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

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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.

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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 high elevations,1200-2200 m, in montanechaparral and pine forests.Thermopsis macrophyllais knownonly from the westernside BIOGEOGRAPHY of Santa Ynez Peak in Santa BarbaraCounty. Our T7hermopsisoccurs not onlyin NorthAmerica; studyof typematerial indicates that this name has 13 species are foundin Asia. The Asian species been erroneouslyapplied to the more commonT. occur at elevationsof 1500-4000 m fromIndia californicavar. californica.Unlike the latter,T. northwardto eastern Kazakhstan and southern macrophyllais highlylocalized, relatively uniform, Russia,eastward across China,Mongolia, and Rus- and strikinglyrobust. Equally impressivein habit sia, and beyondto Japan,far easternRussia, and is Thermopsisrobusta, which is found only in

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TABLE 1. Chromosomenumbers in Thermopsis.Counts listed with voucher only are originalmeiotic counts by the juniorauthor (T); voucherson depositat TEX.

2n number Reference/voucher Asian species T alternifloraRegel 18 Federov(1969) T barbata Benth.ex Royle 18 Chen et al. (1992) T chinensisBenth. ex S. Moore 18 Yeh et al. (1986) T chinensisBenth. ex S. Moore 18 Chen et al. (1992) T lanceolata R. Br. 18 Mesicek & Sojak (1969) T. lanceolata R. Br. 18 Chen et al. (1992) T licentiana E. Peter 18 Chen et al. (1992) T. lupinoides(L.) Link [as T fabacea (Pallas) DC.] 18 Federov(1969) T lupinoides(L.) Link 18 Probatova& Sokolovskaya(1981) T mongolica Czefr. 18 Mesicek & Sojak (1969) T przewalskii Czefr. 18 Chen et al. (1992) Americanspecies T californicavar. argentata 18 California: Modoc Co., T 5813, T 5816 T divaricarpa 36 Colorado: JeffersonCo., T 5636 T divaricarpa 36 Colorado: Las AnimasCo., T 5772 T. divaricarpa 36 Colorado: Saguache Co., Tomb649 T gracilis 36 Oregon: Lane Co., T 5797 T macrophylla 18 California: Santa Barbara Co., T 5820 T montana(var. montana?) 18 Federov(1969) T montana(var. montana?) 18 Colorado: SummitCo., Gillett(1975) T. montanavar. montana 18 Colorado: Saguache Co., T 5775 T montanavar. montana 18 Idaho: OwyheeCo., T 5798 T montanavar. montana 18 Nevada: Elko Co., T 5795, T 5796 T montanavar. montana 18 Utah: Wasatch Co., T 5794 T rhombifolia 18 CANADA: Saskatchewan, Ledingham(1957) T. rhombifolia 18 CANADA: Saskatchewan, Taylor(1967) T rhombifolia 18 CANADA: Saskatchewan, Federov(1969) T rhombifolia 18 CANADA: Manitoba, L6ve & Love (1982) T rhombifolia 18 Colorado: HuerfanoCo., T 5773 T villosa 18 Zhukova(1967)

Humboldtand SiskyouCounties of northwestern centralColorado. Polyploidyis believed to have Californiaalong ridgetopsto 1500 m. Finally, arisen independentlyin the two taxa, largelybe- Thermopsiscalifornica var. semota is restricted cause of theirgeographical isolation from one an- to the woodlandsof San Diego County. other.Relatively few counts have been recorded forthese two taxa, however,and it remainsto be CHROMOSOME NUMBERS 'seen if such countswill hold throughouttheir re- spectivedistributions. The firstchromosome counts for Thermopsis A base chromosomenumber of x = 9 is also were obtainedby Russianworkers in 1931 (Fede- foundin the closelyrelated Baptisia, a genus of rov, 1969). These were obtainedfor the Asian about 17 speciesthat is confinedto lowerelevations species,T. alternifiora,and the Americanspecies, ofthe southeasternUnited States (see Isely, 1981, T. montana; both were found to have somatic for overview).To date, no polyploidshave been countsof 2n = 18. Subsequentcounts for all of reportedfor Baptisia. the species investigatedhave also been on a base of x = 9 (Table 1). SECONDARY CHEMISTRY (MICROMOLECULES) Polyploids(2n = 36) are knownin only two species,T. gracilis and T. divaricarpa,both Amer- Unlikein Baptisia, flavonoidsand alkaloidshave ican, the formeroccurring in northernCalifornia been of limitedusefulness in characterizinginter- and adjacent Oregon,the latteroccurring in the specificvariation in Thermopsis.On the basis of frontrange of the Rocky Mountains,mostly in quinolizidinealkaloid distribution, Salatino & Gott-

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lieb (1980) were able to divide Thermopsisinto do overlap,however, and can be called trendsat twophyletic groups, the Asian and theNorth Amer- best. ican species. A comprehensivephylogeny, how- Vegetativecharacters are oftenquantitative with ever,has not yetbeen proposedsince alkaloidand much variationaround average proportions.Nev- flavonoiddata are insufficient(Turner, 1967; De- ertheless,certain characters have wide diagnostic ment & Mabry, 1975) and contradictory(Balan- applicationwithin the genus.Plant height,leaf size drinet al., 1982). and shape,and pubescence have someutility, though Secondarychemical data have been used, none- thereis overlapbetween taxa, especiallyvarieties. theless,to place the genus withina phylogenetic Althoughthe leaf apices may differon a single reconstructionof a largerscale. Based on quinoli- plant,the numberof leaf veins and theirvisibility zidine alkaloid distributions(Salatino & Gottlieb, are sometimesuseful. The stipulesare oftwo types: 1980), mostof the Thermopsideaeprobably arose the lowerstipules in all taxa are scarious and do from"Sophora-like taxa" (Turner, 1981). Fur- not subtendleaves, the upper are foliaceousand thermore,based on serologicalcross reactivityof blade-bearing.The foliaceousstipules differ diag- seed proteins(Cristofolini, 1987), general mor- nosticallyin shape, rangingfrom lanceolate to phology(Polhill, 1976), and phytochemistry(Fau- broadlyovate, and in size, growingto 9 cm long geras & Paris, 1968; Harborne, 1969; Salatino and 6 cm wide.Inflorescence bracts, likewise, vary & Gottlieb,1983), thewoody Thermopsideae gave in shape and size and also in persistence.Racemes rise in turnto the Genisteae(Bisby, 1981; Cris- may bear shortor longpedicels, and thesemay be tofolini& Feoli-Chiapella,1984; Polhill, 1976; pubescentor not. In addition,the flowersmay be Turner,1981). scatteredalong the axis or may be insertedin whorls,even wherethe flowersare not especially dense. Withinthe flowers,calyx proportionsmay MORPHOLOGY confirmidentity but are notespecially useful alone. Thermopsishas been commonlyreferred to as Most Thermopsis species propagate by rhi- "falselupine," "buckbean," "goldenpea," "golden zomes, sometimes,as in T mollis and T. rhom- banner," and "bush pea." Thoughthe flowerhas bifolia,forming extensive clonal populations.Oth- inspiredseveral of these names, it varies little within erwise,the plants are supportedby thick,perennial the genus and is not especiallyuseful for specific rootstocks.It is oftenimpossible to confirmthe delimitation.There are slightvariations in flower presenceof rhizomeson herbariumspecimens, es- size and petal shape, whichoften appear constant peciallyof large plants,which typically show just withina taxon,but these are so minoras to be of enoughof the apical structuresto fiton a herbarium limitedusefulness for identification purposes. sheet. Yet fieldobservations indicate that a given Fruitcharacters are used to distinguishbetween taxon will have a tendencyto formrhizomes or Thermopsisand Baptisia; compressedpods are not,and thischaracter should be preservedduring mostlyfound in theformer and inflatedin thelatter. collectionand noted foridentification. The postureand shape of the maturepods vary For stems,branches, and fruits,the angle of diagnosticallywithin Thermopsis as well.The fruit insertionis describedas erect, ascending,diver- may be erect, ascending,or divergentfrom the gent,or spreading.Distinctions between these terms raceme axis. There seems to be no geographical may be confusing.In the followingdescriptions, correlationto pod posture;T. californicaand T. "erect" indicatesvertical posture or nearlyso and macrophyllaof California,T. montana of the in- littleor no curvatureat the pointof attachment termountainregion, and T. villosa of the Appa- while"ascending" indicates an angleof attachment lachians all have erect or ascending fruits,and up to 500 fromvertical often with upward curvature irregularlyto stronglydivergent pods are foundin at the pointof attachment.Though "spreading" the remainingspecies. Likewise,pod shape varies and "divergent"are roughlysynonymous, here among species of the same region,though pods "spreading"is used forbranches and "divergent" thatare stronglydivergent will also be arcuate or is used forfruits. annular.Isely (1981) noted that ovule numberis characteristicwithin regions of the genericdistri- SPECIES AND VARIETAL CONCEPTS bution.Indeed, we foundthat in southernCalifornia Thermopsistypically has 5-9 ovulesand in north- As is discussedabove, the primarydifference in ern California8-14 ovules. The Rocky Mountain the varioustreatments of Thermopsishas been in and intermountainspecies have 10-18 ovulesand the rankof the taxa. In thistreatment, the specific theAppalachian species have 12-16. These figures rankhas been chosento expresswhat we perceive

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to be evolutionarilyindependent taxa. We have sent; pedicels 2-17 mm long. Flowerspapiliona- inferredthis frommorphological evidence of re- ceous; calyxcampanulate, appearing 4-lobed, sub- productiveindependence both from herbarium equal, the upperlobe double,broader, and slightly specimensand observationsof populationsin the longerthan the others,truncate to emarginate,the field.For example,although occasional sympatric lower lobes deltoidto lanceolate; corolla yellow; hybridizationis suspectedbetween members of T. bannervery widely ovate, shorterthan wingsand divaricarpa and both T. montana and T. rhom- keel, cleft(emarginate and shortmucronate), claw bifolia, there is no evidence that these hybrids cuneateto oblong,replicate-reflexed; wings usually have stabilizedand persisted.Nevertheless, Isely asymmetric,elliptic to oblong-rhombic,auriculate, occasionally,if not often, saw intergradation,where ? lengthof keel, not adnate to keel, claw narrowly we, like Larisey,see none. We attributethe res- oblong;keel asymmetricallyoblong-elliptic to car- olutionof our perceptionto the extensivefield and inate,auriculate, posteriorly fused, without a beak, herbariumwork that is the basis of thisstudy. claw narrowlyoblong; stamens 10, free,uniform; anthersdorsifixed; pistil short-stipitate; ovary oblong,usually longer than style,villosulous; ovules TAXONOMICTREATMENT 5-15; styleglabrous; stigma minute. Legumes persistent,erect, ascending, or divergent,tan to brown, III. 3. Thermopsis R. Br., HortusKew., ed. 2, narrowlyelliptic, laterally compressed, straight, ar- L. = Ther- 1811. TYPE:Sophora 1upinoides cuate, or rarelyannular, margins straight or, as a = mopsis lanceolata R. Br. ThermopsislN- consequenceof ovule abortion,sinuate or lomen- Link. pinoides (L.) taceous; pericarp chartaceous,not elastic, vari- ThermiaNutt., Gen. N. Amer.P1. 1: 282. 1818. TYPE: ouslypubescent; stipe 2-4 mm long; fruitsslowly Thermiarhombifolia (Pursh) Nutt. dehiscent,valves separatingfrom apex through Scolobus Raf., Amer.Monthly Mag. Crit.Rev. 4: 193. both sutures. Seeds (1-)3-10 per fruit,tan to 1819. TYPE: Scolobus rhombifolius(Nutt. ex Pursh) Raf. brown-black,oblong to reniform,often with a mi- Drepilia Raf.,N. Fl. 2: 52. 1836 [1837]. TYPE: Drepilia nute,membranaceous rim-aril. Base chromosome rhombifolia(Nutt. ex Pursh) Raf. number, x = 9. Perennialherbs, usually rhizomatous; stems erect Diagnosis and relationships. Thermopsisis or ascending, few-branchedto well-branched, a memberof the ThermopsideaeYakovlev (Bot. branchesascending or spreading;plants glabrate Zurn. 57: 585-595, 1972), by virtueof its free to variouslypubescent, unarmed. Leaves petiolate, stamensand palmatelytrifoliate, petiolate leaves. trifoliolate,not glandular-punctate; stipules usually AlthoughThermopsis is looselyallied with all other persistent,foliaceous or nearlyso, dimorphic,the Thermopsideae,it is very similarmorphologically lowermostamplexicaul, scarious, and not blade- to Baptisia, the otherprominent North American bearing,the rest broadlyovate to lanceolate and memberof the tribe.Fruiting Thermopsis can be subtendingleaves, to 6(-9) cm long,to 3(-6) cm distinguishedreadily from Baptisia. The pods of wide, smallerand narroweron branches;leaflets NorthAmerican Thermopsis are flattened,while subpetiolulate,narrowly elliptic to obovate, 2- those of Baptisia are inflated. 8(- 10) cm long,margins entire; estipellate. Inflo- As detailedabove, Thermopsisis foundin Asia rescences terminal(lateral) racemes, commonly and NorthAmerica. In NorthAmerica four species solitary,flowers few to numerous,scattered or -occurin the Pacific coastal states,three occur in whorled;bracts deciduous or persistent,widely ovate the Rocky Mountainsand intermountainregions, to lanceolate,acute to acuminate;bracteoles ab- and threeare foundin the southernAppalachians.

KEY TO SPECIES 1a. Plants of southernAppalachian foothills. 2a. Stipuleson primaryaxis more than 1 cm wide; pedicels less than 3 mm long; pods erect, densely villous...... 10. Termo 2b. Stipuleson primaryaxis less than 1 cm wide;pedicels more than 3 mmlong; pods irregularlydivergent, pubescentor glabrate. 3a. Plantsrhizomatous; pedicels villosulous and shorterthan bracts; racemes terminal; mostly piedmont, 300-800 m ...... 6. Thermopsismollis 3b. Plants froma singlewoody rootstock; pedicels glabrate and at least as long as bracts;racemes terminalor lateral;montane, usually 1000-2000 m...... 3. Thermopsis fraxinifolia lb. Plants of Rocky Mountains,intermountain regions, and Pacificcoastal states. 4a. Plantstomentose or sericeousthroughout; flowers usually in whorlsof 3-5; California.

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5a. Plants mostly3-8 dm tall; leafletsmostly 3-7 cm long; pods ascending,usually with several or more abortiveovules ...... -...... -...... -...... 1. Thermopsis californica 5b. Plantsmostly 10-20 dm tall; leafletsup to 11 cm long;pods ascendingor divergent,usually with well-developedovules. 6a. Pods ascendingand straight;pedicels 2.5-4 mm long; lateral leaf veins 6-8 pairs; Santa Ynez Mts. of California....-...... -...... 5. Thermopsismacrophylla 6b. Pods divergentand arcuate; pedicels4-8 mm long; lateralleaf veins 9-12 pairs; Humboldt and SiskiyouCounties of northwesternCalifornia ...... 9. Thermopsisrobusta 4b. Plantsglabrate to pubescent;flowers usually scattered; Pacific coastal mountains north of 40'N latitude, and more centralportions of the westernUnited States and adjacent Canada. 7a. Pods erect and straight...... -..-..7. Thermopsismontana 7b. Pods divergentand straight,arcuate, or annular. 8a. Branchesspreading, strongly zigzagged; leaflets more than 2 cm wide,sometimes emarginate; petiolesmore than 2.5 cm long; pods straight;western Oregon and northernCalifornia ...... -...... -....-.-.-4. Thermopsis gracilis 8b. Branches ascending,weakly to moderatelyzigzagged; leaflets less than 2 cm wide, not emarginate;petioles less than 2.5 cm long; pods usuallyarcuate or annular;northern and easternRocky Mountainsand adjacent Great Plains. 9a. Plants3-9 dm tall;leaflets more than 3 cm long,lateral veins 7-10 pairs;stipules ovate to narrowlyelliptic; flowers 2.5-3.0 cm long; pods straightto arcuate ...... -...... 2. Thermopsis divaricarpa 9b. Plantsmostly 1-3 dm tall; leafletsless than 3 cm long,lateral veins 5-7 pairs; stipules usuallywidely ovate; flowers1.6-2.1 cm long; pods stronglyarcuate to annular...... -...... -...... -8. Thermopsis rhombifolia

