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In North America Author(S): Chia Jui Chen, Meghan G. Mendenhall and Billie L

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In North America Author(S): Chia Jui Chen, Meghan G. Mendenhall and Billie L Taxonomy of Thermopsis (Fabaceae) in North America Author(s): Chia Jui Chen, Meghan G. Mendenhall and Billie L. Turner Source: Annals of the Missouri Botanical Garden, Vol. 81, No. 4 (1994), pp. 714-742 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/2399917 Accessed: 18-06-2015 19:07 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Missouri Botanical Garden Press is collaborating with JSTOR to digitize, preserve and extend access to Annals of the Missouri Botanical Garden. http://www.jstor.org This content downloaded from 128.83.205.78 on Thu, 18 Jun 2015 19:07:36 UTC All use subject to JSTOR Terms and Conditions TAXONOMYOF THERMOPSIS Chia JuiChen,2 Meghan G. Mendenhall,3 (FABACEAE) IN NORTH and Billie L. Turner4 AMERICA' ABSTRACT Comprehensive reevaluation of both herbarium specimens and fieldobservations of the North American Thermopsis leads to our recognition of 10 species. Three species occur in the southern Appalachians: T. villosa, T. mollis, and T fraxinifolia. The Rocky Mountains and intermountainregions are populated by the relativelyvariable and widespread species T. divaricarpa, T. montana, and T rhombifolia. The Pacific coastal mountains of California are dominated by the variable T californica (with three infraspecifictaxa), which gives way to T gracilis in northern California and western Oregon. Thermopsis macrophylla and T robusta have restricted ranges in California. Widelydistributed throughout the westernand species in the Rocky Mountainand intermountain southeasternUnited States, Thermopsis is a con- regions.The threelong-recognized species of the spicuous,often weedy genus. Thoughit is a her- southeasternUnited States are maintained. baceous plant,its relativelylarge habit and stun- The followingherbaria have generouslysupplied ning racemes of golden or brightyellow flowers the approximately6000 specimensused in this and flattened,rattling pods have no doubtcontrib- study(number of sheetsloaned givenin parenthe- uted to its early and thoroughcollection. These ses): ARIZ (113), CAS (516), COLO (181), F factors,perhaps, should have led to a clear un- (193), GH (511), JEPS (16), LL (15), MO (444), derstandingof its systematics,but a lack of diag- MONTU (91), NCU (112), ND (99), NEBC (3), nosticcharacters within the genus,morphological NY (715), ORE (78), OSC (85), PE (52), PH intergradation,and possible hybridizationhave (193), POM (75), RM (611), RSA (302), TENN complicatedthe taxonomy of Thermopsis. Modern (71), TEX (163), UC (401), US (419), UT (83), treatments(e.g., Isely, 1981) include from4 to WS (161), WTU (200). 10 species in NorthAmerica, though over 20 spe- cificnames have been proposed. HISTORY OF THE GENUS The currentstudy is based on over 20,000 km of roadsidefield study in NorthAmerica (spanning Thermopsis(Greek "thermos" = lupine+ "op- 25 years for B. Turner),greenhouse studies (by sis" = appearance) was originallyincluded in So- M. Mendenhall),and the examinationof approxi- phora by Linnaeus (1753). Willdenow (1799) mately6000 herbariumspecimens (by all of the placed the taxon in Podalyria. In 1811, Robert authors). We find 10 species of Thermopsis in Brownerected Thermopsisbased upon Sophora NorthAmerica. This treatmentdiffers from earlier 1upinoidesL. Brownrenamed the latterThermop- evaluationsprimarily in the rankand affiliationsof sis lanceolata, whichis contraryto the present taxa in the westernUnited States. While an ex- Code of Botanical Nomenclature(Greuter et al., tremeperspective would combine the westerntaxa 1988); thename Thermopsislupinoides (L.) Link, intotwo highly variable species, we recognizefour erectedin 1821 and recognizedstill, precludes the speciesin the Pacificcoastal states,three of which use of thisspecific epithet for the type species of have restrictedranges, and threewidely distributed the genus.In 1818, Nuttallcalled the genus Ther- 'This projectwas partiallysupported by the NationalScience Foundationof Chinaand the May ScholarFellowship at the MissouriBotanical Garden. This studywas undertakenfor the most part at the Plant Resources Center, Departmentof Botany,University of Texas at Austin.We are indebtedto Peter H. Raven at the MissouriBotanical Gardenfor encouraging the seniorauthor to undertakethis study. We also thankGuy L. Nesom, Carol A. Todzia, and Steve Ginzbargat the Universityof Texas at Austin,and Peter C. Hoch at the MissouriBotanical Garden for assistancein the preparationof the manuscript. 