ae HARVARD UNIVERSITY Library of the Museum of Comparative Zoology Tlie Great Basin Naturalist VOLUME 38, 1978 Editor: Stephen L. Wood Published at Brigham Young University, by Brigham Young University TABLE OF CONTENTS Volume 38 Number 1 - March 31, 1978 Endangered and threatened plants of Utah: A reevaluation. Stanley L. Welsh 1 Plants of Navajo National Monument. J. D. Brotherson, G. Nebeker, M. Skougard, and J. Fairchild 19 Environmental interaction in summer algal communities of Utah Lake. Mark C. Whiting, Jack D. Brotherson, and Samuel R. Rushforth 31 Basidiomycetes that decay junipers in Arizona. IL R. L. Gilbertson and J. P. Lindsey . 42 Pinus longaeva in the Stansbury Mountains, Utah. P. A. Kay and C. G. Oviatt 49 Differential parasitism of Peromyscus maniciilatus and Peromyscus truei by Cuterebra larvae. Jeffrey B. Llewellyn 51 An annotated list of the aquatic insects of southeastern Idaho, Part IIL Ephem- eroptera. Robert L. Newell and G. Wayne Minshall 55 Observations on the prey and nests of seven species of Cerceris (Hymenoptera: Sphe- cidae). Howard E. Evans and William L. Rubink 59 Vertebrate fauna of a radioactive leaching pond complex in southeastern Idaho. Douglas K. Halford and Jere B. Millard 64 New synonymy, new species, and taxonomic notes of North American Pityophthorus (Coleoptera: Scolytidae), Part III. Donald E. Bright 71 Fluctuation of a marsh habitat and the reproductive strategy of the Yellow-headed Blackbird. Roger J. Lederer 85 Hosts of juniper mistletoe at Walnut Canyon National Monument, Arizona. Frank G. Hawksworth and Robert L. Mathiasen 89 Allelopathic effects of Bur Buttercup tissue on germination and growth of various grasses and forbs in vitro and in soil. Bruce A. Buchanan, K. T. Harper, and Neil C. Frischknecht 90 New species of Nearctic Neoperla (Plecoptera: Perlidae), with notes on the genus. Bill P. Stark and Richard W. Baumann 97 Morphometry of sculpins (Coitus) in the Clearwater drainage, Idaho. O. Eugene Maughan 115 Correction: Rhamnus cathartica L. is not Primus nigra Ait. from Alberta. R. Keith Shaw 123 Number 2 - June 30, 1978 Kaiparowits flora. Stanley L. Welsh, N. Duane Atwood, and Joseph Murdock 125 Range extension of Onychomys torridus longicaudus (Rodentia: Cricetidae) in north- western Nevada. Mark D. Engstrom and Eugene D. Fleharty 180 The reproductive ecology of the Tahoe sucker, Catostomus tahoensis, in Pyramid Lake, Nevada. Joseph L. Kennedy and Paul A. Kucera 181 Primary productivity in meromictic Big Soda Lake, Nevada. R. P. Axler, R. M. Gersberg, and L. J. Paulson 187 Current status of cutthroat trout subspecies in the western Bonneville Basin. Terry J. Hickman and Donald A. Duff 193 Reproductive biology of the tui chub, Gila bicolor, in Pyramid Lake, Nevada. Paul A. Kucera 203 Distributional records for the Colorado flora. D. Wilken, W. Harmon, C. Feddema, and K. Harrington 208 A comparative analysis of the diatom flora on the snail Ampullaria cuprina from the Goshen Ponds, Utah. Larry L. St. Clair, Lorin E. Squires, and Samuel R. Rush- forth 211 Growth of Dipodomijs ordii (Rodentia: Heteromyidae). H. Duane Smith, Gary H. Richins, and Clive D. Jorgensen 215 Similarity between pronghorn and mule deer fecal pellets. Mark K. Johnson and James G. MacCracken 222 Number 3 - September 30, 1978 Utah flora: Fabaceae (Leguminosae). Stanley L. Welsh 225 Number 4 - December 31, 1978 Effect of irrigation on survival of third-stage Haemonchus contortus larvae (Nema- toda: Trichostrongylidae). Graham R. Bullick and Ferron L. Andersen 369 Mountain ants of Nevada. George C. Wheeler and Jeanette Wheeler 379 New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), Part VII. Stephen L. Wood 397 New range and new subspecies for the snake Eridiphas sleveni. John R. Ottley and Wilmer W. Tanner 406 A new subspecies of the snake Lichantira trivirgata from Cedros Island, Mexico. John R. Ottley 411 Vertical distribution of adult fish in Pyramid Lake, Nevada. Steven Vigg 417 An addition to the amphibian fauna of California. David A. Mullen and Robert C. Stebbins 429 PUptors of the Uinta National Forest, Utah. James A. Mosher, Clayton M. White, Jo- seph R. Murphy, and M. Alan Jenkins 438 Two new species of fleas of the genus Meringis (Siphonaptera: Hystrichopsyllidae). Richard B. Fads ' ' 447 The effects of radiation from uranium mill tailings on Tradescantia. E. A. Rechel and W. F. Campbell 456 Contributions to the biology of the woundfin, Pkigopterus argentissimus (Pisces: Cyprinidae), an endangered species 463 An ecological and taxonomic investigation of Peromijscus maniculatus serratus in Idaho. Thomas L. Thurow 469 The eye fluke disease (Diplostomatosis) in fishes from Utah. Richard A. Heckmann and James R. Palmieri 473 Bite of wolf spider. Dorald M. Allred 478 Index to Volume 38 479 HE GREAT BASIN NATURALIST Iume38 No. 1 March 31 . 1 978 Brigham Young University ■>. COMF". ZOOU, JAN 1 Bt979 UNivsiRttrrv GREAT BASIN NATURALIST MEMOIRS Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agri- cultural Sciences; Ernest L. Olson, Director, Brigham Young University Press, Univer- sity Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only previously unpublished manuscripts of less than 100 printed pages in length and pertaining to the biological natural history of western North America are ac- cepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manu- scripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist is $12 (outside the United States $13). The price for single numbers is $4 each. All back numbers are in print and are available for sale. All matters pertaining to the purchase of subscriptions and back numbers should be directed to Brigham Young University, Life Science Museum, Pro- vo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same of- fice at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts and other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back cover. 10-78 650 34146 ISSN 0017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 38 March 31, 1978 No. 1 ENDANGERED AND THREATENED PLANTS OF UTAH: A REEVALUATION Stanley L. Welsh' Abstract.— Endangered and threatened plants of Utah are reevaluated. The lists are brought up to date, and species now known to be common are recommended to be removed from the proposed lists. Additional species are proposed to be added, including the new species, Astragalus montii Welsh. Passage of the Endangered Species Act of 1973 (Public Law 23-205) provided the le- gal basis for establishment of lists of endan- gered and threatened plant species. Such lists were prepared under the direction of the Smithsonian Institution, and were pub- lished in the Federal Register (40: 27824- 27924. 1975; 41: 24524-24572. 1976). These lists are preliminary and were based on best information available at the time. Much in- formation has been developed since those lists were published, and since the work by Welsh, Atwood, and Reveal (1975) was completed in summary form. This paper is written in an attempt to evaluate the status of rare plants in Utah based on current best information. Several plant species were placed on initial listings which are known to be commonplace. Oth- ers were omitted by error, oversight, or lack of information. Taxa have been described as new which were unknown at the time lists were first compiled. All of these factors must be considered, and the lists should be maintained to be as current as possible. Constraints placed on federal land man- agement agencies provide justification for continued work on the status of rare plants judged to be categorized as either endan- gered or threatened. All such agencies must consider the presence of such plant species before any land use planning can be accom- plished. This constraint applies to all devel- opments which take place on federally con- trolled lands. Information on the nature of endangered and threatened plants in Utah must be ex- plored on a continuous basis, since stability of plant populations is an ideal only; they fluctuate with the many variables of cli- mate, biotic, edaphic, and human devel- opmental factors. The philosophy of human- ity wishing to preserve plant species of unknown value has been questioned by many people. Examples of construction projects being halted when endangered plant or animal species have been found serve to fuel a controversy in our utilitarian society. Idealistically, industrial and other kinds of development should be able to pro- ceed without regard to any living species— whether plant or animal. Since that is not possible in many instances, then which should give way? A "three worms" argu- ment can be brought to focus. Human en- terprise should not be halted, or even se- riously modified because of the presence on the site of anything as insignificant as "three worms." Therein lies the problem. Our basic ignorance of the nature and role 'Department of Botany and Range Science, and Life Science Museum, Brigham Y'oung University, Provo, Utah 84602. Great Basin Naturalist Vol. 38, No. 1 of rare organisms in the ecosystem, and their place in the lives of mankind, should cause us to allow sufficient time to explore the possibilities. The fact that extinction is a reasonable practice in nature is pointed to by knowl- edgeable people as an excuse for the reduc- tion of those species encountered in the course of human expansion onto the natural plant communities of the earth. That plant life has been retreating because of utiliza- tion of lands for agriculture, cities and towns, industrial sites, thoroughfares, and other ancillary features of our culture is hardly in doubt. That certain plant species have been pushed to the point of extinction is apparent, but the question remains as to whether extinction by natural processes and that accelerated by our actions represent two kinds of events. Only the end results are the same. Natural extinction represents the removal, in a selective manner, of or- ganisms which, commonly because of condi- tions of the environment, could not contin- ue to reproduce in sufficient numbers as to maintain a viable population. Forced extinc- tion involves wholesale removal or modifi- cation of indigenous plant communities, re- sulting in the death of populations. This latter type of extinction can involve both inadvertant and purposeful exploitation and has been justified on the basis of necessity— the greatest good for the largest number of human inhabitants. Exploitation without un- derstanding of natural processes has led to catastrophes affecting both human and biot- ic populations. Now, we are at a point in civilization where an awareness has developed that liv- ing things represent a finite resource which cannot be pressed endlessly into a shrinking environment without loss of substantial numbers of species. Lands considered as good for little except limited grazing in the first century of our occupation of Utah are now important as industrial sites and for re- source exploitation, recreation, and other uses. Prior to the establishment of the Endan- gered Species Act, expansion onto wild lands took place without regard to rare or unusual plants. It was easy to work without restrictive regulations, regardless of the con- sequences to plant populations. The cost of such a mode of operation was presumed to be advantageous in our competitive society, but was potentially high in cost of a basic natural resource— our rare plant species. They became endangered from human ex- pansion for two reasons: ignorance of their existence and of their importance. Given the opportunity of passing the costs of expansion on to future generations, or of bearing those costs at the present time, the present generation will likely de- fer the costs to the latter. Cost is not in monies alone, but involves all of our re- sources, including rare plants of unknown value. The Endangered Species Act pro- vides an advocate for present and future generations, allowing all a chance to make determinations and to enable examination of rarities provided in our natural system. The costs of protection will be borne by all gen- erations. Guaranteed is the possibility for pursuit of knowledge about rare plants, now and in the future. All of our indigenous plant species are a part of our heritage. We have driven some vegetative types and spe- cies to the edge of extinction. The loss of a few square kilometers of some widespread species can be tolerated because of the large extent of that type. However, the loss of a few square meters of a rare plant pop- ulation can result in extinction. The Endan- gered Species Act stands to prevent extinc- tion. The lands have become increasingly valu- able, and the indigenous plants will retreat to still smaller areas as demands are made for greater utilization of Utah lands. Rarity of plant species is a concept which requires study. Utah is a large state with a great many habitat types in a vast eleva- tional range. Botanical exploration of its lands was begun seriously only a century ago, and students of its flora have been rel- atively few. Only about a half dozen botan- ists have ever collected more than 5,000 numbers each from the confines of the state. Catalogues of the flora have always been incomplete because of the poorly known nature of the plant species. Numer- ous taxa have been described since 1950, and likely many more remain to be discov- ered. When a plant species is first recog- March 1978 Welsh: Endangered Utah Plants nized, its total range is likely known from a small to a very small area. Because of the small size of the known range, the plant is judged to be rare. Further exploration fre- quently demonstrates that the total range is larger than initially indicated. However, there are those species which occupy nar- row ranges, usually on peculiar substrates, which are actually rare. At the present, the knowledge of Utah plant species is only moderately developed. The distribution of common plants is fairly well documented, but the rare and pur- portedly rare plants require much additional work. The following summary lists represent the current information available on both proposed and candidate endangered and threatened species of plants in Utah. Endangered Plant Lists and Proposed Modifications The current list of proposed endangered plants is presented in Table 1. Some 68 spe- cies, including 4 which are possibly extinct, are included. Evaluation of previous data and of that taken from recent collections in- dicates that the list should be modified. It is therefore proposed that some 12 species be downgraded to the status of threatened and Table 1. Current list of endangered plants as cited in Federal Register publications (1975, 1976), their locality by county in Utah, and recommendations of status based on present data. Endangered Plants Locality Recommendations Allium passeyi Holmgr. & Holmgr. Arctomecon humilis Coville Astragalus cronquistii Bameby Astragalus desereticus Bameby Astragalus detritalis M.E. Jones Astragalus hamiltonii C.L. Porter Astragalus harrisonii Bameby Astragalus iselyi Welsh Astragalus lentiginosus Dougl. var. chartaceus M.E. Jones var. ursinus A. Gray Astragalus loanus Barneby Astragalus lutosus M.E. Jones Astragalus malacoides Bameby Astragalus minthorniae (Rydb.) Jeps. var. gracilior (Bameby) Bameby Astragalus pardalinus (Rydb.) Bameby Astragalus perianus Bameby Astragalus saurinus Bameby Astragalus serpens Bameby Castilleja aquariensis N. Holmgr. Castilleja revealii N. Holmgr. Cryptantha breviflora (Osterh.) Payson Cryptantha grahamii Johnst. Cryptantha johnstonii Higgins Cryptantha jonesiana (Payson) Payson Cryptantha ochroleuca Higgins Cuscuta warneri Yunker Cycladenia humilis Benth. var. jonesii (Eastw.) Welsh & Atwood Cymopterus minimus (Mathias) Mathias Echinocereus engelmannii (Parry) Lemaire var. purpureus L. Benson Box Elder no change Washington no change San Juan no change Sanpete (PoEx) no change Duchesne, Uintah downgrade to T Uintah downgrade to T Wayne no change Grand, San Juan no change Sevier, Sanpete, remove from list Juab, Summit, Daggett Iron (?) no change Garfield, Piute, remove from list Sevier, Wayne Uintah no change Kane, Garfield downgrade to T taxonomic synonym of Astragalus ensiformis M.E. Jones Emery, Garfield, remove from list Wayne Garfield, Piute downgrade to T Uintah downgrade to T Garfield, Iron, remove from list Piute, Wayne Garfield no change Garfield no change Daggett, Uintah remove from list Duchesne Uintah downgrade to T Emery no change Emery downgrade to T Garfield no change Millard (PoEx) no change Grand, Emery no change Iron no change Washington no change Great Basin Naturalist Vol. 38, No. 1 Table 1.— contd. Endangered Plants Locality Recommendations Erigeron flagellaris Gray var. trilohatus Cronq. Erigeron kachinensis Welsh & Moore Erigeron nmguirei Cronq. Erigeron religiosus Cronq. Erigeron zionis Cronq. Eriogonum ummophilum Reveal Eriogonum aretioides Bameby Eriogonum corymbosum Benth. var. davidesei Reveal var. revealianum (Welsh) Reveal Eriogonum cronquistii Reveal Eriogonum ephedroides Reveal Eriogonum humivagans Reveal Eriogonum hylophilum Reveal & Brotherson Eriogonum intermontanum Reveal Eriogonum lancifolium Reveal & Brotherson Eriogonum loganum A. Nels. Eriogonum smithii Reveal Eriogonum zionis J.T. Howell var. zionis Festuca dasyclada Hackel Gilia caespitosa A. Gray Glaucocarpum suffrutescens (Rollins) Rollins Hermidium alipes S. Wats. var. pallidum C.L. Porter Heterotheca jonesii (Blake) Welsh & Atwood Lepidiu7n barnebyanum Reveal Lygodesinia grandiflora (Nutt.) Torr. & Gray var. stricta Maguire Parthenium ligulatum (M.E. Jones) Bameby Pediocactus sileri (Engelm.) L. Benson Penstemon concinnus Keck Penstemon garrettii Pennell Penstemon grahamii Keck Penstemon nanus Keck Phacelia argillacea Atwood Phacelia indecora J.T. Howell Phacelia mammillariensis Atwood Physaria grahamii Morton Psoralea epipsila Bameby Ranunculus acriformis A. Gray var. aestivalis L. Benson Sclerocactus glaucus (K. Schum) L. Benson Sclerocactus wrightiae L. Benson Townsendia aprica Welsh & Reveal Viguiera soliceps Bameby Zigadenus vaginatus (Rydb.) Macbr. Iron San Juan Emery Kane, Washington Washington Millard Garfield Carbon Piute, Garfield Garfield Uintah San Juan Duchesne Grand, Emery, Uintah Carbon Cache Emery Kane, Washington Emery (?), Sanpete (PoEx) Wayne Uintah Duchesne, Uintah Washington, no chang Garfield Duchesne, Uintah Carbon Duchesne, Emery, Uintah Washington Beaver, Millard Duchesne (?), Wasatch Uintah Beaver, Millard Utah San Juan, Wayne Kane Carbon, Duchesne, Emery, Uintah Kane Garfield (PoEx) Duchesne, Uintah Emery, Wayne Sevier Kane Grand, Kane, San Juan no change no change no change no change no change no change no change no change downgrade to T no change downgrade to T no change no change no change no change no change no change downgrade to T no change no change no change remove from list no change check taxonomic status remove from list no change downgrade to T no change no change downgrade to T no change no change no change remove from list no change no change no change no change no change no change downgrade to T March 1978 Welsh: Endangered Utah Plants some 8 be removed altogether from consid- eration as potentially endangered or threat- ened. Two additional species, now listed as threatened, should be upgraded to endan- gered. Those which should be downgraded include the following: Downgraded Species Astragalus detritalis.— This species is no- where abundant in the Uinta Basin, but is broadly distributed, commonly on gravels from the vicinity of Duchesne east into Col- orado. Astragalus hamiltonii.— The Hamilton milkvetch occurs on the Wasatch Formation west of Vernal and alluvium over a relative- ly large area north to the vicinity of Tridel. Astragalus malacoides.— This milkvetch is known from the vicinity of Fifty Mile Mountain on the Straight Cliffs plateau west to Four Mile Bench, and from the Circle Cliffs portion of Garfield County. Astragalus saurinus.— Named from Dino- saur National Monument, where the plant is locally abundant on saline soils of several geological formations, A. saurinus is also known from as far south as Red Wash and from west of Vernal. Cryptantha grahamii.— The Graham cryp- tantha is known from the environs of Wil- low and Hill creeks and from west of the Green River north of the junction of Wil- low Creek. The total range is somewhat more than 30 miles in length. Cryptantha jonesiana.— This showy cryp- tantha is known from the Sinbad member of the Moenkopi Formation from near the San Rafael River south to the middle of the San Rafael Swell. The plants grow on shaly bar- rens, often in the pinyon-juniper or mixed desert shrub communities. Eriogonum corymbosum var. revealia- num.— This wild buckwheat has been shown to occur through the East Fork of the Sevier River and Otter Creek vicinities in both Piute and Garfield counties. Eriogonum ephedroides.— The ephedra wild buckwheat is rare only in collections. It is known to occur in a rather broad area from western Colorado west to the Green River in Uintah County, Utah. The habitat is confined to Green River Shale. Plants oc- cur in peculiar assemblages with Artemisia pygmaea, Glaucocarpum stiffrutescens, Cnjptantha grahamii, Cryptantha hamebyi, and many other unusual plants. Eriogonum zionis var. zionis.— This pretty wild buckwheat is known from the White Cliffs escarpment in western Kane and eastern Washington counties. The total known range is more than 40 miles long, and consists of numerous disjunct popu- lations. Penstemon concinnus.— The range of this Great Basin endemic occurs in western Mil- lard and Beaver counties. Although rarely collected, the plants are rather widely dis- tributed in the Tunnel Springs, Mountain Home, and Needle ranges, and on the ba- hadas in both Pine and Hamblin Valleys. Penstemon nanus.— The dwarf beard- tongue occurs with P. concinnus, but ex- tends east across the Pine Valley into the Wah Wah Mountains. Zigadenus vaginatus.— The sheathed death camus occurs in disjunct populations, commonly in hanging gardens in San Juan and Grand counties. It is less frequent in those mesophytic assemblages than is Pri- mula specuicola or Mimuhis eastwoodae. Delisted Species Plants which should be removed (delisted) from the list of endangered species are as follows: Astragalus lentiginosus var. chartaceus.— The papery speckled millkvetch remained obscure for more than half a century fol- lowing its naming. Recent collections in- dicate the presence of a series of popu- lations with compact inflorescences shorter than the leaves in the near vicinity of the type locality of this variety. They are con- nected to other populations which are un- common but scattered through much of northern Utah. Astragalus loanus.— The Loa milkvetch is another species whose identity long re- mained obscure. Collected first a century ago, the species was not named until the 1940s. The plants occur on volcanic gravels in Sevier, Piute, Wayne, and Garfield coun- ties. Total extent of known range is almost 100 miles in length. 6 Great Basin Naturalist Vol. 38, No. 1 Astragalus pardalinus.— The panther milkvetch is a mirrored image species of both A. pubentissimus and A. sabulonum. Differences were observed by P. A. Ryd- berg, who described A. pardalinus, and were substantiated by R. C. Bameby, who placed it in Astragalus. The habitat occu- pied by this pretty plant is the sandy foot- slope of the San Rafael Swell and Henry Mountains in eastern Emery, Wayne, and Garfield counties. Recently, it has been found along the dip-slopes of the Curtis for- mation near the western margin of the San Rafael Swell. The total length of the area occupied is more than 100 miles long. Astragalus serpens.— The plateau milk- vetch is a handsome plant, long known from the type locality at Loa Pass in west- em Wayne County. It is a plant of the low sagebrush community on volcanic gravels. Recently it has been found in Piute, Gar- field, and Iron counties. The populations, while disjunct, consist of few to very many individuals, which wax and wane with local conditions of weather. The area occupied is very large. Cryptantha breviflora.— The Uinta Basin cryptantha occurs from the west central portion of the basin eastward through cen- tral Uintah County, and into the south- eastern tip of Daggett County. It is one of the common species of the Uinta Basin. Hermidium alipes var. pallidum.— Recent collections from the Uinta Basin indicate a broad range for the pale hermidium. It is at least common, and even locally abundant, in much of its range in lower elevation por- tions of Duchesne and Uintah counties. Parthenium ligulatum.— This caespitose composite is known from the Green River Shale of both Duchesne and Uintah coun- ties, and from the Carmel and Dakota for- mations in the San Rafael Swell in Emery County. It remains obscure, but hardly rare. Physaria grahamii.— This plant has been one of the enigmatic species of the state. Collected by Graham in the 1930s, and named by C. V. Morton during that period, the entity was thought to be extinct when first listed in the Federal Register. It is now known rather widely from the escarpments of the Tavaputs, West Tavaputs, and other western plateaus in Carbon, Emery, Duch- esne, and Uintah counties. Additionally, it occurs in west central Colorado. Upgraded Species There are two plant species which were cited as threatened on the Federal Register list of 1975, but which appear to have the limited range and potential for extinction of endangered plants. Therefore, it is proposed that they be upgraded to that status. In- cluded in this category are the following (see Table 2): Astragalus callithrix.— This obscure spe- cies is known from only a few localities throughout its range in Millard County, Utah, and Nye County, Nevada. The loss of any population might be sufficiently de- structive that the species would be lost. Cryptantha bamebyi.— The total area known to be occupiecj^ by this entity is ap- parently smaller than that in published maps of the distribution. The plant is a Green River Shale endemic and could be eradicated through exploitation of that ma- terial for its oil. Threatened Plant Lists AND Proposed Modifications Threatened plants as currently constituted are listed in Table 2. The present list con- sists of some 85 taxa but includes many taxa now recognized as being common. These were previously poorly known entities, which have been updated by contemporary collections. Because of the accumulated in- formation, it is proposed that some 30 spe- cies be removed from the list of threatened Utah plants. Most of those proposed to be delisted have such broad ranges that they could be threatened only by the most wide- spread destruction of habitat, or by some specific and selective destructive force. Plants which should be delisted include: Astragalus convallarius var. finitimus.— There are a series of populations of this plant extending through northern Washing- ton County, western Iron County, and adja- cent Nevada. This plant is not rare; its in- terpretation has been obscure. Astragalus ensiformis.— When expanded to include the materials known as A. min- March 1978 Welsh: Endangered Utah Plants Table 2. Current list of threatened plants cited in Federal Register publications (1975, 1976), their locality by county in Utah, and recommendations of status based on present data. Threatened Plants Locality Recommendations Arabis demissa Greene var. kinugida Rollins var. riisseola Rollins Asclepias cutleri Woodson Asclepias ruthiae Maguire & Woodson Astragalus ampullaritis S. Wats. Astragalus barnebyi Welsh & Atwood Astragalus callithrix Barneby Astragalus chloodes Barneby Astragalus convallarius greene var. finitirnus Barneby Astragalus cottamii Welsh Astragalus duchesnensis M.E. Jones Astragalus ensifomiis M.E. Jones Astragalus lancearius A. Gray Astragalus oophorus S. Wats. var. lonchocalyx Barneby Astragalus rafaelensis M. E. Jones Camissonia parryi (S. Wats.) Raven Castilleja parvula Rydb. Castilleja scabrida Eastw. Corydalis caseana A. Gray Cryptantha barnebyi Johnst. Cryptantha compacta Higgins Cryptantha elata (Eastw.) Payson Cryptantha semiglabra Barneby Cryptantha striata (Osterh.) Payson Cymopterus basalticus M.E. Jones Cymopterus coulteri (M.E. Jones) Mathias Cymopterus duchesnensis M.E. Jones Cymopterus newberryi (S. Wats.) M.E. Jones Cymopterus rosei M.E. Jones Draba asprella Greene var. zionensis (C.L. Hitchc.) Welsh & Reveal Draba sobolifera Rydb. Draba subalpina Goodman & Hitchc. Epilobium nevadense Munz Erigeron abajoensis Cronq. Erigeron cronquistii Maguire Erigeron garrettii A. Nels. Erigeron mancus Rydb. Eriogonum clavellatum Small Eriogonum eremicum Reveal Daggett Daggett, Uintah Grand, San Juan Emery, Grand, Wayne Kane, Washington Garfield, Wayne Millard Uintah Iron, Washington San Juan Duchesne, Uintah Washington Kane, Washington Beaver, Iron Emery Washington Beaver, Piute Carbon, Duchesne, Emery, Garfield, Grand, Kane, San Juan, Sevier, Uintah, Wayne, Washington Salt Lake, Utah, Wasatch, Weber Uintah Millard Grand Washington Daggett, Summit, Uintah Millard Juab, Sevier, Utah Duchesne, Uintah Garfield, Grand, Kane, Millard, San Juan, Uintah, Washington, Wayne Iron, Sanpete, Sevier, Washington Washington Garfield, Piute Garfield, Iron, Kane Washington Garfield, Kane, San Juan Cache Box Elder, Salt Lake, Utah Grand, San Juan San Juan Millard no change no change no change no change no change no change up grade to E no change remove from list no change no change remove from list no change remove from list no change remove from list no change remove from list remove from list upgrade to E no change no change no change no change remove from list no change no change remove from list remove from list no change no change no change no change no change no change remove from list no change no change no change Great Basin Naturalist Vol. 38, No. 1 Table 2.- contd. Threatened Plants Locality Recommendations Eriogonum jatnesii Benth. var. nipicola Eriogonum nanitm Reveal Eriogonum osthinilii M.E. Jones Eriogonum panguicense (M.E. Jones) Reveal var. alpestre (Stokes) Reveal Eriogonum saiirinwn Reveal Eriogonum thompsonae S. Wats. var. alhiflorwn Reveal var. thompsonae Eriogonum viridulum Reveal Euphorbia nephradenia Barneby Geranium marginale Rydb. Cilia mcvickerae M.E. Jones Haplopappiis scopulorum (M.E. Jones) Blake Hymenopappus filifolius Hook. var. tomentosus (Rydb.) Turner Lesquerella garrettii Payson Lesquerella ruhicundula Rollins Ixrmatium minimum Mathias Lupinus niarianus Rydb. Machaeranthera grindelioides (Nutt.) Shinners var. depressa (Maguire) cronq. & Keck Mertensid viridis A. Nels. var. cana (Rydb.) L.O. Williams var. dilatata (A. Nels.) L.O. Williams Noma retrorsum J.T. Howell Opuntia whipplei Engelm. & Bigel. var. multigeniculata L. Benson Parrija rydbergii Botsch. Penstemon ahietinus Pennell Penstemon acaulis L.O. Williams Penstemon caespitosus Niitt. var. suffruticosus A. Gray Penstemon compavtus (Keck) Crosswhite Penstemon leiophyllus Pennell Penstemon parvus Pennell Penstemon uintahensis Pennell PenstBTUon ivardii A. Gray Peteria thompsonae A. Gray Kane, Washington Box Elder, Weber Piute, Sevier Iron Uintah Washington Kane, Washington Duchesne, Uintah Emery, Kane, Wayne Garfield, Iron, Sevier, Wayne Garfield, Kane, Iron, Piute, Sevier, Uintah Iron, Kane, San Juan Kane, Washington Salt Lake, Utah, Wasatch Garfield, Kane, Piute Garfield, Iron, Kane Garfield, Piute, Sevier Beaver, Millard no change no change no change no change no change no change no change remove from list no change remove from list remove from list remove from list no change no change no change no change remove from list remove from list Summit no change Daggett no change Garfield, Grand, remove from list Kane Washington remove from list Daggett, Duchesne, no change Summit, Uintah Iron, Sevier, remove from list Utah Daggett no change Beaver, Garfield, remove from list Piute Cache no change Garfield, Iron, remove from list Kane, Sevier, Washington Garfield, Piute no change Daggett, Duchesne, no change Uintah Sanpete, Sevier no change Emery, Grand, remove from list Juab, Kane, San Juan, Washington March 1978 Welsh: Endangered Utah Plants Table 2.— contd. Threatened Plants Locality Recommendations Phacelia anelsonii Macbr. Phacelia cephahtes A. Gray Phacelia constancei Atwood Phacelia demissa A. Gray var. hetewtricha J.T. Howell Phacelia howelliana Atwood Phacelia rafaelensis Atwood Phacelia iitahensis E.G. Voss Phlox cluteana A. Nels. Phlox gladiformis (M.E. Jones) E. Nels. Phlox grahamii Wherry Phlox pnesii Wherry Primula 7nagiiirei L.O. Williams Primula specuicola Rydb. Puccinellia parishii A.S. Hitchc. Sclerocactus pubispinus (Engelm.) L. Benson Senecio dimorphophyllus Greene var. intermedins Barkley Silene petersonii Maguire var. petersonii Viola charlestonensis Baker & Clausen Washington Kane, Washington Emery, Garfield, Kane, San Juan Piute, Wayne Sevier Grand, San Juan Emery, Wayne, Washington Sanpete, Sevier San Juan Garfield, Iron, Washington Uintah Washington Cache Grand, Kane, San Juan, Wayne not in Utah Box Elder, Beaver, Sevier, Millard San Juan Sanpete Washington no change no change remove from list remove from list no change remove from list no change no change no change remove from list remove from list no change remove from list remove from list no change no change no change no change thorniae var. gracilior, the species becomes rather broadly distributed in a series of in- terconnected populations, which are hardly threatened. Astragalus oorphorus var. lonchocalyx.— Previously known from only a single local- ity in western Iron County, the taxon is now known from several localities in Iron, and in Beaver County also. Additionally, the species is locally common in Lincoln County, Nevada. Castilleja scabrida.— This species was placed on the list by oversight. It is a wide- ly distributed paintbrush, flowering early in the growing season, and rivaling C. chro- mosa in abundance. Camissonia parryi.— This plant has a re- stricted distribution, occurring only in Washington County, Utah. It is hardly rare, however. When the water regimen is con- ducive, this species grows in great abun- dance. Corydalis caseana.— The distribution of this taxon indicates that it is rather wide- spread, although only locally common. It is not known to be threatened at the present time. Cymopterus basalticus.— While the range of this species is known from Millard Coun- ty only, that county is one of the largest in Utah. And, C. basalticus is broadly dis- tributed in the western portion of that county. Indeed, the plants are common to abundant in much of the region. Cymopterus newberryi.— This is another example of a plant which should not have been placed on the list. The broad range of this common plant has been known for many years. Cymopterus rosei.— The plant remains rather obscure, but even fragmentary rec- ords demonstrate an extensive range for the entity. Erigeron garrettii.— The Garrett fleabane daisy has a wide area of distribution in the northern and central Wasatch Range. Eriogonum viridulum.— The range of E. viriduhim is known to extend for more than 100 miles across the east-west axis of the Uinta Basin. 10 Great Basin Naturalist Vol. 38, No. 1 Geranium marginale,— The plateau gera- nium is a common plant over a broad sec- tion of south central Utah. Gilia mcvickerae.— While occurring in disjunct populations, often on peculiar soils (either modified volcanics or calcareous gravels), the current knowledge indicates that the plants are widespread. Haplopappus scopulorum.— A plant of protected coves in sandstone, this species has a broad range which defies classification as threatened. Lupinus marianus.— This represents a portion of the variation within L. sericeus, with which it grades. The distribution of L. marianus in its narrowest sense is known to be more than 60 miles in length. Machaeranthera grindelioides var. dep- ressa.— This plant is widespread on knolls and ridges in western Millard, Tooele, and Beaver counties, where it is locally common to abundant. Nama retrorsum — The distribution of N. retrorsum precludes classification as threat- ened. Opuntia whipplei var. multigeniculata.— This is a taxonomically questionable and ob- scure unit. The plants should be removed from the threatened category pending addi- tional research. Penstemon abietinus.— Large poulations of this taxon are present within its rather broad range. Penstemon caespitosus var. suffruti- cosus.— The populations of this entity, which has a rather broad distribution on volcanic gravels, seem to be enhanced by disturbance, either natural or artificial. Hence, roadcuts and slide areas support vi- able stands of this pioneer plant. Penstemon leiophyllus.— Collections of P. leiophylliis are so numerous and from such a broad area, that one has difficulty in as- certaining how it could be threatened. Peteria thompsonae.— An unusual and in- teresting plant of broad distribution, this plant is locally common or even abundant. There is no reason to list it as threatened. Phacelia constancei.— Known from divers populations over much of southern Utah, there seems to be little justification for keeping this handsome plant on the list. Phacelia demissa var. heterotricha.— Though disjunct, this variety does not seem to be threatened in any portion of its range. Phacelia rafaelensis.— The problem of range is the same as with P. constancei. It is too broadly distributed to be treated as threatened. Phlox grahamii.— The type, and only known collection of P. grahamii, is a tera- tological specimen of what is presumed to be P. longi folia. Aecial sori of a rust are evident on the leaves and stem of the speci- men, and distortion of vegetative and floral parts are in keeping with that observed in other diseased plants. Phlox jonesii.— The type of P. jonesii ap- pears to represent little more than a shade form of P. austromontana, and certainly be- longs within the limits of that species. Primula specuicola.— While of great in- terest from a phytogeographical sense, P. specuicola is found in too many sites to be classed as threatened. This is true even though one stronghold of the species has been eradicated by the rising waters of Lake Powell. The plant has almost an iden- tical distribution as that of Mimulus east- woodiae. Puccinellia parishii.— There is no evi- dence to support the presence of this entity in Utah. Scelerocactus spinosior.— Indications of the relationship of this plant with S. ivhipplei, in a broad sense, require a reeval- uation of this entity which, though some- what obscure, seems to be rather broadly distributed. Candidate Endangered Species There are four taxa which require consid- eration as endangered species. Of these, only one was known at the time when in- itial lists were prepared (Table 3). That one, Lesquerella tumulosa, had been placed into synonymy with a related species, L. rubi- cundula. In the judgment of this writer, L. tumulosa is distinct morphologically, spa- tially, and substrate-wise from L. rubicun- dula. The two plants are separated by about 900 meters in elevation. They flower at different times, with the low elevation L. tumulosa completing flowering at about the time L. rubicundula begins. Lesquerella March 1978 Welsh: Endangered Utah Plants 11 Table 3. Candidate endangered species not currently on Federal Register lists (1975, 1976), their locality by county in Utah, and justification based on present data. Endangered Plants (Proposed) Locality Justification Astragalus montii Welsh Sanpete Local; restricted to limestone; high elevation Eriogonum corymbosum Benth. Washington Local; restricted var. Matthewsae Reveal to inud-siltstone; low elevation Lesquerella tumulosa (Bameby) Kane Local; restricted Reveal to limy mudstone; low elevation Thelypodiopsis argillacea Welsh & Uintah Local; restricted Atwood To oil shale; moderate elevation tumulosa is known from white calcareous shales of the Winsor member of the Carmel Formation, and L. rubicundula occurs on the Wasatch limestone sequence. Further, the pluricipital, pulvinate growth with a great many inflorescences per plant is not known for L. rubicuncula, but is character- istic of the population of L. tumulosa. The remaining candidate endangered spe- cies are all newly described or undescribed. They are as follows: Astragalus montii Welsh sp. nov.— Astra- galo limnochari Bameby aemulans differt in floribus magnioribus et purpureis et foliolis non ciliatis. Plantae perennes, acaulescentes, 1-5 cm altae, caudicibus ramificantibus ena- nescentibus; pubescentiae basifixae; stipulae 2-4 mm longae totus connatae-vaginantes; foliola 1.3-4.8 cm longa: foliola 5-13, 2-8 mm longa 1-2 mm lata lanceolata oblonga vel elliptica strigosa infra non ciliata ad marginem glabra supra; pedunculi 0.8-4.5 cm longi in fructum reclinatimi; racemi 2- to 8-floribus, floribus ascendentibus vel pat- entibus ad ahthesin, axibus 0.2-0.5 cm long- is in fructum; bracteae 1-3 mm longae; pedicelli 0.8-1.4 mm longi; bracteolae nul- lae; calyx 3.3-4 mm longus, tubo 2.2-2.5 mm longo campanulato strigoso, dentibus 0.6-1.5 mm longis triangularibus vel sub- ulatis; flores 7.2-8 mm longi purpurei, ala apicis albis; legumina patentia sessilia ovoidea vesicaria 11-18 mm longa 8-12 mm lata maculosa uniloculari; ovula 10. Holotype: Utah, Sanpete Co., ca 17 miles due west of Ferron, Flagstaff Limestone, marly barrens, at 3,350 m, S. L. & Jean Welsh 15404, 13 July 1977 (BRY, isotopes to be distributed). Paratypes: Utah, Sanpete Co., M. E. Lewis 4312, 23 July 1976; do, M. E. Lewis 4775, June 1977 (both at BRY). Astragalus montii is a near congener of A. lim,nocharis from which it is isolated geo- graphically by some 225 km. Both are low- growing calciphiles occurring on limestone gravels at high elevations. They agree fur- ther in general aspect and in having small flowers giving rise to bladdery-inflated uni- locular pods. The flowers differ in color, and those of A. montii average larger in size. The striking long hair which margin the leaves of A. limnocharis are lacking in A. montii. The species is named to honor Mont E. Lewis, extraordinary plant collec- tor, and codiscoverer of the plant along with Robert Thompson. Eriogonum corymbosum var. matthew- sae.— The only known population of this buckwheat variety occurs at the mouth of Zion Canyon on the Chinle Formation. Roadways, power lines, and other devel- opments on the slope where the plants oc- cur threaten its existence. Certainly, this plant has the characteristics of an endan- gered species. Thelypodiopsis argillacea.— The Green River Shale formation serves as substrate for many unusual species of plants. Thus, it is not unexpected that still another narrow en- demic be located on that formation. The species was located while searching for Glaucocarpum suffrutescens, another rare plant of Green River Shale. Subsequent in- vestigation of the type area indicated that 12 Great Basin Naturalist Vol. 38, No. 1 T. argillacea is narrowly restricted. The small population is known at present from a single north-facing slope. Based on that in- formation, and given the potential for de- velopment of oil shale, T. argillacaea has the features of an endangered plant. Candidate Threatened Species There are several species and varieties of very limited distribution which were over- looked or unnamed at the time when the original lists of species were proposed as threatened. The candidates for inclusion on the threatened list for Utah is presented in Table 4. Justification for inclusion is pres- ented below. Astragalus castaneiformis var. con- sobrinus.— This is an obscure tiny plant of volcanic gravels with its main locus in Rab- bit Valley, Wayne Co., where it is only lo- cally common. Further materials might dic- tate its removal from the list. Astragalus henrimontanensis (Welsh, nom. nov. based on Astragalus stocksii Welsh Great Basin Nat. 34:307. 1974.- The Dana milkvetch is known from the Henry Mountains only. It is restricted to an eleva- tional range corresponding to the ponderosa pine belt in the Penellen Pass vicinity. To- tal extent of the population based on cur- rent information is through approximately six miles of the eastern flank of Mount Pen- nell. Development of any kind, including T.\BLE 4. Candidate threatened species county in Utah, and justification based on not currently on Federal Register lists (1975, 1976), their locality by present data. Threatened Plants (Proposed) Locality Justification Astragalus castaneiformis S. Wats. var. consobrinus Bameby Astragalus limnocharis Bameby Astragalus henrimontanensis Welsh Astragalus monumentalis Barneby Astragalus sabulosus M.E. Jones Cymopterus hi^insii Welsh Dalea epica Welsh Draba maguirei C.L. Hitchc. Eriogonum natum Reveal Gaillardia flava Rydb. Hedysarum boreale Nutt. var. gremiale (Rollins) Northstrom & Welsh Helianthus deserticolus Heiser Hymenoxys depressa (Torr. & Gray) Welsh & Reveal Lepidium montanum Nutt. var. neeseae Welsh & Reveal var. stellae Welsh & Reveal Lomatium latilobum (Rydb.) Mathias Lupinus jonesii Rydb. Machaeranthera glabriuscula (Nutt.) Cronq. & Keck var. confertifolia Cronq. Machaeranthera kingii (D.C. Eaton) Cronq. & Keck Garfield, Piute, Local; volcanic gravel Sevier, Wayne Iron, Kane Local; calciphile; high elevation Garfield local; montane endemic Garfield, San Juan Local; Cedar Mesa Sandstone endemic Grand Local; Mancos Shale endemic Kane Local; Tropic Shale endemic San Juan Local; Navajo Sandstone endemic Cache, Weber Local; montane endemic Millard Local; lacustrine terrace endemic Grand, Emery Local; Mancos Shale endemic Uintah Local; Bishop Conglomerate endemic Washington Local; sand dune endemic Emery, Garfield, Local; Silicious Uintah (?) conglomerate, very restricted Garfield Local; Navajo Sandstone endemic Kane Local; Windsor endemic Grand, San Juan Local; Entrada Sandstone endemic Washington Local; sandstone and alluvial endemic Garfield, Kane Local; Kaiparowits Endemic Cache, Salt Lake, Disjunct, local; Utah calciphile March 1978 Welsh: Endangered Utah Plants 13 Table 4.— contd. Threatened Plants (Proposed) Locality Justification Mentzelia argillacea Darlington Sevier Local; Arapien Shale endemic Musineon lineare (Rydb.) Mathias Cache Local; lacustrine terrace endemic Najas caespitosa (Maguire) Reveal Sevier Local; endemic to Fish Lake Penstemon atwoodii Welsh Garfield, Kane Local; Kaiparowits vicinity endemic Penstemon bracteatus Keck Garfield Local; Wasatch endemic Penstemon hwnilis Nutt. Washington Local; Navajo Sandstone var. Ohtusifolius (Pennell) endemic Reveal Penstemon tidestromii Pennell Juab, San Pete Local; Sanpitch Mountains endemic Psoralea pariensis Welsh 6f Atwood Garfield, Kane Local; Bryce Canyon— Paria drainage endemic Silene petersonii Maguire Garfield, Iron Local; Wasatch endemic var. minor Hitchc. & Maguire Sphaeralcea caespitosa M.E. Jones Beaver, Millard Local; Wah Wah— Pine Valley endemic Thelypoditim sagittatum (Nutt.) Endl. Garfield, Iron Local; Markagunt var. ovalifolium (Rydb.) Welsh & Plateau endemic Reveal Townsendia mensana M.E. Jones Duchesne, Uintah Local; Uinta Basin endemic Townsendia minima Eastw. Garfield, Kane Local; Wasatch endemic Xanthocephalum sarothrae (Pursh) Uintah Local; Ashley Valley- Shinners var. pomariense (Welsh) Green River endemic Welsh reclamation of lands in range improvement attempts, could result in loss of substantial portions of the population. Astragalus limnocharis.— The Navajo Lake milkvetch is known mainly from the gravelly shore of the lake. Dispersal of the bladdery pods is apparently by wind and by water. The plants tend to occur in rows along minor wave-cut terraces in the beach. A dam extends across the lake basin, limit- ing the water to the west end of the lake, except in peak years. The milkvetch species does not now occur in the portion of the basin from which the lake has been ex- cluded. A secondary location has been re- ported in the Cedar Breaks vicinity, a short distance to the northwest of the Navajo Lake locus. Astragalus monumentalis.— The Mon- ument milkvetch is known from Natural Bridges National Monument westward along the Cedar Mesa Sandstone to where that formation dips beneath other strata west of the Colorado River Canyon in easternmost Garfield County, and north to the Chesler Park region of Canyonlands National Park. Any development on that formation could result in loss of portions of the total popu- lation, and require change of status to en- dangered. Astragalus sabulosus.— This plant is known only from the vicinity of Thompson east to Cisco, and possibly south as far as the banks of the Colorado River in that general vicinity. All modern collections are from Mancos Shale, but early collections by Jones presumably came from Morrison or Cedar Mountain formations near the river. Even in optimum seasons of growth for this handsome large-flowered selenophyte the number of plants in the known populations is small. Only a few dozen plants have ever been observed by the writer in several years of observation. The plant has very large ochroleucous flowers which distinguish this plant easily from the similar A. praelongus, with which it grows. The entire known 14 Great Basin Naturalist Vol. 38, No. 1 range of the species is in northcentral Grand County. Atriplex welshii.— The Welsh saltbush is a local species of the Mancos Shale a few miles southwest of Cisco in Grand County. It is a restricted plant of the Mancos Shale, in a region where few plants occur on the formation. Other local species are known from the vicinity, and perhaps the entire range of the species should be considered for preservation. Cymopterus higginsii.— A plant with lav- ender flowers, C. higginsii, is known from the Tropic Shale (Tununk equivalent) from the vicinity of Coyote Creek east to Smoky Mountain in eastern Kane County. The area is being subjected to heavy use by off- highway vehicles, and other anticipated ac- tivities along with that use might jeopardize portions of the population, which is only lo- cally common in the region. Dalea epica.— The Hole-in-the-Rock prairie clover is known from a small popu- lation on Navajo Sandstone some distance east of Halls Crossing in San Juan County. The population has been interpreted by R. C. Barneby (personal communication) as a portion of D. flavescens, to which it is un- doubtedly closely allied. The population de- serves preservation, even if it is only an ex- clusive phase within the total variation of that species. The taxon is recognized at spe- cies level by Welsh (1977) in a treatment of the legumes of Utah, which is in prepara- tion. Draba maguirei.— This entity, in a broad sense, is very restricted to mountain sum- mits in a small portion of the northern Wasatch Range. Eriogonum natum.— This wild buckwheat is a narrow endemic of Lake Bonneville la- custrine deposits in Millard County. The to- tal known area of the population is very small. It is worthy of being regarded as threatened. Gaillardia flava.— A smelly, resinous plant of canyon bottoms in portions of the Man- cos Shale, this handsome yellow blanket- flower has long remained obscure. Total range is over a region only about 20 miles in length. Hedysarum boreale var. gremiale.— An unusual phase of H. boreale with spinulose projections along the reticulations of the fruit, this plant occurs in west central Uin- tah County. The plants apparently grade with the more typical H. boreale materials in the region. However, this is the only known taxon of American Hedysarum spe- cies with spinulose projections. The popu- lation is poorly known, and further work is needed to elucidate the total area involved. Helianthus deserticolus.— This sand- inhabiting sunflower is known to occur in a limited low-elevation portion of Washington County. More information is necessary to understand the total limits of the taxon and to clarify its relationship to H. anomalus Blake. Hymenoxys depressa.— Although first named by Gray (1849), this plant has re- mained obscure. The type was taken by Fremont on his second expedition, possibly in the Uinta Basin of Utah. Lepidum montanum var. neeseae.— This tiny phase of the polymorphic L. mon- tanum is known from high elevations on Navajo Sandstone only. Lepidium montanum var. stellae.— The Stella Lepidium grows with Lesquerella tumulosa on the Winsor formation in west- em Kane County. Lomatium latilobum.— The broad-lobed biscuitroot is a narrow endemic of sandy soils, usually at the base of monoliths in En- trada Sandstone in south central Grand and north central San Juan counties. The species is only locally common. Development of those regions might cause the species to be- come endangered. Lupinus jonesii.— The Jones lupine in- habits sandy and calcareous soils of central and western Washington County. The spe- cies has long remained obscure, and much more information should be gathered prior to final disposition of this taxon. Machaeranthera glabriuscula var. confer- tifolia.— The total area occupied by this handsome white-flowered daisy is confined to the gray colluvial and in situ soils of the Kaiparowits formation. The plants grow on crests and eroded flats of that formation, generally within the juniper-pinyon wood- land. The formation is local in the northern portion of the Kaiparowits Plateau vicinity. March 1978 Welsh: Endangered Utah Plants 15 and it might be endangered by proposed and future developments. Machaeranthera kingii.— Much more in- formation is required to assess the total range of M. kingii, but indications are that the plant is restricted to limestone outcrops in the mountains of Cache, Salt Lake, and Utah counties. Collections and known local- ities of the plant are few. Mentzelia argillacea.— The clay blazing star is restricted to Arapien Shale and allu- vium derived from that formation and oth- ers adjacent to it. Total area is apparently from near Monroe on the south to Salina on the north. Mining for gypsum and other minerals shrinks the area occupied by this distinctive plant each year. Musineon lineare.— This species is known from a localized portion of Cache County, which is being impressed by construction and other activities. Najas caespitosa.— Fish Lake is the only known habitat of this aquatic plant. The relationship apparently lies with more wide- spread species in the genus. Until the rela- tionship is clarified, it is best to consider these unusual plants as threatened. Penstemon atwoodii.— The Atwood beardtongue has a similar but smaller range than Astragalus malacoides. Both are con- fined to the Straight Cliffs and associated formations on the Kaiparowits Plateau and vicinity. Prospective development of re- sources in the Kaiparowits could result in eradication of portions of the population and might lead to endangered classification. Penstemon humilis var. obtusifolius.— Confined to Navajo sandstone and alluvium derived from that formation, this remark- ably low beardtongue is known only from eastern Washington County. Penstemon tidestromii.— Long obscure, the Tidestrom beardtongue is known only from the Sanpitch Mountains in eastern Juab and Sanpete counties. Psoralea pariensis.— The few known pop- ulations of the Paria scurfpea occur on Wasatch Limestone and alluvium derived from the formation, or less commonly on sandstone and sandy alluvium. Many more collections might indicate a rather broader range and require a reevaluation of the threatened status. Silene petersonii. var. minor.— Endemic to limestone members of the Wasatch For- mation, this variety of the striking S. peter- sonii is known from small populations lo- cated in two main areas. The type locality is at Red Canyon in Garfield County, and the other main locus is at Cedar Breaks. Sphaeralcea caespitosa.— The Jones globe mallow has a similar area of occupation as does Penstemon concinnus and P. nanus, but it does not always occur with those spe- cies. Portions of western Millard and Beaver counties not inundated by Lake Bonneville seem to be the primary areas occupied by this beautiful plant. Thelypodium sagittatum var. ovalifo- lium.— This peculiar plant is known from the vicinity of Panguitch Lake in western Garfield and adjacent Iron counties. The populations are poorly known, indicating a need for much additional information. Townsendia mensana.— The range of T. mensana is only generally known. The stat- us of the species should be indicated as threatened imtil such a time that this entity is demonstrated to be more common than currently indicated. Townsendia minima.— The least town- sendia is known from the white and pink limestone members of the Wasatch Forma- tion. Usually, the plants occur in narrow bandlike strips along the margin of breaks and ridge tops on the formation. Distribution by County Federal agencies and others interested in development of lands in Utah require infor- mation on proposed endangered and threat- ened plant distribution so that decisions on land use can be made. The summary lists presented below will allow interested par- ties to determine whether more specific in- vestigations will be necessary. The lists re- flect the recommended changes indicated in the present paper. Beaver County Castilleja parvula (T) Penstemon concinnus (T) Penstemon nanus (T) Sclerocactus pubispinus (T) Sphaeralcea caespitosa (T) Box Elder County Allium passeyi (E) 16 Great Basin Naturalist Vol. 38, No. 1 Eriogoniim nanum (T) Sclerocactus pubispinus (T) Cache County Draba maguirei (T) Erigeron cronquistii (T) Eriogonum loganum (E) Machaeranthera kingii (T) Miisineon lineare (T) Penstemon compactiis (T) Primula maguirei (T) Carbon County Eriogonum corymbosum var. davidesei (E) Eriogonum lancifolium (E) Lygodesmia grandiflora var. stricta (E) Daggett County Arabis demissa var. lanugida (T) var. russeola (T) Cryptantha stricta (T) Mertensia viridis var. dilatata (T) Parrya rydbergii (T) Penstemon acaulis (T) Penstemon uintahensis (T) Duchesne County Astragalus detritalis (T) Astragalus duchesnensis (T) Cymopterus duchesnensis (T) Eriogonum hylophilum (E) Lepidium bamebyanum (E) Parrya rydbergii (T) Penstemon garrettii ? (E) Sclerocactus glaucus (E) Townsendia mensana (T) Emery County Asclepias ruthiae (T) Astragalus rafaelensis (T) Cryptantha jonesiana (T) Cryptantha johnstonii (E) Erigeron maguirei (E) Eriogonum intermontanum (E) Eriogonum smithii (E) Euphorbia nephradenia (T) Festuca dasyclada ? (PoEx) Gaillardia flava (T) Hymenoxys depressa (T) Sclerocactus wrightiae (E) Garfield County Astragalus bamebyi (T) Astragalus castaneiformis var. consobrinus (T) Astragalus henrimontanensis (T) Astragalus malacoides (T) Astragalus monumentalis (T) Astragalus perianus (T) Castilleja aquariensis (E) Castilleja revealii (E) Cryptantha ochroleuca (E) Eriogeron abajoensis (T) Eriogonum aretioides (E) Eriogonum corymbosum var. revealianum (T) Eriogonum cronquistii (E) Draba sobolifera (T) Draba subalpina (T) Heterotheca jonesii (E) Hymenoxys depressa (T) Lepidium montanum var. neeseae (T) Lesquerella rubicundula (T) Lomatium minimum (T) Machaeranthera glabriuscula var. confertiflora (T) Penstemon atwoodii (T) Penstemon bracteatus (T) Penstemon parvus (T) Phlox gladiformis (T) Psoralea pariensis (T) Silene petersonii var. minor (T) Ranunculus acriformis var. aestivalis (PoEx) Thelypodium sagittatum var. ovalifolium (T) Townsendia minima (T) Grand County Asclepias cutleri (T) Asclepias ruthiae (T) Astragalus iselyi (E) Astragalus sabulosus (T) Atriplex welshii (T) Cryptantha elata (T) Cycladenia humilis var. jonesii (E) Erigeron mancus (T) Gaillardia flava (T) Lomatium latilobum (T) Phacelia howelliana (T) Iron County Astragalus lentigenosus var. ursinus ? (PoEx) Astragalus limnocharis (T) Cymopterus minimus (E) Draba subalpina (T) Erigeron flagellaris var. trilobatus (E) Eriogonum panguicense var. alpestre (T) Lomatium minimum (T) Phlox gladiformis (T) Silene petersonii var. minor (T) Thelypodium sagittatum var. ovalifolium (T) Juab County Cymoptenis coulteri (T) Penstemon tidestromii (T) Kane County Astragalus ampullarius (T) Astragalus lancearius (T) Astragalus limnocharis (T) Cymopterus higginsii (T) Draba subalpina (T) March 1978 Welsh: Endangered Utah Plants 17 Erigeron abajoensis (T) Erigeron religiosus (E) Eriogonum jamesii var. rupicola (T) Eriogonum thompsonae var. thompsonae (T) Euphorbia nephradenia (T) Hymenopappus fiUfohus var. tomentosus (T) Lepidium montanum var. stellae (T) Lesquerella rubicundula (T) Lesquerella tumulosa (E) Lomatium minimum (T) Machaeranthera glabriuscula var. confertiflora (T) Penstemon atwoodii (T) Phaceha cephalote.s (T) Phacelia mammillariensis (E) Psoralea epipsila (E) Psoralea pariensis (T) Townsendia minima (T) Viguiera sohceps (E) Zigadenus vaginatus (T) Millard County Astragalus caUithrix (E) Cryptantha compacta (T) Cuscuta wameri (PoEx) Eriogonum ammophilum (E) Eriogonum eremicum (T) Eriogonum natum (T) Penstemon concinnus (T) Penstemon nanus (T) Penstemon parvus (T) Sclerocactus pubispinus (T) Piute County Astragalus castaneiformis Astragalus perianus (T) Eriogonum corymbosum var. revealianum (T) Castilleja parvula (T) Draba sobolifera (T) Eriogonum ostlundii (T) Lesquerella nibicundula (T) Salt Lake County Lesquerella garrettii (T) Machaeranthera kingii (T) San Juan Count\' Asclepias cutleri (T) Astragalus cottamii (T) Astragalus cronquistii (E) Astragalus iselyi (E) Astragalus monumentalis (T) Dalea epica (T) Erigeron abajoensis (T) Erigeron kachinensis (E) Erigeron mancus (T) Eriogonum clavellatum (T) Eriogonum humivagans (E) Lomatium latilobum (T) Phacelia howelliana (T) Phacelia indecora (E) Senecio dimorphophyllus var. intermedius (T) Sanpete County Astragalus desereticus (PoEx) Astragalus montii (E) Festuca da.syclada (PoEx) Penstemon tidestromii (T) Penstemon wardii (T) Phacelia utahensis (T) Silene petersonii var. petersonii (T) Sevier County Astragalus castaneiformis var. consobrinus (T) Cymopterus coulteri (T) Eriogonum ostlundii (T) Mentzelia argillacea (T) Najas caespitosa (T) Penstemon wardii (T) Phacelia utahensis (T) Sclerocactus pubispinus (T) Townsendia aprica (E) Summit County Cryptantha stricta (T) Mertensia viridis var. cana (T) Parrya rydbergii (T) Uintah County Arabis demissa var. russeola (T) Astragalus detritalis (T) Astragalus duchesnensis (T) Astragalus hamiltonii (T) Astragalus lutosus (E) Astragalus saurinus (T) Cryptantha barnebyi (E) Cryptantha grahamii (E) Cryptantha stricta (T) Eriogonum ephedroides (T) Eriogonum intermontanum (E) Eriogonum saurinum (T) Glaucocarpum suffrutescens (E) Hedysanim boreale var. greiniale (T) Hymenoxys depressa ? (T) Lepidium bamebyanum ? (E) Parrya rydbergii (T) Penstemon grahamii (E) Penstemon uintahensis (T) Sclerocactus glaucus (E) Thelypodiopsis argillacea (E) Townsendia mensana (T) Utah County Cymopterus coulteri (T) Lesquerella garrettii (T) Machaeranthera kingii (T) Phacelia argillacea (E) Wasatch County Lesquerella garrettii (T) Penstemon garrettii (PoEx) 18 Great Basin Naturalist Vol. 38, No. 1 Washington County Arctomecon humilis (T) Astragalus ampullarius (T) Astragalus ensiformis (T) Cryptantha semiglabra (T) Draba zionensis (T) Echinocereus engelmannii var. purpureus (E) Epilobium nevadense (T) Erigeron religiosus (E) Erigeron sionis (E) Eriogonum corymbosum var. matthewsae (E) Eriogonum jamesii var. rupicola (T) Eriogonum thompsonae var. albiflonim (T) var. thompsonae (T) Eriogonum zionis (T) Helianthus deserticolus (T) Heterotheca jonesii (E) Hymenopappus filifolius var. tomentosus (T) Lupinus jonesii (T) Pediocactus sileri (E) Penstemon humilis var. obtusifolius (T) Phacelia anelsonii (T) Phacelia cephalotes (T) Phlox gladiformis (T) Viola charlestonensis (T) Wayne County Asclepias ruthiae (T) Astragalus bamebyi (T) Astragalus castaneiformis var. consobrinus (T) Astragalus harrisonii (E) Euphorbia nephradenis (T) Cilia caespitosa (E) Phacelia indecora (E) Sclerocactus wrightiae (E) Weber County Draba maguirei (T) Eriogonum nanum (T) Literature Cited Gray, A. 1849. Plantae Fendlerianae Nove-Mexicanae. Mem. Am. Acad. 4:1-116. Welsh, S. L. 1976. Proposed threatened, endangered, presumed extinct, or extinct and disjunct relict plants in the Cedar City and Richfield districts, Utah. Final Report, Bureau of Land Manage- ment, unpublihed ms. 205 pp. 1977a. Endangered and threatened plant spe- cies of the Central Coal Land, Utah. Final Re- port, Interagency Task Force on Coal, U.S. Geological Survey. 422 pp. 1977b. Endangered and threatened plant spe- cies of the Southern Coal Lands, Utah. Final Report, Interagency Task Force on Coal, U.S. Geological Survey. 415 pp. Welsh, S. L. and N. D. Atwood. 1977. An unde- scribed species of TJielypodiopsis (Brassicaceae) from the Uinta Basin, Utah. Great Basin Nat. 37:95-96. Welsh, S. L., N. D. Atwood, and J. L. Reveal. 1975. Endangered, threatened, extinct, endemic and rare or restricted Utah vascular plants. Great Basin Nat. 35:327-376. PLANTS OF NAVAJO NATIONAL MONUMENT J. D. Brotherson,' G. Nebeker,^ M. Skougard,' and J. Fairchild' Abstract.— The floristic composition of Navajo National Monument is presented. The flora of the monument consists of 293 species of vascular plants, representing 177 genera and 66 families. The species are scattered throughout 12 plant communities found within the monument. The flora is dominated by forbs (60 percent), fol- lowed by shrubs (16 percent), grasses (12 percent) and trees (5 percent). Betatakin canyon exhibits the greatest floristic richness with 223 species being represented and 123 species found only in that area. This is due primari- ly to two factors: (1) the greater variety of habitats available in the area, and (2) the much longer time the area has been protected from grazing and other manmade disturbance. Navajo National Monument is in north- eastern Arizona about 10 miles north and west of Black Mesa and Arizona Highway 160. The park headquarters are located on the Shonto Plateau near the head of Betata- kin Canyon, a small side canyon of Tsegi Canyon. The principle attractions of the monument are three large Indian "cliff dwellings" of the Anasazi culture. These cliff dwellings are located in three separate canyons. Betatakin and Keet Seel canyons are located in the Tsegi Canyon complex, while Inscription House is situated about 20 miles west of Betatakin in Nitsin Canyon, a branch of Navajo Canyon. All three units lie in pinyon-juniper slick- rock country which tends to influence heav- ily the environment of the monument. In the region are many deep-cut canyons with high-walled sandstone cliffs often reaching heights of 1000 feet above the streambeds. Within these deep canyons and along their walls, water (in the form of springs and seeps) is often present, creating unique habi- tats in which develop plant and animal communities often foreign to the overall pinyon-juniper type. Because of the historical background of the park, almost all previous studies con- ducted at Navajo National Monument have been archaeological in nature. Dean (1969) points out that up to that time no botanical studies of Tsegi Canyon and its environs had been completed. Apparently, he was re- ferring to the vascular flora, since Flowers (1963) published a study on the lichens and mosses of Betatakin Canyon and vicinity. In his introduction. Flowers (1963) briefly lists some of the dominant plant species in and around Betatakin Canyon. Woodbury (1963) at the request of the National Park Service studied the features of Betatakin Canyon and vicinity for their interpretative value and use by the park personnel. His treat- ment is cursory and brief. Climate Climatically, Navajo National Monument lies within the northeast sector of Arizona (Sellers 1964). There is a weather station at the park headquarters. Temperature and precipitation data for this station have ap- peared in Climatalogical Data, Arizona (U.S. Dept. of Commerce), since 1939. The average annual temperature at Nav- ajo National Monument headquarters is 50 F. The yearly maximum temperatures range from 94 to 101 F with an average of 96.1 F. The yearly low temperatures range from -10 to +8 F with an average of 1 F. The frost-free season ranges from 107 to 213 days with an average of 155 days. The total annual precipitation at Betata- kin has historically ranged from a low of 6.84 inches to a high of 18.79 inches with an average of 11.39 inches. The period of greatest precipitation is late summer and early fall. Rainfall within the canyons is variable and spotty, with localized 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 'Department of Botany and Microbiology, Arizona State University, Tempe, Arizona 85281. 'United States Department of Agriculture, Houston, Texas. 19 20 Great Basin Naturalist Vol. 38, No. 1 cloudbursts occurring more often than gen- eral rains. Geology The geology of Tsegi Canyon has been treated by a number of authors (Gregory 1916, 1917; Hack 1942, 1945; Harshbarger, Repenning, and Irwin 1957; Cooley 1958, 1962). Navajo Sandstone is the major geo- logical formation in each of the three local- ities. At Keet Seel and Inscription House the Navajo Sandstone is the only exposed rock formation. At Betatakin the Kayenta formation outcrops beneath the Navajo for- mation and can be observed in some loca- tions in the lower reaches of the canyon. In all three canyons the Navajo Sandstone forms tall, rather sheer cliffs, up to 700 feet in height. At the base of these cliffs in each canyon, talus accumulations can be found, while the canyon bottoms are filled with deep deposits of sandy alluvium (quaternery fill). In Keet Seel and Nitsin Canyons these alluvial deposits are deeply eroded. In Be- tatakin Canyon the alluvial deposits are less extensive and not nearly so thick. They are underlain by the Kayenta formation in the lower half of the canyon and are deeply eroded, as is more characteristic of the can- yon floor throughout the Tsegi Canyon sys- tem. Plant Communities In line with the unique geological and habitat features of Navajo National Mon- ument, the communities within its bound- aries are varied and in some cases unique. Twelve community (association) types (Table 1) are encountered within the mon- ument. Of these only one (Pinyon-Juniper mixed shrub) is found in all segments of the monument. Five of the remaining 11 com- mimity types are found in 2 of the mon- ument areas while 6 of the types are re- stricted to a single segment. Because of the uniqueness of the Betata- kin Canyon area, the vegetation there is highly different and distinct when compared with adjacent Tsegi Canyon and surround- ing areas. Of special note is the presence in the canyon of a complex of species (i.e., Populus tremiiloides, Pseudotstiga menziesii, Mahonio repens, Symphoricarpos vacci- noides, etc.) more often associated with high mountain rather than desert habitat. These species are found in the canyon because of favorable moisture regimes created by the canyon geology and Navajo Sandstone hy- drology. Flora of the Monument The known flora of Navajo National Monument consists of 293 species of vascu- Table 1. Vegetative communities (associations) encountered within the boundaries of Navajo National Monument and the percent of area they occupy. Vegetation type Monument segment Inscription House Keet Seel Batatakin Percent of area Percent of area Percent of area covered covered covered Streamside-tree 1.8 SHckrock-scattered shrub 35.0 12.6 Atriplex-grass 25.9 0.2 Talus slope 11.3 9.3 Mixed weed 13.5 Pinyon-juniper-mixed shrub 26.0 59.6 43.5 Oak 4.8 2.4 Hanging garden OOJ T Pinyon-juniper-sage 49.9 Pinyon-juniper-grass 3.1 Douglas Fir 0.4 Aspen .7 'Total acreage equals: Inscription House = 40 Keet Seel = 40 Betatakin = 452 "T = trace March 1978 Brotherson et al.: Navajo Monument Plants 21 lar plants, representing 177 genera and 66 families. Of these species, 73 percent belong to the plant families listed in Table 2. These figures indicate the relative plant family composition of the monument. When comparisons are made with other studies (Brotherson 1967, Campbell 1977, Allan 1977) done in somewhat comparable areas of the Great Basin (Table 2), 7 of the 17 plant families listed are found to be repre- sented in the floras of all 4 locations. As can be seen from the table. Arches National Park (Allan 1977) is the most similar in its dominant family composition to that of Navajo National Monument, differing only in 5 of the 17 families listed. The studies done in the Uintah Basin and Kaiparowits areas show high degrees of similarity to each other, differing only in one major plant family as well as being dominated by 9 of the 17 plant families listed in the table. In light of these facts it is of interest to note that the Navajo National Monument and Arches National Park areas have as their major geological formations sandstone while the Kaiparowits and Uintah Basin study areas are underlain mainly by geologi- cal formations high in clay. The ecological and/or phytogeographical significance of the dominance of these 17 plant families and/ or their relationship to the general ge- ology of the area is not known; however, further investigation along such lines tends to hold interest. The species richness of the different seg- ments of the Navajo National Monument is shown in Table 3. As can be seen, the Be- tatakin area exhibits more than twice as many species in its vegetation as can be found in the other areas. It also contains the greatest number of species found in only one of the monument areas. This would pri- marily be due to two factors: (1) the mvich longer time the area has been protected from grazing and other man-made disturb- ance, and (2) the greater variety of habitats available in the area. The life-form and /or growth habitat char- acteristics of the flora are summarized in Tables 4 and 5. Native species number 270, 92.2 percent of the total flora. Twenty- three species are introduced. These include such species as Tamarix ramosissima, Elaeagnus angustifolia, Chenopodium al- bum, Salsola iberica. Sisymbrium altissi- mum, Melilotus alba, and other species of a weedy nature. The herbaceous types of growth habit account for almost 75 percent of the species, with shrubs and trees con- tributing together about 20 percent of the total. In area coverage, however, the tree- Table 2. Dominant plant family composition of Navajo National Monument and other great basin sites. Family Navajo National Monument Kaiparowits Arches National Park Uintah Basin Asteraceae Poaceae Brassicaceae Fabaceae Rosaceae Nyctaginaceae Cactaceae Boraginaceae Polmoniaceae Scrophulariaceae Chenopodiaceae Ranunculaceae Solanaceae Polygonaceae Cyperaceae Liliaceae Salicaceae Total 21.2 12.6 4.4 4.4 3.1 3.1 3.1 2.7 2.7 2.7 2.4 2.4 2.4 2.0 2.0 2.0 73.2 21.4 22.0 22.0 15.5 13.7 9.3 4.8 4.5 8.7 4.8 6.9 1.7 1.7 5.8 4.8 2.8 5.7 8.3 4.0 2.8 5.7 10.7 6.6 5.8 4.8 3.3 1.9 1.9 1.7 5.7 75.1 71.5 72.7 22 Great Basin Naturalist Vol. 38, No. 1 and shnib-dominated vegetation types are the most widespread in the monument. CheckHst The present hst of plant species is the re- sult of specimen collection at Navajo Na- tional Monument that has extended over a period of more than 45 years. Personnel as- sociated with the park service began collec- tions during the late 1930s and early 1940s. Scientists from Brigham Young University began work in the monument in the sum- mer of 1972. Since that time regular visits have been made to the area and collections have been made throughout the growing season. Table 3. Floristic richness of the different segments of Navajo National Monument. No. of species found No. of only at area Area of Monument species in question Betatakin 223 123 Keet Seal 108 16 Inscription House 108 31 Table 4. Plant characteristics and generalized life forms of the flora of Navajo National Monument. Category No. of species in category Percent of total Total no. of species Native species Introduced species Perennial species Biennial species Annual species 293 100.0 270 92.2 23 7.8 224 76.5 21 7.2 48 16.4 Table 5. Growth habit characteristics of the flora of Navajo National Monument. Category No. of species in category Percent of total Trees Shrubs Forbs Grasses Sedges Vines Other 14 4.8 49 16.7 176 60.1 37 12.6 4 1.4 1 .3 12 4.1 Over the years the area has been visited and collections made by: M. Wetherill, 1930-1938; J. W. Brewer, 1934-1947; S. P. Brewer, 1941; E. Lehnert, 1960; E. Jackson, 1961; W. S. Phillips, 1964; J. M. Rominger, 1965; M. Skougard and G. Nebeker, 1972-1973; J. Brotherson, 1972-1977; and J. Fairchild, 1977. In the following list the specimens upon which the species are based are designated and the collectors' names ab- breviated as indicated below: MW M. Wetherill JWB J. W. Brewer SPB S. P. Brewer EL E. Lehnert EJ E. Jackson WSP W. S. Phillips JMR J. M. Rominger SN M. Skougard and G. Nebeker JF J. Fairchild JDB J. D. Brotherson Each species entry is also designated as to certain plant characteristics and according to the section of the monument in which it was found growing. These abbreviations are as follows: p Perennial B Biennial A Annual N Native I Introduced T Tree S Shrub F Forb G Grass GI Sedge NV Non-vascular V Vine b Betatakin k Keet Seel i Inscription House The following list of families, genera, and species is arranged in alphabetical order for ease of reference. Plant identification fol- lows Arizona Flora (Kearney and Peebles 1960), Seed Plants of Northern Arizona (McDougal 1973), Manual of the Plants of Colorado (Harrington 1964) and Utah Plants (Welsh and Moore 1973). Taxa Subdivision Lycopsida Selaginella Selaginella mtitica D. C. Eaton (WSP 3617, 19 June 1964) Plant C haracteristics PNNVb March 1978 Brotherson et al.: Navajo Monument Plants 23 Subdivision Sphenopsida Equisetaceae Equisetum hyemale L. (WSP, 19 June 1964) PNFb Subdivision Pteropsida Class Gymnospennae Cupressaceae Junipenis osteosperma (Torr.) Little (SN 156; JF 4, 25 July 1977) PNTbik Ephedraceae Ephedra viridis Coville (SN 125; JF 256, 17 May 1977) PNSbik Pinaceae Pinus edulis Engelm. (SN 94; JMR 389-65, 22 August 1965) PNTbik Pseudotsuga menziesii (Mirb.) Franc (SN) PNTbk Class Angiospeimae Subclass Dicotyledoneae Aceraceae Acer negundo L. (JWB NB 17, 7 June 1934; SN 264, 17 May 1973) PNTbk Amaranthaceae Amaranthus albus L. (JF 130, 25 June 1977) ANFi Amaranthus leucocarpus Wats. (JF 18, 28 August 1977) AIFk Anacardiaceae Rhus trilohata Nutt. (SN 227, 3 July 1972) PNSbik Toxicodendron radicans L. (MW 413) PNSb Apocynaceae Apocynum medium Greene (MW 431, 10 June 1935) PNFb Asclepiadaceae Asclepias subverticillata (Gray) Vail (SN 175, 2 July 1972; JF 62, 1 September 1977) PNFi Funastrum heterophyUum (Engelm.) Standi (MW 221; JF 19, 28 August 1977) PNFk Berberidaceae Mahonia repens G. Don (SN 269, 17 May 1973; MW 459, 21 May 1935; EL, 17 April 1960; JWB E 1, 9 May 1941) PNSbk Betulaceae Betula occidentalis Hook. (SN 145, 1 July 1972; MW 457; WSP 3615, 19 June 1964; JWB NB 33, 17 July 1940) PNTb Boraginaceae Cryptantha confertiflora (Greene) Payson (JWB NB 19, 12 June 1939) PNFb Cryptantha crassisepala (Torr. & Gray) Greene (SN) ANFb Cryptantha flava (A. Nels.) Payson (SN 74, 9 June 1972; JF 93, 11 June 1977) PNFbk Cryptantha jamesii (Torr.) Payson (SN 152, 1 July 1972; JF 15, 30 August 1977; JF 50, 22 July 1977) PNFbi Cryptantha pterocarya (Torr.) Greene (SN, 3 July 1972) ANFb Lappula occidentalis (Wats.) Greene (JF 222, 5 May 1977) AIFb Lappula redowskii (Homem.) Greene (SN 70, 9 June 1972; JF 189, 3 June 1977; MW G-2, 15 May 1937) AIFb Lithospermum multiflorum Torr. (JF 40, 14 July 1977) PNFb Cactaceae Coryphantha vivipara (Nutt.) Britton & Rose (JF 90, 10 June 1977) PNSk Echinocereus fendleri (Engelm.) Rumpler (JWB NB 74, 17 June 1941) PNSbik Echinocereus triglochidiatus Engelm. (SN 325, 19 June 1973) PNSb Opuntia erinacea Engelm. (JWB NB 78, 21 June 1941) PNSbik Opuntia fragilis (Nutt.) Haw. (JWB H-4, 19 June 1941) PNSb Opuntia polycantha Haw. (SN 331, 20 June 1973; JF 91, 10 July 1977) PNSbik Opuntia whipplei Engelm. & Bigel. (JF 277, 30 June 1977) PNSi Sclerocactus parviflorus Clover & Jotter (SN) PNSb Sclerocactus whipplei (Engelm. & Bigel.) Britt. & Rose (JWB NB 75, 19 June 1941; SN) PNSbk Capparidaceae Cleome serrulata Pursh (JWB NB 21, 16 June 1939, MW I-l, 8 August 1937; JF 109, 25 June 1977) ANFbik Caprifoliaceae Symphoricarpos vaccinioides Rydb. (SN 310, 2 June 1973) PNSb Caryophyllaceae Arenaria eastwoodiae Rydb. (JF 145, 30 June 1977) PNFbik Chenopodiaceae Atriplex canescens (Pursh) Nutt. (MW 419, 27 June 1935; JF 203, 21 May 1977) PNSbik Chenopodium album L. (JF 278, 13 August 1977) AIFbik Chenopodium fremontii Wats. (MW K-1; JF 242, 30 June 1977) ANFbik Chenopodium glaucum L. (SN 204, 3 July 1972; JF 136, 30 June 1977) AIFik Chenopodium pratericola Rydb. (JF 160, 30 June 1977) ANFik Kochia scoparia (L.) Schrad. (JF 115, 26 June 1977) AIFi Salsola iberica Sennen & Pau (SN) AIFbik 24 Great Basin Naturalist Vol. 38, No. 1 Compositae (Asteraceae) Achillea millefolium L. (SN .349, 24 July 1973) PNFbk Ambrosia acanthicarpa Hook. (JF 2.5, 19 July 1977; JF 60, 1 September 1977) ANFik Ambrosia artemisiifolia L. (JF 126, 25 June 1977) ANFik Antennaria rrtarginata Greene (JF .39, 14 July 1977) PNFb Antennaria neglecta Greene (SN 150, 1 July 1972) PNFb Antennaria parvifolia Nutt. (SN 120; JWB NB 58) PNFb Artemisia campestris L. (JF 239, 16 June 1977) PNFb Artemisia dracunndus L. (JWB NB .37, 17 July 1940; JF 103, 25 June 1977) PNFbik Artemisia frigida Willd. (SN 359, 24 July 1973; JF 202, 21 May 1977; MW 451) PNSbik Artemisia ludoviciana Nutt. (JF 87, 13 July 1977; JF 107, 25 June 1977) PNFbik Artemisia nova A. Nels. (JF 279, 25 August 1977) PNSbk Artemisia pacifica Nutt. (JF 3, 16 August 1977) PNFb Artemisia tridentata Nutt. (MW 451; SN 360, 24 July 1973) PNSbk Aster arenosus (Heller) Blake (JF 233, 30 June 1977) PNFb Aster hirtifolius Blake (SPB M-.5, 23 June 1941; SN 316, 19 June 1973) PNFb Brickellia californica (Torr. & Gray) Gray (JF 129, 25 June 1977; JF 151, .30 June 1977; PNSbik Brickellia grandiflora (Hook.) Nutt. (JF 228, ,30 June 1977) PNFk Brickellia scahra (Gray) A. Nels. (JF 1.33, 25 June 1977) PNSbik Chaenuctis stevioides H. & A. (SN .327, 20 June 1973) ANSbik Chrysotharnnus depressus Nutt. GF 264, 16 May 1977) PNSi Chrysothamnus nauseosus (Pall.) Britton (JF 121, 25 June 1977) PNSik Chrysothamnus pulchellus (Gray) Greene (JWB, 24 July 1946) PNSbik Chrysothamnus viscidiflorus (Hook.) Nutt. (JF 263, 16 May 1977) PNSbik Cirsium pulchellum (Greene) Wool. & Standi. (JF 174, 30 June 1977) BNFbk Conyza canadensis (L.) Cronquist (JF 1,55, 30 June 1977) ANFb Erigeron concinnus (H. & A.) Torr. & Gray (SN 313, 19 June 1973; JF 88, 13 July 1977; JF 141, .30 June 1977) PNFbk Erigeron divergens Torr. & Gray (SN .304, 1 June 1973; JWB NB-69, 19 June 1941; JWB NB-2, 21 May 1939; JF 13, 30 August 1977) ' BNFbi Erigeron flagellaris Gray (JF 24, 20 July 1977; JF 54, 22 July 1977) BNFbi Erigeron gUdwllus Nutt. (SN 1&3, 1 July 1972) BNFb Erigeron macranthus Nutt. MW M-7; JF 48, 14 July 1977; JF 55, 2 July 1977) PNFb Erigeron utahensis Gray (JF 29, 1 September 1977; JF 56, 22 July 1977; JF 1.38, .30 June 1977) PNFbi Gaillardia pinnatifida Torr. (SN 97, 10 June 1972) PNFb Haplopappus nuttallii Torr. & Gray (SN 1.37; MW 919-3157, 2 September 19.38; JWB M-10; 23 June 1941) PNFbik Haplopappus scopulorum (Jones) Blake (JF 12, 30 August 1977; JF 152, 30 June 1977) PNFbi Heterotheca villosa (Pursh) Shinners (SN 177, 2 July 1972; JF 33, 1 September 1977; JF 68, 25 July 1977; JF 108, 25 June 1977) PNFbik Hymenopappus filifolius Hook. (SN 78, 9 June 1972; JF 44, 7 July 1977; JF 86, 13 July 1977; JF 89, 11 June 1977) PNFbk Hymenopappus lugens Greene (JWB Mil, 17 June 1941) PNFb Hymenoxys acaulis (Pursh) K. F. Parker (JWB NB 23, 12 June 19.39; JF 45, 7 July 1977) PNFb Hymenoxys higelovii (Gray) K. F. Parker (SN ,301, 1 June 1973; JF 83, 13 June 1977) PNFb Hymenoxys ivesiana (Greene) Parker (JWB NB 23, 12 June 19,39) PNFb Hymenoxys richardsonii (Hook.) "Cockerell (JF 235, 13 June 1977) PNFb Lactttca pulchella (Pursh) DC. (SN ,148, 22 July 1973; JWB M-12, 28 July 1940; JF 6, 25 July 1977) PNFbk Machaeranthera grindelioides (Nutt.) Shinners (SN 137; MW; JF 143, .30 June 1977) PNFbk Machaeranthera hansonii Nels. (JF 11, .30 August 1977) PNFi Machaeranthera linearis Greene (JMR ,343 65, 21 August 1965) PNFbk Machaeranthera tephrodes (Gray) Greene (JF 240, ,30 June 1977) BNFb March 1978 Brotherson et al.: Navajo Monument Plants 25 Malacothrix sonchioides (Nutt.) Torr. & Gray (SN) ANFbi Psilostrophe sparsiflora (Gray) A. Nels. (SN 161, 1 July 1972; JF 114, 26 June 1977; MW M-13; JF 139, 30 June 1977) FNFb Senecio longilobattts Benth. (JF 67, 1 September 1977; JF 95, 11 June 1977) PNFbi Senecio multilobattis Torr. & Gray (JWB NB 70, 19 June 194i; JF 186, 3 June 1977) PNFbk Senecio uintahensis (A. Nels.) Greene (SN 100, 10 June 1972) FNFb Solidago canadensis L. (SN 172, 1 July 1972) FNFb Solidago occidentalis (Nutt.) Torr. & Gray (SN 350, 24 July 1973) PNFb Stephanoineria exigua Nutt. (JMR .345-65, 21 August 1965; SN, 3 July 1972; JF 105, 25 June 1977) ANFb Stephanomeria tenuifolia (Torr.) Hall (JDB 2502, 3 August 1977) PNFb Stephanomeria thurheri Gray (JF 53, 22 July 1977) PNFb Taraxacum officinale Weber (SN 273, 17 May 1973) PIFb Tetradymia canescens DC. (SN 358, 24 July 1973; JF 161, 30 June 1977) PNSbik Townsendia incana Nutt. (JWB M-16, 18 June 1941; SN 62, 9 June 1972; JF 206, 21 May 1977) PNFb Verbesina encelioides (Cav.) Benth. & Hook. (JF 1, 1 September 1977) ANFi Xanthocepliahim lucida (Greene) Shinners (JWB NB-30, 3 August 1940) PNSb Xanthocephahim sarothrae (Pursh) Shinners (MW M-9, 2 September 1937; SN 157, 11 June 1972; JF 255, 17 May 1977) PNSbik Cornaceae Cornns stolonifera Michx. (JWB NB 39, August 1940; MW 41, May 1935; SN 308, 2 June 1973) PNSb Crassulaceae Sedttm stenopetahtm Pursh (JWB NB 54, 28 May 1941) PNFb Cruciferae (Brassicaceae) Arabis perannans S. Wats. (JWB NB 46, 19 April 1941; JF 220, 5 May 1977) PNFbi Arabia pidchra Jones (EL, 17 April 1960) PNFb Descurainia pinnata (Walt.) Britton (JWB P-4, 23 June 1941; JF 156, 30 June 1977) ANFbk Descurainia sophia (L.) Webb. (EL, 17 April 1960; SN 73, 9 June 1972) AIFbik Erysimum asperum DC. (JWB NB 52, 30 May 1941; JWB NB 7, 11 April 1939; SN 286, 21 May 1973; JF 225, 5 May 1977) PNFbk Lepidium montanum Nutt. (SN 195, 3 July 1972; JF 63, 1 September 1977) PNFbik Lesquerella ludoviciana (Nutt.) S. Wats. (JWB NB 8, 20 May 1936; SN 303, 1 June 1973) PNFb Lesquerella rectipes Woot. & Standi. (JF 184, 3 June 1977) PNFb Sisymbrium altissimum L. (JR 389-65, 22 August 1965; JF 158, 30 June 1977) AIFbk Streptanthella longirostris (S. Wats.) Rydb. (SN) ANFbik Streptanthus cordatus Nutt. ex Torr. & Gray (SN 75, 9 June 1972; JWB NB 34, 17 July 1940; JWB NB 59, 20 April 1941; EL, 17 April 1960; JF J 35, 30 June 1977) PNFb Thelypodium integrifolium (Nutt.) Endl. (JF 2, 24 July 1977) BNFk Tlikspi fendleri Gray (SN; JWB NB 47, 19 April 1941) PNFb Elaeagnaceae Elaeagnus angustifolia L. (JF 106, 25 June 1977) PITi Shepherdia rotundifolia Parry (SN 281, 17 May 1973; JF 217, 5 May 1977) PNSbik Ericaceae Arctostaphylos pungens H. B. K. (MW 109) PNSb Euphorbiaceae Euphorbia hirida Engelm. (JWB NB 56, 28 May 1941) PNFb Euphorbia micrarnera Boiss. (JF 26, 19 July 1977) ANFi Fagaceae Quercus gambelii Nutt. (SN 160, 1 July 1972; JF 214, 21 May 1977; JWB NB 40, 17 July 1940) PNTbk Quercus turbinella Greene (JF 27, 1 September 1977) PNSi Fumariaceae Corydalis aurea Willd. (SN 201) ANFb Geraniaceae Erodium cicutarium (L.) L'Her (SN 252, 8 May 1973) AIFbik Geranium atropurpureum Heller (SN 116, 10 June 1972; JWB NB 28, 25 July 1940; JWB NB 42, 17 July 1940; JF 75, 12 July 1977) PNFb Hydrophyllaceae 26 Great Basin Naturalist Vol. 38, No. 1 Phacelia corrugata A. Nels. (SN 245, 8 May 1973) ANFb Phacelia ivesiana Torr. (JWB 1080-1723a, 23 June 1941) ANFbik Labiatae (Lamiaceae) Moldovica parviflora (Nutt.) Britt. & Brown (SN 218, 3 July 1973) BNFb Leguminosae (Fabaceae) Astragalus amphioxys Gray (SN) PNFb Astragalus ceramicus Sheld. (SN 103, 10 June 1972; JWB NB 10, 11 May 1939; JF 81, 16 June 1977) PNFb Astragalus lentiginosus Dougl. ex Hook. (SN 251, 8 May 1973; JF 70, 20 July 1977; JF 132, 25 June 1977) ' PNFbik Astragalus mollissimus Torr. var. thompsonae (Wats.) Bameby (SN 101, 10 June 1972; JF 84, 13 June 1977) PNFb Astragalus sesquiflorus S. Wats. (EL, 17 April I960; SN 276, 17 May 1973) PNFb Astragalus zionus Jones (JF 317, 27 July 1977; JF 178, 5 May 1977) PNFbik Lathyrus arizonicus Britton (JF 261, 17 May 1977) PNFb Lathyrus brachycalyx Rydb. (SN 111, 10 May 1972; JF 211, 21 May 1977; JWB X-3, 11 May 1941) PNFb Lupinus argenteus Pursh (JF 35, 27 July 1977) PNFb Medicago sativa L. (SN 105, 10 June 1972) PIFb Melilotus alba Descr. (JF 57, 22 July 1977) BIFb Melilotus officinalis (L.) Lam. (JF 58, 22 July 1977) BIFb Linaceae Linum aristatum Engelm. ex Wisliz. (MW Z-1, 13 August 1937; SN; JF 30, 1 September 1977) ANFi Linum perenne L. (MW 65, 27 May 1935) PNFb Loasaceae Mentzelia alhicaulis Dougl. (JWB NB 5, 11 May 1939; SN .338, 29 June 1973) ANFb Loranthaceae Arceuthobium campylopodium Engelm. (MW AB-1, 23 August 1937) PNFb Phoradendron juniperinum Engelm. (SN 198) PNFb Malvaceae Sidalcea neomexicana Gray (SN) PNFb Sphaeralcea coccinea (Nutt.) Rydb. (SN 335, 29 June 1973; JF 71, 20 July 1977; JF 85, 13 June 1977) PNFbik Sphaeralcea parvifolia A. Nels. (JWB NB 72, 19 June 1941; SN 68, 2 Jime 1972; JF 248, 16 May 1977) PNFbik Nyctaginaceae Abronia elliptica A. Nels. (JWB NB 51, 27 May 1941; JF 187, 3 June 1977) PNFbik Abronia fragrans Nutt. ex Hook. (SN) PNFb Allionia incarnata L. (JF 65, 1 September 1977) PNFi Allionia linearis Pursh (JWB NB 20, 12 June 1939) PNFb Mirabilis comatus Standi. (SN .361, 24 July 1973) PNFk Mirabilis multiflora (Torr.) Gray ex Torr. (SN 174, 2 July 1972; JF 21, 20 July 1977; JF 112, 25 June 1977) PNFi Mirabilis oxybaphoides Gray (JF .34, 27 July 1977) PNFk Oxybaphus linearis (Pursh) Robins (JWB NB 20) PNFb Tripterocalyx wootonii Standi. (JMR ,385-65, 22 August 1965; JWB NB 67, 19 May 1941; JF 31, 1 September 1977) ANFbi Oleaceae Fraxinus anornala Torr. (MW 297) PNTi Onagraceae Epilobiwn hornemanii Reichenb. (JF 74, 24 July 1297) PNFk Oenothera albicatdis Pursh (SN 106, 10 June 1972) ANFbi Oenothera caespitosa Nutt. (SN 194; JF 92, 10 June 1977) PNFbik Oenothera hookeri Torr. & Gray. (JWB, 1941) BNFb Oenothera longissima Rydb. (JF 43, 12 July 1977) BNFb Oenothera pallida Lindl. (JR 398-65, 22 August 1965; SN 217, 3 July 1972; JF 207, 21 May 1977) PNFi Plantaginaceae Plantago purshii Roem. & Schult. (SN 333, 27 June 1973) PNFb Polemoniaceae Gilia aggregata (Pursh) Spreng. (SN 58, 9 June 1972; JF 41, 18 July 1977) PNFbik Gilia leptomeria Gray (SN 283, 17 May 197.3) ANFbik Gilia longiflora (Torr.) G. Don. (JMR 341-65, 21 August 1965; JF 10, .30 August 1977; JF 4, 16 August 1977) ANFbi Gilia scopulorum Jones (SN) ANFb Gilia subnuda Torr. ex Gray March 1978 Brotherson et al.: Navajo Monument Plants 27 (SN) BNFb Leptodactylon piingens (Torr.) Nutt. (SN 126, 10 June 1972; JF 146, 30 June 1977) PNSbk Phlox austromontana Coville (JF 262, 16 May 1977) PNFb Phlox longifolia Nutt. (SN) PNFbk Polygonaceae Eriogonum alatiim Torr. in Sitgr. (SN 197, 3 July 1972; JF 46, 7 July 1977) PNFb Eriogonum cernuum Nutt. (SN 235, 20 July 1972) ANFb Eriogonum corymbosum Benth. in DC (JF 281, 1 September) PNSi Eriogonum microthecum Nutt. (EJ, 16 August 1961; JMR 338-65, 21 August 1965; SN 239, 24 July 1972; JF 244, 8 May 1977) PNSb Eriogonum umbellatum Torr. (JWB NB 35, 17 July 1940; JMR 334-65, 21 August 1965; JF 32, 1 September 1977) PNSi Eriogonum umbellatum Torr. var. cognatum Greene (JF 280, 1 September 1977) PNFb Portulacaceae Portulaca oleracea L. (JF 14, 30 August 1977; JF 69, 20 July 1977) AIFi Portulaca retusa Engelm. (JF 73, 27 July 1977) AIFk Talinum brevifolium Torr. (SN 173, 2 July 1972; JF 16, 30 August 1977) PNFi Primulaceae Androsace septentrionalis L. (JF 47, 14 July 1977) ANFb Ranunculaceae Aquilegia micrantha Eastw. (JWB NAV-B-101-47, 17 June 1947; JWB NB 31, 14 July 1940; SN 93, 10 June 1972) PNFb Clematis ligusticifolia Nutt. (SN 229, 3 July 1972) PNVb Delphinium amabile Tidestr. (SN) PNFb Delphinium nelsonii Greene (JWB 1080-1704, 27 May 1941; JWB NB 6, 11 May 1939; SN 302, 1 June 1973) PNFb Delphinium scaposum Greene (JWB Am-4, 17 June 1941) PNFb Ranunculus cymbalaria Pursh (JF 111, 25 June 1977; JF 125, 25 June 1977; MW AM-5) PNFi Thalictrum fendleri Engelm. (JWB NB 44, 2 September 1940; JWB NAV-B-101-47, 17 June 1947) PNFb Rosaceae Amelanchier utahensis Koehne (SN 129, 10 June 1972; JF 213, 21 May 1977; SPB NB 62, 18 June 1941) PNSb Cercocarpus intricatus S. Wats. (SN 261, 17 May 1973; JWB NB 63, 12 May 1941; MW 452, JF 218, 5 May 1977) PNSbik Cowania mexicana D. Don. (SN 65) PNSb Fallugia paradoxa (D. Don.) Endl. (JF 64, 1 September 1977) PNSi Holodiscus dumosus (Nutt.) Heller (MW 919-3044; JWB 1080-1709, June 1942; MW 434) PNSb Prunus emarginata (Dougl.) D. Dietr. (SN 288, 24 May 1973; JF 163, 30 June 1977) PNTk Prunus virginiana L. (SN 309, 2 June 1973) PNTb Purshia tridentata (Pursh) DC. (SN 307, 1 June 1973) PNSbk Rosa woodsii Lindl. (SN 113, 10 June 1972; JWB NB 16, 7 June 1939; JF 157, 30 June 1977) PNSb Rubiaceae Galium aparine L. (JF 166, 30 June 1977) ANFk Galium triflorum Michx. (SN 154, 1 July 1972) PNFb Kelloggia galioides Torr. (JF 241, 3 June 1977) PNFbk Salicaeae Populus angustifolia James (JF 281, 30 June 1977) PNTi Populus fremontii S. Wats. (SN 179) PNTbi Populus tremuloides Michx. (SN 171) PNTb Salix exigua Nutt. (SN 181, 2 July 1972) PNSbi Salix goodingii Ball (SN; JF 128, 25 June 1977) PNSbk Salix lasiolepis Benth. (JF 208, 21 May 1977) PNTb Santalaceae Comandra pallida DC. (JF 149, 30 June 1977; JF 196, 31 May 1977) PNFbk Saxifragaceae Fendlera rtipicola Gray (SN 289, 24 May 1973; JF 212, 21 May 1977) PNSbik Heuchera parvifolia Nutt. (SN 118, 10 June 1972) PNFb Heuchera rubescens Torr. (SN 311, 19 June 1973) PNFb Ribes cereum Dougl. (JF 142, 30 June 1977; JF 204, 21 May 1977) PNSbk Ribes leptanthum Gray (JF 169, 30 June 1977; JF 205, 21 May 1977) PNSk Ribes viscossissimum Pursh (SN 28 Great Basin Naturalist Vol. 38, No. 1 290, 24 May 1973) FNSk Carex rossii Boott (JF 172, 16 Scrophulariaceae June 1977) PNGlb Castilleja UnarifoUa Benth. Eleocharis nuicrostachya (SN 96, 10 June 1972; JF 49, Britton (JF 28, 1 September 22 July 1977; JF 134, 30 June 1977; JF 1 19, 25 June 1977) PNGLbi 1977) PNFbik Gramineae (Poaceae) CordyUmthns wrightii Gray Agropyron caninurn (L.) Beauv. (JMR .340-65, 21 Avigust 1965; (SN 85, 10 June 1972; JF 182, EJ, 16 August 1961) ANFb 3 June 1977) PNGbk Mimulus eastwoodiae Rydb. (SN Agrostis exarata Trin. (SN 182, 2 221; JF 20, 20 Julv 1977; JF July 1972) PNGi 165, 30 June 1977) PNFbk Agrostis semiverticillata (Forsk.) Pensternon umbiguus Torr. (EL; Christ. (SN 356, 24 July SN; JF 230, 18 May 1977) PNSb 1973; JF 38, 27 July 1977; JF Pensternon barbattis (Cav.) 118, 25 June 1977) Pick Roth (JF 59, 22 July 1977) PNFb Alopecurus aequalis Sobol. (SN Pensternon cotnarrhenus Gray .354, 24 July 1973) PNGk (JWB 27 June 1947; JF 162, Andropogon hallii Hack. (JF 66, 30 June 1977) PNFli 1 September 1977) PNGi Pensternon Eatoni Gray (SN 108, Aristida arizonica Vasey (SN 10 June 1972; JDB 2503, 3 188, 2 July 1973) PNGb August 1974) PNFb Aristida fendleriana Steud. (JF Pensternon virgatiis Gray (JF 52, 100, 25 June 1977; JF 117, 25 12 July 1977) PNFb June 1977) PNGbi Solanaceae Bouteloua barbata Lag. (JF 72, Charnaesarcha coronopus 1 September 1977) ANGi (Dunal) Gray (SN 183, 2 July Bouteloua curtipendula (Michx.) 1972) PNFi Torr. (SN .347, 29 June 1973) PNGi Datura meteloides DC. (JF 282, Bouteloua gracilis (H.B.K.) Lag. 30 June 1977) ANFbik (JF 284, 30 June 1977) PNGb Lyciitrn pallidum Miers. (SN; JF Brornus carinatus Hook. & Am. 98, 26 June 1977) PNSi (SN 224, 3 July 1972) PNGbk Nicotiana attenuata Torr. (JMR Brornus ciliatus L. (SN 84, 10 400-65, 22 August 1975) PNFi June 1972; JF 181, 3 June Physalis fendleri Gray (SN 158, 1 1977) PNGb July 1972; JF 137, .30 June Brornus marginatus Nees (JF 1977) PNFlii 168, 30 June 1977) PNGb Physalis hederaefolia Gray (JF Brornus rubens L. (JF 104, 25 153, .30 June 1977) PNFk June 1977) AIGik Solanum jamesii Torr. (JF 17, Brornus tectorum L. (SN 207, 3 28 August 1977) PNFk July 1973; JF 192, 3 June Tamaricaceae 1977) AlGbik Tarnarix ratnosissirna Ledels. Elymus glaucus Buckl. (JF 42, (JF 2a3, .30 June 1977) PISi 18 July 1977) PNGb Uinbelliferae (Apiaceae) Glyceria striata (Lam.) Hitch. Cymopterus newberryi (S. (JF 7, 25 July 1977) PNGb Wats.) M. E. Jones (JWB, 16 Hordeurn Leporinurn Link (SN) ANGb May 1941) PNFb Lycurus phleoides H.K.B. (JF Urticaceae 23, 20 July 1977) PNGi Parietaria pennsylvanica Muhl. Muhlenbergia andina (Nutt.) (JF 123, 25 June 1977) ANFi A. S. Hitchc. (JF 8, 1 Verbenaceae September 1977) PNGb Verbene bracteata Lag. & Rodr. Muhlenbergia curtifolia (SN a37, 29 June 1973) PNFb Scribn. (JMR .3.32-65, 21 Subclass Monocotyledoneae August 1965) PNGi Comnielinaceae Muhlenbergia pungens Thurb. Tradescantia occidentalis gMR .342-65, 21 August 1965; (Britt.) Smyth (SN 151; JF SN 243, 27 July 1972) PNGb 80, 16 June 1977; JWB NB Muhlenbergia thurheri Rydb. (JF 22, 12 June 1939; JMR 120, 25 June 1977) PNGbi 335-65, 22 August 1965) PNFbik Munroa squarrosa (Nutt.) Torr. Cyperaceae (SN; JF 9, 30 August 1977; JF Carex occidentalis Bailey (JF 51, 61, 1 September 1977) ANGi 12 Julv 1977; JF 173, 16 June Oryzopsis Hymenoides (R. & S.) 1977) ' PNGb Riker (SN'343, 29 June 1973) PNGbik March 1978 Brotherson et al.: Navajo Monument Plants 29 Oryzopsis micrantha (Trin. & Rupr.) Thurb. (SN 86, 10 June 1972; JF 179, 3 June 1977) PNGb Poa fendleriana Vasey (MW 7; MW 28; SN 275, 17 May 1973) PNGbk Poa longiligula Scribn. & Williams (SN; MW 14; JF 94, 13 June 1 977; JF 99, 25 June 1977) PNGbik Poa pratensis L. (SN 357, 24 July 1973; JF 167, 30 June 1977) PlGbk Polypogon monspeliensis (L.) Desf. (SN 351, 24 July 1973; JF 110, 25 June 1977) AIGbik Puccinellia airoides (Nutt.) Wats. & Coult. (JF 170, 30 June 1977) PNGbk Sitanion hystrix (Nutt.) J. G. Smith (SN 323, 19 June 1973; JF 82, 13 June 1977) PNGbik Sporobolus airoides Torr. (JF 22, 19 July 1977) PNGik Sporobolus cryptandrus (Torr.) Gray (JMR .344-65, 21 August 1965; SN 228, 3 July 1972; JF 96, 25 June 1977) PNGbik Sporobolus flexuosus (Thurb.) Rydb. (SN) PNGb Stipa coniata Trin. & Rupr. (SN 80; JF 180, 3 June 1977) PNGbk Vulpia octoflora (Walt.) Rydb. (SN; JF 269, 18 May 1977) ANGbik Juncaceae ]uncus balticus Willd. (JF 171, 16 June 1977) PNGLbi Juncaginaceae Triglochin nuiritima L. (JF 227, 26 June 1977) PNGLi Liliaceae Allium macropetalum Rydb. (JWB NB 11, 20 May 1939; SN 326, 20 June 1973) PNFb Calochortus nuttallii Torr. & Gray (SN 99, 10 June 1972; JWB NB 13, 10 June 1939) PNFb Fritillaria atropurpurea Nutt. (SN 287, 23 May 1973; MW 287, May 1930) PNFb Sinilacina stellata (L.) Desf. (MW Y-4; SN 148, 3 July 1972; SN 223, 3 July 1972; JF 154, 30 June 1977) PNFbk Yucca angustissima Engelm. (SN 56, 9 June 1972; JWB NB 76, 21 June 1941) PNSbik Yucca baccatu Torr. (SN 315, 19 June 1973) PNSb Orchidaceae Habenaria sparsiflora S. Wats. (SN 167, 1 July 1972) PNFb Sparganiaceae Sparganium eurycarpum Engelm. (JF 226, 26 June 1977) PNGLi Literature Cited Allan, J. S. 1977. The plant communities of Arches National Park. Unpublished Ph.D. dissertation, Brigham Young University, Provo, Utah. 98 pp. Brotherson, J. D. 1967. A study of certain community relationships of Eriogonum corymbosum Benth. In: DC in the Uintah Basin, Utah. Unpublished master's thesis, Brigham Young University, Pro- vo, Utah. 100 pp. Campbell, V. O. 1977. Certain edaphic and biotic fac- tors affecting vegetation in the shadscale com- munity of the Kaiparowits area. Unpublished master's thesis, Brigham Young University, Pro- vo, Utah. 58 pp. CooLEY, M. E. 1958. Physiography of the Glen-San Juan Canyon area. Part 1. Plateau 31:21-33. 1962. Late Pleistocene and Recent erosion and alluviation in parts of the Colorado River sys- tem, Arizona and Utah. In: Geological Survey Research 1962: short papers in geology, hydro- logy, and topography, articles 1-59, pp. 48-50. United States Geological Survey Professional Pa- per 450-B. Washington. Dean, J. S. 1969. Chronological analysis of Tsegi phase sites in northeastern Arizona. Papers of the Lab. of Tree-ring Res. No. 3. University of Ari- zona Press. Tuscon, Arizona. 207 pp. Flowers, S. 1963. The lichen and moss flora of Be- tatakin Canyon and vicinity, Navajo National Monument, Arizona. University of Utah Div. of Biol. Sci. Misc. Papers No. 3. 10 pp. Gregory, H. E. 1916. The Navajo country: a geogra- phic and hydrographic reconnaissance of parts of Arizona, New Mexico, and Utah. United States Geological Survey Watersupply paper. Washington. 1917. Geology of the Navajo country: a recon- naissance of parts of Arizona, New Mexico, and Utah. United States Geological Survey Profes- sional Paper 93. Washington. Hack, J. T. 1942. The changing physical environment of the Hopi Indians of Arizona. Papers of the Peabody Museum of American Archaeology and Ethnology, Harvard University. Vol. 35, No. 1. Reports of the Awatovi Expedition, No. 1. Cam- bridge. 1945. Recent geology of the Tsegi Canyon. Ap- pendix 1 in R. L. Beals, George W. Brainerd, and Watson Smith, Archaeological Studies in Northeast Arizona. University of California Press, Berkeley and Los Angeles. Harshbarger, J. W., C. A. Repenning, and J. H. Irwin. 1957. The stratigraphy and the up- permost Triassic and Jurassic rocks of the Nav- ajo country. United States Geological Survey Professional Paper 291. Washington. Harrington, H. D. 1964. Manual of the plants of Col- orado. The Swallow Press, Inc. Chicago, Illinois. 666 pp. 30 Great Basin Naturalist Vol. 38, No. 1 Kearney, T. H., and R. H. Peebles. 1960. Arizona flora. University of California Press. Berkeley, California. 1085 pp. McDouGAL, W. B. 1973. Seed plants of northern Ari- zona. The Museum of Northern Arizona, Flag- staff, Arizona. 594 pp. Sellers, W. D. 1964. The climate of Arizona. Pp. 3-64. In: Christine R. Green and William D. Sellers, eds., Arizona Climate. University of Ari- zona Press, Tucson, Arizona, 175 pp. Welsh, S. L. and G. Moore. 1973. Utah plants- Tracheophyta. Brigham Young University Press. Provo, Utah. 474 pp. Woodbury, A. M. 1963. Biological-ecological aspects of Betatakin Canyon, Navajo National Mon- ument, Arizona. University of Utah Div. of Biol. Sci. Misc. Papers No. 2. 56 pp. ENVIRONMENTAL INTERACTION IN SUMMER ALGAL COMMUNITIES OF UTAH LAKE' Mark C. Whiting,^ Jack D. Brotherson,^ and Samuel R. Rushforth'' Abstract — Utah Lake is a shallow eutrophic lake in central Utah. It is characterized by high nutrient and silt loads and by large algal blooms in late summer and early fall. Phytoplankton samples and environmental data were taken from June through August 1974. Phytoplankton species were identified and then quantified in a Pal- mer counting cell. Environmental continuum theory was employed to describe algal succession, and regression analysis was used to discover interactions between algal communities and the environment. Phytoplankton com- munities in June were characterized by high species diversity. As the lake environment became stressed in late summer due to higher turbidity, nutrient levels, pH, and available inorganic carbon species diversity decreased. By August, the phytoplankton flora was composed essentially of only two species, Ceratium hirundinella and Aphanizovienon flos-aqtiae. Utah Lake is a shallow eutrophic lake in central Utah (Fig. 1). It is the largest natu- rally occurring freshwater lake in the state, covering some 388 km- (Bolland 1974). Wa- ter from the lake is presently used for irri- gation and water regulation as well as for recreational boating and fishing. In the past, commercial fisheries on Utah Lake have been an important resource for the state of Utah. At the time of settlement, the fish population was dominated by a va- riety of Bonneville cutthroat trout {Salmo clarki) which was adapted to the eutrophic conditions of the lake. The trout fishery be- came vital to the survival of the early Mor- mon pioneers during the drought and crop failures from 1855 to 1858. However, with water manipulation for agriculture and the introduction of exotic fish species, the trout rapidly became extinct in subsequent years. During the depression years of 1929 through 1939 commercial fishing for in- troduced species, mainly carp and white bass, became an important industry and food source. Current use of Utah Lake fish- eries is minimal and limited to carp, which are used for fish meal. Utah Lake is characterized by late sum- mer and early fall algal blooms, nutrient en- richment, high silt load, and total dissolved solids and other environmental stresses. Due to the shallowness of the lake (average depth 2.4 m), fine silt-clay sediments are of- ten stirred up by storms, giving the lake water a characteristic gray-green color. The average summer Secchi disk reading is 24 cm, with a range of 12 to 50 cm. In addi- tion, the lake basin receives the flow of nu- merous mineral springs high in carbonates and sulfates. During late summer, when water levels are lowest, the lake approaches a slightly saline ecosystem. According to the U.S. Geological Survey (Hem 1970), lakes with 1,000-3,000 mg/ liter dissolved solids can be considered to be slightly saline. Summer values for dissolved solids in Utah Lake range from 795 to 1,650 mg/1 and are thus in the lower part of this range. Previous algal studies of Utah Lake were done by Tanner (1930, 1931), Snow (1932), Harding (1970, 1971), and Bolland (1974). Tanner's pioneering works listed several of the algae prominent in the lake. Both Har- ding and Snow did taxonomic studies deal- ing with littoral and planktonic algae. Bol- land's work dealt with the fossil diatom flora in the lake sediments. Bolland's re- search indicates that the diatom flora has not changed greatly since presettlement times. Data for this study were gathered during 'This work was financed in part by the National Science Foundation and Brigham Young University. 