1. Thermopsis californica S. Watson, Proc. KEY TO VARIETIES OF THERMOPSIS CALIFORNICA Amer.Acad. Arts11: 126. 1876. TYPE: U.S.A. la. Stipuleswidely ovate to ovate,to 5.0 cm wide; California:Marin Co., Corte Madera, J. M. bractsto 15 mmwide; peduncles usually more Bigelow 1854 (lectotype,selected here, NY; than 6 cm long; pod pubescence of matted probableisolectotypes, NY, PH, US). trichomes .-.-...... -.. lb. Thermopsiscalifornica var. californica Plants robust,(2.5)3-8 dm tall, sericeous or lb. Stipulesovate to lanceolate,to 2.5 cm wide; pedunclesusually less tomentose;stems slender,erect, clustered,well- bracts to 6 mm wide; than 6 mm long; pod pubescenceof straight, branched, arising from rhizomes;branches as- appressedtrichomes. cendingat 20-45?, weaklyto stronglyzigzagged. 2a. Silverysericeous throughout; wing and keel Leafletselliptic or obovate,3-7(-9) cm long, 1.2- petals reniform;northeastern and south- 4(-4.5) cm wide,obtuse or acute, silverysericeous westernCalifornia ...... - la. Thermopsiscalifornica var. argentata or tomentoseto villous,lateral veins 6-9 pairs, 2b. Villousto denselytomentose; wing and keel conspicuouslynet-veined beneath; petioles 1-3.5 petalsoblong-elliptic; restricted to San Di- cm long; stipulespersistent, widely ovate to lan- ego Co., California...... ceolate, 1.5-6.5 cm long,0.7-5 cm wide,cuneate i.. c. Thermopsiscalifornica var. semota to amplexicaul,acuminate. Racemes 7-30(-35) cm long,with 10-40 flowersusually in whorlsof la. Thermopsis californica var. argentata 3-5; peduncles(2-)4-10 cm long; pedicels 2-6 (E. Greene) C. J. Chen & B. Turner,comb. mm long,sericeous or tomentose;bracts semiper- nov. Basionym: Thermopsis argentata E. sistent,ovate to lanceolate,6-15 mm long, 2.5- Greene, Erythea 3: 18. 1895. Thermopsis 15(-20) mmwide. Flowers 1.5-2.0 cm long;calyx gracilis var. argentata (E. Greene) Jepson, 5-10 mm long, 5-9 mm wide at the limb,lobes Man. Fl. P1. Calif. 515. 1925. Thermopsis 2.5-5 mmlong, as long or longerthan tube; wing macrophylla var. argentata (E. Greene) Jep- and keel petals oblong-ellipticto nearlyreniform; son, Man. Fl. P1. Calif. 2: 245. 1936. TYPE: ovaryvelutinous; ovules 5-11. Pods straight,as- U.S.A. California:Modoc Co., "Forestdale," cending (irregularlydivergent when young), 2- Baker 1893 (lectotype,selected here, ND; 6(-7) cm long,(0.5-)0.6-0.9 cm wide,sericeous, isolectotypes,CAS, ND, NY, US). tomentose,or velutinous.Seeds 1-6(-10), oblong In hisprotologue Greene cited a singlecollection witha shortrostrum, 4-5.5 mm long, 2.5-4 mm fromModoc County,without locality, made by Milo wide. S. Baker in 1893. Examinationof materialat ND Three varietiesare recognizedwithin Thermop- revealstwo sheetscollected by Baker in the "Big sis californicaas follows: Valley Mts." during June-August1893. Both

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sheets, which bear printedlabels, were subsequent- californica in its smaller habit and closely ap- ly modified by hand-inkings so as to read August pressed, silvery vestiture. 1894. One sheet is clearly a mixture of plants We are unable to distinguishbetween the north- collected in June (flowering sprig) and fall (late ern and southern populations (Fig. 1). Isely (1981) fruitingsprig). The other sheet is in late fruit.We applied the varietal name argentata (as part of his have designated the latter as lectotype. T. macrophylla) only to the more northern populations and included the southern populations in Plants (2.5-)3-5 dm tall, silvery sericeous his T. macrophylla var. macrophylla. We agree throughout; branches weakly to moderately zig- with the assessment of Abrams (1944) that these zagged. Leaflets obovate to elliptic, 3-6.5 cm long, disjunct interiorpopulations represent a single tax- 1.2-3.5 cm wide, obtuse to acute sometimes with on. At least we can find no characters by which a short mucro; petioles 1.3-2.1 cm long; stipules to distinguishthem though both are readily distin- lanceolate to ovate, 1.5-6.0 cm long, 0.8-2.5 cm guished from varieties californica and semota by wide, cuneate to cordate. Racemes 8-20(-25) cm habit, vestiture, and petal shape. long, with 10-30 flowers;peduncles 4-7(-10) cm long; bracts ovate to lanceolate, 7-15 mm long, 2.5-5 mm wide. Calyx 7-9(-10) mm long, 5-8 lb. Thermopsis californica var. californi- mm wide at the limb, lobes 3.5-4.5(-5) mm long; ca. banner cleft; wing and keel petals asymmetrically Thermopsiscalifornica var. velutinaE. Greene,Erythea to ovules 8-11. Pods 3- oblong-elliptic reniform; 1: 81. 1893. Thermopsisvelutina (E. Greene) E. 6(-7) cm long, (0.5-)0.7-0.9 cm wide. Seeds 1- Greene,Erythea 3: 19. 1895. Thermopsisgracilis 6, oblong, 4-5(-5.5) mm long, (2.5-)3.5-4 mm var. velutina(E. Greene)Jepson, Man. Fl. PI. Calif. wide. Chromosome number, 2n = 18. 515. 1925. Thermopsismacrophylla var. velutina (E. Greene)Larisey, Ann. MissouriBot. Gard. 27: Distribution and ecology (Fig. 1). North- 256. 1940. TYPE: U.S.A. California:Santa Clara Co., MountHamilton, Greene 1891 (lectotype,se- easternmost California (southeastern Siskiyou Co. lectedhere, ND; isolectotype,ND). and closely adjacent Shasta, Modoc, and Lassen cos.) and again in southwestern California (Kern, In his protologue Watson did not cite specimens, Ventura, Santa Barbara, and Los Angeles cos.), but rather referred to the species as occurring in mostly interiormontane chaparral and pine forests "Marin and Napa Counties, California, and prob- from 1200 to 2200 m; floweringMay-June. ably southward." He did say that his Thermopsis californica was the same as what Torrey had re- Representativespecimens examined. U.S.A. Cal- ferredto as "T macrophylla" (Torrey, 1857) and ifornia: Kern Co., Mt. Pinos, June 1905, Hall 6378 T "fabacea" (Torrey, 1859). Watson, in his pro- OwensValley and at FortTejon, Horn 1863 (NY). (UC); tologue, indirectly suggested a lectotype with his Lassen Co., ridgeW of Nubieber,along CaliforniaHwy. 299, 26 June 1941, Heller 16229 (MO, NY, WTU). citation of the Pacific Railroad Report in which Los AngelesCo., Santa Barbara,Liebre Ridge, N slope Torrey cited a Bigelow collection from "Corte Ma- Liebre Mt., 15 Oct. 1923, Hunt 5 (CAS, RM). Modoc dera," which is located in Marin County, California. Co., Forestdale,Baker 1894 (NY, PH, POM, RM, US); We have selected the Bigelow collectionfrom among Forestdale,Baker 1899 (PH, UC); Forestdale,Baker & since the Herbarium Nutting 1894 (CAS, GH, NY, RM, UC); 3 mi. NW of contending lectotypes Torrey Lookout,3 June1968, Turner5813 (TEX); 14 mi. NW collection at NY has the name Thermopsis cali- of Lookout,3 June 1968, Turner5816 (TEX). Santa fornica Watson writtenon it, presumably in Wat- Barbara Co., San Rafael Mt., Ford 1887 (GH); Santa son's hand. Duplicates of the Bigelow collections Ynez, Big Pine Mt., 7 Aug. 1936, Peterson303 (UC). were not found at GH, where Watson worked, nor Shasta Co., ca. 20 mi. W of Fall River Mills, 24 May 1940, Hitchcock 6533 (NY, POM, UC, WTU); 1 mi. were other contending lectotype collections anno- W of Dana, 2 June1968, Turner5809 (TEX). Siskiyou tated by him as belonging to Thermopsis califor- Co., just E of Barnum Flat Reservoir,3 June 1968, nica. Turner5817 (TEX). VenturaCo., Fort Tejon, 19 June 1935, Axelrod 434 (POM, UC); Mt. Pinos region,28 Plants 3-8(-9) dm tall, sparsely to densely to- June 1896, Dudley & Lamb 4745 (CAS, UC); Lakes mentose or villous; branches weakly to moderately Ridge, above Maricopa Road on Matilija-Sespedivide, zigzagged. Leaflets elliptic to obovate or rhombic, Pollard 1955 (CAS, COLO, TEX); 0.25 mi.NW ofReyes Peak, 23 Apr. 1934, Sowder 393 (UC). 3.5-7(-9) cm long, 2-4(-4.5) cm wide, obtuse to acute; petioles 1.5-3.5 cm long; stipules widely Thermopsis californica var. argentata differs ovate to narrowly ovate, 2.5-6.5 cm long, 1.5- from the more widespread, more variable variety 5.0 cm wide, cuneate to amplexicaul. Racemes

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o T. californicavar. argentata

l) T. californicavar. californica A 0*T. califoricavar. semota

*T. 2aii

A T. macrophylla

400 A T. robusta I410

FIGURE 1. Distributionof Thermopsisin California:T californicavar. argentata(small open circles); T californica var. californica(large open circles);T californicavar. semota(large closed circles); T gracilis (smallclosed circles); T. macrophylla(closed triangle);T robusta(open triangle).

10-30(-35) cm long,with 12-40 flowers;pedun- at the limb,lobes 2.5-5 mm long; banner cleft; cles (4-)6-10 cm long; bracts widelyovate to wingand keelpetals asymmetrically oblong-elliptic; nearlylanceolate, 6-15(-30) mmlong, 3-15(-20) ovules5-8. Pods 3-5.5 cm long,0.6-0.7 cm wide. mm wide. Calyx 5-8(-9) mm long,6-9 mm wide Seeds 3-5(-10), oblong,4 mmlong, 2.8 mmwide.

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Distribution and ecology (Fig. 1). Coastal his conceptof T. macrophyllavar. macrophylla, regionsof centralCalifornia from 35 to 400N lat- perceived"a cline towards[T. macrophylla]var. itude,mostly chaparral from near sea levelto about venosa [= T. gracilis in the presenttreatment] 1000 m; floweringMay-June. thatcharacteristically has spreadingand (compar- atively)narrow fruits." This statementwas quali- Representativespecimens examined. U.S.A. Cal- fied by the remarkthat "few specimenspossess ifornia:Alameda Co., hills E ofOakland, 1 Apr.1929, maturepods." Examinationof manydiverse spec- Tebbe62 (UC). Lake Co., 9 mi. fromLower Lake on imensfrom northern California shows that the geo- MonticelloRoad, 23 May 1937, Ewan 10352 (COLO, MO,NY); flatsnear Morgan Valley, 25 May 1920,Jep- graphicalhiatus shown by Isely in mappingthe son 9034 (JEPS).Marin Co., Sausalitoto Larkspur,27 twotaxa is real, and thatthe fewcollections of his Apr.1918, Abrams6882 (CAS,NY, US); San Rafael, T. macrophylla var. macrophylla that he maps dryhillsides, Bolander 1860-1867 (F, MO); AngelIs- withinthe range of T. gracilis (of the present San FranciscoBay, Chestnut1888 (F, UC, US); land, treatment)are mostlynonfruiting forms of T. grac- Lagunitas,Grinnell 1918 (CAS, MO, NY, RM, UC); Tiburon,17 Apr. 1938, Rose 38146 (GH, MO, NY, ilis that are otherwisesuperficially similar to T. WTU, UC). MendocinoCo., 5 mi.E ofPoint Arena, 29 macrophyllavar. macrophylla(sensu Isely). We May 1939, Rose 39184 (GH,MO, NY, UC, US); Little do not see any evidence of clinal intergradation Lake Valley2.5 mi. SE of Willits,Wiggins & Ferris betweenthese two taxa, nor do theyco-occur, as 10183 (CAS,RM, TEX, WS, WTU). MontereyCo., hill nearMonterey, 13 June1861, Brewer704 (UC, US). suggestedin Isely's treatment. NapaCo., Knoxville, 8 May 1903,Baker 3081 (F, GH, LL, MO, NY, UC); 5 mi.S ofCalistoga, 12 Apr.1924, ic. Thermopsis californica var. semota Heller13837 (CAS,F, MO, NY, US). San BenitoCo., (Jepson)C. J. Chen & B. Turner,comb. nov. Aromas,Hickman 1915 (CAS,GH, US). San LuisObispo Co., alongold roadto Pozo, 17 May 1928, Eastwood Basionym:Thermopsis macrophylla Hook. & 15140 (CAS,UC); San LuisObispo, 9 May 1882,Jones Arn. var. semota Jepson,Fl. Calif. 2: 245. 2660 (CAS,MO, NY, PH, RM, UC). San MateoCo., 1936. Thermopsismacrophylla Hook. & Arn. CrystalSprings Lake, May 1903, Elmer4824 (ARIZ, subsp. semota (Jepson)Beauchamp, Phyto- MO,NY, OSC, UC, US, WS); CrystalSprings Lake, CAS, 439. 1986. TYPE: U.S.A. California: 1 May 1902, Greene685 (CAS,GH, LL, MO,NY, UC, logia 59: US,WTU); Mt. Hamilton, June 1903, Elmer 5023 (ARIZ, San Diego Co., Spencer Valley, vicinityof CAS,MO, NY, UC, WS); SmithCreek, foot of Mt. Ham- Julian,Alderson 1898 (holotype,UC). ilton,10 May 1907, Heller8518 (CAS, GH, F, MO, NY, PH, UC, US, WTU); trailto Pine Ridge,N of Plants 3-5(-6) dm tall, denselysilvery tomen- MadroneSprings, 29 May1895, Smith 4075 (GH,OSC, RM,TEX). Santa Cruz Co., Glenwood, Davis 1907 (CAS, tose; branchesmoderately to stronglyzigzagged. GH, RM, UC, US); BearCreek Road about 3 mi.E of Leafletselliptic to narrowlyelliptic, 3-6.5(-7.5) BoulderCreek, 18 May 1946,Ferris 11130 (CAS,GH, cm long, 1.2-3.5(-3.8) cm wide,acute to obtuse RM,TEX, UT, WS). SonomaCo., betweenSanta Rosa witha shortmucro; petioles 1-2 cm long; stipules and Caliente,22 Apr.1902, Heller & Brown5343 Agua ovate to lanceolate,1.5-5.5 cm long,0.7-2.4 cm (CAS,COLO, GH, MO, NY, RM, US); 6.8 mi.E ofSanta Rosaon road to Calistoga, 30 Apr.1949, Wiggins12058 wide,cuneate to cordate.Racemes 7-20(-30) cm (CAS,UC, WTU). long, with10-30 flowers;peduncles 2-6(-9) cm long; bractsovate to nearlylanceolate, 6-11 mm This is the more commonlyencountered, more long, 3.5-6 mm wide. Calyx 5-8(-9) mm long, variabletaxon of the Thermopsiscalifornica com- 5-8 mm wide at the limb,lobes 3-5 mm long; plex. It is distinguishedfrom both T. californica bannershallowly emarginate and shortmucronate; var. semota and T. californicavar. argentata by wingand keelpetals asymmetrically oblong-elliptic; its largerhabit, broader stipules and bracts,and ovules 5-7. Pods 2-4 cm long,0.6-0.7 cm wide. sparselyto denselytomentose vestiture. Seeds 1-2 (fullymatured not seen). Brewer& Watson (1880) correctlynoted that Thermopsismacrophylla is distinctfrom T. cal- Distributionand ecology (Fig. 1). Interior ifornica.Nevertheless, most workers have treated portionsof San Diego Co., California,grassy oak these taxa as synonymouswith the earliername, woodlands,1000-1500 m; floweringApril-June. Thermopsismacrophylla. We view T. macrophylla, like Brewer& Watson, as a localizeden- Representativespecimens examined. U.S.A. Cal- demic thatis restrictedto the Santa Ynez Moun- ifornia:San DiegoCo., 1.7 mi.NW ofJulian, 5 Apr. Bacigalupiet al. 5799 (JEPS,RM, WTU); near tains near Santa Barbara, California.Collections 1957, Julian,Epling 1932 (CAS,NY, POM, UC); Cuyamaca of Thermopsiscalifornica are not sympatricand Reservoir,5 June1946, Fiker3576 (RM, UC, WS); do notintergrade with T. macrophylla.Isely (1981), Julian,Cuyamaca Mts., 28 May1899, Hall 1205 (CAS, who treatedT. californicavar. californicawithin NY, UC); vicinityof Santa Ysabel, Henshaw 1893 (US);

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Laguna Mts., 25 June 1924, Munz 8377 (GH, NY, mm wide. Flowers 2.5-2.9 cm long; calyx 8-11 POM); Pine Hills, 1 June1917, Spencer 504 (GH, NY, mm long, 6-8 mm wide at the limb, lobes 2-5 POM, UC); Ramona,head ofCedar Creek, 27 Mar. 1934, mm long, as long as or shorterthan tube; wing and Wieslander447 (UC). keel petals ellipticto asymmetricallyoblong-elliptic; Isely (1981) treated this taxon as a localized ovary appressed pubescent; ovules 10-18. Pods variety within his concept of Thermopsis macro- straight to arcuate to ca. 450, divergent, 6.5-9 phylla. We agree withhis circumscription.Larisey cm long, 0.6-0.8 cm wide, inconspicuously ap- (1940) inexplicably omitted the taxon from her pressed pubescent. Seeds (6-)10-14, brown-black, treatment. elliptic with a rostrum, 4-5 mm long, 2-2.5 mm wide. Chromosome number, 2n = 36.

