E. tommasiniana A. Bertoloni, Fl. Ital. 5(1):78-79. 1842. Euphorbia graminifolia Villars subsp. zhiguliensis (Prokhanov) Oudejans, comb. nov. Basionym: Galarrhoeus (‘Galarhoeus’) zhiguli- ensts Ya.I. Prokhanov, Tr. Kuibyshevsk. Bot. Sada 1:64-68. 1941. 46 PHYTOLOGIA volume 67(1):43-49 July 1989 Euphorbia hamiltonii Oudejans, nom. nov., pro E. angustifolia Hamilton ex D. Don, Prodr. Fl. Nepal 62. 1825. TYPE: hab. ad Norcotera Nepalensium, Hamilion /s.n.]. Non E. angustifolia (Haworth) E.G. von Steudel, Nomencl. (ed. 1) 323. 1821. Euphorbia heraldiana (Millspaugh) Oudejans, comb. noy. Basionym: Chamaesyce heraldiana C.F. Millspaugh, Field Mus. Nat. Hist., Bot. Ser. 2:409. 1916. Euphorbia herbstii (Wagner) Oudejans, comb. nov. Basionym: Chamae- syce herbsti: W.L. Wagner, Bishop Mus. Occas. Pap. 28:75-76. 1988. Euphorbia hiernii (Croizat) Oudejans, comb. nov. Basionym: Elaeophor- bia hiernit L. Croizat, Desert Pl. Life 8:102-103. 1936. Euphorbia hurusawae Oudejans, nom. nov., pro &. barbellata 1. Hurusa- wa, J. Jap. Bot. 16:571-572. 1940. TYPE: CHINA. hab. in Manshuria, prov. Kwantung, in monte Dai-osyo-zan, M. Nishimura [s.n.]. Non E. barbellata G. Engelmann in W.H. Emory, Rep. U.S. Mex. Bound. 2(1):190. 1858. Euphorbia hurusawae Oudejans var. imaii (Hurusawa) Oudejans, comb. nov. Basionym: EF. 1mazi I. Hurusawa, J. Jap. Bot. 16:576-577. 1940. Euphorbia inaguaensis Oudejans, nom. nov., pro £. abbreviata DS. Correll, Sida 8(4):317-319. 1980. TYPE: BAHAMA ISLANDS. Little Inagua, D.S. Correll 45998. Non E. abbreviata J.L. Thuillier, FI. Paris (ed. 2) 239. 1799. Euphorbia josei Oudejans, nom. nov., pro E. litoralis M. [de] Sessé [y Lacasta] & J.M. Mocino, Fl. Mexic. (ed. 2) 123. 1894. TYPE: habitat ad litora maris insulae de Puerto Rico. Non E. litoralis C.S. Kunth in Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 2:54. 1817. Euphorbia klotzschii Oudejans, nom. nov., pro &. ovalifolia (Engelmann ex Klotzsch) [P.]E. Boissier in A.P. de Candolle, Prodr. 15(2):43. 1862. TYPE: CHILE: hab. in Chili prope Santiago [Santjago] et Atacama, Philippi /s.n.J]. Non E. ovalifolia V.F. Kosteletzky, Allg. Med. Pharm. Fl. 5:1724. 1836. Euphorbia larranagae Oudejans, nom. nov., pro E. rupestris D.A. Larra naga, Escritos D.A. Larranaga 2:164. 1923. TYPE: [Uruguay]. Non E. rupestris C.A. Meyer ex C.F. Ledebour, Icon. Pl. Fl. Ross. 2:26. 1830. Euphorbia leoncroizatii Oudejans, nom. nov., pro E. croizatii (Hurusawa) M. Kitagawa, J. Jap. Bot. 31:304 (as ‘Croizati’). 1956. TYPE: CHINA: hab. Manschuria, prov. Fengtien. Non E. croizatit J. Leandri, Notulae Syst. 12:160-161 (as ‘Croizati’). 1946 Oudejans: New names and combinations in Euphorbia 47 Euphorbia limaensis Oudejans, nom. nov., pro E. inornata (Dressler) A. Radcliffe-Smith, Kew Bull. 32:482. 1978. TYPE: PERU. Dept. & Prov. Lima, Distr. Pachacamac, Atacongo, 195 m, Ynes Mezia 04044. Non E. inornata N.E. Brown in W.T. Thiselton-Dyer, Fl. Cap. 5(2):585. 1925. Euphorbia lutulenta (Croizat) Oudejans, comb. nov. Basionym Chamae- syce lutulenta Croizat, J. Arnold Arbor. 26:195-196. 1945. Euphorbia marianoi Oudejans, nom. nov., pro E. pulchella M. [de] Sessé [y Lacasta] & J.M. Mocino, Pl. Nov. Hisp. (ed. 1) 81. 1888 (‘1887’). TENE: [MEXICO?]. habitat in oppidis Ario et Uruapum. Non B&. pulchella M. Lagasca [y Segura] & J. Rodriguez, Anales Cienc. Nat. (Madrid) 5:71-72. 1802. Euphorbia mazatlamensis Oudejans, nom. nov., pro E. triflora M. {de] Sessé [y Lacasta] & J.M. Mocifio, Pl. Nov. Hisp. (ed. 1) 81. 1888 (‘1887’). TYPE: MEXICO. habitat in Mazatlam. Non E. triflora H. Schott, C.F. Nyman & Th. Kotschy, Analecta Bot. 63-64. 1854. Euphorbia mocinoi Oudejans, nom. nov., pro E. peltata M. [de] Sessé [y Lacasta] & J.M. Mocino, Fl. Mexic. (ed. 2) 122. 1894. TYPE: MEXICO. habitat in Oppido Nandio, prope Zitacuarum. Non E. peltata W. Roxburgh, FI. Ind. (ed. 2) 2:474. 1832. Euphorbia nicaeensis Allioni subsp. maleevii (Tamamshyan) Oudejans, comb. nov. Basionym: FE. maleevitS.G. Tamamshyan, Dokl. Akad. Nauk Armyansk. SSR 1(1-2):45-46 (as ‘Maleevi). 1944. Euphorbia nicholasii Oudejans, nom. nov., pro Elaeophorbia acuta N.E. Brown in W.T. Thiselton-Dyer, Fl. Cap. 5(2):222. 1915. TYPE: SOUTH AFRICA. probably the Transvaal, Burtt-Davy [s.n.]. The name acuta is not available in the genus Euphorbia. Euphorbia ocellata Durand & Hilgard subsp. arenicola (Parish) Oude- Jans, comb. nov. Basionym: FE. arenicola S.B. Parish, Erythea 7:93. 1899. Euphorbia ocellata Durand & Hilgard subsp. rattanii (Watson) Oude- jans, comb. nov. Basionym: E. rattaniz S. Watson, Proc. Amer. Acad. Arts Sci. 20:372. 1885. Euphorbia orbifolia (Alain) Oudejans, comb. nov. Basionym: Chamae- syce orbifolia H. Alain [=A.H. Liogier], Phytologia 47:175-177. 1980. Euphorbia paredonensis (Millspaugh) Oudejans, comb. nov. Basionym: Chamaesyce paredonensis C.F. Millspaugh, Field Mus. Nat. Hist., Bot. Ser. 2:393-394. 1914. Euphorbia parodii Oudejans, nom. nov., pro FE. angustifolia D. Parodi, Anales Soc. Cient. Argent. 11:49-50 (as augustzfolia). 1881. TYPE: PARAGUAY. Igatimi in pratis. Non E£. angustifolia (Haworth) E.G. von Steudel, Nomencl. (ed. 1) 323. 1821. 48 PHY, TOL O:G 1A volume 67(1):43-49 July 1989 Euphorbia porteriana (Small) Oudejans, comb. nov. Basionym: Chamae- syce porteriana J.K. Small, Fl. Southeast. U.S. 711-712, 1333. 1903. Euphorbia porteriana (Small) Oudejans var. keyensis (Small) Oudejans, comb. nov. Basionym: Chamaesyce keyensis J.K. Small, Torreya 28:6. 1928. Euphorbia porteriana (Small) Oudejans var. scoparia (Small) Oudejans, comb. nov. Basionym: Chamaesyce scoparia J.K. Small, Fl. Florida Keys 81. 1913. Euphorbia radlkoferi Oudejans, nom. nov., pro B. gracilis L.[A.] Radl- kofer, Leafl. Philipp. Bot. 5(Art. 79):1606-1607. 1913. TYPE: PHILIPPINES. in Philippinarum insula Mindanao, Prov. Agusan, Ca- badbaran, Mount Urdaneta, 4000 ped, A.D.E. Elmer 13482. Non E. gracilis J.L.A. Loiseleur-Deslongchamps, Fl. Gall. (ed. 1) 2:728. 1807. Euphorbia raphilippii Oudejans, nom. nov., pro E. lanceolata R.A. Phi- lippi, Anales Univ. Santiago 91:508. 1895. TYPE: (CHILE.] Habitat in Andibus Illapelinis loco dicto el Pefioa. Non E. lanceolata K.[P.J.] Sprengel, Fl. Halens. Mant. 41-42. 1807. Euphorbia rochaensis (Croizat) Oudejans, comb. nov. Basionym: Cha- maesyce rochaensis L. Croizat, J. Arnold Arbor. 26:195. 1945. Euphorbia rockii Forbes var. grandiflora (Hillebrand) Oudejans, comb. nov. Basionym: E. clustifolia var. grandiflora W. Hillebrand, FI. Hawaiian Isl. 395 (as ‘clusiaefolia’). 1888. Euphorbia roschianica (Ikonnikov) Oudejans, comb. nov. Basionym: Tithymalus roschianicus S.S. Ikonnikov, Novosti Sist. Vyssh. Rast. 15:220-221. 1979. Euphorbia rothrockii (Millspaugh) Oudejans, comb. nov. Basionym: Chamaesyce rothrockit C.F. Millspaugh, Field Mus. Nat. Hist., Bot. Ser. 2:376-377. 1913. ; Euphorbia serpyllifolia Persoon subsp. hirtula (Engelmann ex Watson) Oudejans, comb. nov. Basionym: £. hirtula [G.] Engelmann ex S. Watson, Bot. Calif. 2:74. 1880. Euphorbia sessei Oudejans, nom. nov., pro E. imbricata M. [de] Sessé [y Lacasta] & J.M. Mogino, Fl. Mexic. (ed. 2) 123. 1894. TYPE: habitat ad litore maris prope Havanam. Non £. imbricata M. Vahl, Symb. Bot. 2:54-55. 1791. Euphorbia smallii Oudejans, nom. nov., pro Chamaesyce pinetorum J.K. Small, Bull. N.Y. Bot. Gard. 3:429-430. TYPE: UNITED STATES. [Florida:] between Cutler and Camp Longview Small & Carter 836. The name pinetorum is not available in the genus Euphorbia. Oudejans: New names and combinations in Euphorbia 49 Euphorbia standleyi (Millspaugh) Oudejans, comb. noy. Basionym: Cha- maesyce standleyi C.F. Millspaugh, Field Mus. Nat. Hist., Bot. Ser. 2:404. 1916. Euphorbia tamaulipasana (Millspaugh) Oudejans, comb. nov. Basionym: Chamaesyce tamaulipasana C.F. Millspaugh, Field Mus. Nat. Hist., Bot. Ser. 2:403-404. 1916. Euphorbia triphylla (Klotzsch & Garcke ex Klotzsch) Oudejans, comb. nov. Basionym: Eumecanthus triphyllus Klotzsch & Garcke ex J.F. Klotzsch, Abh. K. Akad. Wiss. Berlin 1859(1):43. 1860. Euphorbia urbanii (Millspaugh) Oudejans, comb. nov. Basionym: Cha- maesyce urbanii Millspaugh, Field Mus. Nat. Hist., Bot. Ser. 2:412. 1916. Euphorbia waldsteinii (Sojak) Radcliffe-Smith subsp. jaxartica (Prok- hanov) Oudejans, comb. nov. Basionym: Tithymalus jazarticus Ya.I. Prokhanov, Consp. Syst. Tithym. As. Med. 192-197. 1933. Euphorbia wangii Oudejans, nom. nov., pro £. villifera Wentsai [=W.T.| Wang, Acta Bot. Yunnanica 10(1):42. 1988. TYPE: CHINA. Gansu: Zhouqiu, alt. 2000 m, 20 Maj 1959, Jiang Shu & Jin Cun-li 361. Non E. villifera A. Scheele, Linnaea 22:153-154. 1849. Euphorbia wootonii Oudejans, nom. nov., pro Zygophyllidium delicatulum E.O. Wooton & P.C. Standley, Contrib. U.S. Natl. Herb. 16(4):145. 1913. TYPE: UNITED STATES. [New Mexico:] Sierra County, Mineral Creek, 2250 m, O.B. Metcalfe 1414. The name delicatula is not available in the genus Euphorbia. LITERATURE CITED Carter, S. 1988. Flora of Tropical East Africa (R.M. Polhill, ed.). Euphor- biaceae (part 2), tribe Euphorbieae. Koutnik, D.L. 1987. A taxonomic revision of the Hawaiian species of the genus Chamaesyce (Euphorbiaceae). Allertonia 4(6):331-388. Radcliffe-Smith, A. 1986. Flora of Pakistan (E. Nasir & S.J. Ali, eds.). Euphorbiaceae. Sojak, J. 1983. Fragmenta phytotaxonomica et nomenclatorica (4). Cas. Nar. Muz. 152(1):12-25. Phytologia (July 1989) 67(1):50-60. EFFECTS OF METOLACHLOR AND ALACHLOR ON PERMEABILITY AND a SYNTHESIS IN SOME PLANT M. Kord! & T. Hathout? ' Botany Department, Faculty of Science, Cairo University, Cairo, EGYPT ? Botany Department, Girls College, Ain Shams University, Cairo, EGYPT ABSTRACT This study was carried out to investigate the effects of meto- lachlor on root permeability and to determine whether meto- lachlor or alachlor inhibit plant lipid synthesis. Metolachlor at 30 and 40 ppm increased the leakage of pre- viously absorbed *?P-labelled orthophosphate from the roots of onion (a susceptible species) by 14 and 41 times the control values respectively. A significant amount of °?P leaked from the roots of the moderately susceptible species (wheat and tomato) whereas no significant loss of °?P occurred to two tolerant species (Pisum and corn). At 10 or 20 ppm, 1,8-naphthalic anhydride prevented 32P leakage from onion roots in the presence of 30 ppm meto- lachlor. High concentrations of naphthalic anhydride, however, promoted **P leakage and did not protect onion roots from the leakage induced by high concentrations (40 ppm) of metolachlor. Neither metolachlor nor alachlor at 40 ppm, inhibited the uptake of acetate-2-!*C or malonic acid-2-''C into excised wheat root tips. Incorporation of ''C-choline chloride into phosphatidylcholine was not significantly inhibited by metolachlor. KEY WORDS: Plant physiology, metolachlor, alachlor, radioac- “tive labeling. INTRODUCTION Metolachlor is a selective herbicide used for the control of grass weeds, nutsedge and broadleaf species in corn. Other crops showing tolerance include soybean, peanut, potato and certain vegetables. Numerous studies have been conducted to determine the mechanism of ac- tion of a-chloroacetamides. Studies of photosynthesis, respiration, a-amylase 50 Kord & Hathout: Metolachlor & alachlor effects on metabolism Dill synthesis, RNA synthesis, protein synthesis and lipid synthesis have failed, however, to elucidate the primary mechanism of action (Chandler et al. 1972; Devlin & Cunningham 1970; Diner et al. 1978; Duke et al. 1975; Jaworski 1956; Moreland et al. 1969; Sasaki et al. 1966 and Truelove & Diner 1978). There is a considerable literature on the effects of metolachlor on the morphology of higher plants. Dixon (1981), came to the conclusion that 100 ppm of metolachlor reduced shoot growth of maize by 65%, and the uptake of the herbicide was twice in maize as in nutsedge. Shoot and root growth are inhibited by a-chloroacetamides (Deal & Hess 1980; Duke et al. 1975; Keeley et al. 1972 and Pillai et al. 1979). Deal & Hess (1980), found that growth of Pisum sativum and Avena sativa were inhibited by both metolachlor and alachlor. They suggested that growth inhibition by these herbicides resulted from an inhibition of both cell division and cell enlargement. Protein synthesis is inhibited by the effect of certain a-chloroacetamides (Mann et al. 1965). Pillai et al. (1979), found that, with metolachlor, leucine incorporation into protein was inhibited only at concentrations of 1 x 10-*M and higher. Diner et al. (1978), found that a-amylase synthesis was inhibited by alachlor and metolachlor only at concentrations in excess of 1 x 10-°M. While conducting studies on the effects of a-chloroacetamides on root growth, the nutrient solution containing 40 ppm metolachlor (where the onion roots were growing) became turbid after 48 hrs and extensive colonies of fungi and bacteria were associated with the roots. This observation suggested the possibility that the herbicide was stimulating the growth of these organisms by causing the leakage of substance from the roots that stimulated microbial growth. This further suggested that the herbicide may be causing a loss of root cell membrane integrity. Studies of the interaction between herbicide and lipids (one of the two major components of cell membrane) have been carried out by Mann & Pu (1968). Using excised hypocotyls of Hemp sesbania, they demonstrated that a number of herbicides, including the a-chloroacetamide can inhibit lipoxygenases as measured by a reduction in the incorporation of malonic acid-2-''C into lipid. MATERIALS AND METHODS Seeds of Allium cepa L. purchased from The Ministry of Agriculture, Cairo, were surface sterilized, and their basal ends were submerged in beakers containing half-strength nutrient solution (Hoagland & Arnon 1950). Roots were allowed to grow for 10 days at 27°C and constant light intensity of about (6,000 lux) in a growth chamber. Nine onions with a similar number of roots 52 Pen! OU OG 1A. volume 67(1):50-60 July 1989 were transferred to beakers containing 60 ml of phosphorus deficient, half strength nutrient solution containing 5 wCi **P as orthophosphate (sp. act. 0.8 mCi/ml). The bulbs were placed so that only the roots were in contact with the radioactive solution. After 24 hrs, onions were removed from the *?P solution and the roots were washed three times with fresh nutrient solution before transferring them to beakers containing 60 ml of half strength nutrient solution, which contained 100 ug/ml penicillin and 40 xg/ml chloramphenicol to inhibit microbial growth. A stock solution of metolachlor in ethanol was added to give final concentrations of 0, 30 or 40 ppm herbicide in each of three replications. A similar amount of ethanol was present in the no-herbicide controls. Duplicate 0.2 ml samples of the solutions were withdrawn after 0, 4, 8 and 12 hrs and then every 12 hrs for 6 days. These samples were radio assayed by Packard liquid scintillation, type spectrometer series 4000. Similar studies were conducted using roots of Lycopersicon lycopersicum L., Zea mays L., Triticum aestivum L. and Pisum sativum L. To show the effect of naphthalic anhydride on metolachlor-induced °?P leakage from onion roots, the experiments were conducted as described above, but naphthalic anhydride was added to some of the treatments. Preliminary experiments were carried out using aqueous naphthalic anhydride suspensions containing 0.1 and 0.4% respectively, applied both alone and with 40 ppm metolachlor. Later experiments used lower rates of both 10, 20 and 30 ppm metolachlor. Wheat grains were germinated for 72 hrs in the dark at 28°C on filter paper moistened with 1 x 10-3M CaCl). After harvesting, 1 cm long root tips were excised and held in cold aqueous sucrose until a sufficient number had been collected. Groups of 100 root tips were removed from the sucrose and transferred to flasks containing 2 ml of 0.01 M potassium phosphate, 1% sucrose and 10 g/ml chloramphenicol. Herbicide, dissolved in ethanol, was added to give a final herbicide concentration of 40 ppm. Ethanol was present in all treatments at a concentration of 0.5%, 5 microcuries of malonic acid- 2-''C (sp. act. 50.4 mCi/ml) were added as a precursor for lipid synthesis. Flasks were covered with black polyurethane and the root tips were in- cubated for 6 hours at 30°C in a shaking water bath. Following incubation, root tips were removed, washed twice with non labeled malonic-acid (200 pg/ml) and twice with water. The root tips were then frozen at 0°C, freeze dried and weighed before grinding in a mortar. The tissue was transferred into 20 ml of chloroform:methanol (2:1) mechanically stirred for one hour and filtered through a double thickness Whatman No. 1 filter paper. The filtrate was dried under a stream of N». The crude lipid extract was dissolved in 5 ml chloroform, 2 ml H:0 was added, and the mixture was shaken. The lower chloroform layer was removed and passed through a small glass column containing anhydrous sodium sulfate to remove residual water Kord & Hathout: Metolachlor & alachlor effects on metabolism ina} and water soluble non lipid residues. This fraction and the upper, aqueous layer (non incorporated precursor) were both sampled and radio assayed by liquid scintillation. Further studies were conducted in the same manner, but using acetate-2- "IC (sp. act. 59.2 mCi/ml) as lipid precursor instead of malonic acid-2-''C. Similar studies were conducted