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 31 32 Great Basin Naturalist Vol. 38, No. 1 UTAH LAKE Provo ) Salt Lake City Utah Lake S^'P"*^" UTAH Fig. 1. Utah Lake, showing geographical position with respect to the state of Utah, Provo, and the Great Salt Lake. March 1978 Whiting et al.: Utah Lake Algae 33 the summer of 1974. There had been no previous quantitative study of the extant planktonic flora. The study resulted in a flo- ristic paper (Rushforth et al., in press) as well as estimates of productivity and a de- scription of the seasonal succession of sum- mer algal species in Utah Lake to be re- ported herein. Methods Phytoplankton samples and environmental data were collected from June to August 1974. Samples were taken along three tran- sects at nine-day intervals throughout the study period. The transects (Fig. 1) included 14 sample sites permanently marked with buoys. Transects were chosen to cross three major portions of the lake, each with pos- sible differences in ecological conditions. The Geneva transect crossed the northern part of the lake from the outfall of the settling ponds of Geneva Works of the United States Steel Corporation to the west- ern shore. The midlake transect ran west from the Provo Boat Harbor (near the mouth of the Provo River) to the west shore. The southern transect crossed Goshen Bay from Lincoln Beach to the west shore. The Geneva and Boat Harbor transects in- cluded five sampling sites, and the shorter Goshen transect had four sampling sites. Phytoplankton samples were collected by pouring known volumes of lake water through a 67-jLim mesh plankton net. Algae were washed from the net, collected in 30- ml vials, and immediately preserved in for- malinacetic acid (FAA). The vials were later subsampled in the laboratory and individual algae were counted in Palmer counting cells (Palmer and Maloney 1954) at 400X magni- fication, using Zeiss RA research micro- scopes. Individual algae encountered were identified to the species level by two of us (Whiting and Rushforth). An "individual" for filamentous or colonial forms was con- sidered to be a single filament or colony. Tallies were made for each species, as well as for the total number of individuals per subsample. The density of organisms in the original lake water was calculated using multiplication factors determined by the volume of filtered lake water. At least 400 individuals were counted in each sample to reduce sample variance (Clark 1956). Selected water chemistry tests were per- formed in the field using a Hach DR/EL-2 Direct Reading Engineer's Lab. Tests for dissolved oxygen, free carbon dioxide, pH, and Jackson Turbidity readings were per- formed. In addition, a YSI conductivity me- ter was used to measure salinity, con- ductivity, and water temperature. Secchi disk readings and general meterological con- ditions were also recorded. Further water chemistry tests were per- formed in the laboratory. Water samples were collected in opaque Nalgene bottles from approximately 25 cm below the water surface and were refrigerated until ana- lyzed. Laboratory analyses included total al- kalinity, carbonate alkalinity, total hardness, calcivmi hardness, magnesium hardness, ni- trate, orthosphosphate, sulfate, and silica. All tests were performed within 24 hours of collection, using standard methods (Taras 1971). Changes in phytoplankton populations through the summer were evaluated, using the continuum methods of Cvirtis and Mcin- tosh (1950, 1951). Continuum theory is an approach to vegetation and its response to environmental gradients. Continuum study involves the calculation of an index number for each sample which places that sample at some point along an environmental gradient. The index number is considered to reflect the effects of the total environment on a sample expressed in terms of the spe- cies composition and their relative abun- dance. To demonstrate succession, the gradient herein is generated as a time con- tinuum. To generate the continuum index num- bers used, the average density (in numbers of organisms/liter) for each species was cal- culated for all sites on each sample date. Adaptation numbers assigned to each algal species reflect the date at which each achieved maximum abundance. The adapta- tion numbers ranged from 1 to 8, corre- sponding to eight sample dates beginning on 13 June 1974 and ending 15 August 1974. The adaptation numbers for all spe- cies present in any one sample were then summed and averaged. The average adapta- 34 Great Basin Naturalist Vol. 38, No. 1 tion number (continuum index number) for a sample describes its position along the time continuum. Sample index values ranged from 1.58 to 7.00. No sample was found to contain exclusively early summer species or late summer species. All sample index values were plotted along an axis representing the time contin- uum. The continuum was then divided into six "natural" groups of approximately equal length, utilizing naturally occurring breaks as nearly as possible. The six divisions of the continuum allowed averages of environ- mental and biotic parameters to be calcu- lated for each unit. Such average values show the successional trends along the con- tinuum. The parameters plotted included eight major algal species, nine significant environmental parameters, environmental variation (heterogeneity), community varia- tion, and species diversity. A similarity index matrix using environ- mental data for each stand in each unit of the continuum was constructed. Mean sim- ilarity indices and standard deviation values for each unit of the continuum were then computed. A coefficient of variation was then computed from these values (Gilmartin 1974). All data values were adjusted to range from one to 10 to avoid over- weighting some parameters because of their large numerical values. Community variation was measured as above, except the data utilized were taken from the relative densities of the species present. In this instance, community varia- tion is considered to be a measure of the evenness of the contribution each species makes to a sample. Species diversity was calculated, using the Shannon-Wiener formula, as follows: D = -2pi log Pi The term p^ refers to the portion of the sample that each species contributes. The Shannon-Wiener formula expresses diversity in terms of the number of species present as well as the evenness of the contribution that each species makes to the total sample. The response of individual species to single environmental parameters was as- sessed using the linear regression analysis. Those species with significantly similar re- sponses (a = 0.01) to the same environmen- tal parameters were grouped together into "clusters." Results A total of 107 phytoplankton and envi- ronmental samples were taken. Ninety-five species were identified and ranked by im- portance values (average relative density X average percent presence). The six most im- portant species and their importance values were: Ceratiwn hirundinella (3,303), Aph- anizomenon flos-aquae (1,725), Melosira granulata (1,255), Microcystis protocystis (252), Anabaena spiroides (249), and Ana- haena flos-aquae (154) (Fig. 2). The early summer flora can be character- ized by low standing crop (an average of 11,260 ± 25,700 organisms/liter in June) and rich species diversity. The June commu- nities were dominated by several species of green algae, including Ankyra judayi, Schroederia setigeria, Treubaria triappen- diculata, Dictyosphaerium ehrenbergii, Pe- diastrum duplex, and three varieties of An- kistrodesmus falcatus. Associated with these chlorophytes were the blue-greens Ana- baena flos-aquae and Microcystis pro- tocystis, the diatom Melosira granulata, and the dinoflagellate Ceratium hirundinella. By early July, the green algae as a group began to decline in importance and were replaced by Melosira granulata, Ceratium hirundinella, and aphanizomenon flos-aquae (Fig. 3). King (1970) and others have in- dicated that the green algae tend to require free CO2 for maximum growth and are poor competitors for bicarbonate. Our evidence seems to corroborate this conclusion. Free CO2 in Utah Lake declined to levels that were undetectable with Hach chemistry (Fig. 4) at the same time the green algae decreased in importance. The July standing crop averaged 329,425 ± 291,410 organ- isms/liter, almost 30 times the June aver- age. August phytoplankton communities were much reduced in species diversity, often consisting only of two species, Ceratium hi- rundinella and Aphanizomenon flos-aquae. These two taxa were usually 10-50 times March 1978 Whiting et al.: Utah Lake Algae 35 )0 0 Onr OD ood TO oooc tVooooc lOOODQ o o oqpP t)o9o>^ boQOoS^ poV)Oc? 1 ^8 Qd Fig. 2. Dominant phytoplankton species in the Summer Utah Lake flora. 1, Ceratium hiriindinella; 2, Ankyra judayi; 3, Aphanizomenon fhs-aqitae; 4, Melosira granulata; 5, Anabaena flos-aqiiae; 6, Anabaena spiroides; 7, Microcystis protocystis. Each scale equals 10 /xm. 36 Great Basin Naturalist Vol. 38, No. 1 ANABAENA SPIROIDES n — Z UJ ANKYRA JUDAYI oa FLOS -AQUAE 5 6 CURTIS CONTINUUM INDEX Fig. 3. Relative density and percent presence of common algal species in the Utah Lake summer 1974 flora plotted on the Curtis continuum index. March 1978 Whiting et al.: Utah Lake Algae 37 FREE CO2 ^5.0 (0)1.5 •- •■ • • DISSOLVED OXYGEN 5.0 ,- E (0) PHOSPHATES pH SECCHI DISK TEMPERATURE 5 2 3 CURTIS CONTINUUM INDEX 40° C Fig. 4. Selected environmental gradients in Utah Lake in the summer of 1974, plotted on the Curtis contin- uum index. 38 Great Basin Naturalist Vol. 38, No. 1 more abundant than any other species. The decline in other species, especially Melosira granulata, might be attributed to any of a number of factors, such as decrease in avail- ability of inorganic carbon, higher turbidity, higher salinity, or interspecific competition. The average estimate of the August stand- ing crop was 5,405,226 ± 15,719,846 or- ganisms/liter, almost 16 times the average for July and nearly 500 times the average for June (Table 1). Biomass estimations for any given day showed large variation from one sample site to the next. This disjunct distribution was probably due to currents and possibly patchiness of the environment. George and Edwards (1973) have shown that zooplank- ton distribution is strongly correlated with Langmuir circulations. They demonstrated that Daphnia tend to aggregate in up- wellings between foam lines. Presumably, phytoplankton may also be oriented in rela- tion to Langmuir currents. Floating objects, such as blue-green algae with gas vacuoles, should aggregate in the foam lines. Other algae with well developed powers of loco- motion might aggregate between foam lines as do zooplankton. Analysis of phytoplankton using the Cur- tis continuum is summarized in Fig. 3. The six species with the highest importance val- ues are plotted against the continuum. An- kyra jiidayi and "epizooites" (two uniden- tified species of Chrysophyta found on copepods) were also plotted. Although not very important overall, they were included because of their importance in the early summer. Generally, the continuum data in- Table 1. Mean standing crop values of Utah Lake al- gae in the summer of 1974 according to collection date. Mean number of Date organisms per liter 4 June 13,758 13 June 2,230 21 June 20,754 3 July 52,251 10 July 416,128 18 July 541,128 27 July 344,968 7 Aug. 724,061 15 Aug. 10,866,586 dicate similar trends to those already de- scribed. The early summer phytoplankton consisted of a diverse group of diatoms, and green and blue-green algae. Although pres- ent from the beginning of the study, Aph- anizomenon, Ceratium, and Melosira did not become abundant until July. Melosira showed its maximum growth around mid- July. Aphanizomenon and Ceratiwn contin- ued to become more abundant until the end of the study in August, when they made up approximately 90 percent of the total flora. Analysis of environmental parameters us- ing the Curtis continuum is summarized in Figure 4. Generally, the lake became a more stressful system as the season pro- gressed. Availability of inorganic carbon for photosynthesis decreased through the svim- mer to limiting levels (King 1970). Water transparency decreased dramatically in early July and decreased more slowly till the end of the study. Phosphates and nitrates showed maximums in late July, then de- creased slightly in August. Temperature and pH increases were slight. Environmental stresses on the phyto- plankton in the late summer had the effect of reducing biological and environmental di- versity. Figure 5 summarizes diversity trends along the time continuum. Environ- mental variation decreased through the summer, essentially reducing the number of niches available to organisms. Species diver- sity (measured by the Shannon-Wiener in- dex) and community variation (the evenness of the contribution made by each species) also decreased through the summer. The last point on the community variation curve is much higher than the overall trend, due to the fact that in August the algal commu- nities were composed of approximately equal contributions of Ceratium and/or Aphanizomenon, which normally comprised over 90 percent of the algae in a given sample. Simple regression analysis of individual species plotted against environmental gradients shows several significant relation- ships (Table 2). Species showing the same significant trends were grouped into units. The first two groups are essentially early summer species and are predominately green and blue-green algae. As expected. March 1978 Whiting et al.: Utah Lake Algae 39 they generally correlate with environmental parameters that predominate in the early summer, such as high light penetration, free CO2, and low water temperatures. Aphani- zomenon and Ceratium correlate with envi- ronmental parameters that were prevalent in the latter part of the season (i.e., high turbidity, high salinity, high phosphates, basic pH, and higher temperatures). As mentioned previously, increase in temper- ature through the season was slight and, therefore, is probably not a causal relation- ship even though the correlation with Aph- anizomenon and Ceratium was significant. Discussion Communities seldom appear as discrete units. In many cases closely allied commu- nities intergrade one with another both in time and space and often exhibit no distinct boundaries between them. This is especially true of aquatic systems and in instances of biological succession. Therefore, the contin- uum theory as used in this paper is espe- cially useful in the description of the algal communities of Utah Lake. The Curtis con- tinuum has been traditionally used to de- scribe the response of terrestrial vegetation to environmental gradients. However, the principle is just as applicable to aquatic sys- tems where the species involved are mobile and succession is seasonal. The Curtis continuum is also especially useful in environments that are highly fluid, as in the case of planktonic systems. Thus, we have noted in Utah Lake that certain geographical regions maintain early summer floras for more extended periods of time due to local environmental conditions which approximate earlier seasonal conditions. This was noted particularly in the Provo Boat Harbor and Goshen Bay, where spring and river influences are prominent. Summer phytoplankton communities in Utah Lake are marked by decreases in di- versity of the flora as the system becomes more stressed and /or uniform. The phyto- plankton of June are a diverse assortment of species representing several algal divisions. Chlorophyta are important and are mainly associated with Cyanophyta and diatoms. The July phytoplankton are still a diverse group, but are predominately Melosira gran- ulata, two species of Anabaena, and Cera- tium hirundinella. The reduction of species Table 2. Species correlation patterns with respect to environmental parameters as analyzed by regression analy- sis. Species with similar responses are grouped. All correlations listed are at the 0.01 significance level. Correlations Species Clusters Positive Negative Ankistrodesmus falcatus Light penetration Dissolved O2 var. mirabilis Free COg Conductivity Ankistrodesmus falcatus Total alkalinity var. stipatus Si02 Dinobryon divergens Merismopedia glauca Holopedium irregulare Trebaria triappendiculata Anabaena flos-aquae Total alkalinity Water temperature Ankyra judayi Calcium hardness Total hardness Epizooites Microcystis incerta Pediastrum duplex Chlamydomonas globosa Carteria stellifera Total hardness Nitrates Scenedesmus quadracauda Magnesium hardness Aphanizomenon flos-aquae Turbidity Ceratium hinmdinella Salinity Phosphates pH Water temperature 40 Great Basin Naturalist Vol. 38, No. 1 diversity to an almost exclusive Aphani- zomenon-Ceratium community by August is probably due to competition, release of alle- lopathic substances by Aphanizomenon (Pal- mer 1962), and reduction of environmental niches due to decreased variability in the envirionment (Fig. 5). Silica depletion to less than 0.5 mg/liter has been implicated as a factor that is often important in determining succession from diatom-dominated communities to blue- greens (Lund 1965). However, this is not the case in Utah Lake, where silica levels are very high (an average of 19.4 mg/liter for the summer and even higher in August). Water temperature has been shown to be very important in influencing succession from diatom-dominated to blue-green- dominated floras (Patrick 1969). Apparently, this is not the case in Utah Lake. Water temperature is relatively constant through- out the study period and is highest in July when Melosira (the dominant diatom) is most abundant. King (1970) has shown that, imder condi- tions of low alkalinity and high pH, algae may be carbon limited. Blue-green algae seem to be most tolerant to these conditions and Chlorophyta seems to be most sensitive. CURTIS CONTINUUM INDEX In Utah Lake, there was a continuing de- crease in available inorganic carbon (Fig. 6). The disappearance of most chlorophytes corresponds with the period of greatest de- crease in carbon availability. In August, when most of the remaining algal species were replaced by Aphanizomenon and Cera- tiiim, carbon stress was most severe. Many samples had a pH of 8.5 or more and car- bonate alkalinity near 20 mg/liter. King's data (1970) indicate that these conditions are marginal for growth of assorted blue- greens used in his cultures. From this infor- mation, carbon limitation is probably an im- portant factor in determining the composi- tion of the summer phytoplankton communities in Utah Lake. It is important to note that August com- munities in the Lake were dominated by Aphanizomenon flos-aquae and Ceratiiim hi- rundinella, which comprised between 89 and 100 percent of the total algal standing crop. These communities were often com- posed of only Aphanizomenon or Ceratium exclusively. We believe this is strong evi- dence that competitive exclusion is an im- portant factor in regulating the late summer communities of Utah Lake. This hypothesis is presently under investigation in our labo- ratory. 22 ■ 20 - 18 - 16 - O ( 1 S'« - I/) UJ J 12 ■ < IT 10 - • 2 O S8 - 3 o < 6 O 4 - 2 - Fig. 5. Trends in species diversity, community varia- tion, and environmental variation in Utah Lake in the summer of 1974, plotted on the Curtis continuum in- dex. CURTIS INDEX Fig. 6. Partial pressure of CO2 in Utah Lake in the summer of 1974, calculated from disassociation con- stants plotted on the Curtis continuum index. March 1978 Whiting et al.: Utah Lake Algae 41 Literature Cited Holland, R. F. 1974. Paleoecological interpretation of the diatom succession in recent sediments of Utah Lake, Utah. Unpubhshed doctoral dis- sertation. University of Utah, Salt Lake City. Clark, W. J. 1956. An evaluation of methods of con- centrating and counting phytoplankton of Bear Lake, Utah-Idaho. Unpublished master's thesis, Utah State University, Logan. Curtis, J. T., and R. P. McIntosh. 1950. The inter- relations of certain analytic and synthetic phy- tosociological characters. Ecology 31:434-455. 1951. An upland forest continuum in the prairie-forest border region of Wisconsin. Ecolo- gy .32:476-496. George, D. C, and R. W. Edwards. 1973. Daphnia distributions with Langmuir circulations. Lim- nology and Oceanography 18:789-800. GiLMARTiN, A. J. 1974. Variation within populations and classification. Taxon 23(4): 523-536. Harding, W. J. 1970. A preliminary report on the al- gal species presently found in Utah Lake. Great Basin Nat. 30(2):99-105. 1971. The algae of Utah Lake. Part 2. Great Basin Nat. 31(3): 125-134. Hem, J. D. 1970. Study and interpretation of the chemical characteristics of natural waters. 2d ed. Geological Survey Water Supply Paper 1473, U.S. Government Printing Office, Wash- ington, D.C. King, D. L. 1970. The role of carbon in eu- trophication. Journal of Water Pollution Control Federation 42(12):2035-2051. Lund, J. W. G. 1965. The ecology of the freshwater phytoplankton. Biological Review 40:231-293. Palmer, CM. 1962. Algae in water supplies. U.S. De- partment of Health, Education, and Welfare, Amer. Pub. Health Service Pub. 657. Palmer, C. M., and T. E. Maloney. 1954. A new counting slide for nannoplankton. American So- ciety of Limnology and Oceanography, Special Publication 21. Patrick, R., B. Crum, and J. Coles. 1969. Temper- ature and manganese as determining factors in the presence of diatom or blue-green floras in streams. Proc. Nat. Acad, of Sci. 64(2):472-478. Rushforth, S. L., L. L. St. Clair, M. C. Whiting, and T. a. Lewlie. In press. The algae of Utah Lake. Great Basin Naturalist Memoirs No. 3. Snow, E. 1932. A preliminary report on the algae of Utah Lake. Proc. Utah Acad. Sci. 9:21-28. Tanner, V. M. 1930. Fresh water biological studies at Utah Lake. Proc. Utah Acad, of Sci. 7:60-61. 1931. Fresh water studies at Utah Lake. Part 2. Utah Acad. Sci. 8:199-203. Taras, M. J., A. E. Greenberg, R. D. Hoak, and M. C. Rand, eds. 1971. Standard methods for the examination of water and waste water. 13th ed. Amer. Pub. Health Service Pub. BASIDIOMYCETES THAT DECAY JUNIPERS IN ARIZONA. IV R. L. Gilbertson- and J. P. Lindsey^ Abstract.— Five additional species of wood rotting basidiomycetes on Arizona junipers are described and illus- trated. All cause a white rot of dead, fallen trees. Leptosporomyces juniperintis is proposed as a new species. Va- raria fibra is reported from the United States for the first time. Twenty-seven species of wood-rotting basidiomycetes were previously reported on junipers in Arizona (Gilbertson and Lindsey 1975). Since then five additional wood- rotting fungi have been found on Arizona junipers, all on dead fallen trees. This paper presents information on these species. Scanning electron microscopy was done with an ETEC Autoscan. Capitalized color names are from Ridgway (1912). Voucher specimens are deposited in the Mycological Herbarium of the University of Arizona (ARIZ). Hymenochaete arida Karst. Hymenochaete arida Karst., in Sacc, Syll. Fung. 9:228. 1891. Basidiocarps resupinate, effused up to 15 cm, adnate or peeling away at the margin; hymenial surface Buckthorn Brown to Su- dan Brown, smooth, finely setulose under 30X lens, not cracking on drying; margin abrupt, fertile; subiculum thin, up to 200 jum thick, with a single layer of setae from the subhymenial and hymenial hyphae; sub- icular hyphae (Fig. lb) loosely arranged, in- terwoven, simple-septate, moderately thick- walled, pale brown, darkening in KOH, with frequent branching, 3-6 jum diam; setae (Fig. Ic-d) abundant, narrowly sub- ulate, becoming thick-walled, dark brown in KOH, 60-100 X 5-8 /xm, projecting up to 60 jum, straight or slightly curved with a loose membranous sheath (Figs. 2, 3) that disappears or becomes inconspicuous in KOH or Melzer's reagent; ba.sidia (Fig. le) clavate, in loose candelabrums from hyaline subhymenial hyphae (Fig. la), 4-sterigmate, 20-22 X 5-6 ju,m, simple-septate and thick- walled at the base; basidiospores (Fig. If) cylindric, 5-7 X 2-2.5 jum, hyaline, smooth, negative in Melzer's reagent. Hymenochaete arida also decays wood of mesquite (Prosopis jidiflora (Sw.) DC), a common associate of junipers at the lower limits of the pinyon-juniper type (Gilbertson et al., 1975) in Arizona. It is as.sociated with a white rot. When viewed with a scanning electron microscope, the dark, thick-walled setae are seen to have a loose membranous sheath (Figs. 2, 3). Thin-walled, pale brown setae in early stages of development do not have a sheath. Setae are initiated as subulate, thin-walled hyphal ends origina- ting in loose candelabrums with basidia. Secondary wall material is laid down until a narrow lumen remains. During this process the primary wall apparently separates from the secondary wall and forms a loose, mem- branous sheath that hangs loosely on the seta with folds and wrinkles. Reeves and Welden (1967) illustrate setae of Hymeno- chaete luteo-badia (Ft.) Hoehn. et Litsch. and H. berkeleyana (Mont.) Cke. with a sheath. However, they interpret the sheath as being composed of thin-walled, hyaline hyphae. Although occasional setae of H. arida have hyphae appressed to them, the majority have a sheath that is not hyphal in origin. Voucher Specimen: RLG 11321, on al- ligator juniper {Juniperus deppeana Steud.), Gardner Canyon, Santa Rita Mts., Coronado Nat. Forest, Santa Cruz County, AZ. 'University of Arizona Agricultural Experiment Station Journal Article 2794, 'Department of Plant Pathology, University of Arizona, Tucson 85721. 42 March 1978 GiLBERTSON, LiNDSEY: JUNIPER DeCAY 43 Fig. 1. Microscopic characters of Hymenochaete arida (RLG 11321). a, thin-walled, hyaline subhymenial hy- phae; b, thick- walled, pigmented hyphae from lower subiculum; c, setae in early stages of development; d, ma- ture setae, some with membranous sheaths; e, basidia; f, basidiospores. 44 Great Basin Naturalist Vol. 38, No. 1 Figs. 2-5. Scanning electron micrographs of hymenial surfaces of Hymenochaete arida (RLG 11321) and Vararia fihra (RLG 11312). Figs. 2 and 3, setae of H. arida showing membranous sheath; Figs. 4 and 5, mammillate ^oeocystidia projecting through dichohyphidia of V. fibra. March 1978 GiLBERTSON, LiNDSEY: JuNIPER DeCAY 45 Leptosporomyces iitniperinus Gilbn. & Linds., sp. nov. Fructificatio annua, effusa, alba vel cremea, laevi; rhizomorphae albae; hyphae fibulatae, 2-3 ju.m diam; cystidia nulla; ba- sidia clavata, 4-sterigmatibus, 8-15 X 4-5 jLim; basidiosporae subglobosae, hyalinae, laeve, tenuitvinicatae, apiculatae, non- amyloideae, 2.5-3 X 2.5-3.5 juni. Holo- typus: R. L. Gilbertson No. 11325, on Jitni- perus deppeana Steud., Gardner Canyon, Santa Rita Mts., Coronado Nat. Forest, San- ta Cruz County, Arizona; in herb. National Fungus Collections, Beltsville, MD (BPI). Basidiocarps becoming widely effused, ad- nate, cracking into small angular blocks on drying; hymenial surface white to cream- colored, smooth, minutely tomentose; mar- gin thinning out; white mycelial strands present in wood under basidiocarp or at margin; associated with a green alga that becomes incorporated in the subiculum and gives the basidiocarp a greenish tint; hyphal system monomitic; subicular hyphae (Fig. 6a) thin-walled, nodose-septate, 2-3 jLim diam, with abundant coarse crystalline material; cystidia or other sterile hymenial structures lacking; basidia (Fig. 6b) in short candelabrums, clavate, 4-sterigmate, 8-15 X 4-5 jum, with a basal clamp; basidio- spores (Fig. 6c) subglobose, thin-walled, 2.5-3 X 2.5-3.5 jum, hyaline, smooth, nega- tive in Melzer's reagent, with a prominent apiculus. Leptosporomyces juniperinus is associated with a white laminated rot with white my- celial strands developing between the layers of wood. Jiilich (1972) in his monograph of the Athelieae describes an unidentified fim- gus {Leptosporomyces spec. 8650, p. 210) that is similar to L. juniperinus. The sub- globose spores of L. juniperinus distinguish it from the other described species in Lep- tosporomyces. The alga associated with this fungus is a species of the genus Chlorococ- cum Fries. Voucher Specimen: RLG 11325, on al- ligator juniper, Gardner Canyon, Santa Rita Mts., Santa Cruz County, AZ. Stromatoscypha fimbriata (Pers. ex Fr.) Donk Stromatoscypha fimbriata (Pers. ex Fr.) Donk, Rein- wardtia 1:219. 1951. Porotheleiim fimbriatum Pers. ex Fr., Syst. Myc. 1:506. 1821. Basidiocarps annual, resupinate, becoming widely effused, readily separated from sub- stratum; pore surface white to cream- colored, the tubes arising after the devel- opment of apical pores in isolated papillae that become crowded and confluent; pores circular to angular, 3-5 per mm in mature specimens, margin concolorous, soft, rhizo- morphic, usually widely sterile; subiculum white, soft-fibrous, up to 1 mm thick; hyphal system dimitic; subicular generative hyphae (Fig. 7a) thin-walled, nodose-septate, 2.5-3.5 jLtm diam; subicular skeletal hyphae (Fig. 7b) thick-walled, aseptate, with rare branching, 2-2.5 jum diam; cystidia or other sterile hymenial elements lacking; basidia (Fig. 7c) clavate to cylindric, 4-sterigmate, 23-30 X 5-6 jum, with a basal clamp; basi- diospores (Fig. 7d) short-cylindric to ellip- soid, hyaline, smooth, negative in Melzer's reagent, 5-5.5 X 2.5-3 /xm. Stromatoscypha fimbriata has been found on many hardwoods and conifers and causes a white rot. Voucher Specimen: RLG 11324, on al- ligator juniper, Gardner Canyon, Santa Rita Mts., Coronado Nat. Forest, Santa Cruz County, AZ. Trechispora farinacea (Pers. ex Fr.) Liberta Trechispora farinacea (Pers. ex Fr.) Liberta, Taxon 15:318. 1966. Hydnum farinaceum Pers. ex Fr., Syst. Myc. 1:419. 1821. Basidiocarps annual, resupinate, thin and fragile; hymenial surface white to pale buff, grandinioid to hydnaceous; margin floccose, often rhizomorphic; hyphal system mon- omitic; subicular hyphae (Fig. 8a) thin- walled, nodose-septate, frequently ampul- late, 2-3.5 jum diam; cystidia and other ster- ile hymenial elements lacking; basidia (Fig. 8b) short-cylindric to clavate, 4-sterigmate, 3.5-5 jLim diam and up to 15 jum long, with a basal clamp; basidiospores (Fig. 8c) ellip- soid to subglobose, echinulate, hyaline to pale yellow, negative in Melzer's reagent, 3-4 X 2-3 jum. Trechispora farinacea is associated with a white rot of conifer and hardwood logs and slash. 46 Great Basin Naturalist Vol. 38, No. 1 Figs 6-8. Microscopic characters of: 6, Leptosporomyces juniperinus (RLG 11325). a, subicular hyphae; b, ba- sidia c basidiospores; 7, Stronuitoscypha fimhriata (RLG 11324). a, generative subicular hyphae; b, skeletal sub- icular hyphae; cf basidia; d, basidiospores; 8, Trechispora farinacea (RLG 11326). a, subicular hyphae; b, basidia; c, basidiospores. March 1978 GiLBERTSON, LiNDSEY: JuNIPER DeCAY 47 Fig. 9. Microscopic characters of Vararia fibra (RLG 11312). a, thin-walled, simple-septate subicular hyphae; b, thick-walled, aseptate fiber hyphae; c, dichohyphidia; d, gloeocystidia; e, basidia; f, basidiospores. 48 Great Basin Naturalist Vol. 38, No. 1 Voucher Specimen: RLG 11326, on al- ligator juniper, Garden Canyon, Santa Rita Mts., Coronado Nat. Forest, Santa Cruz County, AZ. Vararia fihra Welden Vararia fibra Welden, Mycologia 57:507. 1965. Basidiocarps annual, resupinate, effused up to 5 cm, easily separated from sub- stratum; hy menial surface pale buff, minute- ly tomentose, cracking into small angular blocks on drying, smooth or assuming the configuration of imderlying rhizomorphs; margin thinning out, tomentose to floccose, concolorous, with coarse rhizomorphs that originate in the subiculum and form a net- work under the hymenial layer; subicular hyphae of two types, some simple-septate (Fig. 9a), thin-walled, some partially to completely incrusted, with frequent branch- ing, 2-5 jLim diam; fiber hyphae (Fig. 9b) thick-walled, aseptate, with rare branching, 1.5-4 jum diam; dichohyphidia (Fig. 9c) oc- casional to abundant, with profuse di- chotomous branching, strongly dextrinoid in Melzer's reagent, with ultimate branches slender and recurved, less than 1 jum diam; gloeocystidia (Fig. 9d) abundant, fusoid to capitate or mammillate, thin-walled, with large refractive globules, very weakly posi- tive in sulfuric benzaldehyde, 45-75 X 8-20 jum, with a basal septum; basidia (Fig. 9e) narrowly clavate, 4-sterigmate, 20-28 X 3-3.5 jum, with a basal septum; basidio- spores (Fig. 9f) ovoid, hyaline, smooth, negative in Melzer's reagent, with a promi- nent apiculus, 3-4 X 2-3 ju,m. Vararia fibra has previously been report- ed only from Jamaica. It is associated with a white rot of dead juniper and oak in southern Arizona. Vararia fibra is similar to Vararia ochroleuca (Bourd. & Galz.) Donk. As pointed out by Welden (1965) and Boi- din and Lanquetin (1975), V. ochroleuca lacks rhizomorphs and fiber hyphae and has spores that are more subglobose than those of V. fibra. The scanning electron micro- scope shows the typical catahymenium (Figs. 4, 5) with mammillate gloeocystidia and basidia projecting through the loose surface layer of dichohyphidia. Voucher Specimen: RLG 11317, on al- ligator juniper, Gardner Canyon, Santa Rita Mts., Coronado Nat. Forest, Santa Cruz County, AZ. Acknowledgments This research was supported by Mclntire- Stennis Project 2016-4166-23. We thank Jer- rolyn Campbell of the Department of Anat- omy, University of Arizona College of Med- icine, for help with the scanning electron microscope studies. Literature Cited BoiDiN, J. AND P. Lanquetin. 1975. Vararia subgenus Vararia (Basidiomycetes Lachnocladiaceae): Etude speciale des especes d'Afrique inter- tropicale. Bull. Soc. Mycol. France 91(4):457-513. GiLBERTSON, R. L., H. H. BURDSALL, Jr., AND E. R. Canfield. 1976. Fungi that decay mesquite in southern Arizona. Mycotaxon .3(3): 487-551. GiLBERTSON, R. L. AND J. P. LiNDSEY. 1975. Basidiomy- cetes that decay junipers in Arizona. Great Ba- sin Nat. 35(3):288-304. JuLicH, W. 1972. Monographic der Athelieae (Corti- ciaceae, Basidiomycetes). Willdenowia, Beih. 7, 283 p. Reeves, F. Jr. and A. L. Welden. 1967. West Indian species of Hymenochaete. Mycologia 59:1034-1049. Ridgway, R. 1912. Color standards and color no- menclature. Washington, D.C. Published by the author. Welden, A. L. 1965. West Indian Species of Vararia with notes on extralimital species. Mycologia 57:502-520. PINUS LONGAEVA IN THE STANSBURY MOUNTAINS, UTAH P. A. Kay' and C. G. Oviatt' Abstract.— A new record of Pintts longaeva D. K. Bailey in the Stansbury Mountains, north of the known range of the species, is reported. Bristlecone pine {Pinus longaeva D. K. Bailey) occurs in the Great Basin on high ridges and plateaus, which are often ex- posed. In Utah, the northern-most docu- mented occurrences of Pinus longaeva are at about 40 degrees north latitude in the Deep Creek Mountains, Tooele County, and above the Bad Land Cliffs, Duchesne Coun- ty (Fig. 1). Johnson (1970:25) indicated pres- ence of the species in the western Uinta Mountains, Summit and Duchesne counties, but Bailey (1970, Fig. 3) mapped the Uintas only as a possible habitat without docu- mented occurrence. Bailey (1970, Fig. 3) also showed the Stansbury Mountains, Tooele County, as a possible but unproven location. During recent dendroclimatologi- cal field research, the authors discovered a living bristlecone pine in tlie Stansbury Mountains, well north of the known range limit. Deseret Peak, at 3362 m, is the highest point in the Stansbury Mountains. The mountain bears considerable evidence of Quaternary glaciation, in the form of cirques, U-shaped valleys, and moraines. Contour interval 61 meters 0 1 A Locations with suitable habitats for Pinus longaeva but where the trees have not been found I Fig. 1. Location map. Range of Pintis longaeva from Bailey (1970, Fig. 2). Site location from Deseret Peak quadrangle, U.S. Geological Siu"vey 15-minute topographic series. 'Department of Geography, University of Utah, Salt Lake City, Utah 84112. 49 50 Great Basin Naturalist Vol. 38, No. 1 The bristlecone pine is located northeast of the peak, at about 2960 m on the east side of the arete between Dry Lake Fork and Mill Fork (Fig. 1). The tree is at the head of a steep NNE-trending couloir (slope about 47 degrees) cut into the ESE-facing slope, and is about 25-30 m below the high- est point of the ridge. The site is steep and rocky, and is exposed to direct svinlight and wind. Soil, only locally present, is thin, stony, coarse sand. The pine appears to be rooted directly in the light-colored quartzite bedrock (Cambrian Tintic quartzite; Rigby 1958). Other tree species in the vicinity of the bristlecone pine include Pinus flexilis James, Picea engelmannii Parry, Pseudotsuga men- ziesii (Mirb.) Franco, and Abies lasiocarpa (Hook.) Nutt. Ground cover consists of a few scattered clumps of grasses and com- posites. Two standing snags a few meters uphill from the bristlecone pine have form and bark similar to the live pine, but posi- tive identification was not possible. No oth- er bristlecone pines, dead or living, were found in the immediate area. The tree is of erect form, about 7.5 m tall. The single trunk (55 cm diameter) di- vides about 1 m above the ground into three major fastigate limbs, of which two are dead. Branches are pendulous, but the crown is compact. A major root extends along the ground surface for some 3 m up- hill, perhaps indicating down-slope move- ment. The cambium shows evidence of ex- tensive dieback; two strips of live material cover perhaps one-third of the surface of the trunk. Needles, each about 20-25 mm long, occur in fasicles of five. Each needle bears two resin ducts, but only a very few needles have resin exudations. A single open cone was on the tree and was collected. It is about 90 mm long, has a rounded bottom, bears over 100 scales, and has fine reduced bristles. The morphological characteristics are consistent with classification as Pinus longaeva (Bailey 1970). A voucher specimen of a twig and the cone is housed in the Garrett Herbarium, University of Utah. A preliminary examination of a radial core, obtained with an increment borer, in- dicates 1225 annual rings. The occurrence of Pinus longaeva on quartzite, while not unprecedented, is some- what unusual. The species is usually found on limestone or dolomite, perhaps because of competition experienced on other sub- strates (Bailey 1970). In the Stansbury Mountains, the ridge on the east side of Mill Fork is Cambrian Teutonic limestone (Rigby 1958). A search of that ridge might reveal more bristlecone pine individuals. The authors gratefully acknowledge the assistance in the field of W. E. Riebsame and Jeanne Kay. Additional specimens were found by Riebsame in June 1978. Literature Cited Bailey, D. K. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Ann. Miss. Bot. Gard. 57:210-249. Johnson, C. M. 1970. Common native trees of Utah. Special Report 22, Agricultural Experimental Station, Utah State University, Logan. 109 pp. Rigby, J. K. 1958. Geology of the Stansbury Moun- tains, Tooele County, Utah. Guidebook to the geology of Utah # 1.3, Utah Geological Society, Salt Lake City. 175 pp. DIFFERENTIAL PARASITISM OF PEROMYSCUS MANICULATUS AND PEROMYSCUS TRUEI BY CUTERERRA LARVAE Jeffrey B. Llewellyn' Abstract.— During a 21-month field study of sympatric Peromyscus manic iilatus (Deer Mouse) and Peromyscus truei (pinyon mouse), 12/152 (7.9 percent) P. maniculatus and 1/45 (2.2 percent) P. truei were parasitized by Cut- erebra larvae. Both parasitized P. maniculatus and P. truei and nonparasitized P. maniculatus selected different habitats in the community. Differential parasitism of P. maniculatus and P. truei was possibly caused by host spe- cificity of Cuterebra, whereas differential parasitism of P. maniculatus may have been due to habitat selection. Cuterebrid flies are widely distributed in North and South America where their lar- vae are found embedded in the sub- cutaneous tissues of many rodents (chip- munks, deer mice, wood rats, etc.), rabbits, and sometimes cats, dogs, deer, cattle, hu- mans, and other mammals (Sillman and Smith 1959, Catts 1965). They have also been found in birds (Artmann 1975). Fre- quently, certain species of Peromyscus (deer mice) harbor Cuterebra (botfly) larvae while other species of Peromyscus apparently do not. Why one species is parasitized, but a second closely related species is not para- satized, is not fully understood. During a 21-month field study of sympa- tric Peromyscus maniculatus (deer mouse) and Peromyscus truei (pinyon mouse), differ- ential parasitism of the two species by Cut- erebra larvae was found. Whitaker (1968) suggested that differential parasitism of Per- omyscus may be caused by a species not being in the right place under the proper conditions to be parasitized, or Cuterebra will not parasitize certain species. Previous field investigators have found a correlation between the habitat and parasitism of a spe- cies, and laboratory experiments have shown a strong degree of host specificity by Cuterebra. I suggest that these two factors caused certain P. maniculatus, but not P. truei, to be parasitized. Materials and Methods Sympatric populations of P. maniculatus and P. truei were monitored from April 1975 through December 1976 by live trap- ping in a permanent 1.4 hectare study plot at an elevation of 2025 meters. The study plot was located in the Geiger Grade por- tion of the Virginia Range, which is situated about 18 km SE of Reno, and 10 km NW of Virginia City, Nevada. The Geiger Grade area consists of a pinyon-juniper woodland dominated by single-leaf pinyon {Pinus monophylla) and Utah juniper {Juniperus osteosperma). Be- tween the trees and in the larger open areas a total of 15-20 species of shrubs are found. The most common of these include low sagebrush (Artemisia arbuscula), snakeweed (Gutierrezia sarothrae). Mormon tea (Eph- edra viridis), bitterbrush (Purshia tridentata), and big sagebrush (Artemisia tridentata). Numerous boulder piles and talus rock slides, in which P. truei are abundant, occur throughout the area (for a complete de- scription of the area see Llewellyn 1977). From 25 April 1975 to 21 October 1976, the study plot was sampled for three succes- sive nights on a biweekly basis, using 140 Sherman live traps situated at 10 m inter- vals. Four additional nights of trapping were completed during a rainy period in August 1975; only one three-night sequence was completed in May 1976; and the last sequence in October 1976 consisted of four trap periods. In November and December of 1976, a five-night sequence was com- pleted during the middle of each month. During the 21 -month period, there was a 'Department of Biology, University of Nevada, Reno, Nevada 89557. 51 52 Great Basin Naturalist Vol. 38, No. 1 total of 129 trap periods and 18,060 trap nights. The traps were baited with dry, rolled oats, provided with cotton for insulation during the colder months, and checked for captures in the early morning. All individ- uals captured were sexed, weighed, checked for reproductive condition, marked by toe- clipping, and released at the point of cap- ture. Traps in which a capture was made were thoroughly washed and cleaned before being used again, and all traps were checked for proper working condition as they were being placed at the trap stations. Because of the nature of the long-term study (Llewellyn 1977), no adult parasites were collected. Consequently, the species of Cuterebra was not identified. Results Totals of 152 P. manictilattis (73 females and 79 males) and 45 P. truei (17 females and 28 males) were captured during the 21- month period, and of these, 12 P. manicu- latus (7.9 percent) and one P. truei (2.2 per- cent) were parasitized (Table 1). The in- Table L Incidence of parasitism by Cuterebra larvae in Peromyscus maniculotus and Peromyscus truei from April 1975 through December 1976." Months Number of P. maniculatus captured Number of P. truei captured April 1975 25(0) 0(0) May 42(1) 0(0) June 24 (0) 6(0) July 27(0) 6(0) August 19(0) 7(0) September 12(0) 6(0) October 15(1) 7(0) November 8(1) 7(0) December 5(1) 8(0) January 1976 6(1) 5(0) February 13(2) 6(0) March 9(0) 4(0) April 5(1) 4(0) May 4(2) 5(1) June 5(1) 5(1) July 6(0) 14(0) August 3(0) 10(0) September 9(0) 12(0) October 5(1) 4(0) November 3(0) 3(0) December 7(1) 1(0) 'Numbers in parentheses represent parasitized individuals. fected P. truei in May and June of 1976 represents the same individual (a male), and in both instances one larva was located on the lower left side of the back. The infected P. maniculatus in October and November of 1975 also represents the same individual. Of the 12 parasitized P. maniculatus, 8 were females and 4 were males, 9 individuals contained one larva, 2 contained 2 larvae, and one individual contained 3 larvae. Only one P. maniculatus was parasitized on more than one occasion. Most workers have found the principal site of infestation in the inguinal or scrotal region (Whitaker 1968), although Seaman and Nash (1976) reported that 79.5 percent of the infestations in 87 P. maniculatus oc- curred in the rump region. In this study, all of the larvae were located on the back or on the sides. Most workers have also found that the most important period of infection was during the fall (Whitaker 1968), al- though Brown (1965) found a spring and a fall period in P. boylii (brush mouse), and Wecker (1962) and Hunter et al. (1972) found the highest percentages from July through October in P. leucopus (white- footed mouse) and P. maniculatus. In this study, parasitized individuals were generally found throughout the year, except from July through September of both years (Table 1). Discussion Blair (1941, 1942) found that rodents in- habiting forest environments were para- sitized by Cuterebra, while rodents in- habiting grasslands or more open areas were not parasitized, and he suggested there pos- sibly was a correlation between the habitat and parasitism of a species. In northern Michigan, Blair (1941, 1942) found that P. maniculatus gracilis, Tamias striatus (east- ern chipmunk), and Napaeozapus insignis (woodland jumping mouse), all woodland species, were infested, but in southern Michigan P. maniculatus bairdi, a field form, was not infested, while P. leucopus, a woodland species, was heavily infested. Goertz (1966) also reports differential para- sitism by Cuterebra in conjunction with dif- ferential habitat selection. Goertz (1966) March 1978 Llewellyn: Cuterebra Parasitism 53 found that P. leucopus and Neotoma flori- dana (eastern woodrat), both associated with woodlands, had a high incidence of in- festation, but plains and grassland rodents had exceptionally few parasites. Whitaker (1968) also reports differential parasitism of woodland P. leucopus and grassland P. ma- niculatus bairdi. In this study, parasitized P. maniculatus and P. truei and nonparasitized P. manicu- latus selected different habitats in the study plot. The mean distance to the nearest tree for the 140 trap stations was 1.51 m (some of the stations were located beneath trees, while others were up to and beyond 5 m). During the 21-month period P. maniculatus was captured at 122 stations, with a mean tree distance of 1.44 m, and avoided 18 sta- tions with a mean tree distance of 1.99 m, while P. truei was captured at 82 stations with a mean tree distance of 0.86 m, and avoided 58 stations with a mean tree dis- tance of 2.39 m. The mean tree distance for parasitized and nonparasitized P. manicu- latus was 0.66 m and 1.53 m respectively (p<0.10, Mann- Whitney and Student's T- Tests), while the mean tree distance of par- asitized P. maniculatus and P. trtiei was not significantly different. If differential habitat selection is a cause of differential parasitism by Cuterebra, then such factors as moisture, soil temperature, or vegetation may possibly be responsible. Cuterebra apparently lay their eggs on veg- etation and not directly on the host (Baird 1974); after the larvae leave the host they burrow into the soil and pupate (Sillman and Smith 1959); and Layne (1963) found little or no infestation in P. floridanus (Flor- ida mouse) in drier habitats. Differential parasitism could also be caused by host specificity of Cuterebra. In a field study Seaman and Nash (1976) found that 87/633 P. manictdatus (13.7 percent) and 1/170 P. difficilis (0.6 percent) were in- fested with Cuterebra larvae. These findings are noteworthy because P. difficilis (rock mouse) and P. truei are classified in the same Peromyscus group (Hall and Kelson 1959), and the two species generally select the same type of habitats (Baker 1968). In a laboratory study Penner and Pocius (1956) introduced larvae of C. fontinella (tentative identification) into 10 species of mammals, and development occurred in Mus musculus (house mouse), Rattus rattus (Black rat), R. norvegicus (Norway rat), and P. leucopus, but not in kittens, guinea pigs, wild and do- mestic rabbits, Microtus pennsylvanicus (meadow vole), Sigmodon hispidus (hispid cotton rat), and P. maniculatus. In another laboratory experiment Catts (1965) in- troduced larvae of C. approximata into Cri- cetus auratus (hamster), Neotoma fuscipes (dusky-footed woodrat), and the natural host P. maniculatus. Neotoma fuscipes proved to be refractory; 6.2 percent of the separate exposures in the hamsters resulted in posi- tive infection; and 44 percent of the sepa- rate exposures in P. maniculatus resulted in positive infection. In a second experiment Catts (1965) introduced larvae of C. lati- frons into nine species of rodents and one rabbit, and infection occurred in Cr. aur- atus, R. rattus, M. musculus, and the natu- ral host N. fuscipes. However, only in N. fuscipes was the net production of fully de- veloped larvae high (67 percent), as com- pared with 25 percent in Cr. auratus, 7 per- cent in R. rattus, and 0 percent in M. musculus, and Catts (1965) concluded that susceptibility to infection by Cr. auratus, R. rattus, and M. musculus does not necessarily confirm their suitability as a host. Per- omyscus maniculatus was also infected with larvae of C. latrifrons in the same experi- ment, but appeared refractory. I conclude that differential parasitism of P. maniculatus and P. truei was possibly caused by host specificity of Cuterebra, whereas differential parasitism of P. ma- niculatus may have been due to habitat se- lection. Apparently, P. truei and parasitized P. maniculatus were in the right place un- der the proper conditions to be infected, but only P. maniculatus was infected be- cause of structural, physiological, or behav- ioral differences. Nonparasitized P. manicu- latus evidently were not in the proper place. This hypothesis is offered only as a tenta- tive one, since more experimental evidence is needed before definite conclusions can be made. 54 Great Basin Naturalist Vol. 38, No. 1 Acknowledgments I would like to thank Dr. Stephen H. Jenkins and Dr. Robert W. Mead, Univer- sity of Nevada, Reno, for reading an earlier draft of the manuscript, and Mr. Paul La- zaris, President, Lake Tahoe Recreational Land Co. Inc., for pemission to use the area on which the permanent study plot was lo- cated. Financial support was received from the Department of Biology, University of Nevada, Reno, and from a Merchant Schol- arship awarded by the University of North- ern Iowa, Cedar Falls. Literature Cited Artmann, J. W. 1975. Cuterehra parasitism of an American woodcock. J. Parasitol. 61: 65. Baird, C. R. 1974. Field behavior and seasonal activity of the rodent botfly, Cuterebra tenebrosa, in central Washington (Diptera: Cuterebridae). Great Basin Nat. 34: 247-253. Baker, R. H. 1968. Habitats and distribution, pp. 98-126. In: J. A. King, ed., Biology of Per- omysciis (Rodentia). Spec. Publ. No. 2, Amer. See. Mammal. 593 p. Blair, W. F. 1941. Some data on the home ranges and general life history of the short-tailed shrew, red-backed vole, and woodland jumping mouse in northern Michigan. Amer. Midland Nat. 25: 681-685. 1942. Size of home range and notes on the life history of the woodland deer-mouse and eastern chipmunk in northern Michigan. J. Mammal. 23: 27-36. Brown, L. N. 1965. Botfly parasitism in the brush mouse and white-footed mouse in the Ozarks. J. Parasitol. 51: 302-304. Catts, E. p. 1965. Host-parasite interrelationships in rodent botfly infections. Trans. N. Amer. Wildl. Nat. Resour.' Conf. 30: 184-196. GoERTZ, J. W. 1966. Incidence of warbles in some Oklahoma rodents. Amer. Midland Nat. 75: 242-245. Hall, E. R. and K. R. Kelson. 1959. The mammals of North America. Ronald Press, New York. 2 vol., 1162 pp. Hunter, D. M., R. M. F. S. Sadleir, and J. M. Webster. 1972. Studies of the ecology of cut- erebrid parasitism in deer mice. Can. J. Zool. 50: 25-29. Layne, J. N. 1963. A study of the parasites of the Florida mouse, Peromyscus floridanus, in rela- tion to host and environmental factors. Tulane Studies Zool. 11: 1-27. Llewellyn, J. B. 1977. Competition and coexistence in sympatric populations of Peromyscus manicii- latus and Peromyscus truei. Unpublished Ph.D. Dissertation, Univ. Nevada, Reno. 118 pp. Penner, L. R. and F. p. Pocius. 1956. Nostril entry as the mode of infection by the first stage larvae of a rodent Cuterebra. J. Parasitol. 42: 42. Seaman, R. N. and D. J. Nash. 1976. Variation in the incidence of botfly larvae (Cuterebra) in two sympatric species of Peromyscus in northern Colorado. Great Basin Nat. 36: 481-482. Sillman, E. J. AND M. V. Smith. 1959. Experimental infestation of Peromyscus leucopus with larvae of Cuterebra angustifrons. Science 130: 165-166. Wecker, S. C. 1962. The effects of botfly parasitism on a local population of the white-footed mouse. Ecology 43: 561-565. Whitaker, J. O. Jr. 1968. Parasites, pp. 254-311. In: J. A. King, ed.. Biology of Peromyscus (Rodentia). Spec. Publ. No. 2, Amer. Soc. Mammal. 593 p. AN ANNOTATED LIST OF THE AQUATIC INSECTS OF SOUTHEASTERN IDAHO, PART III. EPHEMEROPTERA Robert L. Newell' and G. Wayne Minshall' Abstract.— The study revealed representatives of 9 families, 22 genera, and 59 species occurring within the study area. No new species and no new state records were collected, but a number of new localities have been added. This paper is the third in a series of pub- hcations designed to describe the aquatic in- sect fauna of southeastern Idaho. A map de- scribing the study area is presented in Newell and Minshall (1977). The Ephemeroptera, or mayflies, are per- haps the best known and most extensively studied group of aquatic insects. Mayflies are often a diverse and abundant com- ponent of the benthic favma of streams in the western United States. In North Ameri- ca north of Mexico there are 60 genera and 622 species of mayflies (Edmunds et al. 1976). Information on the geographical distribu- tion of mayflies is widely scattered through- out published sources. Unfortunately, much of these data are useless for this report since many authors listed species only as "collected in Idaho" and did not supply definite locations. The mayfly fauna of Idaho was poorly known until Jensen's (1966) study "Mayflies of Idaho." Jensen's thorough work reviews the literature and provides keys to nymphs and adults of the 85 species of mayflies known from Idaho. However, the work remains unpublished and therefore is generally inaccessible to a wide number of workers. Hornung and Barr (1970) also reported some mayfly distribu- tion data. A total of 9 families, 22 genera, and 59 species are reported from southeastern Idaho. No new species and no new state re- cords were collected, but a number of new localities were added. An undescribed spe- cies of Stenonema (Edmunds et al. 1976) also is known from southeastern Idaho. The genus Pseudocloeon is reported here and not in the checklist, since the species are not identified. The authors thank the many people who contributed time, effort, and specimens for this study. They also thank Dr. W. L. Pe- ters for identifying some of the specimens. Family Siphlonuridae 1. Amelehis cooki McDunnough.— Ban- nock Co.: Mink Cr. 2. Ameletus oregonensis Eaton.— Ban- nock Co.: Mink Cr. Bonneville Co.: Snake R. (S. Fk.). Butte Co.: Little Lost R. Teton Co.: Teton R. 3. Ameletus sparsatus McDunnough.— Bannock Co.: Rapid Cr. Bonneville Co.: Pine Cr. Custer Co.: Big Lost R. Fremont Co.: Snake R. (Henry's Fk.). 4. Ameletus velox Dodds.— Butte Co.: Little Cottonwood Cr.; Little Lost R. 5. Siphlomirus occidentalis Eaton.— Bear Lake Co.: St. Charles Cr. Blaine Co.: Big Wood R. Bonneville Co.: Rainey Cr. Butte Co.: Big Lost R. Cassia Co.: Cassia Cr. Cus- ter Co.: near Mackay. Fremont Co.: Snake R. (Henry's Fk.). Teton Co.: Teton R. Family Baetidae 6. Baetis bicaudatus Dodds.— Blaine Co.: Big Wood R. Bonneville Co.: Pine Cr.; Rainey Cr. Butte Co.: Little Lost R.; stream near Arco. Custer Co.: Big Lost R. Fremont Co.: Targhee Cr. Teton Co.: Badger Cr. 'Steams-Roger, Inc., P.O. Box 5888, Denver, Colorado 80217. 'Department of Biology, Idaho State University, Pocatello, Idaho 83209. 55 56 Great Basin Naturalist Vol. 38, No. 1 7. Baetis insignificans McDunnoiigh.— Butte Co.: Little Lost R. Gooding Co.: near Tuttle. 8. Baetis intermedius Dodds.— Bannock Co.: Indian Cr.; Mink Cr.; Portneuf R. Bingham Co.: Spring Cr. Gooding Co.: Niagara Springs; Riley Cr. 9. Baetis parvus Dodds.— Bingham Co. Blaine Co.: Big Wood R. Bonneville Co.: Brockman Cr.; Snake R. (S. Fk.). Jefferson Co.: near Roberts. Teton Co.: Teton R. 10. Baetis propinquus (Walsh).— Butte Co.: near Arco. IL Baetis quilleri Dodds.— Gooding Co. 12. Baetis tricaudatus Dodds.— Bannock Co.: Portneuf R.; Rapid Cr. Blaine Co.: Big Wood R. Bonneville Co.: Rainey Cr. Butte Co.: Little Lost R.; near Arco. Clark Co.: Beaver Cr.; Birch Cr.; Camas Cr.; Medicine Lodge Cr. Custer Co.: Big Lost R. Fremont Co.: Snake R. (Henry's Fk.). Franklin Co.: Mink Cr. Gooding Co.: Niagara Springs; Snake R. Jefferson Co.: Birch Cr. Jerome Co. Lincoln Co.: N. Gooding Canal. Oneida Co.: Deep Cr. Twin Falls Co.: Salmon Falls Cr. 13. Callihaetis coloradensis Banks.— Bear Lake Co.: near Montpelier. Blaine Co.: Sawtooth Nat. Forest. Bonneville Co.: Rain- ey Cr. Fremont Co.: near St. Anthony. Franklin Co.: Cub R. Jefferson Co.: near Heise. Lincoln Co.: near Dietrich. 14. Callihaetis montaniis Eaton.— Ban- nock Co.: near McCammon. Franklin Co.: Bear R. Gooding Co.: Riley Cr. 15. Callihaetis nigritus Banks.— Bannock Co.: Mink Cr.; near Arimo; Portneuf R. Bear Lake Co.: near Ovid. Bingham Co.: Clear Cr. Butte Co.: near Howe. Caribou Co.: Portneuf R. Cassia Co.: near Burley. Gooding Co.: Niagara Springs; Riley Cr. Je- rome Co.: near Jerome. Lincoln Co.: near Shoshone. Oneida Co.: Deep Cr. Twin Falls Co.: near Twin Falls. 16. Centroptiliun hifurcatiim McDun- nough.— Gooding Co.: Twin Falls Co. 17. Centroptilurn selanderorum Ed- munds.— Bingham Co.: Aberdeen-Springfield Canal; Snake R. Blaine Co.: Big Wood R. Butte Co.: Big Lost R. Cassia Co.: at Bur- ley. Custer Co.: Big Lost R. Lincoln Co.: N. Gooding Canal. Oneida Co.: Deep Cr. Family Heptageniidae 18. Cinijgma integrum Eaton.— Blaine Co.: ditch near Ketchum. Butte Co.: Little Cottonwood Cr. 19. Cinygmida mimiis (Eaton).— Bannock Co.: Gibson Jack Cr.; Mink Cr. Bonneville Co.: Jensen Cr. 20. Cinygmida par (Eaton).— Butte Co.: Little Lost R. Teton Co.: Badger Cr. 21. Cinygmida ramaleyi (Dodds).— Ban- nock Co.: Mink Cr. Cassia Co.: Cassia Cr. 22. Cinygmiila reticulata McDunnough.— Clark Co.: Birch Cr. 23. Cinygmida tarda (McDunnough).— Custer Co.: near Mackay. 24. Epeorus (Iron) alhertae (McDun- nough).— Bannock Co.: Mink Cr. Blaine Co.: near Ketchum. Custer Co.: Big Lost R. Fremont Co.: Snake R. (Henry's Fk.). 25. Epeorus (Iron) deceptivus (McDun- nough): Blaine Co. Butte Co.: Little Lost R. 26. Epeorus (Iron) longimanus (Eaton).— Bannock Co.: Mink Cr.; Portneuf R.; Rapid Cr. Bear Lake Co.: St. Charles Cr. Bonne- ville Co.: Pine Cr. Fremont Co.: Snake R. (Henry's Fk.). Franklin Co.: Cub R. Teton Co.: Badger Cr. Twin Falls Co.: Rock Cr. 27. Epeorus (Ironopsis) grandis (McDun- nough).— Bonneville Co.: Pine Cr. Teton Co.: Moose Cr. 28. Heptigenia criddlei McDunnough.— Bingham Co.: Clear Cr. Blaine Co.: Rich- field Canal. Butte Co.: Hagerman. Lincoln Co.: Gooding Canal. 29. Heptagenia elegantida (Eaton).— Lin- coln Co.: Gooding Canal. Power Co.: Ban- nock Cr. 30. Heptagenia simplicioides McDun- nough.— Bingham Co.: Aberdeen-Springfield Canal. Caribou Co.: Toponce Cr. Lincoln Co.: Big Wood R. 31. Heptagenia solitaria McDunnough.— Franklin Co.: Bear R. Fremont Co.: Snake R. (Henry's Fk.). Jefferson Co.: Birch Cr. Teton Co.: Teton R. 32. Rhithrogena futilis McDunnough.— Twin Falls Co.: Rock Cr. 33. Rhithrogena luigeni Eaton.— Bannock Co.: Rapid Cr. Bonneville Co.: Pine Cr. Clark Co.: Birch Cr.; Medicine Lodge Cr. Custer Co.: Big Lost R. Franklin Co.: Cub R. Fremont Co.: Snake R. (Henry's Fk.). Jef- March 1978 Newell, Minshall: Idaho Mayflies 57 ferson Co.: Birch Cr. Teton Co.: Badger Cr. 34. Rhithrogeno morrisoni (Banks).— Ban- nock Co.: Mink Cr. Bear Lake Co.: Thomas Fork Cr. Blaine Co.: Big Wood R. Butte Co.: Big Lost R. Fremont Co.: Snake R. (Henry's Fk.). 35. Rhithrogena robusta Dodds.— Bonne- ville Co.: Pine Cr. Butte Co.: Little Lost R. Fremont Co.: Teton R. Teton Co.: Moose Cr. 36. Rhithrogena imdulata (Banks).— Cus- ter Co.: near Mackay. Family Leptophlebiidae 37. Choroterpes albianniilata McDun- nough.— Bingham Co.: Snake R. Franklin Co.: Bear R. 38. Leptophlebia gravastella (Eaton).— Bannock Co.: Mink Cr. Bonneville Co.: Brockman Cr. 39. Paraleptophlebia debilis (Walker).— Blaine Co.: Warm Springs Cr. Butte Co.: Big Lost R. Caribou Co.: Blackfoot R.; Portneuf R. Jerome Co.: near Jerome. Lin- coln Co.: N. Gooding Canal. 40. Paraleptophlebia heteronea (McDun- nough).— Bannock Co.: Mink Cr.; Portneuf R. Bear Lake Co.: Montpelier Cr. Blaine Co.: ditch near Big Wood R. Butte Co.: Little Lost R. Clark Co.: Beaver Cr.; Camas Cr. Custer Co.: Big Lost R. Franklin Co.: Mink Cr. Fremont Co.: Buffalo R. Teton Co.: Teton R. 4L Paraleptophlebia memorialis (Eaton).— Butte Co.: near Howe. Custer Co.: Big Lost R. Fremont Co.: Snake R. (Henry's Fk.). Family Ephemerellidae 42. Ephemerella (Atenella) margarita Needham.— Lincoln Co.: Big Wood R. 43. Ephemerella (Caudatella) hetero- caudata heterocaudata McDunnough.— Blaine Co.: Trail Cr. Custer Co.: Big Lost R. Fremont Co.: Targhee Cr. 44. Ephemerella (Caudatella) hystrix Traver.— Butte Co.: Little Lost R. Fremont Co.: Teton R. (N. Fk.); Targhee Cr. 45. Ephemerella (Dnmella) coloradensis Dodds.— Bannock Co.: Mink Cr. Blaine Co.: Big Wood R. Bonneville Co.: Pine Cr. Butte Co.: Little Lost R. Clark Co.: near Orgona. Franklin Co.: Mink Cr. Fremont Co.: How- ard Cr. Teton Co.: Moose Cr. 46. Ephemerella (Drunella) doddsi Need- ham.— Bannock Co.: City Cr.; Gibson Jack Cr.; Mink Cr.; Portneuf R.; Rapid Cr. Blaine Co.: Big Wood R. Bonneville Co.: Indian Cr.; Jensen Cr.; Pine Cr.; Rainey Cr. Butte Co.: Little Lost R. Franklin Co.: Cub R. Fremont Co.: Targhee Cr. Teton Co.: Moose Cr.; Trail Cr.; Warm Cr. 48. Ephemerella (Drunella) flavilinea McDimnough.— Blaine Co.: Big Wood R. Bonneville Co.: Indian Cr.; Rainey Cr. Butte Co.: Little Lost R. Custer Co.: Big Lost R. Fremont Co.: Warm R. Teton Co.: Teton R. 48. Ephemerella (Drunella) grandis gran- dis Eaton.— Bannock Co.: City Cr.; Gibson Jack Cr.; Mink Cr.; Pocatello Cr.; Portneuf R.; Rapid Cr. Bear Lake Co.: St. Charles Cr. Bonneville Co.: Rainey Cr. Butte Co.: Little Lost R. Clark Co.: Beaver Cr.; Birch Cr.; Camas Cr.; Medicine Lodge Cr. Custer Co.: Big Lost R. Franklin Co.: Cub R. Fre- mont Co.: Snake R. (Henry's Fk.). Teton Co.: Teton R. (N. Fk.). 49. Epheinerella (Drunella) spinifera Need- ham.- Blaine Co.: Big Wood R. Butte Co.: Little Lost R. Fremont Co.: Targhee Cr. 50. Ephemerella (Ephemerella) inermis (Eaton).— Bannock Co.: Mink Cr.; Portneuf R.; Rapid Cr. Bear Lake Co.: Salt Cr. Bing- ham Co.: Snake R. Blaine Co.: Big Wood R.; Silver Cr. Bonneville Co.: Pine Cr. Rainey Cr. Butte Co.: Little Lost R. Cassia Co.: Cassia Cr. Clark Co.: Beaver Cr.; Birch Cr.; Camas Cr.; Medicine Lodge Cr. Custer Co.: Big Lost R. Franklin Co.: Cub Cr.; Mink Cr. Fremont Co.: Warm R. Jefferson Co.: Birch Cr. Lincoln Co.: Little Wood R. Teton Co.: Teton R.; Trail Cr. 5L Ephemerella (Ephemerella) infrequens McDunnough.— Bear Lake Co.: Thomas Fork Cr. Bingham Co.: Clear Cr.; Spring Cr. Blaine Co.: Big Wood R. Bonneville Co.: Rainey Cr.; Snake R. (S. Fk.). Clark Co.: Birch Cr. Custer Co.: Big Lost R. Fre- mont Co.: Warm R. 52. Ephemerella (Serratella) tibialis McDunnough.— Bonneville Co.: Pine Cr. Butte Co.: Little Lost R.; Big Lost R. Cus- ter Co.: near Mackav. Franklin Co.: Cub R. 58 Great Basin Naturalist Vol. 38, No. 1 53. EphemereUa (Timpanoga) heciiba he- cuba (Eaton).— Bingham Co.: Snake R. Cus- ter Co.: Big Lost R. Franklin Co.: Cub R. Family Tricorythidae 54. Triconjthodes minuhis Traver.— Ban- nock Co.: near Pocatello; Portneuf R. Bing- ham Co.: Clear Cr.; Spring Cr. Bonneville Co.: Willow Cr. Butte Co.: Big Lost R.; Little Lost R. Cassia Co.: Snake R. Caribou Co.: Blackfoot R.; Portneuf R. Franklin Co.: Bear R. Gooding Co.: Niagara Springs. Je- rome Co.: at Hazelton. Lincoln Co.: N. Gooding Canal. Minidoka Co.: Snake R. Oneida Co.: Deep Cr. Power Co.: Snake R. Twin Falls Co.: Salmon Falls Cr. mont Co.: Buffalo R.; Snake R. (Henry's Fk.). 58. Hexagenia limbata limbata Serville.— Cassia Co.: near Burley; Snake R. Gooding Co.: at Hagerman; near Gooding. Lincoln Co.: at Shoshone. Minidoka Co.: at Rupert. Power Co.: Snake R. Family Polymitarcidae 59. Ephoron album (Say).— Bannock Co.: Portneuf R. Cassia Co.: Raft R. Franklin Co.: Bear R. Jerome Co.: near Jerome. Lin- coln Co.: near Shoshone. Twin Falls Co.: at Twin Falls; near Murtaugh. Family Caenidae 55. Brachycerctis prudens (McDun- nough).— Lincoln Co.: N. Gooding Canal. 56. Caenis simulans McDunnough.— Ban- nock Co.: Portneuf R. Butte Co.: Craters of the Moon Nat. Monument. Caribou Co.: Portneuf R. Jerome Co.: near Milner. Lin- coln Co.: near Shoshone; N. Gooding Canal. Oneida Co.: Deep Cr. Family Ephemeridae 57. Ephemera simulans Walker.— Fre- Literature Cited Edmunds, G. F., Jr., S. L. Jensen, and L. Berner. 1976. The mayflies of Nortfi and Central Ameri- ca. Univ. of Minn. Press, Minneapolis. 330 p. HoRNUNC, D. S. AND W. F. Barr. 1970. Insects of Cra- ters of the Moon National Monument Idaho. Univ. of Idaho, Coll. of Agric, Misc. Ser. No. 8, 118 p. Jensen, S. L. 1966. The mayflies of Idaho (Ephem- eroptera). Unpublished master's thesis, Univ. of Utah, Salt Lake City. 365 p. Newell, R. L. and G. W. Minshall. 1977. An anno- tated list of the aquatic insects of southeastern Idaho. Part II. Trichoptera. Great Basin Nat. 37:253-257. OBSERVATIONS ON THE PREY AND NESTS OF SEVEN SPECIES OF CERCERIS (HYMENOPTERA: SPHECIDAE)' Howard E. Evans^ and William L. Rubink^ Abstract.— Cerceris species are remarkably constant in their prey preferences. Records are presented from di- verse localities that confirm prey constancy in five species. An apparent exception is noted in the case of C. crotonella Viereck & Cockerell. A mixed aggregation of C. fiimipennis Say and C. californica Cresson is reported, both preying upon Buprestidae and using some of the same species. Nest and prey of C. conifrons Mickel are re- ported for the first time. One of the striking features of the behav- ior of species of Cerceris is their prey con- stancy. Even in widely disparate parts of the range, females of a given species tend to utilize beetles of only one family, often of only a few related genera. Such special- ization doubtless reduces competition among species nesting together. Evans (1971) reported three species nesting togeth- er at Bedford, Massachusetts, each preying upon quite a different kind of weevil; later (1974) he added still a fourth species from this same site, utilizing still a different kind of weevil. Despite their prey specificity, the species of Cerceris are remarkably similar in many aspects of their nesting behavior. In this paper we report still further ex- amples of prey constancy in diverse parts of the ranges of five species. We also report an apparent exception in the case of one species, C. crotonella Viereck & Cockerell. Further, we report a mixed aggregation of two closely related species, C. fiimipennis Say and C. californica Cresson, both prey- ing upon Buprestidae, including several of the same species. The first records of the nest and prey of C. conifrons Mickel are presented, as well as diverse notes on the nests of other species. Cerceris simplex macrosticta Viereck & Cockerell On 20 July 1976, we took a female of this large species flying with her prey over an arroyo 30 km NE of Lordsburg, New Mexico. The prey proved to be a tenebrio- nid beetle, Pechalius siibvittatus Casey. On 24 and 31 August 1976, we studied an ag- gregation of some 40 females of this species about 9 km N of Roggen, Weld Co., Colo- rado. At this site a single species of Te- nebrionidae was being employed as prey, Bothrotes phimbeus plumbeus (LeConte) (N = 38, with several additional sight records and several additional records from the same area in 1977). Since Alcock (1974, 1975) has published two excellent papers on C. simplex macrosticta, we present only a few notes here. The prey at Alcock's site in Arizona also consisted of a single species of Tenebrionidae, in this case Metapoloba pruinosa (Horn) (N = 60, with additional sight records). It is interesting that these three genera, Metapoloba, Bothrotes, and Pechalius, all belong to the tribe Epitragini. Lin (1967) studied C. simplex graphica Smith in Oklahoma and found the prey there to consist of a single tenebrionid spe- cies, Eleodes opaca (Say), a member of a different subfamily. Since the ranges of these two subspecies of simplex appear to overlap broadly, thought might be given to the possibility of their being separate spe- cies. At the Roggen site, nests were located in a steep slope of fine-grained sand, mostly devoid of vegetation, where the chief asso- ciates were species of Bembix, Microbembex, and at least two species of bees. Cerceris 'Research supported by the National Science Foundation, grant BNS76-09319. 'Department of Zoology and Entomology, Colorado State University, Fort Collins, Colorado 80523. 59 60 Great Basin Naturalist Vol. 38, No. 1 nests were situated over a distance of some 30-40 m along the bank, but they were def- initely clumped in certain places, along with those of the bees. In these clumps nest entrances were often no more than 5-15 cm apart, but the clumps themselves were sepa- rated by several meters. Nest entrances were left open while the females were pro- visioning, and each was surrounded by a rim of sand, as is usual in this genus. A male macrosticta was taken on flowers of Croton texensis (Klotzsch) not far away, but no males were seen in the nesting area. During the morning hours, females were frequently seen descending to their nests holding beetles in their mandibles in such a way that they hung down obliquely from the body. Tliey were frequently followed by 1-3 satellite flies, Senotainio sp., and when so followed the wasps would often circle widely before entering their nests, making loops several meters in diameter at different heights and occasionally landing on vegeta- tion. One female spent six minutes making an evasive flight of this nature, then finally entered the nest without any flies following her. Only 1 of the 12 cells eventually exca- vated appeared to have had the contents destroyed by maggots, so we judge that the wasps are often successful in evading satel- lite flies. Nest structure agreed closely with that figured by Alcock (1974) for this species. One of the two nests we excavated had five cells, the other seven. Cells varied in depth from 17 to 32 cm (x = 26, precisely the same figure Alcock obtained in Arizona). In both cases beetles had been stored at vari- ous points in the burrow, chiefly near its terminus. The number of beetles per cell was two in one case, three in nine cases, and seven in one case. Most cells measured 10 x 12 mm, but the one containing seven beetles measured 14 x 23 mm. We suspect this was a "female producing cell", the oth- ers "male" cells, a situation that is probably common if not universal in Cerceris. These figures are also very similar to those ob- tained by Alcock in Arizona. Cerceris hicornuta Guerin This is a relatively well-studied species. and we present only a few notes confirming prey constancy in diverse parts of the range. On 23 July 1975 we took a female C. hicornuta hicornuta 14 km W of La- porte, Colorado, carrying a weevil, Spheno- phorus cicatristriatiis Fahraeus in her man- dibles. She landed on a dirt road, apparently en route to her nest, and rested there for a moment with her wings extend- ed obliquely above her abdomen. During the month of July 1976 we ob- served an aggregation of C. hicornuta fidelis Viereck & Cockerell at the Lajoya Wildlife Preserve, 30 km N of Socorro, New Mexico. On 1 July we counted 44 nests in an area 3 X 5 m in flat, hard-packed sandy loam be- side a dirt road. About an equal number of Tachijtes aurulentus (F.) also nested here, and the nests of the two species were some- what intermingled and rather similar in ex- ternal appearance, both having a large mound at the entrance and the hole near the center of the mound. However, the Tachytes were most active in the early morning, as early as 0630, while the Cer- ceris provisioned their nests over the warm part of the day. The Cerceris nests were well spaced, mostly 0.3-1.0 m apart, but some were very close to Tachytes nests (in one case only 15 cm from a Tachytes nest). Provisioning females entered the area 1-2 m high and plunged directly into the open nest entrances. Prey was carried in the mandibles as usual in the genus, and con- sisted of a single species of weevil, Spheno- phorus austraUs Chittenden (N = 12). Wee- vils were stored in the burrow, as usual in this genus. We dug out only one nest in 1976, early in the season and apparently be- fore any cells had been made. The nest had a mound at the entrance measuring 15 x 17 cm, 2.5 cm high, with the burrow pene- trating the center. Burrow diameter was 8 mm. This burrow was vertical for 29 cm, then leveled off abruptly and extended an- other 27 cm before terminating blindly. Three weevils were found about midway in the horizontal part of the burrow. In July 1977, we made further observa- tions on this same aggregation, though many fewer nests were visible. Three nests were excavated. The first, on 8 July, yielded five weevil prey from the single, recently March 1978 Evans, Rubink: Gerceris Wasp Nests 61 completed cell. Two species, Sphenophortis neomexicanus Chittenden (N = 4) and S. ci- catristriatiis Fahraeus (N = 1) were repre- sented. A second nest, excavated on 12 July, contained no cells and a single weevil, S. ci- catristriatus, midway down the 25 cm long burrow. The third nest, examined on 27 July, yielded more substantial information on nest structure. However, because of sev- eral days of inactivity due to heavy rain, none of the cell contents were suitable for preservation. Seven cells contained from three to seven weevils each, as determined from the prey remains. All cells measured approximately 15 x 30 mm. Two cells, at depths of 11 and 18 cm, contained only in- tact, decomposing weevil remains. Two oth- ers, 13 and 16 cm deep, contained cocoons and remains of weevils. A cell at a depth of 29 cm contained a dipteran larva that died shortly after removal. The two remaining cells, at depths of 24 and 31 cm, contained Cerceris larvae 12 and 21 cm long, respec- tively. Cells were widely spaced, 7-15 cm apart. Krombein (1960) studied C. bicornuta fi- delis in Arizona, where the wasps were also making deep nests along the edge of a road. The prey here consisted of the weevil Eu- pagoderes sp., which he found to be com- mon on the flowers of snakeweed. There are several papers dealing with the biology of C. bicornuta bicornuta in diverse parts of its range, e.g. Rau (1928), Cartwright (1929), Strandtmann (1945), and Krombein (1953). Scullen and Wold (1969) summarize prey records from five different states (New York, Ohio, Missouri, North and South Carolina), in every case a single genus of weevil, Sphenophortis. Evans (1971) added additional records for this genus of weevil from Connecticut, based on unpublished re- cords of Richard Dow. Cerceris crotonella Viereck & Cockerell This small species is a common inhabitant of extensive areas of fine-grained sand in Colorado and New Mexico. We have seen prey-laden females carrying small beetles into open holes in sand on several occasions, but we have had no success in finding cells. On 5 July 1975 we captured a female with prey 4 km S of Caddoa, Bent Co., Colo- rado. The beetle proved to be Pachijbrachys minor Bowditch (Chrysomelidae). This is in contrast to a record from western Texas of this species preying upon a beetle of the imrelated family Nitidulidae (Evans 1971). Clearly this species will bear further study. Cerceris conifrons Mickel This is a small species bearing much re- semblance to crotonclki and belonging to the same species-group (group I of Scullen 1965). On 11 August 1977 a female was seen plunging into an open, oblique burrow along a path at Great Sand Dunes National Monument, Alamosa Co., Colorado. The nest was in fine-grained but rather firm sand in a small open space among low grasses and forbs. There was no mound of soil at the entrance. The female was cap- tured as she left the nest at 1300 hours, and the nest was excavated. The burrow formed about a 30-degree angle with the surface for 17 cm, then went down vertically for another 5 cm, terminating at a vertical depth of 12 cm. There were five very small weevils at the bottom of the burrow, Epi- mechus sp. (Curculionidae). It is probable that these had merely been stored in the bottom of the burrow and that no cell had yet been prepared. Cerceris echo Mickel This species resembles the preceding two in size and color and is a member of the same species-group. Records from widely separate parts of the range demonstrate that this species is a specialist on beetles of the family Phalacridae. Evans (1971) reported beetles of this family as prey of C. echo at- rata Scullen in Lexington, Massachusetts, and as prey of C. echo echo Mickel in Corn- ish, Utah. On 15 August 1977, at the Lajoya State Wldlife Preserve, 30 km N of Socorro, New Mexico, we captured a fe- male C. echo echo as she was hovering above her nest with prey. The nest was lo- cated in a small bare depression in fine aeo- lian sand among scattered Dalea, Atriplex, and bunch grasses, and below the active sand dunes. No tumulus was apparent 62 Great Basin Naturalist Vol. 38, No. 1 aroiind the nest entrance. The burrow pro- ceeded vertically 40 cm through very dry, loose soil, then angled sharply, continuing downward at a 30 degree angle with the horizontal, terminating at a depth of 44 cm. Three beetles were found at the angle in the burrow. Two small cells were found at depths of 54 and 62 cm, but no eggs or lar- vae were visible. These cells contained 19 and 28 prey, respectively. On 1 September, a second nest was found in the same area and excavated. The soil was moist, as a re- sult of recent rains, and the nest was shal- lower. Although it was not possible to fol- low the main burrow, a cache of 4 beetles was found at a depth of 24 cm and a cell containing 17 beetles was found at a depth of 27 cm. All the prey in both nests be- longed to one of two species of Phalacridae, Phalacrus sp. (N = 23) and Olibrus sp. (N = 3). Male C. echo echo were also com- mon in the area of the nest site and were frequently seen entering burrows, presum- ably in search of females. Cerceris fumipennis Say and C. californica Cresson These are both among the best-studied species of this genus, fumipennis being mainly an eastern species, ranging west to Wyoming and New Mexico, californica a common western species ranging east to Utah and Texas. There are many prey re- cords for both species, summarized by Scul- len and Wold (1969); both use a wide varie- ty of Buprestidae. These are related wasps, both belonging to group II of Scullen (1965). It was with considerable interest that we found the two nesting together 11- 12 June 1976 at Monahans Sandhills State Park, Ward Co., Texas. About six nests of C. californica californica were intermingled with about 30 nests of C. fumipennis in slightly sloping soil along the edge of a paved road. The soil was a rather firm, coarse sand containing many small stones; it had apparently been brought from another area into this region of sand dunes to serve as a base for the road. Hoplisoides splendi- dulus (Bradley) was also nesting close beside the two species of Cerceris (Rubink 1977). Females of both species brought in prey at frequent intervals during the day, car- rying the beetles in their mandibles and plunging into open nest entrances. Nests of C. californica had the entrance mounds only on one side of the hole, those of fumipennis surrounding the hole; otherwise there was no difference in external appearance. Nor was there any noteworthy difference in nest structure. We excavated 3 nests of califor- nica. In each case the mound at the en- trance measured about 4x5 cm, about 1 cm high. The burrow penetrated the soil obliquely or nearly vertically, terminating at a depth of only 8-12 cm, and containing 7-32 beetles in storage near the bottom. One nest had only one cell, the other two five cells each, the cells varying in depth from 7 to 13.5 cm (x=10). The number of beetles per cell varied from 7 fairly large ones to 31 very small ones. The two nests of fumipennis we excavated were of vir- tually identical structure. Both had an oblique burrow reaching a depth of 8-9 cm and containing beetles in storage at the ter- minus. One nest had only one cell, the oth- er 11. Cells varied in depth from 7 to 15 cm (x = 10.2). The number of beetles per cell varied from 15 to 16 (N = 3; most of the cells had cocoons and only fragments of beetles). We refrain from further discussion of nesting behavior since both species have been well studied elsewhere, californica by Linsley and MacSwain (1956), fumipennis by Evans (1971) and several others (refer- ences in Scullen and Wold 1969). However, we were especially interested in discovering to what extent females of the two species were competing for prey. Several species of Buprestidae appeared in the nests of both species (Table I). In general, fumipennis fe- males took beetles of medium size (5.5-10.5 mm in length). Females of californica also took beetles of this same size, but 41 per- cent of the prey consisted of a minute bu- prestid, Acmaeodera quadrivittata Horn, not utilized by fumipennis. However, four other species appeared in the nests of both spe- cies. On several occasions satellite flies, Seno- tainia sp., probably rubriventris Macquart, were seen following females carrying prey. Numerous male mutillids flew close above I March 1978 Evans, Rubink: Cerceris Wasp Nests 63 the ground in the morning hours. A number of females were also seen, some of them at- tempting to enter nests of C. fumipennis, though usually being repelled by the resi- dent wasp. The mutillids proved to be males and females of Dasyniutilla snow- oriim Cockerell and Fox. We have no defi- nite information that this species is a para- site of Cerceris, but the species was common in the nesting area and was not noted elsewhere. Acknowledgments We are indebted to the following special- ists for identifying the beetle prey: W. F. Barr(Buprestidae), J. M. Kingsolver (Phala- cridae), T. J. Spilman (Tenebrionidae), R. White (Chrysomelidae), and D. R. White- head (Curculionidae). R. J. Gagne identified the miltogrammine flies. Literature Cited Alcock, J. 1974. The nesting behaviour of Cerceris simplex macrosticta (Hymenoptera: Sphecidae). J. Nat. Hist. 8: 645-652. 1975. Social interactions in the soUtary wasp Cerceris simplex (Hymenoptera; Sphecidae). Be- haviour 54: 142-152. Cartwright, O. L. 1929. The maize billbug in South Carolina (Calendra maidis Chittn.). Bull. So. Carolina Agri. Exp. Sta. no. 257: 31 pp. Evans, H. E. 1971. Observations on the nesting behav- ior of wasps of the tribe Cercerini. J. Kansas Ent. Soc. 44: 500-523. 1974. Digger wasps as colonizers of new habitat (Hymenoptera: Aculeata). J. N.Y. Ent. Soc. 82: 259-267. Krombein, K. V. 1953. Kill Devil Hills wasps, 1952 (Hymenoptera: Aculeata). Proc. Ent. Soc. Wash. 55: 113-135. 1960. Biological notes on several southwestern groundnesting wasps (Hymenoptera: Sphecidae). Bull. Brooklyn Ent. Soc. 55: 75-79. Lin, C. S. 1967. Bionomics of Cerceris simplex graphica at Lake Texoma (Hymenoptera: Sphecidae: Phi- lanthinae). Proc. Ent. Soc. Wash. 69: 312-317. LiNSLEY, E. G. AND J. W. MacSwain. 1956. Some ob- servations on the nesting habits and prey of Cerceris californica Cresson (Hymenoptera: Sphecidae). Ann. Ent. Soc. Amer. 49: 71-84. Rau, p. 1928. Field studies in the behavior of the non- social wasps. Trans. Acad. Sci. St. Louis 25: 319-489. Rubink, W. L. 1977. Nesting behavior and prey re- cords of Hoplisoides splendidulus (Bradley). Southwestern Ent. 2: 1-3. ScuLLEN, H. A. 1965. A review of the genus Cerceris in America north of Mexico (Hymenoptera: Sphecidae). Proc. U.S. Natl. Mus. 116: 3.33-547. ScuLLEN, H. A., AND J. L. WoLD. 1969. Biology of wasps of the tribe Cercerini, with a list of Co- leoptera used as prey. Ann. Ent. Soc. Amer. 62: 209-214. Strandtmann, R. W. 1945. Observations on the nest- ing habits of some digger wasps. Ann. Ent. Soc. Amer. 38: 305-313. Table 1. Prey records for Cerceris fumipennis and C. californica, Monahans, Texas. Beetle species Acmaeodera quadrivittata Horn Acmaeodera variegata Leconte Agrilus pulchelltis Bland Chrysobothris (two species) Total Mean length x width (mm) 6.3 20.3 20.7 38.0 Wasp species fumipennis californica 51 50 11 1 62 58 14 2 125 Total 51 108 25 3 187 VERTEBRATE FAUNA OF A RADIOACTIVE LEACHING POND COMPLEX IN SOUTHEASTERN IDAHO Douglas K. Halford' and Jere B. Millard" Abstract.— An inventory of the terrestrial verte- brate fauna and the seasonal occurrence of each spe- cies was determined for a man-made radioactive leach- ing pond complex in southeastern Idaho. Three reptile, 11 mammal, and 94 bird species were identified from February 1974 through January 1978. Fewest species were observed during the winter and most during the summer. Eight bird species nested at the pond com- plex, while 19 species of birds were common or sea- sonally abundant. It appears that the Test Reactor Area radioactive leaching pond complex is an impor- tant water source and provides habitat for some spe- cies of wildlife. Since 1952, man-made leaching ponds at the Test Reactor Area (TRA) on the Idaho National Engineering Laboratory (INEL) Site in southeastern Idaho (Fig. 1) have been used to dispose of low-level liquid ra- dioactive wastes generated from various test reactors and their support facilities. A varie- ty of wildlife is attracted to the leaching pond complex which is one of the few sources of water available on or near the INEL Site. The Big Lost River flows inter- mittently and sometimes crosses as much as 50 km of the southwestern section of the site and then terminates in the Big Lost River Sinks. The Little Lost River termi- nates in the Little Lost River Sinks east of Howe (Fig. 1) before entering the site. The Big Lost and Little Lost rivers may dry up for a considerable portion of their length during the summer. During the winter they are often frozen. Mud Lake, which is 50 km northeast of the study area, is frozen during much of the winter, as are most other standing bodies of water in the area. How- ever, since the TRA pond complex receives heated effluents, water is available through- out the year. Since the pond complex contains radio- active isotopes, a potential exists for the movement of radionuclides from the area by indigenous animals. Therefore, it is im- portant to document which species use the pond so estimations can be made on the ex- port of radioactive materials and the poten- tial for radionuclide uptake from species taken and consumed by hunters. To determine which species of wildlife used the TRA pond complex, a species in- ventory was conducted. The purpose of this paper is to report species composition, abundance, and seasonal distribution of the vertebrate fauna which occur at the TRA radioactive leaching pond complex. The data from this study will provide informa- tion on wildlife attracted to a man-made ra- dioactive leaching pond and also provide a data base for studies being conducted on the pond ecosystem and the environmental impacts if the pond were drained or other- wise altered. Study Area The study was conducted at the TRA ra- dioactive leaching pond on the INEL Site, Butte Co. in southeastern Idaho (Fig. 1). The INEL Site covers an area of 2315 km-. The site is located on the cool desert shrub biome along the western edge of the upper Snake River Plain. The topography is flat to rolling, with an average elevation of 1470 m. The elevation of the study area is 1500 m. Annual precipitation averages 18-20 cm and temperatures range from -42 to 39 C, with an annual average of 6 C. Vegetation is dominated by desert shrubs, primarily big sagebrush {Artemisia tridentata), which cov- ers about 80 percent of the site; the under- 'Environmental Sciences Branch, Idaho National Enginering Laboratory, U.S. Department of Energy. Idaho Falls, Idaho 83401. 'Department of Radiology and Radiation Biology, Colorado State Uni- versity, Fort Collins, Colorado 80521. 64 March 1978 Halford, Millard: Pond Vertebrates 65 4NC-A-7200 Fig. 1. Location of Test Reactor Area leaching pond complex on the Idaho National Engineering Laboratory Site in southeastern Idaho. 66 Great Basin Naturalist Vol. 38, No. 1 story is composed of grasses and forbs (Har- niss and West 1973). The leaching pond complex is composed of three ponds excavated in gravelly allu- vium in 1952, 1957, and 1964. The low-lev- el liquid radioactive wastes disposed in the ponds percolate in the lithosphere. The ponds constnicted in 1952 and 1957 pres- ently constitute one continuous body of wa- ter 8200 m- in area with a mean depth of 3.0 m. Overflow of water from these ponds enters the adjacent 1964 pond. The 1964 pond is partially covered with shallow wa- ter forming a marshy area of about 1250 m^ A continuous 1.1 m high, 10 cm mesh wire and barbed-wire fence encloses all three ponds, forming a total area of 26,000 m-. Aquatic and littoral vegetation associated with disturbed sites have invaded the area. Littoral plants include thistles {Cirsium ar- venes and C. vulgare), speedwell {Veronica spp.), wild lettuce {Lactuca serriola), cheat- grass {Bromus tectorum), wild barley {Hord- ewn jiibatum), and willow {Salix spp.). The predominant shrub within the fenced area is rabbit brush {Chnjsothamnus nauseosiis and C. vicidiflorus). Sedges {Scirpus spp.) are the predominant macrophytes. A dense mat of periphyton, composed primarily of diatoms, is present on the pond bottom throughout the year. An autochthonous-based food web supports numerous aquatic arthropods. The vegetation outside of the fenced area con- sists primarily of big sagebrush, rabbit bmsh, and cheatgrass. Materials and Methods Birds, reptiles and large mammals were identified and counted inside the fenced area of the pond complex on 849 observa- tion days from 1 February 1974 to 31 Janu- ary 1978. Large mammals observed in close proximity to but outside of the fenced area were also listed on the species inventory. Mammal tracks identified inside the pond fence and along the outside perimeter of the fence were recorded as observations. Observations were made with 8 X 40 bi- noculars and a 20-power spotting scope be- tween 0800 and 1300 from a vehicle driven around the entire perimeter of the pond area. Small mammals were inventoried by trapping inside the pond fence with 70 Sherman live traps. Rodents were ear- tagged, released, and retrapped from July 1976 through November 1976 and May 1977 through October 1977. Trapping was conducted three consecutive nights weekly. Field guides were used to aid in identi- fication (Burt and Grossenheider 1964, Rob- bins et al. 1966, Stebbins 1966). Species abundance was calculated as the percentage of time a species was observed during the total observation period. Species were then placed into arbitrary categories or abundance according to the following criteria: Rare <4 Uncommon 5-19 Common 20-39 Abundant >40 Observed (but not counted) Results and Discussion Three species of reptiles, 11 species of mammals, and 94 species of birds were identified at the TRA pond complex (Table 1). Although not observed, the prairie rat- tlesnake {Crotalus viridis) and short-horned lizard {Phrynosaoma doiiglassii) were prob- ably present because they are known to oc- cur widely over the site and in habitat sim- ilar to that adjacent to the TRA pond complex (Sehman et al. 1976). The only am- phibian known to occur on the INEL Site, the Great Basin spadefoot toad {Scaphiopus intertnontamis), was not observed at the TRA pond complex. Small mammals trapped at the TRA pond complex were resident animals (Halford and Markham 1978). Evidence of small mammal activity, such as burrows, was present. Since trapping was not continued during the win- ter, the seasonal distribution of rodents is not complete. However, the species cap- tvired and observed during spring, summer, and fall are probably representative of the winter occurrence, although some species hibernate. The most abundant small mam- mal at the TRA pond complex was the white-footed deer mouse {Peromyscus maniculatiis) and the least abundant was March 1978 Halford, Millard: Pond Vertebrates 67 the meadow vole (Microtus pennsylvanicus). Pronghorn antelope {Antilocapra ameri- cana) are the most abundant large mammal on the INEL Site and were observed regu- larly outside the wire perimeter on the TRA complex. On one occasion, 20 ante- lope were seen feeding on the vegetation around the marshy portion of the pond. These animals apparently entered the pond complex through an improperly closed gate. No drinking by antelope was noted. Mule deer {Odocoileiis hemionus) were observed drinking and feeding in the TRA pond com- plex on several occasions and may have re- lied on the pond as a water source. Coyotes {Canis latrans), the only large predator noted, were seen infrequently. Most identifications were made from tracks or prey remains, usually sage grouse, found inside the fenced area. A large diversity of birds was observed at the TRA pond complex (Table 1). Of the 94 species identified, 19 were common or abundant seasonally, 19 were sighted once, and 8 nested. Marsh hawks {Circus cyaneus), killdeer {Charadrius vociferus), spotted sandpipers {Actitis tnaciilaria), barn swallows {Hiriindo rustica), and mallards (Anas platyrhynchos) nested each year of the study period, and blue- winged teal (A. discors). Brewer's blackbirds {Euphagus cya- nocephalus), and sage thrashers {Oreoscoptes montanus) were known to nest but once. Bam swallows were the most abundant nester, building over 20 nests each year in a sewage facility 100 m north of the TRA pond complex. Other uncounted nests were located on buildings associated with the TRA complex. Nest material was obtained from the TRA pond by the swallows. Other species, particularly sage {Amphisipiza belli) and Brewer's sparrows (Spizella breweri), probably nest in the area. The most bird species were observed dur- ing summer and the least during winter. Spring and fall counts also showed an abun- dance of birds (Table 2). The Passeriformes was the most frequently represented order, with the highest species counts occurring during spring and summer. Anseriformes was the next most frequently represented order, with most species observed during the spring and fall migration periods. Three of the 15 orders account for more than 75 percent of the species identified (Table 2). The most frequently observed birds were mallards and green-winged teal (Anas crecca). Mourning doves (Zenaida macroura), sage grouse {Centrocercus urophasianus), and wa- terfowl were game birds that frequently used the pond. Other species, such as pheas- ant {Phasianus colchicus) and gray partridge (Perdix perdix), were observed infrequently (Table 1). Mourning doves appeared to use the pond for drinking water and waterfowl used the pond as a resting area. Sage grouse with broods were observed in early summer, and adult flocks were observed during the fall. Previous studies (Halford et al. 1976, Markham 1976) indicate that doves and wa- terfowl ingest or become externally con- taminated with radionuclides from the TRA pond environs. However, these birds have relatively low concentrations of radionuclide contamination. Radionuclide concentrations in sage grouse (Jack Connelly, pers. comm. Dec. 1977), barn swallows, marsh hawks, and American kestrels (Falco sparverius) were also assessed (Millard et al. 1976, Craig et al. 1978) and found to be low. Since none of the birds inventoried stay at the pond for long periods, it is unlikely that they would accumulate sufficient concentrations of ra- dionuclides to be harmful. Studies are cur- rently being conducted to determine what hazard, if any, the TRA pond complex has on fauna utilizing it. Craig (1977) found 12 species of raptors present on the INEL Site, although we ob- served only 4 species at the TRA pond complex. The marsh hawk and American kestrel were the most common species sighted, and both species nested in or near the pond complex. Young marsh hawks and kestrels from these nests had low levels of radionuclides. Nuclides found in nestling raptors were those found in the TRA pond environs, indicating that these birds obtain- ed some prey items from the TRA pond complex (Craig et al. 1978). This species inventory indicated that the TRA radioactive leaching pond complex may provide a source of food, water, and other habitat requirements for several spe- 68 Great Basin Naturalist Vol. 38, No. 1 Table 1. Seasonal occurrence and abundance of reptiles, mammals and birds at the Test Reactor Area leach- ing ponds in southeastern Idaho. Seasonal Occurrence and Abundance Species Spring Summer Fall Winter Reptiles Bull Snake {Pitiiophis melanoleticus) Western Terrestrial Garter Snake {Thamnophis elegans) Sagebrush Lizard {Sceloporus graciosus) Mammals Coyote (Cants latrans) Long-tailed Weasel (Mustela frenata) Least Chipmunk (FAitamias minimus) Ords Kangaroo Rat {Dipodomys ordii) Western Harvest Mouse (Reithrodontomys megalotis) White-footed Deer Mouse (Peroniyscus maniculatus) Meadow Vole (Microtus pennsylvanicus) Mountain Cottontail {Sylvilagus nuttalli) Pygmy Rabbit {Sylvilagus idahoensis) Mule Deer (Odocoileus hemionus) Pronghom {Antihcapra americana) Western Grebe {Aechmophorus occidentalis) Eared Grebe {Podiceps nigricollis) Pied-billed Grebe (Podilymbus podiceps) White Pelican (Pelecanus erythrorhynchos) Whistling Swan {Olor columbianus) Snow goose {Chen caendescens) Canada goose {Branta canadensis) Mallard {Anas platyrhynchos) Pintail {Anas acuta) Gadwall (Anas' strepera) American Wigeon {Anas am.ericana) Shoveler {Anas clypeata) Blue-winged Teal {Anas discors) Cinnamon Teal {Anas cyanoptera) Green-winged Teal {Anas crecca) Wood Duck {Aix sponsa) Redhead {Aythya americana) Canvasback {Aythya valisineria) Ring-necked Duck {Aythya coUaris) Lesser Scaup {Aythya af finis) Common Goldeneye {Bucephala clangula) Barrow's Goldeneye {Bucephala islandica) Bufflehead {Bucephala albeola) Ruddy Duck {Oxyura jamaicensis) Common Merganser {Mergus merganser) American Kestrel {Falco sparverius) Marsh Hawk {Circus cyaneus) Swainson's Hawk {Buteo swainsoni) Ferruginous Hawk {Buteo regalis) Sage Grouse {Centrocercus urophasianus) Ringneck Pheasant {Phasianus colchicus) Gray Partridge {Perdix perdix) Great Blue Heron {Ardea herodias) Green Heron {Butorides virescens) American Bittern {Botaurus lentiginosus) White-faced Ibis {Plegadis chihi) American Coot {Fulica americana) American Avocet {Recurvirostra americana) Killdeer {Charadrius vociferus) Birds r+ + r+ + r+ + r+ + r+ + r+ + March 1978 Halford, Millard: Pond Vertebrates 69 Spotted Sandpiper (Actitis macularia) Solitary Sandpiper {Tringa solitaria) Lesser Yellowlegs {Tringa flavipes) Common Snipe (Capelki gallinago) Willet (Catoptrophoriis semipalmatus) Marbled Godwit {Liinosa fedoa) Wilson's Phalarope (Steganopus tricolor) Herring Gull {Lams argentatus) Ring-billed Gull (Lartis delawarensis) Bonaparte's Gull {Larus Philadelphia) Black Tern {Chlidonias niger) Caspian Tern {Hydroprogne caspia) Mourning Dove {Zenaida macroura) Great Homed Owl {Bubo virginianus) Common Nighthawk {Chordeiles minor) Rufous Hummingbird {Selasphorus rufus) Belted Kingfisher {Megaceryle alcyon) Common Flicker {Colaptes auratiis) Eastern Kingbird {Tyrannus tyrannus) Western Kingbird {Tyrannus verticalis) Say's Phoebe {Sayornis saya) Homed Lark {Eremophila alpestris) Bam Swallow {Hinindo rustica) Cliff Swallow {Petrochelidon pyrrhonata) Violet-green Swallow {Tachycineta thalassina) Tree Swallow {Iridoprocne bicolor) Bank Swallow {Riparia riparia) Rough-winged Swallow {Stelgidopteryx rnficollis) Black-billed Magpie {Pica pica) House Wren {Troglodytes aedon) Sage Thrasher {Oreoscoptes montanus) American Robin {Turdus migratorius) Ruby-crowned Kinglet {Regidus calenditki) Loggerhead Shrike {Lanius ludovicianus) Starling {Sttirnus vulgaris) Yellow-rumped Warbler {Dendroica coronata) Yellow Warbler {Dendroica petechia) Townsend's Warbler {Dendroica townsendi) Western Tanager {Piranga ludoviciana) Western Meadowlark {Sturnella neglecta) Yellow-headed Blackbird {Xanthocephalus xanthocephalus) Red-winged Blackbird {Agelaius phoeniceus) Brewer's Blackbird {Euphagus cyanocephalus) Brown-headed Cowbird {Molothrus ater) House Finch {Carpodacus mexicanus) Dark-eyed Junco {Junco hyemalis) Lapland Longspur {Calcarius lapponicus) White-crowned Sparrow {Zonotrichia leucophrys) Chipping Sparrow {Spizella passerina) Sage Sparrow {Amphispiza belli) Lark Sparrow {Chondestes gammacus) Song Sparrow {Mehspiza melodia) Grasshopper Sparrow {Ammodramus savannarum) Brewer's Sparrow {Spizella breweri) Snow Bunting {Plectrophenax nivalis) r+ + r+ + "*■ r = rare, c = common, u = uncommon, a = abundant, o = observed (numbers undetermined) Single observation ° Known nester 70 Great Basin Naturalist Vol. 38, No. 1 Table 2. Numbers of bird species by order observed seasonally at the Test Reactor Area radioactive leaching pond complex. Bird Orders Number of Species Observed Seasonally Spring Summer Fall Winter Total Species Podicipediformes Pelecaniformes Anseriformes Falconiformes Galliformes Ciconiiformes Grui formes Charadriiformes Columbiformes Strigiformes Caprimulgiformes Apodiformes Coraciiformes Piciformes Passeriformes Totals 2 1 3 1 3 0 0 1 0 1 16 8 16 6 21 1 4 1 0 4 2 1 2 0 3 0 4 0 0 4 1 1 1 1 1 5 9 8 0 14 1 1 1 1 1 0 0 1 0 1 0 0 0 1 0 0 0 1 1 0 0 1 1 0 28 31 16 4 37 57 64 51 13 94 cies of wildlife which may otherwise not be available on the INEL Site. Because the area is protected from public interference, it may also act as a refuge for certain spe- cies. The TRA pond complex provides a unique opportunity to study the movement of radionuclides in an ecosystem and the ef- fects of a radioactive environment on in- digenous species. Acknowledgments We thank B. W. Mortensen for his field assistance and O. D. Markham for critically reviewing the manuscript. Funds for this project were provided by the Division of Biomedical and Environmental Research, U.S. Department of Energy. Literature Cited Burt, W. H., and R. P. Grossenheider. 1964. A field guide to the mammals. Houghton Mifflin Co., Boston. 284 pp. Craig, T. H. 1977. Raptors of the Idaho National En- gineering Laboratory Site. Unpubl. master's thesis. Idaho State University, Pocatello. Craig, T. H., D. K. Halford, and O. D. Markham. 1978. Radionuclide concentrations in nestling raptors near nuclear facilities. Wilson Bulletin (in press). Halford, D. K., J. B. Millard, and O. D. Markham. 1976. Waterfowl study of the Test Reactor Area radioactive leaching pond, pp. 19-34. In: 1975 progress report, Idaho National Engineering Laboratory Site radioecology-ecology programs. IDO-12080. Nat. Tech. Information Service, Springfield, Virginia. Halford, D. K., and O. D. Markham. 1978. Radiation dosimetry of small mammals inhabiting a liquid radioactive waste disposal area. Ecology (in press). Harniss, R. O., and N. E. West. 1973. Vegetation patterns of the National Reactor Testing Sta- tion, southeastern Idaho. Northwest Sci. 47:30-43. Markham, O. D. 1976. Gamma emitting isotopes in mourning doves on the INEL site, pp. 58-64 In: 1975 progress report, Idaho National Engi- neering Laboratory Site radioecology-ecology Programs. IDO-12080. Nat. Tech. Information Service, Springfield, Virginia. MiLi^RD, J. B., F. W. Whicker, and O. D. Markham. 1976. An ecological study of the Test Reactor Area radioactive leaching ponds, pp. 17-18. In: 1975 progress report, Idaho National Engineer- ing Laboratory Site radioecology-ecology pro- grams. IDO-12080. Nat. Tech. Information Ser- vice, Springfield, Virginia. RoBBiNs, C. S., B. Bruun, H. S. Zim, and A. Singer. 1966. A guide to field identification— birds of North America. Racine, Wisconsin: Western Publ. Co. 340 pp. Sehman, R. W., a. D. LiNDER, D. Wyckoff, and W. E. Christopher. 1976. A study of the amphibi- an and reptilian fauna of the INEL Site, pp. 121-131. In: 1975 progress report, Idaho Na- tional Engineering Laboratory Site radioecology- ecology programs. IDO-12080. Nat. Tech. Infor- mation Service, Springfield, Virginia. Stebbins, R. C. 1966. A field guide to western reptiles and amphibians. Boston: Houghton Mifflin Co. 279 pp. NEW SYNONYMY, NEW SPECIES, AND TAXONOMIC NOTES OF NORTH AMERICAN PITYOPHTHORUS (COLEOPTERA: SCOLYTIDAE), PART III Donald E. Bright' Abstract.— New synonymies and new data affecting North American Pityophthortis are proposed as follows: chalcoensis Hopkins { = herrerai Hopkins) and nigricans Blandford ( = chiapensis Bright). Neotypes are designated for the following species described by Eichhoff: concentralis, cribripennis, infans, puchellus, and tuberculatus. New species are: carinatus (Canada), cavatus (Manitoba, Saskatchewan), euterpes (Mexico), hesperius (British Co- lumbia), impexus (Mexico), intentus (Arizona), laticeps (Mexico), rnalleatus (Arizona), mesembria (Guatemala), montezumae (Mexico), scalptus (British Columbia, Colorado), trepidus (California), and vespertinus (Mexico). This is the third paper under this general title wherein various new taxonomic data affecting the classification of the North American Pityophthortis are proposed. A taxonomic monograph of North Ameri- can Pitijophthorus is being prepared, but the data contained herein is needed for other works in preparation. It was therefore de- cided to publish these data now in order that they may be used without delay. The collections where type material is lo- cated are abbreviated as follows: British Museum (Natural History), London (BMNH); Canadian National Collection of Insects, Ottawa (CNC); Karl E. Schedl col- lection, Lienz, Austria (KESC); S. L. Wood collection, Provo, Utah (SLWC), and United States National Museum of Natural History, Washington, D.C. (USNM). Acknowledgments Various individuals assisted in this study by loaning specimens, and I wish to express my appreciation to the following: Dr. D. M. Anderson (USNM); Prof. K. E. Schedl, Lienz, Austria; Dr. R. T. Thompson (BMNH); and Dr. S. L. Wood, Provo, Utah. Dr. S. L. Wood also offered valuable com- ments and opinions concerning the data presented herein. Pityophthortis chalcoensis Hopkins Pityophthorus chalcoensis Hopkins, 1905, p. 73 (Holo- type ? , Mexico, USNM) Pityophthortis herrerai Hopkins, 1905, p. 74 (Holotype ? , Mexico, USNM). New synonymy This species, and its synonym herrerai, un- til recently have remained unknown since their description. References to neither name were found except for listings of the names in catalogs. When reading the descriptions of the two species, it seems obvious that two distinct entities are present. P. chalcoensis is de- scribed as having the second declivital in- terspace deeply impressed and the first and third declivital interspaces with distinct granules. P. herrerai is described as having the second declivital interspace flat, the first declivital interspace with a few irregu- lar granules and the third without distinct granules. With just four specimens of the combined type series of both species to ex- amine, these distinctions can be seen. How- ever, a short series of specimens referred to this species from Hidalgo, Mexico, has also been examined. One female in this series is identical to the female holotype of chal- coensis, one male is identical to the male paratype of herrerai and another female is very close to the female paratype of her- rerai. It seems, therefore, that the differ- ences noted by Hopkins are only variations in the species and only one name should be recognized. The frons of adults of both sexes is almost identical. Both are very broadly flattened and densely pubescent. The only sexual dis- tinction I could detect is the presence of a 'Biosystematics Research Institute, Canada Agriculture, Ottawa, Ontario. 71 72 Great Basin Naturalist Vol. 38, No. 1 slightly elevated, impiinctate callus at the midpoint of the upper margin of the male frons. Unfortimately, this callus is almost al- ways concealed by the anterior margin of the pronotum and is only visible if the head is fully extended or if the anterior margin of the pronotum is broken away. In addi- tion, adults may be recognized by the lack of a pronotal summit, by the densely punc- tured posterior portion of the pronotum, by the large strial punctures which form ir- regular strial rows, and by the steep, bisul- cate elytral declivity which usually bears prominent granules on interspaces 1 and 3. Pityophthorus nigricans Blandford Pitijaphtfwrus nigricans Blandford, 1904, p. 236 (Lecto- type <5 , Guatemala, BMNH) Pityophthorus chiapensis Bright, 1977, p. 522 (Holo- type "i , Mexico, CNC). Neiv synonymy The type material of P. chiapensis was compared to my homotypes of P. nigricans and found to represent the same species. The association of these two names was not apparent until the species described as P. le- pidus Bright was recognized. A series of P. lepidus was previously considered to be P. nigricans and is therefore quite different from specimens subsequently described as P. chiapensis. At the suggestion of S. L. Wood, the specimens under all the above names were reexamined with the result that P. chiapensis must be merged with P. nigr- icans. Neotype Designations W. J. Eichhoff, over a period of years be- tween 1868-1878, described a number of species of North American Pityophthorus. No original type specimens have been found despite the considerable correspond- ence with European museums where the types might have been housed. Presumably the specimens were in the Hamburg Mu- seum (where other Eichhoff types were held) when the museum was destroyed dur- ing World War II. Hans Eggers, a noted authority on the Scolytidae, had the opportunity in 1927 of comparing a number of North American species with the types in European mu- seums. The specimens Eggers compared with the types are in the United States Mu- seum of Natural History in Washington, D.C. Compared specimens representing five of Eichhoff's North American Pityophthorus have been located. These are listed below, along with notes concerning one additional species. Neotypes are now designated for all of Eichhoff's species of North American Pi- tyophthorus except P. hisulcatus. P. concentraUs, 1878, p. 188. No speci- mens bearing Eggers's "compared to type" label have been located. There is a series of four specimens in the USNM that were ex- amined by Eggers and the first of these bears the label "nicht Pityophthorus con- centralis Eichh., Eggers 1927." However, Eichhoff himself identified some specimens in 1892 that had been collected in Florida by Hubbard and Schwarz. Eichhoff, in Eichhoff and Schwarz (1895), states "P. con- centraUs from Florida is correctly deter- mined and does not differ in the least from my typical specimen from Cuba. . . ." The specimens sent to Eggers in 1927 were from the same series that was seen by Eichhoff in 1892. Obviously, either Eichhoff or Eggers made a mistake and I am inclined to follow the o pinion of Eichhoff, the original de- scriber of P. concentraUs. Therefore regard- less of Eggers's label, in the interest of sta- bility of nomenclaure, I designate the first specimen in the USNM series seen by Egg- ers as the NEOTYPE. It bears the data: "Biscayne, Fla., 29.5"/"Coll. Hubbard and Schwarz"/"9" (on a white square) /"nicht Pityophthorus concentralis Eichh." (in Egg- ers's handwriting), Eggers 1927/ "NEOTYPE Pityophthorus concentralis Eichh., D. E. Bright 1977." P. cribripennis, 1869, p. 274. Neotype here designated, bears the data: "Ripley, Mississippi, 4-21-20"/"M.W. Blackman, col- lector"/"N.Y.S.C.F. Lot Mi 261"/"Pityoph- thorus cribripennis Eichh. mit type in coll. Eichhoff verglichten" (in Eggers's hand- writing)/"NEOTYPE Pityonhthorus cribri- pennis Eichhoff. D. E. Brignt 1977." Syn- onym of P. pulhis Zimmermi^n. P. infans, 1872, p. 135. Neotype, here designated, bears the labels: "Syracuse, N.Y., 5-14-18"/"M.W. Blackma.i, collector"/ "N.Y.S.C.F. Lot 1138"/"compared with type of P. infans Eichh. by H. Eggers. Five specimens, all typical" (large red label. March 1978 Bright: American Scolytidae 73 handwriting unknown)/"NEOTYPE Pi- tyophthorus infans Eichhoff, D. E. Bright 1977." Synonym of P. puberulus Le Conte. P. pulchellus, 1869, p. 275. Neotype ( ? ), here designated, is labeled: "Marquette, Mich'VColl. Hubbard and Schwarz'7 " ? '7"13"/"Pityophthorus pulchellus Eichh., m. type in coll. Eichhoff verglichen" (in Eggers's handwriting)/ "NEOTYPE Pityophthorus pulchellus Eichhoff, D. E. Bright, 1977." P. tuberculatus, 1878, p. 498. Neotype {$), here designated, bears the data: "Hopk. U.S. 471a"/"Black Hills, S.D., Hopkins Coir."