2 Instituteof Botany,Academia Sinica, Xiangshan,Beijing 100093, China. 3 Authorfor correspondence, Department of Botany,University of Texas, Austin,Texas 78713, U.S.A. 4 Plant ResourcesCenter, Department of Botany,University of Texas, Austin,Texas 78713, U.S.A. ANN. MISSOURI BOT. GARD. 81: 714--742. 1994. This content downloaded from 128.83.205.78 on Thu, 18 Jun 2015 19:07:36 UTC All use subject to JSTOR Terms and Conditions Volume 81, Number4 Chen et al. 715 1994 Taxonomy of Thermopsis mia. Rafinesqueerected the generaScolobus and the NorthAmerican coast of the BeringSea. Most Drepilia in 1819, both based on specimensof ofthe Asian species were treated by Shishkin (1945). T7hermopsisrhombifolia. Judgingfrom the relativeuniformity of the North Larisey (1940) revised the North American Americanspecies, taxa ancestralto extant Ther- T7hermopsisbecause she saw considerable"taxo- mopsisprobably crossed Beringia into North Amer- nomic confusion,"which she eventuallyascribed ica. to "bibliographical"problems. She distilled22 pub- On the NorthAmerican continent, Thermopsis lishednames to 10 speciesby reducingthe 4 south- is distributedthroughout the westernstates and in easternspecies to 3 and 18 westernspecies to 7. thesouthern Appalachian foothills. Thermopsis vil- St. John(1941) respondedto Larisey's revision losa and T. fraxinifoliaare nativeto Tennessee, withsome notes on nomenclatureand morphology. Georgia, and the Carolinas where they occur in Though variouslytreated in regionalfloras, the woodlandsat elevationsof 700-1600 m. Also in mostsignificant work on T7hermopsissince its re- the east is T. mollis,which is foundin the same vision has been Isely's (1981) treatmentin Le- statesas T. villosa and T. fraxinifoliaas well as guminosae of the UnitedStates. Isely drastically in Virginia,but at lower elevations,typically be- reducedthe number of speciesin Thermopsis,pri- tween300 and 800 m. marilyby loweringthese to varietal status. He In westernNorth America, Thermopsis occurs perceivedvariation in thewestern taxa as tootrivial both in the Rocky Mountainand intermountain to supportthe specificrank. Furthermore,he saw regionsas well as in the Pacific coastal states. that "all taxa are seeminglyconfluent with those ThermopsisMontana is themost widely distributed geographicallycontiguous." His views were not species; it is foundas two intergrading,regional universallyaccepted at the time (Turner,1982), varietiesin the intermountainregion from 300 to and our findingssupport recognition of most of 3200 m. Morphologyand habitat are variable these taxa as separate species. throughoutits range fromMontana southwardto T7hermopsisis the largestand mostwidespread Arizona and New Mexico and northwestwardto memberof the small tribeThermopsideae, which Wyoming,Idaho, Oregon,and Washington.Found consistsof about 46 species in 6 genera (Turner, at relativelylow elevations,1000-2000 m, is T. 1981). All of these are restrictedto temperate rhombifolia.Although this taxon can be found regionsof the NorthernHemisphere. The tribeis fromAlberta and Saskatchewanto New Mexico, characterizedby ten free stamensand petiolate, it is largelyrestricted to alluvialsoils in drygrass- trifoliateleaves. Though most of the genera are lands. Thermopsisdivaricarpa occurs at middle woody,the two herbaceousgenera, Baptisia and to high elevations,2000-3000 m, in the Rocky T7hermopsis,account for most of the speciesin the Mountains from Wyoming to New Mexico, in Thermopsideae.These two genera are unitedby mesophyticconifer forests. many other morphologicalas well as secondary Most of the Thermopsisin the Pacific coastal chemical characteristics.In fact, Larisey (1940) statesis foundin California.In centralCalifornia, referredto Thermopsisas a westerngeographical T. californicavar. californicais commonin chap- supplementof Baptisia, thoughit seems more arral west of the Sierra Nevadas up to 1000 m likelythat one or several lines of Baptisia was elevation.In northwesternCalifornia and southern derivedfrom a Thermopsis-likeancestor (Turner, Oregon,T. gracilis is dominantand occurs at low 1967). In fact,Cranmer & Turner(1967) offered elevations,to 1200 m, in mesophyticforests. The Baptisia and Thermopsisas possibleancestors to remainingtaxa in the extremewest have restricted the Genisteaeon the basis of morphology,alkaloid distributionswithin California. Thermopsis califor- chemistry,and geography.Since then,the woody nica var. argentata has a disjunctrange in north- Thermopsideaehave been favoredas closerelatives easternmostand southwesternCalifornia; this tax- to the Genisteae(Polhill, 1976; Turner,1981). on grows at
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