using a longer incubation period of nine hours. Lipid extracts were spotted on pre-coated TLC plates and separated using petroleum ether:diethy] ether:acetic acid (79:10:1). Radioactive areas were detected by scanning the plate with a TLC radio- scanner. The phospholipids, which remained on the TLC plates, were ex- tracted from the silica gel with chloroform:methanol:water (5:5:1). Extracts were dried under N» and redissolved in chloroform prior to further separa- tion by TLC. The remaining lipid classes on the TLC plates were removed by scraping the radioactive areas of silica gel into vials containing scintillation fluid. The vials were then assayed by liquid scintillation. To study the incorporation of choline chloride into phosphatidylcholine, the experiments were carried out in a similar way to malonic acid-2-!'C and acetate-2-''C incorporation studies with the following exceptions 1.0 uCi choline chloride-1-2-'"C (sp. act. 5.4 mCi/ml) was used as a precursor to phosphatidylcholine and the preliminary TLC of the lipid extract to separate the neutral lipids from the polar phospholipids was omitted. Radioactivity remaining at the origin of the TLC plate after separation by polar solvent system was eluted, concentrated and identified as choline chloride using paper chromatography and a solvent system of n-butanol:ethanol:acetic acid:water (8:2:4:3). RESULTS AND DISCUSSION Preliminary studies with onion had suggested that metolachlor was affect- ing the permeability of root membranes, causing leakage of plant metabolites. To determine whether the effects of metolachlor on permeability were related to the phytotoxicity range of the herbicide, several species differing in sensitivity to metolachlor were tested (Figure 1). In the absence of herbicide, there was very little leakage of *?P from the roots of any of the species, indicating that the antibiotic and alcohol in the nutrient solutions had little if any effect on root permeability. Neither herbicide treated Zea nor Pisum (both metolachlor tolerant species) showed any significant leakage of *?P from the roots compared to untreated plants, even at an herbicide concentration of 40 ppm. The two moderately susceptible species (tomato and wheat) showed three and four times the untreated plant level of ‘*P leakage respectively, after 144 hours treatment with 30 ppm metolachlor. In the presence of 40 ppm metolachlor, however, *?P leakage from these two species was 14 and 11 times the control rates respectively. < July 1989 volume 67(1):50-60 Pare tt OL OG TA 54 J 50 ppm metolachtor AS 40 ppa metoluchior C maze -J) 50 ppm metolucttor KAY. 40 ppm metotachtor 72 time (hours) ( Pisum (Sw /wdp) abox0a; zed time (hours) CD 30 ppm metolachlor GSS 40 ppm metolachlor 9 {omato C30 ppm metolachlor 40 ppm metolachlor wheat 16 vv xn oOo o © vv Nn oO (6w/wdp) aboxoa; zed time (hours) time (hours) onion [) 30 opm metolachlor 40 opm metolachlor 50 40 30 20 10 8) (6w/wdp) e6ox09) zed time (hours) Figure 1. The effect of metolachlor on the leakage of previously absorbed 32P from roots of five plants. Kord & Hathout: Metolachlor & alachlor effects on metabolism 55 The maximum leakage of previously absorbed °’P was from the roots of onion, a species highly sensitive to metolachlor. At 30 ppm metolachlor, *?P leakage from onion roots was 14 times that of untreated roots after 144 hours. At 40 ppm metolachlor, *?P leakage was 41 times the control value. Leakage of 32P by onion roots was significantly different from the control after 72 hours with the 40 ppm metolachlor concentration. At its most extreme, the amount of °?P released from herbicide treated onion roots was about 10% of the total amount absorbed. Because °*’P absorbed by roots is rapidly translocated to the shoot system, however, the amount lost would represent considerably more than 10% of the **P remaining in the roots. The reason for the differing rates of *?P leakage, however, is not known. From these studies, it was not possible to determine whether metolachlor was exerting a direct effect on root membranes, or whether the leakage of exudates was a secondary effect resulting from an inhibition by the herbicide of some metabolic process related to membrane function or to the maintenance of membrane integrity. Seed protectants can confer on germinating seeds a higher tolerance to herbicides and thus widen the margins of safety and selectivity. The potential of using naphthalic anhydride as a safener against metolachlor and alachlor injury has been tested, and it has been shown that naphthalic anhydride can prevent herbicide injury from these herbicides in sorghum (Ahrens & Davis 1978; Jordan & Jolliffe 1971 and Truelove & Davis 1977) and against alachlor injury to Zea (Burnside et al. 1971). These studies were conducted to determine whether naphthalic anhydride could protect onion roots from the permeability changes induced by metolachlor. The results are shown in Figure 2. Metolachlor at a concentration of 30 ppm applied to onion roots via nutri- ent solution caused a 15 fold increase in the leakage of *’P after 144 hours of treatment (Figure 2). When naphthalic anhydride at either 10 or 20 ppm was applied to onion roots with metolachlor at a concentration 30 ppm, however, very little 9*P leakage occurred. Thus, naphthalic anhydride protected the onion roots against permeability changes induced by metolachlor. Although the mechanism of action of metolachlor is unknown, the studies indicate that leakage of plant nutrients and loss of root cell membrane integrity are prob- ably important factors in the mode of action of this herbicide. Naphthalic anhydride applied alone at concentrations of 10 ppm did not cause leakage of °?P from the roots. However, at high rates of application, naphthalic anhydride caused leakage of previously absorbed “*’P from onion roots. Such high rates of naphthalic anhydride did not protect onion roots from leakage induced by metolachlor. In fact, they promoted additional leakage. St. John & Hilton (1973), reported the efflux of electrolytes from roots of wheat seedlings treated with 1 x 10~'M dinoseb. They found that herbi- ial for) PHYTOLOGIA volume 67(1):50-60 July 1989 18 16 14 12 () 48 hours 96 hours CZ) 144 hours MC = Metolachlor ie NA = Naphthalic anhydride 8 6 4 2 (6) ps2 leakage (dpm treatment/dpm control) MC NA MC + NA MC + NA (30ppm) (10ppm) (20ppm) (10ppm) TREATMENT Figure 2. The effect of 1,8-naphthalic anhydride on metolachlor induced °2P leakage from the roots of onion. cide treatment reduced polar lipid levels by 72% and suggested that dinoseb may act by decreasing the levels of polar lipids required for membrane for- mation, hence altering membrane structure and function. Alteration of root membrane permeability characteristics, as shown by the leakage of plant ex- udates, suggested to us that metolachlor might also be acting through some effect on those metabolic processes involving membrane lipids. The effects of metolachlor and alachlor on lipid synthesis were determined by following the uptake and incorporation of radio labeled precursors into the lipids of excised wheat root tips. The uptake of the precursor, acetate-2-''C, by both treated and untreated root tips was much greater than that of malonic acid-2-''C (Figure 3). Between 56 and 63% of the applied acetate-2-''C was absorbed by excised root tips, but only 7 to 9% of the applied malonic acid-2-''C was absorbed. Neither alachlor nor metolachlor, however, affected the uptake of either of these precursors. The radioactivity of the total lipid fraction extracted from herbicide treated tissue was not significantly different from that of untreated tissue, irrespective of which ''C-labeled precursor was used in the study (Figure 3). The amount of radioactivity incorporated into the lipids, however, differed with the precursor used, with 7 to 9% of the applied acetate-2-!'C and 0.6 to 0.8% of the applied malonic acid-2-''C incorporated into the lipids. Kord & Hathout: 100 [-] applied els CX) phospholipids pa) Tottyderds MM triglycerides [XJ unknown lipids SSS SD Se SS SE ez [NNN NS SSS eee SST LT EG LE LG TP [PT EY ! EF LG LPL aS ee Sp a =. PD ED GD > SD Gah ee eG GE eae SS LE LD) EY LD LS LD IEP f PLE LG LF SES LOI, aes. FS ee GP ee Ga ee SD SS 2 eS Ss [Nee SNe Nae Ne SS SS EL ES LY (A LE LSD SEE LEE LEE Se aa a SSD Se SDS = SS SSS _ SD NSN NN ESSE] SD LE LE SP EF LD EL LF LEV LET LOD LS GD iG GS Ge SS Se E22 GBS GPS SD GSS SS DED [ENS ee ee Ne NNN SENS LSE LE ED SES LD (EP LES [I LG, PG, EC LYS (yr ee ee ee ee Pe ee juaoied CO ME (40ppm) (40ppm) AL CO Metolachlor & alachlor effects on metabolism added molonic Cees soda |_ added acetate ele of Figure 3. The effects of metolachlor on the incorporation of acetate-2-!'C and malonic acid-2-!'C into lipids of wheat root tips. 58 PUPPY POLO GTA volume 67(1):50-60 July 1989 After separation of the total lipid extracts into the constituent lipid classes by TLC, the effect of the herbicide treatment on lipid synthesis could be de- termined. The incorporation of radio label into the different lipid classes was not inhibited by herbicide treatment. There were no significant differences in either the radio labeled phospholipid fraction or the neutral lipid classes in the total lipid extract. The phospholipids were eluted from TLC plates and separated into the constituent phospholipid classes by TLC to determine whether alachlor or metolachlor inhibited the synthesis of specific phospholipids. Radio assay indicated that the synthesis of specific phospholipids was not inhibited by either herbicide. No evidence was found to support the contention that a six hour exposure to alachlor or metolachlor inhibited lipid or phospholipid synthesis in excised wheat root tips. The results of the investigation using a nine hour incubation period were similar to those for the six hour incubation, with no evidence of inhibition of either lipid or phospholipid synthesis by alachlor or metolachlor. Earlier work had shown significant inhibition of total lipid synthesis by cotton root tips with a metolachlor treatment, although not with an alachlor treatment. More specifically, we had shown that total phospholipid synthesis was inhibited by both metolachlor and alachlor. Wilkinson (1981), failed to confirm these observations and there was no inhibition of lipid or phospholipid by either herbicide in this investigation. To pursue these investigations further, wheat root tips were incubated with choline chloride-1,2-''C, a precursor for phosphatidylcholine. After sep- aration of the total lipid extract into the constituent lipid classes by TLC, the plates were radio scanned. The scans showed the presence of only two areas of radioactivity on the TLC plates, an area at the origin which ac- counted for 30% of the total radioactivity, and an area that corresponded to the phospholipid standard, phosphatidylcholine. The radioactivity at the origin was eluted from the silica gel and later identified as choline chloride- 1,2''C by paper chromatography. The results presented in Figure 4 show that the incorporation of choline chloride-1,2-''C into phosphatidylcholine was not inhibited by metolachlor. Thus, although metolachlor induces leakage of *?P from the roots of sus- ceptible species and causes subsequent loss of root cell membrane integrity, we have found no evidence, under these experimental conditions, that this loss of membrane integrity is due to the inhibition of total lipid, phospholipid or phosphatidylcholine synthesis by either alachlor or metolachlor. percent Kord & Hathout: Metolachlor & alachlor effects on metabolism 59 100 [_] choline chloride CX) phosphatidylcholine 80 60 40 20 control metolachlor (40ppm) Treatment Figure 4. The effect of metolachlor on the incorporation of choline chloride 1,2-''C into phosphatidylcholine. LITERATURE CITED Ahrens, W.H. & D.E. Davis. 1978. Seed protectant effects on metolachlor absorption and translocation. Proc. South. Weed Sci. Soc. 31:249. Burnside, O.C., G.A. Wicks & E.R. Fenster. 1971. Protecting corn from herbicide injury by seed treatment. Weed Sci. 19:565-568. Chandler, J.M., L.M. Croy & P.W. Santlemann. 1972. Alachlor effects on plant nitrogen metabolism and Hill reaction. J. Agric. Food Chem. 20:66 1-664. Deal, L.M. & F.D. Hess. 1980. An analysis of the growth inhibitory char- acteristics of alachlor and metolachlor. Weed Sci. 28:168-175. Devlin, R.M. & R.P. Cunningham. 1970. The inhibition of GA3 induction of a-amylase activity in barley endosperm by certain herbicides. Weed Res. 10:316-320. Diner, A.M., B. Truelove & D.E. Davis. 1978. Metolachlor effects on lipid metabolism in cotton root tips. Proc. South. Weed Sci. Soc. 31:250. Dixon, G.A. 1981. Uptake translocation, metabolism, and the effects of metolachlor on yellow nutsedge and corn. (Dissertation Abstracts In- ternational, B. 41(11):19-70. Duke, W.B., F.W. Slife, J.B. Hanson & H.S. Butler. 1975. An investigation on the mechanism of action of propachlor. Weed Sci. 23:142-147. 60 PHYTOL O:Gir A volume 67(1):50-60 July 1989 Hoagland, D.R. & D.I. Arnon. 1950. The water-culture method for growing plants without soil. Calif. Agric. Exp. Stn. Circ. #347. 32 pp. Jaworski, E.G. 1956. Biochemical action of CDAA, a new herbicide. Science 123:847-848. Jordan, L.S. & V.A. Jolliffe. 1971. Protection of plants from herbicides with 1,8-naphthalic anhydride as illustrated with sorghum. Bull. Environ. Contam. Toxicol. 6:417-421. Keeley, P.E., C.H. Carter & J.H. Miller. 1972. Evaluation of the relative phytotoxicity of herbicides to cotton and nutsedge. Weed Sci. 20:71-74. Mann, J.D., L.S. Jordan & B.E. Day. 1965. A survey of herbicides for their effect upon protein synthesis. Plant Physiol. 40:840-843. Mann, J.D. & M. Pu. 1968. Inhibition of lipid synthesis by certain herbi- cides. Weed Sci. 16:197-198. Moreland, D.E., S.S. Malhotra, R.D. Gruenhagen & E.H. Shokrau. 1969. Effects of herbicides on RNA and protein synthesis. Weed Sci. 17:556- 563. Pillai, P., D.E. Davis & B. Truelove. 1979. Effects of metolachlor on ger- mination, growth, leucine uptake, and protein synthesis. Weed Sci. 27:634-637. Sasaki, S. & T.T. Kozlowski. 1966. Influence of herbicides on respiration in young Pinus seedlings. Nature (London) 210:439-440. St. John, J.B. & J.L. Hilton. 1973. Lipid metabolism as a site of herbicide action. Weed Sci. 21:477-480. Truelove, B. & D.E. Davis. 1977. A herbicide antidote seed treatment with naphthalic anhydride. Proc. South. Weed Sci. Soc. 30:364. Truelove, B. & A.M. Diner. 1978. Some effects of metolachlor on plant metabolism. Abstr., Weed Sci. Soc. Am. p. 68. Wilkinson, R.E. 1981. Influence of metolachlor on phospholipid synthesis in cotton, soybean and sorghum. Proc. South. Weed Sci. Soc. 34:263. Phytologia (July 1989) G7(1):61-66. THE SEPARATION OF TRIMORPHA (COMPOSITAE: ASTEREAE) FROM ERIGERON Guy L. Nesom Department of Botany, University of Texas, Austin, Texas 78713 USA ABSTRACT The genus Trimorpha Cass. (Erigeron sect. Trimorpha) is re- segregated from Erigeron. The filiform, eligulate pistillate florets and mature pappus that lengthens past the involucre in Trimor- pha are not found in Erigeron. Also, the outer phyllaries with three, orange nerves, which are characteristic of Trimorpha, are known in Erigeron in only the three species of sect. Spinost. In these features of the flowers, pappus and phyllaries, Trimorpha is more similar and apparently more closely related to Conyza than to Erigeron. Six new combinations to Trimorpha are proposed to accommodate the American taxa: T. acris var. asteroides, T. a. var. debilis, T. a. var. kamtschatica, T. elata, T. lon- chophylla and T. scotteri. KEY WORDS: Trimorpha, Erigeron, Asteraceae, New World, systematics. Erigeronsect. Trimorpha(Cass.) DC. includes a group of species set apart from the rest of the genus by the production of two zones of pistillate flowers, an inner zone of ca 1-4 series of eligulate flowers and an outer of 1-3 series of very numerous flowers with short, narrow, often filiform ligules. Cronquist (1943, p. 629) saw sect. Trimorpha as “inextricably bound to Erigeron ... by the obvious evolutionary line of FE. simplez Greene, E. uniflorus L. (sensu lat.) and £. alpinus L., in which E. simplez is true Erigeron, E. alpinus is Trimorpha, and E. uniflorus is somewhat intermediate.” In my view, how- ever, FE. uniflorus is not intermediate between Trimorpha and true Erigeron, as discussed below, and the two groups are best regarded as different genera. Trimorpha is distinguished most conspicuously from all of Erigeron by its dimorphic pistillate flowers. At least in some species of Trimorpha, the ligules loosely coil at maturity. The plants are mostly perennials from short, fibrous-rooted rhizomes and produce few-flowered capitulescences that vary from loosely cylindrical panicles or racemes to corymbs. In a few species the heads are solitary. The leaves are entire and buds erect. The outer phyllaries 61 62 PHY POL/O'G IEA volume 67(1):61-66 July 1989 have three orange-resinous veins, a feature characteristic of Conyza (sensu Nesom [in press]) but found in Erigeron only in the three species of sect. Spinosi (Nesom 1989) and a few others in scattered groups. This feature of nervation is sometimes difficult to observe if the phyllaries are dark-colored, but it is distinctive and occurs in every species of Trimorpha that I have