/ "Pityophthorus tuberculatus Eichh., m. type in coll. Eichhoff verglichen" (in Eggers's handwriting)/ "5" /"NEOTYPE Pi- tyophthorus tuberculatus Eichhoff, D. E. Bright, 1977.") New Species Pityophthorus carinatus, n.sp. Length 1.9-2.2 mm, 3.0 times longer than wide. Female.— Frons moderately concave on each side of a very strongly elevated, longi- tudinal carina, this carina sharp to moder- ately broad on summit, sometimes bearing short setae along summit, the carina usually more strongly elevated on epistomal area and extending to upper level of eyes; con- cave areas extending laterally nearly to eyes, surface moderately shining near eye, weakly and very finely punctured; an oval patch of extremely short setae forms a "spongy" patch in midarea, this patch ex- tending upward on slopes of elevated carina to summit. Antennal club oval, 1.2-1.3 times longer than wide, widest through seg- ment 2; sutures 1 and 2 weakly arcuate, 2 more strongly so; segments 1 and 2 occupy more than half the total club length. Pronotum about 1.2 times longer than wide, widest at level of summit; sides sub- parallel on posterior half; anterior margin broadly rounded, with about a dozen small, basally contiguous serrations; asperities on anterior slope low, small, usually isolated, scattered in no apparent order; summit dis- tinct, high, with transverse impression dis- tinctly indicated; posterior area distinctly punctured, the punctures rather small and shallow, separated by a distance equal to or less than their diameters; interpuncture sur- face moderately shining and rather densely, minutely reticulate; median line narrow, sometimes weakly elevated on anterior half or less. Elytra about 1.8 times longer than wide; apex broadly rounded; striae and interstriae with numerous, intermixed punctures, mak- ing discernment of striae difficult, but striae are discernible because strial punctures are more numerous and arranged in a regular row, the punctures about equal in size and depth to those on posterior portion of pro- notum. Declivity deeply impressed; inter- space 1 weakly, narrowly elevated, deeply impressed below level of interspace 3; inter- space 2 slightly widened, distinctly sulcate, the lateral portions raising abruptly toward interspace 3; interspace 3 distinctly ele- vated, much higher than 1, bearing a row of fine granules on summit; punctures in striae 1 and 2 obsolete. Male.— Frons flattened from eye to eye and from epistoma to upper level of eyes, divided by a strongly elevated, laterally flat- tened longitudinal carina, this carina more strongly elevated on lower portion on epis- tomal region; surface of flattened area strongly punctured, the punctures rather long and deep. Pronotum and elytra as in female. Declivity convex; lateral pro- tuberances strongly to weakly elevated, strongly turned or pushed inward toward sutue. This species occurs in two distinct popu- lations, which are designated as subspecies. 1. Summit of longitudinal, median elevation on female frons sharp, usually devoid of setae; protuberances on male declivity strongly elevated, blunt, extending inward over suture; occurs in NE United States and E Canada P. carinatus carinatus, n. subsp. Summit of median elevation on female frons broad, with sliort setae; protuberances on male declivity not strongly elevated, pushed inward toward sutuj-e; occurs in British Columbia and Alberta P. carinatus monticolae, n. subsp. 74 Great Basin Naturalist Vol. 38, No. 1 Type material.— P. carinatus carinatus, n. subsp. Holotype ( ? ), U.S., New York, Sevey, 8-VIII-1970, D. E. Bright/on Picea/ ? /HOLOTYPE Pityophthorm cari- natus carinatus, D. E. Bright, 1977, CNC No. 15489. Allotype and two paratypes with same data. Nine paratypes are labeled: 64-1340-01, Ex. branch white pine/2 mi. E. of Jemseg, Queen's Co., N.B. 21 May 1965 F.I.S. Paratypes are in the CNC and the SLWC. P. carinatus monticolae, n. subsp. Holo- type (?) is labeled: No. 70-569-01, Date II.VI-1970, F.I.S./P. contorta, Hazelton, B.C./ ? /HOLOTYPE Pityophthorus cari- natus monticolae D. E. Bright, CNC No. 15490. Two paratypes bear the same data. Allotype and three paratypes are labeled: Edmonton, Alta., 5-VI1-1924/A. T. McClay collection. All type material is in the CNC. Remarks.— This species represents a very distinctive form that is easily recognized by the very strongly elevated, median carina on the female frons. The area on each side of the median carina is moderately concave and "spongy" on the inner portions and on the carina. In the eastern population the ca- rina is very sharp along the summit and usually not pubescent, while in the western population the carina is broader and pub- escent on the summit. The male declivity of the eastern popu- lation resembles that of P. cariniceps Le- Conte, but is more variable in the western series. This species probably is related to P. ca- riniceps and could conceivably represent an extreme variation of that species. However, no intergrades between P. carinatus and P. cariniceps have been seen and the size dif- ference seems consistent. Pityophthorus cavatus, n.sp. Length 1.9-2.3 mm, 3.0 times longer than wide. Female.— Frons deeply concave in cen- tral area, concavity fringed by a dense brush of erect, coarse setae, central portion of concavity glabrous but may also bear densely placed, erect setae, these setae usu- ally equal in length to those on periphery; an obscure to distinct, laterally flattened. acute tooth is usually present on midpoint of epistoma; surface above and lateral to concavity deeply punctured, the punctures large, close, space between punctures shin- ing. Antennal club nearly round, about as long as wide; sutvire 1 transverse or very weakly arcuate, 2 more strongly arcuate; segments 1 and 2 occupy more than half the total club length. Pronotum about 1.3 times longer than wide; sides weakly arcuate; anterior margin broadly rounded, bearing about 10 erect, generally isolated serrations; asperities on anterior slope erect, of moderate size, and scattered in no apparent order; summit dis- tinct; transverse depression behind summit distinct; posterior portion densely punc- tured, the punctures rather large, deep, and close; interpuncture space moderately shin- ing, reticulate; median line narrow, impunc- tate. Elytra about 1.8 times longer than wide; apex broadly rounded; discal striae and in- terstriae punctured in regular rows, making discernment of striae or interstriae difficult, strial punctures more numerous and each bears a very short seta, interstrial punctures much less numerous and each bears a longer erect seta; surface of interspaces shining, with numerous fine lines or very faintly ret- iculate. Declivity convex, bisulcate; inter- space 1 weakly elevated, impressed below level of 3, devoid of granules; interspace 2 weakly sulcate, slightly wider than discal width, surface as on disc; interspace 3 weakly elevated, higher than 1, bearing a median row of 2-4 small, rounded granules; punctures in striae 1 and 2 obsolete. Male.— Only two male specimens of this species are known, and each differs consid- erably from the other in the characters of the declivity. Variety I. Frons flattened from upper level of eyes to epistoma, divided by a strongly elevated longitudinal carina, high- est at epistoma and slanting downward on upper areas. Pronotum as on female. Elytra as on female except interstrial punctures more numerous and setae slightly longer. Declivity almost evenly convex, with a me- dian, elongate rounded elevation that ap- pears to arise in interspace 2 and is slightly bent inward over interspace 1; interspace 3 March 1978 Bright: American Scolytidae 75 not elevated or granulate; surface of declivi- ty moderately shining, minutely reticulate with weak strial punctures. Variety 2. Frons distinctly impressed from epistoma to near upper level of eyes, di- vided by a strongly elevated, laterally flat- tened, toothlike carina; generally similar to variety 1. Pronotum and elytra as in variety 1. Declivity convex; interspace 1 weakly impressed, devoid of granules; interspace 2 weakly sulcate; interspace 3 bearing a large, rounded protuberance in middle of declivi- ty, this protuberance directed caudad, inner slope precipitous, outer slope oblique; sur- face of declivity as in variety 1. Type material.— The holotype ( ? ) and four paratypes were collected at Canoe Lake, Saskatchewan, on 21 July 1972, from branches of Finns banksiana, by D. E. Bright (CNC No. 15492). Six additional par- atypes were collected at Charlottetown, Manitoba, on 13 August 1954, from branches of red pine. The holotype and most of the paratypes are in the CNC. Additional paratypes are in the SLWC. Remarks.— This is a unique species that can be easily recognized by the characters given in the description. It is obviously a member of the cariniceps group, but its relationship to other species is not clear. Some female specimens bear a vague resem- blance to some specimens of P. biovalis Blackman, but this does not indicate a true relationship to that species. Pityophthorus euterpes, n. sp. Length 1.5-1.8 mm, 3.0-3.1 times longer than wide. Male.— Frons convex, bearing a very faint, longitudinal carina which extends from the weakly elevated epistoma to the upper eye level; surface shining, closely, deeply punctured, the punctures separated by a distance less than half their own diam- eters; vestiture inconspicuous. Antennal club 1.5 times longer than wide, widest through segment 2; sutures 1 and 2 transverse; seg- ments 1 and 2 occupy about half the total club length. Pronotum 1.2 times longer than wide, widest at about middle; sides weakly ar- cuate; anterior margin rather narrowly rounded, bearing about 10 separated serra- tions; asperities on anterior slope arranged into three distinct, concentric rows, two other indistinct concentric rows at summit, some of those in rows 1 and 2 slightly off- set; summit distinctly but weakly elevated; posterior area strongly punctured, the punc- tures deep, moderately large, separated by a distance equal to about half or less of their diameters; interpuncture space brightly shining, smooth, with scattered fine points; median line broad, scarcely elevated. Elytra 1.7 times longer than wide; apex broadly rounded, weakly produced at su- ture; discal striae punctured in regular rows, the punctures rather large, deeply im- pressed; discal interspaces weakly convex, about 1.5 times wider than striae, the sur- face shining, impunctate, very finely sculp- tured with fine lines and points. Declivity deeply sulcate; interspace 1 weakly elevated above 2, bearing a median row of fine se- taceous granules; interspace 2 not wider than discal width, smooth to apex; inter- space 3 abruptly elevated, much higher than 1, bearing about six prominent, acute granules along summit; punctures of striae 1 and 2 distinct, impressed, somewhat smaller than those on disc. Vestiture mostly con- fined to declivital region, the setae rather long, fine, and sparse. Female.— The form suspected to be the female is nearly identical to the male ex- cept declivital interspace 3 is much less strongly elevated with much finer granules. Type material.— The holotype ( $ ), and five paratypes were collected 21 miles (35 km) west of Lazaro Cardenas, Chiapas, Mexico, on 26 June 1969 from Piniis oo- carpa, by D. E. Bright (CNC No. 15167). Three additional paratypes were collected at Lagos des Colores (Lagunas de Monte- bello National Park), Chiapas, Mexico, on 14 June 1969, by D. E. Bright, and one par- atype is from Ocosingo Valley, Chiapas, Mexico, collected on 7 July 1960, by Stan- nard. The holotype and most of the paratypes are in the CNC; additional paratypes are in the SLWC and the KESC. Discussion.— This species is closely re- lated to P. obtusipennis Blandford, but the 76 Great Basin Naturalist Vol. 38, No. 1 adults of P. euterpes may be distinguished by their smaller, more slender body and by the finer granules on the declivital inter- spaces 1 and 3. Pityophthorus hesperitis, n. sp. Length 2.1-2.3 mm, about 3.0 times long- er than wide. Female.— Frons variable, varying from weakly, longitudinally sulcate in the median area, with the lateral elevations of the sul- cus weakly to moderately elevated and spongy, to frons moderately protuberant in median area and spongy over entire surface; a small, laterally flattened tooth is usually located on epistoma of all forms; surface of remainder of frons moderately deeply punc- tured. Antennal club as in concaviis. Pronotum, elytra, and declivity as in bal- sameus and concavus. Male.— Frons flattened from epistoma to upper level of eyes, divided by a small, lat- erally flattened, distinct carina, the upper margin of flattened area elevated into a transverse, moderately to weakly elevated ridge; surface moderately punctured, shin- ing. Pronotum and elytra as in bahameiis and concavus. Declivity convex; median protuberances arising on interspace 3 pushed inward toward suture or appearing that each interspace 1 is pinched together. Type material.— The holotype (?) is la- beled: No. 74-1571-01, Date: 21-VIII-74, F.I.S. 1974/P. contorta, Hudson Hope, B.C. /Holotype Pityophthorus hesperius D. E. Bright, 1977, (CNC No. 15491). The allo- type bears the same data except the date is 3-IX-1974. Paratypes as follows: 3, same as holotype except dated 6-IX-1974; 5, same except dated 21-VIII-74; 1, same except dated 9-IX-1974; 1, same except dated 16- VIII-74; 1, Hixon, B.C., VII-9-1972, D. E. Bright/Pinus contorta; 1, Prince George, B.C., Pinus contorta/No. 72-1143-01, 14-XI- 72, F.I.S. 1972; 4, Blackwater R., 21-VIII- 57/57-7671-01, Pinus contorta and 1, 57- 7672-02, Pinus contorta/ 158 Mile House, B.C., 15-VIII-57, F.I.S. The holotype, allotype, and most of the paratypes are in the CNC. Additional para- types are in the SLWC and the KESC. Remarks.— This species contains females that resemble both P. balsarneiis Blackman and P. concavus Blackman. In the earlier stages of this study I considered it as a sub- species of P. balsameiis. Wood (pers. comm.) was inclined to associate it with P. concavus. A reexamination of the series showed that forms resembling both species were present, even within populations from one locality. After careful consideration, I decided to treat it as a separate species. More collecting is needed, particularly in British Columbia, Alberta, and the adjacent regions to definitely establish its status. Pityophthorus impexus, n. sp. Length 1.9-2.1 mm, about 3.1 times long- er than wide. Female.— Frons weakly convex on a large semicircular area extending from epis- tomal margin to well above upper level of eyes and laterally nearly from eye to eye; surface shining, bearing large, deep punc- tures on periphery, these becoming smaller toward middle, median portion weakly flat- tened and impunctate on a small area just above epistomal margin; vestiture rather sparse, consisting of long, erect setae scat- tered over the surface, those on periphery somewhat longer and incurved. Antennal club large, about 1.2 times longer than wide, widest through segment 3; segment 1 small, narrower than 2 or 3; suture 1 weak- ly arcuate, 2 more stronglly so; segments 1 and 2 occupy less than half the total club length. Pronotum 1.1 times or less longer than wide; sides weakly arcuate on basal half; anterior margin broadly roimded, bearing about 6 to 8 distinct serrations, these gradu- ally increasing in size toward middle; aspe- rities on anterior slope moderately large, erect, usually isolated but 2 or 3 occasion- ally basally contiguous; summit distinct, high; posterior portion densely punctured, the pimctures large and deeply impressed, separated by a distance equal to about half their diameters; interpuncture surface mod- erately shining, rather densely micro- reticulate; median line broad, not elevated, narrowed just behind summit. Elytra 1.7 times longer than wide; apex broadly rounded; striae punctured in regular March 1978 Bright: American Scolytidae 77 rows, the punctures about equal in size to those on posterior portion of pronotum, rather deeply impressed; discal interspaces about 1.0 to 1.5 times wider than striae, im- punctate (except 1), surface moderately shining, densely microreticulate. Declivity convex, somewhat flattened between the third interspaces; interspace 1 equal in width to its discal width, slightly elevated, bearing a median row of very fine granules and short setae; interspace 2 flat, as wide as discal width, very weakly impressed below level of 1 and 3; interspace 3 very weakly elevated, equal in height to 1, bearing a median row of fine granules and long setae, the granules slightly larger than those in 1, and the setae much longer; remaining alter- nate interspaces bear long setae on posterior third (or less) of elytra; punctures in striae 1 and 2 distinct, smaller than those on disc. Male.— Frons convex, narrowly, trans- versely impressed just above epistoma; this impression divided by a median, longitudi- nal, laterally flattened, small tooth; surface of frons rather densely pubescent and deep- ly, closely punctured, the punctures smaller and finer in transverse impression. Pro- notum as in female except asperities and serrations larger. Elytra essentially as in fe- male. Type material.— The holotype ( ? ), allo- type, and nine paratypes were collected 6 miles (10 km) south of Carapan, Michoacon, Mexico on 18 June 1965 from Pinus sp., by S. L. Wood. Five additional paratypes were collected 3 miles (5 km) west of El Salto, Durango, Mexico, on 17 June 1965 from twigs of Pintis ayacahidte by S. L. Wood, and two paratypes were collected by R. Coronado P. at Tequesquinahua, Mexico, Mexico, on 2 March 1962, from twigs of a Pinus sp. The holotype, allotype, and most of the paratypes are in the SLWC. A pair of para- types is in the CNC (No. 15480). Remarks.— Adults of this species resemble closely those of P. segnis Blackman and P. subopacus Blackman. The adults of P. im- pexus differ from the latter species by the more flattened, more weakly impressed elytral declivity, especially between the third interspaces of each elytron, by the more densely pubescent male frons and the more sharply elevated longitudinal carina and by the more extensive pubescent area on the female frons. Pitijophthorus intentus, n. sp. Length 1.5-1.8 mm, about 2.8 times long- er than wide. Female.— Frons weakly convex, flattened or weakly, transversely impressed from epis- toma to near upper level of eyes, with a weak, longitudinal carina which extends from the epistoma to upper level of eyes, ending at a weakly elevated, impunctate, median elevation; surface deeply, densely punctured, the punctures rather large and close, almost touching; vestiture sparse, scattered and inconspicuous. Antennal club small, oval, about 1.4 times longer than wide; sutures 1 and 2 straight; segments 1 and 2 occupy more than half the total club length. Pronotum 1.1 times longer than wide; sides nearly straight, parallel on basal half; anterior margin broadly rounded, bearing about a dozen very low, basally contiguous serrations; asperities on anterior slope erect, low, generally isolated and arranged in no apparent order; summit distinct; posterior portion finely punctured, the punctures shallow, small, and usually separated by dis- tances greater than their diameters; inter- puncture space brightly shining, generally smooth but usually bearing a few very min- ute points or lines; median line narrow, not clearly evident. Elytra about 1.9 times longer than wide; apex strongly acuminate; striae punctured in regular rows, the punctures much larger than those on posterior portion of pro- notum, deeply impressed; discal interspaces about as wide as striae or slightly narrower, surface moderately shining, with scattered fine points and lines. Declivity bisulcate; in- terspace 1 narrowly, strongly elevated, equal in height to 3, bearing a median row of about four widely separated, acute gran- ules and erect, stout setae; interspace 2 deeply sulcate, distinctly wider than discal width, smooth and shining; interspace 3 dis- tinctly elevated and bearing a median row of about four widely separated, acute gran- ules, these equal in size to those on inter- 78 Great Basin Naturalist Vol. 38, No. 1 space 1; punctures in striae 1 and 2 obso- lete. Male.— Frons almost identical to female except median transverse elevation stronger, longitudinal carina slightly more evident and punctures on surface slightly stronger. Pronotum and elytra as in female except elytral apex less strongly acuminate. Decliv- ity essentially as in female except interspace 2 slightly less deeply impressed and inter- strial setae stouter. Type material.— The holotype ( ? ), allo- type, and 11 paratypes were collected in Bear Canyon, Santa Catalina Mountains, Arizona, on 15 August 1968, from twigs of Pinus ponderosa, by D. E. Bright (CNC No. 15481). Five additional paratypes are from the Santa Rita Mountains, Santa Cruz Co., Arizona, collected on 29 July 1968, from Pinus strobiformis (1) and Pseudotsuga men- ziesii (4), by D. E. Bright. The holotype, allotype, and most of the paratypes are in the CNC; additional para- types are in the SLWC and the KESC. Remarks.— Adults of P. intentus resemble those of P. consimilis LeConte but differ in a number of points. On adults of P. in- tentus, the second declivital interspace is more deeply sulcate, the granules on decliv- ital interspaces 1 and 3 are much larger, the punctures on the frons of both sexes are larger and deeper, and the longitudinal ca- rina and the transverse elevation are much more prominent. In addition, P. consimilis occurs in conifers in eastern North America, and P. intentus occurs in conifers in the southwestern United States. Pityophthorus laticeps, n. sp. Length 2.5 mm about 2.9 times longer than wide. Male.— Frons flattened from epistoma to well above eyes, very weakly impressed in median portion, divided by a proiminent but weakly elevated longitudinal carina which extends from epistomal margin to well above upper level of eyes; surface moderately shining, rather strongly punc- tured, the punctures distinctly impressed, close, separated by a distance equal to or less than their diameters; vestiture short but conspicuous and abundant. Antennal club oval, 1.3 times longer than wide; sutures 1 and 2 very weakly arcuate; segments 1 and 2 occupy more than half the total club length. Pronotum 1.1 times longer than wide, widest at about middle; sides weakly ar- cuate; anterior margin broadly rounded, with about 10 contiguous, rather large ser- rations; asperities on anterior slope erect, rather large, isolated, scattered in no readily apparent order; summit distinct; posterior area distinctly punctured, the punctures of moderate size, usually separated by a dis- tance more than their own diameters; inter- puncture space moderately shining, with nu- merous very fine micropoints; median line broad, impunctate. Elytra 1.6 to 1.7 times longer than wide; apex broadly rounded; discal striae punc- tured in regular rows, the punctures large, almost touching, deeply impressed; discal interspaces about 2.0 times wider than striae, surface brightly shining, densely mi- cropunctate and subrugulose, interspaces 1, 3, 5, 7, and 9 each bearing on disc a me- dian row of about 3 to 4 larger setose punc- tures, these punctures as large and deep as those in striae, the setae arising from these pimctures rather long and stout. Declivity broadly flattened between the fourth inter- spaces; interspace 1 rather deeply impressed below level of 3 on upper half, slightly ele- vated above surface at apex, bearing a me- dian row of very fine granules and short setae; interspace 2 broadly sulcate, slightly wider than discal width, bearing 2 or 3 very small, acute granules at top; interspace 3 weakly elevated on upper half, bearing about 5 rounded granules on the upper por- tion and 1 or 2 more at apex, each granule bearing a moderately long, stout seta; inter- spaces 4 and 5 smooth, bearing a median row of fine setae; punctures of striae 1 large, not impressed, those in striae 2 much smaller in median area of declivital face than those on disc, those at apex nearly as large as those on disc. Female.— Not definitely recognized in material at hand. A female bearing the identical data and apparently associated with the male has the following character- istics: Frons broadly flattened on a large semicircular area, densely pubescent, with March 1978 Bright: American Scolytidae 79 the peripheral setae longer and incurved. Pronotum essentially as in male holotype except interpuncture spaces on posterior portion are more densely micro- punctate /reticulate and the large punctures are larger and deeper. Elytra essentially as in male except strial punctures slightly smaller and shallower. Declivity completely different from male, convex, not flattened; interspace 1 slightly impressed below level of 3, bearing a median row of fine granules; interspace 2 weakly, not broadly sulcate, only slightly wider than discal width; inter- space 3 weakly elevated, distinctly higher than 1, bearing a median row of moderately large, rounded, setose granules; strial punc- tures as in male holotype. Type material.— The holotype ( $ ) was collected 20.5 km. north of Oaxaca, Oaxaca, Mexico, at an elevation of 9000 feet, on 6 June 1971, from Pinits sp., by D. E. Bright (CNC No. 15482). The female specimen de- scribed above bears the same data but is not designated as an allotype. Remarks.— Males of this species are read- ily distinguished by the broadly flattened elytral declivity in which interspace 3 is near the middle of the declivital face and is granulate only on the upper half and again at the apex, and by the finely punctured posterior half of the pronotum which, in the interpuncture spaces, bears dense, fine mi- cropoints. The female associated with the male specimen resembles the male in several fea- tures but I have doubts that it is the actual female of this species. Until more specimens are available, the female must remain doubtful. Pityophthorus malleatus, n. sp. Length 1.9-2.2 mm, about 2.9 times long- er than wide. Female.— Frons flattened from epistoma to above upper level of eye and laterally occupying about 73 percent of distance be- tween eyes; surface of flattened area densely, moderately, and somewhat roughly punctured except on a small, median area just above epistomal margin, and clothed with sparse, moderately long, fine setae, those on periphery of flattened area slightly longer and incurved; surface lateral to and above flattened area smooth, with large, deep, widely separated punctures. Antennal club oval, 1.4 times longer than wide; su- tures 1 and 2 transverse; segments 1 and 2 occupy slightly more than half the total club length. Pronotum about 1.2 times longer than wide; widest just before level of summit; sides straight, parallel on basal half; anterior margin broadly rounded, bearing about 10 or more, broad, prominent, basally con- tiguous serrations; asperities on anterior slope rather large and distinct, scattered in no apparent order; summit distinct; posteri- or portion moderately strongly punctured, the punctures rather large, deep and usually separated by a distance greater than their diameters; interpuncture space brightly shining, smooth, with numerous, minute points. Elytra about 1.7 times longer than wide; apex broadly rounded; striae punctured in even, regular rows, the punctures quite large and deeply impressed; discal inter- spaces about 2.0 times wider than striae, surface brightly shining, smooth, impunctate and usually with numerous, scattered, very minute points. Declivity steep and weakly, broadly bisulcate; interspace 1 moderately elevated, surface more opaque than on disc, bearing a median row of about four widely separated granules; interspace 2 moderately sulcate, distinctly wider than discal width, surface dull, opaque, marked with numerous fine lines; interspace 3 moderately elevated, about equal in height to 1, bearing about four or more widely separated granules, these granules about equal in size to those on interspace 1; punctures in striae 1 and 2 obsolete; setae on interspaces 1 and 3 slen- der, moderately long. Male.— Frons shallowly, transversely im- pressed, upper margin of impression bearing a distinct, arcuate, transverse carina; surface below carina densely, deeply punctured ex- cept on a very weakly indicated, longitudi- nal carina; surface above transverse carina more deeply punctured. Antennal club as in female. Pronotum and elytra essentially as in female except sculpturing slightly strong- er. Declivity as in female except interspace 2 more deeply impressed, interspace 3 more 80 Great Basin Naturalist Vol. 38, No. 1 abruptly elevated, granules on interspaces 1 and 3 slightly larger and setae on inter- spaces 1 and 3 stouter. Type material.— The holotype ( ? ), allo- type, and 6 paratypes were collected at Walker, Yavapai County, Arizona, on 23 August 1968, from twigs of Pseiidotsuga menziesii, by D. E. Bright (CNC No. 15483). The holotype, allotype, and four para- types are in the CNC and two paratypes are in the SLWC. Remarks.— Adults of this species closely resemble those of P. pseudotsiigae Swaine, but the elytral interspaces of P. maUeatus are completely impunctate while in P. pseu- dotsiigae they bear at least a few setose punctures. In addition, the male frons of P. maUeatus is less deeply impressed and the setae on the female frons are much sparser and shorter. In declivital characters, adults of P. mal- leatus also resemble those of P. intextus Swaine and P. cascoensis Blackman. Fe- males and males of P. maUeatus differ in the characteristics of the frons and by the impunctate elytral interspaces. Pityophthorus mesembria, n. sp. Length 2.0-2.2 mm, about 2.8 times long- er than wide. Female.— Frons flattened on a large semicircular area extending from epistomal margin to well above eyes and laterally oc- cupying over 80 percent of distance be- tween eyes; surface shining, moderately densely punctured, largely concealed by the long, incurved setae which extend from per- iphery of flattened area almost to epistomal margin, setae on remainder of flattened area very short, erect. Antennal club oval, about 1.3 times longer than wide, widest through segment 2; sutures 1 and 2 transverse, rather heavily chitinized at lateral margins; segments 1 and 2 occupy more than half the total club length. Pronotum 1.1 times longer than wide, widest on posterior half; sides moderately to rather strongly arcuate; anterior margin nar- rowly rounded, bearing 10 or more erect, basally contiguous serrations of moderate size; asperities on anterior slope erect, usu- ally isolated but some may be basally con- tiguous, scattered in no apparent order; summit prominent; transverse impression behind summit distinctly impressed, divided bv the distinct, narrowly elevated median line; posterior area strongly punctured, the punctures of moderate size, deeply im- pressed, separated by a distance equal to or less than their diameters; interpuncture space brightly shining, with numerous fine points. Elytra 1.6-1.7 times longer than wide; apex moderately acuminate; discal striae punctured in regular rows, the punctures slightly larger than those on posterior por- tion of pronotum and more deeply im- pressed, each puncture bearing a very short, recumbent seta; discal interspaces 1.5-2.0 times wider than striae, surface shining, im- punctate, smooth with numerous very fine lines and/or points. Declivity shallowly bi- sulcate; interspace 1 weakly elevated, bear- ing a median row of 10 or more small, rounded granules, each of these with a longer, erect, moderately stout seta arising from posterior margin; interspace 2 wider than its discal width, shallowly impressed, surface shining, with numerous, scattered, fine points and lines; interspaces 3 slightly elevated, higher than 1, bearing a median row of 8-10 fine, rounded granules, each of these with a stout, erect seta as those on in- terspace 1; punctures in striae 1 and 2 vis- ible but smaller and less deeply impressed than those on disc. Male.— Frons flattened on a semicircular area as in female, punctures on this area very large, deep and close; a weakly ele- vated median, longitudinal carina extends from epistoma to upper level of eyes; vesti- ture moderately long, sparse, and relatively inconspicuous. Pronotum as in female ex- cept asperities larger and the points in the interpvmcture spaces on the posterior por- tion of the pronotum deeper and more con- spicuous, resulting in the large punctures becoming more obscure. Elytra as in female except strial punctures deeper. Declivity as in female except granules on interspaces 1 and 3 larger. Type material.— The holotype ( ? ), allo- type, and five paratypes were collected on Cerro Calel, Quezaltenango, Guatemala, at March 1978 Bright: American Scolytidae 81 an elevation of 10,000 feet, on 26 May 1964, from Abies guatemalensis, by S. L. Wood. The holotype, allotype, and three para- types are in the SLWC; one pair of para- types is in the CNC (No. 15484). Remarks.— Adults of this species are dis- tinguished from other species in the confer- tus group by the arcuate pronotal sides, by the densely punctured interpuncture space on the posterior portion of the pronotum, by the presence of numerous granules on declivital interspaces 1 and 3, by the host, and by the distribution. Pityophthorus montezumae, n. sp. Length 2.5-2.9 mm, about 2.5-2.6 times longer than wide. Female.— Frons flattened on a broad area extending from the epistoma to well above upper eye level and laterally nearly from eye to eye, central portion of flattened area frequently concave; surface moderately shining, densely, finely punctured and marked with numerous fine lines and points; vestiture abundant, those setae in central area short and erect, those on periphery much longer and incurved. Antennal club very large, broad, about 1.1 times longer than wide, widest through segment 3; su- tures 1 and 2 distinctly arcuate; segment 1 small, much narrower than 2 or 3; segments 1 and 2 together occupy less than half the total club length. Pronotum as long as wide, widest on pos- terior one-quarter; sides weakly arcuate to weakly converging; anterior margin broadly rounded, bearing about 10 low, contiguous serrations; asperities on anterior slope nu- merous, low, broad, densely scattered in no apparent order; summit distinct; transverse impression behind summit moderately im- pressed; posterior area densely punctured, the punctures moderately deeply impressed and separated by a distance equal to slightly less than their diameters, the lateral margins of some punctures weakly elevated, giving a weakly, subasperate appearance to the surface, especially on lateral areas; in- terpuncture space dull, finely, minutely reti- culate; median line broad, not elevated, im- punctate. Elytra 1.6-1.7 times longer than wide; apex broadly rounded; discal striae punc- tured in regular rows, the punctures very large, moderately deep and separated by a distance equal to less than half their diame- ters; discal interspaces narrower than or equal in width to striae, surface moderately dull, densely, minutely reticulate, each in- terspace with a median row of large punc- tures equal in size to those in striae, each of these bearing a moderately long, erect seta, the setae longer on posterior portions of each interspace. Declivity convex, moder- ately bisulcate; interspace 1 moderately ele- vated, equal in height to 3 or very slightly lower, bearing a median row of about six widely separated, fine granules, each gran- ule with a long, erect seta arising from lower edge; interspace 2 moderately sulcate, equal in width or slightly wider than discal width, surface as on disc; interspace 3 weakly elevated, bearing a median row of about six or less small granules, these slightly larger than those on interspace 1 and each bearing a long erect seta on lower edge; punctures of striae 1 and 2 obsolete. Male.— Frons flattened to very weakly impressed on each side of a sharply ele- vated, longitudinal carina extending from epistomal margin to upper level of eyes, this carina more strongly elevated on lower half and sometimes nearly toothlike; surface on each side of carina dull, densely punc- tured, and with numerous moderately long setae. Pronotum and elytra essentially as in female. Declivity as in female except gran- ules on interspaces 1 and 3 slightly larger. Type material.— The holotype ( ? ), allo- type, and 22 paratypes were collected 7 miles (11.5 km) east of San Cristobal (de las Casas), Chiapas, on 13 May 1969, from twigs of Pinus montezumae, by D. E. Bright (CNC No. 15485). One additional paratype is from 4 miles (6.5 km) east of San Cristob- al, Chiapas, collected on 26 May 1969, from Pinus sp., by D. E. Bright. The holotype, allotype, and most of the paratypes are in the CNC. Additional para- types are in the SLWC and the KESC. Remarks.— Adults of this species show many of the characteristics of the blandus group except for the remarkable antennal club and the more distinctly elevated carina 82 Great Basin Naturalist Vol. 38, No. 1 on the male frons. Adults are easily recog- nized by the very large, broad, antennal club with the narrow first segment and the strongly arcuate sutures, by the sharply ele- vated but not toothlike carina on the male frons, by the first and third declivital inter- spaces being nearly equal in height, and by the distribution. This is the only species in this group that occurs south of the Isthmus of Tehuantepec. Pityophthorus scalptus, n. sp. Length 2.6-3.0 mm, about 2.7 times long- er than wide. Female.— Frons broadly flattened from eye to eye and from epistoma to well above eyes, slightly but distinctly concave on an almost circular, broad, median area; surface moderately shining, with close, deep, mod- erately large punctures, and bearing short, erect setae over most of the area, the setae on the periphery being much longer, coar- ser, and incurved. Antennal club oval, about 1.3 times longer than wide, widest through segment 3; suture 1 nearly transverse, 2 weakly arcuate; segments 1 and 2 occupy less than half the total club length. Pronotum as long as wide, widest on pos- terior half; sides moderately arcuate; ante- rior margin broadly rounded, with about 10 prominent, basally contiguous serrations, the median pair usually quite large and erect; asperities on anterior slope generally large and erect, usually isolated but several may be basally contiguous, scattered in no ap- parent order; summit distinct; posterior area deeply, closely punctured, the punctures of moderate size, separted by a distance about equal to their diameters, the lateral margin of each puncture elevated into a distinct granule, giving a definite granulate appear- ance to the posterior surface, these granules usually larger and more distinct posteri- orally and laterally to the summit; inter- puncture space moderately shining to dull, bearing minute points or minute reti- culations; median line broad, surface smooth, impunctate and usually shining. Elytra 1.5-1.6 times longer than wide; apex broadly rounded; discal striae punc- tured in regular rows, the punctures small, deep and close, each bearing a minute. erect seta; discal interspaces 2-3 times wider than striae, surface moderately shin- ing, microrugulose or smooth, with numer- ous fine points and microreticulation; inter- spaces 1, 3, 5, and 7 each with a few scattered, median punctures, each of which bears a minute, erect seta, each seta equal in length to those in striae. Declivity broad- ly convex, weakly impressed; interspace 1 narrowly elevated, weakly impressed below level of 3, bearing a median row of close, fine granules; interspace 2 flat, about as wide as on disc, surface as on disc; inter- space 3 not elevated, bearing a median row of extremely fine granules; interspace 9 weakly elevated, joining interspace 3 near elytral apex; punctures in striae 1 and 2 dis- tinct, much smaller than those on disc, striae 1 moderately, narrowly impressed. Male.— Frons broadly, shallowly im- pressed below upper level of eyes, with a distinct, longitudinal carina extending from the epistoma to above upper eye level; sur- face moderately shining, densely, closely punctured. Antennal club narrower than in female. Pronotum and elytra essentially as in female. Declivity essentially as in female except interspace 3 very slightly elevated, granules in interspaces 1 and 3 slightly larger, and junction of interspace 3 and 9 slightly more distinct. Type material.— The holotype ( $ ), allo- type, and 142 paratypes were collected at Aspen Grove, British Columbia, from Pinus ponderosa, by H. Richmond on various dates during July 1931 (CNC No. 15486). Two additional paratypes were collected at Estes Park, Colorado, on 31 August 1938 from Pinus ponderosa (Hopk. US 31541-V). Four additional specimens, not designated as paratypes, are from Stevensville, Mon- tana, collected on 31 August 1967, from Pinus ponderosa, by R. McEwan and M. McGregor (Hopk. US 51945) (USNM). The holotype, allotype, and 10 paratypes are in the CNC, 10 paratypes are in the SLWC, two are in the KESC, two in the USNM, and the remainder are in the CASC and in the University of British Columbia, Vancouver. Remarks.— This species is very closely re- lated to P. scalptor Blackman. The most ob- vious and easily seen character which will March 1978 Bright: American Scolytidae 83 distinguish the two species is the presence of prominent granules on the posterior por- tion of the pronotum of adults of P. scalptus vs. the absence of granules on adults of P. scalptor. Other less obvious dif- ferences that distinguish adults of P. scalptus are the larger and deeper punc- tures on the female frons, the slightly lower carina on the male frons, and the slightly smaller size of the antennal club. Pityophthoriis trepidiis, n. sp. Length 1.6-2.0 mm, about 2.7 times long- er than wide. Female.— Frons generally flattened from well above eyes to epistomal margin and from eye to eye, very weakly transversely impressed at midlevel; sometimes a very weak, longitudinal carina is present; surface densely, closely punctured, the punctures deep, distinct, separated by a distance equal to less than their diameters, surface between punctures smooth, shining; vestiture scat- tered, consisting of moderately long, erect setae. Antennal club oval, about 1.4 times longer than wide, widest through segment 2; sutures 1 and 2 arcuate, 1 slightly more than 2; segments 1 and 2 occupy more than half the total club length. Pronotum about as long as wide, widest just behind middle; sides broadly arcuate; anterior margin broadly rounded, bearing about a dozen, basally contiguous, erect ser- rations; asperities on anterior slope erect, of moderate size, scattered in no apparent or- der; summit distinctly elevated, transverse impression behind summit weak; posterior portion deeply, densely punctured, the punctures rather large, separated by dis- tances equal to less than their diameters; in- terpuncture space brightly shining, smooth. Elytra about 1.8 times longer than wide; apex broadly rounded; discal striae punc- tured in regular rows, the punctures shallow and indistinct, somewhat obscured by the microrugulose sculpturing of surface, each pimcture bearing a very short, erect seta; discal interspaces about 2.0 times wider than striae, surface roughened by micro- rugulose sculpturing, interspaces 1, 3, 5, 7, etc., each bearing a few, scattered, median punctures, each of which bears a very short, erect seta, these setae equal in length to those arising from strial punctures or only slightly longer. Declivity convex, weakly impressed along suture; interspace 1 nar- row, bearing a median row of extremely minute granules; interspace 2 wider than discal width, otherwise as on disc; inter- space 3 essentially as on disc, bearing a few, very fine granules; punctures in striae 1 and 2 obsolete, striae 1 narrowly, weakly im- pressed. Male.— Almost identical to female except longitudinal carina on frons may be more evident. Distinguishable with certainty only by abdominal segmentation. Type material.— The holotype ( ? ), allo- type, and nine paratypes were collected at Ukiah, California, from Piniis radiata, under Hopkins No. 20976. No date or collector is indicated on the specimens. The holotype, allotype, and five para- types are in the USNM. Two paratypes each are in the CNC (No. 15487) and the SLWC. Remarks.— This species is closely related to P. keeni (Blackman), but the adults of P. trepidus can be distinguished by the very short interstrial setae (equal in length to the strial setae) and by striae 1 being weakly impressed on the declivity. The host rela- tionships also are different, because P. keeni is found in various species of pinyon pines in the west and P. trepidus is known from Pinus radiata, a tree restricted to the coast- al and island forests of central southern Cal- ifornia. Pityophthorus vespertinus, n. sp. Length 1.9-2.0 mm, 2.4-2.5 times longer than wide. Female.— Frons weakly flattened on a semicircular area extending from epistoma to slightly above upper level of eyes; sur- face shining, densely, minutely punctured, and clothed with erect, moderately long, yellowish setae. Antennal club nearly circu- lar, widest through segment 3; segment 1 distinctly narrower than others; sutures 1 and 2 arcuate, 2 more strongly so; segments 1 and 2 occupy slightly less than half the total club length. Pronotum about 1.1 times longer than 84 Great Basin Naturalist Vol. 38, No. 1 wide, widest at posterior angles; sides broadly, weakly arcuate; anterior margin broadly rounded, bearing about 10 small to large serrations, these very low and broad laterally, gradually increasing in size toward middle; asperities on anterior slope small, erect, scattered in no apparent order; sum- mit prominent; posterior area bearing large, shallow, close punctures, these separated by a distance equal to or slightly less than their diameters; interpuncture space dull, densely, minutely reticulate; median line impunctate, weakly elevated. Elytra about 1.6 times longer than wide; apex broadly rounded; discal striae punc- tured in fairly regular rows, the punctures rather large, deep and close; discal inter- spaces about 1.5-2.0 times wider than striae, generally impunctate but sometimes bearing a few, widely separated punctures, surface opaque, densely, minutely reticulate. Declivity convex, very weakly impressed; interspace 1 weakly elevated, with a me- dian row of fine granules; interspace 2 as wide as discal width, weakly impressed; in- terspace 3 weakly elevated, about equal in height to 1, bearing a median row of fine granules; punctures in striae 1 and 2 dis- tinct, nearly equal in size and depth to those on disc. Male.— Frons weakly concave or flat- tened on each side of a distinctly elevated, longitudinal carina which extends from epis- tomal margin to near upper level of eyes; surface on each side of carina densely punc- tured, the punctures large and shallow. Oth- erwise similar to female. Type material.— The holotype ( ? ), allo- type, and two paratypes were collected 23 miles (35 km) west of Durango, Durango, Mexico, 6000 feet in elevation, on 4 June 1965, from twigs of Pinus sp., by S. L. Wood. The holotype, allotype, and one paratype are in the SLWC; one paratype is in the CNC (No. 15488). Remarks.— This species is very closely re- lated to P. diglyphiis Blandford and may eventually prove to be a subspecies or may even be the same species. This species is only known from Durango, Mexico, and P. diglyphus is known from quite a few local- ities in southern Mexico and Guatemala. Be- cause of the wide geographical separation and the morphological differences, I have described P. vespertiniis as a distinct spe- cies. Adults of P. vespertiniis are distinguished from those of P. diglyphus by the less strongly elevated carina on the male frons, by the less strongly elevated third and ninth elytral interspaces, by the nearly circular antennal club, and by the more northerly distribution. Literature Cited Blackman, M. W. 1928. The genus Pityophthorus Eichh. in North America: a revisional study of the Pithyophthorini, with descriptions of two new general and seventy-one new species. Bull. New York St. Coll. Forestry, Tech Bull. 25, 159 pp. Blandford, W. F. H. 1895-1907. Fam. Scolytidae. In: Biologia Centrali-Americana 4(6): 81-298. Bright, D. E. 1977. New synonymy, new com- binations, and new species of North American Pityophthorus (Coleoptera: Scolytidae). Part I. Canadian Ent. 109: 511-532. EiCHHOFF, W. J. 1869. Neue Borkenkafer. Berl. ent. Zeitschr. 12: 273-282. 1872. Neue exotische Tomiciden-Arten. Berl. ent. Zeitschr. 15: 131-136. 1878. Ratio, description, emandatio, eorum tomicinonim. Preprint of Mem. Soc. Roy. Sci. Liege, Ser. 2, Vol. 8, 531 pp. EiCHHOFF, W. J. AND E. A. ScHWARZ. 1895. Remarks on the synonymy of some North American scolytid beetles. Proc. U.S. Natn. Mus. 18(1085):605-610. Hopkins, A. D. 1905. Notes on some Mexican Scoly- tidae, with descriptions of some new species. Preprint of Proc. Ent. Soc. Wash. 7(2-3):71-81. FLUCTUATION OF A MARSH HABITAT AND THE REPRODUCTIVE STRATEGY OF THE YELLOW-HEADED BLACKBIRD Roger J. Lederer' Abstract.— The Yellow-headed Blackbird {Xanthocephahis xanthocephalus) nests in marshes and is dependent on emergent vegetation for nest sites. Fluctuating water levels from year to year cau.se an increase or decrease in the amount of emergent vegetation and affect the time required for the vegetation to become suitable for nest- ing. Nests built in marshes are very susceptible to wind, rainfall, vegetation growth, and predation. Many nests are abandoned before being used and the mortality of eggs and young is high. The reproductive strategy of the Yellow-headed Blackbird has been selected for flexibility and opportunism to compensate for the unpredictability of the marsh situation. Fluctuations in the physical environment can often cause dramatic changes in the lo- cal population sizes or reproductive strate- gies of organisms. The more potential the environment has for short-term change, the more flexible the resident-organisms must be. The marsh-nesting Yellow-headed Black- bird (Xanthocephalus xanthocephalus), whose breeding ecology and behavior has been extensively studied by Willson and Orians (1963), Willson (1966), and Orians and Christman (1968), is an organism that clearly demonstrates this flexibility. This study describes the fluctuation in some re- productive characteristics of a population of Yellow-headed Blackbirds in a marsh habitat over a six-year period and expands an ear- lier report (Lederer et al. 1975). tag, and the number of eggs and /or nest- lings in it was recorded. Observations of the movements of the adult males and aggres- sive encounters between males were used to determine territorial boundaries. The size of the female population was estimated from the number of active nests and counting of adults. At the peak of the breeding period the surface area and depth of the marsh were measured. Weekly measurements of emergent vegetation height and density were made in several areas around the marsh. Relationships between variables listed in Table 1 (Dimensions of marsh habitat) and Table 2 (Population and nesting data) were tested with linear regression or linear corre- lation analysis. Study Area and Methods The study area is a marsh 500 m north of the Eagle Lake Field Station on the eastern shore of Eagle Lake, Lassen Co., California. The marsh is dependent on Eagle Lake for its water; a rising water table formed the marsh in 1968 (R. Ediger, pers. comm.). This study was conducted from mid-May to late July from 1972 to 1977, inclusive. The marsh was censused for adult birds, nests, eggs, and young at least five times per week by walking about transects 25 m apart. Each nest was plotted on a map of the study area, marked with a numbered Results Physical dimensions of the marsh.— Table 1 shows that the marsh increased in size and average depth each year from 1972 through 1975, but decreased in both dimen- sions in 1976 and 1977. Adult population and territory SIZE.— Over the six years both the adult population and the number of territories showed an increase. Territory sizes varied. The only significant (p<.05) relationship found was a positive relationship between the surface area of the marsh and territory 'Department of Biological Sciences, California State Universiy, Chico, California 95929. 85 86 Great Basin Naturalist Vol. 38, No. 1 The sex ratio was very consistent over the years, being 2.95 to 3.10 females to males (x = 3.03 ± .07). Nests, clutch sizes, and fledglings.— Active nests (many built were never used) produced a low mean of .03 fledged young per nest in 1977 to 1.70 in 1974. There is a significant (p<.01) negative correlation be- tween the number of active nests and the average number fledged per active nest. There is also a significant (p<.05) negative correlation between the total number of nests and the average number of young fledged per active nest; and the ratio of ac- tive/total nests is significantly (p<.005) ne- gatively correlated with the average num- ber fledged per active nest. If the number of young fledged per active nest is the cri- terion of success, then 1974 was the most successful year. Size of marsh and fledging success.— There is a significant (p<.05) positive cor- relation between the surface area of the marsh and the number of fledglings per adult. Mortality.— Known causes of mortality of eggs and young were abandonment, toppling into water, and predation by the raccoons {Procyon lotor) and coyote {Canis kitrans). Weather-related mortality was ap- parent in 1974 when heavy rains occurred. and severe storms in 1975 and 1976 de- stroyed large numbers of nests which con- tained eggs or young. Willson (1966) report- ed similar results. Mortality of fledglings is not known but may be at least 50 percent (Ricklefs 1973). There was extensive rac- coon predation in 1977. Discussion Nest sites.— The number of Yellow- headed Blackbirds nesting in a habitat is strongly influenced by the adequacy of nest sites (Immelmann 1972, Willson 1966). Nests were placed in emergent vegetation about .3 m above the water and encom- passed 8-70 stalks of Scirpus sp. (bulrush) and/or Jiincus sp. (rush). Because Scirpus stalks are thicker and taller than those of Juncus, only 10-20 stalks of Scirpus could support a nest, but 30-50 stalks of Juncus were required. Often, both species were in- corporated into the nest and its support. Scirpus and Juncus grew in shallower water and thus emerged and were available for nesting before Scirpus. But Juncus frequent- ly proved to be too weak or too sparse, and the nests collapsed or were torn apart as the vegetation grew or the water level changed. Ellarson (1950) and Stevens (1937) (In: Willson 1966) reported Yellow-headed Table 1. Dimensions of marsh habitat. 1972 1973 1974 1975 1976 1977 Surface area (hectares) Average depth (meters) 3.9 1.1 4.0 .98 5.3 1.3 13.4 1.6 8.2 2.5 .43 Table 2. Yellow-headed Blackbird population and nesting data. 1972 1973 1974 1975 1976 1977 Adult population Number of territories Minimum-maximum territory sizes (m^) Active nests Empty nests Total nests Ratio of active /total nests Average clutch size Fledged young Averaged fledged per active nest 25 8 not measured 8 0 8 1 3.0 unknown unknown 85 28 40-101 29 67 96 .30 3.0 47 .6 83 28 40-150 19 110 129 .15 2.7 32 1.7 71 24 804-1500 53 58 111 .48 2.6 63 1.2 137 44 20-225 93 66 159 .58 2.8 90 .9 202 66 7-188 290 30 320 .90 2.5 10 .03 March 1978 Lederer: Blackbird Reproduction 87 Blackbird populations decreasing as the wa- ter rose or fell during the season. Later in the season, Juncus was more suitable for nesting. Annual fluctuation of water levels in the marsh affected the time of emergence and density of the emergent vegetation. If the water level was higher than in the previous years (e.g., 1975), the littoral area of marsh had little or no emergent vegetation since it did not have sufficient time to become es- tablished. Most of the emergent vegetation was in the deeper part of the marsh and had to grow longer before it emerged, re- sulting in a later nesting season. Conversely, if the marsh decreased in size from the pre- vious year, much of what was the littoral zone became dry land and no longer sup- ported emergents. A smaller marsh also meant a shallower one, which allowed all the emergent vegetation to emerge sooner and thus provide for an earlier nesting sea- son. Territories.— The changes in territory size suggest that the territories may have changed in quality with respect to nest sites and/or food (Schoener 1968, Stenger 1958, Willson 1966). Territory sizes indirectly regulate the number of nests that are built because each territorial male will have 2-5 females nesting in his territory (Willson 1966). Nesting success.— Nesting success is typ- ically measured as the percent of eggs which produce fledglings (Lack 1966, Rick- lefs 1969). Using this criterion, the fewer the total number of nests built by the Yel- low-headed Blackbird population, the great- er was the success of the active nests. The assumption is that if fewer nests are built, more energy can be devoted to each nest. Also, a low ratio of active nests to total nests is closely correlated to nest success. Thus it appears that the optimum situation for nesting success is a low number of total nests with a low proportion of them being active. But why are so many never-to-be- used nests built? To imply that the birds actively strive to create this set of circumstances would hint at group selection, which I will not invoke as an explanation. An individual male, seek- ing to attract as many females as possible, will attempt to establish as large a territory as it can defend and include in it as much vegetation edge as possible because females tend to be found in territories with more edge (Willson 1966). An individual female will begin to build a nest as soon as the vegetation can support a nest, but this early nesting leads to many abandoned or dam- aged nests. The weather is unstable early in the nesting season; storms and cold weather can damage or destroy the nests, eggs, or young. Or the nest may be made unusable by the growth of the vegetation in which it was built. The result is that many nests are abandoned and the female builds another elsewhere in the marsh. Orian's (1961) study of Red-winged and Tricolored Blackbirds demonstrated a high mortality of eggs and yoimg, plus nest desertion. Based on the av- erage sex ratio and the total number of nests, I estimate that each Yellow-headed Blackbird female builds two nests. The ef- fect is that many more nests are built than are actually used. Although later nests are more likely to be used than earlier ones, they too suffer high losses. Explanations for the negative correlations between numbers of active nests and total numbers of nests with the success of the ac- tive nests are offered here. A lower number of total nests implies that either (1) there were fewer breeding adults or (2) fewer nests were destroyed and replaced by an- other nest, both signifying less stringent breeding conditions. A lower number of ac- tive nests means fewer breeding adults com- peting. The negative correlation between a low ratio of active/total nests and fledging success can be explained by the movement of some breeding adults out of the marsh after their first breeding attempt. After most of the nests were built and incubation was well along in the active nests, the adult population declined. They apparently moved to areas along the edge of Eagle Lake which are considerably deeper (up to 5 m) and do not become suitable for nesting until mid-July, when the vegetation has emerged. Snelling (1968) found a similar drop in Red-winged Blackbird breeding populations. By this time, the marsh vegeta- tion has grown to its maximal height. A sur- 88 Great Basin Naturalist Vol. 38, No. 1 vey of the lake's shoreline revealed that the blackbirds began to build nests at about the time hatching was occurring in the marsh. Physical dimensions of the marsh and MORTALITY.— Predation becomes an impor- tant factor when the water level is low. The vast majority of losses of active nests in 1977 were due to raccoon predation be- cause the average water level (.43 m) was so low that raccoons had easy access to the nesting areas. Although high water levels are disadvantageous in that they delay the nesting season, low water levels can be ex- tremely detrimental by allowing terrestrial predators access. Ricklefs's (1969) work on the nesting mortality of birds states that (1) marsh (and prairie) nesting birds have the highest mor- tality and (2) the major cause of this morta- lity is predation due to nest accessibility. Starvation rates are also higher than other birds because fields and marshes have more variable food supplies. Conclusions The data seem to imply that the repro- ductive strategy of Yellow-headed Black- birds is inefficient in time and energy. But the situation reported here is probably typi- cal, because the average numbers fledged for all nests were similar to that reported by Willson (1966). The fact that the marsh becomes a suit- able nesting habitat at least a month earlier than other nearby areas, inducing the birds to attempt to nest, probably explains the numerous unused nests. Nest, egg, and young mortality were all high, but some adults were able to bring one or two young to fledging. But whether or not the first nesting was successful, the opportunity to renest in the marsh or in a newly opened habitat was still available. Selection would favor those individuals which attempted to breed in an unstable environmental situation because attempting to nest early (with an apparently tenuous commitment) is prob- ably not much riskier than self-maintenance while waiting for other areas to become suitable; the possibility of bringing off one or two young with only a slight additional risk to the individual bird would confer a selective advantage on those individuals. Additionally, since there is no guarantee that sufficient suitable nest sites will devel- op later, there are selection pressures against those individuals who wait, as well as for those which nest early. Acknowledgments I would like to thank G. Brown, K. Bum- gamer, and T. Tam for considerable field assistance. Parts of this study were funded by a National Science Foundation under- graduate research participation grant awarded to the Eagle Lake Field Station of California State University, Chico. Literature Cited Best, L. B. 1977. Territory quality and mating success in the field Sparrow (Spizella pusilla). Condor 79:192-204. Immelman, K. 1972. Role of the environment in repro- duction. In: D. S. Farner (ed). Symposium on Breeding Behavior and Reproductive Physiology in Birds, pp. 121-157. Lack, D. 1966. Population studies of birds. London, Great Britain: Oxford University Press. Lederer, R. J., W. S. Mazen, and P. J. Metropulos. 1975. Population fluctuation in a Yellow-headed Blackbird marsh. West. Birds 6:1-6. Orians, G. H. 1961. The ecology of blackbird (Agelaius) social systems. Ecol. Monogr. 31:285-312. Orians, G. H. and G. M. Christman. 1968. A com- parative study of the behavior of Red-winged, Tricolored, and Yellow-headed Blackbirds. Univ. Calif. Publ. Zool. 84:1-85. RiCKLEFS, R. E. 1972. Fecundity, mortality, and avian demography. In: D. S. Farner (ed.). Symposium on Breeding Behavior and Reproductive Phys- iology in Birds, pp. 366^34. ScHOENER, T. W. 1968. Sizes of feeding territories among birds. Ecology 49:123-141. Snellinc, J. C. 1968. Overlap in feeding habits of Redwinged Blackbirds and Common Crackles nesting in a cattail marsh. Auk 85:560-585. Stenger, J. 1958. Food habits and available food of ov- enbirds in relation to territory size. Auk 75:3,35-346. Willson, M. F. 1966. Breeding ecology of the Yellow- headed Blackbird. Ecol. Monogr. 36:51-77. Willson, M. F. and G. H. Orians. 1963. Comparative ecology of Redwinged and Yellow-headed Black- birds during the breeding season, (abstr.) Proc. XVI. Intemat. Cong. Zool. 3:342-346. I HOSTS OF JUNIPER MISTLETOE AT WALNUT CANYON NATIONAL MONUMENT, ARIZONA Frank G. Hawksworth' and Robert L. Mathiasen^ Abstract.— Four species of junipers, Jiiniperus osteosperma, J. scopulorum, J. monosperma, and /. deppeana, occur in mixed stands at Walnut Canyon National Monument, Arizona. All are parasitized by Phoradendron juni- perinum, but the mistletoe was most common on /. osteosperma and least common on /. scopulorum. Fernbush {Chamaebatiaria millefolium, Rosaceae) was a very rare host. Phoradendron juniperinum Engelm. ex A. Gray parasitizes several species of south- western junipers (Hedgcock 1915), but no information has been published on the rela- tive susceptibility of its various hosts. The Walnut Canyon National Monument Area, southeast of Flagstaff, Arizona, presents a unique area for host studies as four species of junipers occur together in mixed stands: Utah juniper {Jiiniperus osteosperma (Torr.) Little), Rocky Mountain juniper (/. scopulo- rum Sarg.), one-seed juniper (/. monosperma [Engelm.] Sar.), and alligator juniper (/. dep- peana Steud.). Stands in the vicinity of Monument Headquarters, the Rim Trail, and the Island Trail were studied and each juniper over three meters high was examined for mis- tletoe. In all, 282 junipers of the four spe- cies were examined. Mistletoe was most common on Jiiniperus osteosperma as nearly two-thirds of the trees examined were infected. Perhaps this is be- cause /. osteosperma is the largest of four junipers in this area, and birds which dis- seminate the mistletoe tend to prefer taller trees. Jiiniperus scopulorum had the least mistletoe. Infection of /. monosperma was intermediate, and too few /. deppeana trees were present to evaluate its susceptibility in this area. Phoradendron juniperinum was found once on fernbush {Chamaebatiaria millefo- lium (Torr.) Maxim., Rosaceae) near the Monument Headquarters. Fernbush has also been reported to be a very rare host for this mistletoe at Grand Canyon National Park (Hawksworth 1952). Acknowledgment Thanks are expressed to Mr. James Bail- ey, National Park Service, Walnut Canyon National Monument, for assistance in this study. Literature Cited Hawksworth, G. F. 1952. Unusual hosts for two southwestern mistletoes. Hedgcock, G. G. 1915. Some notes on tree diseases in our national forests. Phytopathology 5:175-181. Table 1. Name of this table. Host Trees examined No. Trees with Phoradendron juniperinum Percent /. osteosperma J. deppeana J. monosperma J. scopulorum 78 6 125 73 64 50 44 22 'U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station, Colorado State University, Fort Collins, Col- orado 80521. 'Department of Plant Pathology, University of Arizona, Tucson, Arizona 85721. 89 ALLELOPATHIC EFFECTS OF BUR BUTTERCUP TISSUE ON GERMINATION AND GROWTH OF VARIOUS GRASSES AND FORBS IN VITRO AND IN SOIL' Bruce A. Buchanan,- K. T. Harper,' and Neil C. Frischknecht^ Abstract.— The allelopathic effects of bur buttercup {Ranunculus testiculatus) tissue on selected grasses and forbs varied according to the substratum for germination and growth. The in vitro effects of an aqueous extract of buttercup tissue on germination and root development of five grasses were strongly inhibitory in all cases. However, in soil the effects of buttercup tissue on germination and growth of seven grasses and two dicotyle- donous herbs were small to nonsignificant. Deleterious effects were less severe in fine- as opposed to coarse- textured soils. Under field conditions, the ability of seedlings of the grasses to compete with buttercup varied with the species. Bur buttercup (Ranunculus testiculatus Crantz), a native of southeastern Europe and central Asia (Benson 1948, Davis 1965) was first collected in North America by A. O. Garrett near Salt Lake City, Utah, in 1932. Since that time, the species has spread throughout the Great Basin, the Snake River Plain, and the Columbia Plateau. In much of that area, the species exists in large populations in native vegeta- tion and waste places of the semiarid zone. Small, isolated populations are also known from northeastern Wyoming, northern Ari- zona, northern California, and interior Brit- ish Columbia (Buchanan 1969). The species is a diminutive annual (aver- aging ca. 5 cm in height) that completes its life cycle during the early spring. In north- ern Utah germination occurs in late fall, but at more northerly latitudes, germination may be postponed until spring (Buchanan 1969). Despite its small size, the species of- ten forms a dense carpet over literally thou- sands of acres of dry farm and range land during the period from March to May in the eastern Great Basin. The invasion of bur buttercup into North America prompted an early interest in its potential as a weed. Two memoranda in the files of the Intermountain Forest and Range Experiment Station (IFRES) of the U.S. De- partment of Agriculture (Ogden, Utah) re- port early observations of bur buttercup in Utah and consider its potential as a weed (Clark 1941, and Stewart 1941). The memos, dated 18 July and 3 September 1941, respectively, discuss the plant's his- tory, spread, importance as a weed, and possible control in the West. However, when our study began in 1966, no aspect of the dispersal or ecology of the species had been reported in the scientific literature (ex- clusive of floras) of North America. Because of rapid spread of bur buttercup and the dearth of information concerning the species, the Intermountain Forest and Range Experiment Station initiated a pro- gram to investigate its life history, ecology, and distribution. Concern over possible deleterious effects of bur buttercup on win- ter wheat crops and on range grass seedings prompted this study of the competitive and allelopathic effects of bur buttercup on wheat and on grasses used in range revega- tation. Germination studies using both glass 'Research supported under contract 12-11-204-30 between the Intermountain Forest and Range Experiment Station, Forest Service, U.S. Department of Agriculture, Ogden, Utah, and the University of Utah. 'Department of Biology. University of Utah, .Salt Lake City, Utah 84112. Current address of Buchanan is Department of Agronomy, New Mexico State University, Las Cruces, 88003. Current address of Harper is Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602, 'Intermotmtain Forest and Range Experiment Station, Provo, Utah 84601. 90 March 1978 Buchanan et al.: Allelopathic Effects 91 and soil systems were utilized so that results might be extrapolated to field situations. Methods In vitro studies in petri dishes compared the effect of a leachate from bur buttercup tissue with both distilled water and manni- tol as media for germination and root devel- opment of five grasses. The leachate was prepared by soaking one part of air-dried whole plant tissue at fruiting state in 30 parts distilled water at room temperature for 24 hours. The air- dried tissue was first ground through 40- mesh screen. Following soaking, the leachate was filtered through #2 filter pa- per and stored at 1 C until used. The lea- chate had an osmotic concentration (O.C.) of 1.8 atmospheres and a pH of 5.5. Seeds of crested wheatgrass {Agropyron desertorum), tall wheatgrass (A. elongatum), common rye (Secale cereale), Delmar fall wheat {Tritictim aestivum), and Gaines fall wheat (T. aestivum) were each subjected to 6 treatments as follows: Treat- O.C. of ment Wetting Agent pH of Effect No. Agent (Atm.) Agent Tested 1 Distilled HgO 0.0 7.0 Control (buffered) 2 Distilled HgO 0.0 5.5 pH 3 Mannitol in HgO 1.8 7.0 Osmotic pressure 4 Mannitol in H,0 1.8 5.5 Osmotic 5 Leachate 6 Leachate pressure and pH 1.8 7.0 Leachate (pH adjusted) 1.8 5.5 Leachate Two pieces of filter paper were placed in each petri dish and 25 seeds were posi- tioned thereon in five rows of five seeds each. Each treatment was replicated 10 times. Dishes were initially wetted with 6 ml of treatment solution. Thereafter, dishes were rewatered daily with 2 ml of distilled water for the duration of the experiment. At the end of the third-to-seventh day (depending on species), percent germination was recorded. Seeds were considered germi- nated if a root of 5 mm or longer had emerged from the seed coat. Root length data were taken by randomly selecting two rows (10 seeds) from each petri dish and re- cording length of all roots. In addition to in vitro studies, the inhib- itory effects of bur buttercup were tested on seven grasses and three forbs in potted silty clay soil. This study involved two lev- els of bur buttercup tissue mixed with soil in imdrained pots: average annual field pro- duction of bur buttercup (0.7g/dm2, IX) and twice average annual field production of bur buttercup (2X). The study was de- signed for twelve species, three treatments (control and two levels of tissue), and nine replicates per treatment. Only ten species or varieties were used, but two species (Gaines wheat and crested wheatgrass) ap- pear twice in the design, the second time in sand. The design was a 3 X 3 trimultiple latin square for each species (Cochran and Cox 1957). Sixteen seeds were planted in each pot by pressing seeds through a tem- plate to insure uniform spacing (four seeds per row in four rows). The silty clay soil (15 percent sand, 43 percent silt, and 42 percent clay) used in this experiment was taken from the top 15 cm of a pasture on the Benmore Experi- mental Range not infested with bur butter- cup. The soil was air dried and passed through a 2 mm sieve before potting. Soil pH was 7.5. A layer of sand (about 1 cm deep) was placed over the soil to reduce crusting. A sand (90 percent sand, 5 percent silt, and 5 percent clay), pH 8.0, was like- wise dried, screened, and potted as a second substratum for Gaines wheat and crested wheatgrass. The sand was taken from a stream channel at the Benmore Range. Plastic pots were filled with 625 g of soil (or 955 g of sand) and topped with 80 g of sand. The experiment was conducted in a greenhouse at the University of Utah. Spe- cies and substrata are listed below. Species Bur buttercup Fairway wheatgrass {Agropyron cristatum) Halogeton (Halogeton glomeratus) Alfalfa {Medicago sativa) Delmar wheat {Triticum aestivum) Substratum Soil 92 Great Basin Naturalist Vol. 38, No. 1 Tall wheatgrass (A. elongatum) Crested wheatgrass (A. desertorum) Gaines wheat {T. aestivum) Common rye (Secale cereale) Western wheatgrass (A. smithii) Gaines wheat Crested wheatgrass Sand The grasses are all often sown in areas in- fested with buttercup, halogeton is an in- troduced annual often associated with bur buttercup in the eastern Great Basin, and alfalfa is a perennial frequently seeded in areas supporting dense stands of buttercup. On 13 January, all pots were watered with distilled water to water-holding capac- ity (WHC) and thereafter to WHC when the soil became noticeably dry. Number of seeds germinated was record- ed after 5, 7, 10, 13, 15, 17, 25, and 28 days from the initial wetting of the pots. Beginning on day 34 and continuing through day 40, the species were clipped, oven dried, and weighed. All replications and treatments of a species were harvested on the same day. Another soil system tested the influence of buttercup on yield of each of 9 grasses sown in the same rows with buttercup, in a cold frame at Benmore. On 5 October, 100 seeds of each grass were planted in each of three rows 150 cm long. Rows planted to the same grass were spaced 2 dm apart, while 4 dm separated rows between species. Fifty and 100 bur buttercup seeds were planted with the grass in the second and third rows, respectively. The first row as- signed to each grass species was maintained as a control. Grasses tested were: crested wheatgrass. Fairway wheatgrass, Russian wildrye {Elymiis junceus), pubescent wheat- grass (Agropyron trichophorum), inter- mediate wheatgrass (A. intermedium), west- ern wheatgrass, common rye, Delmar wheat, and Gaines wheat. Rows were watered 5 October, at plant- ing, and again the following day by rain. Thereafter, lids of the cold frame were closed (except for periodic waterings) until early April. All plants were clipped on 27 and 28 April. Each row was clipped in 15 segments, each 1 dm long. Tissue was bagged and taken immediately to the Uni- versity of Utah laboratory for drying. Sam- ples were dried at 32 C and weighed. To reduce edge effect from the walls of the cold frame, clipping segments 1 and 15 were not used for determination of the mean production per segment. Results In vitro studies in petri dishes indicate that a compound(s) inhibitory to germina- tion and growth of the five species or varie- ties tested is produced by bur buttercup tis- sue or the products of its decay (Table 1). Table 1. Effect of six treatments on germination and root development of two grasses and three cereal grains in vitro.' Treatment Control Control Mannitol Leachate pH Growth Buffered Acidified Mannitol Acidified Adjustment Leachate pH —Osmotic Concentration Species Time (days) 7.0-0.0 5.5-0.0 7.0-0.0 5.5-1.8 7.0-1.8 5.5-1.8 Percent Germination Crested wheatgrass 7 83" 92''b 90"b 93b 0^ 0^ Tall wheatgrass 5 85a 87" 81" 82" Ob Ob Common rye 3 72" 72" 71" 73" 7b 9b Delmar fall wheat 4 83? 81" 78" 80" 12b 9b Gaines fall wheat 3 96 96" 94" 9.5" Root Length (mm) 24b 16b Crested wheatgrass 7 29ab 25" 31"b 37b 0^ 0^ Tall wheatgrass 5 26* 22b 19b 22b 0^ 0^ Common rye 3 75' 63" 84" 80" 8b lib Delmar fall wheat 4 62" 57" 66" 67" 9b 7b Gaines fall wheat 3 92"b 76" 86"b lOOb IQc 13^^ "Means for a given species followed by the same letter, in superscript, do not differ at the 1 percent probability level (t-test). March 1978 Buchanan et al.: Allelopathic Effects 93 Acidic distilled water had no significant ef- fect on germination, but tended to decrease root length of all species below control lev- els, although observed differences were not usually significant. At the level tested, os- motic concentration also appeared not to influence germination of these species. The in vivo study in which buttercup tis- sue was applied to soil in pots showed a different response than was obtained in pe- tri dishes (Table 2). In soil, buttercup tissue significantly retarded germination of only one species, crested wheatgrass. This effect was observed only at the 2X concentration of buttercup tissue. In no case was average weight of individual test plants significantly different than that of control plants. In sand, germination of crested wheat- grass and Gaines fall wheat was significantly reduced by both IX and 2X concentration of buttercup tissue (Table 2 and Fig. 1). Av- erage weight of surviving plants of crested Table 2. Effect of two levels of bur buttercup tissue on germination and dry matter production of seven grasses and two forbs planted in soil or sand. Treatments are: C, control (no tissue added); IX, tissue added equivalent to average production of buttercup per unit area in pastures (0.7 g/dm^), and 2X, tissue added equiva- lent to twice average production of buttercup. Species Days after Planting Growth Treat- Average Weight 5 10 15 25 28 Medium ment per Individual (g) Percent Germination Soil C 1 3^ 9 13 12 12 IX 3 8 15 10 10 2X 1 6 13 14 14 Soil C 0.16 0 3 35 65 67 IX 0.23 0 11 45 61 61 2X 0.23 0 3 51 71 71 Soil c 0.36 4 21 56 67 67 IX 0.32 4 36 53 58 58 2X 0.36 3 42 53 60 60 Soil C 0.30 3 44 89 94 94 IX 0.34 2 62 89 90 90 2X 0.34 0 57 80 87 87 Soil C 0.23 3 38 81 87 89a IX 0.27 13 58 76 80 80ab 2X 0.28 1 37 62 65 65b Soil c 0.36 23 41 69 71 71 IX 0.32 21 46 62 64 64 2X 0.36 15 50 71 71 71 Soil c 0.24 8 19 19 19 19 IX 0.32 17 26 27 27 27 2X 0.24 12 24 25 25 25 Soil C 0.38 3 21 36 37 37 IX 0.41 6 34 49 49 49 2X 0.42 14 35 42 42 42 Soil c 0.39 15 44 53 53 54 IX 0.42 27 53 57 57 57 2X 0.43 25 59 62 62 62 Sand c 0.20a 45« 90a 94a 94a 94a IX 0.20« 3b 67b 76b 76a 76a 2X 0.1 lb Ob 24c 39^^ 41b 41b Sand C 3 94a 96a 96a 96a 96a IX 56b 83b 85b 85b 86b 2X 21^^ 56^ 60C 62^ 62<= Halogeton Western wheatgrass Fairway wheatgrass Tall wheatgrass Crested wheatgrass Alfalfa Common rye Delmar fall wheat Gaines fall wheat Crested wheatgrass Gaines fall wheat 'Plants not harvested. 'Sets differing among themselves at the 5 percent probability level are followed by superscripts. Means followed by the same letter do not differ sig nificantly. 'No data. 94 Great Basin Naturalist Vol. 38, No. 1 wheatgrass was also significantly reduced by the 2X treatment on sand (Table 2). Unfortunately bur buttercup seeds failed to germinate in these greenhouse trials. The seeds apparently require some cold treat- ment for germination, since seed from the same lot did germinate in the cold frame experiment reported in Table 3. Buttercup appeared to significantly inhib- it growth of seedlings of fairway and west- em wheatgrass in the cold frame (Table 3). The results offer no evidence as to whether inhibition is caused by simple competition, allelochemic effects, or both. The cereal grains (common rye and the fall wheats) were far more effective than perennial grass seedlings in suppressing but- tercup growth in the cold frame (Table 4). The cereals germinated within two weeks of planting and attained heights of 15 cm or more prior to the onset of severe cold. Thus buttercup seedlings growing with them were heavily shaded throughout the experi- mental period. Buttercup plants growing with these species never produced seed. Buttercup production was highest in rows seeded to fairway and western wheatgrasses and Russian wildrye. Fairway and crested wheatgrass differed markedly in respect to the amount of competition they offered but- tercup. Although fairway yielded far more growth than crested wheatgrass, buttercup yields were over twice as great in rows sown to fairway as opposed to crested wheatgrass. Discussion and Conclusions Klikoff (1964) has postulated that the ef- fect of tissue of one species on germination and growth of others could possibly be re- duced or eliminated in soil by adsorption of reactive materials on colloidal surfaces, by microbial degradation of the material, by leaching, and /or dilution of concentration by diffusion. In this study, the leaching ef- CRESTED WHEATGRASS GAINES FALL WHEAT 25 30 0--KD CONTROL DAYS AFTER SEEDING • ©IX ^ -^ 2X Fig. 1. Effect of three levels of bur buttercup tissue on germination of crested wheatgrass and Gaines fall wheat on Benmore soil and sand. See text for details. March 1978 Buchanan et al.: Allelopathic Effects 95 Table 3. Average dry weight of grasses (g/dm of row) for three densities of associated bur buttercup plants. Species Seeding Density of Buttercup None 50 Seeds 100 Seeds (Control) per 1.5 m Row per 1.5 m Row F-Value Crested wheatgrass Fairway wheatgrass Intermediate wheatgrass Pubescent wheatgrass Western wheatgrass Russian wildrye Common rye Delmar fall wheat Gaines fall wheat 1.94 2.02 2.40 .30 7.52 3.36 4.26 3.99° 8.24 6.99 7.60 .31 9.21 9.74 8.76 .14 1.47 .50 .14 14.16' 1.6,3 1.06 1.31 .54 .1.07 6.81 10.68 9.53° 6.68 4.74 8.13 2.25 9.67 4.76 10.70 4.80" 'Significant at the 5 percent probability level (analysis of variance) feet was minimized by the use of imdrained pots, but the diffusion of material into the greater vohime of pots as opposed to petri dishes could partially account for the re- duced effect of buttercup tissue on germina- tion and growth of test species in sand or soil. Differential effect of the tissue in sand and soil also implies that adsorption and/or microbial decomposition are operative. Greater adsorption would be expected in silty clay soil than in sand which would have considerably less surface area. Further- more, decomposition may well be more rap- id in this soil, since sands tend to support smaller microbial populations than do heav- ier textured soils (Clark 1957). In view of the effect of soil texture on action of an allelopathic agent as observed in this study, it is of interest that del Moral and Cates( 1971: 1033) conclude that the ca- pacity of soil to detoxify allelopathic com- pounds is unpredictable. Their results are perhaps related to the fact that they used a moistened sponge rather than uncon- taminated soil as a control. A comparison of the germination response of our species on filter paper and in soil (Tables 1 and 2) demonstrates that germination of several species was reduced in soil (as opposed to filter paper) even without the addition of allelopathic material. There is thus a clear need to perform additional experiments be- fore soil texture is dismissed as a significant variable affecting the action of allelopathic compounds. The pot trials (Table 2) demonstrate that buttercup tissue in quantities that might be expected under field conditions is capable of reducing germination and retarding growth of at least some species under natu- ral conditions. These tendencies may well be accumulative under conditions in which comparable amounts of tissue are added to the system each growing season. The cold frame studies also demonstrate that butter- cup seedlings severely suppress the growth of some grasses. Fairway and western wheatgrass, two species having growth significantly inhibited when competing with buttercup in the cold frame trials, were both slow germinators. Rapid germination of buttercvip may give this species a competitive advantage for wa- ter, nutrients, and space. Thus, the suppres- sion of these species by buttercup may be related to slow development and growth habit rather than to allelopathic action. The differing performance of buttercup Table 4. Total top growth (grams dry wt) of bur but- tercup seeded at two rates with various grasses. Competing Species Seeding Density of Buttercup 50 Seeds/ 100 Seeds/ 1.5 m Row 1.5 m Row None 50 80 Crested wheatgrass 17 26 Fairway wheatgrass 43 77 Intermediate wheatgrass 27 34 Pubescent wheatgrass 25 40 Western wheatgrass 60 52 Russian wildrye 31 73 Common rye 0.2 0.3 Delmar fall wheat 0.1 0.1 Gaines fall wheat 0.1 0.01 96 Great Basin Naturalist Vol. 38, No. 1 when grown with fairway and crested wheatgrass, two very closely related species, is remarkable. Although crested wheatgrass produced only about one-half as much bio- mass as fairway (Table 3), buttercup pro- duction was threefold greater when grown with the latter species. Such disparity in performance of buttercup merits further study. Although bur buttercup has been shown to display allelopathic effects against several grasses, the effects may be of limited impor- tance under most field conditions. Any chemical effects which the species might exert are expected to be most pronounced against slow-developing species growing on coarse-textured soils. Since maximum abun- dance of bur buttercup occurs on fine- textured soil (Buchanan 1969), adsorption and decomposition of toxic compounds would be expected to be rapid. Certainly no obvious suppression of associated species is seen in the field. Rapid development in late winter and early spring probably con- tributes more to competitive ability of but- tercup than does allelopathy. Literature Cited Buchanan, B. A. 1969. The life history and ecology of bur buttercup, Ranttnculus testictilatus Crantz. Master's thesis. University of Utah, Salt Lake City. 102 p. Clark, F. E. 1957. Living organisms in the soil, p. 157-165. In: Soil. The 1957 Yearbook of Agri- culture, U.S. Department of Agriculture. U.S. Government Printing Office, Washington. 784 P- Clark, I. 1941. Unpublished memorandum. Files of Intmtn. Forest and Range Exp. Sta., Ogden, Utah. Cochran, W. G. and G. M. Cox. 1957. Experimental designs, 2d ed. John Wiley and Sons, Inc., New York. 611 p. Davis, P. H. 1965. Flora of Turkey and the East Ae- gean Islands, Vol. 1. Edinburgh University Press, Robert Cunningham and Sons Ltd., Great Britain. 567 p. DEL Moral, R., and R. G. Gates. 1971. Allelopathic potential of the dominant vegetation of western Wa.shington. Ecology 52:1030-10,37. Holmgren, A. H., and J. L. Reveal. 1966. Checklist of the vascular plants of the Intermountain Re- gion. U.S. Forest Service Research Paper INT- 32. Intmtn. Forest and Range Exp. Sta., Forest Service, U.S. Dept. Agr., Ogden, Utah. 160 p. Klikoff, L. G. 1964. The toxicity of Beta vulgaris fruits as an inhibitor of germination of grass fruits and as an autotoxin. Northw. Sci. 38(2):43-51. Stewart, G. 1941. Unpublished memorandum. Files of Intmtn. Forest and Range Exp. Sta., Ogden, Utah. Benson, L. 1948. A treatise on the North American Ranunculi. Am. Midi. Nat. 40:1-261. NEW SPECIES OF NEARCTIC NEOPERLA (PLECOPTERA: PERLIDAE), WITH NOTES ON THE GENUS Bill P. Stark' and Richard W. Baumann' Abstract.— The Nearctic Neoperla are reviewed. Eight species, placed in two nominal groups, are recognized. Neoperla carlsoni, new species, N. catharae, new species, N. choctaw, new species, N. freytagi, new species, N. gatifini, new species, and A^. stewarti, new species, are described for male, female, and egg. Male holotypes and female allotypes are designated for each species. Neoperla mainensis Banks and N. clymene (Newman) are redes- cribed and mainensis is raised to species. Descriptions are supported by original drawings and stereoscan photo- micrographs. Relationships of Nearctic species groups to the world fauna are discussed. Needham and Claassen (1925) established the concept of Neoperla clymene (Newman) as a widespread, variable species. Frison (1935) accepted this usage, and Hynes (1952) and Ricker (1952) reiterated varia- bility of the species. Recent studies of Ethiopian Neoperla (Zwick 1973a,b) suggest the need for critical examination of Nearctic populations of the genus. Preliminary results, from material at hand, indicate a minimum of eight species in the complex. Descriptions of these are presented herein. Neoperla carlsoni, n. sp. Figs. 1, 2, 3, 4, 5, 65, 66 Neoperla sp. A: Stewart, Stark & Huggins, 1976:378. Male.— Macropterous. Length of fore- wings 5-7 mm; length of body 5-7 mm. Dorsum of head yellow with brown mark- ings on anterior region of frons. Pronotum brown with darker rugosities and sutures, a narrow pale stripe extending along median suture. Tibiae, tarsi, and apical half of fe- mora dark brown. Wings amber, veins brown; coastal margin pale yellow. Process of tergum 7 slender, ventrally spinulose, ex- tending to near posterior margin of tergum 8. Process of tergum 8 abruptly elevated on posterior margin, slightly spinulose. Spinules absent from membrane of tergum 8. Mesal spinule patch of tergum 9 with 6-10 spin- ules, lateral patches absent. Shaft of ae- deagus with external spines arranged in two 'Department of Biology, Mississippi College, Clinton, Mississippi 39058. 'Department of Zoology, Brigham Young University, Prove, Utah 84602. ventral groups; left grouping with 25-35 spines, right grouping with 5-8 spines. Ex- trusible, apical section of aedeagus heavily spiculate and armed basally with scattered spines. Female.— Macropterous. Length of fore- wings 8-10 mm; length of body 8-10 mm. Color pattern similar to male. Subgenital plate slightly produced, posterior margin rounded and slightly notched. Vagina with- out sclerites; spermatheca short and com- pletely lined with golden-brown spinulae in basal half. Egg.— Outline oval, cross section circular. Collar distinctly stalked. Chorion uniformly, finely punctate. Types.— Holotype ( $ ) and allotype ( $ ) from Rocky Comfort Creek, 7 mi S Quincy, Gadsden Co., Florida, 16-1 V- 1967, W. L. & J. G. Peters. Holotype, # 73355, and allo- type deposited at the U.S. National Mu- seum. Paratypes: FLORIDA: Gadsden Co., Rocky Comfort Creek, 10- VI- 1970, W. L. Peters, et al., 3 5 1 $ (EPS); same data, lO-VIII-1967, G. Cooper, et al., 1 S (FAM). Okaloosa Co., Blackwater River, 8- V-1972, W. L. Peters, et al., 1 ? (FAM). Santa Rosa Co., Blackwater River, Riley Landing, 7- VIII- 1971, W. L. Peters, et al., 2 S (FAM). Walton Co., Alaqua Creek, 21- VI-1971, P. H. Carlson, et al., 3 $ (PHC). LOUISIANA: Grant Par., Pollock, 24-VII- 1967, E. A. Cancienne, 1 $ (NTSU). Liv- ingston Par., Little Natalbany River, Al- bany, 27-VI-1973, B. Stark, 15 3? (BPS). 97 Great Basin Naturalist Vol. 38, No. 1 Vernon Par., Fort Polk, 18-VI-1968, L. D. Newson, 2 S (NTSU). OKLAHOMA: McCurtain Co., Battiest, 14-VI-1972, D. C. Arnold, 3 $ (NTSU). SOUTH CAROLINA: Aiken Co., Three Runs Creek, 21-VIM971, J. W. Richardson, 2 ? (BPS). TEXAS: An- derson Co., Salmon, 7-27-VI-1975, H. R. Burke, 8 5 4 $ (TAM) (NTSU). Nacog- doches Co., Nacogdoches, lO-VI-1969, R. Dorman, 1 5 1 $ (NTSU). Etymology.— This species is named in honor of Mr. Paul Carlson who has made available for our study a large number of Plecoptera specimens from the southeastern United States. Neoperla catharae, n. sp. Figs. 6, 7, 8, 9, 64 Male.— Macropterous. Length of fore- wings 5-7 mm; length of body 5-7 mm. Dorsum of head yellow with brown mark- ings on anterior region of frons. Pronotum brown with darker rugosities and sutures. Tibiae, tarsi and apical half of femora dark brown. Wings amber, veins brown; coastal margin pale yellow. Process of tergum 7 slender, ventrally spinulose, extending to near posterior margin of tergum 8. Process of tergum 8 abruptly elevated on posterior margin, slightly spinulose. Spinules absent from membrane of tergum 8. Mesal spinule patch of tergum 9 with 6-10 spinules, later- al patches with a few scattered spinules. Shaft of aedeagus with external spines ar- ranged in two ventral groups; left grouping with 5-12 spines, right grouping with 12-20 spines. Extrusible, apical section of aedeagus heavily spiculate and armed basally with several scattered spines. Female.— Macropterous. Length of fore- wings 8-10 mm; length of body 8-10 mm. Color pattern similar to male. Subgenital plate slightly produced, posterior margin smoothly rounded to slightly notched. Va- gina without sclerites; spermatheca short and completely lined with golden-brown spinulae in basal half. Egg.— Outline oval, cross section circular. Collar distinctly stalked. Chorion smooth. Figs. 1-5. Neoperla carhoni: 1, Male terrninalia, dorsal; 2, Male terga, 7 and 8, lateral; 3, Female sternum 8; 4, Aedeagus, lateral; 5, Aedeagus, ventral. March 1978 Stark, Baumann: Nearctic Stoneflies 99 Types.— Holotype ( $ ), allotype ( ? ), and 11 $ paratypes from James Creek, Ran- dolph Co., Arkansas, IX- 1969, M. Case. Holotype, #73357, and allotype deposited at the U.S. National Museum. Paratypes: ARKANSAS: Washington Co., Cove Creek, 4- VIII- 1962, 4 $ (INHS). OHIO: Butler Co., Oxford, 30- VI- 1902, 1 $ (UU). Warren Co., Todds Fork, Morrow, 15-VIII-1951, A. R. Gaufin, 2 <5 3 $ (UU) (EPS); same data, 5-VIII-1953, 7 ? (UU) (EPS). Etymology.— This species is named for Dr. Mary Case Cather who collected the type series. Neoperla choctaw, n. sp. Figs. 10, 11, 12, 13, 14, 15, 16, 17, 67, 68 Male.— Macropterous. Length of fore- wings 5-7 mm; length of body 5-7 mm. Dorsum of head yellow, frons without brown markings. Pronotum brown with darker rugosities and sutures. Tibiae, tarsi, and femora dark brown with distal yellow bands on tibiae and proximal yellow bands on femora. Wings and veins brown; coastal margin pale yellow. Process of tergum 7 slender, ventrally spinulose, extending to near posterior margin of tergum 8. Process of tergum 8 abruptly elevated on posterior margin, slightly spinulose. Spinules absent from membrane of tergum 8. Mesal spinule patch of tergum 9 with 5-12 spinules, later- al patches with 0-3 spinules. Shaft of ae- deagus with external spines arranged in two ventral groups; left grouping with 5-10 spines, right grouping with 3-7 spines. Ex- trusible, apical section of aedeagus heavily spiculate and armed with several scattered spines. Figs. 6-9. Neoperla catharae: 6, Male terminalia, dorsal; 7, Female sterna 7 and 8; 8, Aedeagus, lateral; 9, Ae- deagus, ventral. 100 Great Basin Naturalist Vol. 38, No. 1 Figs. 10-17. Neoperla choctaw: 10, Male terminalia, dorsal; 11, Vagina, lateral; 12, Male terga 7 and 8, lateral; 13, Female terminalia, ventral; 14, Aedeagus, lateral; 15, Aedeagus, ventral; 16, Apical section of aedeagus, ven- tral; 17, Aedeagus, apical section extruded, lateral. March 1978 Stark, Baumann: Nearctic Stoneflies 101 Female.— Macropterous. Length of fore- wings 8-10 mm; length of body 8-10 mm. Color pattern similar to male. Subgenital plate slightly produced, posterior margin smoothly rounded to emarginate. Vagina without sclerites; spermatheca short and in- completely lined with golden-brown spin- ulae in basal half. Egg.— Outline oval, cross section circular. Collar absent. Chorion finely punctate, with larger punctations mesally and finer punc- tations near polar ends. Types.— Holotype ( $ ), allotype ( ? ), and 1 (5 3 ? paratypes from Red Oak Creek, Denman, Latimer Co., Oklahoma, l-VIL 1970, B. Stark. Holotype, # 73358, and al- lotype deposited at the U.S. National Mu- seum. Paratypes: OKLAHOMA: Latimer Co., Red Oak Creek, Denman, 1-2-VII-1975, B. Stark, 5 5 12 ? (BPS) (NTSU); Turkey Creek, 6 mi E Red Oak, 2-VII-1975, B. Stark, 30 5 47 ? (BPS) (NTSU) (INHS). WEST VIRGINIA: Berkeley Co., Back Creek, 30- VII- 1946, E. P. Merkel, 3^1? (RWB) (USNM). Etymology.— This species was discovered in the former Choctaw portion of the In- dian Territory and is named in honor of that tribe. Neoperla freytagi, n. sp. Figs. 18, 19, 20, 21, 22, 23, 58, 59 Male.— Macropterous. Length of fore- wings 8-11 mm; length of body 7-10 mm. General color lemon yellow with dark brown markings. Dorsum of head with ocelli covered by a dark brown quad- rangular area. Pronotum light brown with irregular darker rugosities; marginal and median sutures dark brown. Wings sub- hyaline, coastal margin yellow. Process of tergum 7 triangular, ventrally spinulose. Process of tergum 8 moundlike, smoothly roimded in profile and spinulose; membrane of tergum 8 with 0-2 spinules. Tergum 9 with a mesal and two lateral spinule patch- es; anterior margins of each spinule patch with a number of long setae. Shaft of ae- deagus with a dense patch of spinules ven- trally; external spines absent. Extrusible, apical section of aedeagus membranous, bearing a few small spines. Female.— Macropterous. Length of fore- wings 12-14 mm; length of body 12-14 mm. Color pattern similar to male. Sub- genital plate slightly produced, posterior margin straight. Vagina with a pair of scle- rites; spermatheca long, incompletely lined with golden-brown spinulae from near basal third to near apex. Egg.— Outline oval, cross section circular. Collar with two rows of irregular reti- culation. Chorion longitudinally striate with alternating nonpvmctate, elevated bands and depressed finely punctate bands. Micropyles irregularly placed in depressed bands. Polar third of egg with irregular 4-, 5-, or 6-sided reticulation. Types.— Holotype ( $ ), allotype ( ? ), and 7 (5 24 ? paratypes from Larue Co., Ken- tucky, 16-VI-1971, P. H. Freytag & G. Lep- pert. Holotype, # 73356, and allotype de- posited at the U.S. National Museum. Paratypes: ARKANSAS: Benton Co., 24-VI- 1963, O. & M. Hite, 65 ? (INHS). NEW YORK: Greene Co., East Durham, 26-VII- 1971, S. E. Thewke, 2^6? (MU) (BPS). Tompkins Co., Ithaca, 2 1-30- VII- 1929, 2 S (BPS) (DC). OHIO: Warren Co., Little Miami River, Loveland, 2- VII- 1953, A. R. Gaufin, 1 ? (RWB). ONTARIO: Peel Co., Streetsville, 29- VIII- 1952, G. B. Wiggins, 2 ? (ROM); Churchville, Credit River, 14- VII- 1952, G. B. Wiggins, 2 ? (ROM); Erin- dale, Credit River, 16- VI- 1952, G. B. Wig- gins, 7 5 2? (ROM). Halton Co., Norval, Credit River, 14- VII- 1952, G. B. Wiggins, 2 S (ROM). SOUTH CAROLINA: Oconee Co., Brasstown, 29-VI-1969, H. Douglass, 1 ? (BPS). Pickens Co., Wildcat Creek, 25- VI/22-VII-1968, P. Carlson, 31 5 18 ? (PHC) (BPS) (NTSU). TENNESSEE: Sevier Co., Gatlinburg, 13- 19- VI- 1942, N. Dybas, 1 <5 4 ? (FMNH). Etymology.— This species is named for Dr. Paul Freytag who collected the type series. Neoperla gaufini, n. sp. Figs. 24, 25, 26, 27, 28, 69, 70 Male.— Macropterous. Length of fore- wings 6-8 mm; length of body 6-8 mm. Dorsum of head yellow with brown mark- ings on occiput, between ocelli and on lat- 102 Great Basin Naturalist Vol. 38, No. 1 eral margins of frons. Pronotum brown with darker rugosities and sutures. Tibiae, tarsi, and femora dark brown with distal yellow bands on tibiae and proximal yellow bands on femora. Wings and veins brown, coastal margin pale yellow. Process of tergum 7 slender, ventrally spinulose, extending to apical third of tergum 8. Process of tergum 8 abruptly elevated on posterior margin, slightly spinulose. Spinules absent from membrane of tergum 8. Mesal spinule patch of tergum 9 with 5-12 spinules, lateral patches with 3-6 scattered spinules. Shaft of aedeagus with external spines arranged in two ventral groups; left grouping with 2-6 spines, right gruping with 6-10 spines. Ex- trusible, apical section heavily spiculate and armed with several scattered spines. Female.— Macropterous. Length of fore- wings 7-9 mm; length of body 7-9 mm. Color pattern similar to male. Subgenital plate slightly produced, posterior margin smoothly rounded to emarginate. Vagina without sclerites; spermatheca short and completely lined with golden-brown spin- ulae in basal half. Figs. 18-23. Neaperla freytagi: 18, Male terminalia, dorsal; 19, Female sternum 8; 20, Male terga 7 and 8, lat- eral; 21, Vagina, dorsal; 22, Aedeagus, lateral; 23, Aedeagus, ventral. March 1978 Stark, Baumann: Nearctic Stoneflies 103 Egg.— Outline oval, cross section circular. Collar absent. Chorion covered with large regular depressions. Types.— Holotype ( $ ), allotype ( ? ), and 156? paratypes from Little Miami Riv- er, Morrow, Warren Co., Ohio, 26-VI-1952, A. R. Gaufin. Holotype, # 73354, and allo- type deposited at the U.S. National Mu- seum. Paratypes: INDIANA: Jefferson Co., Clifty Falls State Park, 18-VII-1972, A. Pro- vonsha & E. Levine, 1 ? (PU). KEN- TUCKY: Shelby Co., Brashears Creek, 3- VIII-1972, D. White, 1 $ (BPS). Spencer Co., Salt River, 25-26-VI-1971, V. Resh & D. White, 151? (BPS). OHIO: Clinton Co., Cowan Creek, 5-VI-1951, A. R. Gaufin, Figs. 24-28. Neoperla gaufini: 24, Male terminalia, dorsal; 25, Female sterna 7 and 8; 26, Vagina, lateral; 27, Aedeagus, lateral; 28, Aedeagus, ventral. 104 Great Basin Naturalist Vol. 38, No. 1 9 ,? 17 ? (UU) (BPS) (NTSU). Warren Co., Little Miami River, Loveland, 2- VII- 1953, A. R. Gaufin, 5 ^ 10 ? (RWB) (USNM). Etymology.— This species is named in honor of Dr. A. R. Gaufin who collected it and labeled it "Neoperla n. sp." in the early 1950s. Neoperla stewarti, n. sp. Figs. 29, 30, 31, 32, 33, 34, 63 Male.— Macropterous. Length of fore- wings 9-11 mm; length of body 9-11 mm. Dorsum of head yellow with diffuse brown markings on anterior region of frons; ocellar area dark brown. Pronotum brown with Figs. 29-34. Neoperla stewarti: 29, Male terminalia, dorsal; 30, Aedeagiis, apical section extmded, lateral; 31, Female sternum 8; 32, Aedeagus, lateral; 33, Aedeagus, ventral; 34, Vagina, dorsal. March 1978 Stark, Baumann: Nearctic Stoneflies 105 darker rugosities and sutures. Dorsum of tibiae and femora brown becoming lighter on ventral surface and apically; tarsi dark brown. Process of tergum 7 triangular, ven- trally spinulose, extending to near middle of tergum 8. Process of tergum 8 moundlike, smoothly rounded in lateral view and spin- ulose; membrane of tergum 8 with 5 or more spinules on each side of process. Ter- gum 9 with a mesal and 2 lateral spinule patches; anterior margins of each spinule patch with a number of long setae. Shaft of aedeagus with a small dense pad of spinules ventrally; external spines absent. Extrusible, apical section of aedeagus membranous, heavily armed with large spines and a basal group of small spinules. Female.— Macropterous. Length of fore- wings 11-13 mm; length of body 10-12 mm. Color pattern similar to male. Sub- genital plate slightly produced, or unpro- duced. Vagina with paired sclerites; sperma- theca long and lined with golden-brown spinulae in apical third. Egg.— Outline oval, cross section circular. Collar with a single irregular row of reti- culation. Chorion longitudinally striate with alternating nonpunctate, elevated bands and depressed finely punctate bands. Some ele- vated bands branch near midline. Micro- pyles irregularly placed in depressed bands. Polar third of egg with irregular 4-, 5-, or 6-sided reticulation. Types.— Holotype ( ; = 219 .81 .29 .14 .09 .19 .05 .21 .28 .35 .17 .11 .23 .06 .26 .37 .47 .30 .64 R19 R20 R21 R22 R23 R24 R25° R26 R27 R28 R29 R30° R31 R32 R33° R34 R35° R36 C. bairdi N = 219 .23 .67 1.05 .47 .32 .66 .22 .64 .74 .50 1.0 .34 .70 .48 .33 2.14 1.47 .69 C. beldingi N = 546 .31 .62 .96 .48 .28 .72 .32 .64 .76 .44 1.1 .51 .68 .40 .45 1.50 .88 .59 C. confusus N = 227 .30 .63 .99 .49 ..32 .70 .31 .64 .78 .51 1.1 .48 .71 .46 .44 1.62 1.06 .66 C. rhotheus N = 219 .17 .73 1.03 .45 .29 .62 .17 .70 .65 .41 .89 .24 .73 .46 .27 2.75 1.75 .64 'Allow separation of species. canal overlying each mandible and each preopercular bone, respectively. The pores of these canals are the openings of the preoperciilar-mandibular cephalic sensory systems. The head pore system of the genus Coitus has been widely used in systematic studies (Robins and Miller 1957, McAllister and Lindsey 1961, Robins 1954, 1961, McAllister 1964, 1968). Variability is gener- ally in the mental pore condition, with ei- ther the two mental pores fused into a single median chin pore or, more often, the median chin pore split into two mental pores (McAllister 1968). The Clearwater cottids usually had two mental pores and double postmaxillary pores. This condition gave 11 pores on each side of the preopercular-mandibular canal (Tables 4 and 5). Variation from this condi- tion resulted from fusion of the mental pores, absence or addition of one post- Table 3. Extent of body prickles on each of four species of cottids. Species Sample size Mean range Mean" S.E. C confusus C. beldingi C. bairdi C. rhotheus 219 546 227 316 2.0-2.4 1.0- 1.2 4.0- 8.0 11.0-15.0 2.1 1.0 6.0 13.1 0.2 0.1 2.1 2.2 "Based on a scale of 1 to 15. One indicates prickles absent and 15 in- dicates entire body covered with prickles. March 1978 Maughan: Idaho Sculpins 119 maxillary pore, absence of the third pore from the chin midline, an additional pore above the first preopercular spine, or a combination of these factors. Pores other than those mentioned were occasionally ab- sent. Variability in number of preopercular- mandibular pores was high only in South Fork Clearwater C. confusus. About 73 per- cent of the counts differed from 11-11 and ranged from 12-12 to 9-9. The counts were symmetrical (10-10, 9-9, 9-1-9) in about 64 percent of the variant individuals. The 10- 10 counts resulted from nearly 71 percent of the variant fishes having single post- maxillary pores. The postmaxillary pores were double on one side and single on the other in another 21 percent of the variant fishes. The 9-9 and 9-1-9 counts had a more variable basis. The 9-9 condition generally resulted from single postmaxillary pores plus the absence of another pore from each side (commonly the third pore from the chin midline). The 9-1-9 counts were usually a result of fusion of the mental pores into a single median pore plus single postmaxillary pores. Lateral Line.— The degree of com- pleteness of the lateral line is sometimes useful as a taxonomic character (Robins and Miller 1957, Koli 1969a, McAllister and Lindsey 1961). The lateral line was record- ed in this study as complete, incomplete, or interrupted and a number given the position at which the complete line terminated (Table 6). The lateral line in all four species generally extended caudad above the axial line to just slightly behind the extent of the soft dorsal fin. At this point, the lateral line or its remnant was deflected downward to the axial line. If the line were still present beyond this point, it followed the axial line to the hypural plate. Bailey and Bond (1963) found that the lateral line was generally incomplete or in- terrupted in C. confusus. Specimens from the Clearwater generally had the lateral line ending imder the last few soft dorsal rays or slightly behind the extent of the soft dorsal. However, 23 percent of the speci- mens had the lateral line interrupted. There were usually short segments of lateral line along the axial line anterior to the hypural plate in these specimens. Coitus beldingi usually had the lateral line incomplete or interrupted with 15 per- cent of all individuals collected having in- terrupted lateral lines, and less than 5 hav- ing complete lateral lines. However, 70-80 percent of the fish from Elk Creek had complete lateral lines. Cottus bairdi usually has a complete or nearly complete lateral line (Bailey and Table 5. Development of postmandibular pore for each of four species of cottids. Species Sample size Mean range Mean" S.E. C. confusus C. beldingi C. bairdi C. rhotheus 219 546 227 316 4.0-8.2 4.0-6.6 3.9-4.3 4.0-4.7 5.2 2.0 4.2 0.9 4.0 0.1 4.1 0.5 'Based on a scale of 1 through 12. One indicates postmandibular pores were triple on both sides. Twelve indicates postmandibular pores absent on both sides. Four indicates postmandibular pores double on both sides. Table 6. Mean extent of lateral line for each of four species of cottids. Sample Species size Mean range Mean" S.E. C. confusus 219 11.1-18.0 16.0 2.3 C. beldingi 546 10.3-20.3 17.6 1.8 C. bairdi 227 17.3-21.0 19.4 1.6 C. rhotheus 316 18.9-21.0 20.8 1.1 "Based on a scale of 1 to 21. One indicates lateral line ends under first soft dorsal ray. Twenty-one indicates line ends at hypural plate. Table 4. Mean number of preopercular-mandibular canal pores for each of four species of cottids. Sample Right Left Right Left Right Left Species size 219 mean range 9.7-11.2 mean range mean mean S.E. S.E. C. confusus 9.8-11.2 10.7 10.7 0.6 0.6 C. beldingi 546 10.5-11.3 10.2-11.7 10.9 10.9 0.3 0.4 C. bairdi 227 10.8-11.3 10.7-11.2 11.0 11.0 0.2 0.3 C. rhotheus 316 10.2-11.1 10.4-11.1 10.9 10.9 0.4 0.3 120 Great Basin Naturalist Vol. 38, No. 1 Dimick 1949). The lateral line usually end- ed just anterior to the hypural plate in Clearwater specimens but was complete in about 18 percent of the individuals. Coitus rhotheus generally has a complete lateral line except in very small individuals (Bailey and Dimick 1949). The lateral line was complete or nearly complete in all Clearwater specimens. Coitus rhotheus could generally be distin- guished from the other species by the com- pleteness of the lateral line, but the other species could not be distinguished from each other. In all species the development of the lateral line was related to age, with greater lateral line development in larger, mature individuals rather than in smaller fish. Preopercular Armature.