2. Thermopsis divaricarpa Nelson,Bot. Gaz. Distribution and ecology (Fig. 2). South-cen- 25: 275. 1898. Thermopsis rhombifolia var. tral Wyoming (mainly Albany Co.) southeastward divaricarpa (Nelson) Isely, Brittonia30: 470. through the front range of the Rocky Mountains 1978. TYPE: U.S.A. Wyoming: Albany Co., to northeasternmostNew Mexico, mostly pine or Johnson's ranch, Big Laramie River, 8 Aug. spruce forests, usually along streams or in moist 1897, A. Nelson 3903 (lectotype, selected open areas, 2000-3000 m; flowering(April-)May- here, RM; isolectotypes, GH, NY [dated 7 June(-July). Aug. 1897], RM). Representativespecimens examined. U.S.A. Col- orado: Alamosa Co., near S entranceof Great Sand Thermopsispinetorum E. Greene,Pittonia 4: 138. 1900. Dunes NationalMonument, 18 May 1969, Dennam 2950 TYPE: U.S.A. Colorado:Saguache Co., below Mar- (TENN). BoulderCo., Boulder,21 Aug. 1884, Letterman shall Pass, Greene 1896 (lectotype,selected here, 141 (MO, US); plains and foothillsnear Boulder,July ND; isolectotype,ND). 1902, Tweedy5158 (NY, RM). ChaffeeCo., Salida, 18 May 1925, Nelson 10495 (NY); Salida, 7-8 Oct. 1918, Two differentcollections of Thermopsis divar- Eggleston 15019 (US). Clear Creek Co., Idaho Springs, icarpa, E. Nelson 3424 (Pole Creek, 22 July rockyhillsides, 15 May 1916, Clokey2676 (RM, UC); 1897) and A. Nelson 3903, were designated as ChicagoCreek, 11 July1942, Ehlers8398 (COLO, TEX); "type specimens" by A. Nelson, withoutindication Empire,2,576 m, 15-25 July1903, Tweedy5652 (NY, RM). CostillaCo., mountainnear La Veta Pass, 20 June Nelson of the holotype. Isely (1978) designated 1900, Rydberg & Vreeland 5935 (NY). Custer Co., 3903 as the "holotype"; he should have designated NorthCreek Campground,30 May 1954, Douglass 58 this as a lectotype. (COLO); Sangre de CristoCreek, 2 July1900, Rydberg In his protologue for Thermopsis pinetorum, & Vreeland5933 (NY, RM). DouglasCo., alongColorado Hwy. 67, 8.3 mi. N ofjunction of ColoradoHwy. 67 and Greene cited collections made by himself "in fruit, CountyRoad 126, 18 May 1987, Falb & Miercort113 below Marshall Pass," 4 Sep. 1896 and collections (RM). El Paso Co., ColoradoSprings, Jones 1879 (ARIZ, by C. F. Baker at Los Pinos (which is near Bayfield NY, US); near Palmer Lake, 21 June 1936, Rollins in La Plata Co.), 23 May 1899, in flower. The 1187 (GH, MO, ND, NY); Gardenof the Gods, Schallert latter collections appear to belong to Thermopsis 1943 (F, NY, PH, WS). FremontCo., near Hillside,4 June 1949, Gates 22053 (TEX). GarfieldCo., White as montana. The formeris selected the lectotype River,Grizzly Creek, 26 June1932, Ware24 (RM); 38.8 since mature fruitsbest serve to distinguishbetween mi. S of junctionColorado Hwy. 64 on ColoradoHwy. the taxa concerned. 139, 14 June 1971, Wofford1-77 (TENN). GilpinCo., near summitof Black Hawk Pass, Alston 1964 (TEX); Plants delicate, 3-6(-9) dm tall, glabrate or Tolland, Overholts1913 (MO, NY). Grand Co., Trail puberulent; stems slender, erect, clustered, mod- Ridge Road, 11.5 mi. N of GrandLake, 21 July1961, erately branched, arising from a thick woody root- Douglass 320 (TEX); Hot SulphurSprings, 30 May 1940, Pohl 1954 (PH). HuerfanoCo., 8 mi. SW of Cuchara, stock or rhizomes; branches ascending at 20-35?, Bear Lake Camp, 4 Aug. 1970, Stephens & Brooks moderately to strongly zigzagged. Leaflets elliptic 42675 (NY, UC); ErwindCanyon, W ofWalsenburg, 14 to narrowlyelliptic or rhombic, 3-7 cm long, 0.5- June1941, A. & R. A. Nelson 4595 (RM). JacksonCo., 2.5 cm wide, acute to obtuse, glabrate to puber- GouldRanger Station, 23 Sep. 1944, Ginter891 (RM). Vernon ulent, lateral veins 7-10 pairs, conspicuously net- JeffersonCo., Canyon,20 June1921, Bethel & Clokey4186 (CAS, COLO, F, MO, NY, PH, POM, RM, cm veined beneath; petioles 1-2.5 long; stipules UC, US); near Golden,Guy Hill, 2 June 1938, Ehlers persistent,ovate to narrowlyelliptic, 2-3.5(-4) cm 6850 (ARIZ, GH, NCU, ND, PH); ca. 11 mi. W of long, 0.7-3 cm wide, cuneate to cordate, acute. Littleton,20 May 1967, Turner5637 (CAS, RM, TEX). Racemes 7-15 cm long, with 5-15 flowers,usually Lake Co., Everett,7 May 1919, Clokey3588 (CAS, F, GH, MO, POM, RM, TEX, US). LarimerCo., Estes Park, in whorls of 2-4; peduncles 3-5 cm long; pedicels May 1895, Baker 2581 (COLO, MO, NCU, WS). Las 5-8 mm long, glabrate to puberulent; bracts de- AnimasCo., 11 mi.N ofStonewall, 26 May 1968, Turner ciduous, ovate to lanceolate, 7-14 mm long, 3-5 5772 (TEX). Mesa Co., 13 Aug. 1981, Siplivinsky1 961

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250

/400

350~~~~~~~~~~~~~~~

3500

300mi.

FIGURE 2. Distributionof Thermopsisdivaricarpa.

(COLO). Park Co., Pine Junctionto WandcrestPark, 27 POM, RM, UC, US); Mandel, 24 July 1903, Nelson June 1971, Kathleen & Shea 65 (MO); near Jefferson, 8910 (COLO, GH, MO, NY, UC); Fish Creek, 4 July MichiganCreek drainage, 16 July 1960, Walter 369 1917, Nelson9784 (GH, MO, RM, TENN, UC); Pole (COLO). PuebloCo., near Beulah,6 June1908, Robbins Mt. region,6 July1943, Porter 3245 (CAS, GH, MO, 4991 (COLO); W ofPueblo, 8 May 1908, Robbins4991 NY, UC, US, WTU); CrowCreek, Laramie Hills, 11 July (COLO). RouttCo., Elkhead Mts., 10 July1988, Flock 1935, Rollins 977 (GH, MO, NY, WS); Laramie River 2004 (COLO, RM); AnitaPeak, 3 Aug. 1903, Goodding near Laramie,16 June1935, Williams2214 (GH, MO, 779 (CAS, COLO, GH, MO, NY, PH, RM, UC, US). UC, US); nearCrow Creek, Laramie Mts., 20 June1935, Saguache Co., MarshallPass, 19 July1901, Baker 485 Williams 2236 (GH, MO, UC, WS). Carbon Co., Mc- (F, MO, NY, RM, UC, US); 8 mi. E of MarshallPass, -Fadden,in Medicine Bow Mts., 13 June 1972, Jones 27 May 1968, Turner5776 (TEX). SummitCo., near 1085 (MO). Breckenridge,Mackenzie 256 (MO, NY, RM, UC); near Bear River,30 June 1926, Payson 4833 (GH). Teller Isely (1981) treated this taxon withinhis broad Co., S side of Lower Twin Rocks Road, 10 June 1987, Barzee et al. 717 (COLO); 6 mi. NNW of Woodland concept of Thermopsis rhombifolia, noting that it Park, Devender& Kearns 1977 (ARIZ). New Mexico: stands morphologically and geographically some- Colfax Co., CimarronPass, 28 May 1934, McKelvey what between T. rhombifolia and T montana, 2405 (GH); Raton,10 June1887, Evans 59 (NY). Union which it does. Nevertheless, we have retained T. Co., SierraGrande, 19 June1911, Standley 6140 (US); 7 mi. S of Des Moines, 14 May 1974, Stephens 75588 divaricarpa as a distinct species. (NY). Wyoming: AlbanyCo., VedauwooHills E of Lar- We have found that T. rhombifolia, T. divar- amie, 13 June1940, Cain 65 (MO, TENN); LittleLar- icarpa, and T. montana are clearly distinct amie River,15 June1901, Goodding 10 (CAS, F, GH, throughoutmost of their ranges and do not appear MO, NY, POM, RM, UC, US, UT); Rock River,30 June to intergrade as do the varieties in the 1914, Macbride 2403 (RM, WS); Big LaramieRiver, 5 Thermopsis July1900, Nelson 271 (GH, NY, RM); Sand Creek, 2 montana complex. Indeed, in Colorado where the June 1900, Nelson 7031 (CAS, COLO, GH, MO, NY, several taxa are partiallysympatric, T. rhombifolia

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occurs mostlyat lower elevationsin deeper and served at the present time with the taxonomic drieralluvial soils, T. divaricarpa at mid-elevations treatmentadopted here, this being clearly depicted in mesophytichabitats, and T. montanaat higher, to thoseworking with living populations in thefield. subalpineelevations. Isely (1981) maps, but does not cite, at least It is true,however, that occasional intermediates fivecollections of Thermopsisdivaricarpa (as T. between Thermopsisdivaricarpa and T. rhom- Montana var. divaricarpa) fromwestern Mon- bifoliaoccur, but we viewsuch variation as derived tana, but we believethese to be misidentifiedele- fromrelatively isolated hybridization events. Such mentsof T. Montana var. Montana or T. rhom- "hybridized"populations occur in AlbanyCounty, bifolia.In addition,he mapsat leasta fewcollections Wyoming,and probablyoccasioned the description ofT. divaricarpa fromnortheastern Wyoming that of T. arenosa, whichappears to combinesome of we believe to be misidentifiedelements of fairly the featuresof T divaricarpa in the geneticback- typicalT rhombifolia. drop of T. rhombifolia.Likewise, in Colorado at Isely(1981) includedThermopsis pinetorum as MarshallPass along the Saguache-Chaffeecounty a synonymof his T. rhombifoliavar. Montanabut line,T. montanaand T. divaricarpa occurin close examinationof type material, as notedabove, shows proximityand therehas apparentlybeen somegene that the name apparentlywas typifiedby mixed exchangebetween the two.The variantsfrom this collections;the lectotypebelongs to his variety regionhave the generalhabit of T. divaricarpa, divaricarpa. but the ascendingpods approachthose of T. mon- Nelson,in his originaldescription of T. divari- tana. The formmay have precipitatedthe descrip- carpa, providedexcellent illustrations of the fruit- tionof T. pinetorum.The thirdauthor visited the ing branchesof T Montana, T divaricarpa, and generalarea of the typelocality of T pinetorum T rhombifolia,along witha seeminglyinterme- in May 1968 (Turner5776, TEX) and again in diate branch between T rhombifoliaand T di- July1991, and was impressedby the populational varicarpa thathe named T. arenosa. Althoughwe variationin thatvicinity, especially along the east- relegatethe lattername to synonymyunder "typ- ern slopesleading to MarshallPass. But such mor- ical" T. rhombifolia,in truth,T. arenosa is prob- phologicalaberrations are mere wrinklesin the ablya hybridor introgressedindividual between T broad distributionalfabric of T. rhombifolia,T. rhombifoliaand T. divaricarpa. divaricarpa, and T. montanaand do not make a case fortheir taxonomic submergence. If it were 3. Thermopsis fraxinifolia (Nutt. ex Torrey so, then many distinctspecies among the closely & A. Gray) M. Curtis,Amer. J. Sci. 44: 81. relatedgenus Baptisia wouldbe merged,for oc- 1843. Baptisiafraxinifolia Nutt.ex Torrey casional or even rampanthybridization is the rule & A. Gray,Fl. N. Amer.1: 387. 1840. Ther- when species of these taxa occur together(under mopsismollis var. fraxinifolia (Nutt. ex Tor- disturbance).Indeed, interspecific hybridization be- rey& A. Gray)Isely, Brittonia 30: 469. 1978. tweenthe closelyrelated taxa T. rhombifolia,T TYPE: U.S.A. NorthCarolina: Burke Co., Ta- divaricarpa, and T. montanais apparentlymuch ble Mountain("Catawba Ridge" of Nuttall; less commonthan among the more distantly related Graustein,1967), Nuttall1816 (holotype,PH; (amongthemselves) species, Baptisia leucophaea, possibleisotypes, GH, NY). B. sphaerocarpa, and B. leucantha (Alston & Turner,1963). Thereremains a problemwith the correct author It is indeedlikely, however, that T. divaricarpa citationfor Thermopsisfraxinifolia. This is sum- had an amphiploidorigin from T rhombifoliaand marizedby Wilbur(1963) who contendedthat the T. montana. So far as is known,T. divaricarpa correctcitation should be T fraxinifoliaM. Curtis is tetraploidwith 2n = 36 and has morphological ex A. Gray (publishedin 1848). Isely (1978) ac- featuressomewhat intermediate between the dip- ceptedthis, but he subsequentlytook up thecitation loid species T. rhombifoliaand T montana,both T fraxinifoliaNutt. ex Torrey& A. Gray(1840) with2n = 18. Perhaps the formationof amphi- (Isely, 1981). We believethis to be the preferred ploids was preceded by some gene exchange be- citationbecause Torrey & Gray (1840) cited a tweenT. rhombifoliaand T. montanaat thediploid Nuttallspecimen and quoteda descriptionprovided level. The presentranges suggestthat extensive by himin theirdiscussion of T. fraxinifolia.Con- introgressionbetween these taxa was not likely.It traryto Wilbur's observation,the Nuttallname shouldprove interestingto test these varioushy- was not rejectedby Torrey& Gray. They merely pothesesusing molecularcharacters. Regardless, noted that the species had not fallenunder their we believe that the taxonomiccommunity is best observationand thatit was "uncertainwhether it

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350

700

300

MILES 250 0 00o 200 300 400

50 9g0 8 80? 750 FIGURE 3. Distributioof TheXmopss L

FIGURE 3. Distributionof A7ermopsis ftaxinifolia.

[T fraxinifolia]be distinctfrom some of the spe- mm long, 2.5 mm wide. Chromosome number, 2n cies describedabove." We do not view this as a = 18. rejectionof the name, but rathera formof abey- Distribution and ecology (Fig. 3). Southern- ance. most Appalachian Mountains in eastern Tennessee, Plantsdelicate, 5-10 dmtall, glabrate to sparse- western North Carolina and adjacent Georgia and ly puberulent;stems slender, erect, clustered, well- South Carolina, rich woods, mostly upper hillslopes branched,arising from a singlewoody rootstock; along streams and at lower elevations along flood- branchesspreading at 45-80?, stronglyzigzagged. plains, (300-)700-1600 m; flowering(April-)May- Leafletselliptic, 4.5-8 cm long, 2-3.5 cm wide, July. acute to acuminate,glabrous to verysparsely puberulent,lateral veins 9-12 pairs, very conspic- Representativespecimens examined. U.S.A. Geor- gia: Fannin Co., NE of Cisco and NW of Blue Ridge, uouslynet-veined beneath; petioles 1.6-3 cm long; along Jack'sTrail and Jack's River,6 June1983, Coile stipulessometimes persistent, lanceolate to nar- et al. 3610 (RM, TENN). GilmerCo., near StanleyGap, rowlyovate, 1-3.5 cm long, 0.3-0.8 cm wide, 7 June1982, Coile et al. 2844 (POM, TEX). Habersham cuneate, acuminate.Racemes terminalor lateral, Co., Small 1895 (ARIZ, F, NY). Hall Co., near Gaines- ville, Huger 12-25 cm long,with 7-25 flowers, pe- 1897 (NY). MurrayCo., mountainslopes scattered; betweenCohutta and Fort Mountains,27 May 1935, duncles2.5-5.5 cm long;pedicels 7-17 mmlong, Duncan & Hardin 16468 (NCU). OglethorpeCo., ca. glabrate;bracts persistent,lanceolate, 8-12 mm 0.75 mi. S of the Georgia Hwy. 368 bridgeover the long, 1-3 mm wide. Flowers 1.6-1.9 cm long; SavannahRiver, 200 m,7 June1979, Credle863 (NCU); calyx 7-9 mm long, 4-5 mm wide at the limb, near bridgeof GeorgiaHwy. 72 on Savannah River,10 June 1978, Robert & Coile 1860 (NCU). Rabun Co., lobes 2-3 mmlong, much shorter than tube; wings Rabun Bald, 1-4 June1906, House 2266 (US); Tallulah elliptic to asymmetricallyoblong-elliptic; keel Falls, Small 1893 (F, NY). Towns Co., Young Harris, asymmetricallyoblong-elliptic; ovary appressed pu- 10 July1908, Howell 390 (MO, US). North Carolina: bescent;ovules (6-)12-16. Pods straight,irregu- BuncombeCo., Black Mountain,12 Aug. 1905, Biltmore larlydivergent, (3-)5.5-7.5 cm long,0.3-0.5 cm Herbarium 11214 (F, GH, MO, NY, US); Asheville, Kraus 1925 (MO, POM, UC). Burke Co., vicinityof wide, inconspicuouslypubescent. Seeds (5-)10- Table Rock Mt., 2 Aug. 1890, Heller 27 (F, MO, NY, 15, brown,reniform with a veryshort rostrum, 4 POM, UC). HendersonCo., Flat Rock, Gibbes 1854 &