studied. The pappus in plants of Trimorpha also resembles that of Conyza in becoming prominently longer at maturity than the involucre, a feature diagnostic of Conyza but not found in Erigeron. Erigeron lonchophyllus is a fibrous-rooted annual and, alone in Trimor- pha, has only a zone of ligulate, pistillate flowers, lacking the inner zone of eligulate flowers. It clearly belongs with Trimorpha, however, on the basis of its 3-nerved outer phyllaries and elongated pappus. Because of its relatively specialized habit and duration, I believe the lack of the eligulate flowers is a specialization, reflecting a loss rather than a primitive similarity with true Erigeron. With regard to Cronquist’s assertion that Erigeron uniflorus, the gener- itype of Erigeron (Nesom 1989), occupies an intermediate position between Erigeron and Trimorpha, I find that the putative intermediacy of FE. uni- florus lies only in its narrow ligules. In contrast, the ligules do not coil, the outer phyllaries are 1-nerved, and the pappus does not elongate at maturity, features that clearly link it with Erigeron rather than Trimorpha. Trimorpha, then, differs from Erigeron in several characters, and in these same characters it is similar to Conyza. In my opinion, it is set apart as a genus from Erigeron with at least as much justification as Conyza and appears to be more closely related to Conyza. Cronquist (1943, p. 631) noted that “In surveying the numerous species which link true Erigeron to true Conyza, we find that in only one place is there any suggestion of a real break. That is between Trimorphaea and Coenotus.” With Cronquist, I regard Caenotus as true Conyza (Nesom [in press|). Cronquist also noted (1943, p. 630) that “The differences between Coenotusand Trimorphaeaare not great, but the species of the two groups do not seem intimately related. Erigeron canadensis, the most nearly bridging species of the section {Coenotus], is scarcely confusable with any species of Trimorphaea.” The morphologically distinct zones of pistillate flowers are not known from any species of Conyza. In addition to this difference, Conyza is a genus primarily of the southern hemisphere, though some of its species are more widespread; Trimorphais confined to arctic-alpine or temperate regions of the northern hemisphere, and several species are circumboreal. An alternative taxonomic treatment of Trimorpha would be to recognize it as a well-defined section of Conyza. This would emphasize its similarity to Conyza but would require a much greater number of nomenclatural com- binations than the approach taken here, since at least 40 of the Old World Nesom: Segregation of Trimorpha from Erigeron 63 taxa already have names as Trimorpha. TRIMORPHA Cass., Bull. Sci. Soc. Philom. Paris 1817:137. 1817. TYPE: Trimorpha vulgaris Cass. in Cuvier, Dict. Sci. Nat. 55:324. 1828 (=E£. acris L.). Trimorphaea Cass. in Cuvier, Dict. Sci. Nat. 37:462. 1825. Erigeron sect. Trimorpha (Cass. in F. Cuvier) DC., Prodr. 5:290. 1836. Erigeron subg. Trimorpha(Cass.) M. Popov, Acta Inst. Bot. Acad. Sci. URSS, Ser. 1, Fasc. 7:10. 1948. In the original publication of the genus Trimorpha (1817), Cassini cited Erigeron acris L. as its sole constituent. Not until 1828 did he name a species in the former genus, and there he cited EZ. acris as a synonym of T. vulgaris. In 1825, Cassini began using the orthographical variant Trimorphaea, listing the original Trimorpha as a synonym. | have discussed other aspects of the lectotypification of Erigeron and Trimorpha in a separate paper (Nesom 1989). Trimorpha sect. Brachyglossae Vierh., Beih. Bot. Centralbl. 19:423. 1906. LECTOTYPE (designated here): T. acris (L.) S.F. Gray (=Erigeron acris L.). Trimorpha sect. Macroglossae Vierh., Beih. Bot. Centralbl. 19:424. 1906. LECTOTYPE (designated here): T. alpina (L.) S.F. Gray (=Erigeron alpinus L.). Erigeron (sp.-group) Acres Rydb., Fl. Colorado 359.1906, in clave. TYPE: E. acris L. Tessenia P. Bubani, Fl. Pyrenaea 2:264. 1899. LECTOTYPE (designated here): Tessenia alpina (L.) P. Bubani (=Erigeron alpinus L.). Tessenia was a superfluous and substitute name for Erigeron by Bubani. It was later used by Lunnell (1917). There are about 40-45 North American and Eurasian species in Trt- morpha. The whole group is in need of critical taxonomic study. From Eurasia, Vierhapper (1906) treated Trimorpha as a genus and included 26 species; Botschantzev (1959) included 17 species in Trimorpha as a subgenus of Erigeron. In Flora Europaea (Tutin et al., 1976), nine species are treated as Erigeron. From North America, Cronquist (1947) recognized only two species in EF. sect. Trimorpha, E. lonchophyllus and E. acris, the latter with several varieties. In his study of the Alaskan flora, where all the Ameri- can taxa of Trimorpha occur, Hultén (1968b) recognized FE. elatus (E. acris var. elatus sensu Cronquist) as a distinct species and added one taxon (E. acris var. kamtschaticus) known from a single collection on the Alaska- Yukon boundary. I propose combinations to Trimorpha for the American taxa as recognized by Hultén, leaving the taxonomy of the Old World species for botanists better acquainted with those species. 64 PHY POLO GLIA volume 67(1):61-66 July 1989 NEW WORLD AND CIRCUMBOREAL TAXA Trimorpha acris (L.) S.F. Gray, Nat. Arr. Brit. Pl. 2:466. 1821. Erigeron acris L., Sp. Pl. 863. 1753. Trimorpha acris var. debilis (A. Gray) Nesom, comb. nov. Erigeron acris var. debilis A. Gray, Syn. Fl. N. Amer. 1(2):220. 1884. Erigeron debilis (A. Gray) Rydb., Mem. N.Y. Bot. Gard. 1:408. 1900. Trimorpha acris var. kamtschatica (DC.) Nesom, comb. nov. Erigeron kamtschaticus DC., Prodr. 5:290. 1836. Erigeron acris subsp. kamtscha- ticus (DC.) Hara, J. Jap. Bot. 15:317. 1939. Erigeron acris var. kamtschaticus (DC.) Herder, Bull. Soc. Nat. Moscou Sect. Biol., Ser. 2. 38:392. 1865. Trimorpha acris var. asteroides (Andrz. ex Besser) Nesom, comb. nov. Erigeron asteroides Andrz. ex Besser, Enum. Pl. Volh. 33. 1822. Erigeron acris var. asteroides (Andrz. ex Besser) DC., Prodr. 5:290. 1836. Erigeron politus E. Fries, Summa Veg. Scand. 3:184.1845. Erige- ron acris subsp. politus (E. Fries) H. Lindb. f., Enum. Pl. Fennoscand. Orient. 56. 1901; non Schinz & Keller, 1909. Hultén (1968a) suggested that Erigeron asteroides was not known from America. By the publication of his flora (1968b), however, he had apparently changed his mind, because he included F. acris var. asteroides as a synonym of E. acris subsp. politus. Trimorpha elata (Hook.) Nesom, comb. nov. Erigeron alpinus y elatus Hook., Fl. Bor. Amer. 2:18. 1834. Erigeron elatus (Hook.) E. Greene, Pittonia 3:164. 1897. Erigeron acris var. elatus (Hook.) Cronq., Brit- tonia 6:296. 1947. I accept Cronquist’s argument (1947, p. 297) that Hooker’s varietal name is valid and that Greene’s name is homotypic and synonymous with it. Trimorpha lonchophylla (Hook.) Nesom, comb. nov. Erigeron lon- chophyllus Hook., Fl. Bor. Amer. 2:18. 1834. Trimorpha scotteri (B. Boivin) Nesom, comb. nov. Erigeron scotteri B.Boivin, Phytologia 23:52. 1972. According to Boivin (1972), Trimorpha scotteri includes plants that were suggested by Cronquist (1947) to be hybrids between Erigeron humilis Grah. and T. acris var. debilis. REPRESENTATIVE OLD WORLD TAXA Trimorpha alpina (L.) S.F. Gray, Nat. Arr. Brit. Pl. 2:467. 1821. Erigeron alpinus L., Sp. Pl. 864. 1753. Nesom: Segregation of Trimorpha from Erigeron 65 Trimorpha attica(Vill.) Vierh., Beih. Bot. Centralbl. 19:462. 1906. Erzgeron atticus Vill., Hist. Pl. Dauph. 3:237. 1788 (incl. E. villarsiz Bellardi). Trimorpha borealis Vierh., Beih. Bot. Centralbl. 19:447. 1906. Erigeron borealis (Vierh.) Simm., Lunds Univ. Arsskr. n.s. 9:127. 1913. Trimorpha epirotica Vierh., Beih. Bot. Centralbl. 19:446. 1906. Erigeron epirotica (Vierh.) Halacsy, Consp. Fl. Graec., Suppl. 53. 1908. Trimorpha neglecta (A. Kerner) Vierh., Beih. Bot. Centralbl. 19:451. 1906. Erigeron neglectus A. Kerner, Osterr. Bot. Zeitschr. 21:253. 1871. Erigeron orientalis Boiss., Diagn. Pl. Orient. Nov. 3:7. 1856. Apparently no name as Trimorpha. Trimorpha podolica (Besser) Vierh., Beih. Bot. Centralbl. 19:423. 1906. Erigeron podolicus Besser, Enum. Pl. Volh. 76. 1822. ACKNOWLEDGMENTS I appreciate the review and comments of Drs. B.L. Turner, T.P. Ra- mamoorthy and J. Kartesz. LITERATURE CITED Boivin, B. 1972. Erigeron. In Flora of the prairie provinces. Phytologia 23:43-55. Botschantzev, V.P. 1959. Erigeron. Pp. 191-288 in Flora U.R.S.S., vol. 25, ed. V.L. Komarov. U.S.S.R. Acad. Sciences, Leningrad. Cronquist, A. 1943. The separation of Erigeron from Conyza. Bull. Torrey Bot. Club 70:629-632. . 1947. Revision of the North American species of Erigeron, north of Mexico. Brittonia 6:121-302. Hultén, E. 1968. Comments on the flora of Alaska and Yukon. Arkiv Bot., n. ser. 7:1-147. 1968. Flora of Alaska and Neighboring Territories. Stanford University Press, Stanford, California. Lunnell, J. 1917. Enumerantur plantae Dakotae septentrionalis vasculares— XII. Amer. Midl. Nat. 5:55-71. 66 Py TOG. O GT A volume 67(1):61-66 July 1989 Nesom, G.L. 1989. Infrageneric taxonomy of New World Erigeron (Com- positae: Astereae). Phytologia 67:67-93. (in press). Further definition of Conyza (Compositae: Astereae). Phytologia 67:. Tutin, T.G. et al. 1976. Erigeron. Pp. 116-120 in Flora Europaea, vol. 4. Cambridge Univ. Press, Cambridge. Vierhapper, F. 1906. Monographie der alpinen Erigeron—Arten Europas und Vordasiens. Beih. Bot. Centralbl. 19:385-560. Phytologia (July 1989) G7(1):67-93. ant TAXONOMY OF NEW WORLD ERIGERON (COMPOSITAE: ASTEREAE) Guy L. Nesom Department of Botany, University of Texas, Austin, Texas 78713 USA ABSTRACT In a synopsis of infrageneric taxonomy of New World Erigeron, E. uniflorus L. is accepted as the lectotype of sect. Erigeron; other lectotypes are proposed for a number of sections or generic seg- regates by G. Don, Nuttall, de Candolle, Rafinesque, Torrey & Gray, Botschantzev and Vierhapper. Eighteen sections, count- ing sect. Leptostelma of South America, are currently recognized in the New World, eight of which are new: sect. Arenarioides, sect. Imbarba, sect. Cincinnactis, sect. Karvinskia, sect. Os- teocaulis, sect. Scopulincola, sect. Spathifolium and sect. Spinosi. Five of these are segregates from sect. Erigeron sensu Cronquist. Sect. Linearifolii, comb. nov., replaces sect. Pycno- phyllum Cronq. The justification for maintaining Darwiniotham- nus as a distinct genus is weak; it should be returned to Erigeron. Erigeron veracruzensis nom. nov. replaces the later homonym E. scaberrimus (Less.) Nesom. KEY WORDS: Erigeron, Asteraceae, New World, systematics. I present a synopsis of the taxonomy and nomenclature of the New World (primarily North American) species groups of Erigeron. In this, I have at- tempted to list all of the accepted species that occur in North America (in- cluding México) and Central America. I have not dealt with South American species included by Solbrig (1962) in sect. Erigeron; | am studying the species of Erigeron (ca 20) in the Antilles. This infrageneric taxonomy certainly will undergo modifications as concepts of relationships within the genus are de- veloped and refined and as other names may be discovered in surveys of older literature. I believe it is important, however, to point out as clearly as possible the great amount of diversity that exists within Erigeron. This is particularly significant as botanists begin comprehensive investigations of the relationships among genera of Astereae. The approach taken here differs strongly from Cronquist’s more inclusive view (1947) of sect. Erigeron, which I have partitioned into eight sections. Such hypotheses represent a step to- ward understanding phylogenetic patterns within this large group of species, 67 68 PHY T-O2)'0 GIA volume 67(1):67-93 July 1989 and I believe there is at least heuristic value in presenting them as formal taxonomic proposals. The “probable relationships” diagram presented by Cronquist (1947, p. 125) and the comments in the text of his revision have been valuable in indi- cating groups of inter-related species, and his insights remain largely correct. In attempting simply to delimit monophyletic groups, however, I hope to provide a useful and more objective taxonomy. Even then, because of the large degree of suspected morphological parallelism in the genus, hypothe- ses regarding the constitution of single lineages are speculative to a degree, but those regarding the phyletic derivations between species and lineages are even more so. Molecular approaches may be more incisive than strictly morphological ones in discovering patterns of common ancestry. In my opinion, among the most important morphological characters for indicating sectional relationships in Erigeron are the behavior of the ligules of the ray flowers (coiling, reflexing, remaining straight, or closing upwards at night) and the behavior of the buds (erect, sharply nodding or arching- pendant). These are features of necessity observed in the field, because they often are not preserved in pressed and dried specimens. The plant habit and the nature of leaf insertion are also significant. Clasping leaves are almost completely restricted to the plants of sect. Cincinnactis, sect. Fruticosus, sect. Olygotrichium and sect. Polyactis, although this feature is not strictly diagnostic of any of these groups. Leaf lobing or toothing is characteristic of only six sections. Some species of sect. Wyomingia typically have numerous (4-14) achenial ribs in contrast to the usual number in the genus, two per achene. I believe that achene shape will also prove to be a significant feature in assessing relationships, but I have not emphasized that feature in this study. Basally caducous pappus bristles occur in only one section (sect. Polyactis), where they are diagnostic. Asa Gray recognized six sections of Erigeron in 1841 but by 1884 had reduced the number to three (sects. Erigeron, Trimorpha and Caenotus), submerging the others into sect. Erigeron. Cronquist included 99 of the 133 of the North American species (in 1947) in a heterogeneous sect. Erigeron. Solbrig (1960, 1962), too, placed most of the South American species of the genus in sect. Erigeron, recognizing only three species in sect. Leptostelma (see below). In a treatment of the Compositae of the U.S.S.R, Botschantzev (1959) recognized three subgenera: subg. Erigeron (four sections), subg. Trimorpha (two sections), and subg. Conyzastrum (Boiss.) M. Pop. [sect. Conyzastrum and sect. Psychrogeton (Boiss.) Botsch.]. The latter section was recently treated by Grierson & Reichinger (1982) as a separate genus, and I regard Trimorpha as a genus distinct from Erigeron (Nesom 1989b). I attempted to deal with some of Botschantzev’s nomenclature here because a few of the Nesom: Infrageneric treatment of Erigeron 69 Asian and American species and groups cross the Bering Straits and occur on both continents. Conyzastrum and Psychrogeton are not included, because all the species appear to be Asian. Among the first sectional names in Erigeron were those proposed in 1830 by G. Don in Loudon’s Hortus Britannicus (see Sundberg & Jones 1987; 1988). Most of Don’s groups in Erigeron were highly polyphyletic, and I have tried to lectotypify them in a way that will displace as few new names as possible in Erigeronand related genera. The name of one of Don’s sections, however, must supplant a name in current usage, and one of them in Aster, as recently lectotypified with a species of Erigeron by Sundberg & Jones, assumes a position of priority over a name already in use in Erigeron. Nuttall (1818, 1840) proposed names for several sections of Erigeron and I have tried to maintain these with appropriate lectotypes where possible. Such is also true for names proposed by Torrey & Gray (1841). Rydberg (1906, 1918) introduced numerous infrageneric names as sub- headings of his keys to species of Erigeron. In an earlier paper (Nesom 1982), I did not regard these as formal taxonomic proposals because Rydberg gave no indication of the intended rank of these groups and provided neither de- scriptions nor diagnoses. Further, several species of Erigeron key out in more than one subdivision. Here, however, following Jones (1980) in Aster, I have adopted Holmgren’s point of view (1979) in regarding Rydberg’s names as validly published, unranked species groups that may be used as basionyms in future combinations. In contrast, Macbride & Payson (1917) accepted one of Rydberg’s aggregate names (see sect. Tridactylia below) with the following rationale: “... from the fact that the name was given in the plural form and appears to be just above the rank of species it may well be considered as designating a series ... .” Thus, I have attributed this formal taxon primarily to Macbride & Payson, not as a combination, in exception to Holmgren’s view. Rydberg in 1906 clearly had a view of taxonomy in Erigeron that encom- passed more than only the Colorado species, because the names of several species groups (e.g., Acres, Radicati, Decumbentes) were based on epithets of taxa occurring in the Great Plains but not in Colorado. In Rydberg’s broader treatment of 1918, each of these three groups included the species upon which its name was based. I have accepted these species as the types of their nomenclaturally corresponding sections, since I believe that clearly was Rydberg’s intention. The following text presents sections of Erigeron, generic segregates, and groups originally in Erigeron but transferred into other genera. Unless specif- ically stated to the contrary, all infrageneric names cited are in Erigeron. Species marked by an asterisk (*) are ones that occur strictly in México and/or Central America. Names of accepted sections as well as lists of species 70 PHY TOELOCIA volume 67(1):67-93 July 1989 are presented alphabetically. Comments on putative relationships are in the text. I follow Cronquist (1947) and Hultén (1968) in conforming to the use of masculine endings in Erigeron, although Linnaeus regarded the genus as neuter. The International Code of Botanical Nomenclature (1988) notes that Erigeron is a masculine name “for which botanical usage has re-established the classical gender despite another choice by Linnaeus.” I. SECTIONS OF ERIGERON Erigeron sect. Arenarioides (Rydb.) Nesom, comb. et stat. nov. Based on Erigeron(sp.