— The number of preopercular spines is constant enough in some species of cottids to be useful as a tax- onomic character. In other species, this character is highly variable (Bailey and Bond 1963, Robins and Miller 1957). Coitus beldingi was reported to have a single preopercular spine (Eigenmann and Eigenmann 1891, Hubbs and Schultz 1932). This character varied from one to more than three spines in the Clearwater mate- rial. The mode for the species was one spine plus variable combinations of bumps, but the median value was more than two spines. Variability was high at all locations. Variability in spinous condition was also high in C. confusus. Coitus confusus from the Lochsa, Selway, and North Fork Clear- water usually had two or three spines with various combinations of bumps. Few indi- viduals with only one spine (16 percent) were seen. In contrast, individuals from the South Fork Clearwater had no spine, a single spine, or a single spine plus weakly developed bumps. Variation in the preopercular armature of C. rhotheus and C. hairdi was relatively slight. Most specimens of C. rhotheus had three spines plus bumps or four spines. Typ- ically, C. hairdi had three spines and either no bumps or weakly developed bumps. Species did not differ sufficiently in pre- opercular armature to allow identification, but development of the preopercular arma- ture did allow identification of two groups of two species (Table 7). Conclusions Body color was variable within and among species and was generally related to substrate color or breeding condition. No single meristic character allowed dif- ferentiation among these species. Pectoral ray counts allowed separation into two groups of two species but did not separate the species within the groups. Coitus bel- dingi and C. confusus usually had 13 or 14 pectoral rays, whereas C. hairdi and C. rhotheus usually had 15 or 16 rays. The development of palatine teeth usual- ly separated the four species, but unusual specimens could not be identified with this character. Coitus beldingi generally lacked palatine teeth, C. confusus usually had weakly developed palatine teeth, C. bairdi had well-developed palatine teeth with a small separation between the vomerine and palatine teeth, and C. rhotheus had the vomerine and palatine teeth cojoined and well developed. Most body proportions failed to separate species. However, ratios involving caudal peduncle depth usually separated C. rhotheus and C. bairdi from other species and from each other. The development of body prickles gener- ally separated the four species. Coitus bel- dingi lacked prickles, C. confusus had a small patch of axial prickles, C. bairdi had axial prickles plus a row of prickles above the lateral line, and C. rhotheus had prickles over most of the body. However, within each species, some fish showed the condi- tion typical of one of the other species. Table 7. Development of projections from the pre- opercular bone for each of four species of cottids. Species Sample size Mean range Mean" S.E. C. confusus C. beldingi C. hairdi C. rhotheus 219 546 227 316 8.0-28.9 15.9-27.9 30.6-34.5 32.4-34.7 24.3 6.5 23.4 5.8 32.2 1.7 .33.2 1.2 "Based on a scale of 1 to 35. One indicates no projections on pre- opercular bone. Thirty-five indicates four spines on preopercular bone. March 1978 Maughan: Idaho Sculpins 121 Larger C. rhotheus had apparently begun to better developed in C. bairdi and C. resorb prickles and approximated the condi- rhotheus than in C. beldingi or C. confusus. tion typical of C. foairdi. The two groups of two species could not be The development of the lateral line usual- separated using this character, ly separated C. rhotheus from the other spe- ^o single character allowed separation ces if adult specimens were used. The lat- ^ ^^e species of cottids from the era! line was usually complete in C. Clearwater. However, the characters studied tloTevTecr^^'"^ °' '""^"""P '"^ co"ld, when used in combination, give rea- ^ P ' ^■, „ sonable accuracy in identification. Ine preopercular armature was usually Key to the Clearwater Drainage Cottus 1. Palatine teeth generally present. Prickles developed at least in the axial posi- tion. Lateral line variable 2 Palatine teeth usually absent; if present, commonly only developed on one side. Body naked. Lateral line interrupted or incomplete, rarely complete. Found in the main river and lower tributaries C. beldingi 2(1). Prickles covering most of the body except in large specimens. Lateral line usually complete. Palatine teeth well developed, usually conjoined with the vomerine teeth. Caudal peduncle to total length ratio about 0.05. Skin over the eye nubbly. Body saddles extending cephalad at an angle to the long axis of the body. Restricted to the main river and the larger tributaries C. rhotheus Prickles restricted to an axial patch or an axial patch plus a bar paralleling the lateral line. Palatine teeth usually present. Lateral line usually incomplete, rarely complete 3 3(2). Prickles restricted to an axial patch. Palatine teeth usually present and weakly developed, sometimes absent in restricted locations. Lateral line usually ending under the last few soft dorsal rays, sometimes interrupted, with pores on the axial line just anterior to the hypural plate. Caudal peduncle to total length ratio about 0.07. Usually 13 or 14 pectoral rays. Restricted to the headwater areas at the upper portions of the drainage C. confusus Prickles usually an axial patch plus a bar above the lateral line. Palatine teeth strongly developed with only a slight separation between palatine and vomerine tooth patches. Lateral line usually incomplete, ending just anterior to the hypural plate, sometimes complete. Caudal peduncle to total length ratio about 0.06. Usually 15 to 16 pectoral rays. Skin over eye not nubbly. Dorsal saddles perpendicular to the long axis of the body. Restricted to the main river and larger tributaries C. bairdi Acknowledgments gy, University of Idaho, and Dr. Herbert Eastlick, Dr. Charles McNeil, Dr. Vincent Dr. Donald Chapman, Cooperative Fish- Schultz, and Dr. Robert Jonas, Department ery Research Unit, University of Idaho, sug- of Zoology, Washington State University, gested the study, and Dr. I. O. Buss, De- reviewed the work. Maurice O. Nyquist, partment of Zoology, Washington State Larry Gibson, and M. Gary Wickham assist- University, guided it to completion. Dr. ed in collecting, and the Idaho State Fish Carl E. Bond, Professor of Fisheries, Oregon and Game Department supplied necessary State University, gave advice and encour- collecting permits. The U.S. government agement and verified species identifications. (NDEA Grant) and Washington States Uni- Dr. Richard Wallace, Department of Biolo- versity provided funding for the study. The 122 Great Basin Naturalist Vol. 38, No. 1 Department of Zoology, Washington State University, also provided space, equipment, and supplies for the study. The Environ- mental Institute, Oklahoma State University, provided funds for publication. Literature Cited Bailey, J. E. 1952. Life history and ecology of the sculpin Cottiis bairdi ptinctulatus in south- western Montana. Copeia 1952:243-255. Bailey, R. M., and C. E. Bond. 1963. Four new spe- cies of freshwater sculpins, genus Coitus, from western North America. Occasional Papers 643:1-27. Museum of Zoology, University of Michigan. Bailey, R. M., and M. F. Dimick. 1949. Cottits hitbbsi, a new cottid fish from the Columbia River in Washington and Idaho. Occasional Papers 513:1-18. Museum of Zoology, University of Michigan. Eicenmann, C. H., and R. S. Eigenmann. 1891. Coitus beldingi sp. nov. Amer. Nat. 25:1132. HuBBS, C. L., AND L. P. ScHULTZ. 1932. Coitus tubu- latus, a new sculpin from Idaho. Occasional Pa- pers 242:1-9. Museum of Zoology, University of Michigan. Koli, L. 1969a. Geographic variation of Coitus gobio L. (Pisces, Cottidae) in northern Europe. Ann. Zool. Fenn. 6(4):353-390. 1969b. The spiniform armature of some palearc- tic freshwater species of the family Cottidae. Ann. Zool. Fenn. 6(4):428-430. Krejsa, R. J. 1965. The systematics of the prickly scul- pin, Coitus asper; an investigation of genetic and nongenetic variation within a polytypic species. Ph.D. dissertation. University of British Columbia, Vancouver, British Columbia. McAllister, D. E. 1964. Distinguishing characters for the sculpins, Coitus bairdi and Coitus cognatus, in eastern Canada. J. Fish. Res. Bd. Canada 21:1339-1.342. 1968. Mandibular pore pattern in the sculpin family Cottidae. Nat. Mus. Canada Bull. 223:58-59. McAllister, D. E., and C. C. Lindsey. 1961. System- atics of the freshwater sculpins (Coitus) of Brit- ish Columbia. Nat. Mus. Canada Bull. 172:66-89. Maughan, O. E. 1974. Paired fin ray counts of several species and populations of the genus Coitus. Copeia 1974:980. 1976. A survey of fishes of the Clearwater Riv- er. Northwest Sci. 50(2):76-86. Oliva, O., and K. Hensel. 1962. Studies on sculpins {Coitus gobio L.) from the river Pruth. Vestnik. Cechoslov. Spolecnosti Zool. 36:244-249. Robins, C. R. 1954. A taxonomic revision of the Coitus bairdi and Coitus carolinae species groups in eastern North America. Ph.D. dissertation. Cornell University, Ithaca, New York. 1961. Two new cottid fishes from the fresh wa- ters of the eastern United States. Copeia 1961:305-315. Robins, C. R., and R. R. Miller. 1957. Classification, variation, and distribution of the sculpins, genus Coitus, inhabiting Pacific slope waters in Cali- fornia and southern Oregon with a key to the species. Calif. Fish and Game 43:213-233. Smith, R. 1882. Description of a new species of Ura- nidea {Uranidea rhotheus) from the Spokane River, Washington territory. Proc. U.S. Nat. Mus. 1882:347-348. Smyly, W. J. P. 1957. The life history of the bullhead or miller's thumb (Coitus gobio L.). Proc. Zool. Soc. London 128(3):431-453. Williams, J., and C. R. Robins. 1970. Variation in populations of the fish Coitus carolinae in the Alabama River system with description of a new subspecies from below the fall line. Amer. Midi. Nat. 83(2): 368-381. CORRECTION: RHAMNUS CATHARTIC A L. IS NOT PRUNUS NIGRA AIT. FROM ALBERTA R. Keith Shaw' Abstract.— The plant previously reported from Alberta as Canada Plum (Pninits nigra Ait.) flowered and is now recognized European Buckthorn {Rhamnus cathartica L.). The report of Canada Plum (Primus nigra Ait.) in Alberta (Cody and Shaw 1973, Shaw 1976) was based on material collected without flowers or fruit from south Alberta. The preliminary identification was made from floral manuals only, with later veri- fication through comparison with known Prunus nigra material at the Vascular Plant Herbarium of the Canada Department of Agriculture, Ottawa. Several years later these south Alberta plants flowered and fruited. Much to my dismay, it was not Prunus nigra, but an ad- ventive species hitherto unreported for Al- berta to which Prunus nigra bears an un- canny resemblance, Rhamnus cathartica L., the European Buckthorn. This species was introduced from Europe and is now found growing wild as a coarse shrub or small tree in the valley of Lee Creek at Cardston, Al- berta. Two ornamental plantings are now also known for the town of Cardston. When the plants are in flower or fruit no real difficulty is encountered in correctly identifying European Buckthorn, which has flowers small, greenish, perfect, regular, perigynous with an annular disk; sepals 4; petals much reduced to wanting; stamens 4, inserted on the disk opposite the rudimen- tary petals; pistil 1, the ovary inserted in the disk, 4-loculed, each locule 1-seeded; fruit juicy, berrylike, and bitter. However, a comparison of vegetative characteristics will show how readily a per- son unfamiliar with these two species might make an error: spine-tipped branchlets no terminal bud toothed leaves, thin and fragile leaves dark green above, lighter below leaves tend toward opposite or clustered leaf margin glandular scaly winter buds twigs slender, reddish -brown bark smooth, dark greyish-brown, becoming scaly with age short, crooked trunk divided at or just above ground crown irregular; slender zig-zag branches small tree or large shrub thomlike twigs no terminal bud double-toothed leaves, thin and fragile leaves dull dark green above, paler beneath leaves clustered leaf margin glandular scaly winter buds twigs slender, smooth, reddish-brown bark black with greyish lenticels, splitting vertically, scaly with age short, crooked tnmk divided at or just above ground crown narrow and flat- topped; slender, twisted, zig-zag branches small tree Rhamnus cathartica European Buckthorn Prunus nigra Canada Plum I regret any inconvenience this erroneous report of Prunus nigra has caused. I will gladly supply herbarium specimens of south Alberta Rhamnus cathartica on request. Literature Cited Cody, W. J., and K. Shaw. 1973. Canada Plum in southwestern Alberta. Blue Jay 31:217-219. Shaw, R. K. 1976. A taxonomic and ecologic study of the riverbottom forest on St. Mary River, Lee Creek, and Belly River in southwestern Alberta, Canada. Great Basin Nat. 36:243-271. 'Box 364, Cardston, Alberta. TDK OKO 123 I NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Cwtcat Basin Naturalist should be di- rected to Brigham Young University, Stephen L. Wood, Editor, Great Basin Nattiralist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be sim- ilarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. 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Reprint Schedule of the Great Basin Naturalist 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each 100 copies $20 $24 $28 $32 $36 $40 additional 200 copies 28 32 36 40 44 48 2 pp. 300 copies 36 40 44 48 52 56 $4 Great Basin Naturalist Memoirs No. 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodbury, H. H. Frost. $10. No. 2 Intermountain biogeography: A symposium. By K. T. Harper, J. L. Reveal, et al. $15. TABLE OF CONTENTS Endangered and threatened plants of Utah: A reevaluation. Stanley L. Welsh 1 Plants of Navajo National Monument. J. D. Brotherson, G. Nebeker, M. Skougard, and J. Fairchild 19 Environmental interaction in summer algal communities of Utah Lake. Mark C. Whiting, Jack D. Brotherson, and Samuel R. Rushforth 31 Basidiomycetes that decay junipers in Arizona. II. R. L. Gilbertson and J. P. Lind- sey 42 Pinus longaeva in the Stansbury Mountains, Utah. P. A. Kay and C. G. Oviatt 49 Differential parasitism of Peromyscus maniculatus and Peromyscus truei by Cut- erebra larvae. Jeffrey B. Llewellyn 51 An annotated list of the aquatic insects of southeastern Idaho, Part III. Ephem- eroptera. Robert L. Newell and G. Wayne Minshall 55 Observations on the prey and nests of seven species of Cerceris (Hymenoptera: Sphecidae). Howard E. Evans and William L. Rubink 59 Vertebrate fauna of a radioactive leaching pond complex in southeastern Idaho. Douglas K. Halford and Jere B. Millard 64 New synonymy, new species, and taxonomic notes of North American Pityoph- thorus (Coleoptera: Scolytidae), Part III. Donald E. Bright 71 Fluctuation of a marsh habitat and the reproductive strategy of the Yellow-headed Blackbird. Roger J. Lederer 85 Hosts of juniper mistletoe at Walnut Canyon National Monument, Arizona. Frank G. Hawksworth and Robert L. Mathiasen 89 Allelopathic effects of Bur Buttercup tissue on germination and growth of various grasses and forbs in vitro and in soil. Bruce A. Buchanan, K. T. Harper, and Neil C. Frischknecht 90 New species of Nearctic Neoperla (Plecoptera: Perlidae), with notes on the genus. Bill P. Stark and Richard W. Baumann 97 Morphometry of sculpins (Cottus) in the Clearwater drainage, Idaho. O. Eugene Maughan 115 Correction: Rhamnus cathartica L. is not Prunus nigra Ait. from Alberta. R. Keith Shaw 123 FHE GREAT BASIN NATURALIST Volume 38 No. 2 June 30, 1978 Brigham Young University ^^mf^ 2-^ f^^[S ^r .J^^^^ .!^^ MUS. COMP. ZOOL,. LfBRARV y^JIm JAN 2 2 1979 ^^^W HARVARD UrviiVERSJTY # GREAT BASIN NATURALIST MEMOIRS Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agri- cultural Sciences; Ernest L. Olson, Director, Brigham Young University Press, Univer- sity Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only previously unpublished manuscripts of less than 100 printed pages in length and pertaining to the biological natural history of western North America are ac- cepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manu- scripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist is $12 (outside the United States $13). The price for single numbers is $4 each. All back numbers are in print and are available for sale. All matters pertaining to the purchase of subscriptions and back numbers should be directed to Brigham Young University, Life Science Museum, Pro- vo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same of- fice at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts and other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back cover. 11-78 650 34941 ISSN 0017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 38 June 30, 1978 No. 2 KAIPAROWITS FLORA Stanley L. Welsh', N. Duane Atwood,^ and Joseph R. Murdock' Abstract.— The paper presents an annotated list of 851 taxa in 358 genera and 80 families. Threatened and endangered plants are enumerated. Diversity of vegetative types in the Kai- parowits Basin is reflected in the large number of species present in that basin.^ Elevational differences of 2,680 m from the low point at Lee's Ferry to the high point on the summit of Navajo Mountain accotmts for some of the diversity of plant species. Substrate differences in the basin range from saHne clays through all the common particle sizes to boulders and slickrock. The focus of the basin is the Colorado River canyon into which all of the drainages of the basin pour their loads, and plants have followed this feature as a major migrational pathway. This is true of the major commu- nities of plants in the vicinity, and is espe- cially so for those at lower elevations. Lower elevational species are representa- tive of more southern distributional patterns which find the hot dry canyon and the lower features compatible with their needs. Middle and higher elevation species are more representative of boreal vegetation. The two types intertongue in the lower middle elevations where soil conditions are compatible. Specific soil types tend to be occupied by characteristic kinds of vegeta- tion, that which is sufficiently specialized as to survive and compete on the unusual soils. This is demonstrated on the saline clays of such formations as the Tropic Shale which forms the benchland below the Nipple Bench-Smoky Mountain uplands. Amounts of precipitation are roughly cor- rectable with elevation, and the low amotmts at low elevations are reflected in the kinds of plants which can grow there. This is also true for the middle and higher elevations. Vegetation and the diversity of species change from one rainfall regimen to another. Where water is available on a per- manent basis, such as along streams or in hanging gardens, the plant assemblage changes drastically when contrasted to that in nearby arid lands. All of these habitat types combine to set the Kaiparowits Basin apart as a unique entity. The following annotated list of vascular plants contains some 851 taxa (822 species and 29 infraspecific units) placed in 358 genera and 80 families. They represent di- verse phytogeographical provinces from 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 'Bureau of Land Management, Cedar City, Utah 84720. Present address: U.S. Forest Service, Lloyd 500 Building, Suite 1692, 500 NE Multnomah, Portland, Oregon 97232. This is one of a series of papers funded by the Navajo and Kaiparowits Generating Stations participants. 125 126 Great Basin Naturalist Vol. 38, No. 2 cold to warm temperate and even sub- tropical regions of this hemisphere. They are recognized as belonging to montane for- est, ponderosa pine, juniper-pinyon, sage- brush, salt-desert shnib, warm desert shrub, riparian, and hanging garden communities. The list is based upon the plant collec- tions made by the authors and by others at the Navajo-Kaiparowits Project during the period from 1971 to 1975. Additionally, all plants in the herbarium of Brigham Young University have been reviewed, and the per- tinent specimens are cited in the list. Lists published by others have been studied and some species added in that manner, but un- less the reports could be validated, question- able reports have been excluded. All speci- mens cited are in the herbarium of Brigham Young University (BRY) unless otherwise in- dicated. The list should not be thought to be ex- haustive; other species will be foimd in the Basin. This is indicated by species being added to the list in the summer of 1975, following several years of extensive in- vestigation of the region to sample its flora. Many of the plants are recognized as threatened (T), endangered (E), or possibly extinct (PoEx). Designations for these spe- cies is in a paper by Welsh et al. (1975). Some 42 species in the Kaiparowits Basin are considered as belonging to one of these categories. This number represents about 5 percent of the species in the Basin and ap- proximately 20 percent of the threatened, endangered, or possibly extinct species known for Utah. The following species, with status indicated, belong to this unique group of plants. Astragalus desperatus var. conspectus (T) Astragalus bryantii (PoEx) Astragalus emoryanus (T)° Astragalus hallii var. fallax (T)" Astragalus kentrophyta var. coloradoensis (T)" Astragalus lancearius (T) Astragalus malacoides (T) Astragalus striatiflorus (E) Atriplex navajoensis (T) Carex curatorum (T) Castilleja revealii (E) Cirsium rydbergii (T)° Cladium californicum (E)" Draba subalpina (T) Erigeron abajoensis (T) Euphorbia nephradenia (T) Geranium marginale (T) Gilia latifolia (T)" Imperata breviflora (PoEx)° Ivesia sabulosa (T)" Lesquerella rubicundula (T) Lesquerella tumulosa (E) Lomatium minimum (T) Machaeranthera glabriuscula var. confertifhra (T) Menodora scabra (T) Muhlenbergia curtifolia (T)° Nama retrorsum (T) Oenothera megalantha (E)° Oxytropis jonesii (T)" Penstemon leiophyllus (T) Phacelia cephalotes (T) Phacelia constancei (T) Phacelia mammillariensis (E) Phacelia rafaelensis (T) Phlox cluteana (T)° Pinus longaeva (T)° Primula specuicola (T) Psoralea epipsila (E) Ptelea trifoliata ssp. pallida (PoEx)* Townsendia minima (T) Viguiera annua (T) Yucca toftiae (T) Most of these plants occupy only small portions of the Basin, and they must be avoided by any planned development of the region. Generally, the plants are those which occupy some specific habitat or soil types, and they are not known to be cul- tivatable by standard agricultural tech- niques. Thus, the only means of preserva- tion of these unique plants lies in the maintenance of the areas occupied by them. In the annotated list, the collector's names have been appreviated or shortened. The most commonly noted collectors are as follows; a look at collection dates will allow one to determine an outline of the history of floristic investigations of the Basin. Abbreviation Collector's Name Boyle Burkey BW W. S. Boyle N. Burkey B. W. Wood Christensen E. M. Christensen Beck D. E. Beck Buchanan H. Buchanan CAT C. A. Toft Cottam W. P. Cottam DT D. Trotter JRM Gentry Harrison J. R. Murdock J. Gentry, Jr. B. F. Harrison Holmgren Holmgren Karren N. Holmgren P. Holmgren J. B. Karren LCH L. C. Higgins June 1978 Welsh et al.: Kaiparowits Flora 127 « en I 00 128 Great Basin Naturalist Vol. 38, No. 2 NDA N. D. Atwood Nebeker & SkougardG. Nebeker & M. Skougard RA or RWA R. W. Allen Reveal J. L. Reveal SLW S. L. Welsh SL & SLW S. L. & S. L. Welsh Stanton W. D. Stanton Tanner V. M. Tanner Webb M. Webb Woodruff D. W. Woodruff Annotated Checklist AcERACEAE (Maple Family) Acer glabrum Torr. Rocky Mountain Maple Utah: Garfield Co., Bryce Canyon, Boyle 1154. Rare. Acer grandidentatum Nutt. Big-Tooth Maple Utah: Garfield Co., Bryce Canyon, Bu- chanan 379. Kane Co., Kaiparowits Plateau, JRM 399. Rare. Acer negundo L. Box Elder Arizona: Coconino Co., Paria Canyon, NDA & LCH 4015, 20 May 1972. Utah: Kane Co., Escalante Canyon, NDA & RA 3216. Uncommon to locally common along streams and in some hanging gardens. AizoACEAE (Carpetweed Family) Mollugo cerviana (L.) Ser. Utah: Kane Co., Cedar Mt., NDA 3101. Rare. Amaranthaceae (Amaranth Family) Acanthochiton wrightii Torr. Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3346, 3346a. Utah: Kane Co., 2 miles west of Glen Canyon City, NDA & DK 3360, 3360a. The Utah collec- tions are the first known record of this rare plant for Utah. Amaranthus graecizans L. Utah: Kane Co., Cockscomb, NDA & RA 2888b. San Juan Co., Colorado River, across from Last Chance Creek, Gaines (1960). Tidestromia lanuginosa (Nutt.) Standi. Utah: Kane Co., 30 miles east of Glen Canyon City NDA & DK 3335; 18 miles east of Glen Canyon City, NDA 4059. Un- common. Anacardiaceae (Cashew Family) Rhus trilobata Nutt. var. trilobata Skunkbush Utah: Garfield Co., Escalante Canyon, Beck s. n.; Escalante, Beck & Tarmer s. n. Kane Co., 50 miles south of Escalante, Beck & Tarmer s.n.; Nipple Spring, NDA & RA 2825. Rocky slopes and canyons, usually near moist seeps or along drainages; uncom- mon. Rhus trilobata Nutt. var. simplicifolia (Greene) Barkley Arizona: Coconino Co., Lee's Ferry, Cot- tam 2607. Utah: Kane Co., Willow Tank, Beck & Tanner s. n.; 8 miles south of Paria Townsite, Harrison 12046; Four Mile Bench, SLW & NDA 12376; Hole-in-the- Rock, Harrison 12128; Cockscomb, SLW 5330; Tibbet Canyon, NDA 3461; Wah- weap Creek, NDA 3475. Rocky slopes in canyons; common. Toxicodendron rydbergii (Small) Greene Poison Ivy Utah: San Juan Co., one mile east of Hole-in-the-Rock, SLW & CAT 11868. Confined to moist sites, especially in hang- ing gardens, where it is imcommon but lo- cally abimdant. Apocynaceae (Dogbane Family) Amsonia eastwoodiana Rydb. Utah: San Juan Co., east of confluence of San Juan, NDA 4089; Three Garden, SLW & NDA 11699. Sandy sites in warm dry canyons; common. Amsonia palmeri Gray Arizona: Coconino Co., near Lee's Ferry, NDA 4429. Rare. Amsonia tomentosa Torr. & Frem. Utah: San Juan Co., east of confluence of the San Juan, NDA 4090; do, NDA 3182. Dry rocky ground; uncommon. Apocynum androsaemifolium L. Utah: Garfield Co., Bryce Canyon, Cot- tam 2744. Moist canyons and slopes at high- er elevations; imcommon. Apocynum cannabinum L. Dogbane Utah: Garfield Co., Escalante, Cottam 4396. Kane Co., Coyote Creek, Brigham Plains area, NDA & RA 2802; Little Valley June 1978 Welsh et al.: Kaiparowits Flora 129 Canyon, NDA et al. 2718. San Juan Co., Es- calante Canyon, NDA 4116. Seeps, springs, and hanging gardens; common. AscLEPiADACEAE (Milkwccd Family) Asclepias asperula Decne Rough Milkweed Utah: Garfield Co., Escalante Canyon, Woodruff 1164. Kane Co., Dry Rock Creek, Lake Powell, SLW & NDA 11628; head of Last Chance Creek. NDA 4072. San Juan Co., Navajo Mt., NDA et al. 2973; do, NDA 4196; Three Garden, SLW & NDA 11679. Widespread but uncommon. Asclepias cryptoceras Wats. Hidden-horn Milkweed Utah: Garfield Co., 1.3 miles south of Cannonville, Reveal et al. 762. Kane Co., Cottonwood Wash road, LCH & NDA 5258; do, NDA & RA 2907. Clay soils, widespread but uncommon. Asclepias labriformis Jones Utah: Garfield Co., 10 miles southeast of Escalante, Holmgren et al. 2042. Utah en- demic; widespread northward. Asclepias latifolia (Torr.) Raf. Utah: Kane Co., 50 miles south of Esca- lante, Beck & Tanner s. n.; Cedar Mt., NDA 2919; Flat Top, NDA et al. 2959; Es- calante Canyon, SLW & CM 11812; 31.5 miles east of Glen Canyon City, NDA et al. 2700. Sandy sites, especially among boulders on talus; uncommon. Asclepias macrosperma Eastw. Arizona: Coconino Co., Navajo Canyon, NDA 3728. Utah: Kane Co., Cockscomb, SLW 5350; 5 miles north of Glen Canyon Dam, LCH 1006; 13 miles west of Glen Canyon City, NDA & RA 2859; Escalante Canyon, NDA 4215; 2 miles east of Glen Canyon City, NDA 2918; Hole-in-the-Rock, White 212, do, Harrison 12146; Wahweap Creek, Harrison 12089; Paria River, SLW & NDA 9752. Sandy soils at low elevations; common. Asclepias subverticillata (Gray) Vail Utah: Garfield Co., east of Escalante, Holmgren & Holmgren 4723. Kane Co., 30 miles east of Kanab, NDA 5588. Asclepias tuberosa L. var. interior Woodson Utah: Garfield Co., east of Escalante, Holmgren et al. 2117; Escalante, Cottam 4381, Escalante Canyon, Beck s.n. Rare in sandy rim rock situations. Funastrum heterophyllum (Engelm.) Standi. Climbing Milkweed Arizona: Coconino Co., Lee's Ferry, Cot- tam 2617; do NDA 6388. Utah: Kane Co., Hole-in-the-Rock, Harrison 12730. San Juan Co., Three Garden, NDA 4263. Talus slopes, canyon of the Colorado; rare. Berberidaceae (Barberry Family) Mahonia fremontii Torr. Fremont Mahonia Utah: Kane Co., 35 miles southeast of Es- calante, Harrison 12341; Nipple Canyon, RA 257; Tibbet Spring, NDA 3463; 20 miles south of Cannonville, Christensen s.n.; 8 miles south of Paria Townsite, Harrison 12060. Common in drainages and bench- lands through much of the Basin. Mahonia repens L. Creeping Oregon-grape Utah: Garfield Co., Bryce Canyon, Bur- key 124. Kane Co., Brigham Plains vicinity, NDA & RA 2798; Buckskin Mountain, NDA 4658; Kaiparowits Plateau, JRN 398. San Juan Co., Navajo Mt., NDA 4159. Moun- tains and moist drainages; locally common. Betulaceae (Birch Family) Betula occidentalis Hook. River Birch Utah: Garfield Co., Bryce Canyon, Boyle 1155. Kane Co., Escalante Canyon, SLW & CM 11829; head of Wahweap Creek, SLW & BW 12441. Along drainages and seeps and springs; imcommon. Ostrya knowltonii Gov. Knowlton Ironwood Utah: San Juan Co., Rincon Canyon, NDA 4103; do, SLW & CM 11783; do SLW & NDA 11693; do, SLW & NDA 11711; do SLW 11893. Local in Ribbon Canyon, and observed in Escalante Canyon; rare. BoRAGiNACEAE (Borage Family) Coldenia hispidissima (Torr.) Gray Arizona: Coconino Co., upstream from Lee's Ferry, NDA & LCH 4032. Utah: Kane Co., Cedar Mt., NDA 3096; Gunsight Bay, SLW & NDA 11576; Wahweap Creek, 130 Great Basin Naturalist Vol. 38, No. 2 Lake Powell, NDA 3702. Sandy sites at low elevations; common. Cryptantha abata Johnst. Utah: Garfield Co., Bryce Canyon, Boyle 1117. Kane Co., Last Chance Creek, NDA 3747; Collet Wash, NDA 1866. Clay soils; local and uncommon. Cryptantha hakeri Pays. Utah: Garfield Co., Bryce Canyon, Weight s. n., 1934; 4 miles southwest of Cannonville, Reveal et al. 772; Canaan Peak, NDA & DT 5314. Moderate to high elevations; uncommon. Cryptantha capitata (Eastw.) Johnst. Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5253. High elevations; rare. Cryptantha circumscissa (H. & A.) Johnst. Arizona: Coconino Co., one mile west of Glen Canyon Dam, NDA 3447. Utah: Kane Co., East Clark Bench, LCH 4251, 20 May 1971; Paria River, SLW & NDA 9747; Warm Creek, NDA 3712; Grand Bench, NDA 3526; Kaiparowits Plateau, Holmgren et al. 2068; Labyrinth Canyon, Lake Pow- ell, NDA 3737; Cottonwood Wash road. Re- veal & Gentry 809; Cedar Mt., NDA 2736a. Sandy soils, a common spring annual. Cryptantha confertiflora (Greene) Pays. Utah: Garfield Co., 8 miles southeast of Escalante, Holmgren et al. 2036. Kane Co., Paria Townsite, Webb 62; Collet Wash, NDA 1873; Paria River, NDA & LCH 3957; Hackberry Canyon, NDA 3678; Ahl- strom Point road, NDA 3668; Cockscomb, NDA 3613; Tibbett Canyon, NDA 3602. Common yellow-flowered perennial of sandy soils. Cryptantha crassisepala (T. & G.) Greene Arizona: Coconino Co., 3 miles south of Wahweap Marina, SLW & NDA 9768; up- stream from Lee's Ferry, NDA 4593: Paria Plateau, NDA & LCH 3767. Utah: Garfield Co., Willow Tank, Beck & Tanner s. n.; Warm Creek, NDA 3715; Paria River, NDA & LCH 3976; Wahweap Bay, Lake Powell, NDA 3551; Paria River, SLW & NDA 9748. Annual plant of sandy soils; common. Cryptantha fendleri (Gray) Greene Arizona: Coconino Co., Lee's Ferry, NDA 4573; one mile west of Glen Canyon Dam. NDA 3569. Utah: Kane Co., 2 miles west of Glen Canyon City, Nebeker & Skougard 191; Paria River, NDA & LCH 3944; Little Valley Canyon, NDA 3520. Sandy soils; un- common. Cryptantha flava (A. Nels.) Pays. Arizona: Coconino Co., east of Glen Can- yon Dam, NDA 3451. do, SLW & NDA 9778; 9 miles south of Page, RA 218. Utah: Garfield Co., Escalante, Cottam 4408. Kane Co., 31 miles south of Escalante, Karren 89; Cottonwood Wash road, NDA & RA 2807; East Clark Bench, NDA & RA 2677; Kai- parowits Plateau, Woodruff 1147; Paria River, SLW & NDA 9750; Grand Bench, NDA 3528; head of Rock Creek, Harrison 9033; Cockscomb, SLW 5342; Buckskin Gulch, SLW 5319. Widespread and com- mon, yellow-flowered, ubiquitous plant. Cryptantha fulvocanescens (Gray) Pays, var. echinoides (Jones) Higgins Utah: Garfield Co., 1.3 miles south of Cannonville, Reveal et al. 745. Kane Co., Cottonwood Wash road, NDA 3468; 5 miles east of Butler Valley, NDA 3458a; 6 miles southeast of Canaan Peak, SLW & NDA 12372; head of Last Chance Creek, NDA 3752; Paria Townsite, SLW & NDA 9729; East Clark Bench, NDA & RA 2673. Sandy to clay soils; common. Cryptantha gracilis Osterch. Utah: Kane Co., Four Mile Bench, SLW & JRM 12409; Kaiparowits Plateau, NDA & DT 5288; Cockscomb, NDA 4608; Kaiparo- wits Plateau, Harrison 9070a. Understory in pinyon-juniper woodland; common. Cryptantha humilis Gray Pays. var. com- mixta (Macbr.) Higgins Utah: Garfield Co., Bryce Canyon, Bu- chanan 1484 (Bryce). Kane Co., Ahlstrom Point road, NDA 3669; Five Mile Moun- tain, NDA 4642. Rocky soils at middle ele- vations; uncommon. Cryptantha jamesii (Torr.) Pays. var. dis- ticha (Eastw.) Pays. Arizona: Coconino Co., Page vicinity, SLW & NDA 9777, do, NDA 3452. Utah: Kane Co., Paria River, NDA & LCH 3970; 24 miles west of head of Collet Wash, NDA 1873a; Nipple Bench, NDA & RA 2816; Tibbet Canyon, NDA 3601; Crosby Canyon, NDA 3634; Kaiparowits Plateau, NDA & DT 5239; Cockscomb, NDA 3581; Hack- June 1978 Welsh et al.: Kaiparowits Flora 131 berry Canyon, NDA 3677. Ubiquitous; com- mon. Cryptantha jamesii (Torr.) Pays. var. pustu- losa (Rydb.) Harrington Utah: Kane Co., Circle Cliffs, Beck s. n. Cryptantha jamesii (Torr.) Pays. var. setosa (Jones) Johnst. Utah: Kane Co., Cottonwood Wash road, LCH & NDA 5261; 20 miles east of Kanab, NDA 1792a. Uncommon. Cryptantha micrantha (Torr.) Johnst. Arizona: Coconino Co., Lee's Ferry, NDA 4577; 3 miles south of Wahweap Marina, SLW & NDA 9767. Utah: Kane Co., Cedar Mt., NDA et al. 2735; Warm Creek, NDA 3708; Paria River, NDA & LCH 3950; 2 miles west of Glen Canyon City, Nebeker & Skougard 185. Sandy soil; common. Cryptantha nevadensis Nels. & Kenn. Arizona: Coconino Co., upstream from Lee's Ferry, NDA 4587; Lee's Ferry, NDA 4571. Utah: Kane Co., Labyrinth Canyon, Lake Powell, NDA 3736; 6 miles east of Glen Canyon City, SLW & NDA 9810. Lower elevations; uncommon. Cryptantha pterocarya (Torr.) Greene Arizona: Coconino Co., Lee's Ferry, NDA 4572; upstream from Lee's Ferry, NDA 4586; one mile west of Glen Canyon Dam, NDA 3568. Utah: Kane Co., Hack- berry Canyon, NDA 3676; Kaiparowits Plateau, Holmgren et al. 2064; one mile west of Glen Canyon City, Cronquist 10155; Cedar Mt., NDA et al. 2736, 1971. Sandy soils; common. Cryptantha recurvata Gov. Arizona: Coconino Co., Lee's Ferry, NDA 4570. Utah: Kane Co., Little Valley Can- yon, NDA 3518; Nipple Canyon, RA 247; Labyrinth Canyon, Lake Powell, NDA 3738; Kaiparowits Plateau, Harrison 9070 b; 6 miles east of Glen Canyon City, SLW & NDA 9811. San Juan Co., Three Garden, SLW & NDA 11703. Sandy soils; uncom- mon. Cryptantha setosissima (Gray) Pays. Utah: Garfield Co., Bryce Canyon, Bu- chanan 2053 (Bryce). Heliotropium convolvulaceum (Nutt.) Gray Heliotrope Utah: Kane Co., Cedar Mt., NDA & RA 3073; Warm Creek, NDA & RA 2846. Dunes, sandy terraces, and drainages; im- common. Lappula occidentalis (Wats.) Greene Arizona: Coconino Co., upstream from Lee's Ferry, NDA 4584. Utah: Garfield Co., Bryce Canyon, Weight s. n.; 1.3 miles south of Cannonville, Reveal et al. 752. Kane Co.; Willow Tank, Beck & Tanner s. n.; Cocks- comb, NDA 3626, Cottonwood Wash road, NDA & RA 2756; Gunsight Bay Lake Pow- ell, SLW & NDA 11586; Paria River, NDA & LCH 3981. Kaiparowits Plateau, NDA & DT 5286; Four Mile Bench, NDA 3756. Ubiquitous; common. Lithospermum incisum Lehm. Utah: Garfield Co., Bryce Canyon, Bu- chanan 1449, 1454, and 1483 (Bryce); do, Boyle 1112. Kane Co., Grosvenor Arch vi- cinity, NDA 3469; Cottonwood Wash road. Reveal 814; Horse Mt., Four Mile Bench vi- cinity, SLW & NDA 12371. Uncommon, widespread. Lithospermum multiflorum Torr. Stoneseed Utah: Garfield Co., Bryce Canyon, Bu- chanan 1448 (Bryce): do, Boyle 1114. Pectocarya platycarpa Munz Utah: San Juan Co, Three Garden, SLW & NDA 11906. Sandy soils; imcommon to rare. Cactaceae (Cactus Family) Coryphantha vivipara (Nutt.) Britt. & Rose var. arizonica (Engelm.) W. T. Marshall Utah: San Juan Co., Navajo Mt., NDA 4151. Widespread; rare. Echinocereus engelmannii (Parry) Rumpler Hedgehog Cactus Utah: Kane Co., Brigham Plains, NDA & RA 2784a; 36 miles east of Glen Canyon City, NDA et al. 2707. Dry, open slopes, at lower elevations; uncommon. Echinocereus triglochidiatus Engelm. var. melanacanthus (Engelm.) Benson Utah: Garfield Co., 1.5 miles south of Cannonville, Reveal et al. 768. Kane Co., Four Mile Bench, SLW & JRM 12397a; 8 miles south of Glen Canyon City, NDA 3556; Cockscomb, SLW 5364; Willow Tank, SLW s.n. 18 miles south of Paria Townsite, Harrison 12056. Widespread, mostly at middle elevations; uncommon. 132 Great Basin Naturalist Vol. 38, No. 2 Opuntia basillaris Engelm. & Bigel var. aurea (Baxter) Benson Utah: Kane Co., Four Mile Bench, SLW & JRM 12397. Widespread and locally com- mon, middle elevations. Opuntia chlorotica Engelm. & Bigel. Arizona: Coconino Co., Lee's Ferry, Clo- ver and Jotter (1941). Opuntia erinacea Engelm. & Bigel. var. histricina (Engelm. & Bigel.) Benson Utah: Kane Co., Grand Bench, NDA 4258; Little Valley Canyon, NDA et al. 2732. Common low elevations. Opuntia erinacea Engelm. & Bigel. var. utahensis (Engelm.) Benson Utah: Kane Co., Four Mile Bench, NDA 4057. Widespread; locally common. Opuntia macrorhiza Engelm. var. macror- hiza Utah: San Juan Co., Surprise Valley, Clo- ver and Jotter (1941). Opuntia phaecantha Engelm. Utah: San Juan Co., Navajo Mt., NDA 4141. Common in low elevation canyons and terraces. Opuntia polyacantha Haw. var. poly- acantha Utah: Kane Co., 20 miles east of Glen Canyon City, NDA 4063; Fifty Mile Spring, White 155. Widespread and locally com- mon. Opuntia whipplei Engelm. & Bigel. Arizona: Coconino Co., 9.3 miles south of Page, NDA 4083; Kaibito-Tonolea road, NDA et al. 2912. Utah: Kane Co., Cedar Mt., SLW & JRM s.n. Sandy sites at low to moderate elevations in southern portion of basin; uncommon. Sclerocactus whipplei (Engelm. & Bigel.) Britt. & Rose Utah: Kane Co., Four Mile Bench, SLW & JRM 12411; Crosby Canyon, NDA 3636; 8 miles south of Glen Canyon City, NDA 3557. Widespread; uncommon to locally common. Campanulaceae (Bellflower Family) Campanula parryi Gray Utah: Garfield Co., Bryce Canyon, Weight s. n., 1934; Escalante Canyon, Beck 134. Moist sites; uncommon. Campanula rotundifolia L. Utah: Garfield Co., Bryce Canyon, Bu- chanan 1723 (Bryce). Uncommon. Lobelia cardinalis L. ssp. graminea (Lam.) McVaugh Utah: Kane Co., Reflection Canyon, Lake Powell, SLW 11879; Escalante Canyon, NDA & RA 3204. San Juan Co., Rainbow Bridge, NDA & RA 3301; Three Gardens, NDA & RA 3192; do, NDA et al. 3228; east of confluence of the San Juan, NDA & RA s.n. Hanging gardens and seeps; uncom- mon except locally. Nemacladus glanduliferus Jeps. var. orienta- lis McVaugh Arizona: Coconino Co., Lee's Ferry, NDA 4576. Uncommon; southern portion of in- cluded area. Capparidaceae (Caper Family) Cleome lutea Hook. Beeplant Utah: Garfield Co., Escalante, Beck & Tanner s.n.; 5 miles west of Escalante, An- derson 683; 1.3 miles south of Cannonville, Reveal et al. 751; Kane Co., 50 miles south of Escalante, Beck & Tanner s.n., 16 June 1936; Escalante Canyon, SLW & CM 11831; Paria River, SLW & NDA 9742; East Clark Bench, LCH 4246; Left-hand fork of Collet Wash, Woodruff 1160; Cot- tonwood Wash road, NDA & RA 1769; Cockscomb, SLW 5351. Widespread; com- mon. Cleome serrulata Pursh Utah: Kane Co., Paria River, NDA & LCH s.n.; Four Mile Bench, NDA 4303. Widespread; uncommon. Cleomella palmeriana Jones Utah: Garfield Co., one mile east of Hen- rieville, NDA 1877; 18 miles west of Esca- lante, SLW 9411. Kane Co., Crosby Can- yon, NDA 3635; 17 miles northeast of Glen Canyon City, RA 203; Paria River, NDA & LCH 3989; Cottonwood Wash road, NDA & LCH 3830; 18 miles east of Glen Canyon City, NDA 4058; 4 miles east of Glen Can- yon City, NDA 2629. Restricted to clay soils, mainly on the Tropic Shale formation; endemic to the Navajo Basin of Utah and Colorado; common. June 1978 Welsh et al.: Kaiparowits Flora 133 Caprifoliaceae (Honeysuckle Family) Sambucus coerulea Raf. Elderberry Utah: Garfield Co., between Death Ridge and Canaan Peak, SLW et al. 12589. San Juan Co., Navajo Mt., NDA et al. 2979. Canyons and slopes at middle elevations; uncommon. Symphoricarpos longiflorus Gray Snowberry Utah: Kane Co., Brigham Plains vicinity, NDA & RA 2799. San Juan Co., Ribbon Garden, Lake Powell, SLW & NDA 11696. Common in canyons at low to moderate elevations. Symphoricarpos parishii Rydb. Utah: San Juan Co., Navajo Mt., NDA 4146. Uncommon. Symphoricarpos vaccinioides Rydb. Utah: Garfield Co., vicinity of Death Ridge SLW & JRM s.n., June 1975. Kane Co., Bryce Canyon, Boyle 1132; Kaiparo- wits Plateau, NDA & DT 5290. Middle and higher elevations; uncommon. Caryophyllaceae (Pink Family) Arenaria eastwoodiae Rydb. Sandwort Utah: Kane Co., Hole-in-the-Rock, Harri- son 12144; 16 miles east of Glen Canyon City, NDA 3667; Kaiparowits Plateau, NDA & DT 52227. Sandy sites and rimrock; un- common. Arenaria fendleri (Rydb.) Fern. Utah: Garfield Co., Canaan Peak, NDA & DT 5318. Uncommon. Arenaria kingii Wats. Utah: Garfield Co., 2 miles southeast of Escalante, NDA S.N. Kane Co., Four-Mile Bench, NDA 4304. Arenaria nutallii Wats. Utah: Garfield Co., Bryce Canyon, Bur- key 129. Uncommon; higher elevations. Silene antirrhina L. Arizona: Coconino Co., 9.3 miles north of junction of US 89 and 169, NDA 3656. Utah: Kane Co., Cockscomb, NDA 4609. San Juan Co., Three Garden, SLW & NDA 11667, 4 May 1972; Ribbon Canyon, SLW & NDA 11712; Trail Canyon, Lake Powell, SLW 11911. Widespread; uncommon. Silene scouleri Hook. ssp. pringlei (Wats.) Hitchc. & Maguire Utah: San Juan Co., Navajo Mt., NDA 4180. Rare. Stellaria jamesiana Torr. James Starwort Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5239. San Juan Co., Navajo Mt., NDA et al. 2994; do. Gentry & Da- vidse 1770. Uncommon; middle elevations. Celastraceae (Bittersweet Family) Forsellesia spinescens (Gray) Greene Greasebush Utah: Kane Co., Cockscomb, NDA 4613. Uncommon. Pachystima myrsinites (Pursh) Raf. Mountain Lover Utah: Garfield Co., Bryce Canyon, Bu- chanan 1488 (Bryce). Uncommon. Chenopodiaceae (Goosefoot Family) Atriplex canescens (Pursh) Gray Fourwing Saltbush Utah: Garfield Co., Escalante Canyon, Harrison 9142. Kane Co., Four Mile Bench, SLW et al. 12583; Nipple Bench, NDA & RA 2821; Gunsight Bay, SLW & NDA 11592; Hole-in-the-Rock, Hansen 432a; 13 miles east of Warm Creek Junction, NDA 3457. Common throughout the Basin in most vegetative types. Atriplex confertifolia (Torr. & Frem.) Wats. Shadscale Utah: Garfield Co., 3 miles east of Hen- rieville, Hansen 395. Kane Co., Nipple Can- yon, RA 248; Gunsight Bay, Lake Powell, SLW & NDA 11588; 5 miles southeast of Cannonville, Reveal et al. 794; Sit Down Bench, Patton 137; 6 miles east of Glen Canyon City, SLW & NDA 9803. Through- out the Basin; abundant. Atriplex corrugate Wats. Mat-Atriplex Utah: Kane Co., lower Wahweap Creek, Albee 60; 30 miles east of Glen Canyon City, NDA 4545; Ahlstrom road, RA 120; Nipple Canyon, RA 261; Cottonwood Wash road, Reveal et al. 826; 6 miles east of Glen Canyon City, SLW & NDA 9802. Principal perennial vegetation of Tropic Shale forma- tion. Atriplex garrettii Rydb. Utah: Kane Co., Hole-in-the-Rock, Han- sen 432b; San Juan Co., Three Garden, SLW & NDA 11687; east of confluence of 134 Great Basin Naturalist Vol. 38, No. 2 the San Juan, NDA 4088; Ribbon Garden, NDA 4109; between Three Garden and Willow Cove, Lake Powell, SLW 11895. Talus slopes, canyons of the Colorado; un- common. Atriplex graciliflora Jones Utah: Kane Co., Warm Creek, NDA & RA 2839; 29.2 miles southeast of Escalante, Reveal et al. 827. Saline soils; uncommon. Atriplex jonesii Standi. Arizona: Coconino Co., Lee's Ferry, Clo- ver and Jotter (1941). Atriplex navajoensis Hanson Navajo Saltbush Arizona: Coconino Co., Lee's Ferry, NDA 6393; do, Cottam 2620; do, NDA 4351, 4352. Endemic to Coconino County, Ari- zona; known only from Lee's Ferry to Nav- ajo Bridge vicinity; possibly threatened. Atriplex powellii Wats. Utah: Garfield Co., 2 miles east of Hen- rieville, NDA 5359. Kane Co., Cottonwood Wash road. Reveal et al. 828. Locally and periodically abundant, especially on saline clay soils. Atriplex saccaria Wats. Utah: Kane Co., Cockscomb, NDA & RA 2883; Little Valley Canyon, SLW & NDA 11610. Uncommon, saline clay soils. Ceratoides lanata (Pursh) Howell Winterfat Utah: Kane Co., Last Chance Bay, Lake Powell, NDA et al. 3291; one mile east of Church Wells, NDA & RA 3116; 5 miles south of Glen Canyon City, NDA & RA 3116. Common in low elevations, especially on sandy or silty soils. Chenopodium fremontii Wats. Goosefoot Utah: Kane Co., Dry Rock Creek, SLW & NDA 11623; miles south of Glen Canyon City, NDA 3117a. Uncommon. Chenopodium leptophyllum Nutt. Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3340. Utah: Kane Co., near mouth of Kane Creek Gaines (1960). Uncommon to rare. Chenopodium pratericola Rydb. Utah: Kane Co., 5 miles south of Glen Canyon City, NDA 3117. Rare to uncom- mon. Chenopodium sp. Utah: Kane Co., Reflection Canyon, SLW 11876. Corispermum hyssopifolium (Pallas) L. Bugseed Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3343. Utah: Kane Co., Four Mile Bench, NDA 4309. Uncommon. Corispermum nitidum Kit. Arizona: Coconino Co., Teetso Spring, NDA 3420; 25 miles north of Shonto, NDA & DT 3392. Utah: San Juan Co., 5.5 miles below Klondike Bar, Gaines (1960). Uncom- mon. Cycloloma atriplicifolium (Spreng.) Coult. Utah: Kane Co., Willow Tank, Beck & Tanner s.n.; Warm Creek, NDA et al. 3289. Grayia brandegei Gray Utah: Garfield Co., 12 miles east of Hen- rieville, Hanson 396a; 18 miles west of Es- calante, SLW 9140. Kane Co., Hackberry Canyon, NDA 3685; Cottonwood Wash road, Christensen s.n.; do, NDA & DK 3319. Restricted to clay and silty, saline soils where locally abundant. Grayia spinosa (Hook.) Moq. Utah: Kane Co., 31 miles east of Glen Canyon City, NDA et al. 2697; Last Chance Bay, Lake Powell, SLW & NDA 11608; Nipple Canyon, RWA 256. Codomi- nant with Coleogyne ramosissima in por- tions of the desert shrub community; un- common to very common. Halogeton glomeratus (Bieb.) C.A. Meyer Utah: Garfield Co., Pet Hollow, NDA 4323. Observed elsewhere; uncommon in the Basin. Kochia americana Wats. Gray Molly Utah: Kane Co., observed but not collect- ed. Locally common on pediment gravels overlying Tropic Shale formation east from Glen Canyon City. Monolepis nuttalliana (Schult.) Greene Arizona: Coconino Co., upstream from Lee's Ferry, NDA 4583. Utah: Kane Co., Warm Creek, NDA 3600; Tibbet Canyon, NDA 3608. Widespread; uncommon. Salsola iberica Sennen & Pau Russian Thistle Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3348. Utah: Kane Co., Willow Tank, Beck & Tanner s.n.; 6 miles east of Glen Canyon City, NDA & DK 3356; Warm Creek, NDA & RA 2837. Widespread, adventive weed. June 1978 Welsh et al,: Kaiparowits Flora 135 Sarcobatus vermiculatus (Hook.) Torr. Greasewood Utah: Garfield Co., Pet Hollow, NDA 4328. Greasewood is a dominant plant spe- cies in saline soils along drainages through- out middle and lower elevations. Suaeda fruticosa (L.) Forsk. Seepweed Utah: Garfield Co., 18 miles west of Es- calante, SL & SLW 9412. Locally abun- dant; saline clay soils. Suaeda torreyna Wats. Utah: Kane Co., Cottonwood Wash road, NDA & DK 3324. Locally abundant; saline clay soils. CoMMELiNACEAE (Spidcrwort Family) Tradescantia occidentalis (Britt.) Smyth Spiderwort Arizona: Coconino Co., Paria Canyon, Woodruff 1122. Utah: Garfield Co., Esca- lante Canyon, Beck s.n. San Juan Co., For- bidding Canyon, SLW & NDA 11661; Az- tec Canyon, Gaines (1960). CoMPOSiTAE (Composite Family) Acamptopappus sphaerocephalus (Harv. & Gray) Gray Goldenhead Utah: Kane Co., Warm Creek, NDA 3599; Gunsight Bay, SLW & NDA 11589; Dry Rock Creek, SLW & NDA 11624; La- byrinth Canyon, NDA 3731; 14 miles east of Glen Canyon City, NDA 4062. San Juan Co., confluence of the San Juan, SLW 11933. Sandy and gravelly benches at low elevations; locally common. Achillea millefolium L. Utah: Garfield Co., Escalante Canyon, Beck 144. Widespread at higher and middle elevations; uncommon. Agoseris arizonica Greene Mountain Dandelion Utah: San Juan Co., Navajo Mt., NDA & DT 5339. Uncommon. Agoseris aurantiaca (Hook.) Greene Utah: San Juan Co., Navajo Mt., NDA 4168. Uncommon. Agoseris glauca (Pursh) D. Dietr. Utah: Garfield Co., Bryce Canyon, Weight s.n., 1934. Kane Co., Kaiparowits Plateau, Harrison 9045; do, NDA & DT 5233. High elevations; uncommon. Ambrosia acanthicarpa (Hook.) Gov. Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3342. Utah: Kane Co., Cedar Mt., NDA 3120; Paria River, NDA & DK 3374; 6 miles east of Glen Canyon City, NDA & DK 3358; Four Mile Bench, NDA 4306. Sandy and silty soils; wide- spread and common. Antennaria dimorpha (Nutt.) T. & G. Pussy toes Utah: Garfield Co., Bryce Canyon, Bu- chanan 2058 (Bryce). Uncommon. Antennaria parvifolia Nutt. Utah: Garfield Co., Bryce Canyon, Cot- tam 2750. San Juan Co., Navajo Mt., NDA 4183. High and moderate elevations; un- common. Antennaria rosulata Rydb. Utah: Garfield Co., Bryce Canyon, Bu- chanan 1718 (Bryce). Arnica cordifolia Hook. Utah: Garfield Co., Circle Cliffs, Beck s.n. Artemisia bigelovii Gray Bigelow Sagebrush Utah: Garfield Co., west of Escalante, Cottam 6356. Kane Co., Tibbet Canyon, NDA & DK 3326. Rocky slopes and draws; common. Artemisia campestris L. Utah: Garfield Co., Escalante Canyon, Beck 116. Uncommon. Artemisia filifolia Torr. Old Man Sagebrush Utah: Garfield Co., Escalante Canyon Beck 115. Kane Co., 50 miles south of Esca- lante, Beck & Tanner s.n.; 5 miles south of Glen Canyon City, NDA 3118; Paria River, JRM 322; Nipple Bench, NDA & DK 3328. Common on sandy sites, especially at lower elevations. Artemisia frigida Willd. Utah: Garfield Co., Bryce Canyon, Bu- chanan 493; Escalante Canyon, Beck 114. Uncommon. Artemisia ludoviciana Nutt. Utah: Kane Co., Paria River, NDA & DK 3377; Escalante River, NDA 4221. Locally common especially in talus. Artemisia nova A. Nels. Black Sagebrush Utah: Garfield Co., Bryce Canyon, Bu- chanan 73. Abimdant at high elevations, mostly west of our area. 136 Great Basin Naturalist Vol. 38, No. 2 Artemisia pygmaea Gray Utah: Garfield Co., locally common to abimdant, limestone scablands, Bryce Can- yon vicinity; 5 miles west of Widtsoe Junc- tion, SLW & JRM 12563. Artemisia spinescens DC. Bud Sagebrush Utah: Kane Co., one mile northwest of Warm Creek-Last Chance Road, NDA 3438, Ahlstrom road, RA 121; Nipple Can- yon, RA 255. San Juan Co., Rock Creek vi- cinity, Gaines 91960). Locally common; gravelly pediment over saline soils. Artemisia tridentata Nutt. Big Sagebrush Utah: Garfield Co., Escalante, Beck & Tanner s.n. Kane Co., Four Mile Bench, SLW 12367. San Juan Co., Navajo Mt., NDA & DK 3396. Common to abundant in openings in juniper-pinyon woodlands at middle elevations. Aster chilensis Nees ssp. adscendens (Lindl.) Cronq. Aster Utah: Garfield Co., Bryce Canyon, Weight, s.n. Kane Co., Reflection Canyon, SLW 11888. San Juan Co., Surprise Valley, Clover and Jotter (1941). Moist sites; un- common. Aster arenosus (Heller) Blake Arizona: Coconino Co., near Tse' eh' aah, Nebeker & Skougard 208. Utah: Garfield Co., Bryce Canyon, Cottam 2769; Escalante Beck & Tanner s.n. Kane Co., Four Mile Bench, SLW & JRM 12401; Willow Tank, Harrison, 9115; East Clark Bench, Harrison 12062; Paria River, NDA & LCH 4011; Grand Bench, NDA 3527; Nipple Bench, NDA & RA 2808. Ubiquitous; common. Aster commutatus (T. & G) Gray Utah: Kane Co., Four Mile Bench, Wah- weap Creek, SLW 12568. Seeps and springs; rare. Aster frondosus (Nutt.) T. & G. Utah: Kane Co., Four Mile Bench, Wah- weap Creek, NDA 4316. Seeps; rare. Aster glaucodes Blake Utah: Garfield Co., Bryce Canyon, Weight s.n.; Pet Hollow, NDA 4335. Kane Co., Escalante Canyon, NDA 4210. Moist or shaded sites; uncommon. Aster pauciflorus Nutt. Utah: Garfield Co., Escalante Canyon, Beck 1135. Kane Co., Upper Wahweap Creek, NDA 4342. Moist sites; uncommon. Aster spinosus Benth. Arizona: Coconino Co., Lee's Ferry, Cot- tam 2626. Rare. Baccharis emery i Gray Arizona: Coconino Co., 9.3 miles south of Page, NDA 4085. Utah: Kane Co., Drift- wood Canyon, SLW & NDA 11649; Reflec- tion Canyon, SLW 11884; Hole-in-the-Rock, Harrison 12125. Moist sites along major, low elevations drainages; locally commmon. Baccharis glutinosa Pers. Arizona: Coconino Co., Lee's Ferry, Cot- tam 2619. Utah: San Juan Co., 5.5 miles be- low Klondike Bar, Gaines (1960). Uncom- mon. Bahia neomexicana Gray Arizona: Coconino Co., Cockscomb, NDA & DK 3382. Rare. Balsamorrhiza sagittata (Pursh) Nutt. Utah: Garfield Co., Bryce Canyon, Weight s.n. Kane Co., Horse Mt., north of Four Mile Bench, SLW s.n. Gravelly soils at middle elevations; locally common. Brickellia atractyloides Gray Utah: San Juan Co., Three Garden, SLW & NDA 11688. Uncommon to rare. Brickellia calif ornica (T. & G.) Gray Utah: Kane Co., Brigham Plains vicinity, NDA & RA 2794. Rare. Brickellia longifolia Wats. Utah: Kane Co., Tibbet Canyon, NDA & DK 3332; do, NDA & DK 3411. Canyon bottoms and talus slopes; locally common in tributaries of the Colorado. Brickellia oblongifolia Nutt. var. linifolia (DC.) Robinson Utah: Garfield Co., Bryce Canyon, Ma- guire 19105; do Cottam 4363; 3 miles southeast of Escalante, Holmgren et al. 2025. Kane Co., Four Mile Bench. SLW 12832; Willow Tank, Beck & Tanner s.n. Cottonwood Wash road, NDA & RA 2757. Uncommon to locally common at middle and low elevations. Brickellia scabra (Gray) A. Nels. Arizona: Coconino Co., 8 miles southeast of Page, NDA et al. 3242. Utah: Garfield Co., Escalante Canyon, Beck s.n. Locally common on sandstone outcrops. Calycoseris parryi Gray Utah: Kane Co., Hole-in-the-Rock, Gaines (1960). June 1978 Welsh et al.: Kaiparowits Flora 137 Centaurea repens L. Utah: Kane Co., Cottonwood Wash road, NDA & RA 2767; Four Mile Bench NDA 4315. Uncommon; introduced weed. Chaenactis brachiata Greene Utah: Garfield Co., Bryce Canyon, Cot- tam 2573. Rare. Chaenactis douglasii (Hook.) H. & A. Utah: Garfield Co., Bryce Canyon, Weight s.n., 1934. Kane Co., East Clark Bench, JRM 329; Collet Top, Woodruff 1151. San Juan Co., Navajo Mt., NDA et al. 2967. Uncommon to locally common bien- nial. Chaenactis macrantha DC. Utah: Kane Co., 20 miles east of Glen Canyon City, NDA & DT 5071; Gunsight Bay, Lake Powell, SLW & NDA 11585. San Juan Co., 2 miles below Last Chance Creek, Gaines, (1960). Locally common in sandy sites at low elevations. Chaenactis stevioides H. & A. Arizona: Coconino Co., Page vicinity, SLW & NDA 9773. Utah: Kane Co., 3.3 miles southwest of Glen Canyon City, NDA et al. 2745; 1.1 miles west of Glen Canyon City, Cronquist 10158; Gunsight Bay, Lake Powell, SLW & NDA 11575; Grand Bench, NDA 3529. Common in sandy sites at low elevations. Chamaechaenactis scaposa (Eastw.) Rydb. var. parva Preece & Turner Utah: Garfield Co., Red Canyon, Reveal & Reveal 1029. Perhaps outside the limits of the Kaiparowits Basin. Chrysothamnus depressus Nutt. Utah: Garfield Co., Bryce Canyon, Harri- son 11030. Common at moderate elevations. Chrysothamnus linifolius Greene Utah: Garfield Co., Escalante Canyon, Beck 136. Kane Co., Reflection Canyon, CAT 192; Four Mile Bench, SLW et al. 12574; Last Chance Creek, NDA 427a. A tall shrub of drainage margins; locally abun- dant. Chrysothamnus nauseosus (Pall.) Britt. var. alhicaulis (Nutt.) Rydb. Utah: Garfield Co., Bryce Canyon, Bu- chanan 1725 (Bryce). Moderate to high ele- vations; common. Chrysothamnus nauseosus (Pall.) Britt. var. bigelovii (Gray) Hall Arizona: Coconino Co., Lee's Ferry, Cot- tam 2603. Uncommon. Chrysothamnus nauseosus (Pall.) Britt. var. consimilis (Greene) Hall Arizona: Coconino Co., Lee's Ferry, Clo- ver and Jotter (1941). Utah: Kane Co., Tib- bet Canyon, NDA & DK 3408, 3409; Last Chance Creek, NDA 4274. Locally common on terraces and flood plains. Chrysothamnus nauseosus (Pall.) Britt. var. gnaphaloides (Greene) Hall) Utah: Kane Co., Tibbet Canyon, NDA & DK 3410. Locally common along canyon bottoms. Chrysothamnus nauseosus (Pall.) Britt. var. graveolens (Nutt.) Hall Utah: Garfield Co., Bryce Canyon, Cot- tam & Hutchings 2775. Locally common at middle elevations. Chrysothamnus nauseosus (Pall.) Britt. var.« junceus (Greene) Hall Utah: Kane Co., Nipple Bench, NDA & DK 3330. Uncommon. Chrysothamnus nauseosus (Pall.) Britt. var. leiospermus (Gray) Hall Utah: Kane Co., Reflection Canyon, SLW 1182; 2 miles west of Glen Canyon City, NDA & DK 3407. Uncommon. Chrysothamnus parryi (Gray) Greene var. attenuatus (Jones) Kittel in Tidestr. & Kit- tel Utah: Garfield Co., Escalante vicinity, Holmgren et al. 2535; Bryce Canyon, Bu- chanan 1728 (Bryce). Moderate elevations; common. Chrysothamnus viscidiflorus (Hook.) Nutt. var. puberulus (Eat.) Jeps. Utah: Garfield Co., Bryce Canyon, Bu- chanan 1726 (Bryce). Moderate elevations; common. Chrysothamnus viscidiflorus (Hook.) Nutt. var. stenophyllus (Gray) Hall Arizona: Coconino Co., 9 miles southeast of Page, NDA et al. 3240. Utah: Kane Co., 2 miles south of Glen Canyon City, NDA 3097; Buckskin Gulch, SLW 9423; East Clark Bench, NDA & DK 3366. Common at lower elevations on several soil types. Chrysothamnus viscidiflorus (Hook.) Nutt. var. viscidiflorus Utah: Garfield Co., Escalante Canyon, Beck 145. Uncommon. 138 Great Basin Naturalist Vol. 38, No. 2 Cichorium intybus L. Chickory Utah: Kane Co., Four Mile Bench, NDA 4118. Introduced weed; uncommon. Cirsium arizonicwn (Gray) Petrak Utah: Kane Co., Four Mile Creek, SLW et al. 12576. Canyon bottom; uncommon. Cirsium bipinnatwn (Eastw.) Rydb. Utah: Kane Co., Escalante Canyon, SLW & CM 11828. Canyon bottoms; rare. Cirsium neomexicanum, Gray Utah: Kane Co., Driftwood Canyon, SLW & NDA 11648; Dry Rock Creek, Lake Powell, SLW & NDA 11633; Grand Bench, NDA 4261. Sandy sites, especially in can- yons at low elevations; common. Cirsium nidulum (Jones) Petrak Utah: Kane Co., Escalante Canyon, NDA 4120. Uncommon. Cirsium pulchellum (Greene) W. & S. Arizona: Coconino Co., Paria Canyon, Woodruff 1126. Utah: Kane Co., 20 miles east of Glen Canyon City, NDA & DT 5070. Left-hand Fork of Collet Canyon, Woodruff 1154; Wahweap Creek, Four Mile Bench, SLW & BW 12441; Brigham Plains vicinity, NDA & RA 2793; Little Valley Canyon, NDA et al. 2714. Common, especially in canyon bottoms. Cirsium rothrockii (Gray) Petrak Utah: San Juan Co., Navajo Mt., Gentry & Davidse 1975. Uncommon. Cirsium rydbergii Petrak Utah: San Juan Co., east of confluence of the San Juan, NDA & RA 3181. Endemic to the Navajo Basin, restricted mainly to Hanging Gardens; local and possibly threat- ened (Welsh et al, 1975). Cirsium utahense Petrak Utah: Kane Co., Escalante Canyon, NDA 4115. Uncommon. Conyza canadensis (L.) Cronq. Horseweed Utah: San Juan Co., Three Garden vicin- ity, NDA & RA 3182. Rainbow Bridge, Clo- ver and Jotter (1941). Crepis intermedia Gray Utah: Garfield Co., Bryce Canyon, Weight B-31/28. Uncommon. Crepis occidentalis Nutt. Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5341. Uncommon. Crepis runcinata T, & G. var. glauca (Bab- cock) Stebb. Utah: Kane Co., Four Mile Creek, SLW & BW 12443. Seeps and springs margins; rare. Dicoria canescens Gray Utah: Kane Co., East Clark Bench, NDA & DK 3371; 6 miles east of Glen Canyon City, NDA & DK 3371. Common in sandy sites. Dyssodia thurberi (Gray) A. Nels. Arizona: Coconino Co., Lee's Ferry, NDA 4565. Utah: Kane Co., Escalante Canyon, NDA & RA 3215; do, NDA & RA 3200; Dry Rock Creek; SLW & NDA 11626; Tib- bet Canyon, NDA 3609; Rainbow Bridge, SLW & NDA 11652; Hole-in-the-Rock, Beck 141. San Juan Co., east of confluence of the San Juan, NDA & RA 3159. Wide- spread and common. Encelia frutescens Gray Utah: Kane Co., Warm Creek, NDA 3722; Dry Rock Creek, SLW & NDA 11616; Gunsight Bay, SLW & NDA 11579 Last Chance Creek, Woodruff 1142. Wide- spread in canyons at low elevations; uncom- mon. Ericeliopsis nudicaulis (Gray) A. Nels. Arizona: Coconino Co., Lee's Ferry vicin- ity, Cottam 2629. Rare. Erigeron abajoensis Cronq. Utah: Garfield Co., Bryce Canyon, Cot- tam 2754; do NDA 6128. Kane Co., Kai- parowits Plateau, Harrison 9062. A rare and threatened species endemic to Garfield, Kane, and San Juan counties, Utah (Welsh et al., 1975). Erigeron bellidiastrum Nutt. var. bellidiast- rum Utah: Garfield Co., Escalante, Cottam 4394. Kane Co., 45 miles south of Esca- lante, Beck 8278, July 1937; 22 miles south of Cannonville, Reveal et al. 810; Hole-in- the-Rock, Buss 93; 2 miles west of Glen Canyon City, NDA 3637. Common. Erigeron compositus Pursh Utah: Garfield Co., Bryce Canyon, Cot- tam 2768. Uncommon; higher elevations. Erigeron divergens T. & G. Arizona: Coconino Co., upstream from Lee's Ferry, NDA & LCH 4021. Utah: San Juan Co., Three Garden vicinity NDA & RA 3184; Navajo Mt., Gentry & Davidse June 1978 Welsh et al.: Kaiparowits Flora 139 1972. Locally common, especially in moist canyon bottoms. Erigeron eatonii Gray Utah: Garfield Co., Bryce Canyon, Bu- chanan 2049 (Bryce). Kane Co., Kaiparowits Plateau, Harrison 9062. San Juan Co., Nav- ajo Mt., NDA 4156. Erigeron flagellaris Gray Utah: Garfield Co., Bryce Canyon, Cot- tam & Hutchings 2770. Uncommon. Erigeron pumilus Nutt. ssp. concinnoides Cronq, Utah: Garfield Co., 1.3 miles south of Cannonville, Reveal et al. 754. Kane Co., Four Mile Bench, SLW & JRM 124000; Kaiparowits Plateau, NDA & DT 5227; Brigham Plains vicinity, NDA & RA 2795; Fifty Mile Spring, JRM 358. Widespread; uncommon. Erigeron speciosus (Lindl.) DC. var. macranthus (Nutt.) Cronq. Utah: Garfield Co., Bryce Canyon, Weight s.n., 1934. Uncommon. Erigeron utahensis Gray Utah: Garfield Co., Escalante Canyon, McKnight & Beck 199; Escalante, Cottam 4392. Kane Co., 20 miles southeast of Can- nonville, Holmgren et al. 2052; Brigham Plains, NDA & RA 2783; 2 miles west of Glen Canyon City, Cronquist 10167; 37 miles east of G]en Canyon City, NDA et al. 2708; Kaiparowits Plateau, NDA & DT 5270; Willow Tank, Harrison 9029; Drift- wood Canyon, SLW & NDA 11647; Paria River, NDA & LCH 3978; Warm Creek, NDA & RA 2904. Common; especially in rocky canyon slopes. Erigeron vagus Pays. Utah: Garfield Co., Bryce Canyon, Reveal 1019. Rare. Gaillardia gracilis A. Nels. Blanket Flower Utah: Garfield Co., Escalante Canyon, Beck s.n. Kane Co., Cockscomb, SLW 5348; Willow Tank, Harrison 9106; 15 miles west of Glen Canyon City, NDA & RA 2682a. Sandy soils; locally abundant. Gaillardia parryi Greene Utah: Garfield Co., Escalante vicinity, Holmgren et al. 2073; Kane Co., 10 miles southeast of Cannonville, Reveal et al. 804; Buckskin Wash, SLW 9421. Locally com- mon. Gaillardia pinnatifida Torr. Utah: Garfield Co., Escalante Canyon, McArthur & Beck 200, June 1940. Kane Co., 6 miles west of Escalante, NDA 3575; 15 miles west of Glen Canyon City, NDA & RA 2862; Paria River, LCH & NDA 5370a. Locally common. Gnaphalium palustre Nutt. Utah: San Juan Co., near mouth of Aztec Creek, Gaines (1960). Grindelia aphanactis Rydb. Utah: San Juan Co., Three Garden vicin- ity, NDA & RA 3189; do SLW & CAT 11864. Locally common; canyon bottoms. Grindelia squarrosa (Pursh) Dunal Utah: Kane Co., Four Mile Bench, NDA 4321. Locally common. Gutierrezia microcephala (DC.) Gray Arizona: Coconino Co., 8 miles southeast of Page, NDA et al. 3245. Utah: Garfield Co., Escalante Canyon, Beck 120. Kane Co., Tibbet Spring, NDA & DK 3414. Common. Gutierrezia sarothrae (Pursh) Britt. & Rus- by Utah: Garfield Co., Escalante Canyon, Beck 137. Pet Hollow, NDA 4326. San Juan Co., Navajo Mt., Gentry & Davidse 1971. Common at higher and middle elevations. Haplopappus acaulis (Nutt.) Gray Utah: Garfield Co., 1.3 miles south of Cannonville, Reveal et al. 750. Kane Co., Cockscomb, NDA 3621; Cockscomb, NDA 3584; Buckskin Gulch, SLW 5314. Com- mon. Haplopappus acradenius (Greene) Blake Goldenweed Arizona: Coconino Co., Lee's Ferry, Clo- ver and Jotter (1941). Haplopappus armerioides (Nutt.) T. & G. Utah: Garfield Co., Bryce Canyon, Cot- tam 4357. Kane Co., Smoky Mt., RA 20; 51 miles south of Escalante, NDA 517; Four Mile Bench, SLW & JRM 12404. Locally common. Haplopappus drutnmondii (T. & G.) Blake Arizona: Coconino Co., Lee's ferry, NDA 6392. Utah: Kane Co., Kane Creek along Colorado River, Buss s. n., 1935; Hole-in- the-Rock, Beck 140; Cottonwood Wash road, SL & SLW 9417; Tibbet Canyon, NDA & DK 3331; Wahweap Creek, NDA et al. 4299. San Juan Co., east of confluence 140 Great Basin Naturalist Vol. 38, No. 2 of the San Juan, NDA & RA 3188. Locally common; canyon bottoms. Haplopappus scopulorwn (Jones) Blake Utah: San Juan Co., Ribbon Canyon, NDA 4100; Navajo Mt., NDA & DK 3394. Uncommon to rare; threatened (Welsh et al., 1975). Helenium hoopesii Gray Sneezeweed Utah: Garfield Co., north of Escalante, Mc Arthur & Beck 188. Uncommon; higher elevations. Helianthella parryi Gray Utah: San Juan Co., Navajo Mt., NDA et al. 3001; do. Gentry & Davidse 1788. Rare. Helianthus anomalus Blake Sunflower Utah: Garfield Co., Escalante, Beck & Tanner s. n. Kane Co., Dry Rock Creek, SLW & NDA 11615; Escalante Canyon, SLW & CM 11822; Willow Springs south of Escalante, Harrison 9099. Uncommon; sandy drainages and dunes. Helianthus petiolaris Nutt. Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3347. Utah: Garfield Co., Escalante Canyon, McKnight & Beck 201; Cannonville, SLW 9413. Kane Co., 3.3 miles north of Paradise Estates, NDA & RA 2676. San Juan Co., Colorado Canyon, 2 miles below Last Chance, Gaines (1960). Common. Heterotheca villosa (Nutt.) Shinners Golden Aster Arizona: Coconino Co., Paria Canyon, Woodruff 1124. Utah: Kane Co., 43 miles east of Glen Canyon City, NDA et al. 2720; Left-hand fork of Collet Wash, Woodruff 1162; 6 miles west of Glen Canyon City, NDA 2921; Cockscomb, NDA 3615; Hack- berry Canyon, NDA & RA 2804; Smoky Mt., NDA et al. 3296; Cedar Mt., NDA 3129a. San Juan Co., Navajo Mt., NDA et al. 2978; Three Garden, NDA & RA 3179. 3185. Ubiquitous; common. Hymenopappus filifolius Hook. var. cine- reus (Rydb.) Johnst. Utah: Kane Co., Willow Tank, Harrison 9096; 37 miles east of Glen Canyon City, NDA et al. 2710. San Juan Co., Three Gar- den vicinity, SLW & NDA 11664; Forbid- ding Canyon, SLW & NDA 11656; Navajo Mt., NDA 4149. Common. Hymenopappus fillifolius Hook. var. nu- dipes (Maguire) Turner Utah: Garfield Co., Pet Hollow, NDA 4330; Escalante Canyon, Beck 118; Esca- lante, Beck & Tanner s. n. Kane Co., East Clark Bench, Harrison 12071; one mile west of Glen Canyon City, Cronquist 10159; Hole-in-the-Rock, Karren 109; Paria Can- yon, Woodruff 1116; Cottonwood Wash road, NDA & RA 2773. San Juan Co., Rib- bon Canyon, Lake Powell, NDA 4111. Common. Hymenoxys acaulis (Fursh) Parker var. arizonica (Greene) Parker Utah: Garfield Co., 1.3 miles south of Cannonville, Reveal et al. 749; Bryce Can- yon, Cottam 4356; 10 miles south of Esca- lante, Harrison 9121. Kane Co., Cockscomb, NDA 4633; Kaiparowits Plateau, NDA & DT 5249; Paria Townsite, SLW & NDA 9740; Buckskin Gulch, SLW 5315; Nipple Bench, NDA & RA 2812. Common. Hymenoxys acaulis (Pursh) Parker var. ive- siana Greene Utah: Garfield Co., Escalante Canyon, McArthur & Beck s. n., June 1940. Kane Co., Four Mile Bench, SLW & JRM 12405; Little Valley Canyon, NDA 3523a; Dry Rock Creek, SLW & NDA 11620; Four Mile Bench, NDA 5173; Fifty Mile Spring, JRM 363; Kaiparowits Plateau, Harrison 9068. San Juan Co., Navajo Mt., Gentry & Davidse 1786; do, NDA 4150. Common. Hymenoxys cooperi (Gray) Cockerel! Utah: Garfield Co., 10 miles south of Es- calante, Harrison 9122; Escalante, Stanton 873. Kane Co., Left-hand fork of Collet Wash, Woodruff 1155; 12.7 miles southeast of Cannonville, Horse Mt., north of Four Mile Bench, SLW & JRM 12783. Locally common. Hymenoxys richardsonii (Hook.) Cockerel] Utah: Garfield Co., Bryce Canyon, Cot- tam 2756; do, Weight s. n., 1934. Uncom- mon. Iva axillaris Pursh Utah: Kane Co., Four Mile Bench, NDA 4317. Uncommon. Lactuca pulchella (Pursh) Riddel Utah: Garfield Co., Bryce Canyon, Weight s. n. Kane Co., Four Mile Creek, SLW et al. 12587. Uncommon. June 1978 Welsh et al.: Kaiparowits Flora 141 Lactuca scariola L. Prickly Lettuce Utah: Escalante Canyon, NDA 4118. Un- common; introduced weed. Layia glandulosa (Hook.) H. & A, Tidy Tips Arizona: Coconino Co., Cedar Mt., NDA & LCH 3825. Utah: Kane Co., Kane- Garfield county line, south of Escalante, NDA & DT 5299; Cedar Mt., NDA 4667. Uncommon. Lygodesmia grandiflora (Nutt.) T. & G. Rush Pink Utah: Garfield Co., Bryce Canyon, Cot- tam 2671. Kane Co., Paria River, NDA & LCH 3966; Dry Rock Creek, SLW & NDA 11618; Last Chance Bay, SLW & NDA 11599; Nipple Bench, NDA & RA 2811. Uncommon. Lygodesmia spinosa Nutt. Utah: Garfield Co., Escalante Canyon, Beck 146. Kane Co., 30 miles east of Ka- nab, NDA 5586; Four Mile Bench, NDA 4305. Uncommon. Machaeranthera canescens (Pursh) Gray Arizona: Coconino Co., between Page and Kaibito, Shaw 2643; Paria Plateau, NDA 6425. Utah: Kane Co., Cedar Mt., NDA 3130. Common in sandy sites. Machaeranthera glabriuscula (Nutt.) Cronq. & Keck var. confertifolia Cronq. Utah: Kane Co., Four Mile Bench, NDA 5172; Horse Mt., SLW 12819. Endemic, rare, and threatened (Welsh et al., 1975). Machaeranthera grindelioides (Nutt.) Shinners Utah: Garfield Co., Escalante, Cottam 4400; 8 miles southeast of Escalante, Holm- gren et al. 2029. Kane Co., 20 miles south of Cannonville, Christensen s. n., 8 May 1963; Collet Top, Woodruff 1153. Locally common. Machaeranthera linearis Greene Utah: Garfield Co., Escalante Canyon, McArthur & Beck 198. Kane Co., Reflection Canyon, Lake Powell SLW 11887. Uncom- mon. Machaeranthera tanacetifolia (H.B.K.) Nees Utah: Garfield Co., 20 miles southeast of Escalante, Holmgren et al. 2054. Kane Co., Cockscomb, SLW 5349; Paria River, JRM 323; Crosby Canyon, NDA 3633; 6 miles east of Glen Canyon City, SLW & NDA 9759; Sit Down Bench, Olsen 40; Paria Riv- er, NDA & LCH 4009; Tibbet Spring, NDA et al. 4272. Common. Machaeranthera tortifolia (T. & G.) Cronq. Arizona: Coconino Co., upstream from Lee's Ferry, NDA 4588. Utah: Kane Co., Smoky Mt., RA 199; 25 miles south of Esca- lante, Buss 80; Kaiparowits Plateau, Harri- son 9072; Cottonwood Wash Road, Cronquist 10208; Last Chance Bay, SLW & NDA 11600; Nipple Creek, RA 258; Little Valley Canyon, NDA 3535; 6 miles east of Glen Canyon City, SLW & NDA 9801; Wahweap Creek, NDA 3484. Common; es- pecially at lower elevations. Malacothrix glahrata Gray Arizona: Coconino Co., upstream from Lee's Ferry, NDA & LCH 4035. Utah: Kane Co., one mile west of Glen Canyon City, Cronquist 10146; Fifty Mile Spring, JRM 362; Little Valley Canyon, NDA 3524a; Warm Creek, NDA 3719. Wide- spread; common. Malacothrix sonchioides (Nutt.) T. & G. Arizona: Coconino Co., Page vicinity, SLW & NDA 9782. Utah: Kane Co., Wil- low Tank, Harrison 9101; Kodachrome Flat vicinity. Reveal et al. 800, 6 June 1967; Paria Canyon, Woodruff 1117; 13 miles west of Glen Canyon City, NDA & RA 2858; one mile west of Glen Canyon City, Cronquist 10147; 14 miles east of Glen Canyon City, RWA 198. Common. Oxytenia acerosa Nutt. Copperweed Utah: Kane Co., Tibbet Canyon, NDA & DK 3413; Four Mile Creek, SLW et al. 12584. Common along stream courses and seeps in saline sites. Pectis papposa (Harv.) Gray Chinchweed Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3351. Utah: Kane Co., 55 miles east of Kanab, SL & SLW 9419; 3 miles south of Glen Canyon City, NDA 3129; Escalante Canyon, NDA & RA 3213. Uncommon; widespread at lower elevations. Perezia wrightii Gray Utah: Kane Co., Nipple Bench, NDA & RA 2815; Kaiparowits Plateau, Woodruff & Meyer 1144; Dry Rock Creek, SLW & NDA 11641; Smoky Mt., NDA 5204. Along canyons; locally common. Petradoria pumila (Nutt.) Greene Utah: Kane Co., Four Mile Bench, NDA 142 Great Basin Naturalist Vol. 38, No. 2 4307. San Juan Co., Navajo Mt., NDA 4152. Widespread; locally common in rimrock sites. Pluchea sericea (Nutt.) Gov. Arizona: Coconino Co., Lee's Ferry, Cot- tam 2618; do, NDA & LCH 3815. Utah: Kane Co., Hole-in-the-Rock, Harrison 12115. Uncommon along the Colorado Riv- er. Psathyrotes pilifera Gray Arizona: Coconino Co., Lee's Ferry, Cot- tam 2630. Rare. Psilostrophe sparsiflora (Gray) A. Nels. Arizona: Coconino Co., Lee's Ferry, Cot- tam 2614; south base of Navajo Mt., NDA et al. 3006. Utah: Garfield Co., Escalante Canyon, Harrison 9133; Pet Hollow, NDA 4334; 8 miles southeast of Escalante, Holm- gren et al. 2021. Kane Co., Cottonwood Wash, NDA & RA 2776; Paria River, NDA & LCH 3953; Four Mile Bench, SLW & JRM 12406. Common, widespread. Senecio douglasii DG. var. longilobus (Benth.) Benson Utah: Kane Co., Paria River, Holmgren & Holmgren 4710; Cottonwood Wash Road, Reveal & Gentry 832; do, NDA & RA 2770. Uncommon. Senecio multilobatus T. & G. Utah: Kane Co., Four Mile Bench, NDA 3757; head of Last Chance Canyon, SLW & JRM 12784. Uncommon. Senecio spartioides T. & G. Utah: Garfield Co., Bryce Canyon, Weight s. n.; Escalante, Beck & Tanner s. n. Uncommon. Solidago canadensis L. Goldenrod Utah: Kane Co., Escalante Canyon, SLW & GM 11815; Reflection Canyon, SLW & CAT 11847. San Juan Co., Rainbow Bridge, NDA & NW 3304. Local and restricted; un- common. Solidago occidentalis (Nutt.) T. & G. Arizona: Coconino Co., Lee's Ferry, NDA 6390. Utah: Kane Co., Reflection Canyon, SLW & CAT 11861. San Juan Co., east of confluence of the San Juan, NDA & RA 3175. Uncommon. Solidago sparsiflora Gray Utah: Garfield Co., Pet Hollow, NDA 4333. Kane Co., Driftwood Canyon, SLW & GM 11781. San Juan Co., east of confluence of the San Juan, NDA & RA 3160. Hanging gardens; locally common. Solidago trinervata Greene Arizona: Coconino Co., Lee's Ferry, Clo- ver and Jotter (1941). The specimen has not been seen by the present writers, and the report requires confirmation. Sonchus asper (L.) Hill Utah: Kane Co., Escalante River, NDA & RA 3208. San Juan Co., Hanging garden, San Juan River, above the confluence, NDA 4099. Uncommon. Stephanomeria exigua Nutt. Wire Lettuce Utah: Kane Co., Spencer Flat, southeast of Escalante, NDA 483, 16 June 1965; Church Wells, JRM 328; Paria River, north of US Highway 89, NDA & LCH 3955; do, NDA & RA 2761; NDA et al. 2739. San Juan Co., Three Garden, SLW & NDA 11674. Common. Stephanomeria parryi Gray Utah: Kane Co., Nipple Bench, NDA & RA 2822a. Rare. Stephanomeria tenuifolia (Jones) Hall Arizona: Coconino Co., Colorado River channel, north of Navajo Canyon, NDA & RA 3141. Utah: Kane Co., 3 miles south of Glen Canyon City, NDA 3134; Cottonwood Wash, 12 miles north of Highway 89, NDA & KD 3322; Head of left fork of Last Chance Canyon, NDA et al. 2701. Com- mon; rock outcrops and talus slopes. Stylocline micropoides Gray Arizona: Coconino Co., Lee's Ferry, NDA 4568. Utah: San Juan Co., Ribbon Canyon, SLW & NDA 11713. Rare. Taraxacum officinale (L.) Wiggers Dandelion Utah: Kane Co., Nipple Canyon, RWA 246. Introduced, ubiquitous weed. Tetradymia canescens DG. Utah: Kane Co., West side of Cockscomb, NDA 2924; Four Mile Bench, NDA 4068; Buckskin Gulch, NDA 6387. Locally com- mon. Thelysperma subnudum Gray Utah: Garfield Co., 8 miles southeast of Escalante, Holmgren et al. 2026. Kane Co., Nipple Bench, NDA & RA 2810. Uncom- mon. Townsendia annua Beaman Arizona: Lee's Ferry, NDA 4578. Utah: June 1978 Welsh et al.: Kaiparowits Flora 143 Kane Co., Tibbet Canyon, NDA 3605; Tib- bet Bench, NDA & DT 5059; Wahweap Bay, NDA 3552; Grand Bench, NDA 4260. Ca 15 miles west of Glen Canyon City, NDA & T. A. 2870. Common, low eleva- tions. Toumsendia incana Nutt. Utah: Garfield Co., Escalante, Cottam 4402; Ca 8 miles southeast of Escalante, Holmgren et al. 3020; Ca 20 miles southeast of Escalante, Holmgren et al. 2048. Kane Co., Wahweap Creek, Harrison 12087; Kai- parowits Plateau, NDA & DT 5271; Paria River Bridge, SWL & NDA 9753; Paria River, NDA & RA 2777; Glen Canyon City, Cronquist 10156; Cockscomb, SLW 5345; Paria River, NDA & LC Higgins 3968; Paria River, Woodruff 1110; 4 miles south of Cannonville, Cronquist 10082; East Clark Bench, Harrison 12300; Cottonwood Wash, NDA 3589; Cedar Mt., NDA 3114. Abun- dant; widespread. Toumsendia exscapa (Richards.) Porter Arizona: Coconino Co., Buckskin Mt., NDA 4943. Utah: Garfield Co., Bryce Can- yon, Buchanan 1481 (Bryce). Uncommon. Toumsendia minima Eastw. Utah: Garfield Co., Bryce Canyon, Cot- tam 5672. Endemic, rare, and possibly threatened (Welsh et al, 1975). Vanclevea stylosa (Eastev.) Greene Arizona: Coconino Co., 9.3 miles south of Page, NDA et al., 3339; Colorado River Canyon, north of Navajo Canyon, NDA & RA 3144. Utah: Kane Co., 6 miles east of Glen Canyon City, NDA & DK 3355; Fifty Mile Spring, JRM 375; 2 miles south of Glen Canyon City, NDA 3095; East Clark Bench, SL & SLW 9418; Cedar Mt., NDA 6009 A. Abimdant on sandy sites at lower elevations. Verbesina encelioides (Gov.) Benth. & Hook Utah: Garfield Co., Bryce Canyon, Weight B-32/343 (Bryce). Uncommon. Viguiera annua (Jones) Blake Utah: Kane Co., Tropic shale, east of Paria River, NDA & LCH 3993; do, NDA & DK 3320; Cottonwood Canyon, SL & SLW 9416. Uncommon. Viguiera soliceps Bameby Utah: Kane Co., Tropic shale, 4 miles north of Highway 89 along Cottonwood Wash Road, NDA & RA 2759; 20 miles east of Glen Canyon City, NDA & DT 5073; mouth of Paria Canyon, Cottonwood Wash Road, SLW 11056; do, Reveal & Gentry 833. Note: This plant is considered as en- dangered (Welsh et al. 1975). It is endemic to the Tropic Shale formation. Wyethia scabra Hook. Utah: Kane Co., Willow Tank, Beck & Tanner s. n.; Hall Cave, 50 miles south of Escalante, Beck & Tanner s. n.; East Clark Bench, NDA & RA 2684. San Juan Co., Rainbow Bridge, SLW & NDA s. n. Wide- spread, uncommon. Xanthium saccharatum Wallr. Utah: San Juan Co., Three Garden NDA & RA 3187, 23 Aug. 1971; do, CAT & SLW 166. Locally common. Convolvulaceae (Morning Glory Family) Convolvulus arvensis L. Utah: Kane Co., 15 miles south of Can- nonville, NDA & RA 2911. Uncommon. Evolvulus nuttallianus R. & S. Utah: Kane Co., Paria River, NDA & LCH 3962; East Clark Bench, Harrison 12299; do, Cronquist, 10174; do, NDA & RA 2669. Uncommon. Crassulaceae (Stonecrop Family) Sedum lanceolatum Torr. Utah: San Juan Co., Navajo Mt., NDA et al. 2997; do, NDA 4184. Uncommon. Cruciferae (Mustard Family) Arabis fendleri (Wats.) Greene Arizona: Coconino Co., Buckskin Mt., NDA 4941. Utah: San Juan Co., Navajo Mt., NDA 4179. Common. Arabis holboellii Hornem. Utah: Kane Co., Kaiparowits Plateau, Harrison 9056. Uncommon. Arabis lignifera A. Nels. Utah: Kane Co., Four Mile Bench, NDA 5167; west of Cockscomb, NDA 4611. Un- common. Arabis pendulina Greene Utah: Garfield Co., Four miles southwest 144 Great Basin Naturalist Vol. 38, No. 2 of Cannonville, Reveal et al. 776. Kane Co., Four miles south of Cannonville, Cronquist 10080. Uncommon. Arabis perennans Wats. Utah: Kane Co., Kaiparowits Plateau, Harrison 9054; 29 miles south of Escalante, 10031; Five Mile Mt., NDA 4641; Buckskin Mt., NDA 4653; 4 miles south of Cannon- ville, Cronquist 10079. Common. Arabis pulchra Jones. Utah: Kane Co., Smoky Mt., Cronquist 10019; Nipple Bench, NDA 3482; Cock- scomb, NDA 4635. Common. Arabis selbyi Rydb. Utah: Garfield Co., About 4 miles south- west of Cannonville, Reveal et al. 775. Un- common. Brassica nigra (L.) Koch. Black Mustard Arizona: Coconino Co., along U.S. High- way 89, NDA 5197. Uncommon. Capsella bursa-pastoris (L.) Medic Shepherds Purse Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5236. Uncommon. Caulanthus crassicaulis (Torr.) Wats. Utah: Garfield Co., 1.3 miles south of Cannonville, Reveal et al. 760, Kaiparowits Plateau, NDA & DT 5282; Cockscomb, NDA 4612; Last Chance Creek, NDA 3745; Four Mile Bench, NDA 4065; 25 miles southeast of Escalante, Holmgren et al. 2072; Buckskin Gulch, SLW 5313; Brigham Plains, NDA & RA 2784. Common on benchlands. Chorispora tenella (Pall.) DC. Utah: Kane Co., Cockscomb, NDA 3629. Introduced weed, uncommon. Descurainia obtusa Greene Utah: Kane Co., 4 miles below Last Chance Canyon, Gaines (1960). Descurainia pinnata (Walt.) Britt. Arizona: Coconino Co., west side of Cockscomb, NDA 3055. Utah: Kane Co., Little Valley, NDA et al. 2722; Kaiparowits Plateau, Harrison 9044; do, NDA & DT 5287; Smoky Mt. RWA 129; Cedar Mt., RWA 339, 352; Smoky Mt. NDA 3441; Cockscomb, NDA 3588; San Juan Co., Three Garden, SLW 11904. Common to abundant; widespread. Dithyrea wislizenii Engelm. Arizona: Coconino Co., Colorado River between Lee's Ferry and Glen Canyon Dam, NDA 4590. Utah: Kane Co., Wah- weap Creek, NDA & DT 3421; Paria River, NDA & LCH 3949; Warm Creek, RWA 114; 2 miles west of Glen Canyon City, Nebeker & Skougard 184; 5 miles W. of Glen Canyon City, NDA 3115; 13 miles west of Glen Canyon City, NDA & RA 2857; Cockscomb, NDA 3579; V2 mile west of Paria River Bridge, SLW & NDA 9755. Locally common, in sandy sites. Draba cuneifolia Nutt. Utah: Kane Co., Four Mile Bench, NDA 3486; Five Mile Mt., NDA 4412; Kaiparo- wits Plateau, NDA & DT 5272. Uncommon; widespread at lower elevations. Draba subalpina Goodman & Hitchc. Utah: Garfield Co., Bryce Canyon, Bu- chanan 2073, (Bryce); do SLW 12724; do SLW 12808. Rare; threatened (Welsh et al. 1975). Erysimum asperum Nutt. Arizona: Coconino Co., Arizona, 3 miles south of Wahweap Marina, SLW & NDA 9760. Utah: Garfield Co., Bryce Canyon, Boyle 1107; do, Burkey 126. Kane Co., one mile south of Glen Canyon City, NDA 3432; Five Mile Mt., NDA 4683; Kaiparo- wits Plateau, NDA & DT 5234; Cockscomb, NDA 3590; Cedar Mt., NDA 3432; Four Mile Bench, SLW & JRM 12407; Cocks- comb, NDA 3541; 14 miles east of Glen Canyon City, RWA 168. Common. Hutchinsia procumbens (L.) Desv. Utah: San Juan Co., Ribbon Garden, SLW & NDA 11705; Three Garden, SLW 11907. Uncommon. Lepidium densiflorum Schrad. Arizona: Coconino Co., Colorado River, between Lee's Ferry and Glen Canyon Dam, NDA 4585. Utah: Kane Co., Little Valley, NDA 3523; Kaiparowits Plateau, NDA & D. Trotter 4272; Cockscomb, NDA 3587. Common. Lepidium montanum Nutt. var. alyssioides (Gray) Jones Utah: Kane Co., Kodachrome Flats vicin- ity. Reveal et al. 789. Uncommon. Lepidium montanum Nutt. var. glabrum C. L. Hitchc. Utah: Kane Co., Paria River Canyon, JRM M106; Tibbet Spring, RWA 112; June 1978 Welsh et al.: Kaiparowits Flora 145 Smoky Mt., RWA 167; Fifty Mile Spring, JRM 368; Cedar Mt., NDA et al. 1746. Common and widespread. Lepidium tnontanutn Nutt. var. integri- folium (Nutt.) C. L. Hitchc. Arizona: Coconino Co., Lee's Ferry, NDA 6391. Uncommon. Lepidium montanum Nutt. var. jonesii (Rydb.) C. L. Hitchc. Arizona: Coconino Co., Cedar Mt., RWA 170. Utah: Kane Co., 20 miles south of Cannon ville, Christensen s.n.; Willow Tank, Harrison 9028; do. White 112; Tibbet Can- yon, NDA 3444; Nipple Canyon, RWA 244; Collet Canyon, Woodruff 1159; Paria River, NDA & LCH 3975. Common and wide- spread. Lepidium perfoliatum L. Utah: Kane Co., Nipple Rench vicinity, LCH & NDA 5244; Cockscomb, NDA 3617. Uncommon, introduced weed. Lesquerella intermedia (Wats.) Heller Arizona: Coconino Co., Cockscomb, NDA et al. 3008. Uncommon. Lesquerella kingii Wats. ssp. lati folia (A. Nels.) Rollins & Shaw Utah: Garfield Co., Canyon above Trop- ic, Jones 5312d (US); Bryce Canyon, Weight s.n., 1932 (US). Lesquerella ludoviciana (Nutt.) S. Wats. Utah: Garfield Co., Ca 6 miles southeast of Escalante, Karren 121. Uncommon. Lesquerella rectipes Woot. & Standi. Arizona: Coconino Co.; Cockscomb, NDA & RA 2887; do, NDA 3064. Utah: Kane Co., Last Chance Creek, NDA 3749; Cock- scomb, NDA 3580. San Juan Co., Navajo Mt., NDA 4139. Common. Lesquerella rubicundula Rollins. Utah: Garfield Co., Bryce Canyon, ob- served but not collected (see Reveal 1970); rare and threatened (Welsh et al., 1975). Lesquerella tumulosa (Bameby) Reveal Utah: Kane Co., 6.5 miles southeast of Cannonville, SLW 12765. Endemic, rare, and threatened (Welsh et al., 1975). Lesquerella wardii Wats. Utah: Kane Co., Fouj Mile Bench, SLW & NDA 12379. Uncommon. Malcolmia africana (L.) R. Br. Arizona: Coconino Co., Colorado River Canyon, between Glen Canyon Dam and Lee's Ferry, NDA & LCH 4029. Utah: Kane Co., Paria River, NDA & LCH 3963. Uncommon to very common introduced weed. Nasturtium officinale R. Br. Arizona: Coconino Co., Paria primitive area. Woodruff 1133. Rare; watercress. Physaria chambersii Rollins Utah: Kane Co., Last Chance Creek, NDA 3748; Horse Mt., SLW & NDA 12373; Paria River, NDA & LCH 3980; Warm Creek RWA 116; Brigham Plains NDA et al. 3490; Buckskin Gulch 5318. Common. Physaria newberryi Gray Utah: Garfield Co., Bryce Canyon, Bur- key 138; 6 miles north of Escalante, White 145. Kane Co., Kaiparowits Plateau, JRM 403. Common. Schoencrambe linifolia (Nutt.) Greene Utah: Kane Co., Smoky Mt., Cronquist 10018; Paria Townsite, SLW & NDA 9734; Last Chance Creek, NDA et al. 3514; Kai- parowits Plateau, Olson 41; do, NDA & DT 5238; Smoky Mt., NDA 3441; do, RWA 132; Cockscomb, NDA 4623. Common; widespread. Sisymbrium altissimum L. Utah: Kane Co., Buckskin Mt., NDA 4651. Uncommon. Stanleya pinnata (Pursh) Britt. Utah: Garfield Co., 6 miles southeast of Escalante, NDA 466; Escalante, Cottam 4422. Kane Co., 50 miles southeast of Esca- lante, Beck s.n.; Paria Townsite, SLW & NDA 9722; 31 miles east of Glen Canyon City, NDA et al. 2696; Kaiparowits Plateau, Woodruff 1146. San Juan Co., San Juan River Canyon, east of the confluence, NDA 4087. Common selenophyte; widespread. Streptanthella longirostris (Wats.) Rydb. Arizona: Coconino Co., Colorado River between Lee's Ferry and Glen Canyon Dam, NDA 4592; 4 miles southeast of Page, RWA 235. Utah: Kane Co., Warm Creek SLW & NDA 9798; Nipple Bench vicinity, Cronquist 10015; East Clark Bench, Harri- son 12074; 4 miles east of Glen Canyon City, RWA 118; Smoky Mt., RWA 130; Willow Tank, Harrison 9091; 2 miles south of Cannonville, Reveal et al. 779. Common; widespread. 146 Great Basin Naturalist Vol. 38, No. 2 Streptanthus cordatus Nutt. Utah: Kane Co., Grand Bench, NDA 3533; Kaiparowits Plateau, NDA & DT 5281; Five Mile Mt., NDA 4640. Common. Thelypodium integrifolium (Nutt.) Endl. Utah: Kane Co., head of Last Chance Canyon, SLW et al. 12597; Lake Powell, NDA 3137. San Juan Co., San Juan River, east of the confluence, NDA & RA 3150. Uncommon. Thelypodium sagittatum (Nutt.) Endl. var. ovalifolium (Rydb.) Welsh & Reveal Utah: Garfield Co., Bryce Canyon, East- wood & Howell 766 (CAS), fide Al-shebaz (1973). Uncommon. Thlaspi arvense L. Utah: Kane Co., Bryce Canyon, NDA 6125. Introduced weed; uncommon. Thlaspi montanutn L. Utah: Garfield Co., Death Ridge, NDA 5183. High elevations; uncommon. Cupressaceae (Cypress Family) Juniperus communis L. Utah: Garfield Co., Bryce Canyon, Bur- key 128, San Juan Co., Navajo Mt., NDA 4181. Uncommon; higher elevations. Juniperus osteosperma (Torr.) Little Arizona: Coconino Co., 8 miles southeast of Page, NDA et al. 3246. Utah: Garfield Co., Escalante, Beck & Tanner s.n. Kane Co., Kaiparowits Plateau, Harrison 9079; 6 miles east from Glen Canyon City, SLW & NDA 9806; Four Mile Bench, SLW & ERW 12368; Nipple Canyon, RWA 259. San Juan Co., Navajo Mt. NDA et al. 3004; Ribbon Garden, SLW & NDA 11708. Com- mon at middle elevations. Juniperus scopulorum Sarg. Utah: Kane C, Canaan Peak vicinity, common but not collected; San Juan Co., Navajo Mt., NDA et al. 2981. Middle eleva- tions. CuscuTACEAE (Doddcr Family) Cuscuta cephalanthii Engelm. Utah: Kane Co., Hole-in-the-Rock, Beck 138. Rare. Cyperaceae (Sedge Family) Carex curatorum Stacey Utah: San Juan Co., Three Garden, SLW 12425. New Utah Record; rare and threat- ened (Welsh et al., 1975). Carex douglasii Britt. Utah: Garfield Co., Bryce Canyon, Bu- chanan 66. (Bryce). Uncommon. Carex eleocharis Bailey Utah: Garfield Co., Bryce Canyon, Bur- key 144. Uncommon. Carex garberi Fern. Utah: Kane Co., Little Valley Canyon, SLW & NDA 11613; San Juan Co., Ribbon Garden, SLW & NDA 11714. Local in hanging gardens. Carex kelloggii Boott Utah: San Juan Co., Mystery Canyon, Gaines (1960). Carex haydeniana Olney Utah: Garfield Co., Escalante Canyon, Cottam 4426. Uncommon. Carex lanuginosa Michx. Utah: Kane Co., Hole-in-the-Rock, Harri- son 12121. Uncommon. Carex nebraskensis Dewey Utah: Garfield Co., Escalante Canyon, Cottam 4377. Uncommon Carex rossii Boott. Utah: San Juan Co., Navajo Mt., NDA 4161. Uncommon. Carex scirpiformis Mack. Utah: Kane Co., Hd. Mill Fk., east fork of Sevier River, Lewis s.n. Uncommon. Carex vesicaria L. Utah: San Juan Co., Escalante Canyon, NDA 4124. Uncommon. Cladium californicum (Wats.) O'Neill Utah: Kane Co., Driftwood Canyon, SLW & NDA 11709; do, SLW & CM 11780; Hidden Passage Canyon, SLW & CAT, 11870; San Juan Co., Utah, Wilson Creek, SLW & CAT 11874. Rare and threatened (Welsh et al, 1975). Cyperus schwenitzii Torr. Arizona: Coconino Co., Cockscomb, NDA et al. 3009a, do, NDA, 3070. Uncommon. Eleocharis palustris (L.) R. & S. Utah: Garfield Co., Escalante River, Beck s.n. Kane Co., Cottonwood Wash, Reveal et al. 820; Willow Tank, Beck s.n. Locally abundant. June 1978 Welsh et al.: Kaiparowits Flora 147 Eleocharis rostellata Torr. Utah: San Juan Co., Three Garden, SLW & NDA 11669; do, SLW 112437. Local and uncommon. Fimbristylis puberula (Michx.) Vahl Utah: Kane Co., north of Iceberg Can- yon, NDA & NW 3311. Uncommon. Fimbristylis thermalis Wats. Utah: Kane Co., north of Iceberg Can- yon, NDA & NW 3311; Long Garden, Es- calante River, SLW & CAT 11873. Rare. Uncommon. Scirpus acutus Muhl. Utah: Kane Co., Paradise Canyon, Welsh et al. 12601, Escalante Canyon, NDA 4223; first hanging garden mouth of Escalante River, SLW & GM 11814. Uncommon. Scirpus americanus Pers. Arizona: Coconino Co., Paria River, NDA 4008; do. Utah: Kane Co., Ca .6 miles south of Nipple Spring, NDA & RA 2831; do. Four Mile Bench, NDA 4310; Paradise Can- yon, SLW et al. 12595; Willow Tank, Har- rison 9105; Cottonwood Wash Spring, Re- veal et al. 822. Scirpus paludosus A. Nels. Utah: Kane Co., Tibbet Spring, NDA & Kaneko 3415. Rare. Elaeagnaceae (Oleaster Family) Elaeagnus augustifolia L. Russian Olive Utah: Kane Co., 16 miles north of gravel pit in Wahweap Creek, NDA 3471. In- troduced weedy tree. Shepherdia argentea (Pursh) Nutt. Silver Buffaloberry Utah: Garfield Co., 3 miles north of Esca- lante, Maguire s.n., Kane Co., Head of Last Chance Canyon, NDA et al. 3508. Uncom- mon. Shepherdia canadensis (L.) Nutt. Utah: Garfield Co., Bryce Canyon, Weight B-32/256. Rare. Shepherdia rotundifolia Parry Utah: Garfield Co., Escalante, Cottam 4374; do. Beck & Tanner s.n. Kane Co., 50 miles south of Escalante, Beck & Tanner s.n.; 32 miles south of Escalante, Cronquist 10029; 6 miles southeast of Canaan Peak, SLW & NDA 12370; Paria Townsite, Pat- ton 124; do, SLW & NDA 9723; Kaiparo- wits Plateau, JRM 397; do, Harrison 9078; do, NDA & DT 5294; 20 miles south of Cannonville, Christensen s.n., Paria River, NDA & LCH 4002; 31.5 miles east of Glen Canyon City, NDA et al. 2704; 10 miles southeast of Cannonville, NDA et al. 3505b; 8 miles south of Paria, Harrison 12048. Abundant to common; widespread. Ephedraceae (Jointfir Family) Ephedra cutleri Peebles Arizona: Coconino Co., one mile east of Page, NDA & LCH 4019; 4 miles southeast of Page, RWA 215. Utah: Kane Co., Last Chance Bay, Lake Powell, SLW & NDA 11607, 20 miles east of Glen Canyon City, NDA 4061; 7 miles southeast of Glen Can- yon City, RWA 227; one mile west of Glen Canyon Dam, NDA 3573. Ephedra torreyana Wats. Arizona: Coconino Co., Colorado river Canyon, between Glen Canyon Dam and Lee's Ferry, NDA & LCH 4041. Utah: Gar- field Co., 8 miles southeast of Escalante, Holmgren et al. 2037. Kane Co., Wahweap Creek, Harrison 12100; Warm Creek vicin- ity. Lake Powell, NDA 3721; East Clark Bench, NDA & RA 2689; Gunsight Bay, Lake Powell, SLW & NDA; Paria River, NDA & LCH 4016, Hole-in-the-Rock, Har- rison 12141. Widespread; common. Ephedra viridis Coville Green Mormon-Tea Utah: Garfield Co., Escalante, Cottam 4380; 28 miles southeast of Escalante, Har- rison 9119. Kane Co., Willow Tank, Paria Townsite, Olson 27; 4^2 miles south of Glen Canyon City, RWA 331; Fifty Mile Spring, JRM 365; Cedar Mt., NDA et al. 2747a. San Juan Co., Ribbon Garden, SLW & NDA 11697. Abimdant; widespread. Equisetaceae (Horsetail Family) Equisetum arvense L. Utah: Garfield Co., Escalante Canyon, Beck, s.n., Kane Co., Paria River, NDA & LCH 4012; Hackberry Canyon, NDA 3686, 3697. Uncommon. Equisetum hyemale L. Utah: San Juan Co., Rainbow Bridge, SLW & NDA 11660. Uncommon. 148 Great Basin Naturalist Vol. 38, No. 2 Ericaceae (Heather Family) Arctostaphylos patula Greene Manzanita Utah: Garfield Co., Bryce Canyon, White 151; do, Burkey 127; Cockscomb, NDA 3538; do, NDA 3596; Kaiparowits Plateau; Harrison 9082. Local; uncommon. EuPHORBiACEAE (Spurge Family) Croton texensis (Klotzsch) Meuell. Arg. Arizona: Coconino Co., Paria Plateau, NDA 6411. Utah: Kane Co., 6 miles east of Glen Canyon City, NDA & DK 3359; Glen Canyon City vicinity, NDA 2915; Halls Creek Bay, Lake Powell, SLW & CM 11795; Warm Creek Bay, NDA, 3288a; Paria Primitive Area, NDA & RA 2860. Common; sandy sites at low elevations. Euphorbia albomarginata T. & G. Utah: Garfield Co., Escalante Canyon, Harrison 9124. Kane Co., Cedar Mt., RWA 441. Uncommon. Euphorbia fendleri T. & G. Arizona: Coconino Co., mile 6 A on Lake Powell, NDA 3700. Utah: Garfield Co., 8 miles southeast of Escalante, Holmgren et al. 2020; Escalante, Beck & Tanner s.n. Kane Co., Willow Tank, Beck & Tanner s.n.; Ahlstrom Point road, NDA & RA 2894; Kaiparowits Plateau, Woodruff 1148; head of Last Chance Canyon, Cronquist 10027; 4 miles south of Cannonville, Reveal et al. 786. Common. Euphorbia glyptosperma Englem. Arizona: Coconino Co., Paria Plateau, NDA & CM 4278. Utah: Kane Co., Cedar Mt., NDA 3100. Uncommon. Euphorbia nephradenia Barneby Utah: Kane Co., Cottonwood Wash Road, 5.4 miles north of U.S. 89, Reveal and Gen- try 834; Paria Primitive Area, NDA & RA 2865; 6 miles south of confluence of Cot- tonwood Canyon, SLW 11059. Endemic, rare, and threatened (Welsh et al., 1975). Euphorbia robusta (Englem.) Small Utah: Kane Co., Kaiparowits Plateau, Harrison 9037a, 9042; do, NDA & DT 5231. Uncommon. Reverchonia arenaria Gray. Arizona: Coconino Co., Paria Plateau, NDA 6426. Utah: Kane Co., Cedar Mt., NDA 3128; Glen Canyon City vicinity, NDA 2920. Rare. Fag ACE AE (Beech Family) Quercus gambelii Nutt. Arizona: Coconino Co., Paria Plateau, NDA & CM 4293, Utah: Kane Co., Hack- berry Canyon, NDA 3689; Kaiparowits Plateau, NDA & DT 5267; Escalante River, SLW & CM 11820; Hole-in-the-Rock, Har- rison 12732; do, Harrison 12134; Reflection Canyon, SLW & CAT 11854; Escalante River, NDA 4119. San Juan Co., confluence of San Juan Canyon, NDA 4093; Three Garden, SLW & NDA 11684. Common in canyons, and on higher mountain slopes. Quercus turbinella Greene Arizona: Coconino Co., Paria Plateau, NDA & G. Moore 4398. Utah: San Juan Co., Three Garden, SLW & NDA 11684 A. Rare. Quercus undulata Torr. Utah: Kane Co., Dance Hall Rock, JRM 381, 382; 50 miles south of Escalante, Beck & Tanner s.n.; Willow Tank, Beck & Tan- ner s.n. do, Harrison 9087; Escalante Can- yon, SLW & CM 11825, 11826; do, NDA 4220; 60 miles south of Escalante, Harrison 12723; Hole-in-the-Rock, White 125; Cocks- comb, NDA 3544; do, SLW 5334. Common. FuMARiACEAE (Fumitory Family) Corydalis aurea Willd. Arizona: Coconino Co., Paria Plateau, NDA & LCH 3774. Utah: Kane Co., Brig- ham Plains, NDA et al. 2489. San Juan Co., Navajo Mt., NDA 4138. Widespread; un- common. Gentianaceae (Gentian Family) Centaurium exaltatum (Griseb.) Wight Utah: Kane Co., head of Last Chance Canyon, SLW et al. 12593; San Juan Co., Three Garden, SLW & CAT 11866. Rare. Frasera albomarginata Wats. Utah: Garfield Co., Escalante, Cottam 4403. Kane Co., 45 miles east of Glen Can- ■ yon City, NDA et al. 2728; Four Mile | Bench, NDA 4070; Brigham Plains, NDA & June 1978 Welsh et al.: Kaiparowits Flora 149 RA 2786; 25 miles southeast of Escalante, Holmgren et al. 2063. Uncommon. Frasera paniculata Torr. Utah: Kane Co., East Clark Bench, NDA & RA 2687; do, NDA & RA 2876. Rare. Gentiana affinis Griseb. Utah: Kane Co., Wahweap Creek, Four Mile Bench, SLW 12572. Rare. Gentianella tortuosa (Jones) Gillett Utah: Garfield Co., Bryce Canyon, Cot- tam 2765; do Weight s.n. Rare. Geraniaceae (Geranium Family) Erodium cicutarium (L.L) L'Her. Storksbill Utah: Kane Co., Cockscomb, NDA 3628; Little Valley, NDA 3431; Kaiparowits Plateau, NDA & DT 5274. Introduced weed; widespread and locally abundant. Erodium texanum Gray Arizona: Coconino Co., Lee's Ferry, Cot- tam 10004. Geranium marginale Rydb. Utah: Garfield Co., Bryce Canyon, Bu- chanan 1716 (Bryce). Rare and possibly threatened (Welsh et al., 1975). Geranium richardsonii Fisch & Trautv. Utah: Garfield Co., Escalante Canyon, VanCott 982. Kane Co., between Death Ridge & Canaan Peak, SLW et al. 12590. Uncommon. Gramineae (Grass Family) Agropyron caninum (L.) Beauv. var. andi- num (Scribn. & Sm.) C.L. Hitchc. Utah: Garfield Co., 15 miles southeast of Escalante, Hall 13569. Uncommon. Agropyron caninum (L.) Beauv. var. lati- glume (Scribn. & Sm.) C.L. Hitchc. Utah: Garfield Co., 15 miles southwest of Escalante, Hall 13604; Bryce Canyon, Boyle 1141. Uncommon. Agropyron cristatum (L.) Gaertn. Crested Wheatgrass Utah: Garfield Co., 15 miles southwest of Escalante, Hall 13571. Introduced forage grass. Agropyron dasystachyum (Hook.) Scribn. Utah: Garfield Co., Escalante Canyon, Beck s.n. Uncommon. Agropyron intermedium (Host.) Beauv. Utah: Kane Co., east of Kanab; NDA 3279 A. Introduced forage grass. Agropyron repens (L.) Beauv. Quackgrass Utah: Kane Co., Last Chance Canyon, east of Four Mile Bench, SLW et al. 12595. Uncommon. Agropyron smithii Rydb. Western Wheatgrass Utah: Garfield Co., 15 miles southwest of Escalante, Hall 13575; Bryce Canyon, Boyle 1142; do, Cottam & Hutchings 2773. Com- mon at middle elevations. Agropyron spicatum (Pursh) Scribn. & Sim. Utah: Garfield Co., 15 miles southeast of Escalante, Hall 13561. Kane Co., Kaiparo- wits Plateau, NDA & DT 5251. Uncommon. Agrostis alba L. Utah: Garfield Co., 15 miles southwest of Escalante, Hall 13560. Kane Co., Four Mile Bench, SLW et al. 12588; do, NDA 4313. Locally abundant. Agrostis semiverticillata (Forsk.) C. Christ. Utah: Kane Co., Hole-in-the-Rock, Harri- son, 12136; Escalante River, SLW & CM 11823; do, NDA 4120, 4130. San Juan Co., Ribbon Canyon, NDA 4104. Widespread; canyon bottoms. Andropogon barbinodis Lag. Utah: Kane Co., Escalante River, NDA 4219; 2 miles north of confluence of San Juan and Colorado canyons, SL & SLW 11881. San Juan Co., Rainbow Bridge, NDA 4356; east of San Juan confluence, NDA & RA 3165. Uncommon. Andropogon glomeratus (Walt.) B. S. P. Utah: Kane Co., Hole-in-the-Rock, Harri- son 12131; Reflection Canyon, SLW 11889; east of San Juan confluence, NDA & RA 3161. Rare and threatened (Welsh et al, 1975). Andropogon hallii Hack. Utah: Kane Co., Reflection Canyon, ob- served but not collected (SLW). Uncom- mon. Aristida adscensionis L. Arizona: Coconino Co., Cockscomb NDA & DK 3381. Uncommon. Aristida arizonica Vasey Utah: Kane Co., NDA 3131. Rare. Aristida fendleriana Steud. Utah: Kane Co., Escalante River, NDA & RA 3201; Dry Rock Creek, Lake Powell, 150 Great Basin Naturalist Vol. 38, No. 2 SLW & NDA 11637; Willow Tank, Harri- son 9110; Cottonwood Wash, NDA & RA 2774. San Juan Co., east of San Juan con- fluence, NDA et al. 3227. Common. Aristida glauca (Nees) Walt. Utah: Garfield Co., Escalante Canyon, Harrison 9130. Kane Co., 35 miles southeast of Escalante, Harrison 9116; Gunsight Bay, Lake Powell, SLW & NDA 11596; Rainbow Bridge SLW & NDA 11654; Hole-in-the- Rock, Harrison 12143; Willow Tank, Beck & Tanner s.n. San Juan Co., east of the San Juan confluence, NDA 4095; confluence of the San Juan, SLW 11935. Common. Aristida longiseta Steud. Arizona: Coconino Co., Colorado River Canyon north of Navajo Canyon, NDA & RA 3140. Utah: Garfield Co., Escalante Desert, Beck s.n. Kane Co., Dry Rock Creek, Lake Powell, SLW & NDA 11622; Cedar Mt., NDA 3127; Cockscomb, JRM 335. Common. Bouteloua aristidoides (H.B.K.) Girseb. Utah: Kane Co., Cedar Mt., NDA et al. 3254. Rare. Bouteloua barbata Lag. Arizona: Coconino Co., Navajo Canyon, NDA & RA 3221. Utah: Kane Co., East Clark Bench, NDA & DK 3373. Locally common. Bouteloua curtipendula (Michx.) Terr. Arizona: Coconino Co., Colorado River Canyon, north of Navajo Canyon, NDA & RA 3138. Utah: Kane Co., 6 miles east of Glen Canyon City, NDA & DK 3354; Dry Rock Creek, Lake Powell, SLW & NDA 11621. San Juan Co., east of San Juan con- fluence, NDA 4269; San Juan confluence, SLW 11934. Uncommon. Bouteloua eriopoda Torr. Arizona: Coconino Co., Colorado River Canyon, north of Navajo Canyon, NDA & RA 3139; Paria Plateau, NDA 6423. Utah: Kane Co., Cedar Mt., NDA et al. 32521 do, NDA 3104, do, NDA et al. 2961; Nipple Bench, NDA & DK 3327; East Clark Bench, NDA & DT 3363. Common at lower elevations. Bouteloua gracilis (H.B.K.) Lag. Arizona: Coconino Co., 6 miles south of Page, NDA et al. 3238; Paria Plateau, NDA 6422; Cockscomb, NDA & DK 3385. Utah: Kane Co., Cedar Mt., RWA 179; do, NDA et al. 3251; do, NDA 3133. Common. Bromus anomalus Rupr. Utah: Garfield Co., Bryce Canyon, Bu- chanan 54; do, Weight s.n. Uncommon. Bromus carinatus Hook. & Am. Utah: Kane Co., Paria River, LCH & NDA 5393; do, NDA 4004. Uncommon. Bromus inermis Leyss. Utah: Garfield Co., Bryce Canyon, Boyle 1130. Introduced forage grass. Bromus marginatus Nees Utah: Garfield Co., Escalante Canyon vi- cinity, Harrison 9147. Kane Co., Cotton- wood Wash, NDA 3688. Uncommon. Bromus porteri (Coulter) Nash Utah: Garfield Co., 15 miles southwest of Escalante, Hall 13584; Bryce Canyon, Weight s.n.; do, Boyle 1139. Uncommon. Bromus rigidus Roth Utah: San Juan Co., Rainbow Bridge, SLW & NDA 11655. Uncommon. Bromus rubens L. Arizona: Coconino Co., Colorado Canyon, between Lee's Ferry and Glen Canyon Dam, NDA 4594. Utah: Kane Co., Wah- weap Creek, Harrison 12097; Little Valley Canyon, NDA et al. 2716; Fifty Mile Spring, JRM 374; Kaiparowits Plateau, Bingham 32. San Juan Co., Three Garden, SLW et al. s.n. Warm Creek, Olson 31; Dry Rock Creek, Lake Powell, SLW & NDA 11629. Abundant. Bromus tectorum L. Arizona: Coconino Co., Colorado Canyon, between Lee's Ferry and Glen Canyon Dam, NDA & LCH 4040. Utah: Kane Co., 50 miles south of Escalante, Beck & Tanner s.n.; Paria River, NDA & LCH 3995; Little Valley Canyon, NDA & SLW 2717A; Kai- parowits Plateau, NDA & DT 5255. San Juan Co., Rainbow Bridge, SLW & NDA 11651; Navajo Mt., NDA 4147. Abundant. Calamagrostis scopulorum Jones Utah: Garfield Co., Bryce Canyon, Boyle 1153; Kane Co., Last Chance Canyon, NDA et al. 2702. San Juan Co., Rainbow Bridge, NDA & NW 3300; east of San Juan con- fluence, NDA & RA 3164. Local and un- common. Dactylis glomerata L. Orchard Grass Utah: Kane Co., Bryce Canyon, Boyle June 1978 Welsh et al.: Kaiparowits Flora 151 1133; Grand Bench,NDA 4262. Introduced forage grass. Distichlis stricta (Torr.) Rydb. Utah: Kane Co., Hole-in-the-Rock, Harri- son 12126; Paria Primitive area, NDA & RA 2875A. Widespread in saline sites at lower elevations. Elymus canadensis L. Utah: Kane Co., Escalante Canyon, NDA 4260, 4209; mile 47, Lake Powell, NDA et al. 3224. San Juan Co., Three Garden, SLW & CAT 11867; Ribbon Canyon, NDA 4110. Widespread; uncommon. Elymus salina Jones Utah: Garfield Co., Bryce Canyon, Harri- son 11044; Escalante Mts., Stanton 773; 15 miles southwest of Escalante, Hall s.n.; Kane Co., Kaiparowits Plateau, NDA & DT 5252. Widespread; uncommon. Elymus triticoides Buckl. Utah: San Juan Co., Ribbon Garden, SLW & NDA 11695. Rare. Festuca elatior L. Utah: Garfield Co., Bryce Canyon, Weight s.n.; 15 miles southwest of Esca- lante, Hall 13596. Introduced forage grass. Festuca octoflora Walt. Six-weeks Fescue Arizona: Coconino Co., 4 miles southeast of Page, RWA 233; Colorado Canyon, be- tween Lee's Ferry and Glen Canyon Dam, NDA 4596; 1 mile west of Glen Canyon Dam, NDA 3446. Utah: Kane Co., Kaiparo- wits Plateau, Harrison 9036; 14 miles east of Glen Canyon City, RWA 336; Cedar Mt., RWA 340; Grosvenor Arch, Reveal et al. 806. San Juan Co., Three Garden, SLW & NDA 11700; do, SLW 11900. Abundant. Glyceria elata (Nash) Hitchc. Utah: Kane Co., Brigham Plains, NDA & RA 2796. Rare. Glyceria striata (Lam.) Hitchc. Utah: Garfield Co., northeast of Esca- lante, Harrison 12345. Rare. Hilaria jamesii (Torr.) Benth. Galleta Arizona: Coconino Co., Cockscomb, NDA & RA 2882. Utah: Garfield Co., 50 miles south of Escalante, Beck & Tanner s.n. Kane Co., Willow Tank, Beck & Tarmer s.n.; Cedar Mt., NDA 3127A; do, RWA 178; Wahweap Bay, NDA 3554; 14 miles east of Glen Canyon City, RWA 317; Nipple Canyon, RWA 251; Dance Hall Rock, JRM 383; Hole-in-the-Rock, Harrison 12118; Willow Tank, Harrison 9084; Kai- parowits Plateau; Bingham 33, Dry Rock Creek, Lake Powell, SLW & NDA 11636; East Clark Bench, NDA & DK 3369; Warm Creek Olsen 33; Halls Creek Bay, Lake Powell, SLW & CM, 11790; Gunsight Bay, Lake Powell, SLW & NDA 11594; Sit Down Bench, Patton 117, Paria Townsite, JRM 320. Abundant. Hordeum glaucum Steud. Utah: Kane Co., Paria Townsite, JRM 315. Rare. Hordeum jubatum L. Utah: Garfield Co., Escalante, Beck & Tanner s.n. Locally common. Imperata brevifolia Vasey Utah: San Juan Co., Wilson Creek, Lake Powell, SLW 12356, 12356a. Rare, endan- gered, or possibly extinct (Welsh et al., 1975). Koeleria nitida Nutt. Utah: Garfield Co., Bryce Canyon, Boyle 1144; 15 miles southwest of Escalante, Hall 13588; 18 miles southwest of Escalante, Hall 13577. Uncommon. Muhlenhergia andina (Nutt.) Hitchc. Utah: Kane Co., Escalante River, NDA 4226; Four Mile Creek, SLW et al. 12578a; Paradise Canyon, SLW et al. 12598. Un- common. Muhlenhergia asperifolia (Nees & Mey.) Parodi Utah: Kane Co., Tibbet Spring, NDA & DK 3416; Four Mile Creek, SLW et al. 12578. San Juan Co., east of San Juan con- fluence. NDA & RA 3177. Uncommon; sa- line seeps. Muhlenhergia curtifolia Scribn. Utah: Kane Co., Mile 47, Lake Powell, NDA et al. 3233. Local, uncommon, and possibly threatened (Welsh et al., 1975). Muhlenhergia pungens Thurb. Arizona: Coconino Co., between Page and Kaibito, Shaw 2648; Paria Plateau, NDA 6415. Utah: Kane Co., Cedar Mt., NDA 3103; Gunsight Bay, Lake Powell, SLW & NDA 11592. Common. Munroa squarrosa (Nutt.) Torr. Arizona: Coconino Co., Paria Plateau, NDA 6421; Cockscomb, NDA 3062; Lee's Ferry, NDA 4569; 9.3 miles south of Page, 152 Great Basin Naturalist Vol. 38, No. 2 NDA et al. 3352. Utah: Kane Co., Cedar Mt., NDA 3103; Gunsight Bay, Lake Pow- ell, SLW & NDA 11592. Common. Oryzopsis bloomeri (Boland.) Ricker Arizona: Coconino Co., Lee's Ferry vicin- ity, NDA 5210. Utah: Kane Co., Kaiparo- wits Plateau, NDA & DT 5259; Paria Townsite, JRM 313. Rare. Oryzopsis hymenoides (R. & S.) Ricker Arizona: Coconino Co., ca 5 miles south of Page, RWA 230. Utah: Kane Co., 50 miles south of Escalante, Beck & Tanner, s.n.; Cedar Mt., NDA & LCH 3827; do., NDA 3113, Hole-in-the-Rock, Harrison 12130; Willow Tank, Harrison 9086; Dance Hall Rock, JRM 384; Cottonwood Wash Road, NDA & RA 2765; Paria River, NDA & LCH 3948; Four Mile Bench, SLW & JRM 12402; Nipple Canyon, FWA 262; 14 miles east of Glen Canyon City, RWA 326. San Juan Co., Three Garden, SLW & NDA 11680. Abundant. Panicum capillare L. Witchgrass Utah: Kane Co., Escalante Canyon, NDA & RA 3210. Rare. Panicum lanuginosum Ell. Utah: Kane Co., canyon at mile 47, Lake Powell, NDA et al. 3232; Reflection Can- yon, SLW & CAT 11852; Escalante Can- yon, SLW & CM 11824; do., NDA 4123. San Juan Co., east of confluence of San Juan, NDA & RA 3162; Three Garden, SLW 12438. Local; uncommon. Panicum obtusum H.B.K. Utah: Hole-in-the-Rock, Harrison 12133. Rare. Panicum virgatum L. Utah: San Juan Co., east of confluence of San Juan, NDA & RA 3163; Ribbon Can- yon, NDA 4104; Three Garden, SLW & CAT 11863. Rare. Phragmites communis Trin. Utah: Kane Co., Lake Powell, NDA 3136. Widespread; locally common. Poa bigelovii Vasey & Scribn. Utah: Kane Co., Cockscomb, NDA 4625; San Juan Co., Three Garden, SLW & NDA 11701. Rare. Poa canbyi (Scribn.) Piper Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5250. Rare. Poa fendleriana (Steud.) Vasey Utah: Kane Co., Kaiparowits Plateau, Harrison 9041; Cockscomb, NDA 4607; do, NDA 3578; Fifty Mile Spring, JRM 372; Paria Townsite, SLW & NDA 9730; Lake Powell, 2 miles east of Warm Creek, NDA 3720; San Juan Co., Navajo Mt., NDA 4162, 4165; Ribbon Garden, SLW & NDA 11698. Common; widespread. Poa pratensis L. Bluegrass Utah: Garfield Co., Bryce Canyon, Boyle 1124; 15 miles southwest of Escalante, Hall 13586, 13589. Rare; Introduced lawn and forage grass. Poa secunda Presl. Utah: Kane Co., Kaiparowits Plateau, Harrison 9052; do, NDA & DT 5255; Cockscomb, NDA 4624. Uncommon. Polypogon monspeliensis (L.) Desf. Utah: Garfield Co., Escalante, Beck & Tanner s.n.; Kane Co., Willow Tank, Beck & Tanner s.n.; 50 miles south of Escalante, Beck & Tanner s.n.; Hole-in-the-Rock, Har- rison 12110; Warm Creek, NDA & RA 2838; Tibbet Spring, NDA & RA 3417; Es- calante River, NDA 4224. Common; wide- spread. Puccinellia nuttalliana (Schult.) Hitchc. Utah: Garfield Co., Escalante, Stanton; 15 miles southwest of Escalante, Hall 13574. Kane Co., Paradise Canyon, SLW et al. 12594. Uncommon. Schismus arabicus Nees. Arizona: Coconino Co., Navajo Bridge vi- cinity, NDA 4560. Uncommon. Secale cereale L. Rye Utah: Kane Co., 15 miles west of Glen Canyon City, NDA & RA 2878. Introduced weedy species. Setaria viridis (L.) Beauv. Utah: Kane Co., east of Kanab, NDA 3277. Rare. Sitanion hystrix (Nutt.) J. G. Smith Arizona: Coconino Co., Cockscomb, NDA 3780. Utah: Garfield Co., Bryce Canyon, Boyle 1129; do, Cottam & Hutchings 2751; 15 miles southwest of Escalante, Hall 13605; do. Hall 13594. Kane Co., Willow Tank, Harrison 9083; Cedar Mt., NDA et al. 2750; Cottonwood Wash road, NDA & RA 2766a; Kaiparowits Plateau, NDA & DT 5276; Paria Townsite, JRM 316; Drift- wood Canyon, Lake Powell, SLW & NDA June 1978 Welsh et al.: Kaiparowits Flora 153 11643; East Clark Bench, Harrison 12072; Four Mile Bench, NDA 4071; Paria River, NDA & LCH 4001; Cedar Mt., NDA & LCH 3828. San Juan Co., Navajo Mt., Gen- try & Davidse 1779; do, NDA et al. 2970; Three Garden, SLW & NDA 11692. Com- mon to abundant; widespread. Sorghastum nutans (L.) Nash Utah: Kane Co., 2 miles north of con- fluence of San Juan and Colorado canyons, SL & SLW 11880. Rare. Sorghum halipense (L.) Pers. Johnson Grass Utah: Kane Co., east of Kanab, NDA 3279. Rare. Sphenopholis obtusata (Michx.) Scribn. Wedgegrass Arizona: Coconino Co., Navajo Creek, NDA 3724. Utah: Kane Co., Crosby Can- yon, NDA & RA 2890. San Juan Co., Three Garden, SLW & NDA 11670. Sporobolus airoides Torr. Dropseed Utah: Kane Co., Paria Townsite, JRM 317; Cottonwood Wash road, NDA & RA 2766. Common. Sporobolus contractus Hitchc. Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3350A; Paria Plateau, NDA 6424. Utah: Garfield Co., 15 miles southwest of Escalante, Hall 13566. Kane Co., Cottonwood Wash road, NDA & DK 3323; 15 miles west of Glen Canyon City, NDA & RA 2874; Church Wells, NDA & DK 3362. Common. Sporobolus cryptandrus (Torr.) Gray Utah: Garfield Co., 15 miles southwest of Escalante, Hall 13562, 13563. Kane Co., Cedar Mt., NDA 3112; do, NDA et al. 3249, do, NDA 3130; Four Mile Bench, SLW et al. 12586; Willow Tank, Harrison 9104; do. Beck & Tanner s.n.; Church Wells, NDA & DK 3364; Halls Creek Bay, Lake Powell, SLW & CM 11802; Wahweap Creek, NDA 3485. Common. Sporobolus flexuosus (Thurb.) Rydb. Arizona: Coconino Co., Paria Plateau NDA, 6416; 9.3 miles south of Page, NDA et al. 3350; Colorado River Canyon north of Navajo Canyon, NDA & RA 3143. Utah: Garfield Co., 15 miles southwest of Esca- lante, Hall 13603. Kane Co., 50 miles south of Escalante, Beck & Tanner s.n.; Cedar Mt., NDA et al. 3248; Dry Rock Creek, Lake Powell, SLW & NDA 11635. San Juan Co., east of confluence of San Juan, NDA et al. 3230. Common. Sporobolus giganteus Nash Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3345. Utah: Kane Co., Cedar Mt., NDA 3122; Church Wells, NDA & DK 3368; 6 miles northwest of Page, NDA & RA 3075. Uncommon to rare. Stipa arida Jones Utah: Kane Co., Kaiparowits Plateau, Harrison 9060; do, NDA & DT 5298. Rare. Stipa comata Trin. & Rupr. Utah: Kane Co., Hole-in-the-Rock, Harri- son 12111; Kaiparowits Plateau, NDA & DT 5256; Cedar Mt., NDA & LCH 3826; Dry Rock Creek, Lake Powell, SLW & NDA 11642; Paria Townsite, JRM 314; Four Mile Bench, SLW & JRM 12403; Paria River, NDA & LCH 3988; Cock- scomb, SLW 5337; Last Chance Bay, Lake Powell, SLW & NDA 11602. San Juan Co., Three Garden, SLW & NDA 11666; Navajo Mt., NDA 4198. Common. Stipa lettermanii Vasey Utah: Garfield Co., 15 miles southwest of Escalante, Hall 13585. San Juan Co., Navajo Mt., NDA 4187. Uncommon. Stipa neomexicana (Thurb.) Scribn. Utah: Kane Co., Willow Tank, Harrison 9095. Uncommon. Stipa scribneri Vasey Utah: Garfield Co., 20 miles southeast of Escalante, Holmgren et al. 2050. Kane Co., Little Valley Canyon, NDA et al. 2715. San Juan Co., east of confluence of the San Juan, NDA 4092a. Uncommon. Stipa speciosa Trin. & Rupr. Utah: Kane Co., Willow Tank, Harrison 9089; Dry Rock Creek, SLW & NDA 11634; Cockscomb, JRM 339. San Juan Co., Piute Canyon, NDA & DT 5349; Rainbow Bridge, SLW & NDA 11653. Common. Tridens pilosus (Buck!.) Hitchc. Utah: Kane Co., Cockscomb, NDA 4619. Uncommon. Tridens pulchellus (HBK) Hitchc. Arizona: Coconino Co., Lee's Ferry, Cot- tarn & Hutchings 2602. Kane Co., Willow Tank, Harrison 9085; Nipple Canyon, NDA & RA 2835. Uncommon. Trisetum spicatum (L.) Richt. Utah: San Juan Co., Navajo Mt., NDA 4164. Rare. 154 Great Basin Naturalist Vol. 38, No. 2 Hydrophyllaceae (Waterleaf Family) Eucrypta micrantha Terr. Utah: Kane Co., Buckskin Mt., NDA 4648; Cockscomb, NDA 4621; do, NDA 4411. Uncommon. Nama hispidum Gray Utah: Kane Co., Wahweap Bay, NDA 3549; do, SLW & NDA 9770; Gunsight Bay, SLW & NDA 11584; Warm Creek, NDA 3287. Uncommon. Nama retrorswn J. T. Howell Utah: Kane Co., Cedar Mt., NDA et al. 3499; do, NDA 3109, Lake Powell, NDA 3694; Paria River, NDA & LCH 3994. Re- stricted, local, and possibly threatened (Welsh et al., 1975). Phacelia affinis Gray Utah: Kane Co., Cockscomb, NDA 4617. Rare. Phacelia cephalotes Gray Arizona: Coconino Co., Cockscomb, NDA 4924. Utah: Kane Co., Cockscomb, NDA 4663; do, NDA 4603. Restricted, local, and possibly threatened (Welsh et al., 1975). Phacelia constancei Atwood Utah: Garfield Co., 39 miles east of Esca- lante. Reveal & Reveal 2823. Edaphically restricted, local, and threatened (Welsh et al, 1975). Phacelia corrugata A. Nels. Arizona: Coconino Co., Colorado Canyon, between Lee's Ferry and Glen Canyon Dam, NDA 4589; Glen Canyon Dam, NDA 3450; do, SLW & NDA 9781. Utah: Kane Co., Little Valley Canyon, NDA et al. 2723; Brigham Plains, NDA & RA 2789; 15 miles south of Cannonville, NDA & RA 2908; west of Glen Canyon Dam, NDA 3567; Halls Creek Bay, Lake Powell, SLW & CM, 11793; Warm Creek, NDA 3710; East Clark Bench, Harrison 12063; 5 miles southeast of Cannonville, Reveal et al. 797; Grand Bench, NDA 3531; Dry Rock Creek, Lake Powell, SLW & NDA 11617; Gun- sight Bay, Lake Powell, SLW & NDA 11587; Wahweap Bay, SLW & NDA 9771; Paria River bridge, SLW & NDA 9744; do, LCH & DA 5374; Crosby Canyon, NDA 3632; Warm Creek, NDA & RA 2842; 15 miles west of Glen Canyon City, NDA & RA 2863. Abundant. Phacelia crenulata Terr. Arizona: Coconino Co., Lee's Ferry, Lar- son & Anderson 5993. Utah: Kane Co., 10 miles northeast of Glen Canyon City, LCH & NDA 5247; Cedar Mt., NDA 3612, do, RWA 175. Tibbet Canyon, NDA 3603; 14 miles east of Glen Canyon City, RWA 211; Nipple Canyon, RWA 243; 6 miles east of Glen Canyon City, SLW 9791. Phacelia demissa Gray var. demissa Utah: Garfield Co., 1 mile east of Henrie- ville, NDA 1975. Kane Co., 6 miles east of Glen Canyon City, SLW & NDA 9789; Cottonwood Wash road, NDA & RA 2762; 10.5 miles east of Glen Canyon City, NDA 2634; 15 miles west of Glen Canyon City, NDA & RA 2864; 4 miles east of Glen Can- yon City, NDA 2630; Warm Creek, NDA & RA 2898; 20 miles south of Escalante, Holmgren et al. 2057; Paria River, NDA & RA 3991; do, SLW 11060; do, NDA 5163. Common. Phacelia heterophylla Pursh Utah: San Juan Co., Navajo Mt., NDA et al. 2984; do, NDA 4143. Uncommon. Phacelia howelliana Atwood Utah: San Juan Co., Ribbon Garden, SLW & NDA 11702; Three Garden, SLE & NDA 11675; Piute Canyon, NDA & DT 5346. Uncommon. Phacelia integrifolia Torr. Arizona: Coconino Co., 4 miles south of Page, RWA 124; do, NDA 2626; Colorado Canyon, between Glen Canyon Dam and Lee's Ferry, NDA & LCH 4027; between Page and Glen Canyon Dam, SLW & NDA 9780; do, NDA 1539. Utah: Kane Co., Sit Down Bench, Patton 116; Cedar Mt., NDA 3610; do, NDA et al. 3504; 2 miles west of Glen Canyon City, NDA 3638; do, Nebeker & Skougard 182; do, Cronquist, 10170; Warm Creek, NDA 3709; Northwest of Glen Canyon dam, NDA 1536. Common. Phacelia ivesiana Torr. Arizona: Coconino Co., Colorado Canyon, between Lee's Ferry and Glen Canyon Dam, NDA 4591. Utah: Kane Co., Kaiparo- wits Plateau, NDA 1867, 1868; East Clark Bench, NDA & RA 2674; Little Valley Can- yon, NDA et al. 2727; Cedar Mt., NDA et al. 2746A; do, RWA 231, 48 miles southeast of Escalante, Cronquist 10035; Paria River bridge, SLW & NDA 9758; 57 miles east of Kanab, Cronquist 10152; 25 miles southeast June 1978 Welsh et al.: Kaiparowits Flora 155 of Escalante, Holmgren et al. 2065a, Ahl- strom Point road, RWA 122. San Juan Co., Thre dSarden SLW 11896. Common. Phacelia tnammillariensis Atwood Arizona: Coconino Co., 4 miles southeast of Page, RWA 263. Utah: Garfield Co., east of Henrieville, NDA 1874. Kane Co., 10 miles northeast of Glen Canyon City, LCH & NDA 5242; Cottonwood Wash, LCH & NDA 5257; 7 miles east of Glen Canyon City, NDA 2632; Warm Creek, Olsen 34; do, NDA 3597. Brigham Plains vicinity, NDA 3743; Gunsight Bay, Lake Powell, SLW & NDA 11582; 3 miles east of Glen Canyon City, NDA 2628; 16 miles east of Glen Canyon City, NDA 4553; 6 miles east of Glen Canyon City, SLW & NDA 9790. Edaphically restricted and endangered (Welsh et al, 1975). Phacelia rotundifolia Torr. Utah: Kane Co., Cockscomb, NDA 4615 Buckskin Mt., NDA 4652. Uncommon. Phacelia pulchella Gray var. sabulonum Howell Utah: Kane Co., Little Valley Canyon, NDA et al. 2720A; Cottonwood Wash road, NDA & RA 2763a; between Nipple Bench and Lake Powell, Cronquist 10012; Paria River, Cronquist 10201; Warm Creek, NDA & RA 2836; do, NDA & RA 2897; Nipple Canyon, NDA & RA 2827; Nipple Bench, NDA & RA 2809; 18 miles northeast of Glen Canyon City, LCH & NDA 5245. Common. Phacelia rafaelensis Atwood Arizona: Coconino Co., Navajo Bridge vi- cinity, NDA & LCH 3814; do, NDA 4563. Restricted, local and possibly threatened (Welsh et al., 1975). Iridaceae (Iris Family) Iris missouriensis Nutt. Utah: Garfield Co., Bryce Canyon, Boyle 1110. Rare. Sisyrinchium radicatum Bicknell Utah: Kane Co., Cottonwood Wash, NDA 5356; Four Mile Bench SLW & BW 12439. Rare. Juncaceae (Rush Family) Juncus arcticus Willd. var. balticus (Willd.) Trautv. Utah: Kane Co., southeast of Brigham Plains, NDA & RA 2797; Four Mile Bench, SLW et al. 12580; Reflection Canyon, Lake Powell, SLW & CAT 11849; Escalante Canyon, NDA 4213. San Juan Co., Rainbow Bridge, SLW & NDA 11662; east of con- fluence of San Juan, NDA 4667. Locally abundant. Juncus torreyi Coville Utah: Kane Co., Four Mile Bench, SLW et al. 12577; do, NDA 4311; do, SLW et al. 12600. San Juan Co., Three Garden, SLW & CAT 11862; east of confluence of San Juan, NDA 5265. Locally common. Juncus tracyi Rydb. Utah: Garfield Co., Bryce Canyon, Bur- key 156. Kane Co., Escalante Canyon, SLW & CM 11819; do, NDA 4122; Reflection Canyon, SLW & CAT 11850; Cottonwood Wash, Reveal et al. 818. Locally common. Juncus xiphodes E. Meyer Utah: Garfield Co., Escalante vicinity, Beck s.n., Kane Co., Wahweap Creek, NDA 4344, Four Mile Bench vicinity, SLW 12592, 12599. San Juan Co., east of con- fluence of San Juan, NDA 4264. Locally common. Juncaginaceae (Arrowgrass Family) Triglochin maritima L. Utah: Kane Co., seep, on Four Mile Bench, SLW 12571. Rare. Labiatae (Mint Family) Hedeoma drummondii Benth. Utah: Kane Co., Buckskin Mt., NDA 4659. San Juan Co., east of confluence of San Juan, NDA & RA 3149. This specimen is fragmentary, and it might belong to an- other species. Rare. Marrubium valgare L. Utah: Kane Co., east of Kanab, NDA 3282. Rare; introduced weed. Monardella odoratissima Benth. Utah: San Juan Co., Navajo Mt., Gentry & Davidse 1769; do, NDA et al. 2977. Un- common. Poliomintha incana (Torr.) Gray Arizona: Coconino Co., Cedar Mt., NDA et al. 2753; 9 miles south of Page, RWA 156 Great Basin Naturalist Vol. 38, No. 2 229. Utah: Garfield Co., Escalante, Cottam 4384; 18 miles southeast of Escalante, Holmgren et al. 2043. Kane Co., Paria Riv- er, NDA & LCH 3965; 2 miles west of Glen Canyon City, Nebeker & Skougard 190; do, Cronquist 10172; Cockscomb, JRM 338; Wahweap Creek, Harrison 12091; East Clark Bench, NDA & RA 2675A; do, NDA & RA 3370; Warm Creek, NDA & RA 2843. Locally common. Salvia dorrii (Kellogg) Abrams Utah: Kane Co., Cedar Mt., NDA et al. 2747; Cockscomb, SLW 5327; Wahweap Creek, Harrison 12090; Nipple Canyon, RWA 242. Uncommon. Leguminosae (Legume Family) Albizia julibrissin Durazz Silk Tree Arizona: Coconino Co., Page (cultivated), NDA et al. 3247. Astragalus agrestis Nutt. Utah: Garfield Co., Bryce Canyon, Weight s.n. Rare. Astragalus amphioxys Gray var. amphioxys Arizona: Coconino Co., 4 miles southeast of Page, RWA 237; 5 miles southeast of Page, Nebeker & Skougard 196; 3 miles south of Wahweap Marina, SLW & NDA 9765. Utah: Kane Co., 14 miles east of Glen Canyon City, RWA 316; Paria River bridge, SLW & NDA 9757; East Clark Bench, Har- rison 12094; Glen Canyon City vicinity, Cronquist 10145; Warm Creek Bay, CAT 173; do, NDA 3707; Wahweap Bay, NDA 3555; Cedar Mt., RWA 177; Cockscomb, SLW 5333. Common. Astragalus amphioxys Gray var. vespertinus (Sheld.) Jones Arizona: Coconino Co., Buckskin Mt., NDA 4939. Utah: Garfield Co., 17 miles west of Escalante, SLW 1704; 0.6 miles north of Escalante, SLW 1702; 3 miles west of Tropic, Reveal et al. 743. Kane Co., 6 miles east of Glen Canyon City, SLW & NDA 9807; Cottonwood Wash road, SLW 5358; Kaiparowits Plateau, Harrison 9058; East Clark Bench, Harrison 12305; between Nipple Bench and Lake Powell, Cronquist 10012; Gunsight Bay, Lake Powell; SLW & NDA 11590; Fifty Mile Spring, JRM 345, 392; Cockscomb, NDA 4632; head of Col- letts Wash, NDA 306, 307. Paria Townsite, SLW & NDA 9736. Common. Astragalus ampullarius Wats. Arizona: Coconino Co., Cockscomb, NDA & RA 2880; do, NDA 4923. Utah: Kane Co., Cockscomb, NDA 4602; do, 4664. Rare, restricted and threatened (Welsh et al., 1975). Astragalus argophyllus Nutt. var. pan- guicensis (Jones) Jones Arizona: Coconino Co., Buckskin Mt., NDA 4938. Utah: Garfield Co., Canaan Peak, NDA & DT 5315. Rare. Astragalus asclepiadoides Jones Utah: Garfield Co., 3 miles southeast of Escalante, Holmgren et al. 2415; do, NDA 5217. Rare. Astragalus hisulcatus (Hook.) Gray Utah: Garfield Co., 2 miles west of Esca- lante, Karren 125; do, SLW 1703; Alvey Wash, Death Ridge, NDA 5185; southwest of Cannonville, Reveal et al. 770. Kane Co., head of Alvey Wash, NDA 5211; Kaiparo- wits Plateau, Harrison 9063; do, NDA & DT 5242; head of Collets Wash, NDA 1872. Astragalus bryantii Bameby Utah: Kane Co., Glen Canyon (Bameby, 1964). Rare and possibly extinct (Welsh, et al, 1975). Astragalus ceramicus Sheld. Arizona: Coconino Co., Paria Plateau, LCH & NDA 5202. Utah: Garfield Co., 20 miles southeast of Escalante, Holmgren et al. 2046. Kane Co., Kaiparowits Plateau, Harrison 9057; Paria River bridge, SLW & NDA 9749; 12 miles west of Glen Canyon City, Cronquist 10173; East Clark Bench, NDA & RA 2681; Cottonwood Wash road, NDA & RA 2771; 4 miles south of Cannon- ville, Reveal et al. 785; Cockscomb, SLW 5352. Widespread; uncommon. Astragalus calycosus Torr. Utah: Garfield Co., Bryce Canyon, 1708. Kane Co., Five Mile Mt., NDA 4646; Four Mile Bench, SLW 12754. Rare. Astragalus coltonii Jones var. coltonii Utah: Garfield Co., 3 miles northeast of Henrieville, NDA 1248. Kane Co., Kaiparo- wits Plateau, Harrison 9067; do, NDA 7 dt 5297; Paria Townsite, SLW & NDA 9721, June 1978 Welsh et al.: Kaiparowits Flora 157 9733. Utah endemic; locally abundant on the Kaiparowits Plateau. Astragalus desperatus Jones var. conspectus Bameby Utah: Garfield Co., ca 5 miles southeast of Escalante, SLW 1697; do, Isely et al. 8732; do NDA 5218. Rare and local; threat- ened (Welsh et al., 1975). Astragalus desperatus Jones var. desperatus Arizona: Coconino Co., between Page and Glen Canyon Dam, SLW & NDA 9775. Utah: Kane Co., Willow Tank, Harrison 9034; do, SLW 1693; Hole-in-the-Rock, Harrison 12145; do. White 120; Lake Pow- ell, Warm Creek Bay, NDA 3704. Locally common. Astragalus emoryanus (Rydb.) Cory Utah: Kane Co., Cockscomb, NDA 4629 (first Utah record). Rare and endangered (Welsh et al., 1975). Astragalus episcopus Wats. Utah: Kane Co., Nipple Bench, NDA & RA 2817. Rare. Astragalus eremiticus Sheld. Arizona: Coconino Co., Buckskin Mt., NDA 4927. Utah: Kane Co., Cockscomb, NDA 4630. Rare. Astragalus flavus Nutt. var. candicans Gray Utah: Kane Co., Nipple Bench, NDA 3477; Cottonwood Wash road, NDA 3576; Cockscomb, NDA 4662; do, NDA 3630; do. Reveal et al. 835; Paria Townsite, Cronquist 10138; do, SLW & NDA 9738. Restricted to seleniferous mud or siltstone. Astragalus hallii Gray var. fallax Bameby Utah: Kane Co., Buckskin Gulch, SLW 5309; Riggs Creek Canyon, NDA 6130. Rare and local; possibly threatened (Welsh et al., 1975). Astragalus humistratus Gray var. humiva- gans (Rydb.) Bameby Utah: Garfield Co., Bryce Canyon, Weight B31/63. Kane Co., east of Grosve- nor Arch, NDA 3176; Four Mile Bench, SLW & JRM 12392. Rare. Astragalus kentrophyta Gray var. colora- doensis Bameby Utah: Kane Co., Wahweap Creek, Harri- son 12086; 2 miles west of Glen Canyon City, Cronquist 10164; Paria River, NDA & LCH 3969; Last Chance Bay, Lake Powell, SLW & NDA 11609. Rare and restricted, possibly threatened (Welsh et al., 1975). Astragalus kentrophyta Gray var. implexus (Canby) Bameby Utah: Garfield Co., Bryce Canyon, Weight s.n. Rare. Astragalus lancearius Gray Arizona: Coconino Co., Cockscomb, NDA 4927. Utah: Kane Co., Buckskin Gulch, SLW 5305; Smoky Mt., NDA 5202; Tibbet Bench, NDA & DT 5068; Cockscomb, NDA s.n.; East Clark Bench, NDA & RA 2674. Rare and local, possibly threatened (Welsh et al., 1975). Astragalus lentiginosus Dougl. var. albi- florus (Gray) Schoener Utah: Kane Co., east of Glen Canyon City, Webb 63; Cockscomb, SLW 5326; Paria Townsite, Olson 23; Smoky Mt., RWA 131; 6 miles east of Glen Canyon City, SLW & NDA 9808; 31 miles east of Glen Canyon City, NDA et al. 2695; Warm Creek, NDA 3437; Brigham Plains, NDA et al. 3491; 4 miles south of Cannonville, Re- veal et al. 783. Common; middle elevations. Astragalus lentiginosus Dougl. var. palans (Jones) Jones Arizona: Coconino Co., 9.3 miles south of Page, NDA et al. 3341; between Page and Glen Canyon Dam, SLW & NDA 9784. Utah: Kane Co., 2 miles east of Glen Can- yon City, NDA & DT 3433; Little Valley; NDA 3334; Willow Tank, SLW & NDA 11650; Wahweap Bay, NDA 3695; Paria Townsite, SLW & NDA 9727; Nipple Can- yon, RWA 252. San Juan Co., east of con- fluence of San Juan, NDA & RA 3158; Three Garden, SLW & NDA 11665; Navajo Mt., NDA & DT 5334; Trail Canyon, SLW 11909. Common; lower elevations. Astragalus lentiginosus Dougl. var. vitreus Bameby Utah: Kane Co., Four Mile Bench, NDA 4073; do, NDA 4064; do, SLW 12413; do, SLW 12426; do. SLW 12446; do, SLW & JRM 12394; Cathy Flat, SLW & NDA s.n.; Nipple Bench, NDA & RA 2820; Brigham Plains, NDA & RA 2791A; Smoky Mt., Cronquist 10022. Locally common, abun- dant. Astragalus lonchocarpus Torr. Utah: Garfield Co., Death Ridge vicinity. 158 Great Basin Naturalist Vol. 38, No. 2 NDA 5180. Kane Co., 10 miles southeast of Cannonville, SLW 5363; Sheep Creek, Re- veal et al. 778; Little Valley, SLW & NDA 116172. Saline sites; common. Astragalus malacoides Barneby Utah: Kane Co., Smoky Mt., NDA 5203; Tibbet Spring, NDA 4554; Harris Wash Road, NDA & DT 5300; east of Grosvenor Arch, NDA 5171; Tibbet Bench, NDA & DT 5064; Nipple Bench, NDA 3465; Grand Bench, NDA 3532; Brigham Plains, NDA et al. 3492; Nipple Creek, NDA 3476; Kai- parowits Plateau, JRM 394; do, NDA & DT 5243; Four Mile Bench, SLW 12749. This plant is endemic to the Kaiparowits Region; it is considered as threatened (Welsh et al., 1975). Astragalus megacarpus (Nutt.) Gray Utah: Garfield Co., Bryce Canyon, SLW 1707; do. Weight B-31/12. Kane Co., Kai- parowits Plateau, Harrison 9065; Cock- scomb, NDA 4637. Rare. Astragalus miser Dougl. var. oblongifoUus (Rydb.) Cronq. Utah: Garfield Co., Canaan Peak, NDA & DT s.n.; do, SLW 12871. Rare. Astragalus moencoppensis Jones Utah: Kane Co., Brigham Plains, NDA & RA 2787; Nipple Bench, NDA & RA 2819; between Last Chance and CoUett Wash, NDA 5208; Left hand fork of Last Chance Creek, SLW 12748. Uncommon. Astragalus mollissimus Torr. var. thomp- sonae (Wats.) Barneby Arizona: Coconino Co., 3 miles south of Wahweap Marina, SLW & NDA 9761. Utah: Garfield Co., Escalante, Cottam 4405; 5 miles west of Escalante, SLW 1701; 8 miles southeast of Escalante, Holmgren et al. 2041; 1 mile east of Cannonville, Isely et al. 8739. Kane Co., Willow Tank, Harrison 9108; do, SLW 1691; 10 miles southeast of Cannonville, NDA et al. 3505a; Cotton- wood Wash road, SLW 5361; Little Valley Canyon, NDA 3430; Last Chance Bay, SLW & NDA 11603. Abundant. Astragalus tnusiniensis Jones Utah: Garfield Co., vicinity 6 miles south- east of Escalante, SLW 1699; do, Holmgren et al. 22231 do, NDA 5219. Kane Co., Har- ris Wash road, NDA & DT 5301. Rare. Astragalus newberryi Gray Utah: Kane Co., 51 miles south of Esca- lante, NDA 520; Paria Townsite, SLW 5324; Last Chance Creek, Cronquist 10028; Kaiparowits Plateau, JRM 393a; do, NDA & DT 5248; Wahweap Creek, NDA 3480. Common. Astragalus nuttallianus DC. var. micranthi- formis Barneby Utah: Kane Co., Cockscomb, NDA 3619; Tibbet Canyon, NDA 3443; 6 miles east of Glen Canyon City, SLW & NDA 9804; Paria River, Cronquist 10197. San Juan Co., Three Garden, SLW 11902; confluence of San Juan and Colorado canyons, SLW 11908. Common. Astragalus oophorus Wats. var. caulescens Jones Arizona: Coconino Co., Buckskin Mt., NDA 4940. Utah: Garfield Co., Death Ridge vicinity, NDA 5178. Kane Co., Kai- parowits Plateau, Harrison 9073; do, JRM 395; do, NDA & DT 5237; Four Mile Bench, SLW 12762. Rare. Astragalus praelongus Sheld. var. prae- longus Utah: Garfield Co., 10 miles northeast of Henrieville, Harrison 12104; Pet Hollow, NDA 4327, 4329; 1.5 miles south of Can- nonville, Reveal et al. 763; 1 mile east of Cannonville, Isely et al. 8731. Kane Co., Willow Tank, Beck s.n.; do, SLW 1694; Paria Townsite, SLW & NDA 9737; 34 miles southeast of Escalante, Cronquist 10033; Cottonwood Wash, G. Moore 401; 5 miles east of Paria River, JRM 114; Four Mile Bench, SLW & JRM 12393; do, NDA 4074; Wahweap Creek, east of Glen Can- yon City, NDA 3464. Common; seleniferous soils. Astragalus preussii Gray var. preussii Arizona: Coconino Co., Lee's Ferry, LCH 8323. Utah: Kane Co., 6 miles east of Glen Canyon City, SLW & NDA 9792; 30 miles east of Glen Canyon City, NDA 4546. Un- common; seleniferous soils. Astragalus sabulonum Gray Arizona: Coconino Co., 4 miles south of Page, NDA 2627a; between Page and Glen Canyon Dam, SLW & NDA 9776; 9 miles south of Page, RWA 216. Utah: Kane Co., Labyrinth Canyon, Lake Powell, NDA 3733. June 1978 Welsh et al.: Kaiparowits Flora 159 Astragalus sesquiflorus Wats. Utah: Kane Co., Kaiparowits Plateau, Harrison 9035; do, JRM 391; do, NDA 539; Paria Townsite, Webb 64; Buckskin Gulch, SLW 5307; Wahweap Canyon, NDA 3472; Hackberry Canyon, NDA 3683; Last Chance Creek, NDA 3753. San Juan Co., Navajo Mt., NDA 4192; do, NDA et al. 2975. Astragalus striatiflorus Jones Arizona: Coconino Co., Paria River, NDA 4016, NDA 4016; Kane Co., Hackberry Canyon, NDA 3682; Paria River, NDA & LCH 3946. Rare and endangered (Welsh et al., 1975). Astragalus subcinereus Gray Utah: Garfield Co., Weight B031/42; do, Cottam & Hutchings 2759; do, Cottam 4366. Astragalus wardii Gray Utah: Kane Co., 4 miles south of Cannon- ville. Reveal et al. 780; do, Cronquist 10215. Endemic to Utah; rare. Astragalus zionis Jones Arizona: Coconino Co., Glen Canyon bridge. Moore & Tanner 16. Utah: Kane Co., Hackberry Canyon, NDA 3679, 3681; Cedar Mt., NDA 3445; Cockscomb, NDA 3618; Cottonwood Wash road, SLW 5359; do., NDA 3585. San Juan Co., Navajo Mt., NDA 4136, 4167; do, east of confluence of San Juan, NDA & RA 3168. Common on sandstone at lower elevations. Caragana arborescens Lam. Arizona: Coconino Co., Wahweap camp- ground, cultivated, NDA 3453. Cercis canadensis L. Arizona: Coconino Co., Wahweap, culti- vated, NDA 3454. Cercis occidentalis Torr. Arizona: Coconino Co., Colorado Canyon, between Lee's Ferry and Glen Canyon dam, NDA 4597. Utah: San Juan Co., Rainbow Bridge, NDA & NW 3298; east of con- fluence of San Juan, NDA & RA 3171, 3193, 3222; Three Garden, SLW & NDA 11683; do, SLW 12423. Unique plants of drainages in sandstone and hanging gardens; iincommon. Dalea fremontii Torr. Utah: Kane Co., Gunsight Bay, Lake Powell, SLW & NDA 11591; Cottonwood Wash road, Moore 409; Labyrinth Canyon, Lake Powell, NDA 3730; Halls Creek Bay, Lake Powell, SLW & CM 11791; 50 miles south of Escalante, JRM 377; do, Beck & Tanner s.n.; Kaiparowits Plateau, NDA & DT 5296; Hole-in-the-Rock, Harrison 12127; Willow Tank, Harrison 9099. San Juan Co., Three Garden, SLW of San Juan, NDA & RA 3169. Locally common; wide- spread at lower elevations. Dalea lanata Spreng. Arizona: Coconino Co., Kaibito-Tonolea highway, NDA et al. 2913. Utah: Kane Co., Cedar Mt., NDA 3123; do, Shaw 1612. Un- common. Glycyrrhiza lepidota Pursh Utah: Garfield Co., Escalante Canyon, Beck 107. Kane Co., Willow Tank, SLW 1690; Paria River, NDA & LCH 4017; Wahweap Cabin, Four Mile Bench, NDA 4345. Local; uncommon. Hedysarum boreale Nutt. var. boreale Utah: Garfield Co., Bryce Canyon, Cot- tam 2758; Escalante, Cottam 4420; Death Ridge vicinity, NDA 5188. Kane Co., Smoky Mt., NDA 5205; Kaiparowits Plateau, NDA & DT 5290. Uncommon. Lathyrus brachycalyx Rydb. ssp. zionis (C.L. Hitchc.) Welsh Utah: Garfield Co., Bryce Canyon, Harri- son 12318. Kane Co., Kaiparowits Plateau, Harrison 9066; do, JRM 379; dom NDA & DT 5283; Cockscomb, NDA 3620; East Clark bench, NDA & RA 2686. Uncommon. Lathyrus lanzwertii Kellogg sens. lat. Utah: Garfield Co., Death Ridge vicinity, NDA 5187; Escalante Canyon, Cottam 4424. Uncommon. Lotus longebracteatus Rydb. Utah: Kane Co., Cottonwood Wash, Cronquist 10211; Cockscomb, Harrison 12050; do, SLW 5328; Paria River, NDA & LCH 3958. Uncommon; southwestern por- tion. Lotus utahensis Ottley Utah: Garfield Co., Bryce Canyon, Weight B31/66; do, Burkey 166; 15 miles southwest of Escalante, Hall 105. Uncom- mon. Lupinus X alpestris A. Nels. Utah: Garfield Co., Bryce Canyon, Weight B32/302. Uncommon. 160 Great Basin Naturalist Vol. 38, No. 2 Lupinus caudatus Kell. var. caudatus Utah: Kane Co., Four Mile Bench, SLW & NDA 12375; San Juan Co., Navajo Mt., NDA et al. 2971. Local, in drainages. Lupinus kingii Wats. Utah: Garfield Co., Bryce Canyon, Weight s.n., June-July 1934. Rare. Lupinus pusillus Pursh ssp. intermontanus (Heller) Dunn Arizona: Coconino Co., Paria Plateau, NDA & LCH 3775, 3786. Utah: Kane Co., East Clark Bench, Harrison 12069; Hole-in- the-Rock, White 116; 50 miles south of Es- calante, JRM 346, 373; Cockscomb, SLW 5322; Warm Creek, NDA & RA 2845; Ce- dar Mt., RWA 173. Common. Lupinus pusillus Prush ssp. rubens (Rydb.) Dunn Arizona: Coconino Co., 3 miles south of Wahweap Marina, SLW & NDA 9763. Utah: Kane Co., Paria River, Cronquist 10176; 48 miles southeast of Escalante, Cronquist 10026; Dry Rock Creek, Lake Powell, SLW & NDA 11627. Common. Lupinus wyethii Kellogg Utah: Kane Co., Escalante Canyon, SLW & CM 11821. Introduced weed. Medicago lupulina L. Utah: Kane Co., Escalante Canyon, SLW & CM 11821. Introduced weed. Medicago sativa L. Utah: Garfield Co., Bryce Canyon, Rowe 178, 1931. Introduced forage plant. Melilotus alba Descr. Arizona: Coconino Co., Colorado Canyon, between Lee's Ferry and Glen Canyon Dam, NDA & LCH 4033. Utah: Kane Co., Four Mile Bench, SLW 12569. Introduced forage plant. Melilotus officinalis (L.) Lam. Utah: Kane Co., Paria River, NDA & LCH 3961; Escalante River, NDA 4212. In- troduced forage plant. Oxytropis jonesii Barneby Utah: Garfield Co., Bryce Canyon, Cot- tam 2745; Alvey Wash, 8 miles southeast of Escalante, Holmgren et al. 2022. Edaphi- cally restricted, rare, and threatened (Welsh et al., 1975). Oxytropis lambertii Pursh Utah: Kane Co., 1 mile east of Paria Riv- er, SLW 5353; Cockscomb, NDA 3548; Cottonwood Wash road, NDA & RA 2782; East Clark Bench, NDA & RA 2688. Local- ly abundant. Oxytropis oreophila Gray Utah: Garfield Co., Death Ridge, NDA 5186; do, NDA 5215. Rare. Parryella filifolia Gray Arizona: Coconino Co., Lee's Ferry, Cot- tam & Hutchings 2600. Rare and possibly threatened. Petalostemon flavescens Wats. Utah: Kane Co., East Clark Bench, NDA & RA 2675; Cedar Mt., NDA et al. 2735; Little Valley Canyon, NDA et al. 2711; Driftwood Canyon, Lake Powell, SLW & NDA 11646. Common. Petalostemon occidentale Torr. Utah: Kane Co., Reflection Canyon, Lake Powell, SLW & CAT 11858. San Juan Co., east of confluence of San Juan, NDA 4098; Rainbow Bridge, NDA & NW 3299. Com- mon. Petalostemon searlsiae Gray Utah: Kane Co., Canaan Peak vicinity, NDA & DT 5312. Local; uncommon. Peteria thompsonae Wats. Utah: Kane Co., Nipple Bench, NDA & RA 2822; Warm Creek, NDA & RA 2856; Smoky Mt., NDA 5206; Little Valley Can- yon, NDA et al. 2723. Locally abundant, es- pecially at middle elevations. Psoralea epipsila Barneby Utah: Kane Co., west of the Cockscomb, NDA 2668. Rare and endangered (Welsh et al, 1975). Psoralea juncea Eastw. Utah: Kane Co., Last Chance Bay, Lake Powell, SLW & NDA 11605, East Clark Bench, Harrison 12298; Paria River, NDA & LCH 3943; Cottonwood Wash road, NDA & RA 2775; Cockscomb, SLW 5340. Local; common. Psoralea lanceolata Pursh var. lanceolata Arizona: Coconino Co., NDA & RA 2884. Utah: Garfield Co., Kane Co., 19 miles southeast of Escalante, Holmgren et al. 2044; East Clark Bench, NDA & RA 2691; Cockscomb, CAT 136; Fifty Mile Spring, SLW 1696; between Kodachrome Flat and Cannonville, CAT 143; Brigham Plains, NDA & RA 2788. Common. Psoralea lanceolata Pursh var. stenophylla (Rydb.) Toft & Welsh Utah: Kane Co., Fifty Mile Spring, HRM June 1978 Welsh et al.: Kaiparowits Flora 161 371; Escalante Canyon, SLW & GM 11830; Willow Tank, SLW 1689; 50 miles south of Escalante, Beck & Tanner s.n.; Hole-in-the- Rock, Harrison 12114. Common. Psorothamnus polyadenius (Terr.) Rydb. Utah: San Juan Co., east of Navajo Mt., NDA & DT 5350. Rare. Sophora stenophylla Gray Arizona: Coconino Co., Navajo Canyon, NDA 3726. Utah: Kane Co., Hole-in-the- Rock, SLW 1695; Harris Wash road, NDA & DT 5205. Uncommon. Trifolium andinum Nutt. Utah: San Juan Co., Navajo Mt., NDA 4160. Rare. Liliaceae (Lily Family) Agave utahensis Engelm. Arizona: Coconino Co., Paria Canyon, observed but not collected. Allium acuminatum Hook. Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5246. Rare. Allium nevadense Wats. Arizona: Coconino Co., Buckskin Mt., NDA 4944; Paria River, Cronquist 10133; 22.5 miles southeast of Cannonville, Reveal & Gentry 812; Five Mile Mt., NDA 4650. Uncommon. Androstephium breviflorum Wats. Utah: Kane Co., Paria, JRM MHO; 14 miles east of Glen Canyon City, RWA 322; Willow Tank, Karren 97; 30 miles east of Glen Canyon City, NDA 4548; East Clark Bench, NDA et al. 3495; Cedar Mt., RWA 172. Widespread; uncommon. Calochortus aureus Wats. Utah: Kane Co., 28 miles southeast of Es- calante, Harrison 9118; Cottonwood Wash road, Moore 408; between Nipple Bench and Lake Powell, Cronquist 10010; Last Chance Creek, NDA 3744; Nipple Bench, NDA & DT 5066; 6 miles east of Glen Canyon City, SLW & NDA 9793. Edaphi- cally restricted; locally abundant. Calochortus flexuosus Wats. Utah: Kane Co., 40 miles east of Kanab, Cronquist 10140, do, SLW 5343; 16 miles east of Glen Canyon City, NDA 3665; Labyrinth Canyon, Lake Powell, NDA 3729. Low elevations; common. Calochortus nuttallii T. & G. Utah: Garfield Co., Bryce Canyon, Weight s.n. Kane Co., Paria River, Cronquist 10137; Kaiparowits Plateau, Holmgren et al. 2071; Nipple Bench, NDA & DT 5061; Last Chance Creek, ca 3 miles west of jet. with Escalante Road, NDA 3744. Middle elevations; uncommon. Eremocrinum, albomarginatum Jones Utah: Kane Co., Hole-in-the-Rock, Buss 95. Rare. Fritillaria atropurpurea Nutt. Utah: Garfield Co., Death Ridge, NDA 5182; Kane Co., Kaiparowits Plateau, Harri- son 9043; do, NDA & DT 5232; Hackberry Canyon, NDA 3692. Rare. Laucocrinum montanum Nutt. Utah: Garfield Co., Bryce Canyon, Weight s.n. Rare. Smilacina stellata (L.) Desf. Utah: Kane Co., Brigham Plains, NDA & RA 2800. Local; rare. Yucca angustissima Engelm. Utah: Garfield Co., ca 5 miles southeast of Escalante, NDA 5220. Kane Co., Willow Tank, Harrison 9111; Little Valley Canyon, NDA et al., 2731. Uncommon. Yucca baccata Torr. Utah: Kane Co., Four Mile Bench, NDA 5175. Middle elevations; uncommon. Yucca baileyi W. & S. Utah: Kane Co., V4 mile north of Arizona state line, along U.S. 89, NDA et al. 2733; east of Cockscomb, SLW 5347; southeast of Glen Canyon City, RWA 226; 14 miles east of Glen Canyon City, RWA 312; between Glen Canyon City and Paria River, LCH 4254. Common; sandy sites south central basin. Yucca harrim-aniae Trel. Utah: Garfield Co., Escalante, Cottam 4406. Rare. Yucca kanabensis McKelvey Utah: Kane Co., Kanab vicinity, SLW 11065. Rare. Yucca toftiae Welsh Utah: Kane Co., Dry Rock Creek, Lake Powell, SLW & GM 11779. San Juan Co., Ribbon Garden, NDA 4112; Three Garden, SLW 11935a. Endemic, local, and possibly threatened (Welsh et al. 1975). Zigadenus paniculatus Nutt. 162 Great Basin Naturalist Vol. 38, No. 2 Utah: Kane Co., Buckskin Gulch, SLW 5322; Kaiparowits Plateau, NDA & DT 5275. Rare. Zigadenus vaginatus (Rydb.) Macbr. Utah: Kane Co., Reflection Canyon, SL & SLW 11878. San Juan Co., east of con- fluence of San Juan, NDA & RA 3180; do, NDA et al. 3229. Endemic, local, and possi- bly endangered (Welsh et al. 1975). LiNACEAE (Flax Family) Linum aristatum Engelm. Utah: Kane Co., Willow Tank, Harrison 9097; Paria River, NDA & LCH 3954. Widespread; uncommon. Linum kingii Wats. Utah: Garfield Co., Bryce Canyon, Cot- tam 2749. Uncommon. Linum perenne L. Utah: Garfield Co., Bryce Canyon, Weight s.n. Kane Co., Cottonwood Wash road, NDA & LCH 3834; Kaiparowits Plateau, NDA & DT 5280. Widespread; un- common. Linum puberulum (Engelm.) Heller Utah: Garfield Co., 1.5 miles south of Cannonville, Reveal et al. 764. Rare. Linum subteres (Trel.) Winkler Utah: Kane Co., Kodachrome Flat, Holm- gren & Holmgren 4711; 15 miles west of Glen Canyon City, NDA & RA 2871; East Clark Bench, NDA & RA 2690; Four Mile Bench, SLW & JRM 12410. Rare. LoASACEAE (Loasa Family) Mentzelia albicaulis (Dougl.) T. & G. Utah: Garfield Co., 1.3 miles south of Cannonville, Revel et al. 759. Kane Co., Kaiparowits Plateau, Harrison 9074, do, NDA & DT 5289; Little Valley, NDA 3516; 6 miles east of Glen Canyon City, SLW & NDA 9805; Warm Creek, NDA 3718; Cockscomb, SLW 5339; Cedar Mt., RWA 188. Common. Mentzelia integra (Jones) Tidestr. Utah: San Juan Co., Three Garden, SLW 12418. Rare. Mentzelia multiflora (Nutt.) Gray Arizona: Coconino Co., Colorado Canyon, between Glen Canyon Dam and Lee's Fer- ry, NDA & LCH 4036. 9 miles south of Page, NDA et al. 338. Utah: Kane Co., 31.5 miles east of Glen Canyon City, NDA et al. 2706, 20 miles southeast of Escalante, Buss 85; Nipple Canyon, NDA & RA 2834; 15 miles west of Glen Canyon City, NDA & RA 2877; Smoky Mt., NDA & NW 3292; Gimsight Bay, Lake Powell, SLW & NDA 11580; Hole-in-the-Rock, Harrison 12147; Willow Tank, Harrison 9027; Cottonwood Wash road, Reveal et al. 829. San Juan Co., , east of confluence of San Juan, NDA & RA 3166. Common. LoBELiACEAE (Lobclia Family) Lobelia cardinalis L. Utah: Kane Co., Reflection Canyon, SL & SLW 11879; Escalante Canyon, NDA & RA 3204. San Juan Co., Rainbow Bridge, NDA & NW 3301; east from confluence of San Juan, NDA & RA 3170A, 3192; do, NDA et al. 3228. Hanging gardens; uncommon. Malvaceae (Mallow Family) Sphaeralcea coccinea (Pursh) Rydb. Arizona: Coconino Co., Colorado Canyon, between Glen Canyon Dam and Lee's Fer- ry, NDA & LCH 4037. Utah: Garfield Co., Bryce Canyon, Bosworth 24; Escalante Can- yon, Cottam 4372. Kane Co., Last Chance Creek, NDA 3751. Common. Sphaeralcea grossulariaefolia (H. & A.) Rydb. Arizona: Coconino Co., NDA 3727; Page vicinity, SLW & NDA 9785; 9.3 miles south of Page, NDA et al. 3344. Utah: Kane Co., Nipple Canyon, RWA 250; Cedar Mt., RWA 347; Hole-in-the-Rock, Harrison 12112; Little Valley Canyon, NDA et al. 2725; Willow Tank, Harrison 9032; Cotton- wood Wash road, NDA & RA 2764; Grand Bench, NDA 4256, 2 June 1972; Paria Riv- er, NDA & LCH 3956; Last Chance Bay, Lake Powell, SLW & NDA 11597; Esca- lante Canyon, NDA & RA 3211; do, SLW & CM, 11827; 1 mile west of Glen Canyon City, Toft & Jefferies 123; mouth of Esca- lante Canyon, Lake Powell, SLW & CM 11810. San Juan Co., canyon bottom 1 mile east of Hole-in-the-Rock, SLW & CAT June 1978 Welsh et al.: Kaiparowits Flora 163 11869; Three Garden, SLW 12420. Abun- dant. Sphaeralcea leptophylla (Gray) Rydb. Arizona: Coconino Co., Cottam 2624; 2 miles north of Glen Canyon Dam, LCH 1005. Rare. Sphaeralcea parvifolia A. Nels. Arizona: Coconino Co., Colorado Canyon, between Glen Canyon Dam and Lee's Fer- ry, NDA & LCH 4038. Utah: Garfield Co., Escalante, Cottam 4371; do. Beck & Tanner 8240; do, Jefferies 38; Fifty Mile Spring, JRM 354; Cedar Mt., RWA 189; Gunsight Bay, Lake Powell, SLW & NDA 11593; Dry Rock Creek, Lake Powell, SLW & NDA 11625; Cockscomb, NDA 4605; Glen Canyon City vicinity, Cronquist 10149; Four Mile Bench, SLW & JRM 12396; Grosvenor Arch, Jefferies 44. Abundant. Nyctaginaceae (Four-O'Clock Family) Abronia fragrans Nutt. Arizona: Coconino Co., Page vicinity, SLW & NDA 9783. Utah: Kane Co., Beck s.n.; Kaiparowits Plateau, NDA & DT 5229; Cedar Mt., RWA 186; Cockscomb, NDA 3582, Cottonwood Wash road, Reveal et al. 811; Willow Tank, White 90; Hole-in-the- Rock; White 126; Glen Canyon City vicin- ity, Cronquist 10160; 34 miles southeast of Escalante, Cronquist 10032; 8 miles south of Paria Townsite, Harrison 12058; East Clark Bench, Harrison 12065; Warm Creek, NDA & RA 2844; Gunsight Bay, Lake Powell, SLW & NDA 11601. Abundant. Abronia nana Wats. Utah: Garfield Co., Bryce Canyon, Cot- tam 4359. Kane Co., Cathy Flat, SLW & NDA 11716; Five Mile Mt., NDA 4643; Cockscomb, NDA 3623; Alvey Wash, NDA 5213; 4 miles south of Cannonville, Reveal et al. 784; Four Mile Bench, NDA 5174; Last Chance Creek, NDA 3750; Buckskin Gulch, SLW 5308. Local; uncommon. Allionia choisyi Standi. Arizona: Coconino Co., 8 miles southeast of Page, NDA et al. 3244. Rare. Allionia incarnata L. Arizona: Coconino Co., Lee's Ferry, Cot- tam 2621. Utah: Kane Co., east of Twilight Canyon, Lake Powell, NDA & RA 3147. Uncommon. Boerhaavia torreyana (Wats.) Standi. Utah: Kane Co., Cedar Mt., NDA & DK 3389. Rare. Hermidium alipes Wats. Utah: Kane Co., 6 miles east of Warm Creek, SLW & NDA 9788; Cottonwood Wash road, SLW 5355; between Nipple Bench and Lake Powell, Cronquist 10014; 30 miles east of Glen Canyon City, NDA 4550. Edaphically restricted; uncommon. Mirabilis multiflora (Torr.) Gray Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5295; Nipple Bench, NDA & RA 2818A; Warm Creek, NDA & RA 2855; Escalante Canyon, NDA 4114; Four Mile Bench, SLW 12831. Widespread; locally common. Oxybaphus glaber Wats. Utah: San Juan Co., NDA & RA 3153. Rare. Oxybaphus linearis (Pursh) Robins. Utah: Kane Co., Cottonwood Wash road, NDA & RA 2763; Cockscomb, NDA 2923. Uncommon. Tripterocalyx micranthus (Torr.) Hook. Arizona: Coconino Co., Lee's Ferry, Cot- tam 2606. Utah: Kane Co., Willow Tank, Beck & Tanner s.n.; Cedar Mt., RWA 329; Warm Creek, NDA & RA 2848; Paria Riv- er, NDA & LCH 3973; East Clark Bench, NDA & RA 2680. San Juan Co., Three Gar- den, SLW & NDA 11676. Uncommon. Tripterocalyx wootonii Standi. Arizona: Coconino Co., Cedar Mt., NDA et al. 2751. Utah: Kane Co., Willow Tank, Harrison 9026; Hole-in-the-Rock, Harrison 12105; 50 miles south of Escalante, Beck & Tanner s.n.; Cedar Mt., NDA & LCH 3818; Lake Powell east of Wahweap Marina, NDA 3697; Glen Canyon City vicinity Cronquist 10143; Paria River, Cronquist 10178. Locally common. Oleaceae (Olive Family) Fraxinus anomala Torr. Arizona: Coconino Co., Colorado River between Glen Canyon Dam and Lee's Fer- ry, NDA 4598. Utah: Garfield Co., Esca- lante, Cottam 4401; 1.3 miles south of Can- nonville, Reveal 758. Kane Co., Wahweap Creek, NDA 3458; Cedar Mt., RWA 185; 164 Great Basin Naturalist Vol. 38, No. 2 Fifty Mile Spring, JRM 376; Paria Town- site, SLW & NDA 9741; 20 miles south of Cannonville, Christensen s.n.; Smoky Mt., Cronquist 10024; Hole-in-the-Rock, White 131, Cockscomb, SLW 5335. Widespread; uncommon. Menodora scabra Gray Utah: Garfield Co., Holmgren et al. 2033. Rare and possibly threatened (Welsh et al. 1975). Onagraceae (Evening-Primrose Family) Epilobiwn angustifolium L. Utah: San Juan Co., Navajo Mt., NDA et al. 2976. Rare. Epilobium hornemannii Reichenb. Utah: Kane Co., Warm Creek, NDA & RA 2824. Uncommon; wet sites. Gaura parviflora Dougl. Utah: Kane Co., east of Kanab, NDA 3278. Rare. Gayophytum nuttallii T. & G. Utah: San Juan Co., Navajo Mt., NDA 4178; do, NDA & DT 5341. Uncommon. Oenothera albicaulis Pursh Arizona: Coconino Co., Paria Plateau, NDA & LCH 3762. Utah: Garfield Co., 1.5 miles south of Cannonville, Reveal 765; Kane Co., Smoky Mt., NDA 3746; 6.6 miles southeast of Cannonville, Reveal et al. 799; Hole-in-the-Rock, White 93; 6 miles east of Paria, Harrison 12317; East Clark Bench, Harrison 12081; Paria, JRM 116. Common. Oenothera caespitosa Nutt. var. jonesii Munz Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5269; 6.6 miles south of Can- nonville, Reveal 798, no date. Uncommon. Oenothera caespitosa Nutt. var. marginata (Nutt.) Munz Utah: Kane Co., Fifty Mile Spring, JRM 360; Dry Rock Creek, SLW & NDA 11619; 15 miles west of Glen Canyon City, NDA & RA 2868; between Nipple Bench and Lake Powell, Cronquist 10011; Paria Townsite, SLW & NDA 9739; 6 miles east of Glen Canyon City, SLW & NDA 9797; Warm Creek, NDA & RA 2841; Cotton- wood Wash road, LCH & NDA 5251; Ce- dar Mt., NDA 3107. San Juan Co., Navajo Mt., NDA 4193. Common. Oenothera brachycarpa Gray Utah: Garfield Co., 8 miles southeast of Escalante, Holmgren 2027; Bryce Canyon, Cottam 2747. Rare. Oenothera chamaenerioides Gray Utah: Kane Co., Paria River, Cronquist 10198; Cockscomb, NDA 4622. Rare. Oenothera contorta Dougl. ex Hook. Utah: Kane Co., Cedar Mt., NDA 4666; Butler Valley, NDA 5166; Kaiparowits Plateau, NDA & DT 5226. Uncommon. Oenothera coronopifolia T. & G. Utah: Garfield Co., Bryce Canyon, Boyle 1162. Rare. Oenothera decorticans (H. & A.) Greene Utah: Kane Co., Labyrinth Canyon, Lake Powell, NDA 3734; Grand Bench, NDA 3525; 6 miles east of Warm Creek, SLW & NDA 9796; Dry Rock Creek, Lake Powell, SLW & NDA 11632. Local; uncommon. Oenothera eastwoodiae (Munz) Raven Utah: Kane Co., Little Valley Canyon, NDA et al. 2761A; 6 miles east of Glen Canyon City, SLW & NDA 9787; 17 miles east northeast of Glen Canyon City, RWA 204; Warm Creek, RWA 115; Cottonwood Wash road, NDA & RA 2760; Sit Down Bench, Patton 126. Local; common to abun- dant. Oenothera flava (A. Nels.) Garrett Utah: Kane Co., Paria, JRM 117. Rare. Oenothera lavandulaefolia T. & G. Utah: Garfield Co., Bryce Canyon, Cot- tam 2748. Kane Co., Drip Point, NDA 4066; Four Mile Bench, SLW & JRM 12408a. Local; common. Oenothera longissima Rydb. Utah: Kane Co., Wahweap seep, west of Four-Mile Bench, NDA 4343, Escalante Canyon, SLW & GM 11809. San Juan Co., east of confluence of San Juan, NDA & RA 3156; Navajo Mt., NDA et al. 2986. Com- mon; drainages. Oenothera megalantha Munz Utah: Kane Co., Smoky Mt., NDA 5952, 6008. New Utah record; rare and possibly endangered (Welsh et al. 1975). Oenothera multijuga Wats. Utah: Kane Co., Paria River, Woodruff 1121; 14 miles east of Glen Canyon City, NDA & RA 2895A. Uncommon. June 1978 Welsh et al.: Kaiparowits Flora 165 Oenothera pallida Lindl. Arizona: Coconino Co., east side Glen Canyon Dam, NDA 3448; Colorado River between Glen Canyon Dam and Lee's Fer- ry, NDA & LCH 4022. Utah: Garfield Co., Bryce Canyon, Cottam 4355. Kane Co., Fif- ty Mile Spring, JRM 344; 25 miles south of Cannonville, SLW 5360; Church Wells, JRM 331; Harris Wash road, NDA & DT 5307; Paria River bridge, SLW & NDA 9751; Warm Creek, NDA & RA 2840; 14 miles east of Glen Canyon City, RWA 315; 2 miles west of Glen Canyon City, Nebeker & Skougard 181; East Clark Bench, Harri- son 12084; Willow Tank, Harrison 9093; Cockscomb, JRM 333. San Juan Co., Three Garden, SLW 12421. Abimdant. Oenothera pterosperma Wats. Arizona: Coconino Co., Buckskin Mt., NDA 4430. Utah: Kane Co., Cockscomb, NDA 4622a. Uncommon. Oenothera scapoidea T. & G. var. sca- poidea Utah: Kane Co., Cottonwood Wash road, NDA 5161. Rare. Oenothera walkeri (A. Nels.) Raven Arizona: Coconino Co., Lee's Ferry, Cot- tam 2631. Utah: Kane Co., Labyrinth Can- yon, Lake Powell, NDA 3741; Nipple Bench, NDA & DT 5058; Gunsight Bay, Lake Powell, SLW & NDA 11578; Little Valley Canyon, NDA et al. 2726; Warm Creek, NDA & RA 2905; Colorado Canyon between Glen Canyon Dam and Lee's Fer- ry, NDA & LCH 4025; Paria River LCH & NDA 5386; Dry Rock Creek, Lake Powell, SLW & NDA 11639. Local; common. Orchidaceae (Orchid Family) Corallorhiza maculata Raf.? Utah: San Juan Co., Navajo Mt., NDA 4170. Rare. Epipactis gigantea Dougl. Arizona: Coconino Co., Navajo Creek, NDA & RA 3220; Paria Canyon, Woodruff 1128. Utah: Kane Co., Brigham Plains, ND & RA 2792; Four Mile Bench, SLW & BW 12445; do, SLW 12573; Escalante Canyon, NDA & RA 3218; Little Valley Canyon, NDA et al. 2713; Hole-in-the-Rock, Harri- son 12117; Dry Rock Creek, Lake Powell, SLW & NDA 11640. San Juan Co., Three Garden, SLW 12424. Hanging gardens; common. Habenaria sparsiflora Wats. var. laxiflora (Rydb.) Correll Utah: Kane Co., Four Mile Bench, SLW 12577; do, SLW & BW 12444. Rare. Orobanchaceae (Broomrape Family) Orohanche fasciculata Nutt. var. fascicu- lata Arizona: Coconino Co., 9.3 miles south of Page, NDA & LCH 4018; 4 miles south of Page, NDA 2627. Rare. Orohanche fasciculata Nutt. var. lutea (Parry) Achey Utah: San Juan Co., Gardens Cove, con- fluence of the San Juan, SLW 11932. Rare. Orohanche multiflora Nutt. Utah: Kane Co., Pool Garden, CAT 168. San Juan Co., Gardens Cove, confluence of the San Juan, SLW 11931. Rare. Papaveraceae (Poppy Family) Argemone corymbosa Greene ssp. arenicola G. B. Ownbey Utah: Kane Co., 16 miles east of Glen Canyon City, NDA 3664. Common. Argemone munita Dur. & Hilg. ssp. rotun- data (Rydb.) Ownbey Arizona: Coconino Co., Cockscomb, NDA & RA 2886; Kaiparowits Plateau, NDA & DT 5268. Uncommon. PiNACEAE (Pine Family) Abies concolor (Card. & Glend.) Hoopes Utah: Garfield Co., Canaan Peak, abun- dant, not collected. Locally abundant. Abies lasiocarpa (Hook) Nutt. Utah: San Juan Co., Navajo Mt., NDA et al. 2987; do, NDA & SK 3404. Locally abundant. Picea engelmannii Parry Utah: San Juan Co., NDA & DK 3406. Locally abundant. Picea pungens Engelm. Utah: Bryce Canyon, Buchanan 133 (Bryce). Uncommon. 166 Great Basin Naturalist Vol. 38, No. 2 Pinus edulis Engelm. Utah: Garfield Co., Bryce Ganyon, Got- tam 4364. Kane Go., Four Mile Bench, SL & ERW 12369. San Juan Go., Navajo Mt., NDA at al. 3005. Middle elevations; abun- dant. Pinus flexilis James Utah: Garfield Go., Bryce Ganyon, Harri- son. San Juan Go., NDA & DK 3405. High elevations; uncommon. Pinus hngaeva D. K. Bailey Utah: Garfield Go., Bryce Ganyon, Got- tam 4365. Restricted, and possibly threat- ened (Welsh et al. 1975). Pinus ponderosa Laws. Utah: Kane Go., Hackberry Ganyon, NDA 3691. San Juan Go., Navajo Mt., NDA et al. 2992. Widespread; only locally com- mon to abundant. Pseudotsuga menziesii (Mirb.) Franco Utah: Garfield Go., 10 miles northeast of Henrieville, Harrison 12103. Kane Go., Death Ridge vicinity, SLW 12764. Middle and higher elevations; locally abundant. Plantaginaceae (Plantain Family) Plantago purshii R. & S. Arizona: Goconino Go., 4 miles southeast of Page, RWA 348. Utah: Garfield Go., southeast of Escalante, Holmgren et al. 2039. Kane Go., Gedar Mt., NDA & LGH 3823; Fifty Mile Spring, JRM 352; East Glark Bench, Harrison 12082; Paria River, NDA & LGH 3987; Labyrinth Ganyon, NDA 3739. San Juan Go., Ribbon Garden, NDA 4108. Abundant. PoLEMONiACEAE (Phlox Family) Eriastrum diffusum (Gray) Mason Utah: Kane Go., Kaiparowits Plateau, Harrison 9075; Gedar Mt., NDA & LGH 3819. Uncommon. Cilia aggregata (Pursh) Spreng. Utah: Garfield Go., Bryce Ganyon, Boyle 1115; Escalante, Gottam 4393. Kane Go., Four Mile Bench, NDA 4079; do, SLW et al. 12573; Hole-in-the-Rock, Karren 106; 25 miles southeast of Escalante, Holmgren et al. 2069; Buckskin Gulch, SLW 5311; Esca- lante Ganyon, NDA & RA 3207; Gollett Ganyon, Woodruff 1157. San Juan Go., Rainbow Bridge, SLW & NDA 11663. Gommon. Gilia congesta Hook. Arizona: Goconino Go., Gockscomb, NDA 3059. Utah: Garfield Go., Pet Hollow, NDA 4331; Hackberry Ganyon, NDA & RA 2806. Uncommon. Gilia gunnisonii T. & G. Arizona: Goconino Go., Lee's Ferry, Got- tam 2601; do, NDA 4580; 4 miles southeast of Page, RWA 239; 8 miles south of Page, RWA 225. Utah: Kane Go., East Glark Bench, Harrison 12076; Fifty Mile Spring, JRM 355; Willow Tank, Harrison 9102; Hole-in-the-Rock, White 114; Gedar Mt., NDA 3124; Paria River, NDA & LGH 4006; Warm Greek, NDA 3713; Wahweap Bay, NDA 3550; Tibbet Ganyon, NDA 3604; Nipple Bench, NDA & RA 2814; 14 miles east of Glen Ganyon Gity, RWA 319. Gommon. Gilia hutchinsifolia Rydb. Arizona: Goconino Go., Lee's Ferry, Got- tam 2605; Page vicinity, SLW & NDA 9775. Utah: Kane Go., Gunsight Bay, SLW & NDA 11577. Uncommon. Gilia latifolia Wats. Utah: Kane Go., Warm Greek, NDA & RA 2854; do, NDA & RA 2901; Nipple Greek, NDA & RA 2829; Tibbet Ganyon, NDA 5959. Rare and local; possibly threat- ened (Welsh et al. 1975). Gilia leptomeria Gray Arizona: Goconino Go., Lee's Ferry, NDA 4575, Paria Plateau, NDA & LGH 3782. Utah: Kane Go., Kaiparowits Plateau, Harri- son 9076A; do, NDA & DT 5244; 30 miles east of Glen Ganyon Gity, NDA 4547; Ge- dar Mt., NDA et al. 2744; Willow Tank, Harrison 9109; 25 miles southeast of Esca- lante, Holmgren 2066; 6 miles east of Glen Ganyon Gity, SLW & NDA 9794; Gocks- comb, NDA 3595; Nipple Bench, NDA & DT 5065; Paria River, Gronquist 10135; Warm Greek, NDA & RA 2902; 8 miles southeast of Gannonville, Reveal 803; west of Glen Ganyon Gity, Gronquist 10154. Gommon. Gilia longiflora (Torr.) G. Don Arizona: Goconino Go., Gockscomb, NDA 3053; Paria Plateau, NDA & GM 4291. June 1978 Welsh et al.i Kaiparowits Flora 167 Utah: Kane Co., Cedar Mt., RWA 174; East Clark Bench, Harrison 12073. Uncommon. Gilia micromeria Gray Arizona: Coconino Co., Buckskin Mt., NDA 4931. Utah: Kane Co., NDA & DT 5062. Rare. Gilia opthalmoides Brand Utah: Kane Co., Fifty Mile Spring, JRM 356; Kaiparowits Plateau, Harrison 9076; Nipple Canyon NDA & RA 2830; 25 miles southeast of Escalante, Holmgren et al. 2067. San Juan Co., SLW & NDA 11704. Uncommon. Gilia polycladon Torr. Arizona: Coconino Co., Lee's Ferry, Snow 2; do, NDA 4579. Utah: Kane Co., 22 miles southeast of Escalante, Holmgren et al. 2056; Sit Down Bench, Olson 39; Fifty Mile Spring, JRM 357; Nipple Canyon, NDA & RA 2834A; Paria River NDA & LCH 3986; Little Valley, NDA 3519; Tib- bet Canyon, NDA 3607. Low elevations; common. Gilia subnuda Torr. Arizona: Coconino Co., Paria Canyon, Woodruff 1125. Utah: Garfield Co., Esca- lante, Cottam 4389; 20 miles southeast of Escalante, Holmgren et al. 2047; 1.3 miles south of Cannonville, Reveal et al. 753. Kane Co., Harris Wash road, NDA & DT 5306; Cottonwood Wash Road, SLW 5356; Buckskin Gulch, SLW 5321; Four Mile Bench, SLW 12428; Hackberry Canyon, NDA 3680. Common; widespread. Langloisia setosissima (T. & G.) Greene Utah: Kane Co., 20 miles east of Glen Canyon City, NDA & DT 5072. Low eleva- tions; uncommon. Leptodactylon pungens (Torr.) Nutt. Arizona: Coconino Co., Paria Plateau, NDA & LCH 3789. Rare. Leptodactylon watsonii (Gray) Rydb. Utah: Kane Co., Flat Top, NDA et al. 2961. Rare. Linanthus bigelovii (Gray) Greene Utah: Kane Co., Cockscomb, NDA 4616. Rare. Linanthastrum nuttallii (Gray) Ewan Utah: Bryce Canyon, Coltam 4399. Rare. Microsteris gracilis (Dougl.) Greene Arizona: Coconino Co., Buckskin Mt., NDA 4935. Utah: San Juan Co., Navajo Mt., NDA & DT 5342. Uncommon. Navarretia breweri (Gray) Greene Utah: San Juan Co., Navajo Mt., NDA 4175; do, NDA & DT 5336. Uncommon. Phlox austromontana Coville Utah: Kane Co., Cockscomb, NDA 4610. Rare. Phlox cluteana A. Nels. Utah: San Juan Co., Navajo Mt., NDA 4148. Rare and possibly threatened (Welsh et al. 1975). Phlox hoodii Richards. Utah: Garfield Co., Bryce Canyon, Weight s.n.; Escalante, Cottam 4423. Kane Co., Four Mile Bench, NDA 3487; 20 miles south of Cannonville, Christensen s.n.; Round Valley, NDA 3467; Kaiparowits Plateau, Harrison 9071; 4 miles south of Cannonville, Cronquist 10083; Fifty Mile Spring, JRM 370. Widespread; common. Phlox longifolia Nutt. Utah: Kane Co., Buckskin Gulch, SLW 5316; Cockscomb, NDA 3616; Four Mile Bench, NDA 3758. Widespread; uncommon. Phlox stansburyi (Torr.) Heller. Utah: Kane Co., Kaiparowits Plateau, Harrison 9048; do, NDA & DT 5285; 8 miles south of Paria, Harrison 12054; Paria Townsite, SLW & NDA 9735; 29 miles south of Cannonville, Christensen s.n. San Juan Co., Navajo Mt., NDA et al. 2996. Widespread; uncommon. Polemonium delicatum Rydb. Utah: San Juan Co., Navajo Mt., NDA 4154. Rare. Polemonium viscosum Nutt. Utah: Garfield Co., Bryce Canyon, Cot- tam 2783. Rare. PoLYGALACEAE (Milkwort Family) Polygala subspinosa Wats. Utah: Kane Co., Four Mile Bench, SLW 12788. Widespread; uncommon. PoLYGONACEAE (Buckwhcat Family) Chorizanthe brevicornu Torr. Utah: Kane Co., Escalante Canyon, SLW & GM 11804. Rare. Chorizanthe thurberi (Gray) Wats. Utah: Kane Co., left hand fork of Last Chance Canyon, ca 10 miles east of Four Mile Bench, SL & ERW 12721. Rare. 168 Great Basin Naturalist Vol. 38, No. 2 Eriogonum alatum Torr. var. alatum Arizona: Coconino Co., Paria Plateau, NDA & CM 4294; do, NDA 3060. Utah: Garfield Co., Escalante, Cottam 4398. Kane Co., 25 miles southeast of Escalante, Holm- gren et al. 2070; Four Mile Bench, SLW 12431. Widespread; uncommon. Eriogonum cernuum Nutt. Arizona: Coconino Co., Cockscomb, NDA & DK 3388; do, NDA 3069; Colorado Can- yon between Lee's Ferry and Glen Canyon Dam, NDA 4582. Utah: Garfield Co., Pet Hollow, NDA 4341; Cedar Mt., NDA 4057; Nipple Bench, NDA & RA 2813. Common. Eriogonum corymbosum Benth. var. corymbosum Utah: Garfield Co., Pet Hollow, NDA 4336; Kane Co., Smoky Mt., NDA & NW 3294; 15 miles southeast of Cannonville, SL & SLW 9414; Buckskin Gulch, SL & SLW 9422; Four Mile Bench, NDA 4301. Com- mon. Eriogonum corymbosum Benth. var. glu- tinosum Jones Arizona: Coconino Co., Lee's Ferry, NDA 4353. Utah: Garfield Co., 20 miles south- west of Escalante, Holmgren et al. 2413; 12 miles northeast of Henrieville, Reveal & Re- veal 2887; 4 miles east of Henrieville, NDA 4347; Tibbet Canyon, NDA & DK 3325; 6 miles east of Glen Canyon City, NDA & DK 3353. Low elevations; common. Eriogonum corymbosum Benth. var. orbi- culatum (Stokes) Reveal & Brotherson Utah: Garfield Co., 50 miles southeast of Escalante, Beck s.n. Sandstone basins. Can- yon of the Colorado; uncommon. Eriogonum deflexum Torr. var. deflexum Utah: Garfield Co., 7.5 miles southeast of Escalante, Holmgren et al. 2414. Kane Co., Tibbet Canyon, NDA & DK 3333; Nipple Canyon, NDA & RA 2832; Warm Creek, NDA & RA 2900; Buckskin Wash, SC & SLW 9420; Paria River, SLW 5354. Com- mon. Eriogonum flexum Jones Utah: Garfield Co., 20 miles southeast of Escalante, Holmgren et al. 2045; Kane Co., Cottonwood Wash road. Reveal et al. 825; Paria River, Cronquist 10130; Nipple Can- yon, NDA & RA 2828; Nipple Bench, NDA & DT 5069. Common. Eriogonum gordonii Benth. Utah: Cottonwood Wash road, NDA & RA 2755; do. Reveal et al. 824. Common. Eriogonum heermannii Dur. & Hilg. var. subracemosum (Stokes) Reveal Arizona: Coconino Co., Navajo Mt. road, NDA & DK 3391. Uncommon. Eriogonum hookeri Wats. Utah: Kane Co., head of Alvey Wash, NDA 5209. Uncommon. Eriogonum inflatum Torr. & Frem. var. in- flatum Arizona: Coconino Co., Colorado Canyon, between Glen Canyon Dam and Lee's Fer- ry, NDA & LCH 4026. Utah: Kane Co., Willow Tank, Beck & Tanner s.n.; Paria River, NDA & LCH 3951; Warm Creek, NDA 3716; Cedar Mt., NDA 3106. San Juan Co., Three Garden, SLW 12416. Un- common. Eriogonum inflatum Torr. & Frem. var. fusiforme (Small) Reveal Utah: Kane Co., Cottonwood Wash road. Reveal et al. 823; 15 miles west of Glen Canyon City, NDA & RA 2869; 7 miles east of Glen Canyon City, NDA 2631; 10 miles east of Glen Canyon City, Olson 35. Abundant. Eriogonum leptocladon T. & G. var. papi- liunculum Reveal Arizona: Coconino Co., between Page and Kaibito, Shaw 2646; Cockscomb, NDA & DK 3386; Paria Plateau, NDA 6412. Utah: Kane Co., Cedar Mt., NDA 3132; Church Wells, NDA & DK 3367. Endemic to sandy lower portion of Kaiparowits Ba- sin; locally common. Eriogonum microthecum Nutt. var. folio- sum (T. & G.) Reveal Arizona: Coconino Co., Paria Plateau, NDA 6408. Utah: Garfield Co., 1 mile south of Cannonville, NDA & DK 3316. Kane Co., Four Mile Bench, NDA 4302; 8 miles south of Cannonville, NDA & DK 3318; northwest of Cedar Mt., NDA et al. 2960; Smoky Mt., NDA & NW 3295; Nipple Bench, NDA & DK 3329; Cocks- comb, NDA & DK 3378; Ahlstrom Point road, NDA & RA 2892. Widespread; com- mon. Eriogonum microthecum Nutt. var. laxiflo- rum Hook. June 1978 Welsh et al.: Kaiparowits Flora 169 Utah: Kane Co., Fifty Mile Spring, JRM 359; Cockscomb, NDA 4614; Four Mile Bench, SLW & JRM 12398. Common. Eriogonum palmerianum Reveal Arizona: Coconino Co., Colorado Canyon, between Glen Canyon Dam and Lee's Fer- ry, NDA & LCH 4024; Paria Plateau, NDA 6402. Utah: Kane Co., NDA 4259; Cedar Mt., NDA 3105; Little Valley Canyon, NDA et al. 2730; 25 miles southeast of Es- calante, Holmgren et al. 2061; Willow Tank, Harrison 9103. San Juan Co., east of Navajo Mt., NDA & DT 5347. Common. Eriogonum panguicense (Jones) Reveal var. panguicense Utah: Garfield Co., Bryce Canyon, Cot- tam & Hutchings 2762. Rare and local; pos- sibly threatened (Welsh et al. 1975). Eriogonum racemosum Nutt. Utah: Garfield Co., Bryce Canyon, Cot- tam 2755; Pet Hollow, NDA 4340. San Juan Co., Navajo Mt., NDA et al. 3000. Com- mon. Eriogonum salsuginosum (Nutt.) Hook. Utah: San Juan Co., east of Navajo Mt., NDA & DT 5351. Rare. Eriogonum scabrellum Reveal Utah: Kane Co., Warm Creek, NDA & DK 3334; Smoky Mt., NDA 5958; Rock Creek, NDA & DK 3390. Edaphically re- stricted; uncommon (Utah endemic). Eriogonum shockleyi Wats. var. longilobum (Jones) Reveal Utah: Garfield Co., Escalante, Stanton 1047. Kane Co., Willow Tank, Beck & Tan- ner 2086; Harris Wash road, NDA & DT 5302; Ahlstrom Point road, NDA & RA 2892. Uncommon. Eriogonum subreniforme Wats. Arizona: Coconino Co., Cockscomb, NDA & RA 2879. Utah: Garfield Co., Tropic Val- ley, Holmgren et al. 2412; 1 mile east of Henrieville, NDA 1876. Uncommon. Eriogonum umbellatum Torr. var. sub- aridum Stokes Arizona: Coconino Co., Cockscomb, NDA et al. 3007; do, NDA 3071. Utah: Garfield Co., Pet Hollow, NDA 4332. San Juan Co., Iceberg Canyon, Lake Powell, NDA 4133; Navajo Mt., Gentry & Davidse 1777; do, NDA et al. 2966. Uncommon. Eriogonum wetherillii Eastw. Arizona: Coconino Co., Page vicinity, SLW & NDA 9772; do. Reveal 2911. Utah: Garfield Co., 45 miles south of Escalante, Beck & Tanner 8236. Kane Co., Warm Creek, NDA 3717; Paria River, NDA & LCH 3984; Little Valley Canyon, NDA et al. 2729. San Juan Co., east of Navajo Mt., 5348. Common. Polygonum aviculare L. Utah: Garfield Co., 45 miles south of Es- calante, Beck & Tanner s.n. Rare. Polygonum douglasii Greene Utah: Garfield Co., Bryce Canyon, Bu- chanan 232 (Bryce). Rare. Polygonum sawatchense Small Utah: San Juan Co., Navajo Mt., NDA & DK 3400. Rare. Rumex fueginus Phil. Utah: Garfield Co., Escalante Canyon, Beck s.n., without date. Rare. Rumex hymenosepalus Torr. Arizona: Coconino Co., 3 miles south of Wahweap Marina, SLW & NDA 9766; Glen Canyon Dam vicinity, NDA & DT 3426. Utah: Kane Co., Willow Tank, Harri- son 9094; 14 miles east of Glen Canyon City, RWA 323. Common. Rumex venosus Pursh. Utah: Kane Co., 15 miles west of Glen Canyon City, NDA & RA 2867. Rare. PoLYPODiACEAE (Fern Family) Adiantum capillus-veneris L. Arizona: Coconino Co., Paria Canyon, Woodruff 1127. Utah: Kane Co., Hole-in- the-Rock, Harrison 12137; Paria River, LCH & NDA 5388; 31.5 miles east of Glen Canyon City, NDA et al. 2705; Reflection Canyon, SLW & CAT 11859; mile 47, Lake Powell, NDA et al. 3236; Brigham Plains, NDA & RA 2801. San Juan Co., Three Gar- den, SLW & NDA 11673. Hanging gardens; abundant. Cheilanthes feei Moore Utah: Kane Co., Cockscomb, NDA 4628. Rare. Pellaea longimucronata Hook. Utah: San Juan Co., Trail Canyon, Lake Powell, SLW 11910. Rare. 170 Great Basin Naturalist Vol. 38, No. 2 PoRTULACACEAE (PuTslanc Family) Lewisia pygmaea (Gray) Robins. Utah: San Juan Co., Navajo Mt., NDA 4182. Rare. Portulaca mundula Johnst. Arizona: Coconino Co., Paria Plateau, NDA & GM 4281. Utah: Kane Co., Cedar Mt., NDA et al. 3255, do, NDA 3104A. Rare. Portulaca oleracea L. Utah: Kane Co., Cockscomb, NDA & DK 3376. Rare. Talinum breviflorum Torr. Arizona: Coconino Co., north of Shouts, NDA & DT 5330. Rare. Talinum parviflorum Nutt. Arizona: Coconino Co., Cockscomb, NDA 3061. Utah: Kane Co., Cedar Mt., NDA et al. 3253. Rare. Primulaceae (Primrose Family) Androsace septentrionalis L. Utah: Garfield Co., Escalante Canyon, Beck s.n. Kane Co., Cockscomb, NDA 4618. San Juan Co., Navajo Mt., Gentry & Da- vidse 1771. Uncommon. Dodecatheon pulchellum (Raf.) Merrill Utah: Kane Co., Little Valley Canyon, SLW & NDA 11611. Rare. Primula specuicola Rydb. Utah: Kane Co., Hole-in-the-Rock, Karren 102. San Juan Co., Three Garden, SLW & NDA 11668; do, SLW 11905; east of con- fluence of San Juan, NDA & RA 3157; do, NDA 4092. Restricted, local, and threatened (Welsh et al. 1975). PoTAMOGETONACEAE (Pondweed Family) Potam.ogeton pusillus L. Utah: Kane Co., Escalante Canyon, NDA 4208. Rare. Pyrolaceae (Wintergreen Family) Pterospora andromedea Nutt. Utah: Garfield Co., Bryce Canyon, Vogt s.n. (Bryce). Rare. Pyrola secunda L. Utah: San Juan Co., Navajo Mt., NDA 4189; do, Gentry & Davidse 1773. Rare. Ranunculaceae (Buttercup Family) Aconitum, columbianum Nutt. Utah: Garfield Co., Escalante Canyon, Beck s.n., 1936. Rare. Anemone tuberosa Rydb. Utah: San Juan Co., Rainbow Bridge, NDA 3561. Rare. Aquilegia coerulea James Utah: Garfield Co., Buchanan 161 (Bryce). Rare. Aquilegia formosa Fisch. Utah: Kane Co., Four Mile Bench, SLW & BW 12440; do, SLW 12573a. Rare. Aquilegia micrantha Eastw. Arizona: Coconino Co., Paria Canyon, Woodruff 1137. Utah: Kane Co., Four Mile Bench, SLW & BW 12440a; 31.5 miles east of Glen Canyon City, NDA et al. 2698; Last Chance Canyon, 3 miles north of Four Mile Spring, NDA 4076; Escalante River, NDA 4225; do, SLW & GM 11817; Crosby Canyon, NDA & RA 2889. San Juan Co., Three Garden, SLW & NDA 11691; Rain- bow Bridge, SLW & NDA 11658. Abun- dant; hanging gardens. Aquilegia scopulorum Tidestr. Utah: Garfield Co., Bryce Canyon, Cot- tam 2764. Rare. Aquilegia shockleyi Eastw. Utah: Garfield Co., Escalante, Cottam 4383, 4390; north of Escalante, NDA 440. Rare. Clematis ligusticifolia Nutt. Utah: Kane Co., Willow Tank, Beck & Tanner s.n.; Wahweap Bay, NDA 3135. San Juan Co., east of confluence of San Juan, NDA & RA 3191. Abundant. Clematis pseudoalpina (Kuntze) A. Nels. Utah: Garfield Co., Death Ridge, NDA 5184; Canaan Peak, SLW 12829; do, SLW 12878. Rare. Delphinium menziesii DC. ssp. utahense (Wats.) Sutheriand Utah: Kane Co., Kaiparowits Plateau, Harrison 9050; do, NDA 5227. Rare. Delphinium scaposum, Greene Arizona: Coconino Co., Paria Plateau, NDA & LCH 3783. Utah: Garfield Co., 1.5 miles south of Cannonville, Reveal et al. 767. Kane Co., 40 miles east of Kanab, Cronquist 10141; Hole-in-the-Rock, White 115; Kaiparowits Plateau, NDA & DT June 1978 Welsh et al.: Kaiparowits Flora 171 5261; Dry Rock Creek, Lake Powell, SLW & NDA 11614; Cedar Mt., RWA 337. Abundant. Myosurus minimus Pallas Arizona: Coconino Co., Buckskin Mt., NDA 4933. Rare. Ranunculus aquatilis L. Utah: Garfield Co., Escalante, Cottam 4431. Rare. Ranunculus cymbalaria Pursh. Arizona: Coconino Co., Colorado Canyon, between Glen Canyon Dam and Lee's Fer- ry, NDA & LCH 4023. Rare. Ranunculus glaberrimus Hook. var. ellip- ticus Greene Utah: San Juan Co., NDA & DT 5344. Rare. Ranunculus juniperinus Jones Utah: Kane Co., Cockscomb, NDA 3583. Rare. Ranunculus orogenes Greene Utah: San Juan Co., Navajo Mt., NDA 4186. Rare. Thalictrum fendleri Engelm. Utah: San Juan Co., Navajo Mt., NDA et al. 2993. Rare. Rhamnaceae (Buckthorn Family) Ceonothus fendleri Gray. Utah: San Juan Co., Navajo Mt., Gentry & Davidse 1967; do, NDA et al. 2974. Un- common; higher elevations. Ceonothus martini Jones Utah: Garfield Co., Bryce Canyon, Boyle 1113. Uncommon; higher elevations. Rhamnus betulaefolia Greene Arizona: Coconino Co., Navajo Creek, NDA 3723. Utah: Garfield Co., Escalante Canyon, Beck s.n. Kane Co., 31.5 miles east of Glen Canyon City, NDA et al. 2699; Es- calante Canyon, SLW & CM 11813; Hole- in-the-Rock, Harrison 12107; Reflection Canyon, SLW & CAT 11848. San Juan Co., east of confluence of San Juan, NDA & RA 3167; Three Garden, SLW 12422. Canyons and hanging gardens at lower elevations; common. Rosaceae (Rose Family) Amelanchier utahensis Koehne Utah: Garfield Co., Cottam 4375. Kane Co., Tibbet Canyon, NDA 3460; 20 miles south of Cannonville, Christensen s.n.; Kai- parowits Plateau, NDA & DT 5264; Paria Townsite, SLW & NDA 9732; Four Mile Bench, NDA 3759; Cottonwood Wash, NDA 3593; Tibbet Canyon, NDA 3460; Warm Creek, NDA & RA 2853. Common. Cercocarpus intricatus Wats. Utah: Garfield Co., Escalante, Cottam 4416; do. Beck & Tanner s.n. Kane Co., Kaiparowits Plateau, Harrison 9081; Four Mile Bench, NDA et al. 3505. Uncommon. Cercocarpus montanus Raf. Utah: Garfield Co., Bryce Canyon, Harri- son 12322. Kane Co., Kaiparowits Plateau, NDA & DT 5265. Uncommon. Chamaebatiaria millefolium (Torr.) Maxim. Utah: Kane Co., Cockscomb, NDA 4410. Rare. Coleogyne ramosissima Torr. Arizona: Coconino Co., Page vicinity, SLW & NDA 9786; 5 miles southeast of Page, Nebeker & Skougard 197; 8 miles south of Page, RWA 224. Utah: Cedar Mt., NDA et al. 2752; 35 miles southeast of Es- calante, Harrison 9117; Grand Bench, NDA 3536; Hole-in-the-Rock, White 124; 14 miles east of Glen Canyon City, RWA 321; Dry Rock Creek, Lake Powell, SLW & NDA 11638. Codominant in blackbrush communities; abundant. Cowania mexicana D. Don Arizona: Coconino Co., Cedar Mt., NDA et al. 2754. Utah: Kane Co., Willow Tank, Harrison 9112; 50 miles southeast of Esca- lante, Beck & Tanner s.n.; Dry Rock Creek, Lake Powell, SLW & NDA 11630; Cocks- comb, SLW 5336; Cedar Mt., RWA 184; Four Mile Bench, SLW & JRM 12412. San Juan Co., Navajo Mt., NDA 4142; do, NDA et al. 2983. Widespread; uncommon. Fallugia paradoxa (D. Don) Endl. Utah: Kane Co., Paria River, NDA & RA 2779; Kaiparowits Plateau, NDA & DT 5291, Cockscomb, JRM 340; Tibbet Can- yon, NDA 3760; Little Valley Canyon, NDA et al. 2717; 5 miles east of Paria, Harrison 12311. Canyon bottoms; locally common. Holodiscus dumosus (Nutt.) Heller Utah: Garfield Co., Bryce Canyon, Cot- tam 2780; Escalante, Cottam 4373. Kane 172 Great Basin Naturalist Vol. 38, No. 2 Co., Hackberry Canyon, NDA 3690. Rare. Ivesia sabulosa (Jones) Keck Arizona: Coconino Co., Paria Plateau, NDA & CM 4295. Possibly threatened (Welsh et al. 1975). Peraphyllum ramosissinium Nutt. Utah: Garfield Co., Bryce Canyon, Bu- chanan 219. Rare. Petrophytum caespitosum (Nutt.) Rydb. Utah: Kane Co., Reflection Canyon, SLW & CAT 11857. San Juan Co., east of con- fluence of San Juan, NDA et al. 3226. Hanging gardens; locally abundant. Potentilla anserina L. Utah: Garfield Co., Bryce Canyon, Weight s.n. Uncommon. Potentilla concinna Richards. Utah: Garfield Co., Canaan Peak, NDA & DT 5320. San Juan Co., Navajo Mt., NDA & DT 5345; do, NDA 4172, 4173. Higher elevations; uncommon. Prunus virginiana L. Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5230. Rare. Purshia tridentata (Pursh) DC. Utah: Garfield Co., Bryce Canyon, Harri- son 12041. Kane Co., Cockscomb, NDA 3540; Kaiparowits Plateau, Harrison 9080. Uncommon. Rosa neomexicana Cockerell Utah: San Juan Co., Navajo Mt., NDA 4135; do, NDA et al. 2999. Rare. Rosa woodsii Lindl. Utah: Garfield Co., Escalante, Cottam 4421; do, Beck & Tanner 8235. Kane Co., Hole-in-the-Rock, Harrison 12139. San Juan Co., Ribbon Canyon, NDA 4103. Uncom- mon. Rubus neomexicanus Gray Utah: San Juan Co., Ribbon Canyon, SLW & NDA 11710; do, SLW & CM 11782; do, NDA 4102. This is the first known Utah record; rare. RuBiACEAE (Madder Family) Galium bifolium Wats. Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5222. Uncommon. Galium stellatum Kellogg Utah: Garfield Co., Escalante, Cottam 4411. Kane Co., Driftwood Canyon, Lake Powell, SLW & NDA 11644. Rare. RuTACEAE (Rue Family) Ptelea trifoliata L. ssp. pallida (Greene) V.L. Bailey Utah: Garfield and Kane counties, Bailey (1962), possibly extinct. Thamnosoma montana Torr. & Frem. Arizona: Coconino Co., Glen Canyon Dam, NDA & DT 3424; Navajo Canyon, NDA & RA 3145. Kane Co., Cedar Mt., NDA 4425. Rare. Salicaceae (Willow Family) Populus angustifolia James Utah: Garfield Co., Bryce Canyon, Weight B-31/148. Higher elevations; un- common. Populus fremontii Wats. Utah: Kane Co., Little Valley Canyon, NDA 3428, 3419. Drainage bottoms; com- mon. Populus tremuloides Michx. Utah: San Juan Co., Navajo Mt., NDA et al. 2988. Rare to uncommon. Salix amygdaloides Anderss. Utah: Kane Co., Escalante Canyon, NDA & RA 3214; Reflection Canyon, SLW & CAT 11856. San Juan Co., Ribbon Canyon, NDA 4106. Uncommon. Salix bebbiana Sarg. Utah: San Juan Co., Navajo Mt., NDA 4166. Uncommon. Salix exigua Nutt. Utah: Garfield Co., Bryce Canyon, Cot- tam 2779; Escalante, Cottam 4397. Kane Co., Nipple Canyon, NDA & RA 2833; Four Mile Bench, SLW et al. 12585; Paria River, NDA & LCH 4010. Common; drain- ages, seeps, and springs. Salix lutea Nutt. Utah: San Juan Co., Three Garden vicin- ity, SLW 11894. Uncommon. Salix scouleriana Barrett Utah: San Juan Co., Navajo Mt., NDA et al. 2989. Uncommon. Santalaceae (Sandalwood Family) Comandra umbellata (L.) Nutt. var. pallida (A. DC.) Jones Utah: Garfield Co., 1.3 miles south of Cannonville, Reveal et al. 747. Kane Co., June 1978 Welsh et al.: Kaiparowits Flora 173 Kaiparowits, Plateau, NDA & DT 5278; East Clark Bench, NDA & RA 2685; Fifty Mile Spring, JRM 369; Cedar Mt., RWA 187; Cockscomb, NDA 3539, 1972; Four Mile Bench, SLW 12430. Common. Saxifragaceae (Saxifrage Family) Heuchera rubescens Torr. Utah: Garfield Co., Escalante, Cottam 4410. San Juan Co., Navajo Mt., Gentry & Davidse 1781; do, NDA 4188. Rare. Lithophragma tenella Nutt. Arizona: Coconino Co., Cockscomb, NDA 4942. Utah: Kane Co., Kaiparowits Plateau, Harrison 9049. San Juan Co., Navajo Mt., NDA & DT 5337. Uncommon. Philadelphus microphyllus Gray Utah: San Juan Co., Navajo Mt., Gentry & Davidse 1768; do, NDA et al. 2980. Rare. Ribes cerewn Dougl. Utah: Garfield Co., Bryce Canyon, Bu- chanan 105. Uncommon. Ribes leptanthum Gray Utah: Kane Co., Kaiparowits Plateau, NDA 529; do, NDA & DT 5266. San Juan Co., NDA 4134. Uncommon. Ribes montigenum McClatchie Utah: San Juan Co., Navajo Mt., NDA 4153. Rare. Ribes velutinum Greene Utah: Kane Co., Buckskin Mt., NDA 4649. Rare. ScROPHULARiACEAE (Figwort Family) Castilleja chromosa A. Nels. Utah: Garfield Co., Escalante, Cottam 4415; 1.5 miles south of Cannon ville. Re- veal et al. 766; 20 miles southeast of Can- nonville, Holmgren et al. 2049. Kane Co., Kaiparowits Plateau, Harrison 9055; Bryce Canyon, NDA 6127; 14 miles east of Glen Canyon City, RWA 213; Smoky Mt., Cronquist 10020; Cottonwood Wash road. Reveal et al. 808; Paria Townsite, SLW & NDA 9724; Buckskin Gulch, SLW 5310; Nipple Canyon, NDA 3479; Four Mile Bench, SLW & NDA 12377; Brigham Plains, NDA et al. 3488. San Juan Co., Three Garden, SLW 12415; Navajo Mt., NDA 4140. Abundant. Castilleja exilis A. Nels. Utah: Garfield Co., Escalante Canyon, Beck 113; do, Holmgren et al. 2119. Rare. Castilleja linariaefolia Benth. Arizona: Coconino Co., Paria Plateau, NDA 6397. Utah: Garfield Co., 5 miles west of Escalante, Anderson 684. Kane Co., Reflection Canyon SLW 11882; Hole-in-the- Rock, Harrison 12106; Escalante Canyon, NDA & RA 3206; Four Mile Bench, SLW et al. 12579; Cottonwood Wash road, NDA & DK 3321; Hole-in-the-Rock, Karren 105; east of confluence of San Juan, NDA & RA 3174, 3186; Navajo Mt., Gentry & Davidse 1783; do, NDA et al. 2972. Abundant. Castilleja revealii N. Holmgren Utah: Garfield Co., Bryce Canyon, Holm- gren & Reveal 2017. Rare, endemic, and possibly endangered (Welsh et al. 1975). Castilleja scabrida Eastw. Utah: Garfield Co., 12 miles east of Esca- lante, Harrison 9129; southeast of Escalante, NDA 1242. Kane Co., 16 miles east of Glen Canyon City, NDA 3666. Common. Collinsia parviflora Dougl. Utah: San Juan Co., NDA & DT 5340; do, NDA 4185. Rare. Cordylanthus parviflorus (Ferris) Wiggins Utah: Kane Co., 30 miles east of Kanab, NDA 5587. Rare. Cordylanthus wrightii Gray Arizona: Coconino Co., Paria Plateau, NDA 6413. Utah: Kane Co., Escalante Riv- er, NDA 4222; East Clark Bench, NDA & DK 3372; Cedar Mt., NDA 3110. Uncom- mov. Mimulus eastwoodiae Rydb. Utah: Kane Co., Escalante River, NDA & RA 3202, 3217, 3219. San Juan Co., Rain- bow Bridge, NDA & NW 3303; Three Gar- den, SLW & NDA 11677. Hanging gardens; restricted and uncommon. Mimulus glabratus H.B.K. Utah: Kane Co., Cottonwood Wash, SLW 12591; do, NDA & LCH 3833. Uncommon. Mimulus guttatus Fisch. var. guttatus Arizona: Coconino Co., Paria Canyon, Woodruff 1131, 1132. Uncommon. Mimulus guttatus DC. var. depauperatus (Gray) Grant Utah: Kane Co., Escalante Canyon, NDA & RA 3205. Uncommon. 174 Great Basin Naturalist Vol. 38, No. 2 Mimulus rubellus Gray Arizona: Coconino Co., Cockscomb, NDA 4932, 4946. Utah: Kane Co., Cockscomb, NDA 4620; Buckskin Mt., NDA 4655. San Juan Co., Navajo Mt., NDA & DT 5343. Uncommon. Mimulus suksdorfii Gray Utah: Kane Co., Kaiparowits Plateau, NDA & DT 5221. Uncommon. Orthocarpus purpureo-albus Gray Utah: Garfield Co., Bryce Canyon, Weight s.n. Uncommon. Pedicularis centranthera Gray Utah: Garfield Co., Bryce Canyon, Bur- key 135; Death Ridge, NDA 5179. Kane Co., Cottonwood Wash road, LCH & NDA 5254. San Juan Co., Navajo Mt., NDA et al. 2995; do, NDA 4158. Uncommon. Penstemon ambiguus Torr. Arizona: Coconino Co., Glen Canyon Dam vicinity, Reveal et al. 838, Paria Plateau, NDA & CM 4280. Utah: Garfield Co., Escalante, NDA 747. Kane Co., 50 miles south of Escalante, Beck & Tanner s.n.; East Clark Bench, NDA & RA 2683; Paria River, NDA & LCH 3942; Cedar Mt., NDA 3120A. San Juan Co., east of con- fluence of San Juan, NDA 4096. Low eleva- tions; common. Penstemon angustifolius Nutt. ssp. elatus Crosswhite Utah: Garfield Co., Escalante, Cottam 4385a; east of Escalante, Harrison 9136b. Uncommon. Penstemon barbatus (Cav.) Roth. ssp. tor- reyi (Benth.) Keck Utah: Garfield Co., north of Escalante, Holmgren et al. 2423. Rare. Penstemon bracteatus Keck Utah: Garfield Co., Bryce Canyon, Holm- gren et al. 2018. Rare. Penstemon bridgesii Gray Arizona: Coconino Co., Paria Plateau, NDA & CM 4276. Utah: Garfield Co., Pet Hollow, NDA 4322. Kane Co., Cockscomb, NDA 5485. San Juan Co., NDA & DK 3398; do, NDA et al. 2964; do, Gentry & Davidse 1793. Uncommon. Penstemon caespitosus Nutt. ssp. perbrevis Pennell Utah: Garfield Co., Bryce Canyon, Boyle 1138; Utah endemic; restricted edaphically and uncommon. Penstemon comarrhenus Gray Utah: Garfield Co., Bryce Canyon, Cot- tam 2757; Escalante, Cottam 4386; Kane Co., Four Mile Bench, NDA 5357; Collet Wash, NDA 1869; 25 miles southeast of Es- calante, Holmgren et al. 2059. San Juan Co., Navajo Mt., Gentry & Davidse 1780; do, NDA et al. 2965. Middle elevations; widespread and uncommon. Penstemon confusus Jones Utah: Garfield Co., Escalante, White 137. Kane Co., Four Mile Bench, SLW 12427. Uncommon. Penstemon eatonii Gray ssp. undosus (Jones) Keck Utah: Garfield Co., Escalante, Cottam 4386a. Kane Co., Hole-in-the-Rock, Harri- son 12116; Cockscomb, SLW 5344; Kai- parowits Plateau, NDA & DT 5260. Penstemon jamesii Benth. ssp. ophianthus (Pennell) Keck Utah: Garfield Co., 11 miles east of Hen- rieville, Holmgren et al. 2019; Death Ridge, NDA 5177. Kane Co., Collet Wash, NDA s.n. Uncommon. Penstemon laevis Pennell Utah: Garfield Co., near Bryce Canyon, Burkey 165. Kane Co., Cottonwood Wash, NDA & RA 2803; Paria River, NDA & LCH 3998; 3 miles north of Four Mile Spring, NDA 4080. Uncommon. Penstemon leiophyllus Pennell Utah: Garfield Co., Bryce Canyon, Weight 6663. Kane Co., Bryce Canyon, Boyle 1131. Uncommon and possibly threat- ened (Welsh et al. 1975). Penstemon lentus Pennell ssp. lentus Utah: Garfield Co., Escalante, Cottam 4419. Rare. Penstemon pachyphyllus Gray Utah: Garfield Co., Escalante, Cottam 4425. Kane Co., 4 miles south of Cannon- ville, Reveal 782. Uncommon. Penstemon palmeri Gray Utah: Kane Co., Paria River, NDA & LCH 3999; Warm Creek, NDA & RA 2852; Last Chance Creek, 3 miles north of Four Mile Spring, NDA 4081; Collet Wash, NDA 1871. Uncommon. Penstemon thompsoniae (Gray) Rydb. June 1978 Welsh et al.: Kaiparowits Flora 175 Arizona: Coconino Co., Cockscomb, NDA 4945. Utah: Kane Co., Four Mile Bench, RWA 439; do, NDA 5170. Rare. Penstemon utahensis Eastw. Utah: Garfield Co., Bryce Canyon, Cot- tam 4354; 4 miles southwest of Cannonville, Reveal et al. 769. Kane Co., Kaiparowits Plateau, Harrison 9061, 9065; Cockscomb, NDA 3546; Buckskin Gulch, SLW 5306; Little Valley Canyon, NDA 3521; Cotton- wood Wash, Moore 403; Smoky Mt., Cronquist 10021; Paria Townsite, SLW & NDA 9725; 3 miles southeast of Escalante, Buss 107; Fifty Mile Spring, JRM 353. San Juan Co., Navajo Mt., King s.n. Common. Penstemon venosus (Keck) Reveal Utah: Garfield Co., east of Escalante, Harrison 9136a. Uncommon Veronica anagallis-aquatica L. Arizona: between Lee's Ferry and Glen Canyon Dam, NDA & LCH 4020. Utah: Kane Co., Escalante Canyon, NDA 4129. Uncommon. SoLANACEAE (Nightshade Family) Chamaesarcha coronopus (Dunal) Gray Arizona: Coconino Co., Cockscomb, NDA & RA 2888a. Uncommon. Datura meteloides DC. Utah: Kane Co., Willow Tank, Harrison 9092; Hole-in-the-Rock, Harrison 12109; base of Cedar Mt., NDA 2917; Escalante Canyon, NDA & RA 3199; do, SLW & CM 11808. Low elevations; common. Lycium andersonii Gray Utah: Kane Co., Last Chance Creek, NDA et al. 3515; base of Smoky Mt., NDA 3440. Uncommon. Lycium pallidum Miers Arizona: Coconino Co., Paria Plateau, NDA & LCH 3785; Cedar Mt., NDA et al. 2749. Utah: Kane Co., Cedar Mt., NDA 3559. Uncommon. Nicotiana attenuata Torr. Utah: San Juan Co., Navajo Mt., Gentry & Davidse 1789; do, NDA et al. 2968; do, NDA & DK 3397. Uncommon. Nicotiana trigonophylla Dunal Arizona: Coconino Co., east side of Glen Canyon Dam, NDA 3449. Utah: Kane Co., Wahweap Bay, Lake Powell, NDA 3701; Escalante Canyon, SLW & CM 11807; Dry Rock Creek, Lake Powell, SLW & NDA 11631; Gunsight Bay, Lake Powell, SLW & NDA 11583. Uncommon. Physalis fendleri Gray Arizona: Coconino Co., Paria Plateau, NDA & CM 4292. Rare. Solanum elaeagnifolium Cav. Utah: Kane Co., 6 miles west of Glen Canyon City, NDA 2922. Rare. Solanum triflorum Nutt. Utah: Kane Co., Willow Tank, Beck & Tanner s.n.; Escalante Canyon, SLW & CM 11806. Rare. Tamaricaceae (Tamatix Family) Tamarix parvifolia DC. Utah: Kane Co., Paria River, NDA 3577. Uncommon. Tamarix ramosissima Ledeb. Arizona: Coconino Co., between Lee's Ferry and Glen Canyon Dam, NDA & LCH 4034. Utah: Kane Co., Hole-in-the-Rock, Harrison 12120; Paria River, Cronquist 10200; Cottonwood Wash road, NDA & RA 2768; Halls Creek Bay, Lake Powell, SLW & CM 11794; Four Mile Bench, SLW et al. 12582. Abundant. Typhaceae (Cattail Family) Typha domingensis Pers. Utah: Kane Co., Escalante Canyon, NDA 4214; Four Mile Bench, SLW et al. 12602. Uncommon. Ulmaceae (Elm Family) Celtis reticulata Torr. Arizona: Coconino Co., Lee's Ferry, Cot- tam 2625. Utah: Kane Co., Hole-in-the- Rock, Harrison 12108; Escalante Canyon, SLW & CM 11818; Reflection Canyon, SLW & CAT 11860. San Juan Co., Three Garden, SLW & NDA 11689; east of con- fluence of San Juan, NDA & RA 3181A. Canyons; locally common. Umbelliferae (Parsley Family) Cymopterus fendleri Gray Utah: Kane Co., Willow Tank, JRM 387a; 176 Great Basin Naturalist Vol. 38, No. 2 base of Smoky Mt., NDA 3439; East Clark Bench, NDA et al. 3493; 30 miles east of Glen Canyon City, NDA 4549; Little Valley Canyon, NDA 3530. Common. Cymopterus newberryi (Wats.) Jones Arizona: Coconino Co., 3 miles south of Wahweap Marina, SLW & NDA 9769. Utah: Kane Co., 14 miles east of Glen Can- yon City, RWA 119; 6 miles southeast of Glen Canyon City, RWA 133; between Nipple Bench and Lake Powell, Cronquist 10016; Paria River, NDA & LCH 3964; Warm Creek, NDA 3714; Willow Tank, JRM 387. Common. Cymopterus multinervatus (C. & R.) Tidestr. Utah: Kane Co., Cottonwood Wash Road, SLW 5357; Paria Townsite, SLW & NDA 9726; Cockscomb, NDA 4665; East Clark Bench, NDA et al. 3494; 13 miles east of Warm Creek, NDA et al. 3511. Low eleva- tions; common. Cymopterus purpurascens (Gray) Jones Utah: Kane Co., Nipple Canyon, NDA 3481; Cedar Mt., RWA 171; Smoky Mt., RWA 126. Common. Cymopterus purpureus Gray Arizona: Coconino Co., Cockscomb, NDA 4926. Utah: Kane Co., Kaiparowits Plateau, Harrison 9064; do, JRM 406; Buckskin Gulch, SLW 5317; Cottonwood Wash road, Christensen s.n.; Nipple Creek, NDA 3483; Cockscomb, NDA 4630A. Common. Lomatium foenicuUiceum (Nutt.) C. & R. var. macdougalii (C. & R.) Cronq. Arizona: Coconino Co., Cockscomb, NDA 4934. Utah: Kane Co., Four Mile Bench, NDA et al. 3506. Uncommon. Lomatium juniperinum (Jones) C. & R. Utah: Garfield Co., NDA & DT 5319. Rare. Lom^atium minim^um Mathias Utah: Garfield Co., Bryce Canyon, Cot- tam, Harrison & Stanton 4360; do. Weight s.n., Aug. 1934. Endemic, rare, and threat- ened (Welsh et al. 1975). Lomatium nevadense (Wats.) C. & R. Utah: Kane Co., Kaiparowits Plateau, Harrison 9047; do, NDA & DT 5247. Rare. Lomatium, parryi (Wats.) Macbr. Arizona: Coconino Co., 9 miles south of Page, RWA 219. Utah: San Juan Co., Nav- ajo Mt., NDA 4144. Uncommon. Urticaceae (Nettle Family) Parietaria pennsylvanica Muhl. Utah: Kane Co., Escalante Canyon, SLW & CM 11805; Cockscomb, NDA 4413; do, NDA 4627. Uncommon. VioLACEAE (Violet Family) Vto^ canadensis L. Utah: San Juan Co., Navajo Mt., NDA 4155. Rare. Viola nephrophylla Greene Utah: Garfield Co., Escalante Canyon, Harrison 9148. Rare. ViscACEAE (Dwarf Mistletoe Family) Arceuthobium vaginatum (H.B.K.) Eichler Utah: San Juan Co., Navajo Mt., NDA et al. 2991. Uncommon. Phoradendron juniperinum Engelm. Utah: Kane Co., Four Mile Bench, NDA 4308; Cockscomb, NDA 3547. Common. ViTACEAE (Grape Family) Parthenocissus inserta (Kerner) K. Frtsch. Utah: Kane Co., Hole-in-the-Rock, Harri- son 12138; Escalante Canyon, SLW & CM 11816; do, SLW & CAT 11872; do, NDA 4127; Reflection Canyon, SLW & CAT 11853. San Juan Co., Rainbow Bridge, SLW & NDA s.n. Uncommon. Zannichelliaceae (Homed Pondweed Family) Zannichellia palustris L. Arizona: Coconino Co., Lee's Ferry, NDA 4350. Reflection Canyon, SLW & CAT 11855. Uncommon. Zygophyllaceae (Caltrop Family) Tribulus terrestris L. Arizona: Coconino Co., 8 miles southeast of Page, NDA et al. 3241; Paria Plateau, NDA & CM 4289. Uncommon. June 1978 Welsh et al.: Kaiparowits Flora 177 Literature Cited (Plants) Al-shebaz, I. A. 1973. The biosystematics of the genus Thelypodium (Cruciferae). Contr. Gray Herb. 204: 1-148. Bailey, V. L. 1962. Revision of the genus Ptelea (Ru- taceae). Brittonia 14:1-45. Barneby, p. C. 1964. Atlas of North American Astra- galus. Mem. N.Y. Bot. Card. 13:1-1188. Buchanan, H., and G. T. Nebeker. 1971. Checkhst of higher plants of Bryce Canyon National Park, Utah. Weber State College, Ogden, Utah. 30 pp. 1974. Wildflower communities of Bryce Canyon and Cedar Breaks. Bryce Canyon Natural His- tory Association, Bryce Canyon, Utah. 25 pp. Clover, E. U., and L. Jotter. 1941. Cacti of the Col- orado River and tributaries. Bull. Torrey Bot. Club 68:409-419. Eastwood, A. 1893. Plants collected in southeastern Utah. Zoe 4:123-125. Flovvtrs, S. 1959. Vegetation in Glen Canyon, pp. 21-61. In: Ecological studies of the flora and fauna in Glen Canyon, Angus M. Woodbury et al. Univ. Utah Anthro. Papers No. 40, Glen Canyon Ser. No. 7:63-72. Gaines. X. M. 1957. Plants in Glen Canyon Plateau 30(2):31-34. 1960. An annotated catalogue of Glen Canyon plants. Museum of Northern Arizona Technical Series, No. 4. Gregory, H. E. 1947. Colorado drainage basin. Amer. J. Sci. 245:694-705. Haring, I. N. 1961. Mosses of the Glen Canyon area. Plateau 33(4): 120-22. Harrington, H. D. 1954. Manual of the plants of Col- orado. Sage Books, Denver, Colorado. 666 pp. Haskell, H. S. 1958. Flowering plants in Glen Can- yon—late summer aspect. Plateau 31(1): 1-3. Kearney, T. H., and R. H. Peebles. 1964. Arizona flora. University of California Press, Berkeley. 1085 pp. Lindsay, D. S. 1959. Vascular plants collected in Glen Canyon. Appendix A in Ecological studies of the flora and fauna in Glen Canyon, Angus M. Woodbury et al. Univ. Utah Anthro. Papers No. 40, Glen Canyon Ser. No. 25:176-195. Martin, P. S. 1964. Pollen analysis in the Glen Can- yon. Addendum in 1962 excavation. Glen Can- yon area, Floyd W. Sharrock, et al. Univ. Utah Anthro. Papers. No. 73, Glen Canyon Ser. No. 25:176-195. McDouGALL, W. B. 1959. Plants of the Glen Canyon area in the herbarium of the Museum of North- em Arizona. Museum of Northern Ariz., Flag- staff. Mimeographed. Morgan, D. L. 1947. The exploration of the Colorado River in 1869. Utah Hist. Quarterly 15. 270 pp. Plair, T. B. (1939). Vegetative cover of the Utah Parks. Zion and Bryce Nat. Notes 6(3):34-35. Reveal, J. L. 1970. Comments on Lesquerelia hitch- cockii. Great Basin Nat. 30:94-98. Tidestrom, I. 1925. Flora of Utah and Nevada. Contr. U.S. Natl. Herb. 25:1-665. Weight, K. E. 1932. The limber pine. Zion and Bryce Nat. Notes. 4(5):4. 1933. The bristle-cone pine in Bryce Canyon Natl. Park. Zion and Bryce Nat. Notes 5(3):8. 1933. Douglas fir in Bryce Canyon Zion and Bryce Nat. Notes 5(4):4-5. Welsh, S. L. and G. Moore. 1973. Utah plants tracheophyta. Brigham Young University Press, Provo, Utah. 474 pp. Welsh, S. L., N. D. Atwood, and J. L. Reveal. 1975. Endangered, threatened, extinct, endemic, and rare or restricted Utah vascular plants. Great Basin Nat. 35:327-376. Literature Cited (General Biota Including Plants) Adams, E. B., and A. Chavex. 1956. The missions of New Mexico, 1776, a description, with other contemporary documents. University of New Mexico Press, Albuquerque. 387 pp. Arrington, L. J. 1958. Great Basin kingdom, an eco- nomic history of Latter-day Saints, 1830-1900. Harvard Univ. Press, Cambridge, Mass. 524 pp. AuERBACK, H. S. 1943. Father Escalante's journal with related documents and maps. Utah Hist. Quart. 11:1-10. Baker, A. A. 1936. Geology of the Monument Valley- Navajo Mountain Region, San Juan County, Utah. U.S. Geological Bull. No. 865. Washing- ton, D.C. 106 pp. Bancroft, H. H. 1889. History of Utah, 1540-1886. San Francisco History Co., 1889. Amo Press, New York. 808 pp. Benson, S. B. 1935. Biological reconnaisance of Navajo Mountain. Univ. of Calif. Publ. in Zool. 40(14):439-456. Bernheimer, C. L. 1923. Encircling Navajo Mountain with a pack train— a new route to Rainbow Natural Bridge. Natl. Geogr. Mag. 43:197-224. Bolton, H. E. 1950. Pageant in the wilderness, the story of the Escalante expedition to the Interior Basin, 1776, including the diary and itinerary of Father Escalante; translated and annotated. State Historical Society. 265 pp. Brooks, J. 1957. Lee's Ferry at Lonely Dell. Utah Hist. Quarterly 25:283-295. Brigham Young University Navajo-Kaiparowits reference collection. 1971-1974. Brigham Young University, Provo, Utah. Chamberlin, R. V. 1947. Francis Marion Bishop, 1843-1933. Utah Hist. Quarterly 25:155-158. Chidester, L, and E. Bruin. (Daughters of Utah Pio- neers). 1949. Golden nuggets of pioneer days, a history of Garfield County. Garfield County News, Panguitch, Utah. Cleland, R. G. 1963. This reckless breed of men, the trappers and fur traders of the Southwest. Knopf, New York. 361 pp. Crampton, G. C. 1959. Outline history of the Glen 178 Great Basin Naturalist Vol. 38, No. 2 Canyon region. Univ. Utah Papers No. 42, Glen Canyon Ser. No. 9:1-137. Creer, L. H. 1958a. Mormon towns in the region of the Colorado. Univ. of Utah Anthro. Papers No. 32. 26 pp. - 1958b. The activities of Jacob Hamblin in the region of the Colorado. Univ. Utah Anthr. Pa- pers No. 42, Glen Canyon Ser. No. 9:1-137. Danson, E. B. 1958. The Glen Canyon project. Plateau 30(3):75-78. Darrah, W. C. 1947. Biographical sketches and origi- nal documents of the first Powell expedition of 1869, and the second Powell expedition of 1871-72. Utah Hist. Quarterly 15:9-153. 1948. Beaman, Fennemore, Hillers, Del- lenbough, Johnson, and Hottan (and) three let- ters by Andrew Hall. Utah Hist. Quarterly 16-17: 491-408. 1951. Powell of the Colorado. Princeton Univer- sity Press, Princeton, New Jersey. 426 pp. Dellenbaugh, F. S. 1902. The romance of the Colo- rado River. G. P. Putnam's Sons, New York. 399 pp. 1926. A canyon voyage, the narrative of the second Powell expedition down the Green- Colorado River from Wyoming, and the explor- ations on land, in the years 1871 and 1872. 2d edition. Yale Univ. Press. New Haven, Con- necticut. 277 pp. DuTTON, C. E. 1880. Report of the geology of the high plateau of Utah, with atlas. U.S. Geographical and Geological Survey of the Rocky Mountain Region. Washington, D.C. Government Printing Office. 307 pp. Freeman, L. R. 1923. The Colorado River yesterday, today and tomorrow. Dodd, Mead and Co., New York. 307 pp. Freemont, J. C. 1846. Report of exploring expedition to the Rocky Mountains in the year 1842, and to Oregon and North California in the years 1843-44. Gales and Seaton Printers, Washing- ton. 696 pp. Gannett, H. 1900. A gazetteer of Utah. U.S. Geologi- cal Survey Bull. No. 166, 43 pp. Gregory, H. E. 1917. Geology of the Navajo country. U.S. Geological Survey Prof. Paper No. 93. 161 pp. 1938. The San Juan country: a geographic and geologic reconnaissance of southeastern Utah: U.S. Geological Survey Prof. Paper 188. 26 pp. 1939. Diary of Almon Harris Thompson. Utah Hist. Quarterly 7:3-138. 1945. Scientific explorations in southern Utah. Am. Journal of Sci. 243:527-549. 1948. Geology and geography of central Kane County, Utah. Bull. Geol. Soc. of America 59:211-248. . 1949. Stephen Vandiver Jones, 1840-1920, and Journal April 21, 1871-Dec. 24, 1872. Utah Hist. Quarterly 16-17:11-174. 1951. The geology and geography of the Paunsaugunt region, Utah, a survey of parts of Garfield and Kane Counties. U.S. Geol. Survey Prof. Paper No. 226. 116 pp. Gregory, H. E., and R. C. Moore. 1931. The Kai- parowits region. U.S. Geological Survey Prof. Paper No. 164. 161 pp. Gregory, H. E., W. C. Darrah, and C. Kelly, eds. 1948-49. Exploration of the Colorado River and the high plateaus of Utah in 1871-72. Utah Hist. Quarterly 16-17. 540 pp. Hall, A. F. 1934. General report on the Rainbow Bridge-Monument Valley expedition 1933. Univ. of Calif. Press, Berkeley. 32 pp. Hayward, L. C, D. E. Beck, and W. W. Tanner. 1958. Zoology of the upper Colorado River ba- sin. 1. The biotic communities. Brigham Young University Biol. Surv. 1(3): 1-74. Heald, W. F. 1955. Who discovered Rainbow Bridge? Sierra Club Bulletin No. 40:22-28. Hill, J. F. 1930. Spanish and Mexican exploration and trade northwest from New Mexico into the Great Basin, 1705-1853. Utah Hist. Quarterly 3:2-23. HiTE, C. 1939. Colorado River gold. From "Beaver Utonian," January 13, 1893. Utah Hist. Quar- terly 7:139-140. James, G. W. 1922. Utah, the land of blossoming val- leys. Page Company, Boston. 371 pp. Kelly, C. 1947. Captain Francis Marion Bishop's jour- nal, August 15, 1870-June 3, 1872. Utah Hist. Quarterly 15:253-490. 1949. Walter Clement Powell, and journal. ■ April 21, 1871-December 7, 1872. Utah Hist. Quarterly 16-17: 253-490. Kidder, A. V. 1910. Exploration in southwestern Utah in 1908. Am. J. of Archaeology 14(3):337-359. Lawson, a. C. 1913. The gold in the Shinarump at Paria. Economic Geology 8:434-438. Leavitt, F. J. 1943. Steam navigation on the Colorado River. Calif. Hist. Society Quarterly. 22(1-25), 151-174. Miller, D. E. 1958. Discovery of Glen Canyon. Utah Hist. Quarterly 26:220-231. 1959. Hole-in-the-Rock, an epic in the coloniza- tion of the Great American West. University of Utah Press, Salt Lake City, 229 pp. Miser, H. D., K. W. Trimble, and S. Paige. The Rainbow Bridge, Utah. Geographical Review 13:518-531. Parker, T. C. 1933. Game management problems of Utah national parks. Utah State Ag. College Misc. Publ. 10:32-34. PoQUE, J. E. 1911. The great Rainbow Natural Bridge of southern Utah. Natl. Geogr. Mag. 22:1048-1056. Powell, J. W. 1875. Exploration of the Colorado Riv- er of the West and its tributaries; explored in 1869 to 1872. Smithsonian Institution. Govern- ment Printing Office, Washington. 291 pp. Presnall, C. C. 1933. Along nature's highways. Zion- Bryce Nat. Notes 6(6):52-57. 1934. Animals and birds of Bryce Canyon Na- tional Park. Zion-Bryce Nat. Notes 6(6):52-57. Pritchett, C. L. 1962. Vertebrate distribution in rela- tion to certain habitats in central Kane County, Utah. Unpublished M.S. thesis, Dept. of Zoolo- June 1978 Welsh et al.; Kaiparowits Flora 179 gy, Brigham Young University, Provo, Utah. 83 pp. Russell, R. W., and H. E. Thompson. 1964. A listing of common plants and vertebrate animals of Bryce Canyon National Park. Unpublished man- uscript. ScHMECKEBiER, L. F. 1904. Catalogue and index of the publications of the Hayden, King, Powell, and Wheeler surveys. U.S. Geological Survey Bull. No. 222. 208 pp. Tanner, V. W. 1940. A biotic study of the Kaiparowits region of Utah. Great Basin Nat. 1:97-126. Tyler, S. L. 1951. Before Escalante, an early history of the Yuba Indians and the area north of New Mexico. Unpublished Ph.D. dissertation, Univ. of Utah. Salt Lake City. 231 pp. Underbill, R. M. 1956. The Navajos. University of Oklahoma Press, Norman, Oklahoma. 292 pp. Widtsoe, J. A. 1955. A journal of John A. Widtsoe Colorado River Party, September 3-19, 1922, preliminary to the Santa Fe Conference which framed the Colorado River Compact. Edited by A. R. Mortensen with a foreword by G. Homer Durham. Utah Hist. Quarterly 23:195-231. Woodbury, A. M. 1950. A history of southern Utah and its national parks. 2d edition. Salt Lake City (reprinted from Utah State Historical So- ciety Quarterly, July-Oct. 1944: Vol. 12:110-222). Woodbury, A. M., S. Flowers, D. W. Lindsay, S. D. DuRRANT, N. K. Dean, A. W. Grundmann, ]. R. Crook, W. H. Behle, H. G. Higgins, G. R. Smith, G. G. Muser, and D. B. McDonald. 1959. Ecological studies of the flora and fauna in Glen Canyon. Univ. of Utah Anthropol. Pa- per No. 40, Glen Canyon Ser. No. 7:1-225. RANGE EXTENSION OF ONYCHOMYS TORRIDUS LONGICAUDUS (RODENTIA: CRICETIDAE) IN NORTHWESTERN NEVADA Mark D. Engstrom' and Eugene D. Fleharty' Abstract.— Two Onychomys torridus longicaudus Merriam were captured at a site 22 miles W Winnemucca, Humbolt Co., Nevada. This record extends the known range of O. torridus 165 km to the northeast. The distribution of Onychomys torridus generally is restricted to the Lower Sonoran Life Zone of North America (Hall and Kel- son, 1959, The Mammals of North America, Ronald Press, New York). However, there are a few records of O. torridus from the Upper Sonoran Life Zone along the eastern edge of the Sierra Nevada Mountains in northwestern and west-central Nevada (Hall, 1946, Mammals of Nevada, Univ. California Press, Berkeley, p. 495). On 13 August 1964 a field party from the Museum of the High Plains, Fort Hays State University, under the direction of the second author, collected small mammals from a locale 22 miles W Winnemucca, Humbolt Co., in northwestern Nevada. The area was characterized by vegetation typi- cal of the sandy, flat-floored valleys of the region. Dominant plant species, liberally spaced, included Chrysothamnus sp., Atri- plex sp., and Gutierrezia sp., with local pop- ulations of Artemisia tridentata. Subsequently, in conjunction with a forth- coming study on Onychomys leucogaster, examination of the specimens of that species housed in the Museum of the High Plains (MHP) revealed two specimens from the above site which actually pertain to the tax- on O. torridus longicaudus Merriam. This record extends the known range of O. tor- ridus 165 km to the northeast of the north- ernmost locality reported by Hall (1946: 495) which was a site 3 miles NNE Toulon, 3900 ft, in southern Pershing Co., Nevada. Of the two specimens, one (MHP 3998) is an adult male (Age Class V, Van Cura and Hoffmeister, 1966, J. Mamm. 47:613-630), whereas the other (MHP 4446) is a subadult female molting into adult pelage (Age Class n. Van Cura and Hoffmeister 1966). Se- lected measurements (in millimeters unless otherwise noted) of the two (adult, sub- adult), with mean values for O. t. long- icaudus reported by Hall (1946) in parent- heses, are as follows: total length 139, 144 (138); length of tail 43, 44 (46.5); length of hind foot 20, 18 (19.3); weight (in grams) 21.8, 19.5 (22.6); greatest length of skull 25.0, 23.7 (24.9); breadth of braincase 11.4, 19.8 (11.3); length of nasals 9.5, 8.6 (9.3); shelf of bony palate 4.4, 4.2 (4.4); alveolar length of maxillary toothrow 3.7, 3.6 (3.8). The remaining collection of small mam- mals from Humbolt County contained an unusual variety of heteromyid rodents, probably due to an admixture of Lower (Dipodomys deserti and D. merriami) and Upper Sonoran faunal elements. Species and numbers collected were: D. deserti, 6; D. merriami, 15; D. microps, 1; D. ordii, 7; Mi- crodipodops megacephalus, 5; Perognathus longimembris, 1; and P. parvus, 2. Addition- ally, three specimens of Ammospermophilus leucurus were taken. The authors thank Jerry R. Choate and Robert C. Dowler (Fort Hays State Univer- sity) for critically reviewing this manuscript. 'Museum of the High Plains, Fort Hays State University, Hays, Kansas 67601. 180 THE REPRODUCTIVE ECOLOGY OF THE TAHOE SUCKER, CATOSTOMUS TAHOENSIS, IN PYRAMID LAKE, NEVADA Joseph L. Kennedy' and Paul A. Kucera' Abstract.— The Tahoe sucker spawns in Pyramid Lake from April to August at lake temperatures of 11.7 to 22.7 C. The spawning population is comprised of a large lake spawning group and a numerically smaller river running group. The river running group is smaller in length and was not considered during this study. The sex ratio of sampled suckers significantly favored the females. This is the result of the longer life of fe- males and greater mortality of males during spawning. Pyramid Lake Tahoe suckers reach sexual maturity at two to three years of age; however, those in Lake Tahoe do not mature until four or five years of age. The size at sexual maturity is different in both populations, which suggests that size or rate of growth rather than age determines sexual maturity. The fecundity of Tahoe suckers is positively correlated with fork length, weight, and age. Additional analysis showed that a better correlation occurred between fish size (either length or net weight) and total ovary weight. We believe that fish size is primarily correlated with total reproductive tissue produced and secondarily with fe- cundity. A comparison of the Pyramid Lake population and the Lake Tahoe population demonstrated that size, not age, is the most important determinant of Tahoe sucker fecundity. The Tahoe sucker, Catostomus tahoensis, is found in the Lahontan drainage system of western Nevada and eastern California. Within the Truckee River system, it com- prises a major portion of the fish population (LaRivers 1962). It is of little direct eco- nomic value, but it does provide a signifi- cant food source for trout in Pyramid Lake and Lake Tahoe (Miller 1951) and probably contributes to the food of brown trout in the Truckee River. Its current status in rap- idly desiccating Walker Lake is unknown. Although C. tahoensis is one of the most common fishes of the Lahontan drainage, very little is known about its reproductive ecology. Snyder (1917) gave a brief descrip- tion of spawning behavior, and Willsrud (1966) presented fairly extensive data on the reproductive biology of the Tahoe suckers in Lake Tahoe. The objectives of our study were to describe the reproductive ecology of the Tahoe sucker in Pyramid Lake and to compare it with other species of suckers. Special emphasis was given to comparisons between the Lake Tahoe population (Will- srud 1966) and the Pyramid Lake popu- lation. Methods Tahoe suckers were collected monthly from November 1975 through March 1977 through the use of bottom-set 1.83 X 76.2 m variable mesh gill nets. These were com- prised of ten 7.6 m panels of 12.7, 19.05, 25.4, 31.75, 38.1, 44.45, 50.4, 63.5, 76.2, and 88.9 mm bar mesh netting. Fyke nets con- structed of 12.7 mm mesh netting over 1.22 m diameter hoops were also used. The net was 4.8 m long, with a 15.24 m lead. Sex ratios were obtained from a monthly subsample of the Tahoe suckers collected. During the spawning season, secondary sex- ual characters were used after verification by internal examination. Ovaries were removed from 53 freshly killed Tahoe suckers for fecundity determi- nation. Each ovary was weighed to the nearest 0.001 gram, and ten ova diameters were measured to the nearest 0.1 mm. The fork length of each fish was measured to the nearest mm, and each fish was weighed to the nearest gram. Scales were also taken for age determination. Three replicate subsamples of approx- 'W. F. Sigler & Associates Inc., 1005 Terminal Way, Suite 155, Reno. Nevada 89502. 181 182 Great Basin Naturalist Vol. 38, No. 2 imately one gram of eggs were taken from each ovary, weighed to the nearest 0.001 gram, and preserved for later counting. Fe- cundity was then estimated by direct pro- portion per ovary. Linear and nonlinear (logio) regressions were used to examine the interrelationships between fecundity, egg size, body weight, fork length (FL), and age. The analyses that involved fish weight used net weight; that is, body weight minus ovary weight. Results and Discussion Reproductive Cycle.— Nuptial tubercles first appeared on males in December and by January were very pronounced, forming definite rows along the rays of the anal and caudal fin. The females usually had no tu- bercles, but a few had them on the anal fin. The females exhibited a characteristic dis- tended vent. Mature males were collected through the spring and summer and as late as September. During March and April 1976, an esti- mated 5500 Tahoe suckers ascended the Marble Bluff fishway or the Truckee River to spawn. The river spawning nm of Tahoe suckers occurred when the mean lake sur- face temperature was 10 C, and the water temperatiue in the river 9 to 12 C. This corresponded to earlier observations on Pyramid Lake (LaRivers 1962). LaRivers (1962) further reported that C. tahoensis as- cending the Truckee River from Pyramid Lake were all small (< 305 mm) which our data also support. Geen et al. (1966) found that spawning migrations of longnose suck- ers (Catostomus catostomus) began at 10 G and peaked at 12.3 G, and those of white suckers {Catostomus commersoni) peaked at 13,5 G. Spawning runs of C. catostomus, from western Lake Superior, peaked at 13 G over a seven-year period (Bailey 1969). Willsrud (1966) noted separate river and lake spawning populations of Tahoe suckers in Lake Tahoe, Galifomia-Nevada, which suggests they may be genetically different strains. The Utah sucker (Catostomus ar- dens) also exhibits river and lake spawning populations in Bear Lake, Utah-Idaho (McGonnell et al. 1957). Preliminary elec- trophoretic tissue analysis on C. ardens from Bear Lake indicates no significant difference between the two groups (Klar, unpublished data). The river spawning run coincided with the peak of the gonadal somatic index (14 percent) of the suckers collected in the lake. The GSI of the lake spawning fish (Fig. 1) remained high in April and May, and 95.4 percent of the fish collected in June were sexually mature. This decreased to 85.7 per- cent in July and to only 5.8 percent in Au- gust (August GSI = 1.5 percent). Gatch rates reveal that general activity patterns decrease over the spring to a low point in July and August (Fig. 1). In July spent fish were stressed and inactive; therefore, we sampled a higher percent but actually lower numbers of ripe fish, which is reflected in reduced catch rates. As Tahoe sucker activi- ty patterns are associated with spawning be- havior (Vigg, unpublished data), we believe the GSIs and percent ripe fish, in this case, represent biased results. Thus, lake spawn- ing activity commenced in April and was essentially completed by July, at water tem- peratures of 11.7-22.7 G. The protracted spawning period, from April to August, that we observed for C. ta- hoensis has also been found for C. dis- cobolus (Andreasen and Barnes 1975) and by Willsrud (1966) for Tahoe suckers in Lake Tahoe. Spawning activity in Pyramid Lake commences as both water temperature and photoperiod begin to increase and is drawn out over a period of several months. An- dreasen and Barnes (1975) speculated that both water temperature and photoperiod combine to trigger spawning activity, but Kaya and Hasler (1972) reported that fish- ery scientists found a heterogeneity of re- sponses to experimental conditions with dif- ferent species. Ghanges in water temperature and photoperiod are so closely linked in Pyramid Lake that determining the relative importance of either to initia- tion of spawning may be impossible without controlled experiments. Sex Ratio.— Of 1,557 Tahoe suckers sexed from November 1975 through Febru- ary 1977, 740 were males and 817 were fe- males. This ratio, 1:1.10, significantly differs June 1978 Kennedy, Kucera: Tahoe Sucker 183 from the expected 1:1 ratio (P<0.05). The larger number of females in the population is attributable to the longer life of females and greater mortality of males. Age and growth studies have shown that females in Pyramid Lake live to age five, but no males have been collected that were older than four (Robertson, impublished data). A sim- ilar occurrence has been noted in white suckers (Geen et al. 1966, Scott and Cross- man 1973). Willsrud (1966) noted a greater decrease in growth rate and a proportion- ately greater decrease in the number of males in older size groups in Lake Tahoe. Male suckers exhibit reproductive activity for a greater portion of the year in Pyramid Lake and Lake Tahoe and tend to remain in the spawning areas for a greater time span than the females. This may result in higher mortality for males from either stress or predation. Sexual Maturity.— The majority of the male suckers mature at two years of age, and all are mature by age three. A few of the females are mature at two, and the ma- jority mature at three (Table 1). Most indi- viduals were from 212 to 324 mm long at the onset of sexual maturity. Willsrud (1966) reported that Tahoe suckers in Lake Tahoe mature at lengths of 147 to 200 mm and at 250- 200- ^ 150 ^ 100 0) 0) ^ 50 Water temperature GSI Tahoe sucker catch M A — I — M Month A 25 20 15 10 5 0 hl5 -5 u O i_ to -10 "? o E "5 O c o O Fig. 1. Monthly changes in Tahoe sucker catches and gonadal somatic indexes in Pyramid Lake in 1976. Water temperatures are the monthly surface averages. 184 Great Basin Naturalist Vol. 38, No. 2 four or five years of age. These data show that both size and age influence sexual maturity. Since Pyramid Lake suckers reach sexual maturity at a much younger age (and larger size), size seems more important than age in determining the onset of sexual maturity. Egg Size.— The mature egg diameter for Tahoe suckers varied from 1.5 to 2 mm (x = 1.8). Willsrud (1966) reported similar egg size for the Tahoe suckers from Lake Tahoe (Table 1), and Andreasen and Barnes (1975) reported mature ovum diameters of approximately 2 mm for C ardens and C. discobolus. Lack (1948) proposed the exis- tence of an egg size /fecundity relationship that stabilized the average fecundity of the population through a differential survival rate for different size fry. This is supported by Dahl (1918-19, cited by Bagenal 1971) and Gray (1926, 1928, cited in Bagenal 1971), who showed that larger eggs produce larger fry. However, we did not find a sig- nificant relationship between ovum diame- ter and age or fecundity (F = 3.67 and 1.80, respectively). Either our sample size was in- adequate or some other mechanism exists that maintains a stable average fecundity in the Pyramid Lake population. Fecundity.