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1858 (NY). Macon Co., 1.4 mi. N of the Georgiastate cent states, occasionally extending into the moun- line on NorthCarolina Route 28, 1 June 1975, Wood tains, where its range abuts that of T fraxinifolia. & Boufford1389 (A); 2.1 mi. N of theGeorgia state line Indeed, we do not know of any specific sites on NorthCarolina Route 28, 1 June 1975, Boufford where 16075 (MO). Polk Co., Pony Mt., Townsend1897 (US). the two have been collected together, nor do we TransylvaniaCo., WhitewaterRiver, just above Upper perceive a large set of intermediateplants in regions Falls, 7 June1951, Godfrey51298 (MO). Yancey Co., of near contact (or elsewhere) that might justify Mt. Mitchell,Rhoades 1932 (GH, PH, RM); E of Mt. their recognition as regional variants of a single Mitchell,exactly 2.3 mi. SW of RoughRidge Tunnel, 30 species, as June1970, Turner6076 (ARIZ, CAS, F, MO, NY, TEX). treated by Isely (1981). His distribution South Carolina: AbbevilleCo., 3 mi. SW of Calhoun maps of both taxa are fairly accurate, although Falls, near Savannah River, 13 May 1957, Radford these are difficultto evaluate since he did not cite 22768 (NCU, UC). GreenvilleCo., Caesars Head, Smith specimens documenting the distributionsshown. 1881 (F, GH, MO, NY, US). Oconee Co., Boynton1889 Isely (1981) did map a number of specimens (CAS). Pickens Co., 0.7 mi. offU.S. Hwy. 178 on Sas- safras Mt. Road, 30 June 1970, Turner6075 (ARIZ, from easternmost Tennessee that appear to be CAS, COLO, F, GH, MO, NCU, NY, RM, TEX, WTU). anomalous. These are morphologically somewhere Tennessee: BlountCo., ChilhoweeMountain, 10.98 km between Thermopsis mollis and T fraxinifolia and SW of Tennessee Hwy. 73 on FoothillsParkway, 650 were designated "T mollis var.?" on Isely's map. m, 26 May 1982, Boufford& Spongberg 22881 (A, MO, NCU, POM, TEX). GreeneCo., 2.3 road mi. S of We believe that many of these may be derivatives Camp Creek School,just N of HorsehitchGap, 10 June of hybridization between T. mollis and T fraxi- 1980, Boom & Smith547 (TENN); alongKennedy Cabin nifolia. The species occur in close proximity in 11 loop road, June1980, Boom & Perkins554 (TENN); this region and the intermediateshave the habit of mountainslope between Camp Creekand Paint Creekon T mollis but the inflorescence pattern and general GreenMt., 30 July1950, Shanks& Sharp 7455 (TENN). HamiltonCo., LookoutMt., Canby 1869 (F, GH, MO, habitat preference of T fraxinifolia. Both T mollis NCU, NY, PH, US). MonroeCo., Waucheesi Mt. on FR and T fraxinifolia have been collected on Lookout 126-C, 12 June1978, Whitten& Mescall 213 (TENN). Mountain, Hamilton Co., Tennessee, as have sev- Polk Co., Peavine Road, 5.8 road mi. SW of Ocoee No. eral of the putative hybrid intermediates (e.g., 3 Powerhouse,23 May 1980, Boom 471 (TENN); Wolf Churchill Ridge,25 June1983, Z. E. & J.B. Murrell336 (TENN). 1911, TENN; Canby 1869, GH, MO, Cultivated: Massachusetts,Botanical Garden of Cam- NCU, NY, PH, US). bridge,Gray s.n. (NY). More detailed fieldinvestigations may show that the hypothetical hybrids may, in fact, be part of This species is partially sympatric with Ther- a more widespread, highly variable taxon derived mopsis mollis, but in regions of overlap T fraxi- from ancestral hybridizationbetween Thermopsis nifolia usually occurs at higher elevations along mollis and T fraxinifolia. This is inferred from the upper slopes of the mountainous ridges and recent collections by Boufford & Spongberg from hills of the Appalachians. Thermopsis fraxinifolia Blount Co., Tennessee (22881, GH, MO, NCU, may also occur at lower elevations on alluvial de- TEX), and by Whitten from Monroe Co., Tennes- posits along at least some of the riversin this region. see (213, TENN), which seem to reveal stabilized Thermopsis fraxinifolia can be distinguished populations of intermediatemorphology in this area. from T. mollis by its more robust, more branching Some of the characters in at least a few of these habit, lateral (as well as terminal) inflorescences, plants are unique within the American taxa of and longer pedicels. Furthermore, Thermopsis Thermopsis. For example, the Blount County spec- mollis propagates by extensive, vigorous rhizomes, imens cited above have short, squat pods with which are absent in T fraxinifolia. Both species distinct gynophores, much as occurs in some of were well known to Asa Gray, who personally col- the Asian species. On the present information,how- lected T fraxinifolia at its type locality and sub- ever, it is more sensible to include such plants sequently grew this at the botanical gardens at within a variable T fraxinifolia than to accord Harvard. According to Gray (1848b), T mollis these varietal status. was also maintained in the Harvard Botanical Gar- dens at the same time. Living specimens of both 4. Thermopsis gracilis Howell, Erythea 1: 109. were used to make the excellent illustrationsof the 1893. TYPE: U.S.A. Oregon: Josephine Co., two species that accompany his treatmentof Ther- "Mountains near Waldo," 5 July 1887, T J. mopsis of the eastern United States. Howell 669 (holotype, ORE; isotypes, CAS, As noted by Gray (1848b), Wilbur (1963), and GH). Radford et al. (1968), Thermopsis mollis is mostly Thermopsisvenosa Eastw., Bull. TorreyBot. Club 32: a piedmont species, distributed along the middle 198. 1905. Thermopsisgracilis var. venosa(Eastw.) and western portions of North Carolina and adja- Jepson,Man. Fl. P1. Calif.515. 1925. T7ermopsis

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montanavar. venosa (Eastw.) Jepson,Fl. Calif. 2: 245. 1936. Thermopsismacrophylla var. venosa~~1200 ~~~~I25o (Eastw.)Isely, Brittonia 30: 469. 1978. TYPE: U.S.A. California:Shasta (or Trinity)Co., "LewistonTrail near the summitthat dividesTrinity from Shasta County,California," Eastwood 1901 (holotype,CAS). Thermopsissubglabra L. Henderson,Rhodora 32: 26. 1930. TYPE: U.S.A. Oregon:Lane Co., "Shadywoods of Culp Creek, near base of Bohemia Mt. Divide, betweenLane and DouglasCounties, Oregon," 1600 4 50 m, 2 May 1926, J. R. Patterson 9959 (lectotype, selectedhere, ORE).

In his protologue for Thermopsis gracilis, How- ell did not mention a specific type but rather noted the species to occur in "Mountains of southwestern Oregon, from the sources of the Willamette, to northernCalifornia," which suggests that he knew the plant from a number of collections. Actually, there is only a single sheet at ORE that could have 250 served as the voucher for Thermopsis gracilis. L. 4Q0 A. Vorobik, by annotation, selected this as the holotype. A purist might have called this a lecto- 200 mi. type.

Plants robust, 3-8 dm tall, glabrous to sparsely stems slender, erect, single or clustered, villous; FIGURE 4. Distributionof Thermopsisgracilis. well-branched, arising from a woody rootstock or rhizomes, branches spreading at 30-70?, moderately to stronglyzigzagged. Leaflets elliptic to ob- 1935, Parks & Tracy 11227 (CAS, GH, UC, WTU); 2 ovate or rhombic, 3-6(-8) cm long, 2-3.2 cm mi. S of Gasquet on French Hill Road, 1 June 1968, wide, acute, obtuse, or emarginate, glabrous, lat- Turner5801 (CAS, GH, NY, TEX). HumboldtCo., French eral veins 6-9 pairs, very conspicuously net-veined Camp Ridge,in chaparral,6 Aug. 1933, Tracy 12955 (UC, WTU); WillowCreek Canyon,along TrinityHwy., beneath; petioles 2.5-4(-5) cm long; stipules per- rockypoint at RubyCreek, 31 May 1937, Tracy 15326 sistent, asymmetrically ovate-elliptic, 2.5-3.5 cm (UC, WTU). Shasta Co., betweenLa Moineand Williams long, 1.5-3 cm wide, cuneate to cordate, acumi- Ranch, 19 June 1919, Heller 13250 (CAS, GH, MO, nate. Racemes 12-20(-30) cm long, with 7-30 NY, PH, UC, WTU); McCloud Fork, 24 Apr. 1913, flowers, scattered; peduncles 4-9(- 11) cm long; Smith 110 (GH, MO, NY). SiskiyouCo., Dunsmuir,23 July1912, Eastwood 1319 (CAS, GH, MO, NY, US); to vil- pedicels 6-16 mm long, glabrous sparsely ca. 2 mi. S of Shasta Springs,on the old McCloudRoad, lous; bracts semipersistent, ovate, 6-8 mm long, 2 June 1968, Turner5807 (TEX). TrinityCo., E of 3-5 mm wide. Flowers 2-2.3 cm long; calyx 8- Weaver Creek, 21 May 1914, Yates 7-72 (RM, US). 10 mm long, 7-8 mm wide at the limb, lobes 2- Oregon: BentonCo., Mary's Peak Road, ca. 10 mi. SW ofCorvallis, Mills 1973 (UT); Philomath,23 May 1925, 3 mm long, much shorterthan tube; wings elliptic; Stason 2399 (UC). Coos Co., 3 km NE of Remote,7 keel asymmetricallyoblong-elliptic; ovary sparsely May 1980, Sundberg 994 (OSC). CurryCo., 2 mi. SW villous; ovules 9-14. Pods straight,irregularly di- of turn-offfrom road to Agnessalong road to Iron Mt., vergent, 3-6 cm long, 0.4-0.6 cm wide, incon- 16 June1956, Balls & Lenz 21697 (CAS, NY, POM, spicuously pubescent. Seeds 2-8, tan, oblong with RM, UC, UT, WS); Mt. Emily,lower chaparral belt, 10 Aug. 1921, Jepson& Tracy 5909 (TEX, UC). Douglas mm long, 2 mm wide. Chro- a short rostrum, 3-4 Co., Dog Creek drainageof N. Umpqua Riverupstream mosome number, 2n = 36. fromSteamboat River, above LimpyRock, 19 June1976, Chambers4223 (CAS, NCU, POM, WTU); 10 mi. from Distribution and ecology (Figs. 1, 4). North- West Fork,27 June1917, Peck 7255 (F, OSC). Jackson western California and adjacent Oregon, meso- Co., GraveCreek near Winarr,14 May 1889, Hammond phytic lower montane evergreen forests, mostly 74 (MO). JosephineCo., summitof the lower Siskiyou Mts. especiallyN slopes, 3 July1902, Cusick 2927 (F, 100-1200 m; flowering(April-)May-June. GH, MO, NY, OSC, POM, UC, US, WS); Waldo, July 1894, Howell 1495 (COLO, MO, NY, ORE, OSC, PH, Representativespecimens examined. U.S.A. Cal- UC, US, WS, WTU); Onion Mt. Road, 11 May 1956, ifornia: Del NorteCo., Siskiyoulowlands near Gasquet, Steward7059 (CAS, MO, NCU, OSC, PH, POM, TENN); 22 Apr. 1939, Meola 69 (OSC, UC); N side of Middle SextonMt., 9 Apr. 1934, Thompson10209 (CAS, NY, Fork of SmithRiver on Old Gasquet Toll Road, 1 June POM, WYU). Lane Co., Spencer's Butte, 4 mi. S of

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Eugene, 5 May 1946, Baker 2791 (ARIZ, CAS, MO, mopsis macrophylla in connection with his de- NY, OSC, WTU); near CottageGrove, Lorane Road, 17 scriptionof T macrophylla var. agnina, suggesting May 1931, Henderson 13574 (PH). LincolnCo., Alsea a type locality for T macrophylla near Aromas Mts., Lammi 1939 (OSC). Polk Co., 2 mi. W of Grand Ronde, Brehm 1967 (TEX). in San Benito County. But these, too, are merely large-leaved formsof otherwisetypical Thermopsis This species is readily distinguished from the californica. Type specimens of Thermopsis mac- closely allopatric Thermopsis californica and T rophylla match very closely those of T macro- montana by its divergent pods, spreading branch- phylla var. agnina and had Howell examined ma- es, and glabrous to sparsely pubescent foliage. It terial of the Douglas collection we believe he would is most closely related to T robusta, which also not have described the latter. No doubt much of has divergent pods but which differsin its very the confusion regarding recognition of both Ther- robust habit, densely pilose vestiture, and longer mopsis macrophylla and T. californica is due to calyx lobes. Larisey (1940) maintained Thermopsis the fact that T macrophylla is a relatively rare venosa as a variety of T. gracilis, but Isely (1981) localized endemic. This has been confounded by treated both names as synonyms under T. macro- the existence of Thermopsis robusta, a localized phylla var. venosa. If Isely's treatment is ac- robust endemic with divergent pods, which is re- cepted, the correct taxon name under the existing lated to T gracilis but is superficially similar to nomenclatural code would be Thermopsis macro- T macrophylla in habit and vestiture. At least phylla var. gracilis, since that combination was several workers who have recognized Thermopsis validated withthe erection of T gracilis var. veno- macrophylla as distinct from T californica have sa. Regardless, we feel that T gracilis is clearly treated both T macrophylla and T robusta as the distinctfrom T. macrophylla, as discussed in detail same. under the latter. Plants robust, 12-23 dm tall, tomentose; stems thick, erect, solitary or few, moderately branched, 5. Thermopsis macrophylla Hook. & Arn., from a woody rootstock, branches ascending at Bot. Beechey Voy. 329. 1840. TYPE: U.S.A. 450, moderately zigzagged. Leaflets elliptic, 4-10 California: Santa Barbara Co.?, spring of cm long, 2-5 cm wide, villous, acute, lateral veins 1833?, probably Santa Ynez Mt., near Santa 6-8 pairs, not conspicuously net-veined beneath; Barbara, Douglas s.n. (holotype, K; photo of petioles 2-5(-7) cm long; stipules persistent,widely holotype, POM, UC; isotype, GH). ovate, 3-9 cm long, 2-6 cm wide, amplexicaul to cordate, acuminate. Racemes 25-60 cm long, with Thermopsismacrophylla var. agnina J. Howell,Leafl. 30-90 flowers,in whorls of 3-5; peduncles 8-15 W. Bot. 8: 158. 1957. TYPE: U.S.A. California: Santa Barbara Co., CaminoCielo, ca. 1 mi. W of cm long; pedicels 2.5-4 mm long, villous; bracts Santa Ynez Peak. ca. 1060 m, H. M. Pollard 1955, persistent,ovate to lanceolate, 8-10 mm long, 3- plant B, 2 sheets (lectotype,selected here, CAS; 5 mm wide. Flowers 1.7-2.2 cm long; calyx 7-9 isolectotypes,CAS [2 sheetsplant A], US [3 sheets]). mm long, 7-8 mm wide at the limb, lobes 3-4 Many workers have pondered over the type mm long, as long as or shorter than tube; wings locality of Thermopsis macrophylla, equating elliptic; keel obovate; ovary velutinous; ovules 7- Thermopsis macrophylla with Thermopsis cali- 9. Pods straight,ascending, 3.5-5 cm long, 0.5- fornica S. Watson, but examination of type ma- 0.7 cm wide, villous. Seeds 5-7, brown-black,widely terial and knowledge of the variation of Thermopsis elliptic with a short rostrum, 4-5 mm long, 2.5- californica throughout its range strongly suggest 3.0 mm wide. Chromosome number, 2n = 18. that Thermopsis macrophylla was originally col- Distribution and ecology (Fig. 1). Known only lected in the Santa Ynez Mountains near Santa from the Santa Ynez Mountains, Santa Barbara Barbara. Douglas resided for a time in Monterey, Co., California, where it is locally common on the Santa Cruz County, California,from where he made west side of Santa Ynez Peak near the summit, in several overland visits to Santa Barbara Mission. sandy graniticsoils, 1000-1400 m; floweringMay- Eastwood, as noted by Howell (1957), thought she June. might have rediscovered the type locality of Ther- mopsis macrophylla in the Santa Cruz Mountains, Specimens examined. U.S.A. California: Los An- Santa Cruz County, California, near Glenwood, but geles Co., Claremont,weed on the Rancho Santa Ana examination of this material (Davis 1907, CAS, BotanicGarden grounds, "apparently introduced," 9 May 1991, Ross 5047 (POM). Santa Barbara S of GH, RM, UC, US) reveals these to be robust forms Co., slope Santa Ynez Peak, 1130 m, 11 May 1962, Breedlove of otherwisetypical Thermopsis californica. How- 2695 (CAS, POM); Santa Ynez Peak, Fairfax 1977 ell also discussed the possible type locality of Ther- (RM); Santa Ynez Peak, ca. 4000 ft.,Pollard 1956 (CAS,