-group) Arenarioides Rydb., Fl. Rocky Mts. 897. 1918, in clave. TYPE: E. arenarioides (D. Eaton ex A. Gray) Rydb. Additional species in sect. Arenarioides: E. salmonensis Brunsfeld & Ne- som, ined. (Brunsfeld & Nesom [in press]). These two species clearly are closely related between themselves, and I can find no other group whose unity would not be unsettled by their inclusion. Cronquist (1947) has suggested that Erigeron arenartoides may be related to E. oryphyllus (see comments under sect. Ozyphyllum). Erigeron nemato- phyllus shares features of habit with these two species, but it has prominently coiling ligules and J include it in sect. Wyomtngza. Sect. Arenarioides is characterized by the following: tap-rooted perenni- als; caudex branches with densely packed, adherent leaf bases, each branch producing erect, wiry, distally branched stems; leaves ascending-appressed, entire; buds erect; heads turbinate-campanulate; and ray flowers few, with ligules not reflexing or coiling. Sect. Asteroidea Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:308. 1841. LEC- TOTYPE (designated here): E. decumbens Nutt. Nuttall also included E. corymbosus Nutt., E. filifolius Nutt., E. foliosus Nutt., EB. nanus Nutt., E. ochroleucus Nutt., E. pedatus Nutt. (=E. compositus Pursh) and E. radicatus Hook. Erigeron sect. Pseuderigeron Torrey & A. Gray, Fl. N. Amer. 2(1):177. 1841. LECTOTYPE (designated here): EF. caespitosus Nutt. Torrey & Gray also included £. filifolius Nutt., E. douglas Torrey & A. Gray (=E. foliosus Nutt.), E. decumbens Nutt., E. corymbosus Nutt. and BE. ochroleucus Nutt. Erigeron sect. Stenactis Torrey & A. Gray, Fl. N. Amer. 2(1):172. 1841. LECTOTYPE (designated here): E. pumilus Nutt. Other species in- cluded by Torrey & Gray: FE. glaucus Ker Gawler, FE. speciosus (Lind- ley) DC., E. glabellus Nutt., BE. concinnus (Hook. & Arn.) Torrey & A. Gray. See discussion in Nesom (in press). Nesom: Infrageneric treatment of Erigeron (i Asterigeron Rydb., Fl. Rocky Mts. 891. 1918. TYPE: A. watsoni (A. Gray) Rydb. [= Erigeron watsonii (A. Gray) Cronq.]. Erigeron (sp.-group) Caespitosi Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. caespitosus Nutt. Erigeron (sp.-group) Decumbentes Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. decumbens Nutt. Erigeron(sp.-group) Pumili Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. pumilus Nutt. Erigeron (sp.-group) Radicati Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. radicatus Hook. Erigeron(sp.-group) Asperuginei Rydb., Fl. Rocky Mts. 897. 1918, in clave. TYPE: E. asperugineus (D.C. Eaton) A. Gray. Erigeron (sp.-group) Laetevirentes Rydb., Fl. Rocky Mts. 897. 1918, in clave. TYPE: E. laetevirens Rydb. (=E. ochroleucus Nutt.). Erigeron (sp.-group) Filifolii Rydb., Fl. Rocky Mts. 897. 1918, in clave. TYPE: E. filifolius (Hook.) Nutt. The plants of sect. Asteroidea are perennials from thick taproots (except in the E. ursinus group) producing mostly scapose, monocephalous stems. Also, the leaves are entire; the buds are erect; the disc corollas are narrowly tubular; the ligules of the ray corollas often dry dark blue and have a ten- dency (strong in some, weak or not at all evident in others) to coil at the tips, but others appear to reflex (see comments below); and the achenes tend to be narrowly oblong. The plants typically grow in open, level or sloping, some- times rocky sites at medium elevations, often with sagebrush. Both Torrey & Gray and Nuttall saw the reality of this section, because sect. Pseuderigeron and sect. Asteroideaas originally described each included only a single species that I believe is part of a different lineage. Cronquist’s “Group X” of his sect. Erigeron (1947) also included mostly species of sect. Asteroidea. I have divided sect. Asteroidea into several groups and listed the species accordingly. 1. The BE. decumbens group: E. asperugineus (D. Eaton) A. Gray, E. canaani Welsh (=F. eatonii sensu Strother & Ferlatte, 1988), E. decumbens Nutt., B. eatoni A. Gray, E. jonesit Cronq., E. lassenianus E. Greene (incl. E. fleruosus Cronq.), E. nevadincola S.F. Blake, E. sonnei E. Greene, E. wahwahensis Welsh (=E. jonesii sensu Strother & Ferlatte, 1988) and E. watsonit (A. Gray) Cronq. This group is characterized by simple caudices (multicipital caudices occur but uncommonly), strongly decumbent stem bases that are often purplish, and a thickened node of wood and old petiole bases at the root-stem junction. 72 PHY TOWL-O.G LA volume 67(1):67-93 July 1989 The leaves of all species except F. watsonii typically or at least frequently have 3-nerved leaves. Strother & Ferlatte (1988) revised the taxonomy of this group, which they referred to as “Erigeron eatonii and allied taxa” According to them, all of the taxa of this group “... seem to be very closely related; they variously in- tergrade morphologically and may constitute a single, polymorphic species.” I believe two additional species belong with this species group: Erigeron as- perugineous, which previously has been thought to be more closely related to E. clokeyi (Cronquist, 1947; Ake, 1984) and the diminutive FE. watsonit, although it is apparently set apart from the rest. 2. The E£. ursinus group: E. gracilis Rydb. and E. ursinus D. Eaton. These two species are a very closely related pair distinct from the other species of sect. Asteroidea in their somewhat diffuse system of rhizomes. In their purple, decumbent stem bases, they appear to be most similar to the E. decumbens group. 3. The E. caespitosus group: E. abajoensis Crong., E. awapensis Welsh, E. caespitosus Nutt., HE. maguirei Cronq., FE. nauseosus (M.E. Jones) A. Nels., E. ovinus Cronq., E. subglaber Cronq., E. vetensis Rydb. and E£. zothecinus Welsh. This group is characterized by multicipital caudices and relatively broad, often 3-nerved basal leaves. There is some overlap in morphology with the £. decumbens group. Further, the ray corollas of E. vetensis and E. nauseosus have ligules that appear to reflex, setting these two species apart from the others and perhaps linking them with the FE. pumilus group. :In placing LE. zothecinus here, I follow Welsh (1986) who compared it to EH. abajoensis in the diagnosis. 4. The E. corymbosus group: E. corymbosus Nutt., E. filifolus Nutt. and E. ochroleucus Nutt. (incl. E. lackschewttzii Nesom & Weber). These three species have narrowly lanceolate or oblanceolate leaves and the ligules of the ray flowers are typically long and prominently coiling at the tips. Erigeron filifolius and E. corymbosus typically have branched stems; Erigeron ochroleucus is variable in this respect and in other features. 5. The E. radicatus group: E. disparipilus Cronq., E. latus (A. Nelson & Macbr.) Cronq., EF. nanus Nutt., E. parryi Canby & Rose, E. poliospermus A. Gray, E. pygmaeus (A. Gray) E. Greene, 2. radicatus Hook. and £. rydbergit Cronq. Nesom: Infrageneric treatment of Erigeron 73 The Erigeron radicatus group is characterized by multicipital caudices and strictly erect, relatively short, linear to narrowly oblong or narrowly oblanceolate, 1-nerved, basal leaves. 6. The E. pumilus group: FE. aphanactis (A. Gray) E. Greene, FE. clokeyi Cronq., E. concinnus (Hook. & Arn.) Torrey & A. Gray, E. engelmannii A. Nels. and FE. goodrichit Welsh. The species of the Erigeron pumilus group are similar in habit and leaf shape to the EF. radicatus group or perhaps somewhat intermediate to those of the FE. radicatus and E. caespitosus groups. The FE. pumilus group tends to be strikingly hispid in appearance, and most significantly, the ligules of the ray corollas appear to reflex. I have not seen EF. goodrichi but tentatively include it here on the basis of the discussion by Welsh (1983), who noted that its affinities are with FE. clokeyt and E. asperugineous. Groups 4-6 of sect. Asteroidea are much in need of a comprehensive tax- onomic study that includes field observations. The apparent occurrence of two distinctive ligule behaviors suggests that two lineages may be involved and that the groups as I have arranged them may be somewhat artificial. *Erigeron coronarius E. Greene, *E. janivultus Nesom and three unde- scribed species perhaps belong here, near the EF. pumilus group. I am cur- rently studying this group and defer making a formal judgment on its taxo- nomic placement until the completion of the study. Erigeron sect. Cincinnactis Nesom, sect. nov. TYPE: * Erigeron longipes DC. Folia marginibus dentatis, gemmae plus minusve erectae, corol- lae radii numerosae in 1-3 seriebus cincinnatae ad maturitatem plerumque perangustae, achenia brevi-oblongae parvulae costis manifeste aurantiacis, pappus plerumque sine serie externa. Additional species in sect. Cincinnactis: *E. basilobatus S.F. Blake, *E. crenatus Eastw., *E. ezilis A. Gray, *E. narcissus Nesom, *E. oazacanus Greenman, E. procumbens (Houston ex P. Miller) Nesom (=£. myrionac- tis Small), *£. socorrensis ILM. Johnston, *E. tephropodus Nesom, *E. ver- acruzensis Nesom and * EF. stanfordit 1.M. Johnston ex Nesom. The peculiar habit of Erigeron ezilis is ‘similar to that of species of sect. Linearifolw, but the behavior of its ligules allies it with sect. Cincinnactts. In an earlier publication (Nesom & Sundberg 1985), I included Erigeron longipes as a synonym of Erigeron scaberrimus (Less.) Nesom. The two, however, are distinct species. Further, the name FE. scaberrimus had already been proposed for a South American species, making my combination a later homonym. A new name is provided here for the Mexican species, which occurs from Tamaulipas, Veracruz, Puebla and Oaxaca. 74 PHY TOGO GiA volume 67(1):67-93 July 1989 Erigeron veracruzensis Nesom, nom. noy. Based on Aster scaberrimus Less., Linnaea 5:143. 1830. E. scaberrimus (Less.) Nesom, 1985; non E. scaberrimus Gardner, 1848. A group of primarily Caribbean species with tightly coiling ligules appar- ently also belongs in Erigeron sect. Cincinnactis. Two of them, E. bellioides DC. and E£. cuneifolius DC., occur in México, where they probably are ad- ventive. The plants of sect. Cincinnactis are characterized by the following: leaves with toothed margins, buds nodding; rays usually filiform and numerous (up to 300) in 1-3 series, tightly coiling at maturity, or only at the tips in species with longer ligules, usually very narrow; achenes short-oblong and very small, with prominent orange ribs; pappus usually without an outer series. All but one species (FZ. narcissus) are perennials from short, fibrous-rooted rhizomes. All are from México and Central America although E. procumbens ranges northward into the southeastern United States. Sect. Erigeron L., Sp. Pl. 863. 1753. LECTOTYPE (see discussion below): E. uniflorus L. Linnaeus included the following species in Erigeron: E. uniflorus L. E. acris L. E. alpinus L. E. philadelphicus L. E. carolinianus L. (see Fernald 1944, and comments below, following “GENUS UNKNOWN.” E. canadensis L. [=Conyza canadensis (L.) Cronq.] E. bonariensis L. [=Conyza bonariensis (L.) Cronq.] E. gramineus L. [=Arctogeron gramineus (L.) DC.] E. siculus L. [=Pulicaria sicula (L.) Moris] E. camphoratus L. [=Pluchea camphorata (L.) DC.] E. tuberosus L. [=Jasonia tuberosa (L.) DC.] E. viscosus L. [=Inula viscosa (L.) Aiton] Erigeron sect. Uniflori G. Don in Loudon, Hort. Brit. 343. 1830. TYPE: E. untflorus L. Don also listed E. alpinus L., E. composttus Pursh and E. gramineus L. [=Arctogeron gramineus (L.) DC.]. Aster sect. Pauciflori G. Don in Loudon, Hort. Brit. 346. 1830. LEC- TOTYPE (Sundberg & Jones 1987): A. pulchellus Willd. (=Erigeron venustus Botsch.). Nesom: Infrageneric treatment of Erigeron 75 Erigeron (sp.-group) Uniflori Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. uniflorus L. Erigeron sect. Monocephali Vieth., Beih. Bot. Centralbl. 19:492. 1906. LECTOTYPE (designated here): FE. uniflorus L. Erigeron sect. Siphonoglossa Botsch., Bot. Mater. Gerb. Bot. Inst. Ko- marova Akad. Nauk SSSR 16:393. 1954. TYPE: FE. unzflorus L. Erigeron sect. Platyglossa Botsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk SSSR 16:388. 1954. TYPE: Erigeron venustus Botsch. Sect. Platyglassa is tentatively included as a synonym of sect. Erigeron although Botschantzev included in it several species that are members of sect. Fruticosus |e.g., E. peregrinus (Banks ex Pursh) E. Greene, E. thunbergii A. Gray]. Erigeron venustus does not appear to be closely related to these species (Nesom 1982) but rather to be more similar to those of sect. Erigeron. - Additional North American species of sect. Erigeron: E. algidus Jepson (=£. petiolaris E. Greene, non Vierh.), E. alpiniformis Crongq., E. arthuru B. Boivin, EF. aureus E. Greene, E. ertocephalus Vahl, FE. evermannit Rydb., E. flettiti G.N. Jones, FE. grandiflorus Hook., E. humilis Graham, E. hultent Spongberg, FE. hyperboreus E. Greene, FE. lanatus Hook., EF. melanocephalus A. Nelson, E. meziae K. Becker, E. mutrit A. Gray, E. stmplez E. Greene, E. yukonensts Rydb. This view of sect. Erigeron restricts it essentially to monocephalous plants of alpine and subalpine habitats, at least in North America. In addition, these plants are perennials from short, fibrous-rooted rhizomes and have entire leaves, erect buds and coiling ligules. Spongberg (1969), who included most of these species in a biosystematic study, found evidence for reticulate inter- relationships among a number of them. He observed that “in North America, few species at lower altitudes in the Cordillera appear to be truly implicated with arctic-alpine species of Erigeron” and that their nearest relatives are “likely to be found in the mountains of Central Asia and the Caucasus.” Species with a monocephalous habit are found in every section of Erigeron except sect. Arenarioides (two species), sect. Ozyphyllum (two species) and sect. Phalacroloma (two species, perhaps belonging with sect. Olygotrichium). In sects. Fruticosus, Karvinskia, Leptostelma, Olygotrichium and Polyactis, it seems relatively clear that branched capitulescences are primitive, but in the other groups, including sect. Erigeron, it seems likely that monocephaly is primitive and that a branching habit has been derived from ancestors with simple, monocephalous stems. In searching for an area of common ancestry between Erigeron and Aster, Cronquist (1947) emphasized the branching, leafy, Aster-like habit of species of Erigeron sect. Fruticosus. There are boreal, monocephalous species of Aster, however, that also appear to straddle a different, somewhat arbitrary 76 PHY TOMOGt A volume 67(1):67-93 July 1989 morphological boundary between Aster and Erigeron, e.g., A. alpinus L. and A. alpigenus (Torrey & A. Gray) A. Gray. Two lectotypes have been selected for Erigeron by previous botanists. E. acris L. [Britton N.L. & A. Brown, Illus. Fl. N. U.S. (ed. 2) 3:436. 1913]. E. uniflorus L. (Green, M.L., Prop. Brit. Bot. 181. 1929). l accept Erigeron uniflorus as the lectotype and reject the earlier choice for several reasons. Most significantly, the choice of H. acris as the type of Trimorpha and later of Erigeron sect. Trimorpha were made before its se- lection by Britton & Brown as the lectotype of Erigeron (sect. Erigeron). Further, the relatively few (ca 40-45) species of sect. Trimorpha are very distinct in their trimorphic florets from the remainder of the genus. Trz- morpha has been treated as a genus (Vierhapper 1906) and as a subgenus (Botschantzev 1959), and I have resegregated it as a genus (Nesom 1989b). In this case, if EF. acris were the type of Erigeron, more than 300 species would have to be renamed in a genus now understood to be one of the var- ious synonyms of Erigeron. Finally, the choice of the generic name (“early old age”) by Linnaeus probably was intended to reflect the appearance of the densely woolly-villous involucres of some of the species. Erigeron uniflorus shows this feature, &. acris does not. Cronquist (pers. comm.) has added his agreement with this choice of a lectotype with the following comment: “Pennell pointed out to me many years ago that Linnaeus often based his descriptions for Genera Plantarum on a single species, which if it can be identified becomes the logical lectotype. Linnaeus’ description in Genera Plantarum is scarcely compatible with E. acris, although it would fit &. uniflorus and some other species.” Sect. Fruticosus G. Don in Loudon, Hort. Brit. 343. 