— The mean fecundity of Tahoe suckers from Pyramid Lake was 20,550 eggs per female. This is less than was foujid for C. ardens and is roughly comparable to C. discobolus (Andreasen and Barnes 1975). The mean fecundity of the larger C. catostomus was 26,000 (Bailey 1969), but the fecimdity of fish of the same size was greater for Tahoe suckers. The white (C. commersoni) had more eggs per female in all size groups than Tahoe suckers (Vessel and Eddy 1941). The logjo regression for number of eggs produced by a female Tahoe sucker is significantly related to age (F = 47.87, P<0.05) fork length (F = 107.52, P<0.05), and net weight (F = 86.99, P<0.05). There is much disagreement con- cerning the relative importance of each of these variables to the fecundity of fish. The size of the fish, length and weight, deter- mines the amount of energy that can be ex- pended for the production of reproductive tissue, and space available within the body cavity. From a practical point of view, it matters little whether weight or length is used as a predictor variable of fecundity. For Tahoe suckers, both length and net weight are highly correlated with fecundity (r = .79 and .82, respectively, P<0.05). In both cases the relationship had a significant linear fit but was best described by a logjo equation. A reexamination of Willsrud's data (1966) showed a similar high correla- tion between fecundity and fork length (r = .92, P<0.05). He did not present his data in a manner that will allow an analysis of the weight-fecundity relationship. An analysis of the relationship between fork length and weight and ovary weight of Table 1. Comparison of Lake Tahoe suckers from Lake Tahoe, Cahfomia-Nevada (eggs numbers estimated from log equation) and Pyramid Lake, Nevada. X X X Ovum Fork Length Total Weight Diameter X Location Age (mm) (grams) (mm) Fecundity Lake Tahoe IV 147 45 2,510 (from Willsrud 1966) V 170 72 3,502 VI 201 118 5,164 VII 229 172 1.90 7,204 VIII 249 222 8,995 IX 277 304 11,230 X 310 426 14,822 >x 363 680 21,857 Pyramid Lake II 291 304 1.88 13,213 III 330 453 1.87 19,000 IV 360 590 1.80 26,229 V 433 998 1.77 59,268 Jtine 1978 Kennedy, Kucera: Tahoe Sucker 185 Pyramid Lake Tahoe suckers showed that length and weight are highly correlated with ovary weight. In fact, the r values were higher than those for fecundity (r = .85 and .79, respectively). This further supports our belief that fish size is directly related to the total amount of reproductive tissue that can be produced under a given set of envi- ronmental conditions. Fecundity in tiu-n is related to the total ovary weight and ovum size as was shown by Kucera and Kennedy (1977). The interrelationships of fish size (length and weight) and age and fecundity for 62- 57- 52 - 47 42- 37- 32- 27 22 17 12 7 ■ 2- c s? o c 3 0 Tahoe suckers are more difficult to analyze. The Lake Tahoe population has radically different growth rates and age structure. For example, the oldest sucker we have col- lected was 5 years old, and Willsrud col- lected several that were 15 years of age. The length-weight relationship for both populations is similar. A one-way analysis of covariance demon- strated that the slopes of the fork length- fecundity relationships are significantly dif- ferent (F = 9.04, P<0.05). We feel this is due to differences in the growth rate. Ex- amination of these relationships (Fig. 2) O O o: o «^°o°o o o — I 1 1 1 1 1 1 I I 1 I I 1 ■ 140 165 190 215 240 265 290 315 340 365 390 415 440 0 Fork length (mm) Fig. 2. Comparison of Tahoe sucker fecundities from Pyramid Lake (A) and Lake Tahoe (0). 186 Great Basin Naturalist Vol. 38, No. 2 shows there is significant overlap. Further- more, comparison of mean fecimdities over similar ranges in fork length (265 to 440 mm) by a one-way analysis of variance in- dicates no significant differences in the means (F=1.63, P>0.05). Comparison of the mean fecundity for a given weight fish from either population shows close agree- ment (Table 1). This leads us to believe that Tahoe sucker fecundity is related to size and that age is a secondary influence. Acknowledgments We wish to thank the entire crew of W. F. Sigler & Associates Inc., especially Mr. Steven Vigg (distribution and abundance data) and Mrs. Denise Robertson (age and growth data). Literature Cited Andreasen, J. K., AND J. R. Barnes. 1975. Reproduc- tive life history of Catostomus ardens and C. discobolus in the Weber River, Utah. Copeia 1975(4):643-648. Bagenal, T. B. 1971. A short review of fish fecundity, p.89-111. In: S. B. Gerking (ed.). The biological basis of freshwater fish production. Blackwell Sci. Publ., Oxford and Edinburgh. Bailey, M. M. 1969. Age, growth, ayd maturity of the longnose sucker Catostomus catostomus of West- em Lake Superior. J. Fish. Res. Bd. Canada 26(5): 1289-1299. Dahl, K. 1918-19. Studies of trout and trout waters in Norway. Salm. Trout Mag. 17:58-79; 18: 16-33. (Cited in Bagenal 1971). Geen, G. H., T. G. Northcote, G. F. Hartman, and C. C. Lindsey. 1966. Life histories of two spe- cies of catostomid fishes in Sixteenmile Lake, British Columbia, with particular reference to inlet stream spawning. J. Fish Res. Bd. Canada 23(11): 1761-1788. Gray, J. 1926. The growth of fish: I. The relationship between embryo and yolk in Salmo fario. Br. J. Exp. Biol. 4:215-225. (Cited in Bagenal 1971). 1928. The growth of fish: II. The growth rate of the embryo of Salmo fario. Br. J. Exp. Biol. 6:110-124. (Cited in Bagenal 1971). Kaya, C. M., and a. D. Hasler. 1972. Photoperiod and temperature effects on the gonads of green svuifish, Lepomis cyanellus (Rafinesque), during the quiescent winter phase of its annual sexual cycle. Trans. Amer. Fish. Soc. 101(2):270-275. KucERA, p. A., AND J. L. KENNEDY. 1977. Evaluation of a sphere volume method for estimating fish fe- cundity. Prog. Fish-Cult. 39(3): 115-1 17. Lack, D. 1948. The significance of litter size. J. Anim. Ecol. 17:45-50. LaRivers, I. 1962. Fishes and fisheries of Nevada. Ne- vada State Fish and Game Comm. 782 p. Miller, R. G. 1951. The natural history of Lake Tahoe fishes. Ph.D. dissertation. Stanford Univ. 160 p. McCoNNELL, W. J., W. J. Clark, and W. F. Sigler. 1957. Bear Lake: its fish and fishing. Utah State Dept. of Fish and Game, Idaho Dept. of Fish and Game, Wildlife Manage. Dept. of Utah State Agri. Coll. 76 p. Scott, W. B., and E. J. Grossman. 1973. Freshwater fishes of Canada. Fish. Res. Bd. of Canada Bull. 184:538-543. Snyder, J. O. 1917. The fishes of the Lahontan system of Nevada and northeastern California. U.S. Bur. Fish. Bull. 1915-16(35):31-86. Vessel, M. F., and S. Eddy. 1941. A preliminary study of the egg production of certain Minnesota fish- es. Fish. Res. Rep. 26. Minnesota Dept. of Cons. Div. of Game and Fish. 26 p. Willsrud, T. 1966. A study of the Tahoe sucker Cas- tostomus tahoensis Gill and Jordan. M.A. thesis. San Jose State Coll. 96 p. PRIMARY PRODUCTIVITY IN MEROMICTIC BIG SODA LAKE, NEVADA R. P. Axler', R. M. Gersberg', and L. J. Paulson^ Abstract.— In situ radiocarbon uptake measurements conducted at Big Soda Lake, Nevada, indicate that (i) bacterial photosynthesis comprises an important fraction (30 percent) of the lake's total primary production and (ii) bacterial chemosynthesis contributes significantly to organic particle production. The results of nutrient enrichment bioassay experiments support Hutchinson's prediction that availability of inorganic nitrogen, rather than phosphorus, limits primary production in the mixolimnion. Nutrient additions of NO3-N with Fe + '' most stimulated l'*C uptake. Big Soda Lake was described hydro- logically as early as 1885 (Russell 1885) and has received considerable limnological at- tention by virtue of its meromixis (Hutchin- son 1937, 1957, Kimmel et al., in press). However, little biological data has been ac- cumulated on the lake. Hutchinson, in 1933, found a thermally stratified mixolimnion with an oxygen depleted hypolimnion, over- lying an anoxic monimolimnion. He mea- sured concentrations of a number of nutri- ents and deduced that phosphorus was much less likely to limit primary production in the mixolimnion than inorganic combined nitrogen (Hutchinson 1937). Koenig et al. (1971) reported on the limnological status of the lake and noted a stratum of pink- colored water located in the deep hypolim- nion. Photosynthetic purple sulfur bacteria were identified in this water (Rhodothece sp. and Thiothece sp.). However, primary productivity measurements were not made. In a meromictic lake with high concen- trations of sulfide in the chemocline region and an anoxic (but photic) hypolimnion, bacterial photosynthesis may comprise a sig- nificant fraction of a lake's primary produc- tion (Wetzel 1975, Cohen et al. 1977a, b). We have utilized inorganic 14 C to esti- mate the magnitude of algal and bacterial photosynthesis at Big Soda Lake and have conducted nutrient enrichment experiments to investigate the possible nitrogen limita- tion suggested by Hutchinson. Methods Temperature, pH, and dissolved oxygen were measured in situ with a calibrated Hydrolab water quality analyzer. Light pen- etration was measured with a Whitney LMD photometer. Total alkalinity was de- termined by potentiometric titration with O.IN HCl. Chloride concentrations were de- termined by the argentometric method (Am. Public Health Assoc. 1971). Total phos- phorus, reactive iron, and nitrate were de- termined by the methods of Strickland and Parsons (1968) as modified by Fujita (see Goldman 1974). Ammonia was determined as per Solorzano (1969) as modified by Lid- dicoat et al. (1975). In situ incubations of water samples with ^d for determination of primary productivity and nutrient stimu- lation followed the procedures developed and modified by Goldman (1963). Filtered samples were acid fumed (Wetzel 1965) to remove precipitated and adsorbed i-C. Solar insolation was recorded with a Belfort pyr- heliometer. Dissolved inorganic carbon (DIG) available for photosynthesis was ap- proximated from temperature, pH, and al- kalinity data using the data of Saunders et al. (1962). This method probably over- 'Division of Environmental Studies, University of California, Davis, California 95616. 'Department of Biological Sciences, University of Nevada, Las Vegas, Nevada 89154. 187 188 Great Basin Naturalist Vol. 38, No. 2 estimates available DIG in alkaline Big Soda Lake samples (see Hutchinson 1957, Tailing 1973), and thereby results in some overestimation of primary productivity. Results and Discussion On 23 April 1977 the thermocline and chemocline were located at about 5 m and 37.5 m, respectively (Fig. 1). The vertical extinction coefficient was 0.35 m-i and mea- surable light penetrated to 26 m. The vertical distribution of primary pro- ductivity (Fig. 2) was bimodal with peaks at 5 and 25 m. The zone below 17.5 m was oxygen deficient (<1 mg Og*^') and presum- ably, could only support photosynthetic growth by anaerobic bacteria.^ The areal productivity for this zone was calculated to be 317 mg G*m-2'day-i. Similar calculations for the upper, aerobic zone yielded a value of 717 mg G*m-2«day-i (phytoplankton pho- tosynthesis). Therefore, the bacterial contri- bution to the total photosynthetic produc- tivity in the water column was 31 percent. At all depths inorganic carbon uptake in dark bottles represents a large percentage of that in light bottles, thus indicating sig- 65 12 16 20 24 28 32 Q CI"! 10 12 14 16 18 20 22 -I 10 c PH PH Fig. 1. Depth profiles of temperature, pH, and chloride for Big Soda Lake, 23 April 1977. 'Recent evidence indicates that some species of Oscillatoria are able to photosynthesize anaerobically by using H2S as a source of electrons for pho- tosystem II (Cohen et al. 1975). June 1978 AXLER ET AL.: BiG SODA LaKE 189 nificant chemosynthetic activity. In the mix- olimnion this is probably due to both ni- trifying bacteria (which utiUze electrons from NH4+ to reduce CO 2) and sulfur oxi- dizing bacteria (which utilize electrons from H2S to reduce CO2). Concentrations of sul- fide (from Koenig et al. 1971) and ammonia (Fig. 3) in the region of the chemocline are 400 mg S*/-i and 4-34 mg N'/-i, respective- ly, and therefore, the gradual entrainment of monimolimnetic water (see Kimmel et al., in press) must introduce appreciable quantities of these nutrients into the mix- olimnion during the fall-winter overturn. Conditions in the deep hypolimnion are also suitable for chemosynthetic denitrifying sul- fur bacteria, such as Thiobacillus denitrifi- cans. The concentration of nitrate in the ox- ygen depleted hypolimnion does decline from 55 to 9 jUgN-/-i (see Fig. 3). In the strictly anoxic monimolimnion, high dark uptake rates may be due to the presence of anaerobic sulfate reducing bacteria (e.g., Delsulfovibrio desulfuricans) which are fac- ultatively autotrophic and use molecular hy- drogen as an electron donor (Doelle 1975). Our data for total phosphorus, ammonia, and iron (Fig. 3) are similar to those report- ed by Hutchinson (1937). A series of nutri- ent enrichment experiments on epilimnetic water collected on 24 April 1976 provided evidence to support Hutchinson's sugges- tions concerning phosphorus and nitrogen limitation of primary productivity. Max- imum stimulation of i^C uptake occurred in water samples enriched with both nitrate and iron (Fig. 4). Phosphate additions, ei- ther singly or together with nitrate or iron. a> Q mg C m'^hr' 65 4 8 10 12 H — J* — h- 0 10 20 30 40 50 60 70 80 H h + -i h Light Bottle Uptoke. Dork Bottle Uptake 0 2 4 6 8 10 12 mg O2 • I Fig 2. Depth profiles of dissolved oxygen and primary productivity, 23 April 1977 (1000-1400 hrs). Productiv- ity data represents the difference in uptake between light and dark sample bottles where each has been cor- rected for dark-uptake during the time between removal from the lake and filtration. 190 Great Basin Naturalist Vol. 38, No. 2 produced a small effect. Similarly, trace metal additions provided little stimulation imless added in concert with nitrate and iron.^ Primary production in many types of lakes has been shown to be limited by in- organic nitrogen concentration and, in a number of hardwater calcareous lakes, by available iron (Wetzel 1975). Goldman (1972) reported that on water samples from Lake Tahoe, Galifornia, nitrate and iron acting in concert produced a greater stimu- lation of primary productivity than either one added singly. Such an effect may be due to the role of iron in the assimilatory reduction of nitrate by microorganisms. Af- ter reduction to nitrite via the enzyme nit- rate reductase, further reduction to ammo- nium is catalyzed by nitrite reductase. Iron appears to be an integral component of both nitrite reductase and of ferredoxin, the immediate electron donor (Morris 1975, Healey 1973). The data we have accumulated on Big Soda Lake are of a preliminary nature and certainly insufficient to fully delineate the microbial processes occurring in the water column. However, they do indicate a very 0 mg P or /xg Pel mg NH3-N I 0 10 20 30 40 50 60 0 4 8 12 16 20 24 28 32 36 40 ' I, I I I I i I I I I I I I I I .io- 5- 10- 15- 20- ^25- £ -- 30H Q. 35H ^ 40H 45- 50- 55- 60- 65 !.._. OrvT" NO3-N n 1 1 1 1 1 1 1 1 T" 8 16 24 32 40 48 56 64 72 80 fiq NO3-NI Fig. 3. Depth profiles of NO3-N and NH3-N (24 April 1977); total phosphoms, reactive iron, and NH3-N (24 April 1976). 'Unfortunately, the variance in our primary productivity data from April 1976 was generally too high to include in this report (presumably this was due to precipitation of carbonates during the sample filtrations with subsequent clogging of the filtration apparatus). However, the limnological condi- tion of the lake, as determined by vertical profiles of temperature, dissolved oxygen, light attenuation, alkalinity, pH, and chloride and ammonium con- centrations was virtually identical in 1976 to 1977. Furthermore, the primary productivity at 0 m and 5 m (23 April 1976) was 4.5 mg C-m 9.3 mg C'm~'*'hr" , respectively, which were similar to the rates measured at these depths in 1977. -3.hr- 1 and June 1978 AXLER ET AL.: BiG SODA LaKE 191 interesting system in which bacterial photo- synthesis is an important fraction of the to- tal primary productivity, and bacterial chemosynthesis contributes significantly to organic particle production. There was no obvious pink color in deep hypolimnetic water when our measurements were made and, therefore, one might expect the rate of bacterial photosynthesis to be much higher later in the growing season (as when Koenig et al. collected samples in October 1970). However, the fraction of total primary pro- ductivity contributed by the sulfur bacteria seasonally remains to be investigated. We are especially indebted to Dr. Bruce L. Kimmel from the University of Okla- homa Biological Station, who helped with the field and laboratory work and data analysis and interpretation, and who criti- cally reviewed the manuscript. We ac- knowledge the field and laboratory assis- tance of the University of California (Davis) 1976 and 1977 limnology laboratory classes and the financial assistance of the Division of Environmental Studies. Some of the equipment used in this investigation was purchased under the auspices of National Science Foundation grants NSF BM 74- 02246 AOl and NSF DEB 76-19524 to C. R. Goldman. J. Coil contributed significant- ly with the laboratory analyses. M. Smith and G. Malyj aided in manuscript prepara- tion. Literature Cited American Public Health Association. 1971. Stan- dard methods for the examination of water and wastewater. New York. 874 pp. Cohen, Y., B. B. Jorgensen, E. Padan, and M. Shilo. 1975. Sulfide dependent anoxygenic photo- synthesis in the cyanobacterium Oscillatoria lim- netic. Nature 257:489-492. Cohen, Y., W. E. Krumbein, and M. Shilo. 1977a. So- lar Lake (Sinai). 2. Distribution of photo- synthetic microorganisms and primary produc- tion. Limnol. Oceanogr. 22:609-620. 1977b. Solar Lake (Sinai). 3. Bacterial distribu- tion and production. Limnol. Oceanogr. 22:621-634. 5000 r _ 4000 E Q. ^ 3000 o 2000 1000 0 N + Fe + P / ^N+Fe + P + Trace / //N + Fe / '^' // / ,^ ^..;C''..>»Fe + Trace '^'r//:?'^"^'-^::\ Fe + p ^^^ - CON TROL J L 2 4 Days of Incubation Fig. 4. Results of nutrient enrichment bioassays on water collected 24 April 1976. Enrichment concentrations ;re: N = 50 /ig NOg-N-l-i; F = 50 fig P04-3-P-1-1; Fe = 25 /xg Fe + 3l-l; Trace = 3 jug'l"^ each of Co, Mo, were: Zn, Mn. 192 Great Basin Naturalist Vol. 38, No. 2 DoELLE, H. W. 1975. Bacterial metabolism. New York: Academic Press. 738 pp. Goldman, C. R. 1963. The measurement of primary productivity and limiting factors in freshwater with carbon-14, p. 103-113. In: M. S. Doty (ed.), Proc. Conference on Primary Productivity Measu Ement, Marine and Freshwater. U.S.A.E.C, Div. Tech. Inform. Rep. TID-7633. 1972. The role of minor nutrients in limiting the productivity of aquatic systems, pp. 21-38. In: G. E. Likens (ed.). Nutrients and eu- trophication. Amer. Soc. Limnol. Oceanogr. Spec. Symp. No. 1. 1974. Eutrophication of Lake Tahoe empha- sizing water quality. EPA-660/3-74-03. U.S. Gov. Printing Office, Washington, D.C. 408 pp. Healey, F. p. 1973. Inorganic nutrient uptake and deficiency in algae. C.R.C. Grit. Rev. in Mi- crob. 3:69-113. Hutchinson, G. E. 1937. A contribution to the limno- logy of arid regions. Trans. Conn. Acad. Sci. 33:1-132. 1957. A treatise on limnology. I. Geography, physics, and chemistry. New York: John Wiley. 1015 pp. KiMMEL, B. L., R. M. Gersberg, L. J. Paulson, R. P. AxLER, AND C. R. Goldman. In press. Recent changes in the meromictic status of Big Soda Lake, Nevada. Limnol. Oceanogr. KoENic, E. R., J. R. Baker, L. J. Paulson, and R. W. Tew. 1971. Limnological status of Big Soda Lake, Nevada, October 1970. Great Basin Nat. 31:107-108. LiDDICOAT, M. I., S. TiBBITS, AND E. I. BuTLER. 1975. The determination of ammonia in seawater. Limnol. Oceanogr. 20:131-132. Morris, I. 1975. Nitrogen assimilation and protein syn- thesis. In: W. D. P. Stewart (ed.), Algal phys- iology and biochemistry. Univ. of Calif. Press., Berkeley. Russell, I. C. 1885. Geological history of Lake Lahon- tan. U.S. Geol. Surv. Monogr. 11, 287 pp. Saunders, G. W., F. B. Trama, and R. W. Bachmann. 1962. Evaluation of a modified C-14 technique for shipboard estimation of photosynthesis in large lakes. Great Lakes Div., Univ. of Mich., Ann Arbor, Publ. 8, 1-61. SoLORZANO, L. 1969. Determination of ammonia in natural waters by the phenolhypochlorite meth- od. Limnol. Oceanogr. 14:799-801. Strickland, J. D. H. and T. R. Parsons. 1968. A practical handbook of seawater analysis. Bull. Fish. Res. Board Can. No. 167. 311 pp. Talling, J. F. 1973. The application of some electro- chemical methods to the measurement of pho- tosynthesis and respiration in freshwaters. Freshwater Biol. 3:335-362. Wetzel, R. G. 1965. Necessity for decontamination of filters in C-14 measured rates of photosynthesis in freshwaters. Ecology 46:540-542. 1975. Limnology. W. B. Saunders, Phila. 743 pp. CURRENT STATUS OF CUTTHROAT TROUT SUBSPECIES IN THE WESTERN BONNEVILLE BASIN Terry J. Hickman' and Donald A. Duff^ Abstract.— Recent discoveries of native cutthroat trout populations in desert mountain ranges on the western fringe of the Bonneville Basin have prompted intensified management efforts by state and federal agencies. Anal- ysis of Snake Valley cutthroat specimens in Trout Creek, Deep Creek Mountain Range, Utah, indicate this is a pure strain of the trout which once inhabited Pleistocene Lake Bonneville and which was thought to be extinct in Utah. The Snake Valley cutthroat is similar to Salmo clarki Utah of the eastern Bonneville Basin; however, electrophoretic and morphomeristic analysis show unique genetic differences brought about by long-term isolation (8,000 years) from the remainder of the Bonneville Basin cutthroat. This cutthroat is a common ancestor to sever- al other limited cutthroat populations within the basin in Nevada. In May 1977 the BLM withdrew from mineral entry about 27,000 acres within the Deep Creek Mountains for protection of this salmonid cutthroat and other unique resources on the range. Results of 1977 stream surveys on the Pilot Peak Mountain Range, Utah, indicate the presence of the threatened Lahontan cutthroat, Salmo clarki henshawi, in one isolated stream. The ancient Pleistocene Lake Bonneville in the Great Basin once supported a cut- throat trout, native to the Snake Valley area of Utah-Nevada, which abounded in die area's several streams upon the lake's decline (Hickman 1977). Because of deterio- rating habitat the cutthroat population rap- idly diminished in the twentieth century to a point where it was believed to be extinct within its native range (Behnke 1976a) (Fig. In 1953 Ted Frantz, Nevada Fish and Game Department, discovered a cutthroat trout population in Pine Creek on Mt. Wheeler, Nevada (Frantz and King 1958). Samples were sent to Dr. Robert Miller, who indicated they represented pure cut- throat trout. But Dr. Miller was unable to assign them to any described subspecies (let- ter from Dr. Miller to F. Dodge, 26 May 1971). Though it was assumed this cutthroat was introduced from Trout Creek drainage of the Snake Valley area (Miller and Alcorn 1946), this seems imlikely when one consid- ers that there were streams closer to Pine Creek which probably contained cutthroat trout (Lehman, Baker, Snake, and Hendrys creeks). Behnke (1976a) indicates the most logical origin of the Pine Creek cutthroat was from Lehman Creek (Mt. Wheeler tributary of the Snake Valley region) via the Osceola Ditch, constructed as a pioneer waterway. During 1953 the Nevada Fish and Game Department introduced 44 fish from Pine Creek into Hampton Creek, Nevada. A sec- ond transplant of 54 cutthroat from Pine Creek was made into Goshute Creek, Ne- vada, in 1960. The Nevada Fish and Game Department, assuming these were Utah cut- throat, Salmo clarki Utah, closed these streams to fishing and listed S.c. Utah as an endangered species in Nevada. Mr. Frank Dodge, Nevada Fish and Game Depart- ment, in 1972 found a population of cut- throat trout in the headwaters of Hendrys Creek (Mt. Moriah tributary of the Snake Valley region) which resembled those found in Pine Creek. Following this, several un- successful attempts were made by the Ne- vada Fish and Game Department to locate additional pure populations of cutthroat trout in the Snake Valley area of Utah and Nevada. In 1973 the BLM (Utah) began stream habitat surveys in the Deep Creek Moun- ' Department of Fishery & Wildlife Biology, Colorado State University, Fort Collins, Colorado. fisheries Biologist, U.S. Bureau of Land Management, Utah State Office, University Club Building, 136 South Temple, Salt Lake City, Utah 84111. 193 194 Great Basin Naturalist Vol. 38, No. 2 IDAHO '-';,^^'>'>K ;_; Pilot Peak— H^ Deep Creek Mountains UTAH Fig 1. Area map location showing the western Bonneville Basin area. June 1978 Hickman, Duff: Cutthroat Trout 195 tain Range in an attempt to define critical habitats and possible remnant populations of the cutthroat. In the spring of 1974, BLM biologists Don Duff and Josh Warburton discovered cutthroat in the extreme head- waters of Trout Creek, Utah, above a natu- ral barrier falls. Subsequent sampling and analysis by the BLM, Utah Division of Wildlife Resources and Colorado State Uni- versity (under contract funded by BLM) de- termined that Trout Creek specimens were pure strain fish of the Bonneville Basin. In- ventories have continued to date, and the only stream found to contain a pure popu- lation was Trout Creek. Hybridized popu- lations (with rainbow trout) were found in Birch Creek and Johnson Creek (Hickman 1977) (Fig. 2). Reasons for Decline When the Snake Valley arm of Lake Bonneville dried up, there were relatively few perennial streams in the area. In addi- tion to this, since the mid 1800s, in- troductions of nonnative trouts, climatic conditions, irrigation practices, and habitat loss and degradation have been influential in reducing the number of cutthroat popu- lations in the Snake Valley area. Replace- ment and hybridization from introductions of exotic rainbow trout (Salmo gairdneri) have posed the most significant impact to the survival of the Snake Valley cutthroat. Virtually every stream in the Snake Valley region capable of supporting trout has been stocked with rainbows. Brook trout are also capable of replacing the cutthroat through competition because of earlier spawning pe- riods and its ability to become better adapt- ed to life in small spring-fed headwater streams. Exploitation, though not likely a limiting factor by itself, can reduce the number of catchables and may act to favor other exot- ics such as the brooks, browns, and hybrids. It has been documented that cutthroat trout are highly vulnerable to angling mortality (Behnke and Zam 1976). Livestock grazing imposes a subtle but serious threat to the survival of the cut- throat trout in the arid Snake Valley region. Grazing becomes significant when discussing sites for reintroductions, because much of the prime grasslands exist in headwater meadow areas. Livestock interests in the Bonneville Basin have been unconcerned about stream protection of rare trout popu- lations. These problems have made the BLM very cautious in planning for addition- al habitat sites for future reintroductions of the Snake Valley cutthroat. Many studies have shown that livestock grazing destroys and degrades riparian vegetation and streambank soil stability, resulting in altera- tions of channel morphology, loss of cover, and a reduction in numbers and biomass of fish— particularly older and larger trout (Behnke 1977). Studies and management of livestock-impacted areas should be made in order to rehabilitate the grazed areas, either through improvement of the existing graz- ing system or livestock exclusion (Platts 1977). The BLM in Utah and Nevada has been involved in streamside fencing pro- grams to protect the riparian habitat of streams containing sensitive or rare trout populations from continued livestock dam- age (Goshute Creek, Nevada, and Birch Creek, near Beaver, Utah). Droughts and violent thunderstorms may have historically eliminated cutthroat popu- lations from some high gradient streams, be- cause natural recolonization could not be effective after desiccation of the pluvial lake in Snake Valley. This may account for the high number of barren streams found in the Snake Valley region prior to rainbow trout introductions. Past surface disturbance impacts from mining have been slight and of short dura- tion, the main damage resulting from equip- ment movement and road construction to and from the mine site. There exists little room for trails or roads in some of the nar- row canyons; therefore, the streambed may be utilized for such purposes in some areas. Recent uranium mining activities in Utah's Deep Creek Mountains have caused concern over the future impacts of mining to the re- sources of this fragile desert island ecosys- tem environment. The effects of all these environmental im- pacts on the cutthroat trout populations are 196 Great Basin Naturalist Vol. 38, No. 2 NEVADA UTAH!:!:;:::;:: GOSHUTE ELY WHEELER PEAK Fig 2. Local area map showing extent of Lake Bonneville (shaded) in relation to perennial streams having cut- throat trout. June 1978 Hickman, Duff: Cutthroat Trout 197 greatly magnified when considered collec- tively. Many of the streams in the Snake Valley region have been affected by all these major impacts at some time during the recent past history of the area. Uniqueness of Snake Valley Cutthroat Trout Ancient Lake Bonneville went through several periods of fluctuations in which wa- ter levels were closely associated with cli- matic conditions (Gilbert 1879). According to Broecker and Kaufman (1965), four low levels occurred between 8,000 and 22,000 years ago, including one period of complete desiccation followed by refilling that took place about 11,000 years ago. This final desiccation of Lake Bonneville resulted in 10 or 12 independent basins being formed, one of which was the Snake Valley basin (Gilbert 1890). The northern portions of Snake Valley show a lake level elevation of about 5,100 feet. This would have pre- vented water from flowing out of Snake Valley and into the Great Salt Lake basin. In addition to such physical isolation, the cutthroat were forced to seek refuge in the streams to overcome the increased saline conditions brought on by the desiccation (Hunt et al. 1953). Thus, many populations of cutthroat in the Bonneville Basin have been isolated from contact with each other for about 8,000 years. Wydoski et al. (1976) studied the electro- phoretic patterns of proteins in cutthroat lo- cated in the Bonneville Basin, as well as several other groups of cutthroat and rain- bow trout. No protein was unique or dis- tinctive for S.c. Utah specimens, but an un- usual variation for muscle lactate dehydrogenase (LDH) was found in cut- throat from Trout and Goshute creeks, in- dicating a common ancestor. This unusually complex variation seems to indicate the presence of a variant allele. A unique evolu- tionary event, or series of events, occurred in the Snake Valley cutthroat trout LDH, which would indicate long-term isolation from the rest of the Bonneville Basin cut- throat trout. Comparison of samples of the least chub, lotichthys phlegethontis, in the western Bon- neville Basin adds credence to the assump- tion of incipient speciation in fishes isolated in Snake Valley. Samples from Donner Springs (Pilot Peak area) have the typical fin ray counts given by Sigler and Miller (1963). Those found in Snake Valley have one less ray in the dorsal (7), anal (6), and pelvic (7) fins. Smith (1966) stated that the mountain suckers (Catostomus platyrhynchus), of Deep Creek in the Deep Creek Mountain area are different from the typical northern Bormeville form. The Snake Valley cutthroat trout differs from other cutthroat trout of the Bonneville Basin by having more basibranchial teeth and gillrakers and fewer scales in the lateral line series. The spotting pattern is more uniformly distributed over the body and not so concentrated posteriorly as in other Bon- neville Basin cutthroat. The head appears longer and deeper with the body being more compressed and the caudal peduncle deeper, all of which gives it a more chunky body appearance (Behnke 1976 a, b). Status of the Snake Valley Cutthroat Trout Pure populations are found in Pine, Go- shute, Hampton, and Hendrys creeks, Ne- vada, and in Trout Creek, Utah (Fig. 2). Hybridized populations are found in Muncy and Mill creeks, Nevada, and Birch and Johnson creeks, Utah (Behnke 1976a, Hick- man 1977). Goshute Creek probably has the highest nimiber of Snake Valley cutthroat, having about 1,500 in four miles of stream (McLel- land 1975). The Nevada BLM and Nevada Fish and Game Department (NFG) have been instrumental in protecting and enhanc- ing the habitat in Goshute Creek. During the 1977 drought Goshute Creek lost about 38 percent of the cutthroat population per mile. Because of these conditions a con- cerned NFG took 71 cutthroat from Go- shute Creek and transplanted them propor- tionately into Water Canyon Creek (four stream miles habitat) and Clear Creek (one stream mile habitat). Pine Creek, a very small stream with little habitat, has about 100 cutthroats (ex- 198 Great Basin Naturalist Vol. 38, No. 2 eluding fry), as does Hampton Creek, which is also a small stream (McLelland 1975). Pine Creek suffered some mortality as a re- sult of the 1977 drought. Mile Creek, an- other creek with transplanted cutthroat, lost its entire population as the creek dried up from the drought. Hendrys Creek had about 200 cutthroat in the headwater area in 1973. In 1974 eradication of rainbow trout below the bar- rier was conducted on Hendrys Creek to aid the fish's survival. Hendrys, Goshute, and Pine creeks are now closed to angling use. Goshute and Hampton creeks have past histories of losing all their fish from flash floods, and this is the reason they were bar- ren in 1953 and 1960. Because of its small size. Pine Creek is also vulnerable to flash flooding. Therefore, the potential exists that the cutthroat populations in these streams could be lost in the future. During the 1977 drought NFG estimates that 50 percent of the cutthroat populations in Hendrys and Hampton creeks were lost because of dry stream sections. In the interest of managing these unique fish, NFG has identified about 25 streams suitable for reintroductions. They plan to rehabilitate about two to four streams per year in this effort. During 1977 one of the most significant events to take place in the basin for the protection of desert fishes and the environ- ment occurred in the Deep Creek Moun- tains, when the BLM filed for an emergen- cy withdrawal of a 27,000-acre area of critical environmental concern within the moimtain range. Increased uranium mining activity threatened to destroy many of the unique resources of the mountain area. A significant factor in justifying this action was the presence of the rare Snake Valley cutthroat in only about 1V4 miles of critical habitat on Trout Creek, as well as the pres- ence of the rare giant stonefly (Pteronarcys princeps). The area was withdrawn from mineral entry on 3 May 1977 by the Secre- tary of the Interior imder section 204(e) of the Federal Land Policy and Management Act of 1976 (PL 94-579). This withdrawal stays in effect for a three-year period and allows time for study of all resources to as- certain their values. In September 1977, the BLM (Utah) funded a contract to the Utah Division of Wildlife Resources to provide for an inven- tory of all fish and wildlife resources on the mountain range. The contract will last until April 1979 and will provide BLM with in- ventory data necessary to evaluate the fu- ture withdrawal status. It is hoped the con- tract will define possible other streams inhabited by the cutthroat on the mountain. In late October 1977, the Utah Division of Wildlife Resources (DWR) eradicated the rainbow trout below the natural falls barrier on Trout Creek as a start to implement management plans designed to expand the cutthroat population. Future plans call for the transportation of cutthroat from Trout Creek into the headwaters of Red Cedar Creek, a remote stream on the mountain which was given first priority for transplant efforts. The DWR plans to rehabilitate about seven additional east slope streams to enhance cutthroat survival back into their historic range. A habitat management plan (HMP) being developed for the entire mountain ecosystem by the BLM, in coop- eration with the Utah Division of Wildlife Resources, will specify management of all east slope streams for the cutthroat. The complete HMP is scheduled for completion in 1978-79 for all the mountain resources, of which the cutthroat is an integral part. At present the BLM has developed an HMP for Trout Creek, having begun implementa- tion of this plan in 1977 via Sikes Act (P.L. 93-452) authorities. Using Youth Con- servation Corps (YCC) workers, some 75 long-type stream improvement structures were constructed in July in Trout Creek to aid the bank stabilization and pool quality enhancement for the cutthroat. Stream im- provement work is scheduled again in 1978 by BLM using the YCC. Although there are differences in the tax- onomic characters between S. c. Utah and the cutthroat found in Snake Valley, there also exists much overlap. Basibranchial teeth counts, which seem to be a distinctive char- acteristic separating the two forms, were found to be similar in number in one S. c. Utah sample from Willow Creek, Jordan River drainage, Utah (Hickman 1977). With June 1978 Hickman, Duff: Cutthroat Trout 199 the analysis of more samples from the Bon- neville Basin, the degree of overlap be- tween these cutthroat becomes more ob- vious. This overlap is further substantiated through the use of a computer-aided dis- criminant function analysis, which evaluates the similarities and differences between samples (Hickman 1977). Sixteen (16) mor- phomeristic character measurements (Table 1) from samples of various described and undescribed subspecies of cutthroat trout, and one sample of rainbow trout, were compared (Fig. 2). The closer the group centroid (represented by dot in Fig. 3), the more similar the samples. The cutthroat trout in Snake Valley and S. c. Utah are closely situated, indicating a high degree of similarity. Of interest is the similarity de- picted in the discriminate function plot be- tween S. c. pleuriticus (Colorado River Cut- throat) and S. c. stomias (Greenback cutthroat). This supports the taxonomic evaluations of Behnke and Zam (1976) that S. c. pleuriticus gave rise to S. c. stomias via an ancient headwater transfer, and that there exists little taxonomic difference be- tween the two subspecies. To avoid taxonomic confusion, which has led to subspecies classification delays, the cutthroat trout in Snake Valley should be considered a unique form of S. c. Utah. Sal- mo c. Utah is not abimdant in any portion of its native range, and at one point it was thought to be extinct as a pure form (Miller 1950, Cope 1955, Platts 1957, Sigler and Miller 1963). The 1973 version of the U.S. Department of Interior's "Red Book" of en- dangered and threatened species listed S. c. Utah as "status undetermined"; the Inter- national Union for the Conservation of Na- ture (1969) listed it as rare; Holden et al. Table 1. Morphomeristic characters used in the dis- criminant function analysis, 1977. Head length Upper jaw length Snout tip to dorsal fin origin Dorsal fin length Caudal peduncle depth Caudal peduncle length Gillrakers upper Gillrakers lower Gillrakers total Branchiostegal rays right Branchiostegal rays left Scales above latera line Pelvic fin rays Pyloric caeca Basibranchial teet (1974) considered it endangered; the Wyo- ming Game and Fish Department lists it as rare; the Nevada Fish and Game Depart- ment considers it endangered; and Behnke (1973, 1976b) considers it to be rare with a highly restricted distribution. Cutthroat Discovery in the Pilot Peak Range In an effort to locate additional popu- lations of Bonneville Basin cutthroat trout, a survey of the Pilot Peak Range (North of Wendover on the Utah-Nevada border) was conducted in 1977 by the BLM and Colo- rado State University (under a contract fun- ded by BLM). As a result of these surveys, only two streams were found containing sufficient an- nual flows to support trout populations. One stream, to the north of Pilot Peak, Bet- tridge Creek, has an abundant population of rainbow trout which were first stocked by the Utah Division of Wildlife Resources in the 1940s or early 1950s. The other stream, located in the adjacent canyon to the south of Bettridge Creek, is unnamed (for the present we have called it Donner Creek be- cause it historically drained into Donner Springs). The city of Wendover, Utah, ob- tains a portion of its water supply from this creek. Mr. Kent Sumners, Utah Division of Wildlife Resources, discovered the cutthroat in Donner Creek in April 1977 while sam- pling the stream at the request of the BLM. Subsequent specimen collection by the au- thors and their later analysis at Colorado State University confirmed this classifica- tion. Taxonomic analysis of the 17 trout sampled from Donner Creek proved most interesting. They are pure strain cutthroat trout (no sign of hybridization) and have a higher gillraker count than any other cut- throat population (24-29, avg. 26.1). The origin of this cutthroat is uncertain; however, Howard Gibson, retired water master for the city of Wendover, indicated that the cutthroat were in Donner Creek when he commenced work on the stream in 1952 (pers. comm. with H. Gibson, Wendo- ver, Utah). None of the other local residents 200 Great Basin Naturalist Vol. 38, No. 2 contacted could provide any information pertaining to the cutthroat, and most were unaware of its existence in Donner Creek. The Nevada Fish and Game Department has no record of cutthroat stockings in the Pilot Peak Range (letter to Don Duff, BLM, SLC from Pat Coffin, Nevada Fish and Game Dept., Elko, October 1977). The only cutthroat exhibiting such high gillraker numbers is the Lahontan cutthroat trout (S. c. henshawi) (Behnke and Zam 1976). The most probable origin of the Donner Creek cutthroat is Pyramid Lake, because, from the late 1890s to 1930, cutthroat trout from Pyramid Lake were stocked extensively in Nevada. In 1910 Elko County received a large shipment of eggs, but no records exist on where these fish were stocked. Little stocking of Lahontan cutthroat occurred from 1931-1942, but in 1950 Lahontan trout from Summit Lake, Nevada, were used for stocking. After 1930 S. c. henshawi was considered rare, and it seems unlikely that a creek in the Pilot Range would be stocked with this cutthroat subspecies. The discriminant function analysis (Table 1, Fig. 3) indicates that the cutthroat from Donner Creek are the most similar to S. c. henshawi. Summary The Snake Valley cutthroat, a form of S. c. Utah, is a unique desert fish resource lo- cated in the western Bonneville Basin which is worthy of protection and management for the scientific community as well as the American public. S. c. Utah has promising possibilities for enhancing the basin states' fishery programs for wild trout manage- • • DONNER HENSHAWI • HUMBOLT RIVER • STOMIAS SNAKE VALLEY # • UTAH #PLEURITICUS • STOMIAS • GAIRDNERI Fig. 3. Discriminant function plot analysis chart showing relationship to cutthroat subspecies based on mor- phomeristic characters. June 1978 Hickman, Duff: Cutthroat Trout 201 ment. The 1975 listing of endangered and threatened fishes of the western U.S. devel- oped by the Desert Fishes Council did not consider this subspecies in its listing of sen- sitive western fishes. It is hoped that recog- nition of this subspecies for management concern will serve as an aid to organiza- tions and agencies responsible for the man- agement of habitat and this subspecies in the future. The ultimate management design for this subspecies and all others so recog- nized is to provide management to a degree whereby survival and protection of the spe- cies and its habitat are assured, so official status classification by the U.S. Fish and Wildlife Service is not necessary. However, should environmental conditions continue to deteriorate and this subspecies eventually become listed by the U.S. Fish and Wildlife Service, then a classification of "threatened" would provide the necessary protective sta- tus while still allowing for state-federal re- covery programs to function. The interest in desert fishes management has intensified by agencies and the scientific community by the discovery in 1977 of S. c. henshawi in Donner Creek of the Pilot Peak Mountain Range. The major signifi- cance of this find of S. c. henshawi is that it very likely represents the original Pyra- mid Lake genotype— the largest trout native to western North America and long be- lieved to be extinct (Trojnar and Behnke 1975, Behnke and Zam 1976). This find is worthy of intense management effort by the Utah Division of Wildlife Resources (DWR) and the BLM because the existence of this pure strain fish is extremely limited, as in- dicated by its official threatened status by the U.S. Fish and Wildlife Service. Colo- rado State University is continuing contract studies on this mountain range for the BLM. The BLM in Utah plans to imple- ment the Pilot Peak Mountains HMP in 1978 imder Sikes Act authorities in cooper- ation with the DWR. Stream habitat im- provements are being planned for Bettridge Creek, which at present has a natural re- producing population of rainbow trout. This creek could serve in the future as a possible transplant site for the Lahontan cutthroat in Donner Creek. Both creeks have good stream habitat, being in a relatively undis- turbed state from man and livestock activi- ties and located in a remote area adjacent to the arid wastes of the Great Salt Lake desert salt flats. Literature Cited Behnke, R. J. 1973. Status report: Utah or Bonneville cutthroat trout, Salmo clarki Utah. Prepared for Region 2 U.S. Fish and Wildlife Service, Albu- querque, N.M. 4 pp. 1976a. A summary of information on a unique form of cutthroat trout native to the Snake Val- ley section of the Bonneville Basin, Utah and Nevada. Prepared for BLM, Salt Lake City, Utah. 18 pp. 1976b. Summary of information on the status of the Utah or Bonneville Basin cutthroat trout, Salmo clarki Utah. Prepared for Wasatch Na- tional For., Salt Lake City, Utah. 14 pp. 1977. Livestock grazing impact on stream fish- eries: problems and suggested solutions. Paper presented at Livestock and Wildlife-Fisheries Workshop, Sparks, Nevada, May 1977. 10 pp. Behnke, R. J., and M. Zarn. 1976. Biology and man- agement of threatened and endangered western trouts. USDA Forest Service general technical Rept. Rm-28. Rocky Mountain Forest and Range Exp. St., Fort Collins, Colorado. 45 pp. Broecker, W. S., and a. Kaufman. 1965. Radio car- bon chronology of Lake Lahontan and Lake Bonneville II, Great Basin. Bull. Geol. Soc. Amer.:76:537-566. Cope, O. B. 1955. The future of the cutthroat trout in Utah. Proc. Utah Acad. Sci. Arts and Lett.:32:89-93. Duff, D. A. 1974. Reconnaissance survey for Utah cut- throat aquatic habitat in the Deep Creek Mountains. BLM Utah State Office, unpub- lished report. 6 pp. Frantz, T. C, and D. J. King. 1958. Completion re- port, Nevada Fish and Game Commission, Stream and Lake Survey. 188 pp. Gilbert, G. K. 1879. In: J. W. Powell (ed.). Lands of the arid region, pp. 50-65. 1890. Lake Bonneville. Monogr. U.S. Geol. Surv., 438 pp. Hickman, T. J. 1977. Studies on relict populations of Snake Valley cutthroat trout in western Utah 1976. U.S. BLM, Utah State Office, 41 pp. Holden, p. B., W. White, G. Sommerville, D. Duff, R. Gervais, and S. Gloss. 1974. Threatened fishes of Utah. Proc. Utah Acad. Sci. Arts and Lett. 51(2):46-55. Hunt, C. B., H. Varnes, and H. Thomas. 1953. Lake Bonneville geology of northern Utah Valley, Utah. U.S. Geol. Surv. Prof. Pap. 15 pp. McLelland, W. L. 1975. Cutthroat trout in- vestigation, Mt. Wheeler cutthroat trout. White Pine County, Region II. Nevada Fish and Game Department. Prog. No. F-20-11 Job No. 207. 24 pp. 202 Great Basin Naturalist Vol. 38, No. 2 Miller, R. R. 1950. Notes on the cutthroat and rain- bow trouts with the description of a new spe- cies from the Gila River, New Mexico. Occa- sional Pap. Mus. Zool. Univ. Mich. (529):42 pp. Miller, R. R., and J. R. Alcorn. 1946. The in- troduced fishes of Nevada, with a history of their introduction. Trans. Am. Fish. Soc. 73, 1943:173-193. Plaits, W. S. 1957. The cutthroat trout. Utah Fish and Game Magazine, 13(10):4,10. 1977. Livestock interactions with fish and their environment. Paper presented to Calif.- Nevada AFS. 12 pp. SiGLER, W. F., AND R. R. MiLLER. 1963. Fishes of Utah. Utah Dept. Fish and Game. 203 pp. Smith, G. R. 1966. Distribution and evolution of the North American catostomid fishes of the sub- genus Pantosteus, genus Catostomus. Miscl. Publ. Mus. Zool. Univ. Mich. (129): 132 pp. Trojnar, J. R., AND R. J. Behnke. 1974. Management implications of ecological segregation between two introduced populations of cutthroat trout in a small Colorado Lake, Trans. Amer. Fish. Soc. 103:423-430. Wydoski, R. S., G. Klar, T. Farley, J. Braman, Y. Kao, and C. Stalnaker. 1976. Genetic bio- chemical and physiological studies of trout en- zymes. NMFS Final Rept. Prog. No. I-87R. 163 pp. REPRODUCTIVE BIOLOGY OF THE TUI CHUB, GILA BICOLOR, IN PYRAMID LAKE, NEVADA Paul A. Kucera' Abstract.— Sex ratios for tui chubs {Gila bicolor), obtained from a sample of 3,384 fish, deviated significantly from the expected 1:1 ratio (chi-square, P<0.05). Spawning occurred from June to August, with estimated fecun- dities ranging from 6,110 to 68,933 ova. Females spawned after attaining a maximum (average) gonadal somatic index (GSI— percent gonad weight/total body weight) of 9.1 percent and between surface water temperatures of 15.5 to 22.2 C. Males reached sexual maturity at age two, with most females maturing at age three. Consistent increases in fecundity were apparent with increasing length, weight, and age. Linear regressions between fork length and fecundity and weight and fecundity were highly significant (P<0.05). The tui chub (Gila bicolor) is a cyprinid found in the drainage systems on both sides of the Sierra Nevada, from the San Joaquin and Lahontan systems north to the lakes of southern Oregon and the Columbia River (LaRivers 1962). In Pyramid Lake G. bicolor comprises nearly 89 percent of the net catch and represents the main forage base for the Lahontan cutthroat trout (Salmo clarki henshawi). Tui chubs also represent a principal component in the bioenergetics of the Pyramid Lake ecosystem. The present study is a part of the total ecological research and fish life history studies being conducted on Pyramid Lake. Those aspects of the reproductive biology that were studied included: sex ratios, total and by month; changes in fish catch rates and sex ratios; development of the ova and ovaries in relation to the spawning season; age and size at sexual maturity; and number of ova in relation to fish length and weight. Methods Tui chubs were collected monthly from November 1975 through June 1977 in Pyra- mid Lake, Nevada. Two gear types were utilized in the collection of chubs: variable- mesh bottom-set gill nets and fyke nets. Fe- males selected for fecundity studies were collected during Jime and July of 1976 and 1977. These months were selected to ensure collection of ripe females. Ovaries were removed from 23 tui chubs for fecundity determination. Each pair of ovaries was weighed to ± 1 mg, and 10 ovum diameters per ovary were measured to the nearest 0.1 mm with an ocular mi- crometer. The fork length of each fish was measured to the nearest millimeter, and each fish was weighed to the nearest gram. All fecundity determinations were made with fresh ovaries. Fecundity was estimated by the gravi- metric method (Jester 1973; June 1971). Three replicate subsamples of approx- imately 0.5 g were taken from each ovary, weighed to the nearest 1 mg, and preserved for later counting. Fecundity was then cal- culated by direct proportion per ovary. Sex ratios were obtained through internal examination of 3,384 fish. Age groups were assigned by the scale method and length frequency histograms. Linear and logio re- gressions between fecundity and fork length and fecundity and weight were used to ex- amine the relationships between these vari- ables. Discussion Sex ratios for tui chubs were obtained from a sample of 3,384 fish sampled from 'W. F. Sigler & Associates Inc., 1005 Terminal Way, Suite 155, Reno, Nevada 89502. 203 204 Great Basin Naturalist Vol. 38, No. 2 Table 1. Monthly sex ratios of adult tui chubs sampled from November 1975 through February 1977 in Pyra- mid Lake, Nevada. Months N D J F M A M J J A S O N D J F Males eo 74 52 14 76 96 126 462 93 95 79 87 56 31 92 46 Females 91 104 84 51 171 179 134 289 106 75 76 96 100 80 155 54 Ratio M:F 1:1.52 1:1.41 1:1.62 1:3.64 1:2.25 1:1.86 1:1.06 1.6:1 1:1.14 1.27:1 1.04:1 1:1.1 1:1.78 1:2.58 1:1.68 1:1.17 November 1975 through February 1977. The 1:1.20 ratio, 1,539 males to 1,845 fe- males, represents a significant deviation from the expected 1:1 ratio (chi-square, P<0.05). Sex ratios by month (Table 1) present interesting fluctuations. Substantial deviations in sex ratios for duplicated months (December and February) are read- ily apparent and heavily favor the females. This is probably the result of sampling bias. The important trend takes place from April through August, which coincides with the spawning season. In April, the ratio of males to females is almost 1:2. This changes dramatically in May to 1:1.06, in June to 1.6:1, to 1:1.14 in July, and 1.27:1 in Au- Fig. 1. Monthly changes in sex ratios (solid line) and inshore catch rates (dash line) of tui chubs in Pyramid Lake in 1976. Water temperatures are monthly inshore surface averages. gust. Deviations in sex ratios in May and June could be attributed to the movement of males to spawning sites. Of 262 tui chubs sexed from one inshore bottom gill net in early June 1976, 86 percent were males. Data collected from Walker Lake, Nevada (Cooper, personal communication), in mid- June parallel this result (85 percent of the tui chubs sampled in 5-7 m of water were males). Increases in the percentage of males sam- pled from April through June are accom- panied by an increased bottom inshore catch of tui chubs (Fig. 1). Although month- ly sex ratios were shown to be variable (Table 1), they usually favored the females. From May to June, the percent of males in- creased from 49 to 62 percent, indicating a prespawning movement of males inshore. Catch rates also increased dramatically in May, and steadily rose in Jime. An influx of females in July, corresponding to peak spawning, decreased the percentage of males to 46 percent. Increased activity, and thus increased catches, for both sexes, also parallel major periods of zooplankton stand- ing crops, macroinvertebrate abimdances, increasing water temperatures, feeding habits, and growth rates. The overall catch rate inshore in August decreased by 71 per- cent from the bottom inshore average num- bers of June and July. This corresponds with the completion of the spawning season and must result from differential behavior and distribution of mature fish related to spawn- ing. Constant increases in the average female gonadal somatic index (GSI— percent gonad weight /total body weight) were noted from February through May, with a major in- crease of 3.8 percent in June (Fig. 2). This period of major germinal growth (May to June) took place after the major increase in June 1978 KucERA: Tui Chub 205 somatic growth (fork length) in fall and winter. Consequently, tui chubs foraged very actively to support both germinal and somatic increases. Females attained a max- imum average GSI in June of 9.2 percent as inshore water temperatiu-es approached 15.5 C. Zooplankton standing crops and mac- roinvertebrate abundances were both high during this period, thus providing ample food sources. Spawning occurred from June to August, with peak spawning in July when the GSI dropped from 7.3 percent to 1.5 percent in August. Inshore water tem- peratiues in June were 15.5 C and in July were 22.2 C. In Eagle Lake, California, tui chubs spawned when water temperatures reached about 15.5 C (Kimsey 1954). Spawning activity of Mojave chubs, Gila mohavensis, in Soda Lake, California, peaked in mid-March at water temperatures of 18 C (Vicker 1973). LaRivers (1962) states that tui chubs con- gregate along the shoreline during the spawning season. Bottom gill net catches re- veal that almost 97 percent of the tui chubs were inshore in July. This corresponds with the time of peak spawning. Although no spawning activity or spawn was observed during this study, chubs have been seen spawning in shallow water in Pyramid Lake (Galat, pers. comm.). Initiation of sampling during the summer of 1977 to determine vertical distribution patterns suggested the possibility of two separate chub popu- lations: a fine-rakered limnetic population and a coarse-rakered population. The tax- 10 9 8 7 6 5 4 3 2 Water Temperature Spent J A Months - 10 - 4.4 26.6 21.1 n> 15.5 f 1.3 1.2 1.1 1.0 .9 .8 .7 .6 Fig 2. Comparison of monthly tui chub gonadal somatic index values (solid line) with ova diameters (dash line). Water temperatures are monthly inshore surface averages. 206 Great Basin Naturalist Vol. 38, No. 2 onomic controversy concerning the tui chub is well documented (Hubbs and Miller 1943, Hubbs et al. 1974, Kimsey 1954, LaRivers and Trelease 1952, LaRivers 1962). It is be- lieved that the spawning discussion includes the bottom-dwelling chubs, but the paucity of data on pelagic chubs precludes any con- clusion that they also spawn inshore. Varley and Livesay (1976) observed movement of prespawning and spawning populations of Utah chub, Gila atraria, in Flaming Gorge Reservoir, Utah-Wyoming. In the summer these chubs move onshore into littoral areas from mid to late afternoon, apparently to spawn, and then move lakeward into pelag- ic areas by early morning hours. Diel distri- bution patterns of Utah chubs in Fish Lake, Utah paufin 1964), also indicate concentra- tions of chubs in littoral areas during the night. In Pyramid Lake, female tui chubs are sexually mature at two years of age, and all are mature by age three; the majority of males also mature at two and all by three. After attaining sexual maturity, chubs ripen and spawn every year. The oldest mature female sampled was of age group VII, and the oldest mature male was of age group IV. Upon attaining matiu-ity, Utah chub males showed a higher rate of mortality as age group V, and older females out- numbered males almost 10 to 1 (Varley and Livesay 1976). Age group III chubs comprised the ma- jority of females sampled for fecundity stud- ies (Table 2). Three-year-old females aver- aged 232 mm in fork length and 15,135 ova per female. Age group IV females averaged 277 mm in fork length and 31,622 ova per female. One age group V female was 307 mm in fork length and contained an esti- mated 50,032 ova. One age group VII fe- male was 378 mm long and contained an estimated 68,933 ova. Estimated egg pro- duction per female ranged from 6,110-68,933 eggs and averaged 23,292 ova. Olson (1959) found the average number of eggs for Utah chubs from Scofield Reser- voir, Utah, to be 25,282 ova per female. In Hebgen Lake, Montana, Graham(1955) found Utah chub fecundities to average 40,750 eggs per female, although his sample size of n = 7 was small. Estimated fecim- dities for six Mojave chubs averaged 12,687 ova per female (Vicker 1973). Tui chubs ex- hibited consistent increases in fecundity, with increases in length, weight, and age (r values; P<0.05). Ova diameters, from green ovaries, ranged from 1.1-1.7 mm and aver- aged 1.3 mm for all year classes. Kimsey (1954) described newly extruded tui chub eggs as being 1.5-1.9 mm in diameter, and Harry (1951) stated that freshly stripped tui chub eggs were 1.8-2.0 mm in diameter. The linear relationship between fork length and fecundity (Fig. 3) was highly sig- nificant (F = 94.8; P<0.05). A significant (P<0.05) correlation coefficient exists be- tween fork length and fecundity, because 82 percent of the variation in fecundity was accounted for by variation in length. The positive linear relationship between weight and fecundity was also highly significant (F = 211.2; P<0.05), demonstrating an in- crease in fecundity with increasing weight (Fig. 4). A significant (P<0.05) correlation coefficient exists, and 91 percent of the var- iation in fecundity was explained by varia- tion in weight. Both fork length and fectm- dity and weight and fecundity had Table 2. Average lengths, weights, ovum diameters, and fecundities, per age group, for 23 tui chub from Pyra- mid Lake, Nevada, 1976 and 1977. Number Range in of Range in Average Range in Average Egg Average Tui Age Fork Length Fork Length Weight Weight Diameter Diameter Chub Group (mm) (mm) (grams) (grams) (mm) (mm) 15 6 1 UI 210-252 IV 263-323 V - VI VII 232 277 307 378 96-238 196-410 154 317 444 661 1.1-1.5 1.2-1.4 1.3 1.3 1.3 1.35 Range in Number Average of eggs Number 15,135 31,622 50,032 68,933 6,110-31,432 19,307-36,993 June 1978 KucERA: Tui Chub 207 Fig. 3. The relationship between fork length and fe- cundity for 23 tui chubs from Pyramid Lake, Nevada, 1976 and 1977. Fig. 4. The relationship between weight and fecun- dity for 23 tui chubs from Pyramid Lake, Nevada, 1976 and 1977. significant logjo fits but were best described by a linear fit. Acknowledgments I wish to acknowledge the entire crew of W.F. Sigler & Associates Inc., especially Steven Vigg and Denise Robertson, for as- sistance and support in this project. Literature Cited Gaufin, R. F. 1964. Ecology of the Utah chub in Fish Lake. Master's thesis, Univ. of Utah. Ill pp. Graham, R. J. 1961. Biology of the Utah chub in Heb- gen Lake, Montana. Trans. Amer. Fisheries Soc. 90(3):269-276. Harry, R. R. 1951. The embryonic and early larval stages of the tui chub, Siphateles bicolor (Gi- rard), from Eagle Lake, California. California Fish and Game 37(2): 129-132. HuBBS, C. L., AND R. R. Miller. 1943. Mass hybridi- zation between two genera of cyprinid fishes in the Mohave Desert, California, Michigan Acad. Sci., Arts and Letters 28:343-378. HuBBS, C. L., R. R. Miller, and L. C. Hubbs. 1974. Hydrographic history and relict fishes of north- central Great Basin. In: Memoirs of the Califor- nia Acad, of Sci. 259 pp. Jester, D. B. 1973. Life history, ecology, and manage- ment of the smallmouth buffalo, Ictiohus bu- halus (Rafinesque), with reference to Elephant Butte Lake. Agric. Exp. Stn. Res. Rep. 261, New Mexico State University, Las Cruces. Ill pp. June, F. C. 1971. The reproductive biology of northern pike, Esox lucius, in Lake Oahe, an upper Mis- souri River storage reservoir. In: G. E. Hall (ed.), Reservoir fisheries and limnology, pp. 53-71. Spec. Publ. 8, Am. Fish. Soc. KiMSEY, J. B. 1954. The life history of the tui chub, Siphateles bicolor (Girard), from Eagle Lake, California. California Fish and Game 40(4):395-410. LaRivers, I. 1962. Fishes and fisheries of Nevada. Nevada State Fish and Game Comm. 782 pp. LaRivers, I., and T. Trelease. 1952. An annotated checklist of the fishes of Nevada. California Fish and Game 38(1):113-123. Olson, H. F. 1959. The biology of the Utah chub, Gila atraria (Girard), of Scofield Reservoir, Utah. Master's thesis, Utah State University. 34 pp. Varley, J. D., AND J. C. LivESAY. 1976. Utah ecology and life history of the Utah chub, Gila atraria, in Flaming Gorge Reservoir, Utah-Wyoming. Utah Division of Wildlife Resources Pub. No. 76-16. 29 pp. ViCKER, C. E. 1973. Aspects of the life history of the Mojave chub, Gila bicolor mohavensis (Snyder) from Soda Lake, Calif. Master's thesis, Califor- nia State University, Fullerton. 27 pp. DISTRIBUTIONAL RECORDS FOR THE COLORADO FLORA D. Wilken', W. Har^lon^ C. Feddema' and H. Harrington' Abstract.— Nine new records of vascular plants indigenous to western North America, six new records of ad- ventive taxa, and six significant range extensions are reported for the flora of Colorado. The following collections document new records and range extensions for vascular plants as treated by Harrington (1954), We- ber (1976), and Weber and Johnston (1976). Herbarium citations follow the recent treat- ment by Holmgren and Keuken (Index Her- bariorum, Ed. 6, 1974). Indigenous New Records X Agrositanion saxicola (Scribner & Smith) Bowden Mesa Co.: 2 miles south of Monument Hill, 9,700 ft, 28 August 1947, W. A. We- ber 3660 (CS). This specimen possesses the narrow, bifid to trifid glumes, long diver- gent awns, and tardily disarticulating rachis characteristic of Sitanion X Agropyron hy- brids discussed in Cronquist et al. (1977). Astragalus musiniensis Jones Garfield Co.: Badger Wash Experimental Area, 10 miles northwest of Mack, 5,000 ft, 7 June 1973, Reid & Ranck s.n. (USES). Conimitella williamsii (D.C. Eaton) Ryd- berg Summit Co.: along Pass Creek, 2 miles east of State Highway 9, along Ute Pass Road, 8,600 ft, 26 June 1973, D. Walsworth s.n. (CS, USES); 27 June 1975, W. A. We- ber and P. Nelson 15119 (COLO, CS). This site represents a southward range extension of ca. 200 miles from the nearest previously known localities in northern Wyoming. Drymaria depressa Greene Boulder Co.: 2.75 miles west of Copeland Lake, 9,700 ft, 16 August 1976, F. Hermann 27405 (CS). Erigeron ochroleucus Nuttall Rio Blanco Co.: 6.6 miles north of Rio Blanco, 7,200 ft, 4 June 1977, Wilken and Schwab 13058 (CS). This collection docu- ments predictions of its occurrence in Colo- rado by Harrington (1954) and Weber and Johnston (1976). Cilia micromeria A. Gray Mesa Co.: Monument Valley Overlook, Colorado National Monument, 1,700 m, 11 May 1974, Wilken and Kelley 11988 (CS); Moffat Co.: near mouth of Hell Canyon- Yampa Canyon, 5,200 ft, 14 May 1948, Harrington 3906 (CS). Heliotropium convolvulaceum (Nuttall) A. Gray var. californicum (Greene) Johnston Mesa Co.: Rabbit Valley Road, 4 miles south of Highway 50, 1,450 m, 6 July 1976, Wilken, Donahue and Tabar 12684 (CS). This population was represented by only several individuals and should be considered rare in Colorado. The typical variety occurs in local abundance in southeastern Colo- rado. Parthenium alpinum (Nuttall) Torrey & Gray var. alpinum Weld Co.: 3.5 miles north-northwest of junction of County Roads 45 and 122, 5,800 ft, 4 June 1977, Harmon 8919 (CS, GREE), Harmon 8810 (CS, GREE). 'Department of Botany and Plant Pathology, Colorado State University, Fort Collins, Colorado 80523. 'Department of Biological Sciences, University of Northern Colorado, Greeley, Colorado 80639. 'U.S. Forest Service Herbarium, Fort Collins, Colorado 80521. 208 June 1978 WiLKEN ET AL.: COLORADO FlORA 209 Prunus angustifolia Marshall Baca Co.: 33 miles southeast of Springfield, 4,200 ft, 26 July 1956, Harring- ton 8319 (CS). Typha domingensis Persoon Prowers Co.: 5 miles east of Lamar, 6 October 1957, Hotchkiss 7577 (CS); 1 mile south of Bristol, 3,400 ft, 27 July 1959, Har- rington 9102 (CS). Adventive New Records Eragrostis curvula (Schrader) Nees Lincoln Co.: 3 miles south of Hugo, 5,000 ft, 2 October 1957, Hamill s.n. (CS). Fumaria officinalis L. Weld Co.: 2 miles west of Windsor, 4,780 ft, 31 May 1957, Klein s.n. (CS). Recent ob- servations by the senior author indicate that the population is well established and spreading along an irrigation ditch. Anoda cristata (L.) Schlechtendahl Otero Co.: Experiment Station near Rocky Ford, 12 October 1966, Swink s.n. (CS). Montia perfoliata (Donn) Howell Douglas Co.: T. 7 S., R. 69 W., S. 4, 6,000 ft, 14 May 1960, Brunquist s.n. (CS). Efforts to relocate this population have been unsuccessful. Although indigenous to the Pacific Northwest and with nearest known natural populations in northeastern Wyoming, this record is considered to be adventive. Prunus anneniaca L. Garfield Co.: Glenwood Canyon, 4.4 miles east of No Name, 6,200 ft, 26 May 1977, Wilken 12935 (CS). First reported as established in Mesa Verde National Park (Welsh and Erdman 1964), this species is well established and reproducing in Glen- wood Canyon. Puccinellia lemmonii (Vasey) Scribner Boulder Co.: north of Valmont Rd., be- tween the railroad line and Boulder Creek, 2 September 1975, Zanoni 2892 (CS). Ac- cording to Zanoni (pers. comm.) seed of this species was included in a general mix used for local rehabilitation. Range Extensions Ipomopsis roseata (Rydberg) V. Grant Moffat Co.: 3 miles north of Sunbeam, 25 May 1938, Brown s.n. (CS). This site repre- sents a range extension of approximately 100 miles north-northeast of the principal distribution in Colorado National Mon- ument. Lysitnachia thyrsiflora L. Larimer Co.: Wet ground along Poudre River, 23 June 1893, Cowen s.n. (CS). Known previously only from a single site in Colorado near Estes Park (Weber, 1976) this species still occurs as sporadic individ- uals near the crossing of the Poudre River and Highway 14 on the eastern limits of Ft. Collins. Malacothrix torreyi A. Gray Montrose Co.: Naturita, 15 May 1914, Payson 303 (RM); Garfield Co.: Rifle, 25 June 1900, Osterhout 2150 (RM); Eagle Co.: McCoy, 14 June 1903, Osterhout 2758 (RM). This species is treated as rare and oc- curring only in Mesa and Delta Counties (Weber and Johnston 1976). The specimens cited above indicate a wider distribution in western Colorado. Rhododendron albiflorum Hooker Jackson Co.: Lake Katherine, Park Range, 3,000 m, 26 August 1976, Wilken and Painter 12752 (CS); 11 July 1977, Wilken, Painter and Tabar 13178 (CS). Previously known in Colorado only from the head- waters of the South Fork Elk River in Routt Co., this second site represents a range ex- tension east of the continental divide. The population at Lake Katherine is extensive and comprises the dominant understory be- neath Abies lasiocarpa and Picea engel- mannii. Based on specimens at MONT and MONTU, the Colorado populations repre- sent a disjunction of approximately 500 miles southeast of the nearest known sites in western Montana. Ribes americanum Mill. Larimer Co.: Wet seeps along the Poudre River, 20 May 1977, Budzinski s.n. (CS). Known previously only from Roxborough Park in Douglas Co. (Weber, 1976), this species occurs in local abundance near the crossing of the Poudre River and Highway 14 at the present limits of Fort Collins. Salix Candida Fluegge Larimer Co.: 1.5 miles south of the con- 210 Great Basin Naturalist Vol. 38, No. 2 fluence of the Laramie River and Two and One Half Creek, 8,920 ft, 15 July 1974, Phillips 85 (CS). This site represents the second record for Salix Candida in Colo- rado, formerly known only from South Park in Park Co. Literature Cited Harrington, H. D. 1954. Manual of the plants of Col- orado. Denver: Sage Books. Weber, W. A. 1976. Rocky Mountain flora. Boulder: Colorado Associated University Press. Weber, W. A., and B. C. Johnston. 1976. Natural his- tory inventory of Colorado. 1. Vascular plants, Uchens and bryophytes. Boulder: University of Colorado Museum. Welsh, S. L., and J. A. Erdman. 1964. Annotated checklist of the plants of Mesa Verde, Colorado. Brigham Young University Sci. Bull. Biol. Series 4(4): 1-32. A COMPARATIVE ANALYSIS OF THE DIATOM FLORA ON THE SNAIL AMPULLARIA CUPRINA FROM THE GOSHEN PONDS, UTAH Larry L. St. Clair', Lorin E. Squires^-^, and Samuel R. Rushforth' Abstract.— Seventeen genera and 60 species of diatoms were identified from four different substrates in a warm spring-fed pond near Goshen, Utah. The diatom forms sampled included both plankton and periphyton. A minimum of 400 individuals were counted from each sample. Diversity and similarity indices were calculated, and substrate relationships were identified on their basis. Diatoms have long been known to grow on a wide variety of substrates and in a highly diversified range of habitats. Just such a range of distribution has been ob- served to occur in the Intermountain West, where diatom research during the last seven years has revealed over 700 species of dia- toms in many different aquatic and terres- trial ecosystems. During the period between April 1973 and April 1975, the diatom flora of the Goshen ponds and wet meadows, Utah County, Utah (a series of warm, spring-fed ponds), was under investigation (St. Clair and Rushforth 1977). Periodically during this study, several large specimens of Am- pullaria cuprina (mystery snail) were col- lected from the site designated "Lower Pond." A dense growth of algae colonized the shells of these organisms. In May 1974, several large specimens of this snail were collected and taken to the laboratory in or- der to identify and characterize the at- tached algal flora. The present paper treats the diatom species identified from the shell of the snail and compares diatom commu- nities on the snail to planktonic, epiphytic, and epilithic communities in the lower pond. Materials and Methods In early May 1974, four substrates in the Lower Goshen Pond were sampled. The plankton was sampled by filtering pond wa- ter through a 67-jLim mesh plankton net and concentrating it in a 30-ml vial attached to the net. Periphyton on the vegetable mate- rial in the pond was sampled by collecting various reeds, grasses, and filamentous algae from the pond. Several rocks from the bot- tom of the pond were scraped in order to obtain epilithic diatoms. Finally, two speci- mens of Ampullaria cuprina (mystery snail) were collected. All samples were taken to the laboratory, and the diatoms were clean- ed using acid oxidation methods (St. Clair and Rushforth 1976). Cleaned frustules were dried on cover slips and subsequently mounted in Naphrax mountant. Diatoms were identified and counted us- ing a Zeiss RA microscope with Nomarski interference phase-contrast accessories and a lOOx oil immersion objective. A minimum of 400 individual diatoms were counted per sample to obtain relative density informa- tion for each species. Diversity and similarity indices were cal- culated using an IBM 360 model 5 com- puter in order to obtain community struc- ture information for each substrate. The diversity index was calculated using a pro- gram based on the formula d = -2 (Ni/N) Log2 (Ni/N) i in which S = the number of species, Ni == the number of individuals in species i, and 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 'Present address: Vaughn Hansen Associates, Waterbury Plaza, Suite A, 5620 South 1475 East, Salt Lake City, Utah 84121. 211 212 Great Basin Naturalist Vol. 38, No. 2 N = the total number of individuals in S species (Cairns and Dickson 1971, Wilhm and Dorris 1968). This formula is independent of sample size and gives an index which is high when many species are evenly represented in the community and low when most of the indi- viduals occur in only a few species. The similarity index was calculated from Ruzicka's (1958) formula: SI = 2 min (xi,yi) i 2xi + 2yi - 2min (xi,yi) i i i in which xi and yi = relative density for the ith species in the two samples being compared. This index gives a quantitative measure of the similarity between two stands of data. Results The diatom flora from the four sites in- cluded 17 genera and 60 species (Table 1). Of these, 12 genera and 40 species were found on the snail, 11 of the species being unique to the snail's shell. The 5 most com- mon species (Table 2) comprised 61.7 per- cent of the diatom flora from the snail's shell. The remaining species had relative densities below 4.2 percent, and 20 species were below 1 percent. A comparison of the diatom communities from the four habitats suggested some gen- eral relationships. The rock and vegetable substrates had fewer total species and lower diversity (Table 3) than occurred in the plankton or on the snail shells. The lowest diversity occurred on the vegetable sub- strate, which appeared to be the most spe- cialized habitat of the four. It also had the least similarity to the other habitats (Fig. 1), Achnanthes minutissima was the diatom best adapted to the vegetative habitat, com- prising 59 percent of the flora. The higher diversity and total number of species in the plankton and snail samples were due to in- stability in these habitats. Instability in the plankton environment was caused by surface turbulence and the upswelling of subterranean water. As a re- sult, many benthic diatoms were present in the plankton. The similarity of the rock and plankton habitats (Fig. 1) indicates that the influx of epilithic diatoms into the plankton was considerable. Instability in the diatom community on the snail shell was probably a result of the Table 1. Phylogenetic listing of diatom species identified in this study (The species identified with an asterisk were collected from the snail's shell). Order Rhizosoleniales Terpsinoe musica Order Fragilariales Fragilaria brevistriata var. inflata' Fragilaria capucina var. mesolepta' Fragilaria construens'' Fragilaria construens var. venter' Fragilaria crotonensis' Synedra delicatissima' Synedra incisa' Synedra pulchella var. lacerata' Synedra tabulata' Synedra ulna' Synedra ulna var. contracta' Order Eunotlales Eunotia curvata' Order Achnanthales Cocconeis pediculus Cocconeis placentula var. lineata' Achnanthes bottnica' Achnanthes exigua var. heterovalva' Achnanthes exigua Achnanthes lanceolata var. dubia Achnanthes lanceolata' Achnanthes linearis' Achnanthes minutissima' Rhoicosphenia curvata Order Naviculales Anomoeoneis vitrea Diploneis elliptica Navicula cryptocephala var. vetieta Navicula dicephala' Navicula mutica' Navicula pelliculosa' Navicula perpusilla' Navicula pupula Navicula pupula var. rectangularis Navicula radiosa Navicula tripunctata' Navicula sp. 1° Navicula sp. 2 Navicula sp. 3 Pinnularia viridis' Entomoeoneis alata Amphora coffeaeformis Amphora ovalis' Amphora ovalis var. pediculus' June 1978 St. Clair et al.: Diatom Flora 213 Amphora veneta' Cymbella af finis' Cymbella cistula' Cymbella delicatula' Cymbella laevis Cymbella ventricosa' Cymbella sp." Gomphonema angustatum Gomphonema constrictum' Gomphonema gracile' Gomphonema intricatum' Gomphonema parvulum' Order Epithemiales Denticula elegans' Order Nitzschiales Nitzschia denticula Nitzschia dissipata Nitzschia frustulum' Nitzschia frustulum var. perminuta' Nitzschia palea snail's motility. The shell is conducive to di- atom attachment and growth, and, as the snail moves among the aquatic vegetation and stony substrate, it becomes seeded with diatoms which occur in these habitats. The continuous movement creates instability by causing frequent removal and addition of diatom frustules, thus preventing the domin- ance of any one species and providing for colonization of a wider variety of species than commonly occurs on a single station- ary substrate. For this reason the diatom flora on the snail's shell had higher diversity and lower similarity to the rock sample than might be expected. A more complete and replicated sampling program is planned to further investigate Table 2. Diatom species with relative density above 5 percent in at least one habitat. Vegetative Snail Rock Plankton Achnanthes exigua var. heterovalva Achnanthes minutissima Cymbella delicatula Fragilaria brevistriata var. inflata Fragilaria construens Fragilaria construens var. venter Fragilaria crotonensis Synedra ulna 1.4 11.0 7.7 8.3 59.0 20.0 36.6 24.5 .4 1.1 11.7 1.5 2.1 7.5 7.9 3.9 .7 8.2 7.0 5.1 .2 15.0 3.9 3.9 16.2 .9 3.6 11.2 5.2 2.2 2.7 2.9 Percent Similarity 30 40 50 vegetable snail rock plankton Fig. 1. Cluster dendrogram showing similarity of diatom communities from plankton, vegetable, snail, and rock substrates. Similarity indices were computed on the basis of relative density for diatom species. 214 Great Basin Naturalist Vol. 38, No. 2 the differences and relationships which exist in the plankton, rock, snail, and vegetable habitats of Lower Goshen Pond. Table 3. Shannon Weaver diversity index for dia- tom communities from four habitats in Lower Goshen Pond. Sample Diversity Index Total Species Vegetative Snail Rock Plankton 2.42 4.07 3.34 3.82 31 40 29 36 Literature Cited Cairns, J., and K. L. Dickson. 1971. A simple method for the biological assessment of the effects of waste discharges on aquatic bottom-dwelling or- ganisms. JWPCF 43(5):755-772. RuziCKA, M. 1958. Anwendung mathematisch-statistis- cher methoden in der geobotanik (synthetisch e bearbeitung von aufnahmen). Biologia, Bratisl., 13, 647-661. St. Clair, L., and S. Rushforth. 1976. The diatoms of Timpanogos Cave National Monument, Utah. American J. Botany 63(l):49-59. 1977. The diatom flora of the Goshen warm spring ponds and wet meadows, Goshen, Utah, USA. Nova Hedwigia XXVIII 28(2-3):353-425. WiLHM, J. L., AND T. C. DoRRis. 1968. Biological pa- rameters for water quality criteria. Bioscience I8(6):477-481. Figs. 2-3. Representative specimens of Ampullaria cuprina (Mystery snail). GROWTH OF DIPODOMYS ORDII (RODENTIA: HETEROMYIDAE)' H. Duane Smithy Gary H. Richins', and Clive D. Jorgensen^ Abstract.— Growth rates were determined for laboratory-reared Dipodomys ordii pallidas Durrant & Setzer. Instantaneous growth rates were used to express increase of body weight, total length, tail length, ear length and hind foot length as rates between times of measurements and the instantaneous percentage of maximum size. Data were analyzed for growth periods of 1-3, 4-15, 16-29 and 30-70 days. All five parameters provided signifi- cant correlations of growth with age during all growth periods. Even though all of the growth parameters corre- late with age, these parameters cannot be reliably used to predict age. Dipodomys ordii is one of the most wide- ly distributed small mammal species in North American deserts (Hall and Kelson 1959) and as such occupies a vital position in the bioenergetics of desert ecosystems. Their total role within the ecosystem is not clear, but as primary consumers, they pro- vide an important part of the prey base for higher trophic levels. With increased em- phasis being placed on predictive capabili- ties of productivity within ecosystems, it is desirable to know the potential input of all components. Biomass estimates of rodent populations are often calculated but seldom include estimates for animals below trap- pable age. Dipodomys ordii is no exception. Omission of these types of data is under- standable because such data are difficult to obtain, particularly in the field. They are usually provided by laboratory studies, and since D. ordii has proven difficult to breed and rear in captivity (Morse et al., in press), no data of consequence have been reported for this species. Growth rates have been reported for sev- eral small mammals (For example, Meyer and Meyer 1944, Poumelle 1952, Chew and Butterworth 1959, Layne 1959, Goertz 1965, Hayden and Gambino 1966, Lackey 1967, Jones 1967, Homer and Taylor 1968, Richins et al. 1974, Lackey 1976, and oth- ers). These data have aided in imderstanding productivity, but data on D. ordii are una- vailable. The objectives of this study were to characterize growth rates of individual D. ordii from birth to 70 days of age and to provide a data base for predictive purposes in ecosystem management. Methods The initial specimens of Dipodomys ordii pallidus Durrant & Setzer used in this work were live-trapped on the Desert Range Ex- perimental Station, 81 km W of Milford, Millard Co., Utah. Additional animals were obtained from Dugway Proving Grounds, Tooele Co., Utah, and Pahvant Butte, Mil- lard Co., Utah. The laboratory colony was housed at Brigham Young University and maintained at about 85 females and 15-20 males. Animals were caged individually in glass aquaria or in galvanized metal boxes with sand substrate (about 6.0 cm deep). Nest cans with cotton batting were provided. Water and a food mixture of one part each of sunflower seeds, rolled oats, and pigeon mix were supplied ad libitum. Temperature ranged from 20 to 25 C, and the photo- period was manually graduated to simulate a twilight period. 'This study was supported in part by the US-ffiP Desert Biome studies, contracted from Utah State University with Brigham Young University. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. ^Wildlife Biologist, Bums and McDonnel, Kansas City, Missouri. 215 216 Great Basin Naturalist Vol. 38, No. 2 Sexes were kept separate except when mating was attempted. Females were checked daily for the external morphologic changes in the vulva that Pfeiffer (1960) de- scribed as indicators of estrous. The full flowered vulvar condition was presumed to indicate estrous. At this time a male was in- troduced into the female's cage. Males were selected for breeding based on the extent of testicular enlargement. Usually, fighting en- sued; if the male survived the first few min- utes, he was left with the female for 3-4 days. After this period he was returned to a cage with other males. Females were checked for vaginal plugs as an indication of successful coitus. After litters were bom they were not dis- turbed until the following day, at which time the young were toe clipped and mea- sured. Body weight, total length, tail length, ear length and hind foot length measure- ments were taken daily from days 1-29 and then weekly through the remainder of 10 weeks. After eyes of the young opened and individuals became more active, they were anesthetized with Penthrane (Abbott Labo- ratories, North Chicago, Illinois) to facilitate handling and obtaining accurate measure- ments. The instantaneous relative growth rate (IGR) described by Brody (1945) and Lack- ey (1967) was used to express growth as a rate between times of measurements and the instantaneous percentage of maximum size. This rate is expressed as {dW/dt)/^, where W is the parameter measured at the instant the rate of change dW/dt is mea- sured. Since it is not entirely possible to de- velop the "instantaneous" rate of growth, it was necessary to integrate the infinite num- ber of growth rates to derive W = Ae'^'. This may be conveniently rewritten as— lnW,_i = InA, -I- kt where InW = natural logarithm of the var- iable (W), InA = natural logarithm of the variable (A) and k represents the in- stantaneous relative growth rate (when mul- tiplied by 100, k = percentage growth rate). For comparative purposes, the repre- sentative IGR (k) is determined with— k„ = lnW„ - lnW„_, Thus, k is definitive and can be used to compare differences in rates of growth. Correlation coefficients were calculated be- tween age and growth for the five growth variables studied. Results Growth Rates.— Rates of growth along with the instantaneous relative growth rates (k) were determined and are presented for body weight, total length, tail length, ear length and hind foot length (Figs. 1 and 2 and Table 1). When the variables are plotted on a log scale (Figs. 1 and 2), the comparative values of (k) for the growth pe- riods from days 1-70 are illustrated. These results depict growth for animals reared un- der standardized laboratory conditions and analyzed for time growth periods of 1-3, 4-15, 16-29 and 30-70 days. Other growth periods were graphed and analyzed but these represent the most statistically accu- rate analyses of D. ordii growth. The R^ values (Table 1) indicate how much variation is accounted for in the anal- yses, and when converted to r they can be used to determine statistical significance (a = 0.05). A significant r suggests a correla- tion between the appropriate InW (log of the variable) and the age of the growing an- imals when time is partitioned into growth periods. All parameters provided significant corre- lations of growth with age, because r was significant for all growth periods. No indi- vidual parameter consistently provided data with the highest significance throughout all growth periods. The significance levels of all parameters are similar for any given growth period; therefore, it is possible to use any of the parameters as an indicator of growth. Often researchers are more comfortable working with data means (X) than with sta- tistics like InW and k. Because an under- standing of the relationship of InW and X is important in an interpretation of k, means and standard errors of the growth parame- June 1978 Smith et al.: Dipodomys Growth 217 5- 4- 2- Total Length t k Range Total Tail of (days) Length Length Sample Time (mm) (mm) Sizes 1-3 .07158 11471 5657 4 15 .05703 18688 5662 1629 02431 03152 1958 3070 .00285 00303 2538 14 21 28 35 42 49 56 63 70 Age (days) Fig. 1. Means, standard errors (P = 0.95), and growth rates for total length and tail length of Dipodomys ordii reared under standardized laboratory conditions. Growth is expressed in standard measurements described in the text. 218 Great Basin Naturalist Vol. 38, No. 2 Body Weight Ear Length t Time (days) k Range of Sample Sizes Body Weight (gm) Hind Foot Length (mm) Ear Length (mm) 1-3 415 1629 30-70 15987 .07545 .04149 01022 11518 17631 06569 09980 00991 02799 00118 00189 5657 5662 1958 25 38 14 21 28 35 42 49 56 63 70 Age (days) Fig. 2. Means, standard errors (P = 0.95), and growth rates for body weight, ear length, and hind foot length of Dipodomys ordii reared under standardized laboratory conditions. Growth is expressed in standard measurements described in the text. June 1978 Smith et al.: Dipodomys Growth 219 ters are compared to the k values for curves (Figs. 1 and 2). Curves for k and X are al- most identical for all parameters measured, but the confidence intervals of the means for total length and tail length are wider for all growth periods. Since correlations of growth parameters with age were always significant, one might consider using these parameters to predict age. This procedure is important to an eval- uation of the population age structure. Al- though the process seems evident and con- venient, since it would simply involve reading the predicted age from a graph, the results cannot be interpreted with statistical confidence because variation among days is lacking. Calculations of confidence limits about the regression line also present prob- lems; the X axis (age) is a nonrandom vari- able selected by the investigator. Thus, only the regression of Y on X can be estimated with confidence. It is possible that non- parametric procedures could be utilized to provide predictive capabilities (Dapson and Irland 1972). Discussion Ideally, growth data should be collected under field conditions, but the difficulty of such collection for animals below trappable age causes biologists to revert to the labora- tory where organisms can be confined and environmental influences controlled. The growth data analyzed for D. ordii are for animals grown under standardized labora- tory conditions simulating mean temper- atures and photoperiods encountered by an- imals during active periods of growth in western deserts. Although all R^ values were significant (when converted to r) at the P = 0.95 level, one must consider two con- straints in their interpretations: (1) the size of n, which when too large reduces the use- fulness of r, and (2) the percentage of the variation that must be accounted for before significant correlations are considered bio- logically acceptable and k is accepted as a reliable estimate of the instantaneous rela- tive growth rate. When the growth curves (Figs. 1 and 2) are examined, the correla- Table 1. Summary of analyses for growth of Dipodomys ordii pallidus reared under standardized laboratory conditions. Correlation Antilog Age in Days Coefficient Parameter hiA of InW (t=t-l) R^ {r)° Total length 4.09464 74.44 1-3 0.29843 0.54628° 4.15655 149.90 4-15 0.79170 0.88977° 4.65890 210.61 16-29 0.49748 0.70532° 5.27545 237.46 30-70 0.28872 0.53732° Tail length 2.92560 23.31 1-3 0.21841 0.46734° 3.02929 75.18 4-15 0.76737 0.87597° 3.87084 117.92 16-29 0.40309 0.63489° 4.68195 132.95 30-70 0.12449 0.35283° Body weight 1.47177 7.03 1-3 0.25465 0.50462° 1.80782 18.73 4-15 0.64611 0.80380° 2.33647 34.12 16-29 0.44996 0.67079° 3.29270 55.15 30-70 0.60571 0.77827° Ear length 0.45680 2.66 1-3 0.23211 0.48177° 0.71373 9.03 4-15 0.82902 0.91050° 1.79442 13.20 16-29 0.58465 0.76461° 2.48377 13.73 30-70 0.20396 0.45161° Hind foot length 2.40433 16.64 1-3 0.33954 0.58270° 2.61515 36.23 4-15 0.81333 0.90184° 3.37601 38.86 16-29 0.39367 0.62743° 3.59137 39.24 30-70 0.16730 0.40902° 'Significant at a = .05 220 Great Basin Naturalist Vol. 38, No. 2 tions (r) and comparisons of k with the data means and standard errors seem precise within the prescribed growth periods. One is thus inchned to be rather Uberal in set- ting lower limits on R^ and accepting the curves as representing the actual growth of D. ordii for the time intervals under the controlled conditions. It was determined that R^>0.25 should provide enough ac- countability to accept significant correla- tions and to realistically represent the growth rates. This does not mean that the k values for those analyses with R^<0.25 are in error; it means the confidence is not as strong, and caution should be exercised in using these data for actual growth in mod- eling and predictive situations. Total body weight is perhaps the most in- teresting of all parameters measured. It de- termines biomass and relates to secondary production. Because there are such close relationships between the antilogs of InW and means, and because there are such nar- row confidence limits about the means, an- tilogs of InW, means, and k values for body weight should accurately represent the in- stantaneous relative growth rates for periods up to 70 days of age. Therefore, body weight can be used to predict biomass of animals prior to trappable age. Beyond 70 days the correlations of body weight with age become less reliable, but, because the animals are now trappable and have reached adult size, their productivity can be measured in the field. After 70 days, a close correlation is not necessary because k shows very little increase as evidenced by the body weight curves (Fig. 2). Although the growth rates were obtained under standard- ized laboratory conditions, they can be con- sidered generally representative for the growth periods prescribed. These growth periods include times when the animals are undergoing normal active growth. Follow- ing this time period, weight varies in re- sponse to environmental stresses and changes rather than actual growth phenom- ena. One possible weakness of these analyses is an inability to assess k under field condi- tions. Originally, it was assumed that shifts in k imder field conditions would not differ significantly from those established in the laboratory, but analyses of data obtained while experimenting with independent vari- ables (photoperiod, temperature and food) for Peromyscus maniculatus suggest that this assumption may not be valid. Manipu- lation of photoperiod, temperature, and food caused k to vary for growth of P. ma- niculatus, but the k values only varied early in the growth period. By the time the ani- mals reached trappable age k values for body weight, regardless of the independent variable changes, were similar. It is possible that interactions of the variables might compensate for one another and reflect no change in k. If so, estimates of biomass could still be made by correlating body weight with age. There have been several attempts to cor- relate weight with age and other parame- ters (Brody 1945, Dapson and Irland 1972 and others) but, because many of them at- tempted to predict age, the results were not particularly satisfying. Calculation of con- fidence limits about the regression line is not possible because the X axis (age) is a nonrandom variable, selected by the in- vestigator. This study was concerned more with the characterization of growth as far as weight was concerned. Attempts to age organisms should be done with parameters other than body weight, such as dried eye lens weight or perhaps with tyrosine con- tent of lenses (Dapson and Irland 1972). Literature Cited Brody, S. 1945. Time relations of growth of individuals and populations. In: Bioenergetics and Growth. N.Y.: Reinhold, 1023 pp. Chew, R. M., and B. B. Butterworth. 1959. Growth and development of Merriam's kangaroo rat, Dipodomys merriami. Growth 23:75-96. Dapson, R. W., and J. M. Irland. 1972. An accurate method of determining age in small mammals. J. Mamm. 53:100-106. GoERTZ, J. W. 1965. Sex, age, and weight variation in cotton rats. J. Mamm. 46:471-477. Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. New York: Ronald Press, 2 Vol., 1083 pp. Hayden, p., and J. J. Gambino. 1966. Growth and de- velopment of the little pocket mouse, Per- ognathus longimembris. Growth 30:187-197. June 1978 Smith et al.: Dipodomys Growth 221 Horner, B. E., and J. M. Taylor. 1968. Growth and reproductive behavior in the southern grasshop- per mouse. J. Mamm. 49:644-660. Jones, C. 1967. Growth, development, and wing load- ing in the evening bat, Nycticeius humeralis (Rafinesque). J. Mamm., 48:1-19. Lackey, J. A. 1967. Growth and development of Dipo- domys stephensi. J. Mamm. 48:624-632. 1976. Reproduction, growth and development in the Yucatan deer mouse, Peromyscus yucata- nicus. J. Mamm. 57:638-655. Layne, J. A. 1959. Growth and development of the eastern harvest mouse, Reithrodontomys hu- mulus. Bull. Florida State Mus., Biol. Sci. 4(2):61-82. Meyer, B. J., and R. K. Meyer. 1944. Growth and re- production of the cotton rat, Sigmodon hispidus hispidus, under laboratory conditions. J. Mamm. 33:1-20. Morse, E. L., H. D. Smith, and C. D. Jorgensen. A laboratory breeding of Dipodomys ordii. En- cyclia (in press). Pfeiffer, E. W. 1960. Cyclic changes in the morphol- ogy of the vulva and clitoris of Dipodomys. J. Mamm. 41:43-48. PouRNELLE, G. H. 1952. Reproduction and early post- natal development of the cotton mouse, Per- omyscus gossypinus gossypinus. J. Mamm. 33:1-20. RicHiNS, G. H., H. D. Smith, and C. D. Jorgensen. 1974. Growth and development of the western harvest mouse Reithrodontomys megalotis mega- lotis. Great Basin Nat. 34:105-120. SIMILARITY BETWEEN PRONGHORN AND MULE DEER FECAL PELLETS Mark K. Johnson^^ and James G. MacCracken' Abstract.— Botanical compositions and pH values for pronghom (Antelocapra americana) and mule deer {Odo- coileus hemionus) fecal pellets from the Idaho National Engineering Laboratory Site were different. As there was no overlap between ranges of the herbivores' fecal pH values, the fecal pH technique is a valuable tool for dis- tinguishing between fecal pellets of pronghom and mule deer on the study area. Pronghom (Antelocapra americana) and mule deer (Odocoileus hemionus) fecal pel- lets are similar in appearance. On some ranges fecal pH values of pronghom and mule deer did not overlap and it was con- cluded that pH analysis of fecal groups was a legitimate method for distinguishing be- tween the two herbivores for one study area (Howard, J. Wildl. Manage. 31(1): 190-191). Differences in diet and physiology are pos- sible explanations for differences in fecal pH's (Nagy and Gilbert, J. Wildl. Manage. 32(4):961-962). The Idaho National Engineering Labora- tory (INEL) Site occupies about 231,500 ha (894 mi^) of southcentral Idaho and contains a large niunber of pronghom and a small population of mule deer of imknown size. We were studying pronghom food habits using botanical analysis of feces and real- ized that some of our pronghom samples might have been contaminated with those of mule deer. The purpose of this paper is to report our findings as to differences be- tween pH values and botanical composition for mule deer and pronghom fecal pellets from the INEL Site. This research was sup- ported in part by the INEL Ecology Proj- ect, U.S. Department of Energy, under con- tract EY-76-S-07-1526 with Colorado State University. Mule deer pellets were collected in three areas of the INEL Site where deer were lo- cated. Although we did not observe deer depositing pellets which were collected, deer were observed on several occasions in the areas. To our knowledge, no pronghom had been observed in the areas by any per- sons for at least several weeks. Since the pellets collected were fresh, we were con- vinced that they were from mule deer. From each area where deer pellets were collected, they were composited into one sample. From three other portions of the study area herbivore pellets were collected which usually bore closer resemblance to mule deer pellets than to pronghom pellets. Since the identities of these pellets were un- known, they were called imknown Arti- odactyl pellets and were analyzed separate- ly. A composite sample was made for each area sampled. Pronghom pellets were sam- pled from 24 areas of the INEL Site, and a composite sample was made for each area. Pronghom pellets were collected in con- junction with an INEL pronghom ecology study. All pellets used in this study were collected after pronghom were observed de- positing them. Mule deer and unknown Ar- tiodactyl pellets were collected in October 1977. Pronghom pellets were collected dur- ing January, February, and March 1976, July 1976, and July 1977. Fifty pellets were selected from each composite sample and were ground together in a Wiley Mill over a 1.0 mm mesh sieve. 'Department of Range Science, Colorado State University, Fort Collins, Colorado 80521. ■Present Address; Department of Wildlife and Fisheries, P.O. Drawer LW, Mississippi State University, Starkville, Mississippi 39762. 222 June 1978 Johnson, MacCracken: Pellets 223 The botanical composition of each mixture was determined by the method reported by Sparks and Malecheck (J. Range Manage. 21(4):264-265). Similarity in botanical com- positions between samples was estimated us- ing Kulczynski's formula (Costing 1956. The study of plant communities., W. H. Free- man Co. p. 104). One himdred microscope slides were examined for each mixture. Ten different pellets were selected at random from pronghom, mule deer, and unknown Artiodactyl samples for pH analysis. Each pellet was ground in a Wiley Mill over a 1.0 mm mesh sieve and was soaked in 50 ml of deionized water for one hr. The pH was determined with a Sargent- Welch DG recording titrator. Students' tests were used to compare mean pH values among the dif- ferent classes of pellets. Botanical compositions of pronghom and unknown Artiodactyl pellets were about 65 percent similar (Table 1). Botanical compo- sition of mule deer pellets were only about 25 percent similar to pronghom or un- known Artiodactyl fecal pellets. Artemisia, Astragalus, and Sphaeralcea, plus Atriplex, made up more than 70 percent of the plant fragments in pronghom pellets, and Arte- misia and Astragalus, plus Sphaeralcea, made up more than 70 percent of the plant fragments in unknown Artiodactyl pellets. Kochia and Bromus, plus Leptodactylon, made up more than 70 percent of the plant fragments in mule deer pellets. Kochia alone made up more than 50 percent of plant fragments in these pellets. Kochia and Brom.us are common only along roadsides on the INEL Site, where deer had been ob- served feeding on these plants. The average (±SE) pH value for mule deer pellets (9.12 ±0.03: range, 9.05-9.22) was significantly higher (p<0.05) than aver- ages for pronghorn (8.60 ±0.04: range, 8.52-8.72) and unknown Artiodactyl (8.53 ±0.06: range, 8.38-8.70) pellets, which were similar. The range in pH values from the latter two herbivore classes overlapped, but neither overlapped with the range of pH values from mule deer pellets. The range in pH values for pronghom pellets was very narrow, regardless of area or collection date, and range in pH values for mule deer pellets was narrow regardless of area. The major food of pronghom was Artemisia for all collection dates, but Cera- toides made up more than 65 percent of three composite samples. The pH values for these samples were not the highest or the lowest values determined for pronghorn samples. In New Mexico narrow ranges in pH values were found for pronghom and mule deer pellets collected for a whole Table 1. Mean percent (±SE) relative particle densities of plant fragments recovered from pronghom, mule deer, and unidentified Artiodactyl fecal pellets from the Idaho National Engineering Laboratory Site. Unidentified Pronghom Artiodactyl Mule Deer Taxa' No. of Composites Examined = 24 3 3 Artemisia 45.8 ± 1.5 48.9 ± 10.6 18.8 ± 9.3 Astragalus 8.5 ± 3.2 14.8 ± 6.6 2.5 ± 1.1 Sphaeralcea 4.6 ± 2.0 12.2 ± 7.3 3.0 ± 4.7 Atriplex 9.4 ± 4.5 0.7 ± 0.6 Ceratoides 15.0 ± 10.0 0.4 ± 0.4 Kochia 57.8 ± 22.2 Opuntia 4.2 ± 1.8 5.3 ± 3.7 Salsola 6.9 ± 5.1 0.3 ± 0.3 Leptodactylon 1.0 ± 0.9 3.8 ± 4.7 Erigeron 2.7 ± 1.5 1.6 ± 1.0 Bromus 0.4 ± 0.2 10.7 ± 5.6 'Other taxa in herbivore fecal pellets identified in trace (<2 percent) amounts were Sitanion, Koeleria, Spowbolus, Vulpia, Carex, Eriogorum, Crepis, Descurania, Achillea, Lygodesmia, Chrysothamnus, Erysimum. Phlox, Balsamorhiza, Sarcobatus, Convolvulus, CastilUia, Chenopodium, Stipa, Aristida, AgTopymn, Oryzopsis, Saliz, moss, unclassified grasses, unclassified forbs and arthropods. 224 Great Basin Naturalist Vol. 38, No. 2 year, and there was no overlap in the ranges of pH values found for each species (Howard, J. Wildl. Manage. 31(1): 190-191). Evidence suggests that seasonal differences in diets have an insignificant influence on fecal pH's. Differences in fecal pH's be- tween species are probably due to phys- iological differences rather than dietary dif- ferences. Under average circumstances fecal pH's of pronghom and mule deer probably remain within narrow ranges. The similarity in pH values and botanical compositions of imknown Artiodactyl pellets with those of pronghom suggests that the unknown fecal pellets were deposited by pronghom. We conclude that visual exam- ination of Artiodactyl pellets is inadequate for identifying the animal of their origin. However, fecal pH values of pronghom and mule deer on the INEL Site appear to be different. Since mule deer pellets that we collected were from a few areas and repre- sented a limited portion of the year, we recommend further study to corroborate our findings before the fecal pH technique is employed in practical application on the INEL Site. NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be di- rected to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be sim- ilarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuseripts. Two copies of manuscripts are required. They should be typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. Use a recent issue of either journal as a format, and the Couneil of Biology Edi- tors Stifle Manual, Third Edition (AIBS 1972) in preparing the manuscript. An abstract, about 3 percent as long as the text, but not exceeding 200 words, written in accordance with Biological Abstracts guidelines, should precede the introductory paragraph of each ar- ticle. Authors may recommend one or two reviewers for their article. All manuscripts re- ceive a critical peer review by specialists in the subject area of the manuscript under con- sideration. Manuscripts that are accepted and that are less than 100 printed pages in length will automatically be assigned to the Great Basin Naturalist. Those manuscripts larger than 100 printed pages in length will be considered for the Memoirs series. Illustrations and Tables. All illustrations and tables should be made with a view toward having them appear within the limits of the printed page. Illustrations that form part of an article should accompany the manuscript. Illustrations should be prepared for reduction by the printer to either a single-column (2 5/8 inches) or double-column (5 1/2 inches) width, with the length not exceeding 7 1/2 inches. Costs Borne hij Contributor. Contributors to the Great Basin Naturalist should be pre- pared to donate from $10 to $30 per printed page toward publication of their article (in addition to reprint costs outlined in the schedule below). Authors publishing in the Great Basin Naturalist Memoirs may be expected to contribute $35 per printed page in addition to the cost of printed copies they purchase. No printed copies are furnished free of charge. .\ price list for reprints and an order form are .sent with the galley proof to contributors. Reprint Schedule of the Great Basin Naturalist 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each 100 copies $20 $24 $28 $32 $36 $40 additional 200 copies 28 32 36 40 44 48 2 pp. 300 copies 36 40 44 48 52 56 $4 Great Basin Naturalist Memoirs No. 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodbury, H. H. Frost. $10. No. 2 Intermountain biogeography: A symposium. Bv K. T. Harper, J. L. Reveal, et al. $15. TABLE OF CONTENTS Kaiparowits flora. Stanley L. Welsh, N. Duane Atwood, and Joseph R. Murdock.... 125 Range extension of Onychomys torridus longicaudus (Rodentia: Cricetidae) in northwestern Nevada. Mark D. Engstrom and Eugene D. Fleharty 180 The reproductive ecology of the Tahoe sucker, Catostomus tahoensis, in Pyramid Lake, Nevada. Joseph L. Kennedy and Paul A. Kucera 181 Primary productivity in meromictic Big Soda Lake, Nevada. R. P. Axler, R. M. Gersberg, and L. J. Paulson 187 Current status of cutthroat trout subspecies in the western Bonneville Basin. Terry J. Hickman and Donald A. Duff 193 Reproductive biology of the tui chub, Gila bicolor, in Pyramid Lake, Nevada. Paul A. Kucera .'. 203 Distributional records for the Colorado flora. D. Wilken, W. Harmon, C. Fed- dema, and H. Harrington 208 A comparative analysis of the diatom flora on the snail Ampullaria cuprina from the Goshen Ponds, Utah. Larry L. St. Clair, Lorin E. Squires, and Samuel H. Rushforth 211 Growth of Dipodomys ordii (Rodentia: Heteromydiae). H. Duane Smith, Gary H. Richins, and Clive D. Jorgensen 215 Similarity between pronghom and mule deer fecal pellets. Mark K. Johnson and James G. MacCracken 222 rHE GREAT BASIN NATURALIST Volume 38 No. 3 September 30, 1978 Brigham Young University ^L <^^^Mv ^^k MP. zoou JAN 19 1979 ■..3 GREAT BASIN NATURALIST MEMOIRS Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agri- cultural Sciences; Ernest L. Olson, Director, Brigham Young University Press, Univer- sity Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only previously unpublished manuscripts of less than 100 printed pages in length and pertaining to the biological natural history of western North America are ac- cepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manu- scripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist is $12 (outside the United States $13). The price for single numbers is $4 each. All back numbers are in print and are available for sale. All matters pertaining to the purchase of subscriptions and back numbers should be directed to Brigham Young University, Life Science Museum, Pro- vo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same of- fice at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts and other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back cover. 