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TEX); in chaparralW ofSanta Ynez Peak, 17 Oct. 1945, sometimes glabrate above, lateral veins 7-9 pairs, Smith1620 (POM); just W of Santa Ynez Peak, 12 May very conspicuously net-veined beneath; petioles 1957, Smith 5283 (POM); ca. 0.5 mi. due W of Santa 0.7-1.7 cm Ynez Peak, 4 June1968, Turner5820 (GH, NY, TEX); long; stipules caducous, narrowly el- Santa Ynez Mts., along W CaminoCielo Road, 5.5 mi. liptic to linear, 1.5-2 cm long, 0.3-0.5(-0.8) cm E of intersectionwith Refugio Road, 3900 ft., 26 June wide, cuneate, acuminate. Racemes terminal, 7- 1992, Junak 5022 (TEX); Santa Ynez Mts., along W 20 cm long, with 5-17 flowers,scattered; pedun- CaminoCielo Road, 6.2 mi. E ofintersection with Refugio cles 3-7 cm long; pedicels 4-6 mm long, villos- Road, 4020 ft.,26 June 1992, Junak 5020 (TEX). ulous; bracts persistent,lanceolate, 8-13 mm long, The relationship of this localized endemic and 2-4 mm wide. Flowers 1.6-1.9 cm long; calyx its longtime confusionwith Thermopsis californica 7.5-8 mm long, 6-7 mm wide at the limb, lobes is discussed in the above account of its typification. 2-3 mm long, shorter than tube; wings elliptic to The early workers, mainly S. Watson, who pos- asymmetricallyoblong-elliptic; keel asymmetrically sessed an isotype of T macrophylla, maintained oblong-elliptic;ovary densely appressed pubescent; both T. californica and T. macrophylla as con- ovules (8-)12-16. Pods arcuate to straight, di- ceived in the present study. Subsequent workers, vergent, (3-)4-9 cm long, 0.4-0.6 cm wide, mostly Californian, confounded the two taxa. The densely appressed pubescent. Seeds 6-12, oblong taxa were not clarified by Larisey (1940), who witha rostrum,2.5-3.5 mm long, 1.5-2 mm wide. treated both T. californica and T. robusta as syn- Chromosome number, 2n = 18. onymous with her T. macrophylla var. macro- Distribution and ecology (Fig. 5). Mostly mid- phylla (she ignored T. macrophylla var. agnina). elevations of the southern Appalachian Mountains In this Larisey was followed by Isely (1981) who and adjacent piedmont region of northernmost also treated all of these as synonymous with his Georgia, eastern Tennessee, northwestern South concept of T. macrophylla var. macrophylla. To Carolina, western North Carolina, and central Vir- our knowledge, Douglas made only two collections ginia, mostly 300-800 m; flowering(April-)May- of Thermopsis in California, the type of T. mac- June. rophylla and a collection of T californica var. californica. Torrey & Gray (1840) referred the Representativespecimens examined. U.S.A. Ala- latter to T. macrophylla var. fi. This specimen bama: De Kalb Co., Mentone,Loving 1899 (US). Geor- gia: Dade 1 (GH) is a fairlytypical specimen of T. californica, Co., mi. W of Durhamand NE of Cloudland Canyon,12 May 1953, Hardin 15933 (NCU, US); Look- presumably collected in the spring of 1833, per- out Mt., 26 Apr. 1952, Harper 4204 (NY, US). Hab- haps in the vicinity of Monterey, California, to ershamCo., near Cornelia,Huger 1898 (NY). Walker judge from its floweringstate and vestiture. Co., 22.4 mi. SE of La Fayette, 24 Apr. 1961, Turner 4691 (TEX). WhitfieldCo., 4 mi. NW ofDalton, 24 Apr. 1948, Cronquist5055 (NY); near summitof Chattoogata 6. Thermopsis mollis (Michaux) M. Curtis ex Mts., 19 July 1990, Harper 265 (NY, US). North A. Gray, Mem. Amer. Acad. Arts n.s. 3: 47. Carolina: BuncombeCo., [Asheville],Biltmore, 1 May- 10 June 1896, Biltmore1025 (F, MO, NY, PH, RM, 1848. Podalyria mollis Michaux, Fl. Bor.- TENN, UC, US). CaswellCo., Milton,Fomey 1860 (PH). Amer. 1: 264. 1803. Baptisia mollis (Mi- CatawbaCo., 3 mi. SSE of Newton,29 Apr. 1957, Bell chaux) Nutt., Gen. N. Amer. P1. 1: 281. 1818. 6822 (NC, NY); 1.25 mi. SW of Hickoryon U.S. Hwy. TYPE: U.S.A. North Carolina: Mecklenburg 70, 3 May 1953, Radford & Wood 6970 (NCU, US). ChathamCo., of Co., vicinity of Mecklenburg, without date, vicinity Bush Creek,3.9 mi. S ofOrange Co. line on U.S. Hwy. 15 and 0.6 mi. E on paved road, Michaux s.n. (holotype, P not seen; probable 24 Apr. 1957, Ahles 23825 (GH, NCU); Edward's Mt., isotype, PH). 3 mi. E of U.S. Hwy. 501, 6.6 mi. S of Chapel Hill, 11 Apr. 1940, Radford& Steward 77 (NCU). DurhamCo., Baptisia hugeri Small, Bull. TorreyBot. Club 25: 139. Lynn 1933 (ND). ForsythCo., Schallert 1940 (GH); S 1898. Thermopsishugeri (Small) Small, Fl. S.E. edge of Winston-Salem,Wherry 1928 (NY). GastonCo., U.S. 596, 1331. 1903. TYPE: U.S.A. Georgia:Hab- near summitof Kings Mt., 24 Apr. 1968, Leonard & ersham Co., "northernslope on MountGriffin," near Radford 1353 (LL, NCU, OSC, TENN, TEX); Crowders Cornelia,A. M. Huger 1897 (holotype,NY; isotype, Mt., 4 June 1953, Radford & Haesloop 7089 (ARIZ, MO). NCU). GuilfordCo., GuilfordCollege, 30 Apr. 1958, Terrell3205 (NCU). HendersonCo., Flat Rock, Mem- Plants delicate, 3-6 dm tall, sparsely appressed minger1886 (NCU). IredellCo., 3.3 mi. W of Harmony pubescent; stems slender, ascending or spreading, on NorthCarolina Hwy. 901, 21 Apr. 1953, Daggy solitary or clustered, moderately branched, arising 2221 (NCU); Statesville,Hyams 4968 (F, MO, NY, from extensive rhizomes; branches spreading at POM). LincolnCo., Hunter s.n. (GH, NY); near Vale, 28 Apr. 1957, Bell 6654 (NCU). 45-800, stronglyzigzagged. Leaflets elliptic, 3.5- OrangeCo., near Hills- boro, Curtiss.n. (NY). Stokes Co., 1 mi. N of Hanging 8 cm long, 1-4 cm wide, acute to acuminate, Rock, 30 Apr. 1961, Carswell 192 (NCU); above Dan- sparsely appressed pubescent especially on veins, bury,Hunt 1936 (NCU). SurryCo., PilotMt., Ashe 1896

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tY;~~~~~~~0

MILES 0 100 200 300 400

900 850 800 750

FIGURE 5. Distributionof Thermopsis mollis.

(NCU); Pilot Mt., 5 May 1935, Oosting 3519 (NCU). bellCo., SleepyHollow Road, 23 May 1967, Freer5597 Vance Co., Kittrell,Browne 1884 (NY); Kittrell'sSprings, (GH, NCU). CharlotteCo., 4 mi. S of Charlotte,Court Faxon 1874 (GH). Wake Co., 5.5 mi. NE of Holly House township,16 Apr. 1967, Harvill 15436 (NCU). Springs,28 Apr. 1954, Radford& Beard 7984 (NCU); Halifax Co., near Halifax, Rowlett 1921 (US). Prince W of Raleigh,1 mi. N of Tyson's Store,30 Apr. 1951, EdwardCo., HampdenSydney College, Blair 1981 (F, Fox 4580 (MO). WarrenCo., Ridgeway,20 Apr. 1938, GH, MO, NY, US); Farmville,Fattig 1922 (US). Rock- Godfrey3600 (GH, NCU). South Carolina: Greenville bridgeCo., BalconyFalls, 1 May 1947, Freer 1348 (GH). Co., ParisMt., Huger 1903 (NY); ParisMt., Huger 1904 (NY). Oconee Co., Rich Mt., 5.5 mi. W of Walhalla, 16 The relationship of this species to Thermopsis Apr. 1967, Bozeman et al. 8862 (COLO, NCU, POM, fraxinifolia is discussed under T. fraxinifolia. WTU); MountainRest, ca. 0.5 mi. W of West Village Fernald (1950) noted that at least one popula- 258, 30 Apr. 1989, Hill 20511 Creekon CountyRoute tion of Thermopsis mollis from Essex Co., Mas- (NY). Pickens Co., 12.5 mi. S of Rosman along U.S. Hwy. 178, 1 May 1941, McVaugh 5649 (GH, UC). sachusetts, had been introduced and naturalized in Tennessee: BlountCo., ChilhoweeMt., 26 Apr. 1939, the northeastern United States, although he ig- Jennison299 (TENN); ChilhoweeMt., between Millstone nored the more widely cultivated T. villosa. We Gap and Walland, Thomas 1965 (TENN). FranklinCo., have examined material from Essex Co. (GH), pre- edge of Sewanee, 3 May 1941, Sharp & Meyer 1259 sumably that referredto by Fernald, but it appears (MO, PH, TENN, UC); St. Andrews,25 Apr. 1936, Svenson 7592 (GH, MO, PH, UC). GraingerCo., Lea to be T. villosa. Lakes, 3 May 1934, Hesler & Sharp 566 (TENN). HamiltonCo., LookoutMt., Churchill1906 (GH); Look- 7. Thermopsis montana Nutt. ex Torrey & out Mt., Churchill1906 (GH, MO, TENN). MarionCo., A. Gray, Fl. N. Amer. 1: 388. 1840. TYPE: W ofWhitwell, 14 May 1934, Harger 7788 (GH, TENN). Sevier Co., Gatlinburgentrance to Great SmokyMoun- see Thermopsis montana var. montana. tainsNational Park, Rogers 41851 (TENN); Chilhowee Plants delicate, 2-8(-10) dm tall, glabrous to Mt., 7 May 1964, Thomas 32794 (TENN). Virginia: BedfordCo., intersectionof County Route 711 and Coun- appressed pubescent or thinly villous; stems slen- ty Route 811, 11 Aug. 1985, Wright46 9 (MO). Camp- der, erect, solitaryor clustered, moderately or few-

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branched, arising from a woody rootstock or rhi- According to the known distribution of Ther- zomes; branches ascending at 20-45?, weakly to mopsis montana, Nuttall would most likely have moderately zigzagged. Leaflets narrowly elliptic or firstencountered the species in Sweetwater Co., obovate, 3.5-8 cm long, 0.7-3(-5) cm wide, acute Wyoming, at about the time indicated. It is also or obtuse, sometimes short mucronate or emargin- possible that it was firstcollected in southeastern ate, sparsely appressed pubescent to glabrous, lat- Idaho in early July (cf. Graustein, 1967). eral veins 6-1 1 pairs, conspicuously net-veined Plants 2-7 dm tall, appressed pubescent to thin- beneath or not; petioles 1-4 cm long; stipules per- ly villous; stems stifflyerect, solitary or clustered, sistent, widely ovate or elliptic, 1.7-4.5 cm long, few-branched, arising from a woody rootstock or 0.7-2.5 cm wide, oblique to cuneate, acuminate rhizomes; branches moderately zigzagged. Leaflets or acute. Racemes 4-25 cm long, with 6-25 flow- elliptic to narrowly elliptic or obovate, 3.5-8 cm ers, in whorls of 2-3 or scattered; peduncles 2-8 long, 0.7-2.5 cm wide, appressed pubescent, not cm long; pedicels 3.5-5 mm long, villosulous;bracts conspicuously net-veined beneath; petioles 1-2.5 semipersistent, narrowly elliptic to widely ovate, cm long. Racemes 4-18 cm long; bracts elliptic 6-10 mm long, 2.5-6 mm wide. Flowers 1.6-2.2 to widely ovate. Flowers 1.6-2 cm long. Pods cm long; calyx 9-11 mm long, 5-8 mm wide at villosulous. Chromosome number, 2n = 18. the limb, lobes 3-5 mm long, shorter than tube; wing and keel petals asymmetricallyoblong-elliptic; Distribution and ecology (Fig. 6). Widely ovary velutinous or densely appressed pubescent; distributedin the intermountainregion of the Unit- ovules 10-16. Pods straight, erect, 4.5-6.5 cm ed States, from southwestern Montana southward long, 0.4-0.6 cm wide, villosulous or appressed to Arizona and New Mexico and northwestwardto pubescent. Seeds 6-16, brown-black, oblong with southern Idaho and adjacent Wyoming, middle to a rostrum, 3.5-5 mm long, 2.5-3 mm wide. high elevations, 2000-3200 m; floweringMay- June. We recognize two intergradingvarieties under this taxon, as follows: Representativespecimens examined. U.S.A. Arizo- na: ApacheCo., 1 mi. N of Alpine,27 June1944, Pultz KEY TO VARIETIES OF THERMOPSIS MONTANA & Phillips 1030 (ARIZ, NY, UC). CochiseCo., Barefoot Park, Blumer 1907 (ARIZ, F, GH, MO, NY, RM, US). la. Plants less than 7 dm tall, pubescent;stems CoconinoCo., Flagstaff,21 Apr. 1925, Nelson 10200 oftensolitary and few-branched,often arising (GH, MO, NY, PH, POM, RM, UC, WS). Gila Co., Sierra fromrhizomes; leaflets not conspicuouslynet- Ancha,Aztec Peak, 1 May 1931, Harrison 785 7 (ARIZ, veinedbeneath; bracts elliptic to widelyovate US). Graham Co., Graham Mts., Pine Crest, 26 May -...... 7a. Thermopsismontana var. montana 1935, Maguire 12013 (NY, WS, WTU). GreenleeCo., lb. Plants up to 10 dm tall, glabrate;stems clus- near Rose Peak, 43 mi. N of Clifton,7 June 1935, teredand moderatelybranched, arising from a Maguire 11886 (ARIZ, GH, NY, UC). Navajo Co., head woody rootstock;leaflets conspicuouslynet- ofLittle Colorado, 10 July1912, Goodding1147 (ARIZ, veinedbeneath; bracts narrowly elliptic. NY, RM, US). Yavapai Co., S of Prescott,2 June1944, -...... 7b. Thermopsismontana var. ovata Clark 11770 (GH, US). Colorado: ArchuletaCo., Pie- dra, near Piedra River, 18 June 1924, Schmoll 1122 7a. Thermopsis montana var. montana. (RM). ChaffeeCo., Poncha Springs,at intersectionof Thermopsis fabacea var. montana (Nutt. ex railroadand highwayalong creek bed, 27 May 1968, Turner5777 (TEX). Clear Creek Co., Brookvale,Bear Torrey & A. Gray) A. Gray, Proc. Acad. Nat. Creek, Churchill1918 (GH, MO). Conejos Co., SW of Sci. Philadelphia 1863: 60. 1864. Thermop- La JaraReservoir, 19 Aug. 1986, O'Kane & Anderson sis rhombifolia var. montana (Nutt. ex Tor- 2618 (RM). CostillaCo., 61 mi. E ofMontevista, La Veta rey & A. Gray) Isely, Brittonia30: 470. 1978. Pass, 30 May 1934, Frederick& Stone 608 (NY). Custer Co., W of Westcliffe,trail to Lakes of the Clouds, 29 TYPE: U.S.A. Wyoming: "Rocky Mountains" July1950, Weber5894 (COLO). Delta Co., Mule Park, [Sweetwater Co.?], Nuttall 1834 (holotype, 12 July1962, Gierisch2552 (RM). Dolores Co., Lone BM not seen; photo of holotype, POM, UK; Cone, along watercourses, Brewster s.n. (COLO). Eagle isotype, PH; photo of isotype, POM). Co., McCoy,edge ofmeadow, 30 July1898, Shear 5310 (RM). GarfieldCo., Glenwood1910 (GH). GunnisonCo., Thermopsisangustata E. Greene,PI. Baker.3: 34. 1901. Sapinero,19 June1901, Baker 173 (MO, RM, ND, NY, TYPE: U.S.A. Nevada: Elko Co., Star Valley,"foot- POM, UC). La Plata Co., 10 mi. W of Durango,20 June hillsof the Ruby Mountains,"E. L. Greene 1896 1965, Strother389 (TEX). Las Animas Co., Wilkins (holotype,ND). Creek,ca. 0.25 mi. NW ofStonewall Gap, 15 Aug. 1987, Thermopsisstricta E. Greene,PI. Baker. 3: 34. 1901. Neely4635 (RM). Mesa Co., UncompahgrePlateau, head TYPE: U.S.A. Colorado:Gunnison Co., Gunnison,24 ofEscalante Canyon, 19 July1982, Siplivinsky& Beck July1901, 2330 m, C. F. Baker 604 (lectotype, 4251 (COLO, RM). MineralCo., WolfCreek Pass, U.S. selectedhere, ND; isolectotypes,GH [2 sheets],NY Hwy. 160, 3 mi. belowsummit on E side,27 Aug. 1964, RM [2 sheets],UC, US [2 sheets]). Turner5099 (TEX). MontezumaCo., along DoloresRiv-