1830. TYPE: “E. glaucus B.R.” (=E. glaucus Ker Gawler, Bot. Reg., 1815): Monotypic as recognized by Don. Erigeron sect. Pauciflort G. Don in Loudon, Hort. Brit. 343. 1830. LEC- TOTYPE (designated here): JE. bellidifolius Willd. (=E. pulchellus Michx.). Don also included Erigeron caucasicas Stev., E. nudicaulis Michx. (=£.vernus (L.) Torrey & A. Gray), E. glabellus Nutt., E. ja- maicensis L. and E. montevidensis Spreng. {according to Solbrig (1962), this plant probably represents a species of Conyza]. Musteron Rafin., Fl. Tellur. 2:50. 1836. TYPE: “M. bellidifolium” Rafin. Probably =E£. bellidifolius Willd. (=E. pulchellus Michx.). Monotypic as recognized by Rafinesque. Fragmosa Rafin., Fl. Tellur. 2:50. 1836. LECTOTYPE (designated here): E. nudicaults Michx. [=E. vernus (L.) Torrey & Gray]. Also included in Fragmosa by Rafinesque were FE. uniflorus L., E. alpinus L., E. pumilus Nutt., EZ. asper Nutt. (=E. glabellus Nutt.) and FE. glabellus Nutt. Nesom: Infrageneric treatment of Erigeron 77 Woodvillea DC., Prodr. 5:318. 1836. TYPE: W. calendulacea DC. (=E. glaucus Ker Gawler). Monotypic as recognized by de Candolle. Erigeron sect. Phoenactis Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:310. 1840. LECTOTYPE (designated here): E. speciosus (Lindl.) DC. Other species included by Nuttall: FE. macranthus, FE. hispidum Nutt. (=£. glaucus Ker Gawler), E. maritimum Nutt. (=E£. glaucus Ker Gawler). Erigeron sect. Erigeridium Torrey & A. Gray, Fl. N. Amer. 2(1):176. 1841. TYPE: E. vernus (L.) Torrey & A. Gray. Monotypic as recognized by Torrey & Gray (see comments below). Erigeron (sp.-group) Elatiores Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. elatior (A. Gray) E. Greene. Erigeron (sp.-group) Salsuginost Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. salsuginosus (Rich.) A. Gray [=E. peregrinus subsp. cal- lianthemus (E. Greene) Cronq.]. See Cronquist (1947), however, for comments on the misapplication of the name Aster salsuginosus Rich. Erigeron (sp.-group) Macranthi Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. macranthus Nutt. [=E. speciosus (Lindl.) DC.]. Erigeron (sp.-group) Glabelli Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. glabellus Nutt. Erigeron (sp.-group) Asperi Rydb., Fl. Rocky Mts. 897. 1918, in clave. TYPE: E. asper Nutt. (=E. glabellus Nutt.). Erigeron (sp.-group) Verni Small, Man. Southeastern Fl. 1395. 1933, in clave. TYPE: E. vernus (L.) Torrey & A. Gray Erigeron (sp.-group) Pulchelli Small, Man. Southeastern Fl. 1395. 1933, in clave. TYPE: £. pulchellus Michx. Erigeron sect. Peregrinus Nesom, Syst. Bot. 7:463. 1982. TYPE: LE. pere- grinus (Banks ex Pursh) E. Greene. In an earlier paper (Nesom 1982), I included 34 species in this section but now believe that eight (those lacking a pappus of bristles) should be segregated as a distinct group (see sect. Imbarba below). Species of sect. Fruticosus: E. aliceae J. Howell, E. arizonicus A. Gray, E. cascadensis A. Heller, E. cervinus E. Greene, EF. coulterz Porter, E. elatior (A. Gray) E. Greene, EF. ezimius E. Greene, E. formosissimus E. Greene, E. garrettu A. Nelson, E. glabellus Nutt., FE. glaucus Ker Gawler, E. hessit Nesom, EF. howellit A. Gray, E. kuschei Eastw., FE. letbergit Piper, E. oreganus A. Gray, E. peregrinus (Pursh) E. Greene, E. platyphyllus E. Greene, *£. potosinus Standley, E. pulchellus Michx., FE. rusbyi A. Gray, E. rybius Nesom, E. sanctarum S. Watson, FE. speciosus (Lindley) DC., E. subtrinervis Rydb., 78 PMRYTOLIOGLEA volume 67(1):67-93 July 1989 E. supplez A. Gray, E. thunbergit A. Gray “complex,” E. uintahensis Cronq. and the E. palmeri group (see below). Sect. Fruticosus is characterized by the following: perennials mostly from short, thick, fibrous-rooted rhizomes; leaves often 3-nerved, entire or less commonly slightly toothed, the cauline more or less equably distributed, little reduced from the basal, usually at least the upper or lower clasping; buds erect or on slightly curved peduncles; phyllaries in 2-4 equal or nearly equal series, herbaceous, narrowly lanceolate with attenuate, often flexuous tips; ray corollas usually long and coiling at the tips. See Nesom (1982) for comments on problems in circumscribing this group and a discussion on a putative relationship between sect. Erigeron and sect. Fruticosus. The E. palmert group. Four pappose species restricted to México, closely inter-related among themselves, appear to be members of sect. Fruticosus: *E. hintoniorum Ne- som, *E. morelensis Greenm., *E. palmeri A. Gray and *£. wellsii Nesom. Erigeron vernus (L.) Torrey & A. Gray of the southeastern United States, also belongs to the E. palmeri group. This species was the basis for the monotypic sect. Erigeridium of Torrey & Gray, but Cronquist (1947) included it as a member of sect. Olygotrichtum. It has smaller heads on more branched stems than its closest relatives, but the thick, dull green and nearly glabrous basal leaves with remotely and shallowly toothed margins are nearly identical to those of E. palmeri. Further, both species have thick, fibrous roots with no rhizome. The long disc style appendages of FE. vernus, the erect buds and the rays not closing upwards at night are also distinctive. The position of Erigeron pulchellus. The only species of sect. Fruticosus besides Erigeron vernus in the east- ern United States is E. pulchellus. Earlier (Nesom 1982), I was equivocal about its taxonomic placement. It is similar to EF. philadelphicus (sect. Oly- gotrichium) in aspects of its capitular morphology and its numerous, thin- herbaceous, clasping, cauline leaves. Clasping leaves, however, are also typi- cal of sect. Fruticosus, and I believe Cronquist (1947) was correct in noting the distinctness of the scale-leaved, stoloniform rhizomes of FE. pulchellus. Similar rhizomes are not produced in sect. Olygotrichium, but they are found in several species of sect. Fruticosus (e.g., E. rybius, E. ezimius, E. potosi- nus). Further, E. pulchellus has erect buds and large heads with rays that do not close upwards at night, and the ligules of some collections can be seen to be distinctly coiling, all traits typical of sect. Fruticosus but found in no species of sect. Olygotrichium. Erigeron sect. Imbarba Nesom, sect. nov. TYPE: * Erigeron galeottit (A. Gray ex Hemsley) E. Greene. Nesom: Infrageneric treatment of Erigeron 79 Gemmae erectae, phyllaria 2-porcata ad basim, corollae radii cincinnatae ad maturitatem, setae pappo carentes vel paucae dis- persim praesentes. Additional species in sect. Imbarba: * E. astranthioides De Jong & Nesom, *F. forreri(E. Greene) E. Greene, *£. fraternus E. Greene, * FE. guatemalensis (S.F. Blake) Nesom, *£. mimus (S.F. Blake) Nesom and *E£. strigulosus E. Greene. I first included these eight species (Nesom 1982) as members of sect. Peregrinus (=sect. Fruticosus) but now believe there is as much evidence to suggest they represent a sister taxon of the latter as a derivative group of it (Nesom in prep.). These species are distinguished by their toothed leaves, erect buds, basally 2-ridged phyllaries, distally coiling ligules of the ray flowers and essential lack of pappus bristles. Except for EF. strigulosus, a tap- rooted annual, they are all perennials from short, fibrous-rooted rhizomes. - All occur in México and Guatemala. Erigeron sect. Karvinskia Nesom, sect. nov. TYPE: *E. karvinskianus DC. Plantae perennia, folia basalia decidua ab florescentia, fo- lia caulinae obovatae dentatae plus minusve pariter distributae, gemmae erectae, corollae radii nec cincinnatae nec reflexae non clausae sursus nocte. Additional species of sect. Karvinskia: *E. heteromorphus B. Robinson, *F. irazuensis Greenman and * £. pacayensis Greenman. Plants of this section are characterized by the following features: peren- nials from thin, woody taproots or short rhizomes, leaves mostly obovate, toothed, all cauline by flowering and distributed more or less equably on the stems, buds erect, ligules of the ray flowers neither coiling nor reflexing and not closing upwards at night. In addition to those listed above, there are two additional but undescribed species of sect. Karvinskia from México and Central America. The group is presently under study by me. Sect. Karvin- skia may prove to be closely related to sect. Lineartfolii, but the broader, toothed leaves and equal phyllaries of the former and the habit of the latter are distinctive. Erigeron sect. Linearifolii (G. Don) Nesom, comb. nov. Based on Aster sect. Linearifolit G. Don in Loudon, Hort. Bot. 346. 1830. LECTO- TYPE (Sundberg & Jones 1987): A. graminifolius Pursh (=E. hyssopi- folius Michx.). Erigeron sect. Pycnophyllum Cronq., Brittonia 6:141. 1947. TYPE: E. foliosus Nutt. 80 PHY TOLOGTIA volume 67(1):67-93 July 1989 Sundberg & Jones (1987) lectotypified Don’s sectional epithet to preserve an established one in Aster; by doing so, however, they moved Don’s name into a position of priority over Cronquist’s sect. Pycnophyllum. Species of sect. Linearifolii: E. aequifolius H.M. Hall, FE. brewerz A. Gray, *B. chiangii Nesom, E. foliosus Nutt., E. hyssopifolius Michx., FE. tnornatus A. Gray, *E. lepidopodus (B. Robinson & Fern.) Nesom, EF. miser A. Gray, E. petrophilus E. Greene, E. rhizomatus Cronq. and *E. scoparioides Nesom. In earlier publications, I did not clearly recognize the relationships of the three Mexican species of this section. Hrigeron lepidopodus from Chi- huahua and northern Durango, México, however, is clearly the sister species of Erigeron rhizomatus, endemic to Catron and McKinley counties, New Mex- ico. Erigeron chiangii and E. scoparioides, species of northeastern Mexico, are probably sister taxa. The species of sect. Linearifolit are characterized by the presence of nu- merous, linear or narrowly oblong, entire leaves, borne on short internodes and essentially uniform from base to near top of the plant. They are peren- nial, often with long, slender, caudex-like, lower branches. The buds are erect, the phyllaries are usually strongly graduated (though not in the type species) and the rays have ligules straight or slightly coiling at the tips. Sect. Olygotrichium Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:311. 1840. LECTOTYPE (designated here): E. divaricatum Nutt. (=E. divergens Torrey & A. Gray). Nuttall also included E. tenuis Torrey & Gray, E. strigosus Muhl. ex Willd. and E£. “occidentalis” (?=E. strigosus Muhl. | ex Willd.). Heterochaeta DC., Prodr. 5:282. 1836; non Besser ex Schultes & Schultes, 1827. LECTOTYPE (designated here): Erigeron pubescens Kunth. De Candolle also included E. gnaphalioides Kunth (=Conyza confusa Cronq.) and four extra-American species. Erigeron (sp.-group) Philadelphici Rydb., Fl. Colorado 359. 1906, in clave; non Small, 1933. TYPE: EF. philadelphicus L. Erigeron (sp.-group) Bellidiastra Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. bellidiastrum Nutt. Erigeron (sp.-group) Divergentes Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. divergens Torrey & A. Gray. Erigeron (sp.-group) Flagellares Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. flagellaris A. Gray Species of sect. Olygotrichium: E. bellidiastrum Nutt., *E. bigelovii A. Gray, *E. calcicola Greenman, FE. colomezicanus A. Nelson, E. divergens Torrey & A. Gray (incl. E. solisaltator Nesom), *E. dryophyllus A. Gray, E. flagellaris A. Gray, *E. fundus Nesom, F. geisert Shinners, EF. gilensis Nesom: Infrageneric treatment of Erigeron 81 Woot. & Standl., *E. gypsoverus Nesom, E. lemmonii A. Gray, EF. lobatus A. Nelson, *E. metrius S.F. Blake, *£. mihianus S.F. Blake, E. mimegletes Shinners, E. modestus A. Gray, FE. multiceps E. Greene, *E. onofrensis Ne- som, E. philadelphicus L., *E. pinkavi B. Turner, EF. proselyticus Nesom, *E. pubescens Kunth, EF. quercifolius Lam., E. religiosus Crong., E. stonis Cronq., E. tenellus DC., E. tenuis Torrey & A. Gray, *F. turnerorum Ne- som, *E. unguiphyllus Nesom, * EF. velutipes Hook. & Arn. and *£. versicolor (Greenman) Nesom. Erigeron sect. Olygotrichium is a relatively large but well defined sec- tion. The species are primarily from the southwestern United States and Mexico, with a small group from the eastern United States and the Antilles. They are mostly taprooted annuals or biennials, and though perennials are also included, no other section of Erigeron has so many species of plants of annual-biennial duration. They have prominently lobed or toothed leaves, nodding buds, small heads and narrow ligules often with a lilac midstripe on the lower surface. The ligules close upwards at night, neither reflexing nor coiling with maturity. This ligule behavior apparently is unique among the sections of Erigeron, but it is known from other genera of Astereae (e.g., Aphanostephus and at least some species of Townsendia). Four species, E. mimegletes, E. versicolor, E. gilensis and an undescribed species from Chi- huahua, are epappose. A group of inter-related species from the Antilles also appears to belong in sect. Olygotrichium: E. caeruleus Urban, £. dissectus Urban, FE. jamaicensis L. and E. pinetorum Urban. Erigeron psilocaulis Urban differs from these in its non-clasping leaves but probably is part of the same lineage, as is the suffrutescent FE. darrellianus Hemsley. Relationships among the species of sect. Olygotrichium of the eastern and south-central United States are complex. Erigeron philadelphicus, E. quercifolius and E. caeruleus (and its relatives) appear to be closely related. Erigeron tenuis, E. tenellus , E. geiseriand E. turnerorum (northern México) form a closely knit group. Erigeron tenuis, in turn, is very similar to E. strigosus of sect. Phalacroloma. Further comments on the possible relation- ship between sect. Olygotrichium and sect. Phalacroloma are found under the latter. As pointed out by Cronquist (1947), Erigeron glabellus (sect. Fruticosus) is in some ways (particularly its short-lived duration and numerous ray flow- ers with narrow ligules) similar to species of sect. Olygotrichium, and this species may indicate a possible direction of common ancestry for the two sections. In Cronquist’s view, sect. Olygotrichtum comprised 20 species. One of these, E. calvus Coville, I consider to be a synonym of £. divergens and another, FE. plateauensis Cronq., a synonym of E. modestus. Although Cron- 82 PHY TOLO GIA volume 67(1):67-93 July 1989 quist placed FE. multiceps E. Greene in sect. Erigeron because of its perennial duration, it is, as he noted, most closely related to EF. divergens and be- longs in sect. Olygotrichium. Several species cited by Cronquist as members of sect. Olygotrichium are better placed in different groups: EF. myrionactis Small (=E. procumbens, sect. Cincinnactis), E. neomezicanus A. Gray and E. oreophilus Greenman, both of which were included by Cronquist in FE. del- phinifolius (sect. Polyactis), E. glabellus (sect. Fruticosus), E. vernus (sect. Fruticosus) and E. pulchellus (sect. Fruticosus). Comments on the last three species are found under sect. Fruticosus. Erigeron sect. Osteocaulis Nesom, sect. nov. TYPE: E. linearis (Hook.) Piper Bases caulium et foliorum plerumque albido-induratae osseae, gemmae erectae, folia lineares vel oblanceolatae, corollae radii luteae vel caeruleae nec cincinnatae nec reflexae non clausae sur- sus nocte in specie una abscentes differt. Erigeron (sp.-group) Lutet Rydb., Fl. Rocky Mts. 897. 1918, in clave. TYPE: E. luteus A. Nelson [=E. linearis (Hook.) Piper]. Additional species in section Osteocaulis: E. barbellatus E. Greene (rays blue/white), E. bloomeri A. Gray (rayless), E. chrysopsidis A. Gray, E. ele- gantulus E. Greene (blue/pink rayed) and EF. piperianus Cronq. Sect. Osteocaulis is a well defined natural group whose essential composi- tion was recognized by Cronquist in 1947. It is characterized by the following: perennials from caudices with several, short, erect branches; bases of stems and petioles usually whitish-indurated, bony-textured; buds erect; leaves en- tire, linear, varying to narrowly oblanceolate in FE. barbellatus, stiffly erect; ligules of ray corollas not reflexing or coiling, yellow in three species, bluish in three, one species rayless. The only other yellow-rayed species of Erigeron in North America is &. aureus, which is a member of sect. Erigeron. Several species of Erigeron sect. Asteroideaalso have whitish petiole bases (e.g., E. nanus, E. rydbergii) but the margins are ciliate and the cellular texture is different from that in sect. Osteocaulis. Cronquist (1947) did not make any comment regarding the putative re- lationships of Erigeron filifolius, but he did include it as a side branch among the species of sect. Osteocaulis in his “probable relationships” dia- gram. Erigeron filifolius, however, lacks both the yellow rays and strikingly indurated stem/petiole bases of the taxa that make section Osteocaulis un- usual, and [ have placed it in sect. Asteroidea. Sect. Phalacroloma (Cass. in Cuvier) Torrey & A. Gray, Fl. N. Amer. 2(1):175. 1841. Based on Phalacroloma Cass. in Cuvier, Dict. Sci. Nat. 39:404. 1826. TYPE: P. obtusifolia Cass. (=E. strigosus Muhl. Nesom: Infrageneric treatment of Erigeron 83 ex Willd.). Cassini also included P. acutifolia Cass. [=E. annuus (L.) Pers.] in the genus. Diplemium Rafin., Fl. Tellur. 2:50. 