11-78 650 35893 ISSN 0017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 38 September 30, 1978 No. 3 UTAH FLORA: FABACEAE (LEGUMINOSAE) Stanley L. Welsh' Abstract.— A revision of the legume family, Fabaceae (Leguminosae), is presented for the state of Utah. In- cluded are 244 species and 60 varieties of indigenous and introduced plants. A key to genera and species is pro- vided, along with detailed descriptions, distributional data, and pertinent comments. Proposed new taxa are As- tragalus lentiginosus Dougl. ex Hook, var. wahweapensis Welsh; Astragalus subcinereus A. Gray var. basalticus Welsh; Hedysarum occidentale Greene var. canone Welsh; Oxytropis oreophila A. Gray var. juniperina Welsh; and Trifolium andersonii A. Gray var. friscanum Welsh. New combinations include Astragalus bisulcatus (Hook.) A. Gray var. major (M. E. Jones) Welsh; Astragalus consobrinus (Bameby) Welsh; Astragalus pubentissimus Torr & Gray var. peabodianus (M. E. Jones) Welsh; Lathyrus brachycalyx Rydb. var. zionis (C. L. Hitchc.) Welsh; Lathyrus lanzwertii Kellogg var. arizonicus (Britton) Welsh; Lathyrus langwertii Kellogg var. laetivirens (Greene) Welsh; Lupinus argenteus Pursh var. moabensis (Dunn & Harmon) Welsh; Lupinus argenteus Pursh var. ru- bricaulis (Greene) Welsh; Lupinus caudatus Kellogg var. argophyllus (A. Gray) Welsh; Lupinus caudatus Kellogg var. cutleri (Eastw.) Welsh; Lupinus pusillus Pursh var. rubens (Rydb.) Welsh; Lupinus sericeus Pursh var. barbi- ger (S. Wats.) Welsh; Lupinus sericeus Pursh var. marianus (Rydb.) Welsh; Oxytropis besseyi (Rydb.) Blankinship var. obnapiformis (C. L. Porter) Welsh; Psoralea lanceolate Pursh var. stenostachys (Rydb.) Welsh. This paper is the second in a series of works leading to a definitive treatment of the flora of Utah. The first paper dealt with the Brassicaceae (Great Basin Nat. 37:279-365. 1977). The legume family, with 304 taxa, is one of the largest present in the state, apparently ranked second after the Asteraceae, but in third place following the Poaceae if only the 244 specific taxa are counted. Adventive plants account for 45 species, or about 15 percent of the total taxa in the family. This is fewer than the 18 percent determined for the Brassicaceae. Weedy species represent only a small pro- portion of the adventive taxa; the bulk con- sists of cultivated crop and ornamental plants. More than half of the 45 legume genera in Utah are known from introduced plants. The 24 introduced genera account for only 30 species, however. It seems un- likely that other large families will be rep- resented by such a large percentage of ad- ventive genera. The remainder of the introduced species are in genera with in- digenous species in Utah. The largest genus of legumes, and prob- ably of any plant family in Utah, is Astra- galus with 110 species and 36 varieties. This genus has long proved troublesome to taxonomists, partly because of its great size, and partly because of the complex nature of some species. The definitive treatment by Dr. Rupert C. Bameby provides a basis for under- standing of Astragalus, and this treatment is dedicated to him. Generally, the specific and infraspecific lines are rather sharply drawn, but in the Astragalus lentiginosus complex taxa apparently grade through series of morphological intermediates. Some 'Department of Botany and Range Science, and Life Science Museum, Brigham Young University, Provo, Utah 84602. 225 226 Great Basin Naturalist Vol. 38, No. 3 30 taxa within Astragalus, almost a quarter of the total, are endemic to Utah often on highly specialized soil types. The currently proposed lists of endangered or threatened plants are replete with Astragalus taxa. It is difficult to find a portion of Utah where one or more species of Lupinus are not present. Annual species are more com- mon at lower elevations, but some occur at middle and even upper elevations. Specific lines are more easily drawn among annual species. Perennials occur as complex groups which tend to intergrade in endless en- tanglements. Because of these problems, species lines have been subject to inter- pretation, and only the careful work of Dr. David Dunn and his students has allowed meaningful interpretation of Utah lupine materials. Oxytropis is less complex in Utah than elsewhere, primarily due to the low number of species and to the geographical isolation of taxa known to hybridize elsewhere. Only O. oreophila exhibits a wide spectrum of variation in the state. Trifolium consists of both indigenous and introduced species which are markedly dis- tinct, both morphologically and geographi- cally. The Old World introductions listed are among our most important forage plants. The legume family is economically im- portant because of utilization of plants as food for man and animals. Some representa- tives are poisonous and have harmed live- stock in the state. These are noted within the text. Perhaps the most economically important crop derived from this family is alfalfa or lucern (Medicago sativa). Thousands of acres are planted to this forage crop in Utah. The writer follows the example found in Barneby (1964, Atlas of North American As- tragalus) in citing the number of specimens of each species examined. The numbers fol- low the discussion of each taxon, with the Arabic numerals indicating the total number seen and the Roman numerals the number collected by me. Fabaceae Lindley Legume Family Herbs, shrubs, or trees; leaves alternate, pinnately or palmately compound, or simple, stipulate; flowers perfect, irregular or regular, usually borne in racemes; calyx 5-lobed; petals 5 (a banner, 2 wings, and 2 keels) or fewer, less commonly reduced to 1 (banner), or lacking; stamens 10 or 5, or nu- merous, diadelphous, monadelphous, or dis- tinct; pistil 1, the ovary superior, 1- or 2- loculed, 1-carpelled, the style and stigma 1; fruit (pod) a legume or loment, sessile, sub- sessile, stipitate, or with a gynophore, dehis- cent or indehiscent. References IsELY, D. 1973. Leguminosae of the United States: I. Mimosoideae. Mem. N.Y. Bot. Card. 25(1): 1-52. 1975. Leguminosae of the United States: IL Caesalpinoideae. Mem. N.Y. Bot. Card. 25(2): 1-228. Welsh, S. L. 1963. Legumes of Utah L Preliminary report. Proc. Utah Acad. 40: 200-201. 1964. Legumes of Utah IL Con- spectus of the genera. Proc. Utah Acad. 41: 84-86. 1. Flowers regular, in dense heads or compact spicate racemes; stamens numer- ous (Mimosoideae) Key I Flowers irregular (only slightly so in some); stamens 10 or fewer 2 2(1). Corolla not papilionaceous, sometimes nearly regular, the upper petal en- closed by the others; stamens 10 or fewer, commonly distinct (Caesalpi- noideae) Key II - Corolla papilionaceous, the upper petal (banner) enclosing the wing and keel petals in bud, much reduced in Amorpha, or lacking in Dalea, Psorothamnus, and Parryella; stamens 10 or 5 (Papilionideae) ." 3 Sept. 1978 Welsh: Utah Flora, Fabaceae 227 3(2). Plants woody; trees, shrubs, or woody vines Key III — Plants herbaceous perennials or annuals 4 4(3). Leaves even-pinnate Key IV — Leaves odd-pinnate or simple 5 5(4). Leaflets 3 only Key V — Leaflets 5 or more, or the leaves simple Key VI Key I Flowers regular: stamens numerous (Mimosoideae) L Trees, unarmed, cultivated; flowers in umbellate heads; stamens long- exserted, the filaments commonly 20-30 mm long Albizia — Trees or shrubs, armed, indigenous; flowers in spicate racemes; stamens in- cluded or shortly exserted, the filaments less than 5 mm long 2 2(1). Spines recurved; pods flat, 10-20 mm broad, brown at maturity Acacia (greggit) — Spines straight; pods spirally coiled or if flattened then less than 10 mm broad and yellowish to tan at maturity Prosopis Key II Corolla not papilionaceous (Caesalpinoideae) 1. Plants herbaceous perennials; flowers yellow; indigenous to southeastern Utah Hoffmanseggia — Plants trees or shrubs; flowers variously colored; indigenous or introduced 2 2(1). Leaves simple, the blades rotund-ovate; flowers pink, appearing before the leaves Cersis — Leaves once or twice compound; flowers yellow, white, or greenish, appear- ing after the leaves 3 3(2). Shrubs; flowers with yellow petals, the exserted stamens to 50 mm long or more; cultivated and naturalized in Washington Co Caesalpinia — Trees; flowers with white or greenish-yellow petals, the stamens included or not much exserted; distribution broad 4 4(3). Leaves once to twice pinnate; branches often armed; flowers greenish-yellow, borne in spicate racemes; pods long and strap-shaped Gleditsia — Leaves bipinnate; branches unarmed; flower white, long-stalked, in open panicles; pods thick Gymnocladus Key III Trees, shrubs, or woody vines (Papilionideae) 1. Leaves even-pinnate, the rachis produced apically as a bristle; flowers yel- low Caragana — Leaves simple or odd-pinnate; flowers variously colored 2 2(1). Leaves simple or the lower ones 3-foliolate; plants shrubs; flowers yellow; solitary or borne in erect racemes 3 — Leaves compound; plants varying in one or more ways from above 4 228 Great Basin Naturalist Vol. 38, No. 3 3(2). Calyx split above hence 1-lipped, with 5 minute teeth; flowers borne in erect racemes; Washington Co Spartium — Calyx bilabiate, the upper lip 2-lobed, the lower 3-lobed; flowers mostly solitary axillary; Weber Co Cytissus 4(2). Plants twining woody vines; flowers large and showy, borne in terminal, pendulous racemes Wisteria — Plants trees or shrubs; flowers various, usually borne in axillary, erect or pendulous racemes 5 5(4). Leaflets 3; flowers yellow, borne in pendvilous racemes Laburnum — Leaflets 5 or more; flower white, pink, indigo, or yellow, borne in erect or spreading racemes 6 6(4). Herbage glandular-punctate; indigenous shrubs with petals indigo or lacking . 7 — Herbage not glandular-punctate; cultivated or indigenous shrubs or trees; petals white, pink, or yellow, or if indigo (as in Amorpha) then the corolla reduced to a single petal (the banner) 8 7(6). Petals lacking; leaflets linear; plants of Grand and San Juan Co Parnjella — Petals present; leaflets broad; plants of southern and southeastern Utah Psorothamnus 8(6). Petals 1, the banner only present, indigo; sparingly cultivated shrubs Amorpha — Petals 5, white, pink, or yellow; shrubs or trees, sparingly to commonly cul- tivated 9 9(8). Plants shrubs; pods bladdery-inflated; flowers yellov/; ornamental and road- side plants Colutea — Plants trees or shrubs; pods flat or terete; flowers white or pink 10 10(9). Branches armed with stipular spines or internodal hispid processes; staminal filaments diadelphous; petals pink or white Robinia — Branches unarmed; staminal filaments distinct; petals white 11 11(10). Leaf bases hollow, covering superposed buds; pods flat, not constricted be- tween the seeds Cladrastis — Leaf bases solid, not covering buds; pods terete, constricted between the seeds Sophora Key IV Leaves even-pinnate (Papilionideae) 1. Flowers yellow; fruit ripening below ground, tardily dehiscent, constricted between the seeds Arachis - Flowers white, pink, red, lavender, or cream; fruit borne above ground, not constricted between the seeds 2 2(1). Style strongly dilated; sepals foliaceous; plants cultivated Pisum - Style not strongly dilated; sepals not foliaceous; plants indigenous or culti- vated 3 Sept. 1978 Welsh: Utah Flora, Fabaceae 229 3(2). Style bearded down one side; wings of corolla essentially free from the keel . Lathyrus — Style bearded in a tuft or ring at apex; wings of corolla adherent to the keel Vicia Key V Leaflets three (Papilionideae) 1. Leaves palmate, the terminal leaflet neither stalked nor jointed 2 — Leaves pinnate, the terminal leaflet stalked or jointed 4 2(1). Flowers golden-yellow, the banner orbicular, large; legumes narrowly ob- long, erect or ascending; staminal filaments distinct Thermopsis — Flowers ochroleucous to white or pink to pink-purple, the banner not or- bicular, moderate to small in size; staminal filaments diadelphous 3 3(2). Leaflets usually toothed; flowers mostly in heads, commonly pink or white Trifolium — Leaflets entire; flowers not in heads, commonly ochroleucous or pink Astragalus 4(1). Herbage glandular-punctate; indigenous plants with usually linear to oblan- ceolate leaflets Psoralea — Herbage not glandular-punctate; indigenous or cultivated plants with spatu- late to obovate or oblanceolate to ovate leaflets 5 5(4). Leaflets entire 6 — Leaflets toothed (except in some Trifolium species) 8 6(5). Flowers in umbels, loosely capitate, or solitary in leaf axils, yellow or suf- fused with orange Lotus — Flowers in interrupted racemes or panicles, purplish 7 7(6). Leaflets stipellate; pods several seeded, several to many times longer than broad Phaseolus — Leaflets lacking stipels; pods 1-seeded, only somewhat longer than broad Lespedeza 8(5). Flowers usually in heads; corolla persistent, investing the fruit; fruit straight Trifolium — Flowers usually in racemes; corolla not persistent; fruit straight or curved to coiled 9 9(8). Leaflets toothed along the distal V2 or more; racemes elongate, several times longer than broad Melilotus — Leaflets toothed along the distal V3 only (except in some Trigonella); racemes compact or loose, seldom more than twice longer than broad 10 10(9). Fruit straight or falcately curved, prominently veined on the valves; flowers yellow; terminal leaflet with an apical spinose cusp, rarely as much as twice longer than broad; plants rare Trigonella — Fruit coiled or curved, veined or not; flowers pink, lavender, whitish, or yel- low; terminal leaflet seldom strongly cuspidate apically, usually more than twice longer than broad; plants common Medicago 230 Great Basin Naturalist Vol. 38, No. 3 Key VI Leaflets (4) 5 or more, or simple (Papilionideae) 1. Leaves palmately compound, with usually 5-11 leaflets, long-petiolate 2 — Leaves pinnately compound, or if rarely palmately compound (as in some Lotus species), then sessile or with only 4 leaflets 3 2(1). Herbage glandular-dotted; leaflets usually 5, broadly obovate-spatulate; sta- mens usually diadelphous; pods 1-seeded Psoralea — Herbage not glandular-dotted; leaflets usually 7-11, variously shaped; sta- mens monadelphous; pods several-seeded Litpinus 3(1). Herbage glandular-dotted 4 — Herbage not glandular-dotted 5 4(3). Racemes spicate; legumes 1-seeded, not bearing appendages; stamens 5; pet- als (except banner) inserted on staminal tube Dalea — Racemes not spicate; legumes several-seeded, bearing hooked appendages; stamens 10; petals not inserted on staminal tube Glycyrrhiza 5(3). Terminal leaflet of lower leaves several times larger than the lateral; in- florescence of many-flowered head closely subtended by foliose involucral bracts; flowers yellow; introduced, rare Anthyllis — Terminal leaflet not much larger than the lateral; inflorescence a raceme or an umbel, lacking foliose bracts (except in Lotus); flower color various 6 6(5). Margin of leaflets toothed; corolla persistent, investing the fruit Trifolium — Margin of leaflets entire; corolla usually deciduous 7 7(6). Flowers in umbels, loosely capitate, or solitary in leaf axils; petals yellow, often suffused with orange, or pink 8 — Flowers in racemes or cymes; petals usually not yellow 9 8(7). Leaflets 3-5; flowers yellow Lotus — Leaflets 9-23; flowers pink to pink-purple Cownilla 9(7). Keel petals much longer than the wings; fruit a flattened loment 10 — Keel petals subequal to the wings or shorter; fruit a legume (a terete loment in Sophora) 11 10(9). Fruit 4- to several-seeded, not spiny (except in H. boreale var. gremiale); plants indigenous Hedijsarum — Fruit 1- to 2-seeded, more or less spiny-toothed; plants adventive, cultivated and escaping Onobrychis 11(9). Stipules spiny; flowers dirty whitish Peteria — Stipules various, but not spiny; flowers seldom if ever dirty whitish 12 12(11). Staminal filaments distinct; fruit a terete to somewhat flattened loment; plants with blue or white flowers, usually of sandy sites Sophora — Staminal filaments diadelphous or monadelphous; fruit a legume; plants from a caudex and/or taproot, rarely rhizomatous; habitats various 13 13(12). Keel with a porrect beak; ventral suture of legume forming a partial or complete partition; plants usually acaulescent Oxytropis Sept. 1978 Welsh: Utah Flora, Fab ace ae 231 — Keel beakless, or the beak diverging from the floral axis; ventral suture usu- ally not produced internally, the dorsal usually produced in bilocular fruits; plants usually caulescent 14 14(13). Stamens monadelphous; flowers blue Galega — Stamens diadelphous; flowers pink-purple, pink, lavender, ochroleucous, red, white, or variously suffused, but not blue 15 15(14). Flowers red-orange when fresh; plants adventive Sphaerophysa — Flowers pink, pink-purple, lavender, or white to ochroleucous; plants in- digenous, or rarely adventive Astragalus Acacia P. Miller Armed trees; leaves alternate, often clus- tered on short axillary shoots, bipinnate, pe- tiolate, the pinnae bearing several leaflets; internodal spines curved; stipules small and deciduous; flowers numerous, borne in elon- gate spikes; calyx 5-lobed; corolla regular, 5-lobed, inconspicuous; stamens numerous, included, distinct; ovary substipitate; pods flattened, indehiscent. Acacia greggii A. Gray. Catclaw Acacia. Small trees to 4 m tall, the branches armed with curved internodal spines; leaves to about 4 cm long, with 2 pairs of pinnae, each with 4-6 pairs of obovate to oblong leaflets 3-6 mm long, puberulent on both surfaces; petioles 2-5 mm long, bearing a solitary gland between the lower pair of pinnae; spikes mostly 3-6 cm long (includ- ing peduncles); flowers fragrant, 2-2.5 mm long; petals greenish, like the sepals; le- gumes flattened, oblong, usually curved, 50-100 mm long, 10-20 mm wide, con- stricted between the seeds; seeds 5-7 mm broad, nearly circular. Warm desert shrub, drainage-terrace vegetation, at about 870 m in Washington Co.; Nevada, California, Ari- zona, New Mexico, Texas, and Mexico. Our material belongs to var. arizonica Isely. Albizia Durazzini Unarmed trees; leaves alternate, not clus- tered, bipinnate, petiolate, the several pairs of pinnae each with numerous oblique leaf- lets; stipules small and caducous; flowers several to many, in umbellate heads; calyx tubular, 5-lobed; corolla vmited, funnelform, the 5 lobes shorter than the tube; stamens numerous, united into a tube basally, long- exserted; pods flattened, dehiscent. Albizia julibrissin Durazzini. Silk-tree, Mimosa. Small tree to 3 m tall or more and as broad or broader; leaves to 25 cm long or more (including petiole), with 5-10 (15) pairs of pinnae, each with 12-25 (30) pairs of leaflets 7-15 mm long, puberulent, if at all, on rachis and leaflet margins; petioles 3-6 cm long, each with a single large flat- tened gland; calyx 3-3.5 mm long; corolla 7.5-9.5 mm long, cream to greenish; sta- minal filaments exserted 20-30 mm, brightly rose-pink to reddish in color; pods 120-200 mm long, 15-25 mm wide, oblong, flat- tened, membranous. Cultivated ornamental at lower elevations in much of Utah, but frost sensitive; introduced from Asia. Amorpha L. Cultivated shrubs; leaves alternate, odd- pinnate, the leaflets marked with dots, usu- ally with stipels; flowers purple, borne in terminal spicate racemes; calyx 5-toothed, persistent; banner present (wings and keel lacking), wrapped around the stamens and style; stamens 10, monadelphous at the base only, otherwise distinct; pods 1- to 2-seeded, tardily dehiscent. 1. Plants usually less than 1 m tall; petioles short, usually shorter than width of lowest leaflets A. canescens Plants usually more than 1 m tall; petioles elongate, longer than the width of the lowest leaflets A. fruticosa 232 Great Basin Naturalist Vol. 38, No. 3 Amorpha canescens Pursh. Lead Plant. Subshnib, the erect branches, mostly 4-10 dm tall, the herbage densely white-villous; leaves subsessile, 2.5-12 cm long, with 15-51 leaflets, these elliptic to lance-elliptic or oblong, green above, white-hairy be- neath; racemes clustered, paniculate; calyx tube white-villous; pods white-villous, the style almost as "long as the body. Cultivated ornamental in some communities in north- ern Utah (Reimschiissel s.n., BRY); in- troduced from the Great Plains; indigenous from Canada south to Texas and New Mexi- co. Amorpha fruticosa L. False Indigo, Bas- tard Indigo. Shrub to 3 m tall or more, the herbage sparingly pubescent to glabrate; leaves long-petioled, with 13-35 leaflets, these elliptic to oblong, green on both sides, the lower only somewhat paler and strigu- lose; racemes clustered, paniculate; calyx tube glabrous; pods glabrous, the style much shorter than the body. Cultivated ornamen- tal and botanical curiosity in northern Utah (Vickery 1085, 2252, UT); introduced from eastern United States; indigenous in much of eastern North America and south- westward to Arizona. Anthyllis L. Cultivated herbaceous perennial; leaves odd-pinnate; stipules small, adnate the pet- iole, the lowermost somewhat sheathing; flowers many, borne in pedunculate heads or head-like clusters; calyx tubular, 5-lobed; corolla papilionaceous; stamens 10, mon- adelphous; pods invested by the accrescent calyx, 1- or few-seeded. Anthyllis vulneraria L. Kidney Vetch, Woundwort. Stems arising from a caudex, 8-30 cm tall, decumbent to erect; leaves 2-7 cm long, odd-pinnate, with usually 5-9 leaflets, the terminal leaflet of lowermost leaves much larger than the lateral ones; peduncles 5-16 cm long, usually with a fo- liose bract below the inflorescence; heads 1 to few, each closely subtended by foliose bracts; flowers 10-15 mm long, sessile, yel- low (or suffused with red); calyx pilose, much inflated at maturity. Introduced for- age and reseeding plant, known from San- pete Co. (Stevens 364, BRY), but to be ex- pected elsewhere; indigenous to Eurasia. Arachis L. Cultivated annual herbs; leaves even- pinnate, lacking tentrils; stipules prominent, long-attenuate, adnate to the petiole and al- most sheathing the stem; flowers yellow, pa- pilionaceous, few or solitary in the axils, sessile, hypanthium elongating and pushing developing ovary underground; stamens di- adelphous, usually 9 and 1; pods maturing underground, indehiscent, constricted be- tween the seeds. Arachis hypogaea L. Peanut. Stems from a taproot, mostly 20-50 cm tall; leaves mostly 4-15 cm long, even-pinnate, with 2 pairs of leaflets, 2.2-6 cm long, 0.8-2.5 cm wide, entire; stipules 20-35 mm long; flow- ers yellow, soon withering, usually only the lowermost producing fruits. Sparingly culti- vated plants, rarely escaping but not per- sisting, in Utah; introduced from Brazil (?). Astragalus L. Plants annual or perennial, caulescent or acaulescent, from a taproot, a caudex com- monly developed, rarely with a rhizome; leaves alternate, odd-pinnate, trifoliolate, or simple; stipules adnate to the petiole base, sometimes connate-sheathing aroimd the stem; flowers papilionaceous, in axillary racemes, each subtended by a single bract; bracteoles 1 or 2 or lacking, attached at base of calyx or on pedicel; calyx 5-toothed; petals 5, pink, lavender, pink-purple, orcho- leucous, or white, or variously suffused, the keel shorter than the wings, rounded to at- tenuate apically; stamens diadelphous; ovary enclosed in the staminal sheath, the style glabrous; pods variable in size, shape, and dehiscence, unilocular to bilocular, sessile, subsessile, or stipitate (or with a gyno- phore). Note: This is a large and complex genus, certainly the largest genus of flower- ing plants in Utah, and because of this, the keys to species are constructed as to reflect political geographic subdivisions of the state. This makes it possible to identify un- known plants without the effort of strug- gling through a single interminably long key. Keys and descriptions are based on some 3800 specimens from Utah, a quarter of them collected by the author. Sept. 1978 Welsh: Utah Flora, Fabaceae 233 References ican species of Astragalus. By the Au- Barneby, R. C. 1964. Atlas of North Ameri- *«^' ^^^^ ^^^^ ^''y^ ^'^^- ^^^ PP" can Astragalus. Mem. N.Y. Bot. Card. Rydberg, P. A. 1929. Astragalanae. North 13: 1-199. American Flora 24: 251-462. Jones, M. E. 1923. Revision of North Amer- 1. Plants of northwestern Utah (i.e.. Box Elder, Cache, Davis, Juab, Morgan, Salt Lake, Tooele, Utah, and Weber counties) Key I Plants not from northwestern Utah 2 2(1). Plants of south-central and southwestern Utah (i.e., Beaver, Iron, Millard, Piute, Sanpete and Sevier counties) Key II — Plants not from south-central and southwestern Utah 3 3(2). Plants of Washington County Key III — Plants not of Washington County 4 4(3). Plants of northeastern Utah (i.e., Daggett, Duchesne, Rich, Summit, Uintah, and Wasatch counties) Key IV — Plants of east-central and southeastern Utah 5 5(4). Plants of Carbon, Emery, and Wayne counties Key V — Plants of Garfield, Grand, Kane, and San Juan counties 6 6(5). Plants of Grand and San Juan counties Key VI — Plants of Garfield and Kane counties Key VII Key I Plants of Box Elder, Cache, Davis, Juab, Morgan, Salt Lake, Tooele, and Weber counties 1. Plants prostrate; leaves to 1 cm long, with spinulose-tipped leaflets all de- current; usually of high elevations A. kentrophyta — Plants of various habit; leaves often more than 1 cm long, not or rarely spinose, the lateral ones usually jointed to the rachis; distribution various 2 2(1). Leaves all simple; plants strictly acaulescent; dwarf, mat-forming plants of high elevations in Cache Co A- spatulatus — Leaves with (3) 5-25 leaflets or more, or only the uppermost simple; plants caulescent or acaulescent, of low to high elevations 3 3(2). Plants rush-like or sprawling, with slender leaflets, or the uppermost leaves often simple, the terminal leaflet confluent with the rachis 4 — Plants not rush-like, the leaflets various but usually not slender and with the uppermost leaves simple, the terminal leaflet jointed to the rachis 8 4(3). Flowers 15-19 mm long; stems arising from a superficial woody caudex 5 — Flowers less than 10 mm long; stems arising from a slender, sub- rhizomatous, subterranean caudex 6 5(4). Flowers ochroleucous; calyx brown; legumes long-stipitate, pendulous A. lonchocarpus 234 Great Basin Naturalist Vol. 38, No. 3 — Flowers bicolored, pink-purple with white wing-tips; calyx purplish; legumes sessile, erect A. toanus 6(4). Pods bladdery-inflated; pubescence of malpighian (dolabriform) hairs; stems usually sprawling A. ceramicus — Pods oblong, not inflated; pubescence of basifixed hairs; stems erect 7 7(6). Leaflets and leaf rachis commonly expanded into flat, grass-like blades; ovules 10-16; pods mostly 10-17 mm long, 3-3.7 mm broad; plants of moist meadows and streambanks, rare A. diversifolius — Leaflets and leaf rachis usually very narrow; ovules more than 17; pods 25-45 mm long, 2.5-3.3 mm broad; plants of dry hillsides, common A. convallarius 8(3). Calyx tube less than 4 mm long, campanula te or short-cylindric 9 — Calyx tube more than 4 mm long, cylindric to long-cylindric 18 9(8). Plants acaulescent, with a distinctive thatch of persistent leaf bases; pods bladdery inflated; dwarf, at high elevations in western ranges, rare A. platytropis — Plants short- to long-caulescent, not with marcescent leaf bases; pods vari- ous; of low to high elevations 10 10(9). Plants annual, from slender taproots, usually of low elevation arid sites 11 — Plants perennial, with well-developed caudices; commonly montane 12 11(10). Pods bladdery-inflated, unilocular; flowers 3 or more per raceme; plants of sandy sites A. geyeri — Pods curved-oblong, bilocular or nearly so; flowers often 1 or 2 per raceme; plants of various soils A. nuttallianus 12(10). Stipules all distinct, not connate-sheathing around the stem 13 — Stipules connate-sheathing around the stem, at least the lowermost 14 13(12). Pods bladdery-inflated, bilocular; leaflets oblanceolate or broader; flowers pale A. lentiginosus — Pods oblong, not inflated, unilocular; leaflets narrowly oblong; flowers dirty purplish A. pinonis 14(12). Pubescence malpighian; pods falcately curved, bilocular A. falcatus — Pubescence basifixed; pods straight, unilocular 15 15(14). Flowers ochroleucous; pods stipitate, either laterally flattened or bisulcate; inflorescence several times longer than broad 16 — Flowers usually some shade of pink or lavender, sometimes as above; pods stipitate or sessile, neither strongly flattened nor bisulcate 17 16(15). Stipules turning black on drying; flowers usually 15 or fewer; pods strongly laterally compressed A. tenellus — Stipules not turning black on drying; flowers usually many more than 15; pods bisulcate A. bisulcatus 17(15). Keel with a prominent upturned beak; pods and ovaries sessile, laterally compressed; plants common A. miser Sept. 1978 Welsh: Utah Flora, Fabaceae 235 — Keel merely rounded apically; pods and ovaries stipitate, 3-angled; plants rare A. alpinus 18(8). Plants acaulescent or subacaulescent, the internodes seldom apparent, the stems then prostrate; herbage usually grayish-hairy 19 — Plants caulescent, the internodes not obscured by leaf bases and stipules (subacaulescent in A. megacarpus); stems usually erect or ascending; herbage commonly green 26 19(18). Wing tips bilobed; plants with malpighian hairs; flowers commonly 12 mm long or less A. calycosus — Wing tips entire; pubescence basifixed; flowers usually 15 mm long or more 20 20(19). Plants strictly acaulescent, clothed below with a persistent thatch of leaf bases and stipules 21 — Plants either not strictly acaulescent or the thatch not, or only poorly, developed 22 21(20). Flowers ochroleucous, the keel purple-tipped; pods thinly long-pilose, the valves apparent through the pubescence A. eurekensis — Flowers pink-purple throughout; pods densely woolly-villous, the valves obscured by hairs A. newberryi 22(20). Leaves very densely hirsute-tomentose, with the longer hairs straight and spirally twisted; pods bilocular, densely woolly-hairy A. mollissimus — Leaves variously pubescent, but if densely tomentose, the hairs all fine, sinous, and cottony, none straight and spirally twisted 23 23(22). Pubescence of leaves, and commonly of entire plant, softly villous-tomentose, consisting of fine, cottony, contorted or entangled hairs; pods both villous- tomentose and hirsute 24 — Pubescence of leaves various but not of extremely fine entangled hairs; pods merely strigose, or both villous-hirsute and tomentose 25 Leaflets mostly obovate and obtuse; flowers bright pink-purple A. utahensis Leaflets various, but where the range of this and the preceding overlap (in Box Elder County), either elhptic or the petals whitish A. purshii 25(23). Pods hirsute and tomentose, the valves obscured by the long hairs; plants uncommon in northwestern Utah A. marianus — Pods strigillose to strigose, the valves not obscured by the short hairs; plants common in northwestern Utah A. argophyllus 26(18). Plants subacaulescent; inflorescences with 1-4 pink-purple flowers, soon sur- passed by the leaves; pods unilocular, 25-60 mm long, bladdery-inflated A. megacarpus — Plants caulescent; inflorescences with 5 to many ochroleucous to pink-purple flowers, often surpassing the leaves; pods uni- or bilocular, often less than 25 mm long, or flowers not pink-purple, bladdery-inflated or not 27 27(26). Stems arising from slender, rhizome-like caudex branches; flowers subsessile in head-like racemes, erect or ascending; pods erect, long-pilose, less than 12 mm long A. agrestis 236 Great Basin Naturalist Vol. 38, No. 3 — Stems arising from a woody caudex; flowers variously arranged, but if head- like then commonly spreading; pods various, but if erect then not long- pilose and often over 12 mm long 28 28(27). Pubescence of herbage consisting largely or entirely of malpighian hairs; pods erect, oblong-cylindric, fully bilocular; flowers ochroleucous; plants flowering in June and July A. canadensis — Pubescence of herbage consisting of basifixed hairs; pods not as above; flow- ers variously colored; plants flowering in springtime 29 29(28). Stipules connate-sheathing, at least some 30 — Stipules all distinct 32 30(29). Stems and leaves long-hairy; plants with nodding white flowers; pods pend- ulous, stipitate, the body narrowly oblong, straight glabrous, bilocular A. drummondii — Stems and leaves merely strigose; flowers ochroleucous; pods differing 31 31(30). Pods and ovaries glabrous, the body more than 12 mm long when mature, curved, trigonous, bilocular; plants of foothills and mountains, not with odor of selenium A. scopulorum — Pods and ovaries usually strigose, the body often less than 12 mm long when mature, straight, bisulcate, unilocular; plants of low elevations in clay soils A. hisulcatus 32(29). Flowers small, the banner 12 mm long or less 33 — Flowers larger, the banner 12.5-28 mm long 34 33(32). Pods narrowly lanceolate to lance-elliptic in outline, never inflated, semi- bilocular; plants of western Box Elder Co A. iodanthus — Pods greatly inflated, bilocular; plants of various distribution A. lentiginosus 34(32). Flowers ochroleucous, or less commonly pink-purple; pods either bladdery- inflated or leathery and dorsiventrally compressed, borne on a stipe-like gynophore (a stalk of receptacular origin), usually jointed to the pod 35 — Flowers pink-purple or bicolored, not or seldom ochroleucous (except in A. adanus); pods bladdery inflated or not, sessile or nearly so 36 35(34). Pods bladdery-inflated, 1-loculed, commonly more than 30 mm long, the stipe (gynophore) more than 2 mm long in flower A. oophorus — Pods leathery, subunilocular, never bladdery, dorsiventrally compressed, 15-30 mm long, the stipe (gynophore) 2 mm long or less in flower A. beckwithii 36(34). Flowers ochroleucous; stem and pods erect; plants rare in Utah A. adanus — Flowers pink-purple or bicolored; pods usually spreading; plants common to abundant 37 37(36). Flowers usually bicolored, borne in compact racemes; pods oblong in out- line, leathery, unilocular A. cibarius — Flowers usually pink-purple, borne in short to elongate racemes; pods blad- dery-inflated, membranous to papery, bilocular A. lentiginosus Sept. 1978 Welsh: Utah Flora, Fabaceae 237 Key II Plants of Beaver, Iron, Millard, Piute, Sanpete, and Sevier counties 1. Plants mat-forming to erect, with leaflets all spinulose-tipped and decurrent along the rachis A. kentrophyta — Plants various but not with both spinulose tips on leaflets and the leaflets all decurrent 2 2(1). Leaves simple, the blades oval to orbicular A. asclepiadoides — Leaves plurifoliolate, or if some simple then not as above 3 3(2). Plants rush-like or sprawling, the terminal leaflet confluent with the rachis and the upper leaves often simple 4 — Plants various, but seldom msh-like, the terminal leaflet jointed to the rachis and all leaves commonly plurifoliolate 9 4(3). Flowers less than 10 mm long; stems arising from a slender subterranean, subrhizomatous caudex 5 — Flowers 15-20 mm long; stems arising from a superficial caudex 6 5(4). Pods bladdery-inflated; pubescence of malpighian hairs; stems usually sprawl- ing A. ceramicus — Pods narrowly oblong, not inflated; pubescence basifixed; stems erect A. convallarius 6(4). Flowers ochroleucous or very pale fresh pink 7 — Flowers pink-purple or bicolored 8 7(6). Racemes commonly shorter than the subtending leaves; calyx not brown; pods sessile, curved; plants in rounded clumps, not especially rush-like A. tetrapterus — Racemes much longer than subtending leaves; calyx brown; pods long-stipi- tate, pendulous, straight; plants rush-like A. lonchocarpus 8(6). Uppermost leaves often simple; flowers bicolored; pods sessile, erect; plants of western Millard County A. toanus — Uppermost leaves usually with tiny leaflets; flowers uniformly pink-purple; pods stipitate, pendulous; plants of Sevier Co A. coltonii 9(3). Plants annual, usually growing in sand; pods bladdery-inflated A. geyeri — Plants perennial, of various soils; pods various 10 10(9). Pubescence of herbage consisting largely or entirely of malpighian hairs 11 — Pubescence of herbage consisting of simple basifixed hairs 15 11(10). Plants acaulescent; stipules all distinct 12 — Plants caulescent; stipules connate, at least those at the lowermost nodes 13 12(11). Wing-tips deeply cleft apically; flowers usually bicolored, or varicolored in populations; pods bilocular, oblong A. calycosus — Wing-tips entire; flowers ochroleucous suffused with purple; pods unilocular, obliquely ovoid; plants of eastern Sevier Co A. consobrinus 238 Great Basin Naturalist Vol. 38, No. 3 13(11). Flowers yellowish; pods erect; plants erect, with odor of selenium A. flavus — Flowers pink-purple or ochroleucous; pods ascending; plants spreading, not with odor of selenium 14 14(13). Racemes 1- to 3-flowered; flowers pink-purple; leaflets 7-11; plants rare in Sanpete Co A. sesquiflorus — Racemes 7- to many-flowered; flowers ochroleucous or suffused with dull purple; leaflets 11-17; plants of Beaver and Iron cos A. humistratus 15(10). Flowers small, the banner 12 mm long or less 16 — Flowers larger, the banner 12.5-20 mm long or more 29 16(15). Stipules all distinct, not even the lowermost connate 17 — Stipules connate into a sheath, at least at the lowermost nodes 21 17(16). Flowers 5.5-8.5 mm long, bicolored; pods narrowly oblong, 3-angled, stipi- tate, the stipe 1.4-2 mm long A. straturensis — F owers over 8.5 mm long, or if shorter then not bicolored; pods either bladdery-inflated or sessile, or both 18 18(17). Flowers dull purplish; pods oblong in outline; caudex usually subterranean A. pinonis — Flowers ochroleucous to pink-purple; pods bladdery-inflated; caudex super- ficial 19 19(18). Flowers very small, ochroleucous, the banner 5.2-8 mm long; pods di- aphanous, unilocular; plants of the Sevier Valley A. wardii — Flowers larger, or purplish, or the pods not especially diaphanous, or else bi- locular; plants of various distribution 20 20(19). Pods with a stipe 1-2.5 mm long, unilocular; flowers usually pink-purple; plants of plateaus in Iron and Piute cos A. serpens — Pods sessile, bilocular; flowers ochroleucous tinged purplish; plants mainly of western Beaver and Iron cos A. lentiginosus 21(16). Stipules turning black on drying; flowers ochroleucous; pods stipitate, the body strongly laterally flattened A. tenellus — Stipules not turning black on drying; flowers variously colored; pods stipitate or sessile, the body not as above 22 22(21). Plants subacaulescent, the internodes obscured by stipules and leaf bases 23 — Plants short- to long-caulescent, the internodes apparent though sometimes short 25 23(22). Flowers ochroleucous; plants of lake shores and ridges in eastern Iron Co A. limocharis — Flowers pink-purple; plants of ridge tops in western Beaver and Sanpete cos 24 24(23). Leaflets glabrous above; pods unilocular; plants of Sanpete Co A. montii — Leaflets strigose above; pods semibilocular; plants of western Beaver Co. A. platytropis 25(22). Plants erect or ascending; pods stipitate, pendulous 26 Sept. 1978 Welsh: Utah Flora, Fabaceae 239 — Plants prostrate to decumbent or erect; pods short-stipitate to subsessile, usu- ally not pendulous 27 26(25). Flowers deflexed, numerous; plants with odor of selenium A. hisulcatus — Flowers ascending to spreading; plants lacking odor of selenium A. australis 27(25). Plants high alpine dwarfs with bladdery-inflated pods; known from Tertiary igneous gravels in Piute Co A. perianus — Plants of high to low elevations, spreading, the pods oblong or merely tur- gid 28 28(27). Plants erect; keel with an elongate erect beak, pods oblong, not at all inflated A. miser — Plants spreading-decumbent, keel-tip merely rounded; pods inflated A. subcinereus 29(15). Plants acaulescent or subacaulescent; herbage grayish or silvery-pubescent 30 — Plants caulescent (except subacaulescent in some A. megacarpus); herbage usually green 39 30(29). Plants strictly acaulescent; leaflets 3-11, or flowers 1-6 per raceme, or both; thatch of persistent leaf bases and stipules often obscuring the caudex branches 31 — Plants not strictly acaulescent, or if so then leaflets more than 11, or flowers more than 8, or both; thatch of persistent leaf bases and stipules poorly de- veloped or lacking 33 31(30). Herbage pubescent with malpighian hairs (even though the attachment just above the base); flowers ochroleucous, tinged with purple, the keel purple- tipped; plants local in Piute and Sevier cos A. loanus — Herbage pubescent with basifixed hairs; flowers ochroleucous or pink-purple, the keel purple-tipped; plants more widely distributed 32 32(31). Petals commonly ochroleucous, sometimes faintly suffused with purple; valves of pod scarcely obscured by curly hairs A. eurekensis — Petals pink-purple; valves of pod obscured by contorted underhairs A. newberryi 33(30). Herbage pubescent with malpighian hairs; pods strigose to strigillose; plants of Iron Co A. cmiphioxys — Herbage pubescent with basifixed hairs; pods variously pubescent; plants of broad or other distribution 34 34(33). Leaves very densely hirsute with the longer hairs spirally twisted; pods bi- locular, densely long-hairy A. moUissimus — Leaves variably pubescent, if densely tomentose, then the hairs all extremely fine, sinuous, and cottony, none straight and spirally twisted; pods unilocu- lar, variously hairy 35 35(34). Pubescence of leaves (and commonly of entire plant) softly villous-tomen- tose, composed of extremely fine, cottony, or contorted and entangled hairs; pods both villous-tomentose and hirsute A. utahensis — Pubescence of leaves various, composed either of straight, appressed of nar- rowly ascending hairs, or of spreading-incurved and^ sometimes sinuous and contorted hairs; pods strigulose, villous, or hirsute ^.:. 36 240 Great Basin Naturalist Vol. 38, No. 3 36(35). Pods strigose to strigulose, the valves not obscured by the pubescence A. argophyllus — Pods hirsute or tomentose or both, the valves usually obscured by the pu- bescence 37 37(36). Valves of pod hirsute with lustrous hairs, not obscured by the hairs; plants of sandy sites at low elevations in Millard Co A. callitlirix — Valves of pods shaggy-hirsute and tomentose, almost or quite concealed by the hairs; plants of various distribution .38 38(37). Leaf pubescence appressed or nearly so; petals not very strongly graduated, the banner 17-21 and the keel 15-19 mm long; pods 20-35 mm long; ovules 27-36; plants widespread A. marianus — Leaf pubescence mostly ascending; petals strongly graduated, the banner 18-22.5, the keel 12-13.5 mm long; pods to 12 mm long; ovules 14-16; plants local in Sanpete Co A. desereticiis 39(29). Plants subacaulescent; inflorescences with 1-4 pink-purple flowers, soon sur- passed by the leaves; pods unilocular, 25-60 mm long, bladdery-inflated A. megacarpus — Plants caulescent; inflorescences with 5-many ochroleucous to pink-purple flowers, often surpassing the leaves; pods uni- or bilocular, often less than 25 mm long, bladdery-inflated or not 40 40(39). Stems arising from slender, rhizome-like caudex branches; flowers subsessile, in head-like racemes, erect or ascending; pods erect, long-pilose, less than 12 mm long A. agrestis — Stems arising from a woody caudex; flowers variously arranged, but if in head-like racemes then usually spreading; pods various but if erect then not long-pilose and often over 12 mm long 41 41(40). Stipules connate into a sheath, at least at the lowermost nodes (Note: go to couplet 29, Key I) — Stipules all distinct 42 42(41). Plants odoriferous selenophytes of claysoils; flowers ochroleucous; pods leath- ery-woody, cylindric to ovoid, spreading to ascending 43 — Plants not with odor of selenium, of various soils; flowers ochroleucous, pink-purple, or bicolored; pods inflated, often bladdery and membranous, but if leathery then of different shape 44 43(42). Calyx ochroleucous or whitish, as pale as the petals; pods cylindric, steeply ascending; plants of eastern Sevier Co ^- pattersonii — Calyx of somewhat different hue from the petals; pods normally spreading- ascending and ellipsoid or broadly cylindric; plants of western Sevier, Mil- lard, Beaver, and Iron cos .'. A. praelongus 44(42). Pods and ovaries long-hairy, at maturity plumply ovoid; flowers ochro- leucous, steeply ascending; plants introduced in Sanpete Co., but to be ex- pected elsewhere A- cicer — Pods and ovaries strigose to glabrous, at maturity variously shaped; flowers ochroleucous, pink-purple, or bicolored, seldom steeply ascending; plants in- digenous and widespread '^^ Sept. 1978 Welsh: Utah Flora, Fabaceae 241 45(44). Flowers ochroleucous 46 — Flowers pink-purple or bicolored 47 46(45). Pods bladdery-inflated, unilocular, usually more than 30 mm long at matu- rity; stipe more than 2 mm long subequal to the calyx tube A. oophorus — Pods leathery, subunilocular, never bladdery, dorsiventrally compressed, 15-30 mm long; stipe 2 mm long or less in flower A. beckwithii 47(45). Flowers bicolored, the wing-tips white, borne in subcapitate racemes; pods oblong, leathery, sessile A. cibarius — Flowers pink-purple, borne in open racemes; pods various, but not as above 48 48(47). Pods and ovaries bilocular, sessile or nearly so; plants widespread and com- mon A. lentiginosus — Pods and ovaries subunilocular or unilocular, shortly to moderately stipitate; plants of rather restricted range in western Millard, Iron, and Beaver cos 49 49(48). Pods bladdery-inflated, unilocular, usually more than 30 mm long; stipe sub- equal to the calyx tube; calyx tube more than 7 mm long A. oophorus — Pods leathery, subunilocular, never bladdery, dorsiventrally compressed 15-30 mm long; stipe shorter than the calyx tube; calyx tube less than 7 mm long A. beckwithii Key III Plants of Washington County I. Plants rush-like or sprawling, the terminal leaflet confluent with the rachis, and the upper leaves often simple 2 Plants various, but seldom rush-like, the terminal leaflet jointed to the rachis and all leaves plurifoliolate 5 2(1). Flowers more than 12 mm long, the petals ochroleucous or suffused with pale pink; pods curved, dorsiventrally compressed A. tetrapterus Flowers less than 12 mm long, the petals variously colored; pods neither curved nor dorsiventrally compressed 3 3(2). Pods and ovaries stipitate, at maturity bladdery-inflated; pubescence malpi- ghian; stems from elongated rhizome-like caudex branches A. ceramicus - Pods and ovaries sessile or subsessile, oblong in outline, not inflated; pu- bescence basifixed; stems from superficial to deep-seated caudex 4 4(3). Stipules connate into a sheath, at least at lowermost nodes; plants of north- western Washington Co A. convallarius - Stipules all distinct; plants of eastern Washington Co A. lancearius 5(1). Plants slender, diminutive annuals with tiny flowers and curved bilocular pQ^g A. nuttallianus — Plants perennial, and otherwise commonly differing from above 6 6(5). Pubescence of herbage consisting largely or entirely of malpighian hairs 7 — Pubescence of herbage consisting of basifixed hairs 11 242 Great Basin Naturalist Vol. 38, No. 3 7(6). Flowers pink-purple, more than 12 mm long; pods 20-30 mm long or more A. amphioxys — Flowers variously colored, but if pink-purple then usually less than 12 mm long or pods shorter than 12 mm long 8 8(7). Plants acaulescent; wing petals deeply cleft apically; pods bilocular A. calycosus — Plants with well-developed stems; wing petals entire apically; pods uni- or bilocular 9 9(8). Flowers ochroleucous, nodding at an thesis; pods erect, bilocular, oblong- cylindric A. canadensis — Flowers ochroleucous to pink-purple, ascending to anthesis; pods erect to spreading, variously shaped, unilocular 10 10(9). Stems prostrate-spreading; pods curved, not bisulcate ventrally; plants not with odor of selenium A. humistratus — Stems erect or ascending; pods straight, bisulcate ventrally; plants with odor of selenium A. flavus 11(6). Flowers small, the banner 12 mm long or less 12 — Flowers larger, the banner 12.5-25 mm long or more 15 12(11). Flowers 6.5-8.5 mm long, bicolored; pods narrowly oblong, 3-angled, stipi- tate, the stipe 1.4-2 mm long A. straturensis — Flowers over 8.5 mm long, or if shorter then not bicolored; pods bladdery- inflated, sessile or subsessile 13 13(12). Plants subacaulescent, usually less than 5 cm tall; flowers 1-5; rare in sandy sites in eastern Washington Co A. striatiflorus — Plants caulescent, usually over 20 cm tall; flowers numerous; plants of vari- ous habitats and distribution 14 14(13). Pods bilocular, diaphanous; lowermost stipules distinct A. lentiginosus — Pods unilocular, opaque and usually mottled; lowermost stipules shortly con- nate-sheathing A. siibcinereus 15(11). Plants acaulescent or subacaulescent; herbage grayish or silvery hairy 16 — Plants caulescent; herbage usually green 22 16(15). Plants strictly acaulescent, the caudex branches obscurred by a thatch of persistent leaf bases; leaflets 11 or fewer A. newberryi — Plants subacaulescent, the caudex branches not obscurred by a thatch of leaf bases; leaflets commonly more than 11 17 17(16). Leaflets more than 21 on at least some mature leaves; pods strigose or strigulose A. tephrodes — Leaflets fewer than 21 on all leaves, or the pods densely villous 18 18(17). Leaves very densely hirsute with the longer hairs spirally twisted; pods bi- locular, densely long-hairy A. mollissimus — Leaves variably pubescent, if densely tomentose, the hairs all extremely fine, sinuous, and cottony, none straight and spirally twisted; pods unilocular, variously hairy 19 Sept. 1978 Welsh: Utah Flora, Fabaceae 243 Pods densely long hairy 20 Pods merely strigose or strigulose 21 Leaflets oval to orbicular, rounded apically, white cottony-hairy; calyx tube more than 4 mm wide (when pressed) A. utahensis — Leaflets elliptic to obovate, obtuse or acute to rounded apically, silvery stri- gose; calyx tube less than 4 mm wide A. marianus 21(19). Leaflets mostly acute; pods brightly mottled; rocky ledges and talus of sand- stone canyons and escarpments A. zionis — Leaflets obtuse to acute; pods not mottled; plants mostly of humus in moun- tain brush and upwards A. argophyllus 22(15). Stipules connate-sheathing, at least at the lowermost nodes 23 — Stipules all distinct 24 23(22). Petals ochroleucous, the keel purple-tipped; pods pendulous, the body ob- long-cylindric, bisulcate; plants with odor of selenium A. bisulcatus — Petals pink-purple or ochroleucous; pods erect, the body ovoid, not bisul- cate A. ampullarius 24(22). Plants odoriferous selenophytes, usually of clay soils; flowers ochroleucous or pink-purple with white wing-tips; pods oblong-cylindric, ascending or spreading 25 — Plants not odoriferous selenophytes, of various soils, flowers variously col- ored; pods various 26 25(24). Flowers bicolored; calyx suffused dark-purple; pods erect-ascending A. preussii — Flowers ochroleucous; calyx greenish; pods ascending-spreading A. praelongus 26(24). Pods and ovaries stipitate, the stipe at maturity subequal to or surpassing the calyx 27 — Pods and ovaries sessile or nearly so 28 27(26). Stems decumbent to ascending; pods bladdery-inflated, unilocular, the body usually over 25 mm long; flowers ochroleucous or pink-purple A. oophorus — Stems erect; pods not or only somewhat inflated, bilocular, the body usually less than 20 mm long; flowers ochroleucous or pink-purple A. eremiticus 28(26). Pods bladdery-inflated, ovoid or merely curved-oblong, bilocular, sessile; flowers variously colored, but not usually bicolored A. lentiginosus — Pods not bladdery, oblong, usually curved, bi- or unilocular; flowers com- monly bicolored 29 29(28). Flowers borne in subcapitate racemes; pods dorsiventrally compressed, imi- locular A- cibarius — Flowers borne in elongate racemes; pods laterally or trigonously compressed, bilocular ^- ensiformis Key IV Plants of Daggett, Duchesne, Rich, Summit, Uintah, and Wasatch Counties 1. Plants mat-forming or erect, with leaflets all spinulose-tipped and decurrent 244 Great Basin Naturalist Vol. 38, No. 3 along the rachis A. kentrophyta — Plants various, but not with the leaflets both spinulose-tipped and decurrent on the rachis 2 2(1). Leaves simple, oval to orbicular; plants with odor of selenium A. asclepiadoides — Leaves various, but if simple then linear to linear-oblanceolate; plants with or without odor of selenium 3 3(2). Leaves simple (rarely with leaflets on some leaves), the blades grass-like; plants acaulescent 4 — Leaves plurifoliolate, or rarely trifoliolate, or the uppermost only simple; if acaulescent never as above 5 4(3). Leaves not over 8 cm long; racemes with flowers usually fewer than 8, less than 3.5 cm long in fruit A. spatulatus — Leaves usually more than 8 cm long; racemes with flowers usually more than 8, more than 4.5 cm long in fruit A. chloodes 5(3). Plants rush-like or sprawling, the terminal leaflet confluent with the rachis, and some of the upper leaves often simple (see also A. duchensnensis) 6 — Plants various, but seldom nish-like, the terminal leaflet jointed to the rachis and all leaves plurifoliolate (except in some A. detritalis, A. aretioides and A. gilviflorus) 10 6(5). Flowers 10 mm long or less; stems arising from a slender subterranean subrhizomatous caudex 7 — Flowers 15-20 mm long or more; stems arising from a superficial caudices 8 7(6). Pods bladder-inflated; pubescence of malpighian hairs; stems usually sprawl- ing A. ceramicus — Pods narrowly oblong, not inflated; pubescence of basifixed hairs; stems commonly erect A. convallarius 8(6). Flowers pink-purple; pods leathery-woody, laterally compressed; plants se- lenophytes of clays and silts, restricted to Uintah Co A. saimnus — Flowers ochroleucous to yellow; pods various, but not as above; plants se- lenophytes or not, from Uintah or Daggett cos 9 9(8). Pods stipitate, pendulous; plants not with odor of selenium, usually 3.5 dm tall or more, known only from Uintah Co A. hamiltonii — Pods sessile or subsessile, ascending; plants seleniferous, less than 3.5 dm tall, known only from Daggett Co A. nehonianus 10(5). Plants annual, usually growing in sand; pods bladdery-inflated A. geyeri — Plants perennial, of various soils; pods various 11 11(10). Pubescence of herbage consisting largely or entirely of malpighian hairs 12 — Pubescence of herbage consisting of basifixed hairs 18 12(11). Plants acaulescent or subacaulescent, the herbage grayish or whitish pub- escent 13 — Plants caulescent, the herbage commonly green 16 Sept. 1978 Welsh: Utah Flora, Fabaceae 245 13(12). Flowers ochroleucous, borne sessile among the leaves; leaves trifoliolate; plants of Summit Co A. gilviflorus — Flowers pink-purple or pale and suffused with purple, pedunculate; leaves with 5 or more leaflets on at least some leaves; plants of Daggett, Duchesne, and Uintah cos 14 14(13). Leaves with leaflets 3, silvery strigose; flowers 6-8 mm long; plants local in Daggett Co A. aretoides — Leaves simple or with 3-17 leaflets, strigose but not silvery; flowers 13-20 mm long or more; plants more widely distributed 15 15(14). Plants strictly acaulescent; leaflets narrowly oblong, spinulose-tipped; pods linear-oblong A. detritalis — Plants subacaulescent; leaflets obovate, rounded to obtuse apically; pods el- lipsoid A. chamaeleuce 16(13). Plants selenophytes, usually of clay soils; flowers yellowish; pods unilocular A. flavus — Plants not selenophytes, of various soils; flowers ochroleucous or pink- purple; pods bilocular 17 17(16). Flowers pink-purple, erect or steeply ascending at anthesis; stems from a caudex; plants known from Daggett Co A. adsurgens — Flowers ochroleucous, nodding at anthesis; stems from creeping rhizomes A. canadensis 18(12). Flowers small, the banner 12 mm long or less 19 — Flowers larger, the banner 12.5-25 mm long or more 30 19(18). Stipules all di^inct, not even the lowermost connate 20 — Stipules connate-sheathing, at least at the lowest nodes 24 20(19). Plants dwarf, arising from a deeply seated caudex and elongate rhizome-like branches; flowers ochroleucous; pods bladdery-inflated, strigose, unilocular; plants of Green River Shale, Uintah Co A. lutosus — Plants differing from above, the caudex superficial; flowers pink-purple or ochroleucous; pods various, but if bladdery-inflated then bilocular or spread- ing-hairy 21 21(20). Flowers ochroleucous, suffused faintly with purple; pods bladdery-inflated, strigose to glabrous; plants of Wasatch and Summit cos A. lentiginosus — Flowers pink-purple or bicolored; pods not inflated or if so then spreading- hairy; plants of Summit, Duchesne, and Uintah cos 22 22(21). Calyx more than half as long as the petals; pods sessile, straight, spreading; plants of moderate elevations, north slope of Uinta Mountains A. eucosmus — Calyx less than half as long as the petals; pods stipitate or sessile, straight and pendulous or spreading and curved; plants of low elevations in the Uinta Basin 23 23(22). Leaflets linear to narrowly oblong; calyx teeth broadly triangular, 1 mm long or less; pods pendulous, straight, strigose A. duchesnensis — Leaflets oblanceolate to obovate; calyx teeth narrowly subulate, more than 2 246 Great Basin Naturalist Vol. 38, No. 3 mm long; pods sessile, inflated, spreading, curved, long-pilose A. pubentissimus 24(19). Plants dwarf, less than 6 cm tall; caudex branches with a persistent thatch of marcescent stipules; flowers minute, 6.5 mm long or less; pods bladdery- inflated; known from Rich Co A. jejunus — Plants taller; caudex branches not as above; flowers commonly larger; pods not bladdery-inflated; distribution various 25 25(24). Stipules turning black on drying; flowers ochroleucous; pods short-stipitate, strongly laterally flattened A. tenellus — Stipules not turning black on drying; flowers pink-purple or ochroleucous; pods various, but if as above then flowers pink-purple 26 26(25). Plants odoriferous selenophytes; flowers pink-purple or ochroleucous, the keel purple-tipped, numerous, nodding at anthesis; pods bisulcate ventrally A. hisulcatus — Plants not smelling of selenium; flowers pink-purple or ochroleucous; pods not bisulcate ventrally 27 27(26). Flowers ochroleucous; pods long-stipitate more than 20 mm long at matu- rity, laterally compressed but not flattened; plants known from Wasatch Co. A. australis — Flowers pink-purple or if stipitate then less than 15 mm long and flattened laterally 28 28(27). Pods and ovaries sessile or substipitate, the body flattened; plants known from Duchesne Co A. wingatanus — Pods and ovaries sessile or substipitate, the body various, but not strongly flattened; distribution various 29 29(28). Plants with sprawling stems, usually of pinyon-juniper and desert shrublands; pods terete, mostly 12 mm long or less; rare in Uintah Co A. flexuostis — Plants with upright stems, usually of aspen or spruce-fir woodlands; pods lat- erally compressed, usually 15 mm longer more A. miser 30(18). Plants acaulescent or subacaulescent; herbage grayish or silvery hairy 31 — Plants caulescent (subacaulescent in some A. megacarpus); herbage usually green 34 31(30). Leaves very densely hirsute-tomentose, with the longer hairs straight and spirally twisted; pods bilocular, densely woolly-hairy A. mollissimus — Leaves variously pubescent, but if densely tomentose, the hairs all fine, sinuous, and cottony, none straight and spirally twisted 32 32(31). Pubescence of leaves various, but not of extremely fine entangled hairs; pods merely strigose A. argophyllus — Pubescence of leaves softly villous-tomentose, consisting of fine, cottony, contorted or entangled hairs; pods both villous-tomentose and hirsute 33 33(32). Leaflets mostly obovate and obtuse; flowers bright pink-purple A. utahensis — Leaflets either elliptic and acute or petals ochroleucous A. purshii 34(30). Plants subacaulescent; flowers pink-purple, 1-5 per raceme; pods bladdery- Sept. 1978 Welsh: Utah Flora, Fabaceae 247 inflated, unilocular, commonly over 30 mm long A. megacarpus — Plants differing in one or more ways from above 35 35(34). Plants odoriferous selenophytes, usually of clay soils at lower elevations; flowers nodding at anthesis 36 — Plants not with odor of selenium, usually of loamy soils at moderate eleva- tions 37 36(35). Pods and ovaries strigose, at maturity ventrally bisulcate, mostly less than 15 mm long A. bisulcatus — Pods and ovaries glabrous, at maturity trigonous, not bisulcate, mostly over 15 mm long A. racemosus 37(35). Stems arising from slender, rhizome-like caudex branches; flowers subsessile in head-like racemes, erect or ascending; pods erect, long-pilose, less than 12 mm long A. agrestis — Stems arising from a caudex; flowers variously arranged, but if in head-like racemes then spreading; pods not erect, usually more than 15 mm long 38 38(37). Flowers in compact, head-like racemes; petals bicolored; pods oblong, dorsi- ventrally compressed A. cibarius — Flowers in elongate racemes, or if somewhat shortened then petals pink- purple or the pods bladdery-inflated 39 39(38). Flowers ochroleucous; pods stipitate, pendulous A. scopulorum — Flowers pink-purple; pods sessile, spreading A. lentiginosus Key V Plants of Carbon, Emery, and Wayne Counties 1. Plants mat-forming or erect, with leaflets all spinulose-tipped and decurrent along the rachis A. kentrophyta — Plants various, but not with both leaflets spinulose-tipped and decurrent along the rachis 2 2(1). Leaves simple, oval to orbicular; plants with odor of selenium A. asclepiadoides — Leaves usually plurifoliolate, at least some (simple in A. spatulatus); plant with or without odor of selenium 3 3(2). Leaves simple; plants pulvinate caespitose; plants of Emery Co. A. spatulatus — Leaves plurifoliolate, at least some; plants of various distribution 4 4(3). Plants rush-like or sprawling, the terminal leaflet confluent with the rachis, some of the uppermost leaves sometimes simple 5 — Plants various, but not or seldom rush-like, the terminal leaflet jointed to the rachis and all leaves plurifoliolate (rarely the lowermost simple) 14 5(4). Pubescence of herbage consisting largely or entirely of malpighian hairs 6 — Pubescence of herbage consisting of basifixed hairs 7 6(5). Flowers 10 mm long or less; stems arising singly from elongate, rhizome-like 248 Great Basin Naturalist Vol. 38, No. 3 caudex branches; pods bladdery-inflated, stipitate; plants not with odor of selenium A. ceramicus — Flowers 15 mm long or more; stems arising several together from a sub- terranean caudex; pods laterally compressed, sessile; plants with odor of sele- nium A. woodruff ii 7(5). Flowers 10 mm long or less 8 — Flowers 11-25 mm long or more 10 8(7). Flower numerous; calyx densely pilose; pods short, spreading-ascending A. moencoppensis — Flowers 12 or fewer; calyx merely strigose; pods narrowly oblong, declined to pendulous 9 9(8). Pods and ovaries stipitate; flowers bright pink-purple; plants of sandy wash- es; known only from Capitol Reef National Park A. harrisonii — Pods and ovaries sessile; flowers dull purplish; plants of different distribu- tion A. convallarius 10(7). Flowers ochroleucous; calyx brown; pods stipitate, pendulous; plants often over 50 cm tall A. lonchocarpus — Flowers pink to pink-purple; calyx often cyaneus, not brown; pods various, usually sessile, or if stipitate then less than 40 cm tall 11 11(10). Plants tall selenophytes; flowers 20 mm long or more; plants of clay or silt in San Rafael Swell A. rafaelensis — Plants not with odor of selenium; flowers 17 mm long or less; plants of vari- ous soils and distribution 12 12(11). Pods and ovaries sessile or nearly so; petals pale pink or tinged with pur- plish; plants of San Rafael Swell and southward A. episcopus — Pods and ovaries stipitate, at maturity the stipe 3-6 mm long or more; pet- als mostly bright pink-purple; plants usually west or east of the San Rafael Swell proper 13 13(12). Caudex superficial; flowers spreading to declined at anthesis; plants mostly west of the San Rafael Swell proper A. coltonii — Caudex subterranean; flowers ascending at anthesis; plants east of the San Rafael Swell proper A. nidularius 14(4). Plants definitely annual; flowers usually less than 8 mm long; pods bladdery- inflated and unilocular, or curved-oblong and bilocular 15 — Plants perennial, though sometimes flowering the first year, but the flowers then mostly over 8 mm long; pods various 16 15(14). Pods bladdery-inflated unilocular; stems and peduncles often 1 mm wide or more A. geyeri — Pods curved-oblong, bilocular; stems and peduncles filiform, mostly less than 1 mm thick A. nuttallianus 16(14). Herbage pubescent largely or entirely with malpighian hairs 17 — Herbage pubescent with basifixed hairs 25 17(16). Plants caulescent; pods short, ascending; stipules connate, at least the lower- Sept. 1978 Welsh: Utah Flora, Fab ace ae 249 most 18 — Plants acaulescent or subacaulescent, not smelling or selenium; pods various; stipules all distinct 19 18(17). Plants odoriferous selenophytes, of low elevations, pods unilocular A. flavus — Plants not smelling of selenium, of moderate to high elevations; pods bilocu- lar ; .^ A. adsurgens 19(17). Wing tips deeply cleft apically; flowers usually bicolored or varicolored in populations; pods bilocular, oblong A. calycosus — Wingtips entire; flowers various in color; pods unilocular, variously shaped 20 20(19). Leaflets (1) 3-5 (rarely more in A. loanus); plants strictly acaulescent, the caudex branches obscurred by a thatch of persistent leaf bases; pods spread- ing-hairy 21 — Leaflets mostly more than 5, at least on some leaves; plants various, but if strictly acaulescent the thatch poorly or not developed; pods strigose 22 21(20). Leaflets 1-3 on most leaves; flowers pink-purple; pods with hairs to 2 mm long; plants rather widespread in the region A. musiniensis — Leaflets usually 5 on most leaves; flowers ochroleucous or tinged purplish; pods with hair 2-2.5 mm long; plants of western Wayne Co A. loanus 22(20). Flowers ochroleucous, tinged purplish, 11-12.5 mm long; pods laterally com- pressed only near the apex; plants of western Emery and Wayne cos. A. consobrinus — Flowers pink-purple or bicolored, mostly 15-25 mm long; pods laterally compressed throughout or dorsiventrally so 23 23(22). Pods laterally compressed, straight; persistent; flowers often pale or dull purplish; common in the vicinity of the San Rafael Swell A. cymboides — Pods dorsiventrally compressed, usually curved, deciduous; flowers pink- purple or bicolored; common to uncommon in the region 24 24(23). Flower usually bicolored; walls of pod at least 1 mm thick, the endocarp and exocarp separated by a thick mesocarp; plants rare in the region A. chamaeleuce — Flowers usually bright pink-purple; walls of pod less than 1 mm thick, be- coming leathery when ripe; plants common A. amphioxys 25(16). Flowers small, the banner 12 mm long or less 26 — Flowers larger, the banner 12.5-15 mm long or more 39 26(25). Stipules all distinct, not even the lowermost connate 27 — Stipules connate-sheathing at least at the lowermost nodes 32 27(26). Flowers 5 mm long or less, 2-5, born on linear-filiform peduncles; plants sprawling, usually of volcanic gravels A. brandegei — Flowers commonly 7 mm long or more, usually more than 5, borne on sub- stantial peduncles; plants ascending to erect, never really sprawling, seldom, if ever, of volcanic soils 28 28(27). Plants subacaulescent, the stems shorter than the inflorescence; stipules 250 Great Basin Naturalist Vol. 38, No. 3 prominent A. desperatus — Plants caulescent, the stems longer than the inflorescence; stipules hardly prominent 29 29(28). Pods strongly mottled, bladdery-inflated, merely strigose; plants known from western Wayne Co A. serpens — Pods not or slightly mottled, inflated but hardly bladdery; villous with spreading hairs; plants mostly along the sandy eastern portion of the region 30 30(29). Racemes with 10 or more flowers; plants rare in the region A. pubentissimus — Racemes with 10 or fewer flowers; plants rare to locally common 31 31(30). Pods 5-8 mm in diameter, curved-oblong; ovules 10-19; plants rare in the region A. sabulonum — Pods 8-11 mm in diameter, ovoid-ellipsoid; ovules 20-28; plants locally common A. pardalinus 32(26). Stipules turning black on drying; flowers ochroleucous; pods short-stipitate, strongly laterally flattened; plants of the high plateau sections A. tenellus — Stipules not turning black on drying; flowers various; pods various, but if as above then flowers pink-purple or plants of low elevations 33 33(32). Stems shorter than the inflorescences; flowers pink-purple, numerous; pods erect; sessile A. moencoppensis — Stems longer than the inflorescences, or if shorter than the flowers 10 or fewer per raceme; pods seldom erect 34 34(33). Plants odoriferous selenophytes; flowers ochroleucous, numerous, nodding at anthesis; pods bisulcate ventrally A. bisulcatus — Plants not smelling of selenium; flowers pink-purple or sometimes ochro- leucous, ascending at anthesis; pods not bisulcate ventrally 35 35(34). Pods and ovaries stipitate, the body flattened or triquetrous 36 — Pods and ovaries sessile or subsessile, the body various but not strongly flat- tened 37 36(35). Pods not strongly compressed; wing petals bilobed; plants of high elevation in Emery Co A. australis — Pods strongly compressed; wing petals entire; plants of lower elevations in Carbon and Emery cos A. wingatanus 37(35). Plants with elongate clambering stems, known from moist meadows in west- ern Wayne Co A. bodinii — Plants with short to elongate, erect to sprawling stems of different distribu- tion or habitat 38 38(37). Plants with sprawling stems, commonly of pinyon-juniper and desert shrub- lands; pods mostly 15 mm long or less; known from lower elevations Carbon and Emery cos A. flexuosus — Plants with upright stems, usually of aspen or spruce-fir woodlands; pods lat- erally compressed, usually 15 mm long or more A. miser 39(26). Plants acaulescent or subacaulescent; herbage grayish- or silvery-hairy 40 Sept. 1978 Welsh: Utah Flora, Fabaceae 251 — Plants caulescent (subacaulescent in A. megacarpus); herbage usually green 43 40(39). Herbage merely strigose; flowers 13-15 mm long; stipules prominent; pods hirsute A. barnebyi — Herbage tomentose to strigose; flowers 15-25 mm long; stipules not promi- nent; pods variously pubescent 41 41(40). Leaves very densely hirsute-tomentose, with the longer hairs straight and spirally twisted; pods bilocular, densely woolly-hairy A. mollissimus — Leaves variously pubescent, but if densely tomentose, the hairs all fine, sinuous, and cottony, none straight and spirally twisted 42 42(41). Pubescence of leaves softly villous- tomentose, consisting of fine, cottony, contorted or entangled hairs; pods both villous-tomentose and hirsute A. utahensis — Pubescence of leaves strigose, not of fine entangled hairs; pods strigose A. argophyllus 43(39). Plants subacaulescent; flowers pink-purple, 1-5 per raceme; pods bladdery- inflated, unilocular, often more than 30 mm long A. megacarpus — Plants differing in one or more ways from above 44 44(43). Plants odoriferous selenophytes, usually of clay soils at low elevations; flow- ers ochroleucous and nodding or the calyx purple and flowers ascending 45 — Plants not smelling of selenium, of various soil types and elevations; flowers not with either of the combinations noted above 48 45(44). Flowers ochroleucous, nodding at anthesis; calyx whitish, cream-colored or greenish; pods woody 46 — Flowers pink-purple or bicolored, ascending at anthesis; calyx purple; pods leathery 47 46(45). Calyx and petals concolorous both whitish to cream-colored; pods cylindric, sessile, steeply ascending; plants of Carbon Co A. pattersonii — Calyx and petals discolorous, the calyx often greenish, the petals ochro- leucous; pods spreading ascending or if steeply ascending, then stipitate A. praelongus 47(45). Pods horizontally spreading or declined, borne on ascending to reclinding peduncles; stems seldom more than 10 cm long; plants rare A. eastwoodae — Pods erect or steeply ascending, born on erect peduncles; stems mostly much longer than 10 cm long; plants common A. preussii 48(44). Stems arising from slender, rhizome-like caudex branches; flowers subsessile in head-like racemes, erect or ascending; pods erect, long-pilose, less than 12 mm long A. agrestis — Stems arising from a caudex; flowers variously arranged, but not or rarely head-like, spreading at anthesis; pods not as above 49 49(48). Flowers pink to pink-purple; pods sessile, bilocular A. lentiginosus — Flowers ochroleucous; pods stipitate, uni- or bilocular 50 50(49). Stipules connate-sheathing at least some; pods bilocular, triquetrous, 3-5 mm wide A. scopulorum 252 Great Basin Naturalist Vol. 38, No. 3 — Stipules all distinct; pods unilocular, bladdery-inflated, 10-30 mm wide A. oophorus Key VI Plants of Grand and San Juan Counties 1, Leaves simple, oval to orbicular; plants with odor of selenium A. asclepiadoides — Leaves usually plurifoliolate (simple in A. spattilatus); plants with or without the odor of selenium 2 2(1). Leaves simple; plant pulvinate-caespitose, of Grand Co A. spatulatus — Leaves usually plurifoliolate; plant seldom as above, of various distribution 3 3(2). Plants rush-like or sprawling, the terminal leaflet of at least uppermost leaves confluent with the rachis, some of the uppermost leaves often simple . 4 — Plants various but seldom rush-like, the terminal leaflet jointed to the rachis in all leaves, and all leaves plurifoliolate (rarely the lowermost simple) 8 4(3). Pubescence of herbage consisting of basifixed hairs; pods not bladdery- inflated , A. ceramicus — Pubescence of herbage consisting of basifixed hairs; pods not bladdery- inflated 5 5(4). Flowers 10 mm long or less 6 — Flowers 12 mm long or more 7 6(5). Stems shorter than the inflorescence; flowers numerous, pink-purple; pods spreading-ascending A. moencoppensis — Stems longer than the inflorescences; flowers usually 15 or fewer, dull pink- purple; pods spreading-pendulous A. convallarius 7(5). Flowers ochroleucous; calyx brown; plants commonly 50 cm tall or more A. lonchocarpus — Flowers pink-purple; calyx greenish or blackish; plants commonly less than 45 cm tall, local in White Canyon, San Juan Co A. nidularius 8(3). Plants definitely annual (see also A. sabulonum); flowers usually less than 8 mm long; pods bladdery-inflated and luiilocular, or curved-oblong and bilocular 9 — Plants perennial, though sometimes flowering the first year, but the flowers then mostly more than 8 mm long; pods various 10 9(8). Pods bladdery-inflated, unilocular; stems and peduncles often 1 mm thick or more A. geyeri — Pods curved-oblong, bilocular or nearly so; stems and peduncles filiform, mostly less than 1 mm thick A. nuttallianus 10(8). Herbage pubescent largely or entirely with malpighian hairs 11 — Herbage pubescent with basifixed hairs 18 11(10). Plants caulescent selenophytes; pods erect, less than 12 mm long A. flavus — Plants acaulescent or subacaulescent, not smelling of selenium; pods various . 12 Sept. 1978 Welsh: Utah Flora, Fabaceae 253 12(11). Stems diffuse and prostrate, sometimes matted; racemes 1- to 3-flowered A. sesquiflorus — Stems various, but not as above; racemes commonly with more than 3 flow- ers 13 13(12). Plants strictly acaulescent; flowers less than 12 mm long, or leaflets 5 or fewer, or both 14 — Plants subacaulescent; flowers various; leaflets 5 or more on at least some leaves 15 14(13). Flowers 12 mm long or less; wing petals bilobed apically; pods bilocu- lar A. calycosus — Flowers 18-25 mm long; wing petals entire apically; pods unilocular A. musiniensis 15(13). Pods narrowly oblong to oblong-ellipsoid, straight, laterally compressed when ripe A. cymboides — Pods obliquely ovoid to ellipsoid, mostly curved, if straight then dorsally compressed 16 16(15). Walls of pod at least 1 mm thick, the exocarp and endocarp separated by a thick mesocarp; petals mostly bicolored A. chamaeleuce — Walls of pod much less than 1 mm thick, becoming leathery when ripe; petals pink-purple or bicolored 17 17(16). Pods persistent or tardily deciduous, mostly lance ovoid in outline; plants rare A. missouriensis — Pods readily deciduous, ellipsoid in outline; plants common A. amphioxys 18(10). Flowers small, the banner 12 mm long or less 19 — Flowers larger, the banner 12.5-25 mm long or more 32 19(18). Stipules all distinct, not even the lowermost connate 20 — Stipules connate-sheathing at least at the lowermost nodes 25 20(19). Plants subacaulescent, the stems shorter than the inflorescences; stip- ules prominent 21 — Plants caulescent, the stems longer than the inflorescences; stipules in- conspicuous 22 21(20). Pods spreading hairy, unilocular, dorsally compressed; plants wide- spread A. desperatus — Pods strigose, bilocular, laterally compressed; plants of the Cedar Mesa Sandstone, San Juan Co A. monumentalis 22(20). Stems arising from elongate rhizome-like caudex branches; pods narrowly oblong, pendulous; plants of Cutler formation. Comb Wash, San Juan Co A. cronquistii — Stems from a superficial caudex; pods ovoid to ellipsoid, mostly spread- ing; plants of various distribution 23 23(22). Petals whitish, with pink veins; pods ovoid-acuminate, strongly beaked, strigose; plants of Colorado River Canyon east from Moab A. wetherillii 254 Great Basin Naturalist Vol. 38, No. 3 — Petals pink-purple or less commonly ochroleucous; pods ellipsoid, to ovoid-ellipsoid, spreading hairy; plants of various distribution 24 24(23). Flowers 8.8-11.7 mm long, commonly more than 10 per raceme; pods most- ly more than 8 mm in diameter; plants of the Tavaputs escarp- ment A. puhentissimus — Flowers 5.8-8.2 mm long, commonly fewer than 10 per raceme; pods less than 8 mm in diameter; plants of San Juan Co A. sahulonum 25(19). Stipules turning black on drying; flowers ochroleucous; pods short-stipi- tate; strongly laterally flattened A. tenellus — Stipules not blackening on drying; flowers various; pods various; but if as above then flowers pink-purple 26 26(25). Stems shorter than the inflorescences; flowers pink-purple, numerous; pods erect, sessile A. moencoppensis — Stems longer than the inflorescences of if shorter, then the flowers 10 or fewer per raceme; pods seldom erect 27 27(26). Plants odoriferous selenophytes; flowers ochroleucous, numerous, nod- ding at anthesis; pods bisulcate ventrally A. bisulcatus — Plants not smelling of selenium; flowers pink-purple or sometimes ochroleucous, ascending at anthesis; pods not bisulcate ventrally 28 28(27). Pods sessile or subsessile, the body oblong 29 — Pods stipitate, or the body bladdery-inflated 30 29(28). Plants with sprawling or slender and erect stems; pods mostly less than 18 mm long, subterete; mostly of mountain brush and pinyon-juniper communities A. flexuosus — Plants with erect stems; pods laterally compressed usually 19 mm long or more; mostly in aspen or spruce-fir communities A. miser 30(28). Flowers 8 mm long or less; pods strongly laterally flattened; plants mostly in pinyon-juniper woodlands A. wingatanus — Flowers 8.5-11.5 mm long; pods various, but not as above; plants of various habitats 31 31(30). Herbage fresh green; racemes usually with fewer than 10 flowers; pods nar- rowly oblong, not inflated; plants of the Tavaputs Plateau, Grand Co A. alpinus — Herbage often cinereous; racemes with more than 10 flowers; pods bladdery-inflated; plants of lower elevations in San Juan Co A. fucatus 32(18). Plants acaulescent or subacaulescent; herbage often grayish- or silvery- hairy 33 — Plants caulescent; herbage often green 36 33(.32). Flowers 15 mm long or less; pods merely strigose, linear-oblong; plants of sandstone formations; San Juan Co A. cottamii — Flowers more than 15 mm long; pods variously pubescent, ovoid-ellip- soid to ellipsoid; plants of various habitats and distributions 34 34(33). Leaflets mostly 17 or more per leaf, densely hirsute-tomentose, with longer hairs straight and spirally twisted; pods bilocular, densely woolly- Sept. 1978 Welsh: Utah Flora, Fabaceae 255 hairy A. mollissimus — Leaflets mostly 15 or fewer, mostly strigose; pods unilocular 35 35(34). Leaflets mostly acute; pods brightly mottled; rocky ledges and talus of sandstone canyons and escarpments A. zionis — Leaflets obtuse to acute; pods not mottled; plants mostly of humus in mountain brush and upwards A. argophyllus 36(32). Plants odoriferous selenophytes, usually of clay soils 37 — Plants not smelling of selenium, usually not of clay soils 43 37(36). Stipules connate into a bidentate sheath, at least at the lowermost nodes; flowers pink-purple, declined at anthesis; pods stipitate, pen- dulous, bisulcate A. hisulcatus — Stipules all distinct, even at the lowermost nodes; flowers variously colored, but if declined at anthesis then ochroleucous; pods sessile or stipitate, ascending to spreading, not bisulcate 38 38(37). Calyx tube purple; petals pink-purple or bicolored; pods leathery-inflated 39 — Calyx tube green to whitish, not purplish; petal ochroleucous to whitish 40 39(38). Pods horizontally spreading or declined, borne on ascending to reclin- ing peduncles; stems seldom more than 10 cm long; plants rare A. eastwoodae — Pods erect or steeply ascending, borne on erect peduncles; stems mostly much longer than 10 cm; plants locally common A. preussii 40(38). Flowers declined at anthesis; pods ascending to erect, 9 mm in diam- eter or less, leathery-woody in texture 41 — Flowers spreading to ascending; pods spreading to declined, usually over 10 mm in diameter, leathery in texture 42 41(40). Calyx and petals concolorous, both whitish to cream-colored; pods cylindric, sessile, steeply ascending; plants not definitely known from the region but nearby in Colorado A. pattersonii — Calyx and petals discolorous, the calyx greenish, the petals ochroleucous; pods spreading-ascending or if steeply ascending then stipitate; plants common in the region A. praehngus 42(40). Flowers 17-18 mm long, the petals whitish; calyx tube 5.5-6.3 mm long; plants of Paradox and Morrison formations; La Sal Mts A. isleyi — Flowers 28-31 mm long, the petals ochroleucous; calyx tube 11.5-14 mm long; plants of the Mancos Shale formation, north of the Colorado River A. sabulosus 43(36). Stems arising from slender, rhizome-like caudex branches; flowers subsessile in head-like racemes, erect or ascending; pods erect, long-pilose, less than 12 mm long A. agrestis — Stems arising from a caudex; flowers variously arranged, but seldom head- like and ascending to spreading; pods more than 12 mm long, spreading to pendulous 44 44(43). Pods and ovaries sessile, spreading, either bladdery-inflated or curved oblong to straight, bilocular 45 — Pods and ovaries stipitate, descending to pendulous, not much inflated or 256 Great Basin Naturalist Vol. 38, No. 3 curved, uni- or bilocular 46 45(44). Pods trigonous, the sides flattened, straight, 3-5 mm thick; plants rare or extinct in Glen Canyon A. hryantii — Pods terete or dorsiventrally compressed to inflated, usually over 5 mm thick; plants common A. lentiginosus 46(44). Flowers ochroleucous; pods bilocular; plants fresh green A. scopulorum — Flowers pink-purple; pods unilocular; plants more or less cinereus A. coltonii Key VII Plants of Garfield and Kane Counties 1. Plants mat-forming or erect, with leaflets all spinulose-tipped and de- current along the rachis A. kentrophijta — Plants various, but not with leaflets both spinulose-tipped and decurrent along the rachis 2 2(1). Leaves simple, oval to orbicular; plants with odor of selenium A. asclepiadoides — Leaves usually plurifoliolate, at least some; plants with or without the odor of selenium 3 3(2). Plants annuals (see also A. subulonum); flowers commonly less than 8 mm long; pods curved-oblong and bilocular or bladdery-inflated and ovoid 4 — Plants perennial, though sometimes flowering the first year, and then differing in one or more respects from above 6 4(3). Pods bladdery-inflated, unilocular; plants of eastern Garfield Co A. geyeri — Pods curved oblong, bilocular; plants of Kane and Garfield cos 5 5(4). Keel tips roimded; leaflets all truncate-retuse on all leaves; pods de- ciduous, dehiscing at both ends after falling A. emoryanus — Keel tips pointed; leaflets not tnmcate-retuse on all leaves; pods per- sistent, dehiscent at tip only A. nuttallianus 6(3). Plants with terminal leaflets confluent with the rachis, the uppermost leaves sometimes simple 7 — Plants with terminal leaflets jointed to the rachis, the leaves all plurifo- liolate (or the lowermost simple) 16 7(6). Pubescence of herbage consisting largely or entirelv of malpighian hairs ' 8 — Pubescence of herbage consisting of basifixed hairs 9 8(7). Flowers 10 mm long or less; stems arising singly from elongate, rhizome-like caudex branches; pods bladdery-inflated, stipitate A. ceramicus — Flowers 15 mm long or more; stem arising several together from a sub- terranean caudex; pods laterally compressed sessile A. woodniffii 9(7). Flowers ochroleucous, or whitish tinged with pink, 15 mm long or more 10 Sept. 1978 Welsh: Utah Flora, Fabaceae 257 — Flowers pink-purple bicolored, or pale, 12 mm long or less 11 10(9). Inflorescences usually shorter than the subtending leaves; calyx not brown; pods sessile, curved; plants clump-forming not slender and tall, of western Kane Co A. tetrapetrus — Inflorescence much longer than the subtending leaves; calyx brown; pods long-stipitate, pendulous, straight; plants tall and slender, wide- spread A. lonchocarpus 11(9). Stems shorter than the inflorescences; flowers numerous; calyx densely pilose; pods short, spreading-ascending A. moencoppensis — Stems shorter or longer than the inflorescences; flowers few to numer- ous; calyx merely strigose; pods various but not as above 12 12(11). Flowers dull pink-purple, usually 10 mm long or less; pods narrowly oblong, sessile, the body 4 mm wide or less A. convallarhis — Flowers pale to bright pink-purple, usually over 10 mm long, or pods more than 5 mm wide 13 13(12). Pods and ovaries sessile; petals pale; plants west and south of the Henry Mountains 14 — Pods and ovaries stipitate, at maturity the stipe 3-6 mm long or more; plants west or east of those mountains 15 14(13). Calyx short-cylindric, the tube longer than broad, suffused purplish or very pale, white-strigose; ovules 16-26 A. episcopus — Calyx campanulate, the tube about as long as broad, not purplish, black-strigose; ovules 8-14 A. lancearius 15(13). Caudex superficial; flowers spreading or declined at anthesis; plants west of the Henry Mountains A. coltonii — Cadex subterranean; flowers ascending at anthesis; plants east of the Henry Mountains A. nididarius 16(6). Herbage pubescent, largely or entirely, with malpighian hairs 17 — Herbage pubescent with basifixed hairs 25 17(16). Plants caulescent; pods short, ascending 18 — Plants acaulescent or subacaulescent; pods various 20 18(17). Plants erect or descending; flowers mostly more than 12 m long; odor- iferous selenophytes A. flavus — Plants prostrate-spreading; flowers less than 12 mm long; not smelling of selenium 19 19(18). Racemes 1- to 3-flowered; flowers pink-purple; leaflets 7-11 plants of sand- stone escarpments and ledges, mostly at lower elevations A. sesqtiiflorus — Racemes 7- to many-flowered; flowers dull purplish; leaflets 11-17; plants mostly of higher plateaus A. humistratus 20(17). Plants subacaulescent, with one or more internodes usually apparent; caudex branches only rarely with persistent leaf bases 21 — Plants strictly acaulescent, the caudex branches usually with a per- 258 Great Basin Naturalist Vol. 38, No. 3 sistent thatch of leaf bases 22 21(20). Flowers usually bicolored; walls of pod at least 1 mm thick, the exocarp and endocarp separated by a thick mesocarp; plants rare in the region A. chamaeleuce — Flowers usually bright pink-purple; walls of pod less than 1 mm thick, becoming leathery when ripe; plants common A. amphioxys 22(20). Flowers mostly less than 12.5 mm long; pods less than 17 mm long 23 — Flowers more than 15 mm long; pods more than 20 mm long 24 23(22). Flowers pink to pink-purple, or bicolored, or ochroleucous, the wing petals emarginate apically; pods linear-oblong, bilocular A. calycosus — Flowers ochroleucous, the wing petals entire apically; pods ovoid-ellip- soid, unilocular A. consohrinus 24(22). Leaflets 1-3 on most leaves; flowers pink-purple; pods with hairs to 2 mm long; plants mostly east of the high plateaus A. musiniensis — Leaflets 5 on most leaves; flowers ochroleucous or tinged purplish; pods with hairs 2-2.5 mm long; plants of western Garfield Co A. loanus 25(16). Flowers small, the banner 12 mm long or less 26 — Flowers larger, the banner 12.5-25 mm long or more 40 26(25). Flowers 6 mm long or less; pods unilocular, borne on slender peduncles, re- supinate; stems sprawling; plants usually of volcanic gravels A. brandegei — Flowers over 6 mm long, and the plants differing in other ways, usually not of volcanic gravels (or seldom so) 27 27(26). Plants with stipules all distinct, not even the lowermost connate 28 — Plants with stipules connate-sheathing, at least the lowermost 33 28(27). Plants subacaulescent; stipules prominent; stems shorter than the in- florescences 29 — Plants caulescent; stipules not conspicuous; stems longer than the in- florescences 30 29(28). Pods spreading-hairy imilocular, dorsally compressed; plants widespread along canyons of the Colorado A. desperatus — Pods strigose, bilocular, laterally compressed; plants of Cedar Mesa Sand- stone, eastern Garfield Co A. monumentalis 30(28). Pods bladdery-inflated, diaphanous, strigose to glabrous; plants of moderate to high elevations in western Kane and western to eastern Garfield cos 31 — Pods only moderately inflated, opaque, spreading-hairy; plants of low eleva- tions in eastern portions of Kane and Garfield cos 32 31(30). Flowers ochroleucous; pods sessile, glabrous, or nearly so, usually not mot- tled A. wardii — Flowers pink-purple; pods strigose, short-stipitate, strongly mottled A. serpens 32(30). Pods 5-8 mm in diameter, curved-oblong; ovules 10-19; plants of eastern Kane Co A. sabulonum — Pods 8-11 mm in diameter, ovoid-ellipsoid; ovules 20-28; plants of eastern Sept. 1978 Welsh: Utah Flora, Fabaceae 259 Garfield Co A. pardalinus 33(27). Plants subacaulescent, the stems shorter than inflorescences; pods bladdery- inflated; growing in sand, on beaches, or volcanic gravels 34 — Plants caulescent, the stems shorter or longer than the inflorescences, when shorter, the pods not inflated; growing in various soil types and habitats 36 34(33). Flowers ochroleucous; plants of beaches in western Kane Co A. limnocharis — Flowers pink-purple to whitish; plants of sandy sites at low elevations or of volcanic gravels at high elevations 35 35(34). Flowers 8-9 mm long, the banner not strongly veined; plants of volcanic gravels at high elevations in western Garfield Co A. perianus — Flowers 10-12 mm long, the banner strongly veined; plants of sandy sites at low elevations in Kane Co A. striatiflorus 36(34). Stipules turning black on drying; flowers ochroleucous; pods short-stipitate, strongly laterally flattened A. tenellus — Stipules not blackening on drying; flowers various; pods various, but not as above 37 37(36). Stems shorter than the inflorescences; flowers pink-purple; numerous, calyx densely villous; pods spreading-ascending, sessile A. moencoppensis — Stems longer than the inflorescences, or if shorter, the flowers 10 or fewer, or the pods not spreading-ascending; calyx merely strigose 38 38(37). Pods oblong, not inflated; plants of aspen and spruce-fir communities at higher elevations A. miser — Pods inflated to bladdery-inflated; plants commonly of ponderosa pine, pin- yon-juniper, or desert shnib communities at moderate to lower elevations 39 39(38). Calyces and stems silvery-caulescent, the hairs appressed; pod bladdery- inflated, more than 12 mm in diameter; ovules 21-32; plants of sandy sites at low elevations in eastern Garfield Co A. fucatus — Calyces and stems not silvery-caulescent, the hairs spreading; pods moder- ately inflated less than 13 mm in diameter; ovules 10-20; plants of moder- ate elevations in Kane and Garfield cos A. suhcinereus 40(25). Plants acaulescent or subacaulescent; herbage often grayish or silvery-hairy ... 41 — Plants caulescent; herbage often green 48 41(40). Flowers 15 mm long or less, pink-purple; pods curved, 10-12 mm long, densely spreading-hairy; plants rare and local, Garfield Co A. barnebyi — Flowers more than 15 mm long; pods curved or straight, more than 12 mm long, variously hairy; plants of various distribution 42 42(41). Plants strictly acaulescent; caudex branches with a thatch of persistent stip- ules 43 — Plants subacaulescent; caudex branches with thatch poorly developed or lacking 45 43(42). Flowers ochroleucous, 15-23 mm long; pods lance-ovoid, strigose; plants known only from the Henry Mountains A. henrimontanensis 260 Great Basin Naturalist Vol. 38, No. 3 — Flowers ochroleucous or pink-purple, 18-25 mm long or more; pods spread- ing-hairy or villous; plants of various distribution but not as above 44 44(43). Flowers ochroleucous, sometimes suffused with purple; valves of pod scarcely obscured by the curly hairs A. eurekensis — Flowers pink-purple; valves of pod obscured by contorted under hairs A. newberryi 45(42). Leaflets mostly 17 or more per leaf, densely hirsute-tomentose, with longer straight hairs spirally twisted; pod bilocular, densely woolly hairy A. moUissimus — Leaflets mostly fewer than 17 per leaf, variously hairy, but not with longer straight hairs spirally twisted; pod unilocular, variously hairy 46 46(45). Leaflets oval to obovate, rounded apically; pods woolly hairy A. utahensis — Leaflets oblanceolate to elliptic, obtuse to acute apically; pods merely stri- gose 47 47(46). Leaflets mostly acute; pods brightly mottled; plants of rocky ledges and talus of sandstone canyons and escarpments, mostly at lower elevations A. zionis — Leaflets obtuse to acute; pods not mottled; plants of loamy soils in moun- tain brush and upwards A. argophyllus 48(40). Plants odoriferous selenophytes, usually of clay soils 49 — Plants not smelling of selenium, usually not of clay soils 52 49(48). Calyx tube purple; petals pink-purple or bicolored; flowers erect-ascending at anthesis; pods leathery-inflated, ascending A. preussii — Calyx tube greenish or whitish, not purplish; petals ochroleucous, or keel-tip purple, descending at anthesis; pods various 50 50(49). Stipules connate into a bidentate sheath, at least at the lowermost nodes; pods stipitate, pendulous, bisulcate A. bisulcatus — Stipules all distinct, even at the lowermost nodes; pods sessile or shortly stipitate, ascending to spreading, not bisulcate 51 51(50). Calyx and petals concolorous, both whitish to cream-colored; pods cylindric, sessile, steeply ascending; plants rare in Henry Mountains A. pattersonii — Calyx and petals discolorous, the calyx greenish, the petals ochroleucous; pods broadly ellipsoid, sessile or shortly stipitate, spreading-ascending; plants common A. praelongus 52(48). Plants subacaulescent; flowers pink-purple, 1-5 raceme; pods bladdery-inflated, unilocular, often more than 30 mm long A. megacarpus — Plants differing in one or more respects from above 53 53(52). Stems arising from slender, rhizome-like caudex branches; flowers subsessile in head-like racemes, erect or ascending; pods erect, long-pilose, less than 12 mm long A. agrestis — Stems arising from a superficial or subterranean caudex; flowers variously ar- ranged, but seldom head-like and ascending to spreading; pods more than 17 mm long, spreading to pendulous 54 54(53). Stipules connate-sheathing, at least at the lowermost nodes; caudex sub- Sept. 1978 Welsh: Utah Flora, Fabaceae 261 terranean 55 — Stipules all distinct, even at the lowermost nodes; caudex superficial to sub- terranean 56 55(54). Pods and ovaries long-stipitate; plants known from clay soils of the Chinle formation A. ampullarius — Pods and ovaries sessile or subsessile; plants known from sandy aluvium A. hallii 56(54). Pods and ovaries stipitate, the stipe at maturity subequal to or surpassing the calyx 57 — Pods and ovaries sessile or nearly so 58 57(56). Stems decumbent to ascending; pods bladdery-inflated, unilocular, the body mostly more than 25 mm long; flowers ochroleucous; plants commonly of higher middle elevations A. oophorus — Stems erect; pods not or only somewhat inflated, bilocular, the body usually less than 20 mm long; flowers ochroleucous or pink-purple; plants of lower elevations A. eremiticus 58(56). Flowers borne in subcapitate racemes; pods curved-oblong, dorsiventrally compressed, unilocular A. cibarius — Flowers usually in elongate racemes; pods ovoid to curved-oblong, bilocular, compression variable 59 59(58). Plants from a subterranean caudex, usually less than 25 mm tall; pods curved-oblong, laterally or trigonously compressed; known from the Kaiparo- wits and Circle Cliffs regions, at moderate elevations A. malacoides — Plants from a superficial caudex, often more than 25 cm tall; pods curved- oblong and dorsiventrally compressed, or bladdery-inflated and ovoid; wide- spread A. lentiginosus Astragalus adanus A. Nels. Boise Milk- Gierisch 288, UTC); Idaho; 1 (0). vetch. Perennial, caulescent, 20-45 cm tall. Astragalus adsurgens Pallas. Standing from a branching caudex; pubescence basi- Milkvetch. [A. striatus Nutt. ex torr. & fixed; stems erect or ascending; stipules 4-9 Gray; A. nitidus Douglas ex Hook.; A. ni- mm long, all distinct; leaves 7-13 (16) cm tidus var. robustior (Hook.) M. E. Jones.], long; leaflets 13-27, ovate to oblong, lance- Perennial, caulescent, 15-45 cm tall, from a oblong, or elliptic, 6-26 mm long, retuse to branching caudex; pubescence malpighian; obtuse, glabrous above, strigose below; pe- stems decumbent-ascending to erect; stip- dimcles (4) 10-18 cm long; racemes 4- to ules 6-10 mm long, at least the lowermost 27-f lowered, the flowers ascending to connate-sheathing; leaves 6-12 cm long; spreading at anthesis, the axis 2-3 cm long leaflets 15-23, 13-28 mm long, 3-9 mm in fmit; bracts lanceolate, 1.5-3.5 mm long; wide, oblong to elliptic, acute or obtuse; pedicels 1-5 mm long; bracteoles usually 2; peduncles 6-16 cm long; racemes 16- to 50- calyx 6.9-10.3 mm long, often black-pilose, flowered, the flowers ascending at anthesis, the tube 5.2-8 mm long, short-cylindric, the the axis 1.5-13 cm long in fruit; bracts 2-5 teeth 1.6-2.8 long, triangular-subulate; flow- mm long; pedicels to 0.8 mm long; brac- ers ochroleucous, 15.5-22 mm long; pods teoles 0; calyx 5.8-10.5 mm long, the tube erect, sessile, oblong-ovoid, 3-18 mm long, 4.4-7 mm long, short-cylindric, strigulose, 5-6 mm thick; ovules 14-18. Known in the teeth 1.4-4.2 mm long, subulate; flowers Utah from a single collection at the base of 13-16 mm long, pink-purple; pods erect, Mt. Nebo, Juab Co. (Mt. Nebo R.S., R. K. sessile, ovoid-oblong, 7-12 mm long, 2.3-3.8 262 Great Basin Naturalist Vol. 38, No. 3 mm thick bilocular; ovules 9-16. Juniper- sagebrush community at 2000 m in Daggett Co. and 3175 m in Emery and Sanpete cos.; Alaska east to Manitoba and south to Wash- ington, Idaho, Colorado, Nebraska, and Iowa; Eurasia. Our material belongs to ssp. robustior (Hook.) Welsh (A. adsurgens var. robustior Hook.); 3 (i). Astragalus agrestis Douglas ex G. Don. Field Milkvetch. [A. dasyglottis Fisch, ex DC, not Pallas; A. goniatus Nutt. ex Torr. & Gray; A. hypoglottis polyspermus Torr. & Gray; A. agrestis var. polyspermus (Torr. & Gray) M. E. Jones.] Perennial, caulescent, 9-43 cm tall, from a subterranean caudex and long rhizome-like caudex branches; pubescence basifixed; stems erect to decum- bent-clambering; stipules 4-11 mm long, at least the lowermost connate-sheathing; leaves 2-10 cm long; leaflets 13-23, 4-18 mm long, 2-5 mm wide, narrowly elliptic to lance-oblong, obtuse to retuse or acute, strigulose above and below; peduncles 1.5-15 cm long; racemes subcapitate, 5- to 15-flowered, the flowers ascending-erect at anthesis, the axis 0.5-2.5 mm long in fruit; bracts 3-7 mm long; pedicels 0.5-1.5 mm long; bracteoles 0; calyx 7-12.5 mm long, the tube 5-7.8 mm long, cyclindric, villous, the teeth 2.5-5.5 mm long, linear; flowers 17-24 mm long, pink-purple, ochroleucous, or almost white; pods short-stipitate, the stipe 0.3-1 mm long, the body 7-10 mm long, 2.8-4.5 mm thick, unilocular, oblong- ellipsoid, silky-villous; ovules 14-26. Mead- ows and openings in sagebrush and aspen at 1850 to 3050 m in Box Elder, Cache, Car- bon, Daggett, Duchesne, Emery, Garfleld, Juab, Kane, San Juan, Sanpete, Sevier, Sum- mit, Tooele, Uintah, Utah (US) Wasatch, and Wayne cos.; Yukon east to Ontario, and south to California, Nevada, New Mexico, Nebraska, and Iowa; 54 (xvi). Astragalus alpinus L. Alpine Milkvetch. Perennial, caulescent, 2-30 cm tall, from a subterranean caudex and rhizomatous cau- dex branches; pubescence basiflxed; stems decumbent to ascending; stipules 1.5-8 mm long, at least the lowermost connate-sheath- ing; leaves 3-15 cm long; leaflets 15-26, 6-20 mm long, 2-10 mm wide, ovate to el- liptic or oblong, retuse to roimded, strigu- lose above and below; peduncles 3-15 cm long; racemes 5- to 17-flowered, the flowers erect to declined at anthesis, the axis 0.5-5 cm long in fruit; bracts 1-2.5 mm long; pedicels 0.5-2.3 mm long; bracteoles 0; ca- lyx 3.2-6.3 mm long, the tube 2-3.5 mm long, campanulate, strigulose, the teeth 1-3.2 mm long; flowers 9-12 mm long, pink-purple; pods pendulous, stipitate, the stipe 2-5 mm long, the body oblong- lanceolate in outline, 10-17 mm long, 1.5-4 mm thick, strigulose, semibilocular; ovules 5-11. Aspen and coniferous woods at 2430 to 2730 m in Grand (on Tavaputs Plateau) and Salt Lake (at Brighton) cos., and to be sought elsewhere; Alaska to Nova Scotia and south to Oregon, Nevada, New Mexico, Wisconsin and Vermont; Eurasia; 2 (0). Astragalus amphioxys A. Gray. Crescent Milkvetch. Perennial (rarely flowering the first year), subacaulescent to shortly caules- cent, 2-20 cm tall, from a weak caudex; pubescence malpighian; stems lacking or up to 20 cm long, the internodes often con- cealed by stipules; stipules 2-13 mm long, all distinct or the lowermost sometimes connate-sheathing; leaves 2-13 mm long; leaflets (1) 5-21, 3-20 mm long, 1-9 mm wide, elliptic to obovate or oblanceolate, obtuse to acute, strigose on both sides; pe- duncles 2-15 (20) cm long; racemes 2- to 13-flowered, the flowers ascending at an- thesis, the axis 1-6.5 cm long in fruit; bracts 2.5-8 mm long; pedicels 0.6-2.5 mm long; bracteoles 0-2; calyx 6.3-14.2 mm long, the tube 5.2 mm long cylindric, strigose, usually purplish; the teeth 1.1-4.5 mm long, sub- ulate; flowers 16.5-31 mm long, pink- purple, rarely white; pods ascending, sessile, 1.5-5 cm long, 5-12 mm thick, usually curved, mostly dorsiventrally compressed, unilocular, strigose; ovules 42-70. Two rather feeble and sympatric varieties are present. 1 Banner twice as long as the calyx or less; keel 14.3-19 mm long A. amphioxys var. amphioxys Banner more than twice as long as the calyx; keel 17-25 mm long A. amphioxys var. vesperttnus Sept. 1978 Welsh: Utah Flora, Fab ace ae 263 Var. amphioxys. [A. shortianus var. (?) minor A. Gray; Xyhpfiacos arnphioxys (A. Gray) Rydb. X. aragalloides Rydb.; X. mela- nocalyx Rydb.; A. amphioxys var. melanoca- lyx (Rydb.) Tidestr.; A. marcus-jonesii Munz].— Desert shrub, pinyon-juniper, and less commonly in mountain brush commu- nities at 670 to 1830 m in Emery, Garfield, Grand, Iron, Kane, San Juan, Washington, and Wayne cos.; Nevada, Arizona, and New Mexico; 133 (xxxiii). Var. vespertinus (Sheldon) M. E. Jones [A. vespertinus Sheldon]. Desert shrub, pinyon-juniper, and rarely in mountain brush communities at 670 to 1530 m in Emery, Garfield, Grand, Kane, San Juan, Washington, and Wayne cos.; Colorado, Arizona, Nevada, and New Mexico; 104 (xl). Astragalus ampullarius S. Wats. Gumbo Milkvetch. [Tragacantha ampullaria (S. Wats.) Kuntze; Phaca ampullaria (S. Wats.) Rydb.]. Perennial, shortly caulescent, 2-28 cm tall, from a deep subterranean caudex; pubescence basifixed; stems prostrate- ascending, radiating; stipules 2-6 mm long, at least the lowermost connate-sheathing; leaves 3-14 cm long; leaflets 7-15 (19), 4-15 mm long, 2-12 mm wide, obovate, rounded to emarginate, strigose on both sides or glabrous above; peduncles 0.5-9.5 cm long; racemes 5- to 30-flowered, the flowers ascending at anthesis, the axis 1.2-13 cm long in fruit; bracts 1.5-3 mm long; pedicels 1-3 mm long; bracteoles 2; calyx 4.8-7.5 mm long, the tube 4.2-6 mm long short-cylindric, black strigose, the teeth 0.5-1.5 mm long, broadly triangular; flow- ers 13.5-22 mm long, pink-purple with white wing-tips, or ochroleucous; pods as- cending-erect, stipitate, the stipe 9-19 mm long, the body ovoid to subglobose, inflated, 12-20 mm long, 8-11 mm thick, sub- unilocular, glabrous or nearly so; ovules ca. 12. Clay soils of the Chinle and Tropic (?) Shale formations at 970 to 1650 m in Kane (west of the Cockscomb) and Washington COS.; Mohave and Coconino cos., Arizona. The gumbo milkvetch is certainly one of the most unusual of the vast array of spe- cies in Utah. Its propensity for the variegat- ed shales of the Chinle Formation near Ka- nab (the type locality) has long been known. The hypogeous caudex and short, prostrate-ascending stems, which persist in rosette form with marcescent stems and pods of previous years circular-reclining, bleached and skeleton-like, are quite unlike any of the taxa within the region. Phases of A. eremiticus, q.v., with inflated ovoid-ob- long pods resemble A. ampullarius, but the caudex is superficial, the stipules are all dis- tinct, and the stems commonly are erect; 11 (iii). Astragalus argophyllus Nutt. ex Torr. & Gray. Silver-leaved Milkvetch. Perennial, acaulescent to subacaulescent, 1.5-12 cm tall, arising from a superficial caudex; pu- bescence basifixed; stems obsolete or to 10 cm long, prostrate; stipules 2-10 mm long, all distinct; leaves 1.5-12 cm long; leaflets 7-21, 2-12 mm long, 1-6 mm wide, elliptic, oblanceolate or obovate, acute to obtuse, pilose above and below; peduncles to 9 cm long; racemes 1- to 6-flowered, the flowers ascending at anthesis, the axis httle elongat- ing in fruit; bracts 1.8-6.5 mm long; pedi- cels 1.2-3.8 mm long, bracteoles 0-2; calyx 9-16.8 mm long, the tube 6.5-11.8 mm long, cylindric, pilose with mixed black and white hairs, the teeth 1.5-6 mm long, lin- ear; flowers 15-25 mm long, pink-purple; pods ascending, sessile, 1.5-3.7 mm long, 5-13 mm thick, ovoid-acuminate, unilocu- lar, strigose or rarely villous, the valves not obscured; ovules 25-43. Three rather dis- tinctive varieties are present. 1. Flowers 15-17.5 mm long, the keel 12-15.2 mm long; pods curved-ellipsoid, densely silky strigose, 3-4 times longer than broad A. argophyllus var. panguicensis Flowers 18-25 mm long, the keel 15.9-21 mm long; pods various, variably pubescent, mostly 2-3 times longer than broad 2 2(1). Petals bright pink-purple, the flowers 22-25 mm long; plants of grasslands, streambanks, and lake shores A. argophyllus var. argophyllus 264 Great Basin Naturalist Vol. 38, No. 3 Petals dull pink-purple or pale, the flowers 18-22.5 mm long; plants of sage- brush and mountain brush communities A. argophylltis var. martinii Var. argophyllus. [Xylophacos argophxjUus (Nutt.) Rydb.; A. uintensis M. E. Jones, in part; X. uintensis (M. E. Jones) Rydb.]. Meadows, stream banks, and lake shores at 1400 to 1970 m in Box Elder, Cache, Car- bon, Salt Lake, San Juan, Sanpete, Summit, Utah, Wasatch, and Weber cos.; Nevada, Idaho, and Wyoming; 22 (ii). Var. martinii M. E. Jones. [A. argo- phyllus var. cnicensis M. E. Jones; A. agr- ophyllus var. pephragmenoides Barnebyj. Sagebrush, mountain brush, aspen, and spruce-fir communities at 1700 to 2600 m in Beaver, Carbon, Duchesne, Emery, Grand, Iron, Juab, Millard, Piute, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, and Wayne cos.; Idaho, Wyoming, and Colorado; 54 (xi). Var. panguicensis (M. E. Jones). M. E. Jones; [A. chamaeleuce var. panguicensis M. E. Jones; A. panguicensis (M. E. Jones) M. E. Jones; Batidophaca sabinarum Rydb.; A. sahinarum (Rydb.) Barneby]. Ponderosa pine, aspen-douglas fir-limber pine, white fir, sagebnish, and pinyon-juniper commu- nities at 2130 to 2900 m in Garfield, Iron, Kane and Washington cos.; Arizona; 50 (ix). Astragalus aretioides (M. E. Jones) Barn- eby. Cushion orophaca. [A. sericoleucus var. aretioides M. E. Jones]. Perennial, pulvinate caespitose, from a branching caudex; pu- bescence malpighian; stems almost entirely concealed by stipules; stipules 3.5-7 mm long, connate-sheathing, hyaline, glabrous dorsally or nearly so; leaves 0.6-2 cm long, palmately trifoliolate, the leaflets 3-7.5 mm long, 1.2-1.8 mm wide, spatulate to elliptic, acute, silvery-strigose on both sides; pe- duncles 7-15 mm long; racemes 2- or 3- flowered, the flowers ascending, the axis very short; bracts 2-3 mm long; pedicels 1-1.5 mm long; bracteoles 0; calyx 3.3-4.2 mm long, the tube 2.1-2.3 mm long, camp- anulate, densely long-strigose, the teeth 1.2-2 mm long; flowers 6-8 mm long, pink- purple (rarely white); pods ascending, ses- sile, 4-5 mm long, 1.2-2 mm thick, densely hairy, unilocular. Volcanic ash "barrens" at 1769 m in Daggett Co.; Montana and Wyoming; 2 (0). Astragalus asclepiadoides M. E. Jones. Milkweed Milkvetch. [Jonesiella asclepia- doides (M. E. Jones) Rydb.]