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er, EscalanteSector, Bye et al. 1978 (COLO). Montrose Mescalero,26 May 1934, Ferguson& Ottley5223 (UC). Co., 30 mi. SW of Montrose,28 Aug. 1937, Rollins Rio ArribaCo., Ortega Mts., ca. 2 mi. W of Hopewell 1978 (CAS, GH, MO, ND, NY, RM, US). Ouray Co., Lake, 17 Aug. 1984, Hill 15167 (NY). San MiguelCo., Clements1907 (NY). Rio Blanco Co., 2 mi. E of Meeker, 8 mi. E of Glorietta,along the Pecos River,9 June1897, 18 Aug. 1935, Maguire & Piranian 12844 (MO). Park A. A. & E. G. Heller 3681 (MO, NY, US). Sandoval Co., JeffersonSouth Park, 9 Sep. 1942, Ewan 14706 Co., 24 mi. N of JemezPueblo, 6 June 1974, Atwood (COLO). Rio GrandeCo., alongSouth Fork of Rio Grande 6249 (NY). Santa Fe Co., nearSanta Fe RecreationArea, River, 27 Aug. 1964, Turner5095 (TEX). Saguache just offNew Mexico Route 475, 16 June1964, Perdue Co., MarshallPass, 21 July1960, Barrell 127-60 (US). 6047 (US). Taos Co., Penasco, Santa Barbara Canyon, Idaho: Bannock Co., E of Lava Hot Springs,Everson 7 July1936, Marcelline 1989 (F). Oregon: CrookCo., 1946 (NY). Bear Lake Co., 1.3 mi. N of St. Charles,25 5 mi. W of Prineville,15 June1928, Pech 15964 (OSC, June 1967, Faircloth 4548 (NCU). BinghamCo., Ft. PH). GrantCo., 3 mi. E of CanyonCity, 5 June 1953, Hall, Everson 1942 (TEX). CaribouCo., 8 Mile Canyon, Cronquist7087 (GH, NY, WS, WTU). HarneyCo., 23 12 mi. S of Soda Springs,28 June1960, Cottamet al. mi.S ofFrenchglen, 16 May 1959, Cronquist8245 (GH, 16280 (UT). Cassia Co., 20 mi. S ofOakley, along Goose NCU, NY, POM, RM, TEX, UC, WS, WTU) [approaches Creek,2 July1950, J. H. & C. B. Christ19580 (WS). varietyovata]. MalheurCo., 3 mi. W of McDermitt,25 Clark Co., Kaufman,near Birch Creek, 19 July1934, June1934, Peck 21734 (OSC, UC). Utah: Beaver Co., Christ 5818 (NY). Elmore Co., Moore's Creek, Boise Wah Wah Mts.,Birch Creek, 4 May 1937, Cottam7109 Rivertributary, Grandjean 155 (RM). FranklinCo., 11 (UT). Box Elder Co., BrighamCity, 9 May 1910, Smith mi. NE of Preston,5 mi. up MillcreekCanyon, 14 May 2120 (GH, NY). Cache Co., 0.25 mi. E of Wellsville,30 1966, G. & G. Davidse 391 (UC, US, WTU). Lemhi May 1940, Maguire 18600 (NY, WTU). DaggettCo., Co., 20 mi. E of Gilmore,along Birch Creek, 25 June vicinityof FlamingGorge, 29 May 1932, Williams432 1947, Hitchcock 15766 (CAS, COLO, MO, NY, PH, (GH, MO, NY, RM). DuchesneCo., 5 mi. N ofDuchesne, POM, RM, UC, WS, WTU). Oneida Co., dividebetween 6 June1979, Neese & Welsh 7503 (NY). GarfieldCo., Marshand Malad Valleys,Coulter s.n. (PH). OwyheeCo., 7 mi. SE of Boulder,Deer Creek, 4 May 1977, Foster ca. 39 mi. S of Bruneau,30 May 1968, Turner5798 3661 (NY, UT). GrandCo., 2 mi. N of WarnerRanger (NY, RM, TEX). Twin Falls Co., Twin Falls, 26 May Station,La Sal Mts., 8 July1933, Maguire 313 (UC, 1912, Bennett 88 (RM). Montana: Beaverhead Co., WTU). Iron Co., ca. 15 mi. E of Cedar City,just W of Armstead,20 June 1920, E. B. & L. B. Payson 1734 the entranceto Dixie National Forest, 13 June 1969, (CAS, GH, MO, RM). BroadwaterCo., 4 mi. SE ofTown- Meyer 356 (UT). MillardCo., Simonsen'sRanch, along send,10 June1960, SchmautzJES-345 (CAS, RM, WS). irrigationstream, 16 June 1933, Maguire & Becraft DeerlodgeCo., Fishtrap,ca. 25 mi. S of Anaconda,20 3932 (GH, UC). MorganCo., flatarea belowdam by old June1944, Hitchcock& Muhlick 9173 (CAS, GH, MO, gaugingstation, 17 May 1987, Peterson 14 (NY, UT). NY, RM, UC, WS, WTU). GallatinCo., Belgrade,low Piute Co., 5 mi. E of Kingstonon Utah Hwy. 62, 3 May ground,2 June1906, Blankenship 681 (F, MO, POM, 1977, Foster 3613 (NY, RM). Rich Co., W shore of RM, UC, US). JeffersonCo., Boulder,24 Apr. 1939, Bear Lake, 0.7 mi. E of Pickleville,5 July1985, Thorne Rose 709 (RM). Madison Co., VirginiaCity, Swingle 3938 (GH, MO). Salt Lake Co., vicinityof Salt Lake City, 1933 (RM, WTU). Powell Co., vicinityof Deer Lodge, Clements1908 (F, GH). Sevier Co., 3 mi. S of Sevier, Fitch s.n. (RM). SilverBow Co., LowerMoose Creek,ca. along stream,8 May 1937, Hitchcock2947 (GH, RM, 1 mi. N of Moose Creek ParkingArea, 28 June 1979, UC, WS, WTU). San JuanCo., ca. 80 mi. S of Moab, Lowry2270 (RM). Nevada: Elko Co., 3 mi. N of Ruby NW flankof Abajo Mts., 12 June 1961, Cronquist& Valley on E side of Ruby Mts., 29 May 1968, Turner Holmgren9259(GH, NY, NCU, POM, TEX, WS, WTU). 5795 (TEX). EurekaCo., Eureka,2 July1904, Kennedy SummitCo., GoodmanRanch near Bear River,30 June 838 (ARIZ, MO, RM, UC). HumboldtCo., Santa Rosa 1926, E. B. & L. B. Payson 4833 (MO, PH, POM, Range, NE of Winnemucca,29 June 1967, Gentry& RM, UC, US); UintahCo., E of Hayden, along Uintah Davidse 1674 (ARIZ, COLO, GH, MO, NY, OSC, POM, River, 2 June 1933, Graham 8044 (F, GH, NY, US). UC, WTU). LanderCo., Reese RiverValley, U.S. Hwy. Utah Co., neartop of Price Canyon, Malburtt 1959 (UT). 50, along edge of Reese River, 13 June 1964, Spellen- Wasatch Co., 6 mi. N of Heber City,along highway, 28 berg 370 (F, NY). LincolnCo., GeyserRanch, 28 May May 1968, Turner5794A & 5794B (TEX). Washington 1945, Ripley & Barneby6436 (NY). Nye Co., Darrough Co., 5 mi. SW of Enterpriseat Aspen Park Spring,10 Hot Springs,Smoky Valley, 11 May 1939, Train 2743 June1983, Atwood9366 (NY, RM). Wayne Co., Fruita, (MO, NY, POM, TEX, UC, WTU). White Pine Co., 18 May 1956, Vickery587 (ARIZ, GH, POM, UT, canyonE of Mt. Grafton,11 Aug. 1969, Holmgren& WTU). Weber Co., Huntsville,26 May 1940, Brizzee Bethers 3861 (NY, POM, US, WTU). New Mexico: 7794 (UT). Wyoming: CarbonCo., Baggs,4 July1926, BernalilloCo., 8.5 mi. fromSan Antonito,0.7 mi. N of Nelson 10720 (GH, MO, RM, UC). Park Co., McCosh CapulinSpring, 21 May 1967, Turner5649 (CAS, NY, 1878 (NY). SweetwaterCo., Uinta Mts., HenrysFork, TEX). CatronCo., MogollonMt., near West Fork of the riverbottoms, 26 June1902, Goodding 1192 (CAS, F, Gila River,23 Aug. 1903, Metcalfe593 (MO, ND, NY, GH, MO, NY, POM, RM, UC). Uinta Co., 1 mi. S of RM, US). ColfaxCo., 1 mi. NW of Ute GulchCamp, 17 Lonetree,27 July1939, Rollins & Munoz 2903 (CAS, June1968, Hartman 1973 (RM, TEX). GrantCo., Pinos GH, US). Altos,2.6 mi. by road to Signal Peak E of New Mexico Hwy. 15, 25 May 1985, Worthington13180 (NY, RM). Thermopsis montana var. montana is wide- LincolnCo., Sierra Blanca, valleyof CarrizoCreek, 11 spread and highly variable but can usually be dis- June1932, Wilkins2402 (PH, US); McKinleyCo., Long from its smaller habit, Canyon, 6 mi. S of Crownpoint,Jump 1974 (ARIZ). tinguished variety ovata by Mora Co., 1 mi. E of La Cueva, Stiteler 1949 (PH). narrower leaflets and bracts, and greater tendency Otero Co., near summitof pass betweenHollywood and to spread by rhizomes. In southwesternOregon the

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500 125" ~~~1200C 1150 1100 1050

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taxa appear to intergrade.Isely (1981) recognized The strictlyerect pods distinguishThermopsis a "Lemhi race" (LemhiCounty, Idaho) withinthe montana fromT. gracilis and fromthe superfi- taxon we call T. montana var. montana. Though ciallysimilar T. divaricarpa. Isely (1981) treated withsomewhat broader leaflets, most of those spec- T. montana and T. divaricarpa as varietiesof a imensare fairlytypical of T. montana var. mon- widespread,extremely variable T. rhombifolia.We tana. have foundthat T. rhombifolia,T. divaricarpa, In Arizonaand mostof New Mexico(from below and T. montana are clearly distinctthroughout San Juanand Rio Arribacounties) there also occurs mostof their ranges and do notappear to intergrade a seriesof populationsthat are composedof rather as do T. montana var. montana and T. montana broad-branchingplants withleaves more or less var. ovata. Intermediatepopulations and possible like Thermopsis montana var. ovata. Unlike Isely hybridsare discussed under Thermopsisdivari- (1981), we do not findsuch plantsin Colorado. carpa.

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7b. Thermopsis montana var. ovata (Rob- Representativespecimens examined. U.S.A. Idaho: inson ex Piper) St. John,Torreya 41: 112. Ada Co., 13 mi. E of Boise, Moore's Creek Canyon,14 (NY). Adams ovata June 1947, J. H. & C. B. Christ 16858 1941. Thermopsis montana subsp. Co., Davis 1939 (WS); Hornet,13 May 1915, Mangun Robinsonex Piper, Contr. U.S. Natl. Herb. 139 (RM). BoiseCo., 15 mi.W ofLowman, above Payette 23: 49. 1906. Thermopsisovata (Robinson River, on road betweenBanks and Lowman, 31 May ex Piper)Rydberg, Bull. TorreyBot. Club 40: 1944, Hitchcock & Muhlick 8563 (NY, WS, WTU). 43. 1913. Thermopsisrhombifolia var. ovata ButteCo., Christ3386 (NY). CanyonCo., Falk's Store, 10 May 1911, Macbride 800 (F, GH, MO, NY, POM, (Robinsonex Piper) Isely, Brittonia30: 470. RM, UC, US, WS, WTU). ClearwaterCo., Clearwater 1978. TYPE: U.S.A. Idaho: Latah Co., "Cedar Forest,Lolo Creek, 16 Oct. 1934, Christ6322 (NY); 5 Mountain,"4 July1893, C. V. Piper 1489 mi. W of Orofino,S side of the ClearwaterRiver, 5 May (lectotype,selected here, WS; isolectotypes, 1951, Roeder 241 (WS). Idaho Co., 0.5 mi. fromPilot Rock, Elwood 51 (CAS, NY, POM, UC, WS, WTU); 7 GH, US, WS). Trii.SE of Elk City,14 June1950, Jones 14 (CAS, GH, POM, WS, WTU); 2 mi. S ofGrangeville, 29 May 1944, Thermopsisxylorhiza Nelson, (xylorrhiza auct.), Bot. Hitchcock& Muhlick 8455 (CAS, NY, RM, UC, WS, WTU). Latah Co., woodedhills just W of Troy,24 May Gaz. 52: 265. 1911. TYPE: U.S.A. Idaho:Canyon Co., Falk's Store,667 m, 24 May 1910, J. F. 1964, Hitchcock 23465 (CAS, NY, UC, WS, WTU). Macbride99 (holotype,RM; isotypes,GH, MO, Lewis Co., MissionCreek, 30 May 1924, John et al. NY, RM,UC, WS). 6334 (UC, WS, WTU). Nez Perce Co., Aug. 1898, Thermopsismacrophylla var. hitchcockii Isely, Brittonia Brown 6 (CAS, MO, ND, NY, PH, UC, US); 3 mi. below Lowell,30 May 1936, Constance & Rollins 1621 (F, 30: 469. 1978. TYPE: U.S.A. Washington:Grays HarborCo., roadside near Humptulips, 10 July 1931, GH, NCU, MO, NY, RM, UC, PH, WS, WTU); near J. W. Thompson7342 (holotype,WTU; isotypes, Lewiston,7 May 1896, A. A. & E. G. Heller 3035 (CAS, GH, OSC, PH, POM, UC). F, MO, ND, NY, PH, UC, WTU); valleyof Hatwai Creek, 28 Apr. 1892, Sandberg et al. 68 (ARIZ, F, GH, MO, NY, POM, US). ShoshoneCo., 8 mi. N of Calder,along Piper's collectionnumber 1489 was apparently St. JoeRiver, 19 May 1940, Christ10969 (NY); Clear- a mixtureof plantscollected on 16 June 1893, 4 waterCamp on the LittleNorth Fork of the Clearwater 4 1941, Wilson 467 (WS). Valley Co., 5 21 1894 (in flower). River, Aug. July1893 (in fruit),and June mi. W of McCall, 24 June1946, Hitchcock& Muhlick We have selected the fruitingspecimen as the 13937 (CAS, MO, NY, POM, RM, UC, WS); 9 mi. E lectotype.It is assumedthat all of the collections ofMcCall, 11 July1953, Holmgren& Tillett9542 (GH, came fromthe same site,although this is notcertain NY, OSC, UC, WTU). WashingtonCo., S of Council, because some of the labels (21 June 1894, NY) wet meadowsalong highway,11 May 1939, Cronquist 1151 (NY). Montana: Sanders Co., Dixon, 31 May givethe locality as "along streamsin woods." The 1937, Brome 28 (MO, PH, RM, WTU); 2 mi. W of publishedtype locality is givenby Peck as "Cedar Dixon,Wolf Point, 13 May 1936, Hitchcock2877 (MO, Mountain"; this localitywas not on any of the ND, OSC, RM, WS, WTU). Oregon: Clackamas Co., labels numbered1489 thatwe examined. Eagle Creek,high hills, 24 May 1884, Henderson 179 (OSC, PH, WS); near base of Table Rock, Leach 1927 Plants 5-8(-10) dm tall, glabrateto sparsely (WTU). Linn Co., MonumentPeak, Gilkey 1947 (OSC, appressedpubescent; stems erect, clustered, mod- POM). MarionCo., Silverton,Spring 1871, Hall 83 (F, POM); 6 mi. E of Brownsville,Cascade Mts., 18 June eratelybranched, arising from a woodyrootstock; 1918, Lawrence 1751 (OSC, US); SilverCreek Falls, 19 branchesweakly to moderatelyzigzagged. Leaflets June 1918, Nelson 2257 (GH). Polk Co., 2 mi. W of broadlyto narrowlyelliptic or obovate,4.5-6(-8) GrandRonde, Brehm 1967 (TEX). UmatillaCo., 1 mi. cm long, 1.5-3(-5) cm wide, sparselyappressed SE of Meacham, 23 June 1946, Hitchcock & Muhlick NY, OSC, UC, WS, WTU); glabrous above, conspicuously 13841 (CAS, GH, MO, pubescentbelow, betweenPendleton and LaGrande,near summitof Blue net-veinedbeneath; petioles3.2-4 cm long. Ra- Mts., 14 June 1928, Thompson4723 (CAS, GH, MO, cemes 12-25 cm long; bracts narrowlyelliptic. US, WTU). Union Co., Kamela, 1 June 1910, Heller Flowers1.9-2.2 cm long. Pods appressedpubes- 10145 (CAS, GH, PH, UC); 2 mi. S of Elgin, 20 May cent. 1975, Watkins& Dunn 95 (MO, NY). Wallowa Co., along summitof road betweenElgin and LangdonLake, Distributionand ecology (Fig. 6). Columbia 28 July1950, Kruckeberg2487 (CAS, COLO, GH, NY, and Snake Riverdrainages of northernIdaho and RM, UC); WalupiCanyon, 18 Aug. 1897, Sheldon 8724 (NY, US). Washington: AsotinCo., Snake River,just adjacentstates of Wyoming,Washington, and Or- below the mouthof the Grande Ronde, 17 Apr. 1949, egon, withisolated populationsalong the Pacific Cronquist5701 (GH, NY, POM, UC, US, WS, WTU); coast of Washingtonand northwesternOregon, SW of Anatone,along Grande Ronde River,just above 300-2000 m in the upper river drainages but RattlesnakeCreek, 22 Apr. 1949, Cronquist5709 (GH, Columbia be- lowerelevations m) about the NY, POM, UC, US, WTU). Co., Dayton, mostlyat (10-200 tweenhospital and Touchet River,3 May 1983, Baird mouthof the Columbia River and its immediate 51 (NY); Blue Mts., 4 mi. N of Table Rock, 20 July tributaries;flowering May-June. 1935, Constance et al. 1278 (MO, WS). GraysHarbor