1836. LECTOTYPE (designated here): Erigeron strigosus Muhl. ex Willd. Rafinesque also included Erigeron nervosus Willd. [=Heterotheca graminifolia (Michx.) Elliott], Erigeron quercifolius Lam. and Erigeron carolinianus L. Erigeron (sp.-group) Ramosit Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. ramosus (Walter) B.S.P. (=£. strigosus). Erigeron (sp.-group) Annut Small, Man. Southeastern Fl. 1395. 1933, in clave. TYPE: E. annuus (L.) Pers. Cassini was not sure whether Phalacroloma obtusifolia might be the same species as Erigeron hyssopifolius Michx. or FE. carolinianus L. “This confusion commenced with Pursh, who erroneously adduced the figure of Dillenius and the E. Carolinianus as synonyms of the E. hyssopifolium of Michaux” (Gray 1841). This putative synonymy was repeated by Cassini, de Candolle and Lessing. Cassini’s description of the pappus and discussion of the characters of Phalacroloma leave little doubt, however, as Gray (1841) early realized, that the type was E. strigosus (see Fernald 1944, for further comments on the identity of Erigeron carolinianus). Torrey & Gray (1841) included as members of sect. Phalacroloma: E. strigosus, E. annuus, E. tenuis and £. divergens, the last two of sect. Oly- gotrichium. Cronquist (1947) included only the two species lacking pappus bristles on the ray achenes, Erigeron strigosus and FE. annuus, but, as he noted, EF. tenuis is very similar to E. strigosus and sect. Phalacroloma may be only artificially separated from sect. Olygotrichium. Both species of sect. Phalacroloma are annuals with slightly to prominently nodding buds. The leaves of EF. annuus are strongly toothed; those of EF. strigosus vary from entire to toothed. Chromosome counts of Erigeron annuus have all been triploid; diploid and various polyploid populations of FE. strigosus have been reported. I spec- ulate that E. annuus may be of hybrid origin between FE. strigosus and some other species of the eastern United States, E. philadelphicus being the most conspicuous candidate. Sect. Polyactis (Less.) Nesom, Phytologia 66:416. 1989. Based on Polyactis Less., Syn. gen. Comp. 188. 1832. TYPE: Erigeron delphinifolius Willd. Polyactidium DC., Prodr. 5:281. 1836. Stenactis Cass. in Cuvier, Dict. Sci. Nat. 37:485. 1825. TYPE: Erigeron delphinifoltus Willd. Non Stenactis sensu Nees, 1832; non sensu Less., 1832; non E. sect. Stenactts Torrey & A. Gray, 1841. Achaetogeron A. Gray, Mem. Amer. Acad. Arts, n. ser. 4 (Pl. Fendl.):72. 84 Pane YE OG OLGA. volume 67(1):67-93 July 1989 1849. TYPE: Achaetogeron wislizeni A. Gray [=Erigeron wislizeni (A. Gray) E. Greene]. Species of sect. Polyactis: *E. annuactis Nesom, *E. basaseachensis Ne- som, *E£. caulinifolius Nesom, *E. circulis Nesom, *£. coroniglandifer Nesom, *&. dactyloides (Greenm.) Nesom, * £. delphinifolius Willd., *E. eruptens Ne- som, *E. griseus (Greenm.) Nesom, *£. inoptatus A. Gray, *E. nacoriensis Nesom, FE. neomezicanus A. Gray, E. oreophilus Greenman, *E. podophyllus Nesom, *E. polycephalus (Larsen) Nesom, *E. rhizomactis Nesom, *E. see- mannii (Schultz-Bip.) E. Greene, *Z. subacaulis (McVaugh) Nesom and *£. wislizeni (A. Gray) E. Greene. These species are characterized for the most part by pinnatifid or coarsely toothed leaves, arching-pendant buds, ligules that sharply reflex at the tube- ligule junction and a pappus of bristles that are basally caducous or com- pletely lacking. All occur in western and south-central México; two of them, E. oreophilusand E. neomezicanus, also are found in the southwestern United States. See Nesom (1989a) for details and a discussion of typification of the generic synonyms noted here. Erigeron sect. Scopulincola Nesom, sect. nov. TYPE: E. scopulinus Ne- som & Roth Plantae parvulae rhizomatosae scopuli habitantes, folia oblanceolati- spathulata, corollae radii reflexae. Additional species in sect. Scopulincola: E. kachinensis Welsh & Moore, E. leiomerus A. Gray and E. pringlei A. Gray. These species from the southwestern United States appear to form a well defined natural group. The habitat of each is in cracks and crevices of rock faces and cliffs. All are rhizomatous perennials with entire, oblanceolate- spatulate leaves, erect buds and ray flowers with prominently reflexing ligules. In their leaves and habit, these plants are similar to those of sect. Spatht- folium, but the peculiar habitat, distinctive rhizomes (vs caudex branches) and reflexing ligules set them apart. Erigeron sect. Spathifolium Nesom, sect. nov. TYPE: EF. tener (A. Gray) A. Gray Plantae parvae monocephalae ramis brevibus caudicis, indu- mentum strigillosum trichomatum alborum arcte adpressorum brevium rigidorum acutorum, folia oblanceolati-spathulatae, gem- mae erectae, et corollae radii nec cincinnatae nec reflexae non clausae sursus nocte. Additional species in sect. Spathifolium: E. acomanus Spellenberg & Knight ined, &. cavernensis Welsh & Atwood, FE. cronquistit Maguire, E. tener (A. Gray) A. Gray, E. tweedyi Canby and E. uncialis S.F. Blake. Nesom: Infrageneric treatment of Erigeron 85 These six species comprise a relatively well defined natural group of mostly small perennials with long or short caudex branches, ultimately from a taproot, entire, oblanceolate-spatulate leaves, a strigillose indument of closely appressed, white, short, stiff and sharp-pointed hairs; erect buds and ligules that neither coil nor reflex. The stem pubescence of EF. uncialis is longer and spreading but otherwise similar. The stems of these plants are simple and monocephalous, except for those of EF. tweedy1, which usually are few branched. Earlier, Nesom & Roth (1981) tentatively placed Erigeron cronquisti closer to sect. Scopulincola. Ake (1984), however, as well as Maguire (1944) in its original description, correctly recognized the relationship of E. cronquisti with E. tener. Atwood & Welsh (1988) noted in their discussion that E. cavernensis is “evidently allied to E. simplez,” but they correctly contrasted it with E. uncialis, to which it is most similar, in the diagnosis. The habit and peculiar pubescence of plants of sect. Spathifolium are similar to those in sect. Wyomingia, and there is a possibility of a close relationship between the two groups. Erigeron sect. Spinosi (Alexander in Small) Nesom & Sundberg, comb. nov. Based on Aster (sp.-group) Spinos: Alexander in Small, Man. Southeastern Fl. 1365. 1933, in clave. Aster sect. Spinosi (Small) A.G. Jones, Brittonia 32:233. 1980. TYPE: Aster spinosus Benth. (=Erigeron ortegae S.F. Blake) Additional species in sect. Spinosi: E. ozyphyllus E. Greene. Sundberg (1986) concluded that A. spinosus is better placed in Erigeron than Aster, but he did not discover that it has such a close relative already placed in Erigeron. The similarity, however, between E. ozyphyllus and A. spinosus is striking. Plants of both produce stems that are 5-25 dm tall, strictly erect, green but somewhat ligneous, glabrous and striate. Further, both are perennial and usually produce many simple stems from a woody, thizomatous base. The entire leaves are produced early in the season and are quickly deciduous, usually leaving only the green stems by flowering. In both species, the buds are erect, the phyllaries strongly graduated in 4-5 series and the heads solitary or in loose corymbs. On the other hand, the achenes of E. oryphyllus are strigose and have 2-4 thick, orange, resin-filled ribs, while those of A. spinosus are glabrous, have 5, thinner, light-colored nerves and are not so strongly compressed. The phyllaries with three, orange-resinous veins emphasized by Sundberg as characteristic of A. spinosus are apparent on some plants of E. ozyphyllus (e.g., Peebles & Harrison 5270 (LL) but not on others. Plants of E. ozyphyllus are not spiny, but neither are some forms of A. spinosus. Cronquist (1947) described Erigeron ozyphyllus as a “peculiar and well- marked species ... taxonomically somewhat isolated” but probably related to 86 PHY TOO aA volume 67(1):67-93 July 1989 E. arenarioides. Plants of the latter have erect buds and strongly graduated phyllaries, but in habit they are much smaller with thinner stems and have persistent basal and cauline leaves, and I have placed them in a different section. The nomenclature and infraspecific taxonomy of Aster spinosus will be treated by Sundberg (in prep.). The addition of a third species, which is from México and yet unde- scribed, will somewhat alter the circumscription of sect. Spinos:. This will be discussed in the formal proposal of the species (Sundberg & Nesom [in prep.]). Besides Aster spinosus, Jones (1980) also included A. intricatus (A. Gray) Benth. (=Leucosyris E. Greene) in her Aster sect. Spinosi. See Nesom et al. (in press) for additional comments on the latter species, which is not part of the same group as A. spinosus. Sect. Tridactylia Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:310. 1841. TYPE: E. compositus Pursh. Monotypic as recognized by Nuttall. Erigeron series Multifidi Rydb. ex Macbride & Payson, Contr. Gray Herb. 49:73. 1917. Based on Erigeron (sp.-group) Multifidi Rydb., Fl. Col- orado 359. 1906, in clave. LECTOTYPE (designated here): E. com- positus Pursh. Erigeron (sp.-group) Compositi Rydb., Fl. Rocky Mts. 896. 1918, in clave (Rydberg changed the name of this group in 1918). Macbride & Payson also included FE. pinnatisectus (A. Gray) A. Nelson. Additional species in sect. Tridactylia: E. allocotus S.F. Blake, E. basalti- cus Hoover, E. flabellifolius Rydb., E. mancus Rydb., E. pallens Cronq., E. pinnatisectus (A. Gray) A. Nelson, E. purpuratus E. Greene, E. salishiu Douglas & Packer, E. trifidus Hook. and E. vagus Payson. Becker (1976) described EF. meziae Becker, which sometimes has apically tridentate leaves, as a “connecting link” between sect. Erigeron and sect. Tridactylia and it may be a member of the latter. Most of the species of sect. Tridactylia have pinnatifid leaves; in several, the leaves are apically trilobed. In all species the leaves are non clasping, the buds are erect and the ligules neither coil nor reflex. Also, they are perennials with a strong tendency to produce rhizome like caudex branches. Sect. Wyomingia (A. Nelson) Cronq., Brittonia 6:140. 1947. Based on Wyomingia A. Nelson, Bull. Torrey Bot. Club 26:249. 1899. TYPE: E. pulcherrimus Heller [= Wyomingia pulcherrima (Heller) A. Nelson]. Erigeron (sp.-group) Cani Rydb., Fl. Colorado 359. 1906, in clave. TYPE: FE. canus A. Gray Erigeron (sp.-group) Tetrapleuri Rydb., Fl. Rocky Mts. 897. 1918, in clave. TYPE: E. tetrapleuris (A. Gray) Heller (=F. utahensis A. Gray). Nesom: Infrageneric treatment of Erigeron 87 The species of sect. Wyomingia are taprooted perennials, and they have entire, more or less linear leaves, a strigillose vestiture of very short and closely appressed hairs, erect buds and prominently coiling ligules. I recognize two groups within the section and list the species accordingly. 1. The E. pulcherrimus group: FE. argentatus A. Gray, EF. canus A. Gray, E. parishtt A. Gray, FE. pulcherrimus and EF. utahensis A. Gray. These species have multi-nerved [(2-)4-10(-14)] achenes, a feature re- stricted mostly to this group. The achenes of Erigeronsect. Spinosi, however, also may have 4 to 5 ribs, as may those of E. peregrinus (sect. Fruticosus) and E. lepidopodus and E. rhizomatus (sect. Linearifolit). 2. The E. compactus group: E. carringtonae Welsh, FE. compactus S.F. Blake, EF. consimilis Cronq., E. nematophyllus Rydb. and E. untermanniz Welsh & Goodrich. Erigeron compactus, E. consimilis and E. nematophyllus were noted by Cronquist (1947) to be “apparently not far removed from the line of descent of the section Wyomingia.” They lack the multinerved achenes characteris- tic of the EF. pulcherrimus group and tend to be smaller in stature but are otherwise very similar. Some plants of EF. pulcherrimus, however, are small and nearly identical in habit to those of E. compactus. Erigeron nematophyl- lus is obviously more similar to FE. utahensis in its habit of relatively tall, branching stems, which suggests that the taxonomic use of achenial nerves may result in an artificial division, at least in this section. Welsh (1983) hy- pothesized that E. untermanniiand E. carringtonae are most closely related to E. compactus, and they are included here on that basis. Erigeron rhizomatus, which was included by Cronquist (1947) in sect. Wyomingia, is placed in sect. Lineartfolii in the present treatment with its sister species, FE. lepidopodus. SOUTH AMERICAN GROUPS - POSITION IN ERIGERON UNKNOWN Erigeron sect. Leptostelma (D. Don) Benth. & Hook., Gen. Pl. 2:280. 1873. Based on Leptostelma D. Don in Sweet. TYPE: Leptostelma mazimum D. Don [E. mazimus (D. Don) DC|]. Additional species of sect. Leptostelma (see Solbrig 1962): EB. tucumanen- sts Cabrera and FE. tweediet Hook. & Arn. Darwiniothamnus Harling, Acta Horti Berg. 20(3):108. 1962. TYPE: D. tenuifolius (Hook. f.) Harling (=Erigeron tenuifolius Hook. f.). 88 PU Ys ITROPLNOLGsICA volume 67(1):67-93 July 1989 The two species of Erigeron that occur on the Galapagos Islands (E.tenu- ifolius and E. lancifolius Hook. f.) were segregated as the genus Darwinio- thamnus by Harling (1962). These plants were said to differ from Erigeron in the following features: habit completely fruticose; involucre obconic to narrowly campanulate; phyllaries strongly imbricate in 4-6 unequal series, the outer grading into peduncular bracts; ray flowers more numerous than the disc flowers; achenes “slightly but distinctly” dimorphic, the ray achenes longer with fewer and weaker nerves than the disc achenes; and the embryo sac development monosporic. The putatively distinctive features in Darwiniothamnus of phyllary ar- rangement and involucral shape can be found in many species in different sections of Erigeron. Examples of slight achenial dimorphism can be found in sect. Fruticosus, sect. Polyactis and probably others, although the usual case is for the ray achenes to have more numerous and thicker ribs than those of the disc. The reduced number of disc flowers in the Galapagos plants is unusual in continental American species of Erigeron, but a number of Caribbean species have very small heads and a correspondingly reduced number of flowers. The embryology of relatively very few species (ca 25, none from South America) of Erigeron has been investigated (Harling 1951). Two of these, however, were shown by Harling himself to have strictly monosporic embryo sac development [E. philadelphicus (sect. Olygotrichium) and E. pere- grinus (sect. Fruticosus]; other species are variably mono-, bi- or tetra-sporic. In summary, the primary difference separating the two Galapagos Islands species from others of the genus is the peculiar, shrubby habit, and in my estimation this is inadequate justification for segregating them as a different genus. A case might be made, however, for regarding them as a section, but I leave this to someone more familiar with South American taxa. The tendency for the evolutionary development of a woody habit in insu- lar plants is well documented. Carlquist (1974) provided examples of Com- positae and many other families in which genera with herbaceous mainland species have developed woody, insular species. Among those in the Com- positae are insular species of Bidens, Centaurea, Dubautia, Perityle, Remya, Robinsonia, Senecio, Sonchus and Stephanodoria. As in Darwiniothamnus, most of these are rosette trees or rosette shrubs with long, mostly unbranched stems and leaves clustered near the stem tips. Besides those of the Galapagos Islands, suffrutescent species of Erigeron with similar habits also have developed on the Revillagigedo Islands (noted below), the Bahama Islands (£. darrellianus Hemsl., tentatively placed in sect. Olygotrichium) and the Juan Fernandez Islands (see comments following Terranea). Among these, the species of the Revillagigedo Islands are most similar in habit and other characters to those of the Galapagos Islands, and in contrast to I.M. Johnston’s original hypothesis of the relationship of the Nesom: Infrageneric treatment of Erigeron 89 latter with sect. Caenotus, I place them in Erigeronsect. Cincinnactis, which otherwise comprises species of the Mexican mainland. The habital similarity in these insular taxa almost certainly has resulted from convergent evolution. Terranea Colla, Mem. Reale Accad. Sci. Torin. 38:11. 1835. TYPE: T. fernandezia Colla [=£. fernandezia (Colla) Harling; non E. fernan- dezianus Solbrig]. Erigeron fernandezia was included by Solbrig (1962) in his broad concept of sect. Erigeron. Three other closely related species of Erigeron occur in the Juan Fernandez Islands and probably evolved from a single common ancestor from the mainland, perhaps E. leptorhizon DC. (Valdebenito et al. 1985). Astradelphus Remy, Ann. Sci. Nat. Bot. ser. 3. 12:185. 1849. (=Gusmania Remy in C. Gay, Fl. Chile 4:12. 1849; non Guzmania Ruiz & Pavon, 1802). TYPE: A. chilensis (Remy) Remy (=Gusmania chilensis Remy; =Erigeron remyanus Wedd., Chlor. And. 1:195. 1857). : According to Solbrig, Erigeron remyanus is perhaps a member of sect. Erigeron (sensu Solbrig 1962). GROUPS TRANSFERRED FROM ERIGERON TO OTHER GENERA 1. CONYZA Erigeronsect. Caenotus Nutt., Gen. Plant. 2:148. 1818. TYPE: E. canaden- sis L. [=Conyza canadensis (L.) Cronq.]. Caenotus (Nutt.) Rafin., FI. Tell. 2:50. 1836. Conyza sect. Caenotus (Nutt.) Cronq. ex Cuatr., Webbia 24:211. 