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Co., N of Aberdeenand Hoquiam along U.S. Hwy. 101, fornicaalso as partof the T. rhombifoliacomplex, 31 May 1987, Boufford& E. W. Wood 23757 (A, CAS, themorphological boundaries between T. montana MO, NCU, NY); ca. 20 mi. N of Hoquiam, 24 June, Hitchcock 19872 (COLO, OSC, POM, UC, WTU); be- var. ovata and T. californica being at least as tweenNew Londonand Humptulips,28 June1933, Jones weak as those betweenT. rhombifoliaand T. di- 3937 (CAS, POM, RM, UC, WTU); open burns near varicarpa or T. montana. Humptulips,30 May 1940, Thompson14606 (CAS, GH, MO, NY, PH, POM, RM, UC, US). LewisCo., Chenalis, OxfordPrairie, 12 June1897, Lamb 1197 (CAS, F, MO, NY, PH, UC, WS). Walla Walla Co., 2 mi. S of Walla 8. Thermopsis rhombifolia (Nutt. ex Pursh) Walla, 24 May 1944, Hitchcock& Muhlick8274 (CAS, Richardson,Bot. App. 737. 1823. Thermia NY, RM, UC, WS, WTU); 1 mi. NE of Walla Walla, rhombifolia(Pursh) Nutt., Gen. N. Amer.P1. flatsnear MillCreek, 2 May 1936, Moore 23 (UC, WS). 1: 282. 1818. Scolobus rhombifolius(Nutt. WhitmanCo., 1 mi. N of Colfax,18 May 1935, Hitch- ex Pursh) Raf., Amer. MonthlyMag. Crit. cock & Samuel 2594 (OSC, RM); Kamiack, 15 June 1893, Moore 1489 (US, WTU); Dusty, 3 mi. W of Rev. 4: 193. 1819. Drepilia rhombifolia Colfax, 14 May 1922, Parker 355 (NY, OSC, POM, (Nutt. ex Pursh) Raf., N. Fl. 2: 53. 1837. WS). TYPE: U.S.A. NorthDakota: MercerCo., near FortMandon, Bradbury 1810 (lectotype,se- This variety differsfrom Thermopsis Montana lected here, PH, A. B. Lambertherbarium). var. Montana in its relatively robust habit, moderately branched stems, and broader leaflets and 7Thermopsisarenosa Nelson, Bot. Gaz. 25: 276. 1898. bracts. Thermopsisrhombifolia var. arenosa (Nelson) Lar- isey,Ann. Missouri Bot. Gard. 27: 251. 1940.TYPE: Plants similar to Thermopsis Montana var. ova- U.S.A. Wyoming:Albany Co., LaramieHills, 16 ta also occur as populational variants in Arizona, June1897, A. Nelson3182 (holotype,RM; iso- most of New Mexico, parts of southern Utah, and types,ND, RM, [2 sheets]UC, US). occasionally in Colorado, as discussed under T. 7Thermopsisannulocarpa Nelson, Bull. Torrey Bot. Club Montana var. Montana. Our treatmentis like that 26: 239. 1899. 7Thermopsisrhombifolia var. an- nulocarpa(Nelson) L. Williams,Ann. Missouri Bot. of Isely (1981), who positioned variety ovata within Gard.23: 450. 1936. TYPE: U.S.A.Wyoming: Car- his broad T. rhombifolia, except that we would bonCo., FerrisMountains, vicinity of FortSteele, include his T. macrophylla var. hitchcockii within "amongthe rocks on thenaked slopes near their T. Montana var. ovata. We do not believe that summits,"25 July1898, Elias Nelson4971 (holotype,RM; isotype,NY). the Pacific coast plants (= T. macrophylla var. hitchcockii, sensu Isely) differappreciably from Larisey (1940) designateda Nuttallcollection plants of the interior montane regions that Isely (PH) as the typeof Thermopsisrhombifolia; how- places within T. rhombifolia. The disjunct distri- ever, Pursh citedonly a Bradburycollection, list- bution of the Pacific coastal populations is thought ing Cytisusrhombifolius Nutt. as a nomennudum to represent founderpopulations, perhaps dispersed fromFraser's Catalogue withoutciting a Nuttall by periodic floodingevents and subsequent estab- collection. lishment of propagules from populations along the upper Columbia and Snake River drainages. One Plantsdwarf but robust, 1.2-3 dm tall,glabrate may assume that these northernPacific coast pop- to appressed pubescent;stems slender,erect, or ulations are more closely related to the southern ascending, solitary or clustered, few-branched, populations of T californica, as suggested by Ise- arising from a woody rootstock or rhizomes; ly's treatment. Although both taxa have upright branchesascending at 25-50?, weaklyzigzagged. pods, the distribution of T. californica is inter- Leafletselliptic to obovateor narrowlyelliptic, 1.7- rupted in Oregon by the quite distinctive T. grac- 3.3 cm long, 0.6-2 cm wide, acute to obtuse, ilis, which has divergentpods. All of the specimens sometimesshort mucronate, appressed pubescent of T. Montana var. ovata that we have examined to sparselytomentose or glabratebelow, glabrous (including the Pacific coastal populations) have above, lateralveins 5-7 pairs, conspicuouslynet- strictly erect pods, yet have other characters of veinedbeneath; petioles 0.8-2.2 cm long; stipules the Rocky Mountain species, namely T. Montana persistent,ovate to verywidely ovate or ellipticto var. Montana. If one accepts the taxonomic con- widelyelliptic, 1-2.6 cm long, 0.5-1.5(-2.2) cm cepts espoused by Isely (1981) in which T divar- wide,cuneate to oblique,acute to obtuseand short icarpa and T. Montana are reduced to varietal mucronate.Racemes 4-12 cm long, with 5-15 entities of a widespread, highly variable T. rhom- flowers,in whorlsof 2-3 or scattered;peduncles bifolia, one is compelled by morphological consid- 1-4 cm long; pedicels 3-7 mm long, glabrateto erations to include both T. gracilis and T cali- denselyappressed pubescent; bracts deciduous, el-

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FIGURE7. Ditribuiono Theropsisrhombfolia

liptic to obovate, 5-9 mm long, 3-5 mm wide. Representativespecimens examined. CANADA. Al- Flowers 1.9-2.1 cm long; calyx (6-)8-10(-12) berta: ca. 26 km NE of Edmonton,1 km SW of Fort mmlong, 4.5-6 mmwide at thelimb, lobes (3-)4- Saskatchewan,20 June 1959, Shetler & Stone 3034 (US); 20 km W of Cochrane,1 July1979, Watton24 5 mm long,as long as or shorterthan tube; wings (COLO, MO, NY, OSC, PH). Manitoba: on side of asymmetricallyoblong-very widely ovate; keel asym- ManitobaHwy. 1 in the sand hills,9 June 1950, Dore metricallyoblong-widely elliptic; ovary glabrateto 10658 (GH, NY). Saskatchewan: CypressHills Proven- appressedpuberulent; ovules 12-14. Pods arcuate cial Park, 1 July1947, Breitung4200 (MO, RM). U.S.A. Colorado: ArapahoeCo., near Denver, 13 May 1917, to annular,divergent, 3-7 cm long, 0.4-0.7 cm Clokey2793 (ARIZ, F, GH, RM, TEX, UC, US). Boulder wide,glabrate to sparselytomentose. Seeds 2-10, Co., 10 mi. NE of Boulder,White Rocks, on Gunbarrel brown,elliptic with a rostrum,3.5-4 mmlong, 2.5 Hill, 18 May 1967, Turner5634 (RM, TEX). Chaffee mm wide.Chromosome numbers 2n = 18. Co., 5 mi. SW of Buena Vista, 13 June 1967, Johnson 80 (MONTU, NCU, TEX). Douglas Co., 2 mi. W of Distributionand ecology (Fig. 7). Central Franktown,27 June1937, Beetle 1477 (CAS, NY, WTU). ElbertCo., 2 mi. N of Simla, 23 June 1937, Ownbey GreatPlains of Canada and the UnitedStates from 1296 (COLO, GH, MO, MONTU, NY, RM, UC, WS). central Alberta and Saskatchewanto northeast- El Paso Co., near Ramah, 25 May 1947, Moldenke ernmostNew Mexico,largely confined to relatively 19401 (PH, RM). JeffersonCo., E of Golden, 8 May xericgrasslands in deep alluvialsoils but extending 1941, Ehlers 8064 (COLO, TEX). Lake Co., Tennessee Fork, 16 June1933, Applegate 8524 (F). LarimerCo., into the lower canyons of the frontrange of the 7 mi.S of FortCollins, 3 May 1965, Irving6511 (TEX). RockyMountains, 1000-2000 m,to 3000 m along Las AnimasCo., N of Mesa de Maya, 28 Apr. 1948, roadsides;flowering May-June. Rogers 5664 (COLO, TEX). LincolnCo., 2.5 mi. E of

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Lincoln,7 May 1987, Falb & Miercort23A (RM). Moffat Ullin,May 1891, Holzinger 5 (GH). MountrailCo., 12 Co., Blue Mountain,8.2 mi. N of U.S. Hwy. 40, 6.5 July1960, Stevens& Moir 2289 (NCU, UC). Slope Co., airlinemi. NNE of Dinosaur,11 June 1971, Holmgren Atkinson1935 (F, NCU, RM, UC). StarkCo., Dickinson, et al. 5162 (CAS, NY, US, WTU). Park Co., Como, 15 Whitman 1949 (US). Ward Co., Haigh 1900 (NY). June1892, Hughes 42 (GH). Saguache Co., 2 mi. S of WilliamsCo., Williston,Stevens 1918 (F). Oklahoma: Villa Grove on U.S. Hwy. 285, 27 May 1968, Turner CimarronCo., N of Mesa, 28 Apr. 1948, Rogers 5664 5774 (TEX). Weld Co., Pawnee Buttes, 1 July1906, (US). South Dakota: ButteCo., 5 mi.N ofBelle Fourche, Dodds 2111 (COLO). Montana: Big Horn Co., road to 23 May 1970, Stephens 38357 (NY). CusterCo., head visitorcenter at YellowtailDam, 16 May 1983, Lichvar of BlocktailGulch, 12 Apr. 1910, Murdoch 4020 (F, 5666 (RM). Blaine Co., Chinook,26 May 1937, Wel- NY). Fall RiverCo., Ochich,8 Apr. 1911, Nelson 2724 lington 1049 (MONTU). Carter Co., Medicine Rocks (F, RM). JacksonCo., Cedar Pass, 17 July1913, Over StatePark, 28 Apr.1986, Mooerset al. 1085 (MONTU). 6113 (COLO). HardingCo., 4 Aug. 1927, Over 17438 Cascade Co., ca. 14 mi. W ofGreat Falls, 11 May 1956, (US); RabbitButte, 1 May 1912, Vesher553 (F). Law- Booth 56224 (GH, MONTU). ChouteauCo., FortBenton, rence Co., near Lead, 17 June 1929, Palmer 37352 June1883, Anderson547 (US). CusterCo., 5 Woolfolk (GH, MO, NY, US). Meade Co., 40 mi. NW of Rapid 1936 (GH, PH, POM, RM). Daniels Co., VonKuster 12 City,9 July1947, Crider& Atkins44 (RM). Pennington (RM). DawsonCo., 5 mi. SW of Glendive,20 May 1965, Co., Black Hills, Harney Peak, 8 July1927, Hayward McMullin 308 (UC). FergusCo., 3 mi. up Fords Creek 1961 (F, RM). PerkinsCo., Lemmon,Halse 1969 (OSC). turnoff, 19 May 1988, Lipscomb 1 (MONTU). Gallatin Utah: DaggettCo., 6 air mi. N of Dutch John,7 June Co., benchgravel, 6 July1910, Spragg 245 (MONTU). 1979, Atwood 7284 (NCU). UintahCo., ca. 14 air mi. GarfieldCo., 15 mi. N of Jordan,13 May 1968, Wood- NW of Vernal along Tridell-DryFork Road, 31 June land 1632 (CAS). GlacierCo., S of Hudson Bay Divide, 1983, Neese 14116 (MONTU, NY). Wyoming: Albany 31 May 1958, Harvey 66750 (MONTU). JudithBasin Co., Laramie Mts., Telephone Canyon, 30 May 1936, Co., Long'sRanch, 8 June1900, Wilcox388 (US). Lewis Ownbey1027 (COLO, MO, MONTU, RM, WS, WTU). and ClarkCo., ca. 4 mi. W of WolfCreek, 2 July1948, CampbellCo., ca. 8 air mi. SE of Reno Junctionat Little Hitchcock17984 (CAS, COLO, GH, MONTU, NY, POM, Thunder Creek, 3 June 1978, Hartman & Dueholm UC, WS). McCone Co., WolfPoint, 19 May 1937, Buck- 5896 (NY, RM). CarbonCo., Como Bluff,25 May 1947, of 546 (MONTU, WTU). MusselshellCo., Roundup,11 Porter4162 (CAS, MO, NY, PH, RM, TEX, UC, WTU). May 1937, Syblon 522 (CAS, RM, UC, WTU). Petro- ConverseCo., near the head of La Prele Creek,27 June leumCo., Flatwillow,28 May 1936, Wickland18(MON- 1950, Porter 5374 (CAS, COLO, GH, MO, NY, RM, TU). Powder River Co., 18 May 1975, Lackschewitz TEX, WTU). Crook Co., 8 mi. NW of Hulett,6 June 5937 (MONTU). PrairieCo., 10 mi. S of Brockway,5 1935, Ownbey 646 (COLO, MO, MONTU, NY, RM, May 1941, Koch 4 (MONTU). RosebudCo., along U.S. UC, WS, WTU). FremontCo., ca. 20 mi. SW of Lander Hwy. 212, ca. 10-15 mi. W of Ashland,18 July1967, alongWyoming Hwy. 28, 2 July1983, Evert5282 (NY, Hess 762 (MO). SheridanCo., Westby, 17 July1927, RM). Goshen Co., sandy hills betweenWheatland and Larsen 20 (GH, MO, PH, RM). Sweet Grass Co., Big Veteran,19 June1953, Porter6235 (RM, WTU). Hot Coulee Creek,Blankinship 1902 (MONTU, WS, WTU). SpringsCo., 20 mi. NW of Thermopolisalong Wyoming TetonCo., 22 mi.SW ofChouteau, Finley 1963 (MONTU); Hwy. 120, 3 July1983, Evert5312 (NY, RM). Johnson Collins,Jones 1904 (UC). Toole Co., Sweetgrass,Recup Co., BighornMts. W of Buffalo,Circle Park, 23 July 1918 (NY). ValleyCo., ForkPeck, 26 May 1919, Rams- 1958, C. L. & M. W. Porter 7562 (CAS, RM, UC, den & Lackschewitz345 (MONTU). WibauxCo., Hodg- WTU). LaramieCo., Pine Bluffs,2.5 mi. S of U.S. Hwy. es, Griffins1936 (MONTU, RM, TEX, WTU). Yellow- 30 and InterstateHwy. 80, 6 June 1969 Cox & Dunn stoneCo., 10 mi. NW of Billingson Lavina Road, 3 June 1091 (MO). NatronaCo., 5 mi. W of PowderRiver, 5 1973, Doles 563 (UT). Nebraska: Box ButteCo., Hem- July1979, Dueholm 7760 (NY, RM). NiobraraCo., 22.5 ingford,1 June 1950, Kiener 24623 (US). Cheyenne air mi. NW ofLusk and 14.5 air mi. SW ofLance Creek, Co., 2 mi. W of Sunol, 18 May 1970, Stephens38111 2 June 1978, Nelson 1591 (RM). Platte Co., 21.5 air (NCU). Deuel Co., RushCreek, Rydberg 1891 (GH, US). mi. N of Guernsey,31 May 1978, Nelson 1540 (NY, Keith Co., E of Cedar Point BiologicalStation, 7 June RM, UC). SheridanCo., 9 mi. S of Sheridan,30 June 1983, Sutherland5598 (NY). KimballCo., 8 mi. SW 1979, Dueholm 7542 (NY, RM). SweetwaterCo., 16 and 1.5 mi.S ofBushnell, 18 May 1970, Stephens38143 mi. W of WyomingHwy. Rt. 789, 15 July1973, Hill (GH, NY). Garden Co., Ash Hollow State Park, W of 985 (RM). WashakieCo., S of Ten Sleep, 25 July1901, Lewellenon U.S. Hwy. 26, 19 May 1976, Churchill Goodding353 (F, GH, MO, NY, RM, US). WestonCo., 7323 (MO, NY). SheridanCo., 13 mi. N of Hay Springs, 3.5 air mi. NW of Osage, 22 May 1984, Marriott6214 5 May 1964, Nixon 6 (RM). Sioux Co., 7.5 mi. S of the (NY, RM). junctionof Nebraska Hwys. 2 & 71 on Hwy. 71, 27 May 1972, Churchill 77 (MO, NY). New Mexico: Larisey (1940) accepted three varieties under Union Co., Capulin MountainNational Monument,17 this species, but all of them were reduced to Ther- July1941, A. & R. A. Nelson 4650 (MONTU). North mopsis rhombifolia var. rhombifolia by Isely Dakota: BillingsCo., Cedar Canyon,9 June1952, Clau- sen 8194 (GH). BurleighCo., Bismarck,Lanterman 1891 (1981). Although Isely's treatment extended the (MO, NY, WTU). Burke Co., Portal, 18 June 1903, parameters of Thermopsis rhombifolia to include Barber 360 (GH). Divide Co., 25 May 1961, Stevens the long-recognized species T. montana and T. 2377 (UC, US). Dunn Co., KilldeerHistorical Site, Ste- divaricarpa, which we do not accept, we agree that vens 1951 (US). GrantCo., PrettyRock NationalWildlife recognition of varieties within our Refuge,23 Aug. 1907, Bell 1197 (UC). Mercer Co., narrower con- MissouriRiver Valley, 8 mi. N of KnifeRiver, 29 June cept of T rhombifolia is not justified. Thermopsis 1941, Stevens 905 (NCU, UC, US). MortonCo., Glen rhombifolia does not appear to intergrade with