1969. Erigeron sect. MultifloriG. Don in Loudon, Hort. Brit. 343. 1830. LECTO- TYPE (designated here): E. canadensis L. [=Conyza canadensis (L.) Cronq.]. Don also included 22 other species of a number of disparate lineages. I regard sect. Caenotus as a synonym of sect. Conyza, which includes C. canadensis (L.) Crong., C. bonariensis (L.) Cronq., C. ramosissima Cronq,., C. primulaefolia (Lam.) Lourteig & Cuatr. [=C. chilensis Spreng.] and others. See Nesom (in press) for further comments and synonymy. 2. CELMISIA Pappochroma Rafin., Fl. Tellur. 2:48. 1836. TYPE: P. uniflora Rafin. (=Erigeron pappochroma Labill., an Australian species, perhaps Cel- mista Cass., 1825). 3. ORITROPHIUM Erigeron sect. Oritrophium (Kunth) Benth. & Hook., Gen. Pl. 2:280. 1873. Based on Aster sect. Oritrophium Kunth. LECTOTYPE (Cuatre- casas 1961): Aster pellitus Kunth {[=Oritrophium peruvianum (Lam.) 90 PHY POLO GLA volume 67(1):67-93 July 1989 Cuatr.]. Celmisia sect. Oritrophium (Kunth) Solbrig. Oritrophium (Kunth) Cuatr. 4. TRIMORPHA Erigeron sect. Trimorpha (Cass. in F. Cuvier) DC., Prodr. 5:290. 1836. Trimorpha Cass., Bull. Sci. Soc. Philom. Paris 1817:137. 1817. TYPE: Trimorpha vulgaris Cass. in Cuvier, Dict. Sci. Nat. 55:324. 1828 (=£. acris L.). Trimorphaea Cass. in Cuvier, Dict. Sci. Nat. 37:462. 1825. Erigeron subg. Trimorpha (Cass.) M. Pop. Erigeron (sp.-group) Acres Rydb., Fl. Colorado 359. 1906, in clave. TYPE: E. acris L. I regard Trimorpha as a distinct genus more similar to Conyza than Erigeron (Nesom 1989b). 5. GENUS UNKNOWN Erigeron sect. Submultiflori G. Don in Loudon, Hort. Brit. 343. 1830. LECTOTYPE (designated here): EF. carolintanus L. Don also included Erigeron villarsti Bell. (=Erigeron atticus Villars of sect. Trimorpha in his sect. Submultzflorz. Fernald (1944) discussed the identity of Erigeron carolinianus L. but could only conclude that he could not identify it “unless it was something not orig- inally from Carolina or, as Dillenius thought, an abnormal individual.” It is certainly not an Erigeron, and I agree with Fernald that it is unlike any species of Euthamia. It is more Solzdago-like, but it is likely that the identity of this Linnaean epithet will remain equivocal. Because Don’s sect. Sub- multiflori was essentially meaningless in comprising two extremely divergent species from different genera, I feel justified in attaching it to a species name that itself has little chance of being clearly identified. To choose FE. villarsiias the lectotype would necessitate replacing the widely used and well established name of sect. Trimorpha. Tanazion Rafin., Fl. Tellur. 2:51. 1836. TYPE: T. tomentosum Rafin. (=£rigeron bahamensis Scop.). I do not know the identity of this plant. ACKNOWLEDGMENTS I thank Art Cronquist, John Kartesz, John Strother, Scott Sundberg and Billie Turner for their helpful comments and criticisms of the manuscript. Nesom: Infrageneric treatment of Erigeron 91 LITERATURE CITED Ake, K.M. 1984. Systematics of six species of Erigeron L. section Erigeron (Asteraceae). M.S. Thesis, University of Montana, Missoula. Atwood, N.D. & S.L. Welsh. 1988. An Erigeron from Nevada and a Pen- stemon from Idaho. Great Basin Naturalist 48:495-498. Becker, K.M. 1976. A new species of Erigeron (Asteraceae) from Alaska and Yukon. Brittonia 28:144-146. Bentham, G. & J.D. Hooker. 1873. Genera Plantarum, Vol. 2(1). L. Reeve & Co., London. Botschantzev, V.P. 1959. Erigeron. Pp. 191-288 in Flora U.R.S.S., vol. 25, ed. V.L. Komarov. U.S.S.R. Acad. Sciences, Leningrad. Brunsfeld, S.J. & G.L. Nesom. (in press). Erigeron salmonensis (Aster- aceae), a rare new species from Idaho. Brittonia 41:. Carlquist, S. 1974. Island Biology. Columbia Univ. Press, New York. Cronquist, A. 1947. Revision of the North American species of Erigeron, north of Mexico. Brittonia 6:121-302. Fernald, M.L. 1944. Is Erigeron carolintanus a valid American species? Rhodora 46:323-330. Gray, A. 1884. Synoptical Flora of North America. Vol. 1, part 2. Ivison, Blakeman, Taylor & Co. New York. Grierson, A.J.C. & K. H. Reichinger. 1982. Compositae V — Astereae. Flora Iranica 154:1-70 (+76 figures). Harling, G. 1951. Embryological studies in the Compositae:III. Astereae. Acta Horti Berg. 16:73-120. . 1962. On some Compositae endemic to the Galapagos Islands. Acta Horti Berg. 20:63-120. Holmgren, N.H. 1979. Subgeneric and sectional names for Intermountain Penstemon (Scrophulariaceae). Brittonia 31:358-364. Hultén, E. 1968. Flora of Alaska and Neighboring Territories. Stanford University Press, Stanford, California. Jones, A.G. 1980. A classification of the New World species of Aster (Aster- aceae). Syst. Bot. 32:230-239. Loudon, J.C. 1830. Loudon’s Hortus Britannicus Printed for Longman et al., London. Macbride, J.F. & E.B. Payson. 1917. A revision of the Erigerons of the series Multifidt. Contr. Gray Herb. 49:72-79. 92 PHYTOLOGIA volume 67(1):67-93 July 1989 Maguire, B. 1944. Great Basin plants—VIII. New species in Carez and Erigeron. Brittonia 5:199-202. Nesom, G.L. 1976. A new species of Erigeron (Asteraceae) and its relatives in southwestern Utah. Brittonia 28:263-272. 1982. Systematics of the Erigeron rusbyi group (Asteraceae) and delimitation of sect. Peregrinus. Syst. Bot. 7:457-470. 1989a. Taxonomy of Erigeron sect. Polyactis (Compositae: Astereae). Phytologia 66:415-455. 1989b. The separation of Trimorpha (Compositae: Astereae) from Erigeron. Phytologia 67:61-66. . (in press). Further definition of the genus Conyza (Compositae: Astereae). Phytologia 67:. & V.D. Roth. 1981. Erigeron scopulinus (Compositae) an en- demic from the southwestern United States. J. Arizona Nevada Acad. Sci. 16:39-42. & S. Sundberg. 1985. New combinations in Erigeron (Aster- aceae). Sida 11:249-250. , L. A. Vorobik & R. L. Hartman. (submitted). Recent collec- tions, identity, and classification of Aster blepharophyllus (Asteraceae: Astereae). Syst. Bot. Nuttall, T. 1818. Genera of North American plants. Reprint by Hafner Publ. Co., New York, 1971. . 1840-41. Descriptions of new species and genera of plants in the natural order of the Compositae, etc. Trans. Amer. Philos. Soc. ser. 2, 7:283-453. ; Rafinesque, C.S. 1836. Flora Telluriana. Philadephia: H. Probasco. (Fac- simile reprint, Arnold Arboretum, 1946). Rydberg, P.A. 1906. Erigeron. In: Flora of Colorado. Agric. Exp. Sta. Agric. Coll. Colorado Bull. 100:359-366. . 1918. Erigeron. In: Flora of the Rocky Mountains and Adjacent Plains. Pp. 896-911. Hafner Publishing Co., New York. Solbrig, O.T. 1960. The South American sections of Erigeron and their relation to Celmisia. Contr. Gray Herb. 188:65-86. 1962. The South American species of Frigeron. Contr. Gray Herb. 191:3-79. Spongberg, S.A. 1971. A systematic and evolutionary study of North Amer- ican arctic and alpine monocephalous species of Hrigeron (Compositae). Ph.D. dissertation, University of North Carolina, Chapel Hill. Nesom: Infrageneric treatment of Erigeron of Strother, J.L. & W. J. Ferlatte. 1988. Review of Erigeron eatonii and allied taxa (Compositae: Astereae). Madrono 35:77-91. Sundberg, S.D. 1986. The systematics of Aster subg. Ozytripolium (Com- positae) and historically allied species. Ph.D. dissertation, University of Texas, Austin. & A.G. Jones. 1987. Loudon’s Hortus Britannicus (1830): An early source of sectional names, necessitating nomenclatural changes in many genera—Aster: A case in point. Taxon 36:97-98. . 1988. Loudon’s Hortus Britannicus: A correction. Taxon 37:151. Torrey, J. & A. Gray. 1841. A Flora of North America, vol. 2, part 1. New York: Wiley & Putnam. Valdebenito, H., T.F. Stuessy & D.F. Crawford. 1985. Evolution of the genus Erigeron in the Juan Fernandez Islands, Chile. Amer. J. Bot. (Abstracts) 72:974. 1985. Vierhapper, F. 1906. Monographie der alpinen Erigeron—Arten Europas und Vorderasiens. Beih. Bot. Centralbl. 19:385-560. Welsh, S.L. 1983. A bouquet of daisies (Hrigeron, Compositae). Great Basin Naturalist 43:365-368. 1986. New taxa in miscellaneous families from Utah. Great Basin Naturalist 46:261-264. Phytologta (July 1989) 67(1):91-99. DOS NUEVOS REGISTROS DE ARISTOLOCHIA (ARISTOLOCHIACEAE) PARA VERACRUZ, MEXICO José Ortega Ortiz Universidad Veracruzana, Xalapa, Veracruz, MEXICO RESUMEN Aristolochia arborea Linden y A. pilosa H.B.K. se registran por primera vez para el estado de Veracruz, la primera de la region de Uxpanapa y la segunda cerca de Las Choapas. Se discuten las distribuciones, habitats, afinidades y relaciones de estos taxa. PALABRAS CLAVES: Aristolochiaceae, Aristolochia arborea, A. pilosa, Uxpanapa, Las Choapas, Veracruz. ABSTRACT Aristolochia arborea Linden and A. pilosa H.B.K. are recorded for the first time for the state of Veracruz, the former in the Rio Uxpanapa watershed, and the latter near Las Choapas. Their known distributions, habitats and relationships are discussed. KEY WORDS: Aristolochiaceae, Aristolochia arborea, A. pilosa, Uxpanapa, Las Choapas, Veracruz. Durante el estudio taxondmico de la familia Aristolochiaceae que el au- tor realizé para la Flora de Veracruz (Ortega Ortiz 1988), se encontraron ejemplares de Aristolochia arborea y A. pilosa, los cuales constituyen nuevos registros para el estado. Aristolochia arborea (Figure 1) se describid tomando como base los es- pecimenes colectados por Ghiesbreght en el estado de Chiapas, de locali- dad desconocida. Esta especie se encuentra distribuida en las selvas altas perennifolias del sureste de México (Chiapas, Tabasco, Veracruz y proba- blamente Oaxaca), Guatemala y El Salvador. Pfeifer (1966) incluyo a A. steyermarkiu Stand]. como sindnimo de A. arborea, aunque posteriormente Barringer (1983) encontré diferencias que justifican su reconocimiento como una especie distinta. Aristolochia steyermarkii es un arbusto con flores axi- lares, y el limbo de la flor tiene una protuberancia sésil y rugosa (la ampolla), 94 Ortega Ortiz: Nuevos registros para Aristolochia en México 95 ria Hy Ey Fig. 1. Anrtstolochia arborea. a, Forma biolégica; a, recuadro del ‘| tallo, mostrando las flores caulinares; b, rama con hojas; c, flor ' (en vista lateral y frontal); d, corte transversal de la flor mos | trando el ginostemo y la ampolla; e, recuadro de la flor, mostran | do en detalle la ampolla; f, ginostemo; g, fruto; h, semilla. Ilustraci6n por Seatiel Guiochin, basada en fotografias de Dorantes et al. 2957 (a, a') y en ejemplares de J. Ortega y R. Ontega 348 (b, c, d, e, f, g, h). es ae i wae pee Se 96 PAHAYsE: Oo; O:GubiA volume 67(1):94-99 July 1989 a diferencia de A. arborea que presenta flores caulinares y el limbo posee una ampolla estipitada. Las especies anteriores estan estrechamente relacionadas con A. impudica Ortega (Ortega, 1987), la cual se distingue por la presencia de una inflorescencia de 32-50 cm de largo, los pedicelos geniculados de 6-13 cm de largo, una ampolla de 2.5-4 cm de largo, 2-5 cm de ancho, y el margen revoluto de los lobulos laterales del limbo del caliz. Aristolochia arborea ha sido escasamente colectada. Posee una distribucion que corresponde, en parte, a un area de alta precipitacion que se extiende desde el S de Veracruz y S de Tabasco al N de Chiapas, pero que no se continua hasta Guatemala. Esta zona corresponde a un refugio de elemen- tos tropicales durante el Pleisticeno tardio (Wendt 1985). Sin embargo, esta especie continua su distribucién hasta Guatemala y El Salvador (Figure 2). Aristolochia pilosa es una liana de amplia distribucion desde México hasta el norte de Sudamérica. Recientemente ha sido encontrada cerca de Las Choapas, Veracruz, entre Nanchital y Cuichapa en altitudes ca de 50 m. Ocurre principalmente cerca de arroyos o a la orilla de carreteras sobre pequenos arbustos, en suelos arenosos con vegetacion secundaria de selva baja caducifolia. Aristolochia pilosa se reconoce facilmente por el indumento abundante y ferrugineo que se presenta en toda la superficie externa de la planta, asi como su flor, cuyo limbo es espatulado-oblongo con papilas moreno oscuras en el apice, y por el fruto de 5-7.5 cm de largo. A continuacion se presenta una descripcion breve de A. arboreay A. pilosa basada en los ejemplares de Veracruz. Aristolochia arborea Linden, Cat. Pl. Exot. 13:6. 1858. TIPO: MEXICO: Chiapas, Ghiesbreght s.n. (Holotipo BM). Arbustos pequenos, perennes, de 3-5 m de alto; tallo erecto. Hojas al- ternas, aromaticas, lanceoladas, 8-26 cm de largo, 2-6.5 cm de ancho; apice acuminado o caudilado, base redondeada y obtusa. Inflorescencia cauliflora, racemosa, 5-8 cm de ancho; flores bisexuales, caliz geniculado, de color purpura-guinda, utriculo ovoide-elipsoide, siringe ausente, tubo doblado, lim- bo trilobado, lobulo central mas largo, con una mancha blanca en el apice, una ambolla purpurea, globosa-capitada, esponjosa y con pelos glandulares por arriba del orificio del tubo, de 1 cm de largo y un estipite de 8-9 mm de largo; estambres 6, en un ginostemo con 3 ldbulos estigmaticos y 2 anteras por lobulo. Fruto una capsula cilindrica, ligeramente doblada, 10-13 cm de largo, 2.5-3.5 cm de ancho; semillas cordiforme-triangulares, 5-7 mm de largo y ancho. Ejemplares examinados. MEXICO: Veracruz. Mun. Hidalgotitlan, km 0-2 al S del campamento Hnos. Cedillo, camino a Rio Alegre, desviacién al E, 140 m, 23 May 1974, Dorantes et al. 2957 (ENCB,MEXU,XAL); km 3 del campamento Hnos. Cedillo-Rio Alegre, 150 m, 30 Ago 1974, Dorantes et al. 3557 (XAL). Ortega Ortiz: Nuevos registros para Aristolochia en México 97 Golfo de Mexico Honduras A. arborea * A. pilosa e Fig. 2. Mapa de la distribucién de Aristolochia arborea Linden y A. pilosa H.B. & K., basado en ejemplares examinados en este trabajo. 98 PAAET O.L-OsG tA volume 67(1):94-99 July 1989 Nombre Local. Flor de Chapo, Pachuli (Tabasco), Tecolotillo (Guatemala). Usos. El tallo se utiliza como febrifugo. Ecologia: Selva alta perennifolia, cerca de rios, arroyos y lugares mon- tanosos, sobre suelos calcareos y pedregosos. Aristolochia pilosa H.B.K., Nov. Gen. Sp. 2:156. 1819. TIPO: ECUADOR: ca Guayaquil, Humboldt & Bonpland s.n. (Holotipo B; foto XAL!). Liana, perenne, 2-5 m de largo, el tallo piloso o hirsuto. Hojas cordi- formes, 9-14.5 cm de largo, 7-10 cm de ancho, membranaceas, la haz glabras- cente, el envés densamente piloso o hirsuto, el margen entero, el apice redondo u obtuso, la base cordada; peciolo cilindrico, 3-4 cm de largo, piloso o hir- suto. Inflorenscia axilar, solitaria, 6-14.5 cm de largo, 1-2 cm de ancho; flores bisexuales, de color verde amarilento y purpura; caliz arcuado, 6-8.5 cm de largo, 1-2 cm de ancho, piloso-hirsuto en la superficie externa, el utriculo verde amarillo, ovoide o subpiriforme, 1.8-2.5 cm de largo, 1-1.5 cm de an- cho, lanuloso-flocoso en la parte interna, el siringe de color crema, un ligero borde inequilatero en el interior, ca 1 mm de largo, 5-7 mm de ancho, opaco y glabro, el tubo verde amarillo, rectilineo-arcuado, 1.8-2.5 cm de largo, 0.3-0.5 cm de ancho, opaco, con pelos estrigosos en la superficie interna; el limbo con un solo lobulo, de color pupura y amarillo, espatulado-oblongo, 3-5.5 cm de largo, 1-2 cm de ancho, brillante, esparcidamente fimbriado en la superficie interna, el margen involuto, la base redondo-cordada; estambres 6, insertos en un ginostemo capitado, 5-8 mm de largo, 3-6 mm de ancho, anteras 1 por lobulo. Fruto capsular, lenoso, elipsoide, 5-7.5 cm de largo, 1.5-2 cm de ancho, septifraga, piloso-hirsuto; semillas de color moreno claro-moreno oscuro, planas, triangular-cordiformes, 4-5 mm de largo, 3-4 mm de ancho y 0.5-0.7 mm de grueso, el margen entero, el apice agudo, la base cordada, la testa delgada y aceitosa, granulosa, pubérula; endospermo abundante. Ejemplares examinados: MEXICO: Veracruz. Mun. Coatzacoalcos, 10 km WNW of Las Choapas on road to Nanchital, 50 m, 2 Ene 1986, Nee 32464 (NY,XAL); Carretera Nanchital-Las Choapas, 2 km antes de llegar a la desviacion a Chuichapa, 20 Jun 1987 Ortega, McDonald & Aguilar 507 (XAL). Nombre local: Huehuecho, huehueche, pecho de paloma, curanina (Chia- pas); sombrerito, hediondilla (Guatemala). Ecologia: Selva baja caducifolia, vegetacion secundaria de selva alta peren- nifolia, ruderal. Floracion. Julio a Enero. AGRADECIMIENTOS Se agradece al Dr. Kerry Barringer por la bibliografia proporcionada para Ortega Ortiz: Nuevos registros para Aristolochia en México 99 la realizacion de este trabajo, al Dr. Andrew McDonald por su ayuda en mi formacion profesional y revision critica del manuscrito. Asimismo a las siguientes personas que me apoyaron en la elaboracion de este trabajo y sus valiosos comentarios al manuscritos: Dr. Andrew Vovides, Biol. Rocio Jiménez, Biol. Hector Narave y Biol. Roberto Ortega. A mi amigo Seatiel Guiochin H., por la elaboracion de las figuras. Se agradece a los curadores de los siguientes herbarios: CSAT, ENCB, F, IBUJAT, GH, MEXU, MO, US y XAL por las facilidades que brindaron para la revision de material. Tambien, se agradece a la Dra. Linda Escobar por su revision critica del manuscrito. REFERENCIAS Barringer, K. 1983. Aristolochiaceae. En: Flora Costaricensis. Field Mus. Nat. Hist., Bot. 13:79-87. Ortega Ortiz, J. 1987. Notas del herbario XAL. IX. Dos especies nuevas hexandreas de Aristolochia (Aristolochiaceae) de Veracruz, México. Bidtica 12:209-216. 