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eitherT. divaricarpa or T. montana,but the oc- Vollmer& Beane 146 (CAS, POM, WTU); near Orleans, casionalhybrid or hybridswarm is believedto occur 19 May 1931, Wieslander 7 (RM, UC). SiskiyouCo., 11 mi. N of Somes Bar, shadyforest near Camp Three, betweenat least T. rhombifoliaand T. divaricar- 22 June 1954, Hitchcock20223 (NY, WTU); 9.5 mi. pa, especiallyin AlbanyCounty, Wyoming. The N of KlamathRiver, Cedar Camp Road, 11 July1950, name T. arenosa is probablybased on such hybrid Hoffman3643 (UC); Prospect Hill, 1220 m, 18 July variants(see discussionunder T. divaricarpa). 1950, Tracy 19220 (UC); ProspectHill, 20 July1950, Tracy 19213 (POM, UC, WTU); near State Creek, 12 June 1928, Wieslander129 (JEPS). Countyunknown, 9. Thermopsis robusta Howell,Erythea 1: 109. coast of California,Vasey 1876 (GH). 1893. TYPE: U.S.A. California:"California Thermopsis robasta is a localized, robust en- Coast Range, near the Oregonboundary," T. demic of northwesternCalifornia. Because of its Howell 1884 (holotype,ORE; isotypes,F, habit, large leaflets,and densely pilosulous vestiture GH, NY [2 sheets],US, WS, WTU). it is superficiallysimilar to T. macrophylla. Both Some ofthe apparent isotypes are labeled"Coast Larisey (1940) and Isely (1981) included T. ro- Mts.,Curry Co., Oregon"(GH) and "Oregon,Coast busta withintheir concept of T. macrophylla var. Mountains,fortieth parallel" (F, NY, US, WTU), macrophylla. We believe it is more closely related whileanother is labeled"Oregon, Coast Mountains, to T. gracilis, to judge fromits divergent pods and forty-secondparallel" (WS). We have accepted distribution. the location as given in the protologue(above), whichwould place the actual typelocality in Cal- 10. Thermopsis villosa (Walter)Fernald & B. ifornia,probably Humboldt or SiskiyouCounty to G. Schubert,Rhodora 50: 201. 1948. So- judge fromthe specimenswe have examined. phora villosa Walter,Fl. Carol. 134. 1788. Plantsrobust, 8-18 dm tall,densely tomentose; TYPE: U.S.A. North Carolina: without definite stemsthick, erect, clustered, well-branched, arising localityor date, Walter s.n. (holotype,BM froma woody rootstock;branches ascendingat not seen). Typificationof this plant is thor- 30-500, stronglyzigzagged. Leaflets elliptic to oughlydiscussed by Fernald & Schubertin widelyelliptic or ovate, 6.5-11 cm long, 2.5-6.5 theirtransfer of the species fromSophora to cm wide, acute to acuminate,short mucronate, Thermopsis. tomentose,lateral veins 9-12 pairs, not conspic- Thermopsiscaroliniana M. Curtis,Amer. J. Sci. 44: 80. uously net-veinedbeneath; petioles 2.2-3.5 cm 1843. TYPE: U.S.A. NorthCarolina: Haywood Co., long; stipulespersistent, widely ovate to nearly "in sylvismontanis," M. A. Curtis1839 (holotype, lanceolate, 1.5-5.5 cm long, 0.7-3 cm wide, GH; possibleisotypes, PH, NY). oblique to cuneate, acuminate. Racemes 20-45 Plantsdelicate, 6-18 dm tall,glabrate (densely cm long,with 20-40 flowers,in whorlsof 2-5 or villous inflorescences); stems thick, erect, solitary scattered;peduncles 7-10 cm long; pedicels4-8 or clustered, few-branched, arising from a woody mm long, densely tomentose;bracts persistent, rootstock; branches spreading at 45-60?, moder- ovate,8-14 mmlong, 3-8 mmwide. Flowers 1.6- ately zigzagged. Leaflets elliptic, 4-9.5 cm long, 1.8 cm long; calyx 7-9 mm long, 5-8 mm wide 1.8-4.4 cm wide,acute to obtuse,glabrate above, at the limb, lobes 2-4 mm long, as long as or sparselyvillous below, lateral veins 6-8 pairs,con- shorterthan tube; wingand keel petals asymmet- spicuously net-veined beneath; petioles 2-5(-6) cm ricallyoblong-rhombic; ovary denselytomentose; long; stipulespersistent, elliptic to ovate, 1.5-4.5 ovules 10-13. Pods arcuate, divergent,4.5-6.5 cm long,0.9 to 2.4 cm wide,oblique to cuneate, cm long, 0.4-0.5 cm wide, densely tomentose. acute to obtuse. Racemes 15-50 cm long, with Seeds 5-10, brown,reniform without a rostrum, 10-50 flowers,scattered or in whorlsof 2-3; pe- 3.5-4 mm long, 2 mm wide. duncles 6-13 cm long; pedicels 2-3 mm long, Distributionand ecology(Fig. 1). Knownonly denselyvillous; bracts deciduous, widely ovate to fromnorthwestern California, Humboldt and Sis- widely obovate, 5-7 mm long, 5-6(-8) mm wide. kiyou cos., along ridgetops,150-1500 m; flow- Flowers1.7-1.9 cm long; calyx 7-8 mmlong, 5- eringMay-June. 6.5 mm wide at the limb, lobes 2-3 mm long, much shorterthan tube; wing and keel petals asym- Specimensexamined. U.S.A. California: Hum- metricallyoblong-rhombic; ovary velutinous; ovules 59- boldtCo., ca. 12 mi.SW ofOrleans, Bee Mt.,Quick 12-16. Pods straightto slightlyarcuate, erect, 4- 42 (CAS, COLO, TEX); 11 mi. fromOrleans on Bar ShantyRoad, 6 July1956, Tillett620 (POM); E of 5.5 cm long,0.4-0.6 cm wide,densely tomentose Korbel,Bald Mt., on trailto Blohn'scabin, June 1951, to villous.Seeds 7-12, olive brown,elliptic with a

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900 850 80750 700 650

350

300 mi.

FIGURE 8. Distributionof Thermopsis villosa: collectionsthought to be native(closed circles);collections thought to be introducedor cultivated(open circles). rostrum, 3-3.5 mm long, 1.8-2 mm wide. Chro- Northampton,railroad right-of-way, Ahles 78850 (TENN). mosome number, 2n = 18. MiddlesexCo., Framingham,Pike exit to Massachusetts Hwy. 9, 27 June 1975, Ahles 80370 (NEBC); Wil- Distribution and ecology (Fig. 8). Native to mington,W side of InterstateHwy. 93, 22 June 1986, eastern Tennessee, western North Carolina, north- Boufford& Wood 23650 (CAS, NEBC, MO, PH). New Hampshire: CheshireCo., Boufford6134 eastern Alabama, northern Georgia, and closely (MO). New Jersey: Sussex Co., Edwards 1971 (NCU). Union Co., adjacent South Carolina, cultivated northward to Moldenke et al. 30109 (ARIZ, LL). North Carolina: Maine, scattered localized populations in open ar- AveryCo., 0.5 mi. N of Cranberryon U.S. Hwy. 19E, eas of deciduous oak woodlands, mostly 1000- 17 June 1958, Ahles & Duke 43574 (NCU). Cherokee 1600 m; floweringMay-June(-July). Co., NotlelyRiver near NorthCarolina Hwy. 60, 6 June 1953, Radford& Haesloop 7187a (NCU); Rugel 1842 (NY). Clay Co., base of Doe Knob, Buck Creek,24 July Representativespecimens examined. U.S.A. Ala- 1951, Godfrey51485 (TENN, US); NE of ChunkyGal bama: DeKalb Co., 6 mi. N of Dawson, 2 July1943, Mt., along Buck Creek Road, 26 June 1970, Leonard Harper 3941 (F, GH, MO). Georgia: FanninCo., near et al. 3314 (NCU). Haywood Co., 1.6 mi. N of Cove Three ForksMountain, 28 June1983, Coile et al. 3736 Creek,5 June1958, Ahles & Duke 42242 (NCU). Hen- (RM), 23 July1909, Smith2475 (F). RabunCo., Tallulah dersonCo., ca. 1 mi. N of Lake Logan, 12 June 1964, River, Small 1893 (F, GH, MO, NY, PH, UC, US). Wilbur7142 (GH, UC). JacksonCo., near Soco Falls, 6 Union Co., BrasstownBald Mt., 15 July1908, Howell July1940, Hood 109 (MO); Soco Gap, ca. 3 mi. W of 393 (US). Maine: Hancock Co., Reef Point Gardens, U.S. Hwy. 19, 1 July1954, Sargent 6851 (POM, RM, Spaulding 582 (GH). Maryland: Prince GeorgesCo., WTU). Macon Co., near Highlands,Biltmore Herbarium Hermann18409 (RM). Massachusetts: HampshireCo., 1332b (F, GH, MO, NCU, NY, US); 3 mi.SE ofSassafras

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Gap, 18 June 1969, Leonard & Russ 2512 (COLO, ALSTON, R. E. & B. L. TURNER. 1963. Natural hy- NCU, POM, TENN, UC, WTU); 6.7 mi. W of Wayah bridizationamong four species of Baptisia (Legu- Valley Ranch, ArrowoodGlade, 29 June 1970, Turner minosae).Amer. J. Bot. 50: 159-173. 6072 (F, GH, NY, RM, TEX, WTU). MadisonCo., Ahles BALANDRIN, M. F., E. F. ROBBINS & A. D. KINGHORN. & Duke 46380 (NCU, NY). Swain Co., along stream, 1982. Alkaloiddistribution in some species of the 12 June 1982, Churchill 82698 (RM); 1.3 mi. from Papilionaceoustribes Thermopsideae and Genisteae. NorthCarolina Hwy. 28 on Mica Knob Road, 6 June Biochem.Syst. & Ecol. 10(4): 307-311. 1956, Bell 3200 (GH, NCU). TransylvaniaCo., U.S. BISBY, F. A. 1981. Genisteae(Adans.) Benth. Pp. 409- Hwy. 276 near Pink Beds, 23 Sep. 1957, Freeman 425 in R. M. Polhill& P. H. Raven (editors),Ad- 57819 (NCU); along motorroad in Pink Beds, Ripper vances in Legume Systematics.Royal BotanicGar- 307 (RM). Pennsylvania: McKean Co., Fogg & Wher- dens, Kew. ry 9818 (PH). Tennessee: Blount Co., Cades Cove, BREWER, W. H. & S. WATSON. 1880. Polypetalae.In: near Crib Gap, Hoss 1954 (TENN). CumberlandCo., Botanyof California,Vol. 1. Welch, Bigelow,Cam- Adams 1962 (NCU, TENN, US). Davidson Co., NW bridge,Massachusetts. Nashville,at intersectionof Marrowbone and Eaton'sCreek BROWN, R. 1811. P. 3 in W. T. Aiton(editor), Hortus Roads, 17 June1968, Kral 31489 (NCU, TENN). Maury Kewensis,ed. 2, Vol. 3. Longman, Hurst, Rees, Co., just S of and above Rockdaleon U.S. Hwy. 43, 26 Orme & Brown,London. June 1970, Kral 39741 (GH, MO, NY); Harpethhills CHEN,C. J., X. Y. ZHU & Y. M. YUAN. 1992. Cyto- near Nashville,Gattinger 1879 (GH, MO, NY, US). Polk logicalstudies on thetribe Thermopsideae (Fabaceae) Co., ca. 1.5 km SE of summitof Big Frog Mt., 14 July I. Report on karyotypesof eleven species of four 1983, Z. E. & J. B. Murrell389 (TENN). Vermont: genera. Cathaya 4: 103-116. WindhamCo., Charette3345 (MO), and 3349 (NEBC). CRANMER, M. F. & B. L. TURNER. 1967. Systematic Virginia: Roanoke Co., Uttal 6228 (NCU). significanceof lupine alkaloidswith particular reference to Baptisia (Leguminosae).Evolution 21: 7Thermopsisvillosa is a distinctive,stiffly erect 508-517. species of the southernAppalachian ridges and CRISTOFOLINI, G. 1987. Serologicalrelationships among upper valleys.Czefranova (1970) treatedit (as T Sophoreae, Thermopsideae,and Genisteae (Faba- ceae). Bot. J. Linn. Soc. 94: caroliniana) as the only member of series Caro- 421-432. & L. FEOLI-CHIAPELLA. 1984. linianae.It is Originand di- superficiallysimilar to the Californian versificationof the Genisteae. Webbia 38: 105-122. T macrophylla, as noted by Gray (1848a), in CZEFRANOVA, Z. 1970. Seriesnovae generis Thermopsis having denselyvillous inflorescences and closely R. Br. NovitatesSystematicae Plantarum 7: 213- compactederect pods but differsmarkedly from 216. DEMENT, W. A. & T. J. MABRY. 1975. that species in its much more slender,strict habit Biologicalim- plicationsof flavonoidchemistry in Baptisia and and glabratefoliage. Thermopsis.Biochem. Syst. & Ecol. 3: 91-94. Sporadic,locally naturalized populations of this FAUGERAS, C. & R. PARIS. 1968. Chemiotaxinomiedes taxonoccur from Virginia, northeastward to Maine. Papilionacees-Genistees.Bull. Soc. Bot.France 1965: Fernald (1950) did not account for T villosa in 75-102. FEDEROV, A. A. (editor). 1969. ChromosomeNumbers his treatment of 7Thermopsisfor the northeastern of FloweringPlants. Acad. Sci. USSR, Moscow. UnitedStates, nor did he commenton the clearly FERNALD, M. L. 1950. Gray's Manual, 8th ed. Amer- naturalized populations of this species outside of its ican Book Company,New York. naturaldistribution (Fig. 8). GILLETT, J. M. 1975. In: A. Lhve (editor),IOPB Chro- mosomeNumber Reports L. Taxon 24: 671-678. The species was taken into cultivation early on. GRAUSTEIN, J. E. 1967. Thomas Nuttall,Naturalist, Gray (1848a) stated "I raised the plant in the Explorationsin America,1808-1841. HarvardUniv. Cambridge Botanic Garden (where it is perfectly Press, Cambridge. hardy) fromseeds . .. communicated by Mr. Buck- GRAY, A. 1848a. Chloris Boreali-Americana.Mem. Amer.Acad. Artsn.s. 3(1): 1-56. ley." No doubt the species is still retained in the 1848b. A Manual ofthe Botany of the North- occasional garden or has persisted in gardens long ern UnitedStates. JamesMonroe, Boston. vacated. One collection is known from Hawaii GREUTER, W., ET AL. 1988. InternationalCode of Bo- (Oahu, "Mauka of Kawaiiki ditch intake," 30 Apr. tanicalNomenclature. Regnum Veg. 118. 1939, Degener 17363, GH). We also believe that HARBORNE, J. B. 1969. Chemosystematicsof the Le- guminosae.Flavonoid and isoflavonoidpatterns in the numerous collections near Nashville, firstmade the tribeGenisteae. Phytochemistry 8: 1449-1456. by Gattinger in the Harpeth hills in 1875 (MO, HOWELL, J. T. 1957. The Santa Ynez Thermopsis. TENN, US), may also be locally naturalized pop- Leafl. W. Bot. 7(6): 158-159. ulations. ISELY, D. 1978. New varietiesand combinationsin Lotus, Baptisia, Thermopsis,and Sophora (Legu- minosae).Brittonia 30: 466-472. LITERATURECITED - 1981. Leguminosaeof the UnitedStates, III. ABRAMS,L. 1944. Fabaceae. Pp. 480-627 in An Il- SubfamilyPapilionoideae: Tribes Sophoreae, Poda- lustratedFlora of the Pacific States, Washington, lyrieae,and Loteae. Mem. New York Bot. Gard. Oregon,and California,Vol. 2. StanfordUniv. Press, 25(3): 1-264. Stanford,California. LARISEY, M. M. 1940. A revisionof the North American

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speciesof thegenus Thermopsis.Ann. Missouri Bot. SHISHKIN, B. K. 1945. Thermopsis.Pp. 30-36 in V. Gard. 27: 245-258. L. Komarov (editor),Flora of the USSR, Vol. 11. LEDINGHAM,G. F. 1957. Chromosomenumbers of some [Englishtranslation.] Israel Programfor Scientific SaskatchewanLeguminosae with particular reference Translation.Jerusalem. 1971. to Astragalus and Oxytropis.Canad. J. Bot. 35: ST. JOHN,H. 1941. Notes on NorthAmerican Ther- 657-666. mopsis.Torreya 41: 112-115. LINK, H. F. 1821. Icones PlantarumSelectarum. Pub- TAYLOR, R. L. 1967. In: A. L6ve (editor),IOPB Chro- lishedby the author,Berlin. mosomeNumber Reports XIII. Taxon 16: 445-461. LINNAEUS,C. 1753. Species Plantarum,ed. 1. Holmiae TORREY, J. 1857. Explorationsand surveysfor a rail- [Stockholm],(impensis Laurentii Salvii). road. PacificRailroad Rep. 4: 81. LOVE, A. & D. LOVE. 1982. In: A. Love (editor),IOPB . 1859. Botany of the boundary.In: W. H. ChromosomeNumber Reports LXXV. Taxon 31: Emory(editor), Rep. U.S. Mex. Bound. Surv., Vol. 342-368. 2, pt. 1. A.O.P. Nicholson,Printer, Washington. MESICEK, J. & J. SOJAK. 1969. Chromosomecounts of & A. GRAY. 1840. A Flora of NorthAmerica. some Mongolianplants. Folia Geobot. Phytotax.4: Wiley & Putnam,New York. 55-86. TURNER,B. L. 1967. Plantchemosystematics and phy- NUrTALL, T. 1818. The Genera of NorthAmerican logeny.Pure Appl. Chem. 14: 189-213. Plants. Printedfor the authorby D. Heart, Phila- . 1981. Thermopsideae.Pp. 403-407 in R. delphia. M. Polhill & P. H. Raven (editors),Advances in POLHILL, R. M. 1976. Genisteae(Adans.) Benth.and Legume Systematics.Royal BotanicGardens, Kew. relatedtribes (Leguminosae). Pp. 143-368 in V. H. . 1982. Review of: D. Isely, Leguminosaeof Heywood(editor), Botanical Systematics, Vol. 1. Ac- the United States. III. SubfamilyPapilionoideae: ademic Press, New York. Tribes Sophoreae, Podalyrieae,Loteae. Syst. Bot. PROBATOVA,N. S. & A. P. SOKOLOVSKAYA. 1981. Ka- 7(3): 350-352. riologicheskoeissledovanie sosudistykh rastenij os- WILBUR, R. L. 1963. The Leguminousplants of North trovovDal'nevostocnogo gosudarstvennogo morsko- Carolina.Technical Bulletin No. 151. NorthCarolina go sapovednika.Sb. CvetkayeRastenija Ostrovov AgriculturalExperiment Station. Raleigh, North Car- DalnevostochnogoMorskogo Sapovednika. Pp. 92- olina. 114. WILLDENOW, C. L. 1799. Species Plantarum.Berlin RADFORD, A. E., H. E. AHLES & C. R. BELL. 1968. (impensisG. C. Nauk). Manual ofthe Vascular Flora ofthe Carolinas.Univ. YEH, M. S., H. YUASA & F. MAEKAWA. 1986. Chro- NorthCarolina Press, Chapel Hill, NorthCarolina. mosomenumbers in theLeguminosae. Sci. Rep. Res. RAFINESQUE,C. S. 1819. J. Phys. Chim. Hist. Nat. Inst. Evol. Biol. 3: 57-71. Arts.89: 259. ZHUKOVA,P. G. 1967. Karyologyof some plants,cul- SALATINO,A. & 0. R. GOTTLIEB. 1980. Quinolizidine tivatedin the Arctic-AlpineBotanical Garden. Pp. alkaloidsas systematicmarkers of the Papilionoideae. 139-149 in N. A. Arorin(editor), Plantarum Zonam Biochem.Syst. & Ecol. 8: 133-147. PolaremTransportatio. II. Leningrad. & . 1983. Chemogeographicalevo- lutionof quinolizidinesin Papilionoideae.P1. Syst. Evol. 143: 167-174.

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