1988. Aristolochiaceae. En: Flora de Veracruz (en comité edito- rial). Pfeifer, H.W. 1966. Revision of the North and Central American hexandrous species of Aristolochia (Aristolochiaceae). Ann. Missouri Bot. Gard. 53:115-196. Wendt, T. 1985. A new simple-leaved species of Recchia (Simaroubaceae) from Southeastern Mexico. Brittonia 37:219-225. Phytologia (July 1989) 67(1):100-103. CONTRIBUTION TO THE LICHEN FLORA OF VE LA, VII ah NEZUELA, Manuel Lopez-Figueiras y Antonio Morales Méndez Instituto de Investigaciones, Facultad de Farmacia, Universidad de Los Andes, Mérida, VENEZUELA ABSTRACT Several new distribution records for Pyzine are reported from Venezuela. KEY WORDS: Floristics, lichens, Pyzine, Venezuela, Neotropics. Study of Pyzine in Venezuela has been superficial, mainly due to the small number of collectors and the lack of correctly identified herbarium specimens. Vareschi (1973) and Lopez-Figueiras (1986), respectively mentioned seven and five species present in Venezuela. Later relationships of one of us (A.M.M.) with Dr. K. Kalb [monographer of the Brasilian pyxines (1987)], has allowed an increase in accurate herbar- ium specimens by his kindly help in botanical identification of our material. The results of these studies are reported as follows: Pyzine berteroana (Fée) Imsh. Estado Falcon: Alrededores de Sinamaica, proximidades de Cerro Azul (Sierra Ziruma o Empalado), Lopez-F. 21571. Estado Mérida: Mocomboco, Aricagua, Lopez-F. 8969, 12607. Pyzine caesiopruinosa (Nyl.) Imsh. Estado Lara: En Pico-Pico (Sierra de Bobare), Lopez-F. & R. Smith 20768-B. Estado Mérida: El Pedregal de Jali, junto a La Carbonera, Lopez-F. 18145, 18174. Estado Tachira: Proximidades de Providencia, cercanias de Rubio, car- retera (vieja) San Cristoébal-Rubio, Lopez-F. 25613. Estado Trujillo: Las Palmas, carretera Carache-Agua de Obispo, Lépez-F. 28020, Cercanias de Santa Ana de Trujillo, carretera Bocon6é-Valera, Lopez- F. 28130; Vertiente oriental del paramo del Guache, Los Cortijos, cercanias de Carache, Lopez-F. 28398,28411. Pyzine cocoes (Sw.) Nyl. 100 Lopez-Figueiras y Morales Méndez: Pyzine in Venezuela 101 Estado Falcon: Paraguana: vi Coro-Adicora, Lopez-F. 19190; entre El Hato y Pueblo Nuevo, Lopez-F. 19217, el Balsamal, via Pueblo Nuevo hacia el Vinculo, Lopez-F. 21242,21244; alo largo de Monte Cano, Lopez-F. 21262; Cerro Santa Ana, a lo largo del arroyo Santa Ana, Lopez-F. 21345; a lo largo del arroyo Siraba, Lépez-F. 21349, NW del cerro Santa Ana frente a Bella Vista, Lopez-F. 21743, Punta Prudencio, al suroeste de Tacuato, Lopez-F., R. Wingfield & A. Morales 32495. Inmediaciones del rio Ricoa, via Coro- Moron, Lopez-F. 21691. Entre La Cruz y La Goya, vi Coro-Churuguara, Lopez-F. 21878. Sierra San Luis, alrededores de la Piedra Campana, entre Carrizalito y Cucaire, Lépez-F. & R. Wingfield 22405,22406. Proximidades del rio Japure, entre Paso Calderas y Buruica, via Pedregal-Tapure, Ldpez-F. & R. Wingfield 22510. Estado Lara: Sierra Portuguesa, Loma El Leon, cercanias de Barquisimeto, Loépez-F. & R. Smith 21168. Estado Merida: Arriba de Las Gonzalez, carretera Mérida-Lagunillas, Lopez-F. 16294; zona xerofila de Lagunillas, Lopez-F. 18031. Estado Trujillo: Entre Carache y La Cuchilla, Lopez-F. 22278. Pyzine cognata Stirt. Estado Merida: Alrededores de Minzal, cercanias de Mesa Quintero, via hacia Pico de Horma, Lopez-F. 23238; La Sabana, area de Pico de Horma, al suroeste de Mesa Quintero, Lopez-F. & H. Rodriguez 23331, Parque Na- cional Sierra Nevada, alrededores de La Mucuy, Lépez-F. & H. Rodriguez 27184,27236. Estado Tachira: Margenes del rio Pedernales, Laguna Garcia, zona de Pregoneros, Lopez-F. & A. Morales 32461. Estado Trujillo: Paramo de Cende, Lopez-F. 12992. Pyzine coralligera Malme Estado Lara: Parque Nacional Yacambu, Sierra Portuguesa, Lopez-F. S29 AN agnG: Estado Mérida: Finca Los Topes, entre Chiguara y La Trampa, Lopez-F. & A. Morales 31005. Estado Tachira: Margenes del rio Pedernales, Laguna Garcia, cercanias de Pregoneros, Lopez-F. & A. Morales 32454-A. Estado Trujillo: Entre El Filo de San Isidro y La Becerrera, a 25 km de La Concepcion de Carache, por la carretera nueva, Lopez-F. & H. Rodriguez 28063. Pyzine daedalea Krog & R. Sant. Estado Mérida: Mocomboco, Aricagua, Lopez-F. 12602; San Juanito, Chiguara, A. Morales 116. 102 PRY DO DOG TA volume 67(1):100-103 July 1989 Pyzine endolutea Kalb Estado Mérida: El Maciegal, cuenca del rio La Pedregosa, cercanias de Mérida, Lopez-F. 10643; Finca Los Topes, cercaniias de Chiguara, Lopez-F. & A. Morales 32473. Estado Trujillo: Carretera (vieja) Trujillo-La Cristalina~-Bocono, Lopez- F. & M. Keogh 11240,11248,11249,11257, Los Cortijos, vertiente oriental del paramo del Guache, cercanias de Carache, Lopez-F. 28388. Pyzine eschweileri (Tuck.) Vain. Estado Lara: Sabanas El] Altar-Yaritagua, proximidades de un puente sobre el rio Turbio, Lépez-F. & R. Smith 16479. Estado Mérida: Sierra Nevada de Mérida, quebrada Fafoy, cercaniias de El Carrizal, Lopez-F. & M. Hale 20123,20174,20274. Pyzine limbulata Mull. Arg. Estado Mérida: Paramo de Pinago, via Pinago, a lo largo de la carretera entre las quebradas Las Tapias y Pinago, Lopez-F. 27745. Esta especie es primera cita para el hemisferio occidental. Pyzine microspora Vain. Estado Lara: Sierra Portuguesa, Loma El Leon, proximidades de Bar- quisimeto, Lopez-F. & R. Smith 21148. Estado Mérida: El Pedregal de Jali, junto a La Carbonera, Lopez-F. IROL fle Pyzine petricola Nyl. in Cromb. Estado Lara: Hacienda Los Cristales, via Barquisimeto-Sarare, Ldpez-F. 16089, El Gamelotal, via Barquisimeto-El Altar-Yaritagua, Lopez-F. 16130- B; entre Barbacoas y San Pedro, Loépez-F. 19025,19039,19064,19067,19072, 20813, Sierra Portuguesa, Loma El Leon, proximidades de Barquisimeto, Lopez-F. & R. Smith 21180,21210, Sierra Ziruma o Empalado, en el Carron, Lopez-F. & R. Smith 20813. Estado Mérida: Arriba de Las Gonzalez, carretera Mérida-Lagunillas, Lopez-F. 16292,16299-B,16300-B,16303-A,16303-B; Zona xerofile de Lagu- nillas, Lépez-F. 18053,18055-A; El Pedregal de Jali, junto a La Carbonera, Lopez-F. 18156; Arriba de Tovar en la carretera hacia el Paramo de Marino, Lépez-F. 24695, Proximidades de Mesa Bolivar, cercanias del entronque de la carretera Mesa Bolivar-El Vigia, Lopez-F. 25540-B. Estado Tachira: Las Coloradas, zona xerofila de La Grita, Lopez-F. 14409; alrededores de Salomom, via San Cristébal-Cordero-Alto de El Zum- bador, Lopez-F. 24803. Estado Trujillo: Paramo de La Nariz, alrededores de T.V., Lopez-F. 16680-A; entre Carache y La Cuchilla, Lopez-F. 22271; En Florencia, paramo El Turmal, inmediaciones del camino hacia el paramo Cendé, Lopez-F. 30734. Lopez-Figueiras y Morales Méndez: Pyzine in Venezuela 103 Pyzine pungens Zahlbr. Estado Lara: Sierra de Barbacoas entre San Pedro y Barbacoas, Lopez-F. 18983-B. Estado Mérida: Mocomboco, Aricagua, Lopez-F. 12610; El Pedregal de Jali, junto a La Carbonera, Lopez-F. 18169,26795, La Carbonera, cercanias de Mérida, Lépez-F. 22058; El Valle cercanias de Mérida, Lopez-F. 29295. Pyzine pyzinoides (Mull. Arg.) Kalb Estado Mérida: El Paramito, un sector de La Carbonera, Lopez-F. 17523. Pyzine rhodesica Vain ex Lynge. Estado Mérida: Mocomboco, Aricagua, Lopez-F. 12608; Finca San Isidro, La Carbonera, via La Azulita, Lopez-F. 26748; La Carbonera, Lopez-F. 30086. Pyzine subcinerea Stirt. Estado Falcon: Sierra de San Luis, La Chapa, via Las Negritas-Uria, Lopez-F. 19241. Estado Lara: A lo largo de la carretera entre Guarico y Chubasquin, Lopez-F. 17207-B, 17208. ACKNOWLEDGMENTS The authors are grateful to Dr. K. Kalb by his invaluable aid in the revision of the material and to C.D.C.H.T. of Universidad de Los Andes, by the economic help to the Fa-91-88 project. LITERATURE CITED Kalb, K. 1987. Brasilianishe Flechten. 1. Die Gattung Pyzine. Bibl. Lichenol., J. Cramer, Berlin-Stuttgart. Lopez F., M. 1986. Censo de Macroliquenes Venezolanos de los Estados Falcon, Lara, Mérida, Tachira y Trujillo. U.L.A., Facultad Farmacia 1-521. Vareschi, W. 1973. Catalogo de Liquenes de Venezuela. Acta Bot. Venez. 8(1-4):177-245. Phytologia (July 1989) 67(1): 104-106. NEW COMBINATIONS IN Te (COMPOSITAE: ASTEREAE) Guy L. Nesom Department of Botany, University of Texas, Austin, Texas 78713 USA ABSTRACT Two new combinations in Ericameriaare proposed: E. palmeri var. pachylepis and E. parishii var. peninsularis. KEY WORDS: Ericameria, Haplopappus, Asteraceae, México, systematics. In the preparation of a taxonomic treatment of the Mexican species of Ericameria, two new combinations are necessary. Ericameria palmeri (A. Gray) H.M. Hall var. pachylepis (H.M. Hall) Nesom, comb. nov. Based on Haplopappus palmeri subsp. pachylepis H.M. Hall, Carnegie Inst. Washington Publ. No. 389:267. 1928. Hap- lopappus palmeri var. pachylepis (H.M. Hall) Munz, Man S. Calif. 522. 1935. Ericameria palmeri is recognized by its strictly erect stems and ovate- cylindric capitulescences with radiate heads. Var. pachylepis, the northern variant of the species, which ranges from Riverside County, California, north- ward to Ventura County, differs from var. palmert primarily in its larger leaves (5-16 mm vs 10-40 mm long) and the broader, more definite gland on the apical half of the phyllaries. Differences noted by Munz (1974) in the in- volucre height and number of ray and disc flowers separate the two taxa inconsistently. Ericameria parishii (E. Greene) H.M. Hall var. peninsularis (R. Moran) Nesom, comb. nov. Based on Haplopappus arborescens subsp. peninsu- laris R. Moran, Trans. San Diego Soc. Nat. Hist. 15:152. 1969. The three taxa of the Ericameria arborescens (A. Gray) H.M. Hall group occupy three separate geographic zones. Moran (1969) recognized each of the three as a subspecies of £. arborescens (subsp. arborescens, subsp. parishii (E. Greene) R. Moran and subsp. peninsularis R. Moran), but all other studies of the California flora have maintained EF. arborescensand E. parishiias distinct species. Ericameria arborescens, the northernmost form, apparently does not 104 Nesorn: New combinations in Ericameria 105 intergrade morphologically with FE. parishiz, which is restricted to southern California, where their ranges meet in southern California. On the other hand, Ericameria parishi var. parishiz and var. peninsularis, which is endemic to Baja California Norte, are similar in all respects except leaf shape, and according to Moran (1969), they intergrade in this character. “Since some specimens are virtually identical, [var. peninsularis] can scarcely be treated as a species separate from [E.] parishit” (Moran 1969, p. 154). Hall’s concept of E. parishit (1928) also included var. peninsularis, because he cited as E. parishi a collection from Sierra San Pedro Martir (Robertson 48, UC) that was included by Moran as var. peninsularis. Moran’s solution in identifying all three taxa as subspecies of one species emphasized their close relationship but disregarded the clear discontinuity between FE. arborescens and E. parishii as well as the greater similarity between var. parishit and var. peninsularis. In my view, these taxa are best identified by the following synoptic key. 1. Heads on ultimate peduncles (4-) 7-15 mm long, in loose cymes; disc flowers 18-25 per head, prominently hairy on the tube and lower OLR CLD1 fe Site, Sees eee eens, « Sa eae Sanh ae Meee ea es eae E. arborescens 1. Heads on ultimate peduncles 0-5 mm long, in compact cymes; disc flowers 8-18 per head, glabrate to very sparsely papillate or minutely hairy on tities Shave ater he crap ctor ace eaalese aes = asJne hake xe os 3S) aR (2) 2. Leaves linear-oblanceolate to linear, 1.5-2.0 (-3.5) mm | ek Sata aa gy E. parishi var. pentnsularis 2. Leaves narrowly elliptic to elliptic-oblanceolate, 3-10 mm NGC Sita cea ge ct tats pst oid Bat, Sex we ctuatoysh nes E. parishi var. parish ACKNOWLEDGMENTS I thank Drs. Billie Turner and Marshall Johnston for their comments on the manuscript. LITERATURE CITED Hall, H.M. 1928. Sect. Ericameria, in The Genus Haplopappus—A phylo- genetic study in the Compositae. Carnegie Inst. Washington, Publ. 389:258-288. 106 PRY TOL Oe'G TA volume 67(1):104-106 July 1989 Munz, P.A. 1974. Haplopappus. Pp. 174-181 in Flora of Southern Califor- nia. Univ. California Press, Berkeley. Moran, R. 1969. Five new taxa of Haplopappus (Compositae) from Baja California, Mexico. Trans. San Diego Soc. Nat. Hist. 15:149-164. Urbatsch, L.E. & J.R. Wussow. 1979. The taxonomic affinities of Haplopap- pus linearifolius (Asteraceae — Astereae). Brittonia 31:265-275. Phylologta (July 1989) 67(1):107- 108. A NEW SPECIES OF VERBESINA SECTION VERBESINARIA FROM THE DOMINICAN REPUBLIC John Olsen Department of Biology, Rhodes College, Memphis, Tennessee 38112 USA ABSTRACT Verbesina howardiana is described from Hispanola. KEY WORDS: Asteraceae, Verbesina, Lesser Antilles, systemat- ics. During the preparation of the Flora of the Lesser Antilles, the following new taxon was revealed. Verbesina howardiana Olsen, sp. nov. TYPE: DOMINICAN REPUB- LIC: St. Paul Parish. Morne Trois Piton, 4400 ft, barren area on moun- tain top, 23 Oct 1964, D.H. Nicolson 1815. (holotype: GH). Frutex ca 1 m altus; cauli exalati, valde cicatricibus foliorum notati. Folia fasciculata sub capitulis, laminae obovatae, apex acute latus ad rotundatem, folia margine minutissime serrata; supra glabrescentia, subtus resinose pubescentia dense in nervis. Capitula ca 1.5 cm diametro; flores radii 19-21, ligulae luteae, ca 11-15 mm longae; flores disci numerosi corollae luteae, tubi glabri. Achaenia nigra, glabra, alae non latae; pappus biaristatus, inaequalis. Shrub to 1.0 m tall; stems terete, pithy, unwinged, nearly glabrous below and strongly marked by prominent leaf scars, becoming densely pubescent above. Leaves clustered below the capitulescence, obovate, to 12.0 cm long, 7.0 cm wide, apex rounded to broadly acute, base narrows to a short resinous pubescent petiole, < 1.0 cm long, margins revolute, minutely serrate, blade glabrous above or with a few resinous hairs along the midvein, densely resinous pubescent along all veins below, very sparsely pubescent on the blade surface. Capitulescence a loose corymb of 12-20 heads on densely pubescent peduncles to 6.0 cm long; heads to 1.5 cm in diameter, excluding the rays. Involucre 2-3 seriate, outer series oblong, 3.0-4.5 mm long, 1.2-1.5 mm wide, inner series oblanceolate, 5.0-5.8 mm long, 1.2-1.5 mm wide, apex rounded to broadly acute, pubescent at the base only, margins not ciliate. Pales 6.5-8.0 107 108 PAY TOLOGIA volume 67(1):107-108 July 1989 mm long, 1.7-2.0 mm wide, glabrous, apex herbaceous, margins not ciliate. Rays 19-21, yellow, ligules 11-15 mm long, 3.2-3.5 mm wide, apex minutely 3- dentate, styles well exserted from the tube; tube 1.8-2.0 mm long, 0.4-0.5 mm wide, pubescent. Disc florets numerous, ca 75, corolla yellow, tubular, 3.8-4.0 mm long, 1.4-1.5 mm wide, tube 1.5 mm long, 0.5 mm wide, glabrous. Ray achenes black, glabrous, 5.0-5.5 mm long, 1.3-1.5 mm wide, thinly winged, wings < 0.2 mm wide; awns unequal, to 4.5 mm long. Disc achenes black, glabrous, 5.3-5.5 mm long, 1.7-2.0 mm wide, thinly winged, wings < 0.3 mm wide; awns unequal, to 5.0 mm long. Additional specimens examined (all come from Morne Trois Piton): K.L. Chambers 2588 (GH); W.H. Hodge 1397(GH); C. Kimber 975 (GH [2 sheets]). V. howardiana is a member of section Verbesinaria. All collections of this taxon have been from elevations above 4000 ft, at or near the summit of Morne Trois Pitons in areas described as rain forest (400 inches per year) where it grows in open areas. V. howardiana is related to several Caribbean taxa including V. guade- loupensis Urb., V. karsticola Proctor and V. caymanensis Proctor. V. howar- diana is easily differentiated from V. guadeloupensis on the basis of its corky stem with prominent leaf scars and obovate leaves. It also has a completely glabrous tube on the disc floret. The Urban species has lanceolate leaves, lacks the prominent leaf scars and has a pubescent tube on the disc florets. Both V. caymanensis and V. karsticola are members of section Ochractinia. V. caymanensis has the prominent leaf scars seen in V. howardiana, but has white florets and the leaves are serrate to weakly lobed, especially at the apex. V. karsticola has a leaf morphology which more closely matches V. howardiana, but again has white florets and lacks the prominent leaf scars. It is a pleasure to name this taxon for Dr. Richard Howard who brought it to my attention during his preparation of the Flora of the Lesser Antilles. ACKNOWLEDGMENTS I thank Dr. Tom Jolly who assisted with the Latin description and Drs. B.L. Turner and G.L. Nesom who reviewed the manuscript. Ra Information for Authors Articles from all lines of botany, biographical sketches, critical reviews and summaries of literature will be considered for publication in PHYTOLOGIA. Manuscripts may be submitted either on computer diskette, or as typescript. Diskettes may be 5.25 inches or 3.5 inches but must be written in PC/DOS (MS/DOS) format or as flat ASCII files. Typescript manuscripts should be single spaced and will be read into the computer using a page scanner. The scanner will read standard typewriter fonts but will not read dot matrix print. Manuscripts submit- ted in dot matrix print cannot be accepted. Use underscore (not italics) for scientific names. Corrections made on typescript manuscripts must be complete and neat as the scanner will not read them otherwise. Language of manuscripts may be either English or Spanish. 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