. Perennial, cau- lescent, 7-62 cm tall, arising from a usually superficial caudex; pubescence basifixed; stems glabrous, erect; stipules 2-15 mm long, all distinct; leaves simple, 1.5-6.5 cm long, 1-5.5 cm wide, ovate, orbilcular, or cordate, obtuse to rounded or retuse, glabrous; peduncles 0.5-4.5 cm long; ra- cemes 2- to 12-flowered, the flowers as- cending, the axis 0.4-2.5 cm long in fruit; bracts 1-5 mm long; pedicels 1-5 mm long; bracteoles usually 2; calyx 10-17 mm long, the tube 8.3-13 mm long, cylindric, strigose with black hairs, the teeth 1.5-3.8 mm long, linear to subulate; flowers 17-27 mm long, suffused purple or almost ochroleucous; pods erect-ascending, stipitate, the stipe 10-21 mm long, the body ovoid or ovoid-el- lipsoid, inflated, 25-35 mm long, 11-16 mm thick, unilocular, glabrous; ovules ca 40. Sa- line desert shrub vegetative types on Man- cos Shale, Tropic Shale, Carmel, Moenkopi, Arapien, and Duchesne River formations at 1250 to 1900 m in Carbon, Duchesne, Emery, Garfield, Grand (type from Cisco), Sanpete, Sevier, Uintah, and Wayne cos.; Colorado. This singular selenophyte was first collected by M. E. Jones (1923) ". . . on sand-bars along the Price River. . . ." in Sep- tember of 1888, but the locality is appar- ently unique, for it is commonly distributed on the clay soils of the Mancos Shale in that vicinity; 42 (xvi). Astragalus australis Fischer. Subarctic Milkvetch. (A. aboriginum Richards., for a complete list of synonyms, see Barneby 1964.) Perennial, caulescent, 6-20 cm tall, from a superficial caudex; pubescence basi- flxed; stems pubescent, erect; stipules 2-7 cm long, at least the lowermost connate- sheathing; leaves 2-7 cm long, sessile; leaf- lets 7-15, 3-22 mm long, 1-7 mm wide, el- liptic, acute, villous to glabrate on both sides; peduncles 2-8.5 cm long; racemes 2- to 30-flowered, compact and ascending at anthesis, the axis 1-10 cm long in fruit; bracts 1.2-5 mm long; pedicels 0.8-3.5 mm Sept. 1978 Welsh: Utah Flora, Fab ace ae 265 long; bracteoles 0; calyx 3.7-6.4 mm long, the tube 2.1-3.9 mm long, campanulate, vil- lous, the teeth 1-3 mm long, subulate; flow- ers 7.5-12.6 mm long, ochroleucous or suf- fused with pink, the wing petals bilobed apically; pods pendulous, stipitate, the stipe 2.5-6 mm long, the body obliquely and nar- rowly elliptic in outline, 13-27 mm long, 3-6 mm wide, semibilocular, glabrous; ovules 8-16. The subarctic milkvetch is one of the poorly known species within Utah. It is recorded from only three collections. The plants grow on calcareous ridges with grasses and shrubs at elevations of from 1750 to 3050 m in Piute (Marysvale, 10000 ft., Jones in 1899, POM), Emery, and Wasatch cos.; Alaska east to Gaspe, and south to Oregon, Nevada, Colorado, and South Dakota; 3 (ii). Astragalus barnebyi Welsh in Welsh, At- wood, & Reveal. Bameby Milkvetch. [A. desperatus var. conspectus Bameby]. Pe- rennial, acaulescent or subacaulescent, 1.5-5 cm tall, from a branching caudex; pu- bescence basifixed; stems 0-5 cm long, mostly obscured by stipules; stipules 2-7 mm long, at least some connate-sheathing; leaves 1.5-5 cm long; leaflets 7-17, 3-9 mm long, 0.9-3.2 mm wide, elliptic to oblan- ceolate, acute to obtuse, strigose on both sides; peduncles 0.5-5.2 cm long; racemes 2- to 8-flowered, the flowers ascending at anthesis, 0.5-2.5 cm long in fruit; bracts 2-4 mm long; pedicels 0.5-1.5 mm long; calyx 6.1-7.7 (8.4) mm long, the tube 5.2-6.5 mm long; short-cylindric, pilose with mixed black and white hairs, the teeth 0.9-1.7 mm long, subulate; flowers 12.2-15 mm long, pink-purple or bicolored; pods declined, sessile or short-stipitate, ovoid-el- lipsoid, curved, 12-19 mm long, 5-6 mm thick, subunilocular, long silky-pilose; ovules ca. 20. Pinyon-juniper woods and mixed desert shrublands on platy shales of the Car- mel or on sandstones of Jurassic and Cre- taceous ages at 1430 to 1830 m in eastern Garfield and Wayne cos.; Navajo Co., Ari- zona. The Bameby milkvetch is a near con- gener of A. desperatus, q.v., from which it differs in the larger size of flowers and parts, and in the usually more compact habit of growth; 10 (iv). Astragalus beckwithii Torr. & Gray. Beckwith Milkvetch. Perennial, caulescent, 5-40 (70) cm tall, from a branching caudex; pubescence basifixed; stems decumbent to ascending or erect; stipules 2-10 mm long, all distinct; leaves 2-15 cm long; leaflets (7) 11-27, 3-17 (25) mm long, 2-9.6 mm wide, orbicular to obovate, obtuse to refuse, glabrous to glabrate on both sides; pe- duncles 3-15 cm long; racemes 7- to 16- flowered, the flowers ascending at anthesis, the axis 1-7 cm long in fruit; bracts 1-7 mm long; pedicels 1-3.5 mm long; brac- teoles 2; calyx 7-9.5 mm long, the tube 3.5-5.7 mm long, short-cylindric, sparingly strigose to glabrous, the teeth 2.5-4.4 mm long, subulate; flowers 14.5-21 mm long, ochroleucous to whitish or pink-purple; pods ascending to declined, stipitate, the stipe (gynophore) 1.5-5 mm long, the body obliquely leathery, unilocular; ovules 18-41. Two distinctive varieties are known in Utah. 1. Flowers pink-purple or bicolored; plants of western Beaver, Juab, and Millard cos ^- beckwithii var. purpureus Flowers ochroleucous, concolorous; plants of central to northwestern Utah A. beckwithii var. beckwithii Var. beckwithii. [Tragacantha beckwithii (Torr. & Gray.) Kuntze; Phaca beckwithii (Torr. & Gray.) Piper; Phacomene beck- withii (Torr. & Gray.) Rydb.] Juniper- pinyon, sagebrush, bvmch-grass, and moun- tain brush communities at 1330 to 2300 m in Beaver, Carbon, Box Elder, Davis, Mil- lard, Morgan, Piute (US), Salt Lake, Sevier, Tooele, Utah, and Weber cos.; Nevada. Plants of var. beckwithii are in- distinguishable in anthesis from those of A. oophorus var. caulescens, with which it is sympatric in part of its range (q.v.); 61 (v). Var. purpureus M. E. Jones. [A. arte- misiarum M. E. Jones; Phaca artemisiarum (M. E. Jones) Rydb.; Phacomene artemisia- 266 Great Basin Naturalist Vol. 38, No. 3 rum (M. E. Jones) Rydb.]. Pinyon-juniper bracteoles 0-2; calyx 3.5-9.6 mm long, gib- and cool-desert shrublands at 1370 to 2200 bous-saccate, the tube 2.8-5.7 mm long, m in western Beaver, Juab, Millard, and obliquely campanulate, sparingly strigose, Tooele cos.; Nevada; 24 (5). the teeth 1-3 mm long, subulate; flowers Astragalus hisukatus (Hook.) A. Gray. 8-18 mm long, ochroleucous, whitish, or Two-grooved Milkvetch. Perennial, caules- pink-purple, the keel-tip usually purple; cent, 15-70 cm tall, from a caudex; pu- pods pendulous, stipitate, the stipe about bescence basifixed; stems erect; stipules equaling the calyx tube, the body ellipsoid, 2.5-10 mm long, at least the lowermost 6.5-20 mm long, 2-4.5 mm thick, dorsiven- connate-sheathing; leaves 3-13.5 cm long; trally compressed, bisulcate, unilocular, stri- leaflets (7) 15-35, 5-33 mm long, 1.5-11 gose or glabrous; ovules 5-15. The two- mm wide, lance-oblong to oblong, elliptic grooved milkvetch is an ill-scented primary or oblanceolate 2-16 cm long; racemes 25- indicator of selenium in most areas, but to 80-flowered, the flowers declined at an- non-scented populations are known. Three thesis, the axis 3-25 cm long in fruit; bracts distinctive varieties are present in Utah. 2.5-6 mm long; pedicels 1-3.5 mm long; 1. Flowers bright pink-purple; calyx purple; plants of Daggett, Uintah and Grand cos A. hisulcatus var. hisulcatus — Flowers ochroleucous or whitish, the keel often purple-tipped; calyx ochro- leucous, whitish, or greenish; distribution various 2 2(1). Flowers 8-11 mm long; body of pod 6.5-9.5 mm long, prominently retic- ulate; plants usually montane in Utah A. hisulcatus var. haydenianus — Flowers 10-18 mm long; body of pod 10-17 mm long, smooth or faintly reticulate; plants usually not montane A. hisulcatus var. major Var. hisulcatus. [Phaca hisulcata Hook.: Tragacantha hisulcata (Hook.) Kuntze; Di- holcos hisulcatus (Hook.) Rydb.]. River ter- races at 1530 to 2130 m in Daggett, Grand, and Uintah cos.; Alberta east to Manitoba and south to Idaho, New Mexico, and Kan- sas; 6 (ii). Var. haydenianus (A. Gray) Bameby. [A. haydenianus A. Gray ex Brand.; Trag- acantha haydeniana (A. Gray) Kuntze; Di- holcos haydenianus (A. Gray) Rydb.]. Sage- brush-mountain brush communities at 2130 to 2600 m in Carbon, Duchesne, San Juan, Sanpete, Sevier, Uintah, and Utah cos.; Wyoming, Colorado, New Mexico, and Ari- zona; 16 (v). Var. major (M. E. Jones) Welsh comb, nov. based on A. haydenianus var. major M. E. Jones Zoe 2: 240. 1891. [A. scohatina- tulus Sheldon; Diholcos scohatinatulus (Sheldon) Rydb.]. Pinyon-juniper, sagebrush, mountain brush, and salt desert shrub at 1530 to 2300 m elevation in Beaver, Gar- field, Juab, Kane, Rich, Sanpete, Sevier, and Washington cos.; Wyoming, Colorado, and Arizona. This taxon was included within an expanded var. hisulcatus by Bameby (1964), who considered it as being "taxonomically negligible." However, var. major differs in about the same manner and degree as do other varieties in Astragalus. It is the domi- nant type within Utah. A population of var. major from north of Glendale in western Kane Co., lacks the characteristic odor of selenium commonly associated with all vari- eties of A. hisulcatus; 24 (vii). Astragalus bodinii Sheldon. Bodin Milk- vetch. [Phaca hodinii (Sheldon) Rydb.; A. dehilis sensu authors, not A. Gray (?); A. yukonis M. E. Jones; P. yukonis (M. E. Jones) Rydb.; A. hodinii var. yukonis (M. E. Jones) B. Boi.]. Perennial, caulescent, 15-80 cm long, from a superficial or buried cau- dex; pubescence basifixed; stems straggling on other plants; stipules 1-7 mm long all connate-clasping; leaves 2.5-8.5 cm long; leaflets 7-17, 2-12 mm long, 1-7 mm wide, oblanceolate to obovate, ovate or elliptic, rounded to emarginate or acute, strigose be- neath, glabrous above; peduncles 1.5-10 cm Sept. 1978 Welsh: Utah Flora, Fabaceae 267 long; racemes 3- to 15-flowered, the flowers ascending at anthesis, the axis 0.5-9 cm long in fruit; bracts 0.5-2.5 mm long; pedi- cels 0.7-2.2 mm long; bracteoles 0; calyx 4.5-5.2 mm long, the tube 3.4-3.8 mm long, campanulate, black-strigose, the teeth 1.5-1.8 mm long, subulate; flowers 9.5-10.2 mm long, pink-purple; pods ascending to spreading, stipitate, the stipe (gynophore) to 1 mm long, the body ellipsoid, 5.5-10 mm long, 3-4.5 mm thick, somewhat trigonous, unilocular, strigose; ovules 2-10. Wet mead- ows at 2000 to 2200 m in Rabbit Valley near Loa, Wayne Co. and along Otter Creek in Piute and Sevier cos.; Alaska east to Newfoundland and south to Alberta and Manitoba, and from Wyoming, Nebraska, and Colorado. Of the great number of spe- cies present in Utah, only a handful are true mesophytes. Included in that category are A. agrestis, A. argophyllus var. argophyllus, A. bodinii, A. canadensis, and A. diver- sifolius. They grow almost exclusively in meadows or other sites which are moist through much of the year. The habitat has been exploited to an extent almost unknown in more arid situations and is seldom ex- plored botanically because of the exploita- tion, fencing, and private ownership. De- spite that exploitation, A. bodinii has persisted from its initial discovery by Lester F. Ward (602, US, BRY) in Rabbit Valley, Utah on 18 August, 1875. Possibly A. bo- dinii will yet be found in other meadow- lands of the state; 8 (iii). Astragalus brandegei T. C. Porter in Port. & Coult. Brandegee Milkvetch. Pe- rennial, though sometimes flowering as an annual, caulescent, 5-35 (40) cm long, from a branching caudex; pubescence basifixed; stems prostrate-spreading, very slender; stip- ules 1.5-5 mm long, at least the lowermost usually connate-clasping; leaves 2-11.5 cm long; leaflets 5-15, 6-27 mm long, 0.5-2.6 mm wide, linear-filiform to narrowly ob- long, acute to obtuse, strigose beneath, glabrous above; peduncles 2.5-14 cm long, very slender; racemes 1- to 5-flowered, the flowers ascending at anthesis, the axis 0.5-6 cm long in fruit; bracts 1-2 mm long; pedi- cels 1.2-4 mm long; bracteoles 2; calyx 2.7-4 mm long, the tube 1.8-2.5 mm long. campanulate, black-strigose, the teeth 0.9-2 mm long, subulate; flowers 4.5-6 mm long, ochroleucous or tinged violet; pods pen- dulous to ascending, sessile or subsessile, the body obovoid to oblong-ellipsoid, 10-18 mm long, 3.5-5 mm thick, slightly dorsi- ventrally compressed, semibilocular, stri- gose. Volcanic gravels in mixed shrublands or pinyon-juniper at 1650 to 2430 m in Carbon, Emery (US), Garfleld (US), Piute, and Wayne cos.; Colorado, New Mexico, and Arizona. This is a cryptic plant that is seldom collected, probably because of its in- conspicuous, tiny flowers, slender peduncles, and slender prostrate stems; 4 (iii). Astragalus bryantii Barneby. Bryant Milkvetch. Perennial, caulescent, 35-80 cm tall, from a caudex; herbage basifixed; stems erect; stipules 3-8 mm long, all distinct; leaves 4-12 cm long; leaflets 11-21, 5-15 mm long, 2-12 mm broad, ovate, obovate, or broadly elliptic, emarginate to truncate, strigose beneath, glabrous above; peduncles 4-10 cm long; racemes 12- to 22-flowered, the flowers spreading-declined at anthesis, the axis 6-13 cm long in fruit; bracts 2-3.5 mm long; pedicels 0.5-3 mm long; brac- teoles 0-2; calyx 7-9.3 mm long, the tube 5.1-6.4 mm long, short-cylindric, strigose, the teeth 1.6-3.4 mm long, subulate; flowers 14-19.5 mm long, pink-purple or bicolored; pods spreading-ascending, sessile or sub- sessile, the body linear to linear-lanceolate in outline, 1.5-3 cm long, 3-5 mm thick, laterally compressed, bilocular, strigose, to glabrous; ovules 22-27. Talus slopes in mixed desert shrub communities at 770 to 1070 m in Lost Eden and West Creek canyons. Glen Canyon, San Juan Co.; Grand Canyon, Arizona. The low elevation reaches of Glen Canyon are flooded almost to the 1128.5 m level by the water of Lake Powell, and it seems probable that the Bryant milkvetch is extinct in Utah. The plants simulate A. len- tiginosus var. palans, but differ in pod char- acters; 1 (0). Astragalus callithrix Barneby. Callaway Milkvetch. Perennial, subacaulescent, 2-11 cm tall, from a caudex; pubescence basi- fixed; stems lacking or to 10 cm long, pros- trate, the internodes often concealed by stipules; stipules 2-5 mm long, all distinct. 268 Great Basin Naturalist Vol. 38, No. 3 leaves 2-11 cm long; leaflets 9-21, 2-13 mm long, 1.5-10 mm wide, obovate, sub- orbicular, or lanceolate, obtuse to trtmcate or emarginate, villous on both sides; pe- duncles 2-8 (12) cm long; racemes 5- to 15- flowered, the flowers ascending at anthesis, the axis 0.5-6 cm long in fruit; bracts 3-7.5 mm long; pedicels 1-1.5 mm long; brac- teoles 0-2; calyx 6.8-13.3 mm long, the tube 5.5-10.8 mm long, cylindric, villous- pilose, purplish, the teeth 1-3.2 mm long; flowers bright pink-purple, 16-26 mm long; pods ascending-spreading, sessile, oblong- ovoid, 10-20 mm long, 5-7.5 mm thick, dorsiventrally compressed, curved, long hairy (the valves not obscured), unilocular; ovules 24-34. Sandy flats and dunes in mixed desert shrublands at 1550 to 1625 m in western Millard Co.; Nye Co., Nevada; a Great Basin endemic; 3 (0). Astragalus calycosus Torr. ex S. Wats. Torrey Milkvetch. Perennial, low, acaules- cent, 1-12 cm tall, from a branching cau- dex; pubescence malpighian; stems lacking or to 2 cm long, the internodes concealed by stipules; stipules 1.5-6 mm long, all dis- tinct; leaves 1-8 (12) cm long; leaflets (1) 3-13, 2-19 mm long, 1-7 mm wide, ob- ovate, oblanceolate, or elliptic, obtuse to acute, silvery strigose on both sides; pe- duncles 0.5-10 cm long, rarely longer; ra- cemes 1- to 8-flowered, the flowers ascend- ing to spreading at anthesis, the axis 0.2-2.5 cm long in fruit; bracts 0.5-2 mm long; pedicels 0.7-3 mm long; bracteoles 0-2; ca- lyx 5-8.5 mm long, the tube 4-6.7 mm long, campanulate to shortly-cylindric, stri- gose, the teeth 1-4.2 mm long, subulate; flowers 10-16.5 mm long, varicolored, och- roleucous to shades of pink and purple, with white or pale wing-tips, the wings bi- lobed apically; pods ascending, sessile, nar- rowly oblong, usually curved, 8-25 mm long, 3-45 mm thick, laterally compressed, bilocular, strigose. Rather widespread and distinctive, the plants of Torrey milkvetch are separable into three varieties, two rare and one common. 1. Leaves with 5-13 leaflets along a rachis usually more than 1 cm long; scapes erect-ascending, usually over 7 cm long; raceme axis usually more than 2 cm long; plants of Washington Co A. calycosus var. scaposus Leaves with 3-13 leaflets along a rachis less than 1 cm long; scapes ascend- ing to decumbent, 1-7 cm long; raceme axis less than 2 cm long; distribu- tion various 2 2(1). Leaflets 7-13, mostly 2-6 mm long; alpine plants of extreme west-central Utah rare A- calycosus var. mancus Leaflets 3-7, mostly 5-19 mm long; plants of lower elevations, widespread, except in the northeastern third of Utah A. calycosus var. calycosus Var. calycosus. [Tragacantha calycosa (Torr.) Kuntze; Hamosa calycosa (Torr.) Rydb.; A. brevicaulis A. Nels.]. Mixed desert shrublands, pinyon-juniper, and ponderosa pine communities at 1430 to 2730 m in Beaver, Box Elder, Emery, Garfield, Iron, Juab, Kane, Millard, San Juan, Sanpete, Se- vier, Tooele, Utah, Washington and Wayne COS.; Wyoming, Idaho, Nevada, California, and Arizona; 92 (xviii). Var. mancus (Rydb.) Barneby. [Hamosa manca Rydb.; A. mancus (Rydb.) Wheeler.] Ridgetops at 2650 to 3660 m in Deep Creek Mts., Juab Co.; Nevada; 5 (i). Var. scaposus (A. Gray) M. E. Jones. (A. scaposus A. Gray; H. scaposa (A. Gray) Rydb.; A. candicans Greene. Ridgetops at ca. 2730 m in Beaverdam Mts., Washington Co.; Nevada, Arizona, Colorado, and New Mexico; 1 (0). Astragalus canadensis L. Canada Milk- vetch. Perennial, caulescent, 15-120 cm tall, erect or ascending; stipules 3-12 mm long or more, at least the lowermost connate- sheathing; leaves 5-30 cm long; leaflets 13-35, 10-52 mm long, 6-16 mm wide, lan- ceolate, lance-oblong, or elliptic, obtuse to emarginate, strigose on both sides or Sept. 1978 Welsh: Utah Flora, Fabaceae 269 glabrous above; peduncles 4-22 cm long; racemes many-flowered, the flowers spread- ing-declined in anthesis, the axis 2.5-16 cm long in fruit; bracts 1.5-10 mm long; pedi- cels 0.5-3.5 mm long; bracteoles 0-2; calyx 4.6-10.5 mm long, the tube 4-8.5 mm long, short-cylindric, strigose, the teeth 1.2-4.4 mm long, subulate or triangular; flowers 13.5-17.5 mm long, ochroleucous; pods erect, sessile or subsessile, cylindroid, 10-20 mm long, 2.9-5.2 mm thick, bilocular, stri- gose or glabrous; ovules 16-28. Two vari- eties of Canada milkvetch are known from Utah. These are peculiar among our many low-elevation species in flowering in mid- summer. 1. Pods and ovaries glabrous, terete at maturity, not sulcate dorsally; calyx teeth 2.5-4.1 mm long; plants mostly 40-120 cm tall A. canadensis var. canadensis Pods and ovaries pubescent, sulcate dorsally at maturity; calyx teeth mostly 1-2.5 mm long; plants 10-50 cm tall A. canadensis var. brevidens Var. brevidens (Gand.) Barneby. [A. mor- tonii f. brevidens Gand.]. Meadows, stream banks, lake shores, and hillsides at 1830 to 2300 m in Box Elder, Cache, Daggett, Rich, Summit, and disjunctly in Washington cos.; Montana, Idaho, Oregon, California, Ne- vada, Wyoming, and Colorado; 10 (iii). Var. canadensis. [A. carolinianus L.]. Stream terraces and lake shores at 1370 to 1600 m in Cache, Piute (Rydberg & Carlton 6967, 20 July 1905, Sevier River below Marysvale, US), Salt Lake, Utah, and Wasatch cos.; British Columbia east to On- tario and south to Washington, New Mexi- co, Texas, Louisiana, Alabama, and South Carolina; 6 (i). Astragalus ceramicus Sheldon. Painted Milkvetch. [Phaca picta A. Gray; A. pictus (A. Gray) A. Gray, not Boiss. & Gaill.; A. pictus var. foliolosus A. Gray; Tragacantha picta (A. Gray) Kuntze; A. angustus var. pictus (A. Gray) M. E. Jones; A. angustus var. ceramicus (Sheldon) M. E. Jones; A. pictus var. angustus M. E. Jones, type from Montezuma Canyon; A. ceramicus var. jonesii Sheldon; A. angustus (M. E. Jones) M. E. Jones; A. pictus var. magnus M. E. Jones, type from Silver Reef.]. Perennial, caulescent, 3-40 cm tall, from elongate rhi- zome-like caudex branches and deeply-bur- ied caudex; pubescence malpighian, stems sprawling to erect; stipules 1.5-9 mm long, at least some connate-sheathing; leaves 2-17 cm long; leaflets 3-13 or only one, the ter- minal continuous with the rachis, 3-30 mm long, 0.5-3 mm wide, filiform to narrowly oblong, obtuse to retuse or acute; peduncles 0.7-7.5 cm long; racemes 2- to 15-flowered (rarely more), the flowers ascending to de- clined at anthesis, the axis 1-12 cm long in fruit; bracts 1-2.5 mm long; pedicels 0.7-3.1 mm long; bracteoles 0; calyx 3.1-4.2 mm long, the tube 2.1-3.3 mm long, camp- anulate, strigose, the teeth 1-2.4 mm long, subulate; flowers 6.3-9.5 mm long, dull purplish to pink, or rarely whitish; pods pendulous, stipitate, the stipe 1-3.3 mm long, the body bladdery-inflated, ellipsoid to ovoid, 12-30 mm long, usually mottled, glabrous, imilocular; ovules 12-29. Sandy soils in pinyon-juniper, sagebrush, stream bank, grassland, and mixed desert shrub communities at 1270 to 2360 m in Beaver, Emery, Garfield, Grand, Juab, Kane, San Juan, Tooele, Uintah, Wayne, and Washing- ton COS.; Colorado, New Mexico, Arizona. Our materials belong to var. ceramicus, 40 X. Astragalus chamaeleuce A. Gray in Ives. Cidada Milkvetch. [Phaca pygmaea Nutt.; Tragancantha pyginaea (Nutt.) Kuntze; A. cicadae var. laccoliticus M. E. Jones, type from Henry Mts.; A. pijgmaeus var. laccoli- ticus (M. E. Jones) M. E. Jones.]. Perennial, acaulescent to subacaulescent, 2-10 cm tall, from a caudex; pubescence malpighian; stems lacking or to 6 cm long and prostrate, the intemodes mostly obscured by the sti- pules; stipules 2-7 mm long, all distinct; leaves 2-10 cm long; leaflets 5-17, 4-15 mm long, 2-10 mm wide, obovate to oblan- ceolate, obtuse to truncate or emarginate. 270 Great Basin Naturalist Vol. 38, No. 3 strigose on both sides; peduncles 1-8 cm long; racemes 2- to 11-flowered, the flowers spreading-ascending, the axis 0.9-2 cm long in fniit; bracts 2-5 mm long; pedicels 1-3.5 mm long, short-cylindric, strigose, the teeth 1.5-2.9 mm long; flowers ochroleucous or tinged pm-plish to pink-pm-ple; pods ascend- ing, sessile, oblong-ovoid or ellipsoid, 2-4 cm long, 7-16 mm thick, the fleshy valves ca. 3 mm thick, shrinking in ripening, the papery exocarp ultimately lustrous, separat- ing from the veins beneath and appearing quite smooth, mottled, strigose, unilocular; ovules 37-46. Juniper-pinyon, sagebrush, mixed desert shrub, and grassland commu- nities at 1530 to 2130 m in Daggett, Du- chesne, Emery, Garfleld, Grand, and Uintah COS.; Colorado and Wyoming. The cicada milkvetch is the only subacaulescent species of Astragalus of its type with malpighian hairs known from the Uinta Basin. To the south of there it is rare, but mingles with both A. amphioxys and A. cymboides, and mature fruit is necessary for positive identi- fication of A. chamaeleuce from those en- tities; 29 (x). Astragalus chloodes Barneby. Grass Milk- vetch. Perennial, acaulescent or sub- acaulescent, 5-24 cm tall, from a branching caudex; pubescence malpighian; stems ob- scured by stipules; stipules 2-8 mm long, all usually connate-sheathing; leaves simple, 1-13 (17) cm long, 1-3 mm wide, flat or in- volute, strigose on both sides; peduncles 2-9 cm long; racemes loosely 7- to 23-flowered, the flowers ascending at anthesis, the axis 4.5-24 cm long in fruit; bracts 2-4.5 mm long; pedicels 1-2.5 mm long; bracteoles 0; calyx 4.5-8.5 mm long, the tube 2-3 mm long, campanulate, strigose, the teeth 2.5-5.2 mm long, rigid-spreading; flowers 6.2-8.2 mm long, pink-purple; pods erect or ascending, sessile, obliquely lanceolate or oblong in outline, curved, 8-12 mm long, 1.7-3 mm wide, glabrous or strigose, uni- locular; ovules 4-8. Entrada Sandstone (or less commonly on Navajo Sandstone) hog- backs and cuestas in pinyon-juniper and mixed desert shrub communities at 1450 to 1700 m in Uintah Co.; endemic. The grass milkvetch simulates a grass, not only in its narrow leaves, but also in the flowers in bud which resemble grass spikelets; 11 (ii). Astragalus cibarius Sheldon. Browse Milkvetch. [A. webberi var. cibarius (Shel- don) M. E. Jones; Xylophacos cibarius (Shel- don) Rydb.; A. arietinus M. E. Jones, type from Cedar City.]. Perennial, caulescent, 6-30 cm tall, from a branching caudex; pubescence basifixed; stems decumbent to ascending; stipules 3-8 mm long, all dis- tinct; leaves 3.5-10 cm long; leaflets 11-19, 4-17 mm long, 2-13 mm wide, obovate, ob- long, oblanceolate, obtuse or retuse, strigose beneath, glabrous above; peduncles 3-8 cm long; racemes 4- to 14-flowered, subcapitate at early anthesis, the flowers spreading- ascending, the axis 0.5-2.7 cm long in fruit; bracts 2-4 mm long; pedicels 1-2.5 mm long; bracteoles 0-2; calyx 6.4-9.2 mm long, the tube 5-7 mm long, cylindric, stri- gose, the teeth 1.4-2.5 mm long; flowers 15-19 mm long, pink-purple with white wing-tips or whitish to ochroleucous and variously tinged; pods ascending, subsessile, ellipsoid to oblong, 17-32 mm long, 7-10 mm thick, curved to almost straight, stri- gose, unilocular, woody or stiffly leathery; ovules 27-32. Mountain brush, sagebrush, juniper-pinyon, and mixed desert shrub communities at 1630 to 2430 m in Beaver, Box Elder, Cache, Davis, Garfield, Iron, Juab, Millard, Morgan, Salt Lake, Sanpete, Sevier, Summit, Tooele, Utah, Wasatch, Washington, and Weber cos.; Nevada, Idaho, Montana, Wyoming, and Colorado. The browse milkvetch is allied to A. ensi- fonnis and A. malacoides, but is easily dis- tinguished from them both morphologically and geographically; 108 (xiv). Astragalus cicer L. Chickpea Milkvetch. [Cystiu7n cicer (L.) Stev.]. Perennial, caules- cent, 20-70 cm tall or more, from a branch- ing caudex; pubescence basifixed; stems pro- strate to ascending; stipules 2-8 mm long, at least the lowermost connate-sheathing; leaves 4-21 cm long; leaflets 17-27 (31), 5-40 mm long, 2-14 mm wide, lance-ellip- tic to oblong, acute to obtuse, strigose on both sides or glabrous above; peduncles 4-12 cm long; racemes densely 10- to 30- flowered, the flowers ascending at anthesis, the axis 2-7 cm long in fruit; bracts 2-6.5 mm long; pedicels 0.3-1.5 mm long; brae- Sept. 1978 Welsh: Utah Flora, Fabaceae 271 teoles 0; calyx 6-9 mm long, the tube 5-6 mm long, short-cylindric, strigulose, the teeth 1.6-3 mm long; flowers 12.5-16.5 mm long, ochroleucous; pods ascending, or by crowding, spreading, subsessile, the body ovoid or subglobose, strongly inflated, 6-14 mm long, 6-10 mm thick, pilose, bilocular. Introduced forage plant, escaping and per- sisting in pinyon-juniper, sagebrush, and as- pen communities at 1770 to 2170 ni in San- pete Co., and to be expected elsewhere; indigenous to Europe; 3 (0). Astragalus coltonii M. E. Jones. Colton Milkvetch. Perennial, caulescent, 10-75 cm tall, from a branching caudex; pubescence basifixed; stems erect or ascending; stipules 1-7 mm long, all distinct; leaves 2-10 cm long; leaflets 3-19, or the uppermost leaves simple, 3-20 mm long, 0.3-3 mm wide, lin- ear, narrowly oblong, or ovate, strigose on both sides; peduncles 4-30 cm long; ra- cemes loosely 5- to 30-flowered, the flowers spreading-declining at anthesis, the axis 3-28 cm long in fruit; bracts 0.5-3.2 mm long; pedicels 0.8-2.5 mm long; bracteoles 0; calyx 4.5-8 mm long, the tube 4-6.7 mm long, cylindric, strigose, purplish, the teeth 0.6-2.3 mm long, broadly subulate; flowers 12-19 mm long, pink-purple; pods pend- ulous, stipitate, the stipe 4-11 mm long, the body oblong to oblanceolate in outline, 19-35 mm long, 3-6 mm wide, strongly lat- erally flattened, glabrous, unilocular; ovules 14-20. Two allopatric and distinctive varie- ties of Colton milkvetch occur in Utah. 1. Leaves all odd-pinnate, with 9-19 leaflets, the terminal one jointed on all leaves; plants of Grand and San Juan cos A. coltonii var. moabensis Leaves odd-pinnate, or the uppermost simple, with 3-9 leaflets, the terminal one continuous with the rachis; plants not from Grand or San Juan cos. A. coltonii var. coltonii Var. coltonii. [Homalobus coltonii (M. E. Jones) Rydb.; A. coltonii var. aphijUus M. E. Jones, type from Richfield.]. Bunchgrass, salt desert shrub, pinyon-juniper, and mountain brush communities at 1470 to 2300 m in Carbon (type from Castle Gate), Emery, Garfield, Kane, Sevier, and Wayne cos.; en- demic; 38 (ix). Var. moabensis M. E. Jones. [A. coltonii var. foliosus M. E. Jones ex A. Eastwood; Homalobus canovirens Rydb., type from LaSal Mts.; A. canovirens (Rydb.) Barneby.]. Pinyon-juniper and less commonly mountain brush communities at 1400 to 2300 m in Grand and San Juan (type from Monticello) COS.; Colorado, New Mexico, and Arizona; 21 (viii). Astragalus consobrinus (Barneby) Welsh comb. nov. based on A. castaneifonnis var. consobrinus Barneby Amer. Midi. Nat. 41: 496. 1949. Bicknell Milkvetch. Perennial, sometimes flowering the first year, acaules- cent, 1-5 cm tall, the caudex branches ob- scured by persistent leaf bases and stipules; herbage malpighian; stems essentially lack- ing; stipules 3-7 mm long, all distinct; leaves 1-5 cm long; leaflets 3-11, 2-8 mm long, 1.5-4.2 mm wide, obovate to oblan- ceolate or orbicular, rounded to obtuse or acute, strigose on both surfaces; peduncles 0.5-3 cm long; racemes 2- to 10-flowered, the flowers ascending at anthesis, the axis to 1 cm long in fruit; bracts 1.5-3.5 mm long; pedicels 1-2 mm long; bracteoles 0; calyx 5.5-8.9 mm long, the tube 4.1-6.8 mm long, cylindric, strigose, the teeth 1.4-1.7 mm long, subulate; flowers 10-15.5 mm long, ochroleucous suffused with purple; pods as- cending, sessile, obliquely ovoid or lance- ovoid, 11-19 mm long, 3-8 mm thick, stri- gose, unilocular; ovules 18-33. Sagebrush- grasslands and pinyon-juniper communities at 1830 to 2200 m in Emery, Garfield, Piute, Sevier, and Wayne cos.; endemic. The Bicknell milkvetch has been compared with and treated within an expanded A. castaneifonnis, with which it is similar in some salient morphological features. The similarity appears to be at least in part coincidental, and the true relationship of the Bicknell plants might lie with A. cym- boides with which it is apparently allied. In any event, it seems best to treat A. con- sobrinus at specific level; 7 (v). 272 Great Basin Naturalist Vol. 38, No. 3 Astragalus convallarius Greene. Lesser Rushy Milkvetch. Perennial, caulescent, 10-60 cm tall, from a subterranean caudex; pubescence basifixed; stems erect or ascend- ing; stipules 2-7 mm long, at least the lowermost connate-sheathing; leaves 2-11 cm long; leaflets, when present, 3-13, the uppermost leaves reduced to the rachis, mostly 5-30 mm long, 0.5-4 mm wide, lin- ear to oblong or oblanceolate; peduncles 1-14 cm long; racemes 3- to 25-flowered, the flowers ascending to declined at an- thesis, the axis 2-20 cm long in fruit; bracts 0.5-2.3 mm long; pedicels 1-8 mm long; bracteoles 0-2; calyx 4-6.3 mm long, the tube 3.4-5.4 mm long, campanulate, black strigose, the teeth 0.5-1.4 mm long, triangu- lar-subulate; flowers 6.5-12 mm long, och- roleucous or variously tinged or veined with purple; pods pendulous to spreading, sessile, linear to narrowly oblong, straight, 13-50 mm long, 2.3-4 mm thick, laterally com- pressed, strigose, unilocular; ovules 11-26. This is possibly the most widespread of the milkvetch species within Utah, definitely known from all but two counties. Two allo- patric varieties are present. Pods 25-50 mm long, 2.3-3.3 mm wide; plants widespread but not in west- ern Iron and Washington cos A. convallarius var. convallarius Pods 13-26 mm long, 3.4-4 mm wide; plants of western Iron and Washing- ton cos A. convallarius var. finirimus Var. convallarius. [Homalobiis campestris Nutt. ex Torr. & Gray: Tragacantha camp- estris (Nutt.) Kuntze; A. serotinus var. campestris (Nutt.) M. E. Jones Phaca con- vallaria (Nutt.) Greene; H. junceiis Nutt. ex Torr. & Gray; A. junceus (Nutt.) A. Gray, not Ledeb. ex Spreng.; T. juncea (Nutt.) Kuntze; A. diversifolius var. junceus (Nutt.) M. E. Jones; A. diversifolius var. roborum M. E. Jones; A. junciformis A. Nels.; H. jun- ciformis (A. Nels.) Rydb.; A. junceus var. at- tenuattis M. E. Jones, type from Price]. Mixed desert shrub, sagebrush, pinyon- juniper, mountain brush, ponderosa pine, and aspen communities at 1400 to 2900 m in all except Grand and Rich cos., and likely there also; Idaho and Montana south to Nevada and Colorado; 139 (xxiv). Var. finitimus Barneby. Pinyon-juniper and sagebnish communities at 1700 to 2270 m in western Iron and Washington cos.; Lincoln Co., Nevada; 7 (ii). Astragalus cottamii Welsh. Cottam Milk- vetch. Perennial, sometimes flowering the first year, acaulescent or subacaulescent, 1.2-8 cm tall, from a branching caudex; pubescence basifixed; stems lacking, or 0.5-6 cm long, the internodes mostly ob- scured by stipules; stipules 2-6 mm long, all distinct; leaves 1.2-8 cm long; leaflets (5) 9-19 (21), 2-9 mm long, 1-4.2 mm wide. elliptic to oval or oblanceolate, acute to ob- tuse, strigose on both sides or glabrate above; peduncles 0.7-7 cm long; racemes 3- to 9-flowered, the flowers ascending at an- thesis, the axis 0.5-2 cm long in fruit; brac- teoles 0-2; calyx 6.2-8 mm long, the tube 4.8-6.7 mm long, cylindric, strigulose, purplish, the teeth 1.2-2 mm long, subulate; flowers 11-17 mm long, pink-purplish or bicolored; pods spreading-descending, ses- sile, curved, oblong to oblong-lanceolate in outline, triquetrous, the dorsal suture sul- cate, bilocular, strigose, usually purple- blotched. Rimrock and ledges of Cedar Mesa, Kayenta, and Entrada sandstones, and in the sandy canyons cut from them in pinvon-juniper woods and blackbnish at 1300 to 1400 m in San Juan Co.; Navajo Co., Arizona. The Cottam milkvetch was not distinguished among the limited collec- tions available to Barneby (1964), where the few specimens were thought to represent a portion of the range variability within A. monurnentalis. The rather larger set cur- rently available allows segregation on the basis of easily discernible diagnostic features of flowers and fruit; 11 (ix). Astragalus cronquistii Barneby. Cronquist Milkvetch. Perennial, caulescent 1.5-4 cm long, from a subterranean caudex; pubescence basifixed; stems decumbent- Sept. 1978 Welsh: Utah Flora, Fab ace ae 27.'^ ascending; stipules 2-6 mm long, all dis- tinct; leaves 1.5-4.5 cm long; leaflets 7-15, 6-23 mm long, 1.5-4 mm wide, oblong to narrowly elliptic, retuse to truncate, strigose beneath, glabrate above; peduncles 2-6.5 cm long; racemes 6- to 20-flowered, the flowers declined at anthesis, the axis 1.5-8.5 cm long in fruit; bracts 0.6-1.2 mm long; pedicels 1.5-2.5 mm long; bracteoles 0; ca- lyx 3.8-5.3 mm long, the tube 3.3-4 mm long, campanulate, strigose, the teeth 0.5-1.3 mm long, triangular; flowers 8-9 mm long, pink-purple; pods declined-pend- ulous, sessile or subsessile, the body narrow- ly elliptic to oblanceolate in outline, 13-30 mm long, 3-4.8 mm wide, trigonous, grooved dorsally, strigose, semibilocular. Blackbnish community, on Cutler Forma- tion, at 1430 m in San Juan Co.; endemic; 4 (ii). Astragalus cymboides M. E. Jones. Ca- noe Milkvetch. [Xylophacos cijmhoides (M. E. Jones) Rydb.; A. amphioxijs var. cym- bellus M. E. Jones]. Perennial, acaulescent or subacaulescent, 2.5-8 cm tall, from a simple or branched caudex; pubescence malpighian; stems lacking or 0.5-3 cm long, the internodes mostly obscured by stipules; stipules 3-8 mm long, all distinct; leaves 2.5-8 cm long; leaflets (1-3) 5-13, 3-13 mm long, 2-9 mm wide, obovate, elliptic or oblanceolate, obtuse to acute, pubescent on both surfaces; peduncles 2-8 cm long; ra- cemes 3- to 9-flowered, the flowers ascend- ing at anthesis, the axis 0.5-2 cm long in fruit; bracts 1-4 mm long; pedicels 0.7-2.5 mm long; bracteoles 0; calyx 7.6-10.2 mm long, the tube 5.9-8 mm long, cylindric, strigose, the teeth 1-2.3 mm long, subulate; flowers 15-18.5 mm long, ochroleucous or suffused purplish, or pink-purple; pods as- cending, sessile, oblong to oblong- elliptic in outline, straight, 17-30 mm long, 6-9.5 mm wide, laterally compressed, the valves stiffly papery or cellular-spongy, the exocarp in time exfoliating, strigose, uni- locular; ovules 39-57. Salt desert shrub and pinyon-juniper communities at 1600 to 2300 m in Carbon, Emery, and Grand cos.; en- demic. The small-flowered phases of canoe milkvetch approach A. consobrinus and the large-flowered phases are difficult to dis- tinguish from A. amphioxys. Further work is indicated; 36 (vii). Astragalus desereticus Bameby. Deseret Milkvetch. Perennial, acaulescent or sub- acaulescent, 4-11 cm tall, from a caudex; pubescence basifixed; stems to 5 cm long, the nodes mostly obscured by stipules; stip- ules 3.5-7 mm long, all distinct; leaves 4-11 cm long; leaflets 11-17, 2-10 mm long, el- liptic to obovate, short-acuminate to acute, strigulose-villosulous on both sides; pe- duncles 2-5.5 cm long; racemes 5- to 10- flowered, the flowers ascending at anthesis, the axis 0.5-2 cm long in fruit; bracts 3-6 mm long; pedicels 2-3 mm long; bracteoles 0-2; calyx 8.4-11.5 mm long, the tube 6.2-7.5 mm long, cylindric, villous, the teeth 2-4 mm long, subulate; flowers 18-22.5 mm long, ochroleucous (?), the keel purple-tipped; pods ascending, sessile or substipitate, ovoid-ellipsoid, curved, 10-12 mm long, 4-5 mm thick, densely hirsute with lustrous hairs; ovules 14-16. Sagebrush or pinyon-juniper communities at 1830 to 2000 m at Indianola, Sanpete Co.; endemic. This is an obscure taxon, apparently with affinities to A. argophyllus var. martinii, but differing in its densely long-hirsute and small pods. The Deseret milkvetch differs further in its more strongly graduated petals (banner 18-22.5 mm, but the keel 12-13.3 mm long). More material is necessary to make adequate predictions as to the true nature of this entity. It has not been col- lected in more than half a century; 1 (0). Astragalus desperatus M. E. Jones. Rim- rock Milkvetch. [Batidophaca desperata (M. E. Jones) Rydb.; Tiiim desperatum (M. E. Jones) Rydb.; A. desperatus var. petrophilus M. E. Jones, type from San Rafael Swell; B. petrophila (M. E. Jones) Rydb.]. Perennial, acaulescent or subacaulescent, 1-12 cm tall, from a branching caudex; pubescence basi- fixed; stems to 8 cm long, the internodes of- ten obscured by stipules; stipules 1.5-7 mm long, at least some connate-sheathing; leaves 1-12 cm long; leaflets 7-17, 2-13 mm long, 1-5 mm wide, elliptic to oblanceolate or obovate, acute to obtuse, strigose on both sides or glabrate above; peduncles 0.5-13 cm long; racemes 3- to 18-flowered, the flowers ascending to declined at anthesis. 274 Great Basin Naturalist Vol. 38, No. 3 the axis 0.4-13 cm long in fruit; bracts 1.5-5 mm long; pedicels 0.5-1.4 mm long; bracteoles 0-2; calyx 3.5-6 mm long, the tube 2.5-4 mm long, campanulate, strigose- pilose, the teeth 0.8-2.6 mm long, subulate; flowers 6-9 mm long, pink-purple or bico- lored; pods declined to deflexed, sessile or short-stipitate, the stipe (gynophore) to 1.2 mm long, the body obliquely ovoid to lance-ellipsoid, curved, 6-19 mm long, 3-6 mm thick, hirsute with lustrous hairs, uni- locular; ovules 16-28. Mixed desert shrub and pinyon-juniper communities, often on rimrock, at 1130 to 1900 m in Emery, Gar- field, Grand, Kane, San Juan, and Wayne COS.; Colorado and Arizona; Colorado Plateau endemic. There is some variation within A. desperatus. The deep pink-purple, small flowers with dark purplish calyces dis- played by plants of the interior and north- ern half of the San Rafael Swell are the most distinctive of the variant types. These apparently form the basis of var. petrophila M. E. Jones. Further collections might dem- onstrate the necessity for recognition of that variety; 66 (xxi). Astragalus detritalis M. E. Jones. Debris Milkvetch. [Hornalohus detritalis (M. E. Jones) Rydb.; A. spectabilis C. L. Porter, not Schischk., the type from southwest of Vernal]. Perennial, acaulescent, 0.5-8 cm long, from a branching caudex; pubescence malpighian; stems essentially lacking; stip- ules 3-10 mm long, at least some (usually all) connate-sheathing; leaves 0.5-8 cm long, simple or leaflets 3-7, 3-30 mm long, and 0.5-2.5 mm wide, narrowly oblanceolate to linear, spinulose-tipped, strigose on both sides; peduncles 1-9 cm long; racemes 2- to 8-flowered, the flowers ascending at an- thesis, the axis 0.2-3.8 cm long; bracts 2.5-7 mm long; pedicels 0.5-2.5 mm long; brac- teoles 0-2; calyx 5-9.6 mm long, the tube 3.1-5.4 mm long, campanulate, strigose, the teeth 1.6-4.7 mm long, subulate; flowers 13-20 mm long, pink-purple; pods erect to steeply ascending, sessile, narrowly oblong, straight to curved, 15-38 mm long, 2-3.5 mm wide, laterally compressed, mottled, strigose, unilocular; ovules 15-24. Pinyon- juniper and mixed desert shrub communities at 1650 to 1950 m in Duchesne (type from southwest of Duchesne) and Uintah cos.; Rio Blanco Co., Colorado; a Unita Basin en- demic; 10 (ii). Astragalus diversifolius A. Gray, Mead- ow Milkvetch. [Hornalohus orthocarpus Nutt. ex Torr. & Gray, not Boiss.; A. camp- estris var. diversifolius (A. Gray) Macbr.; A. junceus var. orthocarpus (Nutt.) M. E. Jones; A. junceus var. diversifolius (A. Gray) M. E. Jones; A. convallarius var. diversifolius (A. Gray) Tidestr.; A. ihapensis M. E. Jones; Atelophragma ibapense (M. E. Jones) Rydb.]. Perennial, caulescent, 20-50 cm long, from a subterranean to superficial caudex; pu- bescence basifixed; stems prostrate to as- cending; stipules 1-3 mm long, the lowest connate-sheathing; leaves 1.5-7 cm long; leaflets 1-7, 4-47 (67) mm long, narrowly elliptic, linear, oblanceolate or lanceolate, the uppermost often simple, strigose on both sides; peduncles 2-15 cm long; race- mes 3- to 8-flowered, the flowers ascending at anthesis, the axis 0.5-3 mm long in fruit; bracts 0.7-2.5 mm long; pedicels 1.8-4 mm long; bracteoles 0; calyx 4.4-6.7 mm long, the tube 3.2-4.7 mm long, campanulate, strigose, the teeth 1-2 mm long, subulate; flowers 8-13.5 mm long, greenish- white, of- ten tinged with purple; pods ascending to declined, sessile or substipitate, the body narrowly oblong, 10-17 mm long, 2.7-4 mm wide, strongly compressed, strigose, unilocular; ovules 10-16. Moist, often saline, meadows at 1340 to 1700 m in Juab (Juab, Goodding 1084, GH, NY, US, June 10, 1902), and Tooele (type of A. ihapensis from Deep Creek Mts., M. E. Jones s.n., 23 June 1891, POM); Idaho. This is a poorly known entity in Utah, and it is not known from recent collections. Possibly it is extinct in Utah; 1 (0). Astragalus drummondii Douglas ex Hook. Dnimmond Milkvetch. [Tragacantha drummondii (Douglas) Kuntze; Tium drum- mondii (Douglas) Rydb.]. Perennial, caules- cent, 25-60 cm tall, from a branching sub- terranean caudex; pubescence basifixed; stems erect or ascending; stipules 3-12 mm long, at least some connate-sheathing; leaves 4-13 cm long; leaflets 17-33, 4-33 mm long, 2-12 mm wide, oblong to oblanceo- late or obovate, obtuse to truncate or Sept. 1978 Welsh: Utah Flora, Fab ace ae 275 emarginate, villous-pilose beneath, usually glabrous above; peduncles 4-12 cm long; racemes 14- to 30-flowered, the flowers spreading-declined at anthesis, the axis 3-22 cm long in fruit; bracts 2-5 mm long; pedi- cels 1.5-5 mm long; bracteoles 0-2; calyx 7-12.5 mm long, the tube 4.7-8 mm long, short-cylindric, sparingly villous, the teeth 1.7-4.5 mm long, subulate; flowers 18-26 mm long, whitish to ochroleucous, the keel purple-tipped; pods pendulous, stipitate, the stipe 5-11 mm long, the body narrowly ob- long to oblanceolate in outline, 17-32 mm long, 3.5-5.5 mm thick, trigonous, sulcate dorsally, glabrous, bilocular; ovules 14-30. Pinyon-juniper, ponderosa pine, and moun- tain brush communities at 1530 to 2130 m in Beaver, Sevier, and Utah cos.; Alberta and Saskatchewan south to New Mexico; 12 (ii). Astragalus duchesnensis M. E. Jones. Duchesne Milkvetch. [Lonchophaca duchesnensis (M. E. Jones) Rydb.j. Per- ennial, caulescent, 15-35 cm tall, from a branching caudex; pubescence basifixed; stems straggling to ascending or erect; stip- ules 3-8 mm long, all distinct; leaves 2-10 cm long; leaflets 5-15, 3-20 mm long, 0.5-3 mm wide, linear to oblong or narrowly ob- lanceolate, obtuse to retuse, strigose on both sides or glabrate above, the uppermost leaf- let sometimes continuous with the rachis; peduncles 3-10.5 cm long; racemes 6- to 22-flowered, the flowers ascending at an- thesis, the axis 2.5-12 cm long in fruit; bracts 0.7-2 mm long; pedicels 0.8-2.2 mm long; bracteoles 0; calyx 3.6-5.5 mm long, the tube 3.1-4.3 mm long, campanulate, usually purple, the teeth 0.4- 1mm long, triangular; flowers 8.5-12.5 mm long, pink- purple with white wing-tips; pods declined sessile, oblong to narrowly oblanceolate in outline, 20-35 mm long, 3.3-5 mm thick, dorsiventrally compressed in the lower half, becoming laterally compressed in the distal portion, strigose, unilocular; ovules 21-31. Sand to heavy clay soils in mixed desert shrub and pinyon-juniper communities at 1450 to 1750 m in Duchesne and Uintah COS., endemic; 7 (ii). Astragalus eastwoodae M. E. Jones. East- wood Milkvetch. [A. preussii var. sulcatus M. E. Jones; Phaca eastwoodae (M. E. Jones) Rydb.; A. preussii var. eastwoodae (M. E. Jones) M. E. Jones.]. Perennial, short caulescent, 8-20 cm tall, from a branching caudex; pubescence lacking except on calyx, basifixed; stems 2-14 cm long, decumbent to ascending; stipules 2-6.5 mm long, all distinct; leaves 3-13 cm long; leaflets 13-25, 1-15 mm long, 1-5 mm broad, ellip- tic to lance-elliptic, oblanceolate or obo- vate, obtuse to truncate-emarginate, glabrous; pedimcles 2-10.5 cm long; bracts 1.5-4.5 mm long; pedicels 1.5-3.5 mm long; bracteoles 2; calyx 10-12.2 mm long, the tube 8-9.5 mm long, cylindric, purple, spar- sely black-strigose, the teeth 1.3-2.7 mm long, subulate; flowers 18-22 mm long, pink-purple; pods spreading to declined, stipitate, the stipe 1.5-4.5 mm long, the in- flated body oblong-ellipsoid, 14-26 mm long, 7-14.5 mm thick, the valves papery and straw-colored, unilocular, glabrous; ovules 20-38. Mixed desert shrub and pinyon- juniper communities at 1330 to 1830 m in seleniferous soils in Emery, Grand, San Juan, and Wayne cos.; Colorado. The East- wood milkvetch is closely allied to A. pre- ussii It differs mainly in the shorter stems and spreading-descending, thin-textured pods; 6 (ii). Astragalus emoryanus (Rydb.) Cory. Emory Milkvetch. [Hamosa emoryana Rydb.]. Annual or winter annual, caulescent, 4-45 cm long, from a slender taproot; pu- bescence basifixed; stems prostrate; stipules 1.5-3.6 mm long, all distinct; leaves 1-4.5 cm long; leaflets 11-19, 2-10 mm long, 1-6 mm wide, oval-obovate to obcordate or ob- lanceolate, obtuse to retuse on all leaves, sparingly strigose on both sides or glabrate above; peduncles 2-6 cm long; racemes 2-to 10-flowered, the flowers spreading at an- thesis, the axis 0.3-2.5 cm long in fruit; bracts 0.5-2 mm long; bracteoles 0; calyx 3.6-6 mm long, the tube 1.9-3.5 mm long, campanulate, strigose, the teeth 1.3-2.5 mm long; flowers 7.3-11 mm long, pink-purple; pods declined to ascending, sessile, narrowly oblong, curved, 0.8-2.2 cm long, 2.2-4.3 mm wide, trigonous, glabrous, bilocular; ovules 10-16. Pinyon-juniper community at 1700 m in Kane Co. (Cockscomb); Arizona, 276 Great Basin Naturalist Vol. 38, No. 3 New Mexico, Texas, and Mexico. This is an obscure entity in Utah. It is very similar in most salient features with A. niittallianiis, from which it can be distinguished by the deciduous, straw-colored pods, merely stri- gose calyx teeth, retuse-obtuse leaflets on all leaves, and rounded keel-tip. This latter fea- ture is shared with A. nuttallianus var. niicranthifomiis, but that entity is known to occur in Kane Co. only east of the Cocks- comb; 1 (0). Astragalus ensiformis M. E. Jones. Pa- gumpa Milkvetch. [A. ursinus M. E. Jones, not A. Gray; Hamosa ensiformis (M. E. Jones) Rydb.; A. ensiformis var. gracilior Barneby; A. minthorniae var. gracilior (Bameby) Barneby.]. Perennial, caulescent, 8-45 cm tall, from a superficial to sub- terranean caudex; pubescence basifixed; stems decumbent to erect; stipules 4-10 mm long, all distinct; leaves 4-15.5 cm long; leaflets (5) 11-23, 6-24 mm long, 1.5-13.5 mm wide, ovate to oblong, obovate, or ob- lanceolate, obtuse to retuse, strigose (some- times sparsely so) beneath, strigose to glabrous above; peduncles 2.5-13 cm long; racemes 12- to 30-flowered, the flowers as- cending to declined at anthesis, the axis 3.5-13.5 cm long in fruit; bracts 2-6 mm long; pedicels 1-3.5 mm long; bracteoles 0-2; calyx 5.2-7.8 mm long, the tube 4.5-6.5 mm long; short-cylindric, pilosulose with black hairs, the teeth 1.2-2.5 mm long, subulate; flowers 13-17 mm long, purplish to pink-purple, the wing-tips pale to white; pods ascending to descending, sessile or sub- stipitate, the body narrowly oblong, curved, 15-30 mm long, 4-6 mm thick, subterete (compressed laterally when pressed), bilocu- lar, strigose to strigulose; ovules 24-36. Pinyon-juniper, sagebrvish, and blackbnish- larrea communities at 1230 to 2350 m in Washington Co.; Mohave Co., Arizona. The materials included herein as portions of an expanded A. ensiformis have been treated as belonging to A. ensiformis sens. str. and to A. ensiformis or A. minthorniae as var. gra- cilior. The var. gracilior was named by Barneby (Calif. Acad. Sci. Proc. 25:158. 1944) as a variety of A. ensiformis on the basis of plants taken near Vevo. Later Barn- eby (Amer. Midi. Naturalist' 55:493. 1956) transferred the variety to A. minthorniae (Rydb.) Jeps. The var. gracilior is supposed to differ fom A. ensiformis sens. str. by its erect (not short decumbent) stems, reflexed pedicels (not spreading-ascending), and by the sparingly long strigose (not shortly pilo- sulous) essentially pendulous (not ascending to erect) pods. With a rather substantial number of specimens now available the sup- posed diagnostic criteria fail as differential features. Stems are decumbent to ascending or erect, varying with each separate popu- lation. Peduncle position does not appear to be taxonomically important because of lack of correlation with other features. The posi- tion of the fruit is likewise not correlated with other characteristics. Pubescence size and shape forms a continuvun. The final fea- tures, involving the pods, are tenuous at best and almost impossible to demonstrate at worst. Each sub-population varies, the whole consisting of apparent recombination types, with A. ensiformis sens. str. the ex- treme in one series and var. gracilior, the extreme in another. Thus, A. ensiformis is treated herein as consisting of a series of polymorphic populations, not readily sepa- rable into infraspecific taxa. The status of A. minthorniae, shorn of this taxon, is beyond the scope of this work; 18 (iv). Astragalus episcopus S. Wats. Bishop Milkvetch. [Tragacantha episcopa (S. Wats.) Kuntze; Homalobus episcopus (S. Wats.) Rydb.; A. kaibensis M. E. Jones; Loncho- phaca kaibensis (M. E. Jones) Rydb.]. Pe- rennial, caulescent, rush-like, 20-45 cm tall arising from a subterranean caudex; pu- bescence basifixed; stems erect or ascending; stipules 2-13 mm long, all distinct; leaves 2-10 cm long, most of them reduced to the rachis, some with leaflets 3-13 in number, the leaflets 1-15 mm long, 0.5-2 mm wide, linear to elliptic or oblong, acute to obtuse or emarginate, strigose on both sides; pe- duncles 6-23 cm long; racemes very loosely 6- to 30-flowered, the flowers ascending in anthesis, the axis 3-30 cm long in fruit; bracts 1.3-3 mm long; pedicels 1.5-3.5 mm long; bracteoles 0-2; calyx 4.1-7 mm long, the tube 3.4-5.2 (6) mm long, short cylin- dric, always much longer than broad, suf- fused purplish or very pale, white-strigose, Sept. 1978 Welsh: Utah Flora, Fabaceae 277 the teeth 0.6-2.2 mm long, triangular to subulate; flowers 10-15.5 mm long, pale pink or whitish to pink-purple; pods pend- ulous, sessile or subsessile, the body oblong to lance-elliptic in outline, slightly curved to straight, 14-32 mm long, 4-8 mm wide, laterally compressed, glabrous to strigose, straw-colored or tinged or mottled purple, imilocular; ovules 16-26. Mixed desert shrub and pinyon-juniper communities often in clay or silty soils at 1270 to 1700 m in Emery, Garfleld, Kane, and Wayne cos. (type from southern Utah); Arizona. The Bishop milkvetch closely simulates A. lan- cearius q.v., with which it is sympatric in Kane Co. and in Arizona. The calyx of A. lancearius is black-strigose, not or seldom suffused with purple, with the tube cam- panulate and as broad as long or only some- what longer than broad, and the flowers are broader in proportion to length than in A. episcopus; 38 (xii). Astragalus eremiticus Sheldon. Hermit Milkvetch. [A. arrectus var. eremiticus (Shel- don) M. E. Jones]. Perennial, caulescent, 20-45 cm tall, from a branching commonly superficial caudex; pubescence basifixed; stems erect or ascending; stipules 3-11 mm long, 1.5-12 mm wide, ovate to oblong, el- liptic, or narrowly oblong, obtuse toretuse, strigose beneath, glabrate to glabrous above; peduncles 2.5-15 cm long; racemes 10- to 26-flowered, the flowers ascending to de- clined at anthesis, the axis 5-17 cm long in fruit; bracts 1.5-4 mm long; pedicels 0.7-3.5 mm long; bracteoles 0-2; calyx 4.5-8.7 mm long, the tube 4-6.8 mm long, short-cylindric, strigose, the teeth 0.7-2 mm long, triangular to subulate; flowers 12-18 mm long, ochroleucous, pink-purple, or merely tinged purplish; pods erect, stipitate, the stipe 6-15 mm long, the body oblong to ellipsoid or obliquely ellipsoid, 12-27 mm long, 3.5-8 mm thick, trigonous, glabrous, bilocular; ovules 17-32. Juniper-pinyon, live oak, and sagebrush communities at 1130 to 1830 m in Kane (west of Cockscomb) and Washington cos.; Arizona, Nevada, and Idaho. Plants with inflated and proportion- ately shorter pod bodies simulate the rare A. ampullarius which occupies clay soils at lower elevations; 13 (i). Astragalus eucosmus B. L. Robins. Ele- gant Milkvetch. [A. oroboides var. anierican- us A. Gray, not A. americanus (Hook.) M. E. Jones; Phaca elegans Hook.; A. elegans (Hook.) Sheldon, not Bunge; Atelophragma elegans (Hook.) Rydb.]. Perennial, caules- cent, 10-40 cm long, from a branching cau- dex; pubescence basifixed; stems erect to as- cending; stipules 1.5-6 mm long, at least some of the lower ones usually connate- sheathing; leaves 2-10 cm long; leaflets 9-15, 10-22 mm long, 4-7 mm broad, ob- tuse, strigose beneath, glabrous or glabrate above; peduncles 5-18 cm long; racemes 7- to 25-flowered, the flowers declined at an- thesis, the axis 3.5-15 cm long in fruit; bracts 1-3.5 mm long; pedicels 0.5-3 mm long; bracteoles usually 0; calyx 3.3-5.4 mm long, the tube 2.5-3.5 mm long, cam- panula te, strigulose, the teeth 0.9-1.6 mm long, subulate; flowers 6-8 mm long, purple or whitish; pods spreading to deflexed, ses- sile or subsessile, 8-12 mm long, 2.5-5 mm thick; laterally compressed, pilose, semi- bilocular; ovules 4-8. Woods, rare, north slope of Uinta Mountains, Summit Co., Utah; Alaska east to Nova Scotia and south to Colorado and Maine; 1 (0). Astragalus eurekensis M. E. Jones. Eu- reka Milkvetch. [Xylophacos eurekensis (M. E. Jones) Rydb.]. Perennial, acaulescent, 2-15 cm tall, the caudex branches obscured by persistent leaf bases; pubescence basi- fixed; stipules 3-11 mm long, all distinct; leaves 2-15 cm long; leaflets (3) 5-19, 3-35 mm long, 2-8 mm wide, elliptic to oblong, acute, gray or silvery strigose on both sur- faces; pedimcles 1-13 cm long, pilose; ra- cemes 3- to 7-flowered, the flowers ascend- ing at anthesis, the axis 0.2-2 cm long in fruit; bracts 4-8 mm long; pedicels 1.2-3 mm long; calyx 10.5-15.5 mm long, the tube 8.5-10.5 mm long, cylindric, pilose-vil- lous, the teeth 2.5-5.7 mm long, subulate; flowers 22-27 mm long, ochroleucous, faintly to strongly suffused with purple or rarely pink-purple; pods ascending, sessile, obliquely lance-ovoid, 15-40 mm long, 5-9 mm thick, villous-hirsute, unilocular; ovules 26-36. Sagebrush, pinyon-juniper, and mountain brush communities at 1370 to 2135 m in Beaver, Garfield, Iron, Juab, Mil- 278 Great Basin Naturalist Vol. 38, No. 3 lard, Sanpete, Sevier, Tooele, Utah, and Wasatch cos.; endemic. When, as rarely, the flower color is bright pink-purple this en- tity, lacking fruit, is exceedingly difficult to separate from A. newbernji (q.v.); 51 (xiv). Astragalus falcatus Lam. Russian Sickle Milkvetch. Perennial, caulescent, 40-90 cm tall, from a branching caudex; pubescence malpighian; stems ascending to erect, form- ing large clumps; stipules 2-12 mm long, at least some connate-sheathing; leaves 5-22 cm long; leaflets 19-37, 6-35 mm long, 1.5-10 mm wide oblong to elliptic or oblan- ceolate, acute to apiculate, strigose below, glabrous above, green on both sides; pe- duncles 6-17 cm long; racemes 20- to 70- flowered, the flowers delined at anthesis, the axis 3-20 cm long in fruit; bracts 2-5 mm long; pedicels 2-5 mm long, recurved in fruit; bracteoles 2; calyx 3.6-4.7 mm long, the tube 3-3.5 mm long campanulate, strigose, the teeth 0.5-1.2 mm long, triangu- lar; flower 9-11 mm long, greenish-white, sometimes suffused with purple; pods de- curved, subsessile, curved-oblong, 13-23 mm long, 2.5-4.5 cm wide, triangular, strigulose, bilocular, ovules 12-14. Introduced soil stabilization plant, established in Juab Co., and to be expected elsewhere in Utah; southeastern Europe. This is a robust per- ennial capable of surviving in harsh, heavy soils in the mountain brush zone of the state; 2 (ii). Astragalus flavus Nutt. ex Torr & Gray. Yellow Milkvetch. Perennial, caulescent, 5-30 cm tall, from a branching caudex; pubescence malpighian; stems decumbent to ascending or erect; stipules 2-10 mm long, all connate-sheathing; leaves 3-15 (18) cm long; leaflets (5) 9-21, 3-31 mm long, 0.5-6 mm wide, linear, narrowly oblong, or oblan- ceolate to ovate, obtuse to acute, silvery strigose (greenish) on both sides or glabrate to glabrous above; peduncles 3-23 cm long; racemes 6- to 30-flowered, the flowers as- cending at anthesis, the axis 2-12 cm long in fruit; bracts 1.5-5 mm long; pedicels 0.7-1.2 mm long; bracteoles 0; calyx 5.5-9.5 mm long, the tube 3-5.2 mm long, cam- panulate, strigose to pilose, the teeth 2-6 mm long, subulate; flowers 9-17 mm long, yellow to ochroleucous, whitish, lilac, or pink-purple; pods erect, sessile, oblong, 7-13 mm long, 3.5-5 mm thick, straight, dorsiventrally compressed, strigose, unilocu- lar; ovules 6-17. Three more or less dis- tinctive varieties, all primary selenophytes, are present in Utah. 2(1) Calyx shaggy long villous, the teeth 3-6 mm long, equaling or longer than the tube; flowers pink-purple; plants of Emery, Grand, Wayne, and Garfield (•OS A. flavus var. argiUosus Calyx strigose to short villous, the teeth 2-3 (4) mm long, shorter than the tube; flowers yellow to white or tinged purplish, rarely pink-purple; plants of various distribution 2 Flowers whitish or yellowish, sometimes tinged with purple, rarely pink- purple; keel 6-8 mm long; plants from central and southern counties A. flavus var. candicans Flowers cream to yellow; keel 8-10 mm long; plants from east-central to northeastern Utah ^- /^uus var. flavus Var. argiUosus (M. E. Jones) Barneby. [A. argillosus Jones; Cnemidophacos argil- losus (M. E. Jones) Rydb.] Mancos Shale, Summerville, Cedar Mountain, and Morri- son formations, on saline clays and silts with salt desert shrubs at 1230 to 1600 m in Emery (type from Green River), Garfield, Grand, and Wayne cos.; endemic; 23 (vi). Var. candicans A. Gray. [A. confertiflorus A. Gray, type from near St. George; Cne- midophacos confertiflorus (A. Gray) Rydb.]. Mancos and Tropic shales, Moenkopi, Chinle, and Kaiparowits formations and oth- er saline clays and silts at 900 to 2130 m in Garfield, Kane, San Juan, and Washington COS.; Arizona and Nevada. The var. candi- Sept. 1978 Welsh: Utah Flora, Fab ace ae 279 cans passes into var. flavus in central east- em Utah; 43 (vii). Var. flavus. [Cnemidophacos flavus (Nutt.) Rydb.; Tragacantha flaviflora Kiintze; A. confertiflorus var. flaviflorus (Kuntze) M. E. Jones]. Mancos Shale, Chinle, Moenkopi, Duchesne River, Uinta, and other formations composed of saline silts and clays in salt desert shrub and pin- yon-juniper communities at 1230 to 1730 m in Carbon, Daggett, Duchesne, Emery, Grand, San Juan, Sanpete, Sevier, and Uin- tah COS.; Arizona, New Mexico, Colorado, and Wyoming. Sheep poisoning attributable to var. flavus is known from the lower ele- vation portions of the Uinta Basin; 40 (xiii). Astragalus flexuosus (Hook.) Don. Pe- rennial, caulescent, 10-60 cm tall, from a branching caudex; pubescence basifixed; stems decumbent or ascending; stipules 1-7 mm long, at east the lowermost connate- sheathing; leaves 1.5-9 cm long; leaflets (5) 9-25, 3-19 mm long, 1-8 mm wide, linear or oblong to oblanceolate or obovate, ob- tuse to truncate or retuse, strigose to gla- brate beneath, usually glabrous above; pe- duncles 1.5-14 cm long; racemes 7- to 26- flowered, the flowers spreading at anthesis the axis 2.5-13 cm long in fruit; bracts 0.6-4.5 mm long; pedicels 0.7-3.5 mm long; bracteoles 0-2; calyx 3.3-5.8 mm long, the tube 2.4-4.3 mm long, campanulate, stri- gose, the teeth 0.5-1.7 mm long, subulate; flowers 7-11 mm long, pink-purple to dull purplish; pods descending to spreading, ses- sile or short-stipitate, the stipe 0.5-1.3 mm long, the body oblong to oblanceolate or el- liptic in outline, 8-24 mm long, 2.7-4.8 mm thick, subterete or variously somewhat flat- tened, strigose to glabrous, unilocular; ovules 12-25. Two rather distinctive varie- ties are present, both confined to eastern Utah. Calyx tube 2.4-2.7 mm long; pods sessile or nearly so; plants spreading- decumbent, in pinyon-juniper and mixed desert shrublands A. flexuosus var. diehlii Calyx tube 2.7-4.3 mm long; pods subsessile to shortly stipitate; plants of pinyon-juniper and mountain brush communities A. flexuosus var. flexuosus Var. diehlii (M. E. Jones) Barneby. [A. diehlii M. E. Jones; Phisophaca diehlii (M. E. Jones) Rydb.]. Salt desert shrub and pinyon-juniper communities at 1370 to 1670 m in Carbon, Emery, Grand, and Uintah COS.; Colorado; 12 (vii). Var. flexuosus. [Phaca flexuosa Hook.; Homalobus flexuosus (Hook.) Rydb.; Piso- phaca flexuosa (Hook.) Rydb.]. Pinyon- juniper and mountain brush communities at 1675 to 2135 m in Grand and San Juan COS.; British Columbia east to Ontario and south to New Mexico, Nebraska, and Min- nesota; 2 (0). Astragalus fucatus Barneby. Hopi Milk- vetch. [A. subcinereus sensu M. E. Jones, not Gray, q.v.]. Perennial, caulescent, 7-45 cm tall, from a subterranean to superficial caudex; pubescence basifixed; stems ascend- ing to erect or sprawling; stipules 1-5.5 mm long, the lowest connate-sheathing; leaves 2-12.5 cm long; leaflets 9-17, 3-20 (25) mm long, 0.5-2.5 mm wide, obtuse to re- tuse, strigose beneath, glabrous above; pe- duncles 1-6.5 cm long; racemes 9- to 22- flowered, the flowers ascending to declined at anthesis, the axis 2-11.5 cm long in fruit; bracts 0.8-2 mm long; pedicels 0.7-3.5 mm long; bracteoles 0; calyx 3.3-5.4 mm long, the tube 2.3-3.3 mm long, campanulate, strigose, the teeth 0.8-2.2 mm long; flowers 6.4-8.7 mm long, pink-purple; pods spread- ing to declined, sessile, bladdery-inflated, ovoid, ellipsoid or subglobose, 17-.32 mm long, 12-20 mm wide (when pressed), mot- tled, strigose, unilocular; ovules 21-32. Mixed desert shrub communities, usually in sandy soil, at 1330 to 1830 m in Garfield and San Juan cos.; Colorado, New Mexico, and Arizona; 18 (iv). Astragalus geyeri A. Gray. Geyer Milk- vetch. [Phaca annua Geyer, not A. annuus DC.]. Annual (rarely biennial), caulescent, 6-27 cm long, from a slender taproot; pu- 280 Great Basin Naturalist Vol. 38, No. 3 bescence basifixed; stems prostrate to as- cending or erect; stipules 1.5-4 mm long, all distinct; leaves 2-10.5 cm long; leaflets 3-13, 3-17 mm long, 1-5.2 mm wide, linear to oblong or narrowly elliptic, obtuse to re- tuse, strigose beneath and strigose to glabrous above; peduncles 0.6-1.5 cm long; racemes 2- to 8-flowered, the flowers as- cending at anthesis, the axis 0.8-1.5 cm long in fruit; bracts 0.7-2 mm long; pedi- cels 0.6-1.5 mm long; bracteoles 0-1; calyx 2.7-3.8 mm long, the tube 1.6-2.4 mm long, campanulate, strigose, the teeth 0.6-1.5 mm long; flowers 5-7.6 mm long, pale, suffused with purple, or pink-purple; pods spreading to declined, bladdery-inflated, obliquely ovoid, 15-24 mm long, 6-12 mm wide (when pressed), strigose, unilocular; ovules 7-18. Sandy soil in mixed desert shrub com- mimities at 1370 to 1830 m in Beaver, Dag- gett, Duchesne, Emery, Garfield, Grand, Juab, Millard, Salt Lake, Tooele, Uintah, and Wayne cos.; Oregon east to Wyoming and south to California, Nevada, and Colo- rado. Plants grown from seed in a green- house produced mature fruit and seeds in 58 days; 32 (iv). Astragalus gilviflorus Sheldon. Plains Orophaca. [A. triphtjllus Pursh, not Pallas; Tragacantha triphylla (Pursh) Kuntze; Oro- pliaca triphijlla (Pursh) Britt; Phaca caespi- tosa Nutt., not A. caespitosus Pallas; Phaca argophylla Nutt., not A. argophyllus Nutt. ex Torr. & Gray.]. Perennial, acaulescent, 1.5-13 cm tall, from a branching caudex; pubescence malpighian; stems entirely ob- scured by stipules; stipules 6-18 mm long, connate-sheathing; leaves 1-13 cm long, palmately trifoliolate, the leaflets 7-20 mm long, 2-7 mm wide, spatulate to elliptic, acute to obtuse silvery strigose on both sides; peduncles obsolete; racemes capitate, 1- to 3-flowered, the flowers erect, the axis very short in fruit; bracts 4.5-7.6 mm long, tridentate; pedicels 0-1.6 mm long; brac- teoles 0; calyx 9.3-18 mm long, the tube 10-14 mm long, cylindric, loosely villous, the teeth 1.6-4 mm long; flowers 17-28 mm long, whitish to ochroleucous; pods erect, sessile, ovoid-ellipsoid, 6-10 mm long, 2.5-5 mm thick, densely hairy, unilocular. Sagebrush community at about 2130 m in Summit Co.; Alberta east to Ontario and south to Wyoming and Nebraska; 1 (0). Astragalus hallii A. Gray. Hall Milk- vetch. [A. fallax S. Wats.; A. familicus Shel- don, not A. fallax Fischer; A. gracilentus var. fallax (S. Wats.) M. E. Jones; Pisophaca familica (Sheldon) Rydb.]. Perennial, caules- cent, 12-50 cm tall, from a subterranean caudex; pubescence basifixed; stems decum- bent to ascending or erect; stipules 1-7 mm long, at least the lowermost connate- sheathing; leaves 2-9 cm long; leaflets 11-2.3, 2-11 mm long, 1-7 mm wide, obo- vate to oblanceolate or elliptic, retuse to tnincate or obtuse, strigulose beneath, spar- ingly hairy or glabrous above; peduncles 3-9.5 cm long; racemes 9- to 25-flowered, the flowers spreading-declined at anthesis, the axis 1-7 cm long in fruit; bracts 1.5-5 mm long; pedicels 1.2-4 mm long; brac- teoles 0-2; calyx 6-7 mm long, the tube 5.2-6.2 mm long, short-cylindric, villosulous, the teeth 0.7-1.2 mm long, triangular; flow- ers 12.8-15 mm long, pink-purple; pods spreading to declined, short-stipitate, the stipe 1.5-3.5 mm long, the inflated body cylindroid to obliquely ovoid-ellipsoid, 19-27 mm long, 8-12 mm thick, strigulose, imilocular; ovules 20-37. Pinyon-juniper and mountain brush communities at 1600 to 2130 m in Garfield (?) and Kane cos.; Ari- zona and New Mexico. Our plants belong to var. fallax (S. Wats.) Bameby. They seem not to differ in any significant way from those of the Flagstaff Plateau in north-cen- tral Arizona; 3 (i). Astragalus hatniltonii C. L. Porter. Ham- ilton Milkvetch. Perennial, caulescent, 25-60 cm long, from a shallowly sub- terranean caudex; pubescence basifixed; stems erect; stipules 1.5-9.5 mm long, all distinct or rarely some shortly connate- sheathing; leaves 3-8 cm long, the up- permost (and sometimes the lowermost) simple, the others with leaflets 3-7, 10-47 mm long, 2-7 mm wide, elliptic to narrow- ly oblanceolate, obtuse to retuse, strigose on both sides, the terminal leaflet continuous with the rachis; peduncles 2.5-15.5 cm long; racemes 7- to 30-flowered, the flowers spreading-declined at anthesis, the axis 2-11 cm long in fruit; bracts 1-2.5 mm long; Sept. 1978 Welsh: Utah Flora, Fabaceae 281 pedicels 1.2-3 mm long; bracteoles 0-2; ca- lyx 8.2-11 mm long, light brown, the tube 6.5-9.2 mm long, cylindric, gibbous, stri- gose, the teeth 1.7-2.6 mm long, subulate; flowers 20-24 mm long, ochroleucous, con- colorous; pods pendulous, stipitate, the stipe 9-12 mm long, the body ellipsoid, 25-35 mm long, 4-7.5 mm thick, dorsiventrally compressed, the valves often brownish, stri- gose, unilocular; ovules 16-22. Duchesne River and Wasatch formations at 1600 to 1770 m in the juniper-pinyon community of western Uintah Co.; endemic. This is a mir- rored-image cogener of A. lonchocarpus (q.v.), which is not known from the Uinta Basin; 8 (iii). Astragalus harrisonii Barneby. Harrison Milkvetch. Perennial, caulescent, rush-like, 40-70 cm tall, from a subterranean caudex; pubescence basifixed; stems diffusely inter- branched, in clumps to 1 m wide or more; stipules 1-5 mm long, all distinct; leaves 1.5-6.5 cm long, the uppermost simple, with the terminal leaflet expanded and con- fluent with the rachis, the others with leaf- lets 3-9, 2-11 mm long, 0.5-1.5 mm wide, linear-elliptic, acute, strigose on both sides; peduncles 6-19 cm long; racemes loosely 3- to 15-flowered, the flowers ascending at an- thesis, the axis 5-40 cm long in fruit; bracts 0.5-1.1 mm long; pedicels 1.5-5.5 mm long; calyx 2.7-4.6 mm long, the tube 1.5-3.7 mm long, campanulate, strigose, the teeth 0.5-1.9 mm long, triangular; flowers 9-10.5 mm long, pink-purple; pods pendulous, stipitate, the stipe 3-4 mm long, the body narrowly ellipsoid, straight or curved, 17-28 mm long, dorsiventrally compressed, strigose to glabrous, unilocular; ovules 10-12. Pinyon-juniper community at 1650 m on Navajo Sandstone, Capitol Reef National Park, Wayne Co.; endemic. The Harrison milkvetch is a near cogener of A. nidularius (q.v.); 7 (iii). Astragalus henrimontanensis Welsh, nom. nov. based on Astragalus stocksii Welsh Great Basin Nat. 34:307. 1974, not A. stock- sii Benth. ex Bunge Astragal. Geront. 1:6. 1868. Dana Milkvetch. Perennial, acaules- cent, 4-15 cm tall, from a branching cau- dex, the branches clothed with coarse, per- sistent leaf bases; pubescence basifixed; stipules 3-8 mm long, all distinct; leaves 2.7-12.5 cm long; leaflets 7-17, 3-13 mm long, 1.5-6 mm wide elliptic to oblanceo- late, mucronate, acute to obtuse to truncate, strigose on both sides; peduncles 1.1-8 cm long; racemes 2- to 11-flowered, the flowers ascending at anthesis, the axis 0.3-2.2 cm long in fmit; bracts 1.8-5.5 mm long; pedi- cels 1.3-2.5 mm long; bracteoles 0-2; calyx 10.2-15 mm long, the tube 8.2-11.5 mm long, cylindric, strigulose, the teeth 1.9-3.5 mm long, subulate; flowers 15-23 mm long, ochroleucous, the wings and keel purple- tipped; pods ascending, sessile, lance-ovoid, slightly incurved, 22-35 mm long, 5-11 mm thick, somewhat dorsiventrally compressed, strigose, unilocular. Ponderosa pine, pinyon- juniper, and sagebmsh communities at about 2430 m in the Henry Mountains, Garfield Co.; endemic. The Dana Milkvetch was named to honor Dana L. Stocks, late chair- man of the Department of Botany at Brig- ham Young University. The species com- bines features of A. argophtjlhis var. panguicensis, as to pod shape and pu- bescence, and of A. eurekensis, as to habit and flower color; 11 (iv). Astragalus humistratus A. Gray, Ground- cover Milkvetch. [Batiodophaca humivagans Rydb.]. Perennial, caulescent, 7-80 cm long, radiating from a caudex; pubescence mal- pighian; stems prostrate; stipules 2-9 mm long, 0.5-6 mm wide, elliptic to oblong or oblanceolate, acute, strigose on both sides or glabrate above; peduncles 2-9 cm long; racemes 3- to 20-flowered, the flowers as- cending at anthesis, the axis 1-12 cm long in fruit; bracts 1.5-7 mm long; pedicels 0.4-2.2 mm long; bracteoles 0-2; calyx 4.5-8.8 mm long, the tube 2.4-4.1 mm long, campanulate, strigose, the teeth 1.4-5 mm long; flowers 7-11.5 mm long, greenish to ochroleucous, often suffused or veined purplish, or pink-purple; pods ascending to spreading, obliquely ovoid or oblong-ellip- soid, 6-14 mm long, 3.5-5.7 mm wide, vari- ously compressed, strigose, unilocular. Mountain brush, cool desert shrub, pinyon- juniper, and ponderosa pine communities at 1600 to 2430 m in Beaver, Garfield, Iron, Kane, and Washington cos.; Nevada, Ari- zona, and New Mexico. Our material be- 282 Great Basin Naturalist Vol. 38, No. 3 longs to var. humivagans (Rydb.) Barneby; 34 (xi). Astragalus iodanthus S. Wats. Humboldt River Milkvetch. [Tragacantha iodantha (S. Wats.) Kuntze; Xylophacos iodantha (S. Wats.) Rydb.; A. iodanthus var. dia- phanoides Barneby.]. Perennial, caulescent, 8-35 cm long, from a branching caudex; pubescence basifixed; stems prostrate to de- cumbent; stipules 2-6 mm long, all distinct; leaves 2-8 cm long; leaflets 7-21, 3-18 mm long, 2-12 mm wide, obovate to oblong or oblanceolate to elliptic, tnmcate to retuse, obtuse or mucronate, sparingly strigose to glabrous on both sides; peduncles 1-4.5 cm long, shorter than the leaf; racemes 7- to 17-flowered, the flowers ascending to spreading at anthesis, the axis 0.5-4.5 cm long in fruit; bracts 1-3 mm long; pedicels 0.3-2 mm long; bracteoles 2; calyx 5-8 mm long, the tube 3.4-5 mm long, short- cylindric, strigose, the teeth 1..3-3 mm long; flowers 12-15.5 mm long, pink-purple to pale; pods ascending to declined, sessile, the body curved through a half-circle, 20-40 mm long, 5-8 mm thick, dorsiventrally or trigonously compressed, obliquely lanceo- late, unilocular or semibilocular, strigose; ovules 14-30. Juniper-sagebrush community at about 1670 m in Box Elder Co.; Oregon, California, and Nevada; 1 (0). Astragalus iselyi Welsh. Isely Milkvetch. Perennial, caulescent, 8-25 cm tall, from a branching caudex; pubescence basifixed; stems ascending to erect; stipules 3-9 mm long, all distinct; leaves 3.2-8.5 cm long; leaflets (3) 5-13, 7-23 (37) mm long, 3-9 (16) mm wide, elliptic to rhombic, acute to mucronate, sparsely strigose to glabrate on both sides; peduncles 1.7-10 cm long; ra- cemes 7- to 20-flowered, the flowers spread- ing at anthesis, the axis 1-3 cm long in fruit; bracts 2-4.5 mm long; pedicels 0.8-2.5 mm long; bracteoles 0; calyx 6.7-10 mm long, the tube 5.5-6.3 mm long, cylin- dric, strigulose, the teeth 1.8-3.1 mm long, subulate; flowers 17-18 mm long, ochro- leucous, concolorous; pods spreading- declined, sessile or subsessile, inflated, sub- cylindric, 25-38 mm long, 10-15 mm thick, strigose, leathery, unilocular; ovules 38-44. Morrison and Paradox formations in pinyon- jimiper and salt desert shrub communities at 1530 to 1830 m on western foothills of the LaSal Mountains in Grand and San Juan COS.; endemic. The Isely milkvetch is closely allied to A. sahulosus (q.v.). Both are obli- gate selenophytes of gypsiferous clays and silts of the Grand River Valley; 7 (iii). Astragalus jejunus S. Wats. Starveling Milkvetch. [Tragacantha jejuna (S. Wats.) Kuntze; PJiaca jejuna (S. Wats.) Rydb.]. Pe- rennial, acaulescent, 1-4 cm tall, from a much-branched caudex; pubescence basi- fixed; stems obsolete, obscured by stipules and leaf bases; stipules 1.5-3 mm long, all connate-sheathing; leaves 1-4 cm long; leaf- lets 9-15, 1-5 mm long, 0.5-1 mm wide, linear to narrowly elliptic, obtuse to acute, involute, strigose on both sides; peduncles 0.5-3.5 cm long; racemes 3- to 7-flowered, the flowers spreading at anthesis, the axis 0.2-1 cm long in fruit; bracts 1-1.5 mm long; pedicels 1-2.5 mm long, very slender; bracteoles 0; calyx 2.3-3 mm long, the tube 1.5-2 mm long, campanulate, strigose, the teeth 0.5-1 mm long subulate; flowers 5-6.5 mm long, pink-purple; pods spreading, ses- sile, bladdery-inflated, subglobose, 10-17 mm long, 7-11 mm thick, mottled, strigose, unilocular; ovules 10-14. Sagebrush and sagebrush-juniper communities, often on windswept ridgetops at 1830 to 2300 m in Rich Co.; southwestern Wyoming and east- central Nevada. This tiny plant has as its nearest congener A. Umnocharis (q.v.), a plant of higher elevations in central south- ern Utah; 2 (0). Astragalus kentrophyta A. Gray. Kent- rophyta. Perennial, caulescent, mat-forming to erect, 5-45 cm long, from a caudex and stolon-like creeping stems; pubescence basi- fixed or malpighian; stems prostrate to erect, compact to elongate; stipules 1.5-5 mm long, at least some connate-sheathing; leaves 0.4-2.6 cm long; leaflets 3-9, 3-13 mm long, 0.5-1.5 mm wide, linear to nar- rowly elliptic or lanceolate, all continuous with the rachis and spinulose-tipped, stri- gose on both sides; peduncles to 1.5 cm long; racemes 1- to 3-flowered, the flowers declined at anthesis, the axis to 0.5 cm long in fruit; bracts 0.8-3.5 mm long; pedicels 0.5-2 mm long; bracteoles 0; calyx 2.4-8.3 Sept. 1978 Welsh: Utah Flora, Fab ace ae 283 mm long, the tiibe 1.3-3.3 mm long, cam- panulate, strigose, the teeth 1.5-5 mm long, subulate; flowers 4.5-10 mm long, pink- purple or whitish, ochroleucous, or purplish- tinged; pods declined or spreading, sessile, elliptic to oblong or lance-acumuminate in outline, usually curved, 4-10 mm long, 1.3-4 mm wide, strigose, unilocular; ovules 2-8. Three rather distinctive varieties occur in Utah. 1. Pubescence entirely of basifixed hairs; plants prostrate, of barrens at high elevations A. kentrophyta var. implexus Pubescence mostly of malpighian hairs; plants prostrate to erect, of low to high elevations 2 2(1). Calyx 6-8.3 mm long, the teeth 3.4-5 mm long; pods 7-10 mm long; plants prostrate, of sandy sites in canyons of the Colorado A. kentrophyta var. coloradoensis Calyx 3.4-5.5 mm long, the teeth 1.5-2.4 mm long; pods 4-7 mm long; plants erect or prostrate, usually of clay or silty soils at low to higher eleva- tions 3 3(2). Plants prostrate; pods 3-4.5 mm long, beakless or nearly so; known from Daggett Co A. kentrophyta var. jessiae - Plants erect or prostrate; pods 4-7 mm long, beaked; known from Duchesne Co., southward A. kentrophyta var. ekitus Var, coloradoensis M. E. Jones. Canyon Kentrophyta. [A. montanus var. colora- doensis (M. E. Jones) M. E. Jones; Kent- rophyta coloradoensis (M. E. Jones) Rydb.]. Wash bottoms and rimrock in mixed desert shrub communities at 970 to 1630 m in Emery, Garfield, Kane, and Wayne cos.; Coconino Co., Arizona; a Colorado Plateau endemic; 15 (v). Var. elatus Wats. Tall Kentrophyta. [A. viridis var. impensns Sheldon; A. kent- rophyta var. impensns (Sheldon) M. E. Jones; A. impensus (Sheldon) Woot. & Standi.; A. montanus var. impensus (Sheldon) M. E. Jones; A. tegetarius var. elatus (S. Wats.) Barneby]. Juniper-piny on, ponderosa pine, pine-spruce, mixed (often salt) desert shrub, and floodplain communities at 1530 to 2600 m in Beaver, Duchesne, Emery, Garfield, Iron, Kane, Piute, Sevier, and Wayne cos.; California, Nevada, Arizona, Colorado, and New Mexico. Both erect and prostrate phases are known, the presence of malpi- ghian hairs is diagnostic; 45 (xvii). Var. implexus (Canby) Barneby. Moun- tain Kentrophyta. [A. tegetarius var. im- plexus Canby; A. tegetarius S. Wats.; Trag- acantha tegetaria (S. Wats.) Kuntze; Homalobus tegetarius (S. Wats.) Rydb.; A. montanus var. tegetarius (S. Wats.) M. E. Jones; A. tegetarius var. rotundus M. E. Jones, type from Loa; A. kentrophyta var. rotundus (M. E. Jones) M. E. Jones]. Ridge- tops and breaks, commonly in barrens, at 2130 to 3500 m in Box Elder, Cache, Gar- field, Iron, Juab, Kane, Salt Lake, Sanpete, Sevier, Summit, Tooele, and Utah cos.; Ore- gon to Montana and south to California, Nevada, and New Mexico; 48 (vii). Var. jessiae (Peck) Barneby. Jessie Kent- rophyta. [A. jessiae Peck]. White volcanic ash "barrens" at 1770 m in Daggett Co.; eastern Oregon, Idaho, and Wyoming; 1 (0). Astragalus Uincearius A. Gray. Lancer Milkvetch. [Homalobus lancearius (A. Gray) Rydb.]. Perennial, caulescent, rush-like, 15-55 cm tall, arising from a subterranean caudex; pubescence basifixed; stems erect or ascending; stipules 2-7 mm long, all dis- tinct; leaves 1.5-10.5 cm long, some or most of them reduced to the rachis some with leaflets 3-7 in number, 2-14 mm long, 0.5-1.5 mm wide, linear to oblong, acute to obtuse, strigose on both sides; peduncles 4-23 cm long; racemes very loosely 6- to 25-flowered, the flowers ascending at an- thesis, the axis 3-19 (26) cm long in fruit; bracts 1-1.5 mm long; pedicels 1-3 mm 284 Great Basin Naturalist Vol. 38, No. 3 long; bracteoles 0-2; calyx 3.5-5.2 mm long, the tube 2.8-4.2 mm long, cam- panulate, about as broad as long, cam- panulate, black-strigose, the teeth 0.5-1.2 mm long, triangular; flowers 8.8-11.5 mm long, pink-purple or merely tinged purplish; pods deflexed, sessile or subsessile, the body lance-oblong to oblong or lance-elliptic in outline, slightly curved or straight, 20-35 mm long, 5-9 mm wide, laterally com- pressed, glabrous to strigose, brown to straw-colored, unilocular; ovules 8-14. Mixed desert shrub and pinyon-juniper com- mimities at 1270 to 1730 m in Kane and Washington cos.; Coconino and Mohave COS., Arizona; a Mohave-Virgin endemic. The lancer milkvetch is very similar to A. episcopus in most salient features (q.v.); 7 (i)- Astragalus lentiginosus Douglas ex Hook. Freckled Milkvetch. Perennial, caulescent, mostly 1.5-60 cm tall, from a caudex; pu- bescence basifixed; stipules 1.5-7 mm long or more, all distinct; leaves 2.4-15 cm long; leaflets 9-23,, 2-23 mm long, 1-13 mm wide, elliptic to ovate or lanceolate, obtuse to rounded, emarginate, or acute, pubescent to glabrous on one or both sides; peduncles 1-14 cm long, sometimes more; racemes (5) 11- to 30-flowered, the flowers ascending to declined at anthesis, the axis 1-18 cm long in fruit; pedicels 1-4 mm long; bracts 1.5-6 mm long; bracteoles 0-2; calyx 3.5-11.6 mm long, the tube 3-9 mm long, cylindric to short-cylindric, strigose, the teeth 0.6-2.5 mm long, subulate or triangular; flowers 8.4-22 mm long, pink-purple, ochroleucous, whitish, or variously suffused with pink or purple; pods ascending to declined, sessile, variable in outline, either inflated and ovoid (12-26 mm long, 5-20 mm thick) or not in- flated and oblong in outline (15-25 mm long, 3-7.5 mm thick), strigose or glabrous, mottled or not, leathery to membranous, bi- locular; ovules 16-38. Some 10 varieties of freckled milkvetch are known to occur in Utah. The complex in Utah has been re- vised by Schoener (1975). A revision of the Astragalus lentiginosus complex for the state of Utah. (Unpublished MS thesis, BRY). 1. Flowers small, the keel 5.5-8.5 mm long 2 Flowers larger, the keel 9-16 mm long or more 5 2(1). Raceme axis little elongating in fruit, not over 4 cm long 3 Raceme axis much elongating in fruit, 4-15 cm long or more 4 3(2). Pods opaque, stiffly papery; rare plants of extreme west-central Utah A. lentiginosus var. scorpionis — Pods transparent to translucent, thinly papery; plants locally common and rather widespread in western to central Utah A. lentiginosus var. salinus 4(2). Pods bladdery-inflated, the body globose or ovoid; plants of Washington Co. A. lentiginosus var. frernontii — Pods not inflated, the body oblong to narrowly ellipsoid; plants of Iron (?) Co A. lentiginosus var. ursinus 5(1). Raceme axis much elongating in fruit, usually more than 8 cm long 6 — Raceme axis not much elongating, seldom as much as 7 cm long in fruit 9 6(5). Flowers ochroleucous or faintly suffused with purple; pods diaphanous, glabrous; plants of Washington Co A. lentiginosus var. vitreus — Flowers ordinarily pink-purple to pink; pods usually opaque or strigose or both; distribution various 7 7(6). Pods bladdery-inflated; herbage cinereus, the stems canescent; plants known from the western slope of Beaverdam Mountains in Washington Co. A. lentiginosus var. stramineus Sept. 1978 Welsh: Utah Flora, Fab ace ae 285 — Pods not or scarcely inflated; herbage greenish, the stems not or seldom canescent; plants of various distribution, but not as above 8 8(7). Flowers relatively large, the keel 10-15 mm long; plants from the Virgin and Colorado River canyons A. lentiginosus var. pakins — Flowers smaller, the keel 8.5-10 mm long; plants rare and obscure, known only from "Bear Valley," Iron Co A. lentiginosus var. ursinus 9(5). Pods thinly papery, diaphanous; plants of eastern Kane Co. A. lentiginosus var. wahweapensis — Pods stiffly papery to leathery, opaque or nearly so; plants of various distri- bution 10 10(9). Leaflets 9-17; peduncles 1-5 cm long, shorter than the subtending leaves; flowers pale pink to whitish, or less commonly pink-purple; plants uncom- mon in northern Utah A. lentiginosus var. platyphyllidius — Leaflets 15-23; peduncles 2.5-8 cm long, mostly longer than the subtending leaves; plants variously distributed 11 11(10). Pods commonly strongly curved, with a long, lance-acuminate beak; plants common in Utah A. lentiginosus var. araneosus — Pods commonly little curved, with a broad triangular beak; plants uncom- mon in eastern Garfield and San Juan cos A. lentiginosus var. albiflorus Var. albiflorus (A, Gray) Schoener. Blad- der Milkvetch. [A. diphysus var. albiflorus A. Gray; A. diphysus A. Gray; A. lentigi- nosus var. diphysus (A. Gray) Jones; Trag- acantha diphysa (A. Gray) Kuntze; Cystium diphysum (A. Gray) Rydb.] Pinyon-juniper and mixed desert shrub communities at 1470 to 1970 m in the Henry Mountains, Garfield Co., and in San Juan Co.; Colo- rado, Arizona, and New Mexico. The Utah materials are transitional to var. araneosus on the one hand and to var. palans on the other. The name is a misnomer, since the flowers are ordinarilv a bright pink-purple; 3 (ii). Var. araneosus (Sheldon) Bameby. Cob- web Milkvetch. [A. araneosus Sheldon, type from Frisco; A. palans var. araneosus (Shel- don) M. E. Jones; Cystium araneosum (Shel- don) Rydb.; A. lentiginosus var. chartaceus M. E. Jones, type from Ephraim]. Sage- brush, mixed desert shrub, pinyon-juniper, and less commonly, in mountain brush com- munities at 1270 to 2430 m in Box Elder, Beaver, Garfield, Iron, Juab, Kane, Millard, Piute, Sanpete, Sevier, Tooele, and Wayne COS.; Nevada. The var. araneosus is the common short-racemed milkvetch with bright pink-purple flowers in much of south-central to western Utah. It approaches var. albiflorus in some of its technical fea- tures. Acquisition of specimens connecting the ranges of vars. araneosus and chartaceus demonstrate complete intergradation of sup- posed diagnostic features and dictate that the two should be combined. Because the leaflet number appeared to be fewer than that acceptable for var. araneosus, the var. chartaceous was construed as to include var. platyphyllidius by Schoener (1975). That re- mainder of the materials treated previously as var. platyphyllidius is maintained as such, even though it is hardly uniform; 123 (xxxii). Var. fremontii (A. Gray) S. Wats. Fre- mont Milkvetch. [A. fremontii A. Gray ex torr.; A. coulteri var. fremontii (A. Gray) M. E. Jones; Cystium fremontii (A. Gray) Rydb.] Braided stream gravels and slopes in creosote bush, Joshua tree, and pinyon- juniper communities at 670 to 1230 m in western Washington Co.; California and Ne- vada; 11 (ii). Var. palans (M. E. Jones) M. E. Jones. Straggling Milkvetch. [A. palans M. E. Jones, type from Montezuma Canyon; Tium 286 Great Basin Naturalist Vol. 38, No. 3 palans (M. E. Jones) Rydb.]. Salt desert shrub, blackbnish, juniper, pinyon-juniper, and mixed desert shrub communities at 1130 to 1900 m in Carbon, Emery, Grand, Kane, San Juan, Washington, and Wayne COS.; Colorado and Arizona. The stragghng milkvetch is the common phase of A. len- tiginosus in the canyons of the Colorado. It is distinctive in having oblong, usually curved pods, seldom more than 7 mm thick, and usually pale pink-purplish flowers; 88 (xxv). Var. platyphyllidius (Rydb.) Bameby. Broad-leaved Milkvetch. [Cystium platy- phyllidium Rydb.] Sagebrush, pinyon- juniper, and mountain brush communities at 17-0 to 2135 m in Daggett, Juab, Summit, and Tooele cos.; Oregon and California east to Wyoming and Colorado. The peduncles and commonly the fruiting racemes are sur- passed by the subtending leaves. This fea- ture, coupled with the leaflets which aver- age fewer per leaf seem to distinguish var. platyphyllidius from most of var. araneosus. Flower color varies from whitish with keel- tip purple to pink-purple throughout; 9 (ii). Var. salinus (Howell) Barneby. Salt Milk- vetch. [A. salinus Howell; Cystium salinum (Howell) Rydb.]. Pinyon-juniper, mountain brush, and sagebrush communities at 1770 to 2430 m in Beaver, Iron and Summit cos.; California, Oregon, Nevada, Idaho, and Wyoming. There appear to be two main centers of distribution of this variety in Utah, one in Iron and Beaver cos., and one in the area of contact of the Uinta and Wasatch mountains in Summit Co. Schoener (1975) reported that salt milkvetch from Salt Lake, Wasatch and Morgan cos., pre- sumably on the basis of plants collected by her, but there are no specimens in the her- barium at BRY from those localities. The tiny flowers and thin diaphanous pods are diagnostic; 12 (vii). Var. scorpionis M. E. Jones. Scorpion Milkvetch. [Cystium scorpionis (Jones) Rydb.] Sagebrush community upwards to timberline at 2130 to 3350 m in western Juab Co.; Nevada. This rare, small-flowered plant differs from var. salinus mainly in its opaque, leathery pods; 1 (0). Var. stramineus (Rydb.) Barneby. Straw Milkvetch. [Cystium stramineum Rydb., type from southern Utah]. Mixed warm desert shrub community at 900 to 1000 m on the western slope of the Beaverdam Mountains, Washington Co.; Arizona and Nevada. This is the only member of the species having moderate sized flowers known to occur in southwestern Washing- ton Co.; it is a Beaverdam-Virgin endemic; 5(0). Var. ursinus (A. Gray) Bameby. Bear Milkvetch. [A. ursinus A. Gray, type from Bear Valley, south-central Utah]. Habitat and elevation and specific locality are un- known. Presumably, the material was col- lected in Bear Valley, northeastern Iron Co.; endemic. The variety has been com- pared with var. palans but the small flowers and presumed locality each preclude associ- ation with the straggling milkvetch. Possi- bly, the Bear milkvetch is extinct, but that seems unlikely when one considers the abundant material in the type collection. A catastrophe might account for its dis- appearance, however; 1 (0). Var. vitreus Bameby. Glass Milkvetch. Creosote bush and mixed warm desert shrub community at 830 to 1430 m in central and eastern Washington Co.; Mohave and Coco- nino COS., Arizona. In its typical phase the flowers are very pale, either ochroleucous or white tinged with a faint pink to blue- purple. Southward and eastward the flower color becomes a bright pink-purple. The large, glassy, diaphanous pods are dis- tinctive; 20 (iii). Var. wahweapensis Welsh var. nov. As- tragalo lentiginoso var. araneoso aemulans, differt leguminibus diaphanis et stramineis. Holotype: Utah, Kane Co., pinyon-juniper community at ca 1900 m in lower Kaiparo- wits Formation on Four Mile Bench, Welsh 12426, 8 May 1974 (BRY, nine isotypes). Additional materials; Utah, Kane Co., ca. 5 mi south of Cannonville, Welsh 5364, 6 May 1966; do, 4 mi south of Cannonville, Reveal et al 783, 4 June 1967; do, Wah- weap Creek, Atwood 3474, 22 Mar. 1972; do. Four Mile Bench, Welsh 12413, 12 Apr. 1974; do. Smoky Mt., Cronquist 10022, 5 May 1965; do, Escalante road. Woodruff 1139, 12 June 1971; do. Four Mile Bench, Sept. 1978 Welsh: Utah Flora, Fab ace ae 287 Welsh & Murdock 12394, 12395, 17 May 1974; do. Four Mile Bench, Atwood 4064, 1 June 1972; do. Nipple Bench, Atwood & Allen 2920, 14 June 1971; do, Brigham Plains, Atwood & Allen 2791a, 13 June 1970; do, Four Mile Bench, Welsh 12446, 16 June 1974; do. Four Mile Bench, Welsh & Moore 13631, 29 June 1976 (all at BRY). The Wahweap milkvetch is an inhabitant of sandy soils mainly in pinyon-juniper and sagebRish communities at 1670 to 1900 m in the Paria, Wahweap, and Last Chance drainages west of the Kaiparowits Plateau proper. It is compared in the diagnosis to var. araneosus which surrounds the region, mainly at lower elevations, but shares fea- tures with var. vitreus, a low elevation plant to the south and west of the Wah- weap centrum, which has more strongly graduated petals and usually larger and pa- ler flowers; 16 (vii). Astragalus limnocharis Barneby. Navajo Lake Milkvetch. Perennial, subacaulescent, 1-5 cm tall, arising from a branching cau- dex; pubescence basifixed; stems prostrate to erect; stipules 2-4 mm long, all connate- sheathing; leaves 1.5-7 cm long; leaflets (5) 7-13, 2-9 mm long, 1-2 mm wide, lanceo- late to elliptic or oblong, obtuse, strigose beneath, long-ciliate on the involute margin, glabrous above; peduncles 2-5 cm long, re- clining in fruit; racemes 2- to 10-flowered, the flowers spreading to declined at an- thesis, the axis 0.2-0.5 cm long in fruit; bracts 1-3 mm long; pedicels 0.8-1.5 mm long; bracteoles 0; calyx 2.8-3.6 mm long, the tube 2-2.5 mm long, campanulate, stri- gose, the teeth 0.7-1.6 mm long, subulate; flowers 6.2-7.5 mm long, ochroleucous, con- colorous; pods spreading, sessile, ovoid, bladdery-inflated, 9-18 mm long, 7-13 mm thick, mottled, strigose, unilocular; ovules 10-12. Lake shores and limestone breaks of Wasatch Formation at 2670 to 3400 m in Iron and Kane; endemic. This "lake beauty" has a principal locality on the beach at Navajo Lake where it is evidently dispersed by wave action. Plants can be found along the terraces below the high water line as the lake recedes. Evidently, the nearest al- lies of the Navajo Lake milkvetch is A. je- junus (q.v.), known in Utah only from Rich Co. and A. montii of the Wasatch Plateau; 10 (iii). Astragalus loanus Barneby, Loa Milk- vetch. [A. newberryi var. wardianus Bar- neby, type from east of Glenwood]. Pe- rennial, acaulescent, 3-19 cm tall, the caudex branches clothed with a thatch of persistent leaf bases; pubescence malpi- ghian; stipules 5-10 mm long, all distinct; leaves 2-18 cm long; leaflets (1) 3-11, 6-21 mm long, 2-12 mm wide, obovate to ellip- tic or lanceolate, obtuse to acute, densely strigose on both sides; peduncles 0.5-10 cm long; racemes 2- to 7-flowered, the flowers erect-ascending at anthesis, the axis 0.2-0.8 cm long in fruit; bracts 2.5-6 mm long; pedicels 1.2-1.8 mm long; bracteoles 0-2; calyx 10.7-18.5 mm long, the tube 8.5-14 mm long, cylindric, loosely strigulose, the teeth 1.7-4.5 mm long; flowers 20-28 mm long, ochroleucous or greenish-white often tinged faintly purplish, the keel-tip purple; pods ascending; sessile, inflated, ovoid, 17-30 (40?) mm long, 8-19 mm wide (when pressed), the valves red-purple to straw- colored, hirsute with lustrous long hairs, imilocular; ovules 28-38. Sagebrush and pinyon-juniper communities exclusively on igneous gravels at 1830 to 2570 m in Gar- field, Piute, Sevier, and Wayne cos.; endem- ic. This remarkable species, known until re- cently from only four collections, apparently consists of two phases, separable inter alia by leaflet shape. The features are apparently not constant, and more speci- mens are required before the taxonomic status, if any, of the variants will be clari- fied; 14 (ix). Astragalus lonchocarpus Torr. Great Rushy Milkvetch. [Phaca macrocarpa A. Gray; Tragancantha lonchocarpa (Torr.) Kuntze; Homalohus macrocarpus (A. Gray) Rydb.; Lonchophaca macrocarpa (A. Gray) Rydb.; A. macer A. Nels.; L. macro (A. Nels.) Rydb.]. Perennial, caulescent, 30-84 cm tall, from a shallowly subterranean cau- dex; pubescence basifixed; stems erect, often in dense clumps; stipules 1-9 mm long, all distinct; leaves 2-13 cm long, the up- permost and sometimes all simple, the lower with leaflets 3-9, 2-36 mm long, 0.5-4 mm wide, linear to narrowly ob- 288 Great Basin Naturalist Vol. 38, No. 3 lanceolate, obtuse to acute, strigose on both sides or glabrous above; peduncles 6-24 cm long; racemes loosely 7- to 40-flowered, the flowers spreading-declined at anthesis, the axis 3.5-45 cm long in fruit; bracts 0.8-2.5 mm long; pedicels 1.3-4.5 mm long; brac- teoles; calyx 5.8-10.3 mm long, usually brown, the tube 5-8 mm long, cylindric, gibbous, strigose, the teeth 0.6-2 mm long, subulate; flowers 13-20 mm long, ochro- leucous to almost white, concolorous; pods pendulous, stipitate, the stipe 3-15 mm long, the body elliptic to oblong in outline, 22-50 mm long, 3.3-6.2 mm wide, dorsi- ventrally compressed, the valves often brownish, strigose, unilocular; ovules 12-26. Salt desert shrub, pinyon-juniper, floodplain, and seep communities at 1170 to 2530 m in Beaver, Carbon, Emery, Garfield, Grand, Iron, Juab, Kane, Millard, Piute, San Juan, Sevier, Tooele, and Wayne cos.; Colo- rado, New Mexico, Arizona, and Nevada; 85 xxvu Astragalus lutosus M. E. Jones. Dragon Milkvetch. Perennial, short-caulescent, 2-10 cm tall, from a subterranean caudex; pu- bescence basifixed; stems prostrate to as- cending, radiating; stipules 2-5 mm long, all distinct or some shortly connate-sheathing; leaves 1-5.5 cm long; leaflets 15-27, 1-9 mm long, 1-5 mm wide, obovate to elliptic, obtuse to retuse, gray-strigulose on both sides or glabrous above; peduncles 0.5-2 cm long; racemes 6- to 10-flowered, the flowers ascending-spreading at anthesis, the axis 0.3-1 cm long in fruit; bracts 1.5-2.5 mm long; pedicels 1.2-3 mm long; bracteoles 0; calyx 4.8-10.4 mm long, the tube 3.6-7.4 mm long, short-cylindric, strigulose, the teeth 1.2-3 mm long, subulate; flowers 9-17 mm long, white to ochroleucous, the keel- tip purplish; pods spreading to ascending, stipitate, the stipe (gynophore) 1-4.5"^ mm long, the body ovoid or ellipsoid bladdery- inflated, 15-37 mm long, 8-17 mm thick, strigose, imilocular. Bare places on white (Green River) shale ridges and talus slopes in mixed desert shrub community in south- eastern Uintah Co. and in adjacent Rio Blanco Co., Colorado; a Uinta Basin En- demic. The Dragon milkvetch is one of the rarest of our species of milkvetches; 2 (0). Astragalus malacoides Barneby. Kaiparo- wits Milkvetch. Perennial, caulescent, 7-25 cm tall, from a slightly subterranean cau- dex; pubescence basifixed; stems decumbent or prostrate to ascending; stipules 2-7 mm long, all distinct; leaves 4.5-14 cm long; leaflets 15-29, 3-25 mm long, 2-12 mm wide, obovate to oblong or elliptic, obtuse to emarginate, hirtellous beneath, glabrous above; peduncles 4-12 cm long; racemes 10- to 24-flowered, the flowers ascending- spreading at anthesis, the axis 1.2-10 cm long in fruit; bracts 3-5 mm long; pedicels 1-2.5 mm long; bracteoles 0-2; calyx 10-15 mm long, the tube 7-11 mm long, cylindric, hirsutulous, the teeth 2-6.2 mm long, linear- subulate; flowers 16-22 mm long, pink- purple; pods declined to ascending, short- stipitate, the stipe 2-3 mm long, the body oblong, curved, 25-40 mm long, 5-8 mm wide, laterally compressed, hirsutulous, bi- locular; ovules 24-30. Straight Cliffs, Wah- weap, Kaiparowits, and Moenkopi forma- tions, usually in clay or silty soils, in juniper-pinyon and mixed desert shrub com- munities at 1600 to 2330 m in Garfield (Circle Cliffs and Tarantula Mesa) and Kane (Kaiparowits vicinity) cos.; endemic; 21 (iv.) Astragalus marianus (Rydb.) Barneby. Sevier Milkvetch. [Xylophacos marianus Rydb., type from Marysvale]. Perennial, acaulescent or subacaulescent, 3-10 cm tall, from a branching caudex; pubescence basi- fixed; stems 0-6 cm long, the internodes mostly concealed by the stipules; stipules 2-7 mm long, all distinct; leaves 1.2-8.5 cm long; leaflets 7-17, 3-11 mm long, 1-4 mm broad, obovate to oblanceolate, obtuse to emarginate or acuminate to acute, strigose on both sides; peduncles 1-8 cm long; ra- cemes 2- to 10-flowered, the flowers as- cending at anthesis, the axis 0.2-3.5 cm long in fruit; bracts 2-4.5 mm long; pedi- cels 0.8-3 mm long; bracteoles 0-2; calyx 10-13.2 mm long, the tube 7.3-9.4 mm long, cylindric, pilosulous, the teeth 2-3.5 mm long, subulate; flowers 17-24 mm long, pink-purple, often pale; pods spreading- ascending, sessile or nearly so, 10-23 mm long, 7-12 mm thick, ovoid-acuminate, densely shaggy-hirsute, imilocular; ovules 27-36. Oak-sagebrush, mixed warm and cool Sept. 1978 Welsh: Utah Flora, Fabaceae 289 desert shrub, pinyon-juniper, and aspen- white fir communities at 900 to 2430 m in Beaver, Garfield, Iron, Juab, Millard, Piute, Sevier, Washington and Wayne cos.; Ne- vada. The Sevier milkvetch resembles A. ar- gophijUus var. martinii but the shaggy- hirsute pods and flowers that average small- er are diagnostic; 36 (xviii). Astragalus megacarpus (Nutt.) A. Gray. Great Bladdery Milkvetch. [Tragacantha megacarpa (Nutt.) Kuntze; A. megacarpus var. prodigus Sheldon.]. Perennial, short- caulescent or subacaulescent, 3-15 (17) cm tall, arising from a caudex; pubescence basi- fixed; stems 1-5 cm long, in dense, leafy clumps, the internodes mostly obscured by stipules; stipules 2-7 mm long, all distinct; leaves 2-15 (17) cm long; leaflets 7-19, 3-19 mm long, 2-12 mm wide, obovate, ovate, elliptic, or suborbicular, obtuse to re- fuse and mucronate, strigose to glabrous be- neath, glabrous above; peduncles 0.5-6 cm long, much shorter than the leaves; racemes 1- to 7-flowered, the flowers ascending in anthesis, the axis 0.2-2.5 cm long in fruit; bracts 2-5 mm long; pedicels 3.5-8 mm long; bracteoles 0-2; calyx 7-10.2 mm long, the tube 5.2-8.5 mm long, cylindric, stri- gose, the teeth 1.8-3.5 mm long; flowers 15-23 mm long, pink-purple; pods ascend- ing to descending, stipitate, the stipe (gyno- phore) 2-4 mm long, the body bladdery- inflated, ellipsoid, 35-65 (70) mm long, 15-31 mm wide (when pressed), commonly mottled, strigose, unilocular; ovules 38-54. Commonly on clay soils, these often saline or calciferous, in salt desert shrub, sage- brush, oakbrush, pinyon-juniper, and ponderosa pine communities at 1650 to 2430 m in Carbon, Duchesne, Emery, Gar- field, Iron, Juab, Kane, Millard, Sanpete, and Sevier cos.; Nevada, Wyoming, and Colorado. The great bladdery milkvetch is among our most distinctive species of Astra- galus. A plant with its large pods declined on the ground around the base is a surpris- ing sight. Our plants belong to var. parryi A. Gray ex S. Wats., sens, str., but this vari- ety apparently differs mainly in flower col- or (though our plants seem to have smaller calyces too) from var. megacarpus, sens, str.; 44 (xii). Astragalus miser Douglas ex Hook. Weedy Milkvetch. Perennial, caulescent or short-caulescent, 3-35 cm tall, from a branching caudex; pubescence basifixed; stems decumbent to erect, 1-25 cm long or more; stipules 1.5-7 mm long, at least some connate-sheathing; leaves 1.5-2.0 cm long or more; leaflets 3-21, 3-35 mm long, 0.5-7 mm wide, linear to oblong, elliptic, or oval, acute to obtuse or emarginate, strigose be- neath, glabrous or glabrate above; peduncles 2-14 cm long; racemes 3- to 20-flowered, the flowers spreading-declined at anthesis, the axis 1-10 cm long in fruit; calyx 2.1-5.2 mm long, the tube 1.9-2.9 mm long, cam- panulate, strigose, the teeth 0.5-2.5 mm long, subulate; flowers 5.9-10 mm long, pink-purple, ochroleucous, or whitish, often suffused or veined with purple; keel with an upturned usually purple beak; pods declined-pendulous, sessile or nearly so, nar- rowly oblong or oblanceolate in outline, 12-25 mm long, 2-4 mm wide, strigose, imilocular; ovules 8-19. The weedy milkvetch is a widespread poisonous plant of middle and upper eleva- tions in much of Utah. Both cattle and sheep are poisoned by this plant. Two rather tenuous varieties are known from Utah, one common and the other evidently rare. Leaflets 3-11; flowers 6-8 mm long; pods narrowly oblong in outline; plants rare, scattered in northern Utah A. 7niser var. tenuifolius Leaflets mostly 11-21; flowers 8-11 mm long; pods oblanceolate in outline; plants common in Utah A. miser var. oblongifolius Var. oblongifolius (Rydb.) Cronq. Ryd- berg Weedy Milkvetch. [Homalobos ob- longifolius Rydb.; A. hylophilus var. ob- longifolius (Rydb.) Macbr.; A. decumbens var. oblongifolius (Rydb.) Cronq.; Homa- lobus humilis Rydb., type from the Tushar 290 Great Basin Naturalist Vol. 38, No. 3 Mts.; A. carltonii Macbr., not A. humilus MB.] Aspen and mixed-aspen conifer, and coniferous woodlands, and in sagebrush, mountain brush, and alpine meadow com- munities at 1830 to 3500 m in all Utah counties except Box Elder, Davis, Millard, Morgan, San Juan, Tooele, Washington, and Weber cos., (and to be expected in some of them); Nevada, Arizona, Wyoming, and Colorado; 141 (xxvi). Var. tenuifolius (Nutt.) Barneby. Gar- rett's Weedy Milkvetch. [Homalobus tenui- folius Nutt.; H. pauciiugus Rydb.; A. gar- rettii Macbr., not A. paucijiigus Schrenk.]. Sagebrush upwards to mountain summits at 1970 to 3050 m in Rich and Salt Lake (and likely in Summit) cos.; Idaho, Nevada, and Wyoming; 2 (0). Astragalus missouriensis Nutt. Missouri Milkvetch. Perennial, subacaulescent, 3-12 cm tall, from a caudex; pubescence malpi- ghian; stems prostrate, to 8 cm long, the in- ternodes often concealed by stipules; stip- ules 2-9 mm long, all distinct; leaves 1.5-12 cm long; leaflets 5-15, 3-15 mm long, 1-8 mm wide, elliptic to obovate, acute to mucronate or obtuse, strigose on both sides; peduncles 1.5-8 cm long; racemes 3- to 12- flowered, the flowers ascending at anthesis, the axis 0.4-3 cm long in fruit; bracts 2.5-8 mm long; pedicels 1-3.5 mm long; brac- teoles 0-2; calyx 8.5-13 mm long, the tube 7-10 mm long, cylindric, strigose, the teeth 1.5-3 mm long, subulate; flowers 15-22 mm long, pink-purple, rarely white; pods as- cending to descending, sessile, ellipsoid, 15-25 mm long, 7-9 mm thick, curved, dor- siventrally compressed, strigose, unilocular; ovules 35-55. Pinyon-juniper and sagebrush communities at 1600 to 2430 m in Grand and San Juan cos.; Colorado. Our materials are assignable to var. amphibolus Barneby; 2 (0). Astragalus moencoppensis M. E. Jones. Moenkopi Milkvetch. [Cnemidophacos moencoppensis (M. E. Jones) Rydb.]. Pe- rennial, caulescent, 9-60 cm tall, from a branching caudex; pubescence basifixed; stems erect or ascending, commonly shorter than the longest peduncles; stipules 1.5-7 mm long, at least some connate-sheathing; leaves 4-16.5 cm long; leaflets 5-15, 2-23 mm long, 0.3-2 mm wide, filiform to linear or narrowly elliptic, acute to obtuse, the terminal often contiguous with the rachis, strigose below, glabrous on the involute up- per surface; peduncles 4-25 cm long; ra- cemes 6- to 34-flowered, the flowers as- cending at anthesis, the axis 3-25 cm long in fruit; bracts 1.5-3.5 cm long; pedicels 0.3-2 mm long; bracteoles 0-1; calyx 5-7.5 mm long, the tube 3-4 mm long, cam- panulate, white-pilose, the teeth 1.8-3.5 mm long, lance-subulate; flowers 8-11 mm long, pink-purple; pods ascending-spreading, sessile, ovoid to ellipsoid, 6-7 mm long, 2.3-3.4 mm thick, strigulose, unilocular. Salt desert shrub, mixed desert shrub and pin- yon-juniper communities at 1330 to 2130 m, usually in clay or silty soils, in Emery, Gar- field, Grand, Kane, San Juan and Wayne COS.; Arizona. The Moenkopi milkvetch is a primary selenium indicator of distinctive mein and odor; 44 (xi). Astragalus mollissimus Torr. Woolly Lo- coweed. [A. thompsonae S. Wats.; Trag- acantha thoiyipsonae (S. Wats.) Kuntze; A. bigelovii var. thompsonae (S. Wats.) M. E. Jones; A. stjrticolus Sheldon]. Perennial, acaulescent, 6-45 cm tall, from a caudex; pubescence basifixed; stems mostly obscured by stipules; stipules 4-13 mm long, all dis- tinct; leaves 2-28 cm long; leaflets 15-35, 2-18 mm long, 1-14 mm wide, obovate to suborbicular or elliptic, obtuse to retuse oracute, densely wooly-tomentose on both sides; pedimcles 2.5-24 cm long; racemes 7- to 20-flowered, the flowers ascending at an- thesis, the axis 1.5-18 cm long in fruit; bracts 2.5-8 mm long; pedicels 0.5-3 mm long; bracteoles 0-2; calyx 11-15.5 mm long, the tube 7.7-23 mm long, cylindric, villous, the teeth 2-4.2 mm long, subulate; flowers 18-25 mm long, pink-purple; pods descending, sessile, ovoid, 11-23 mm long, 6-11 mm thick, curved, densely villous- tomentose, bilocular; ovules 28-38. Salt desert shrub, mixed desert shrub, grassland, and pinyon-juniper communities at 1130 to 2330 m in Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Iron, Kane, Mil- lard, San Juan, Tooele, Uintah, and Wash- ington COS.; Nevada, Colorado, Arizona and New Mexico. The woolly locoweed is one Sept. 1978 Welsh: Utah Flora, Fabaceae 291 of the earliest of the species of Astragalus to flower in Utah. It has been collected in anthesis as early as February in southern counties. The species as a whole is recog- nized as being poisonous. The plants are seldom abundant, except along roadsides, and this might account for the lack of re- ports of poisoning from this entity in Utah; 157 (xxxvi). Astragalus montii Welsh. Heliotrope Milkvetch. Perennial, subacaulescent, 1-5 cm tall, arising from a branching caudex; pubescence basifixed; stems ascending to erect; stipules 2-4 mm long, all connate- sheathing; leaves 1.3-4.8 cm long; leaflets 5-13, 2-8 mm long, 1-2 mm wide, lanceo- late to oblong or elliptic, strigose beneath, not ciliate on the involute margin, glabrous above; peduncles 0.8-4.5 cm long, reclining in fruit; racemes 2- to 8-flowered, the flow- ers ascending to spreading at anthesis, the axis 0.2-0.5 cm long in fruit; bracts 1-3 mm long; pedicels 0.8-1.5 mm long; bracteoles 0; calyx 3.3-4 mm long, the tube 2.2-2.5 mm long, campanulate, strigose, the teeth 0.6-1.5 mm long, triangular-subulate; flow- ers 7.2-8 mm long, pink-purple, the wing- tips white; pods spreading, sessile, ovoid, bladdery-inflated, 11-18 mm long, 8-12 mm thick, mottled, strigose, unilocular; ovules 10. The heliotrope milkvetch is known only from Flagstaff Limestone at one locality at 3350 m on the Wasatch Plateau in Sanpete Co.; endemic. The species is a near con- gener of A. limnocharis from the Markagunt Plateau, differing inter alia in the pink- purple flowers which are larger and in the merely strigose (not long ciliate) leaf mar- gins; 4 (i). Astragalus monutnentalis Barneby. Mon- imient Milkvetch. Perennial, acaulescent or subacaulescent, 3-18 cm tall, from a branching caudex; pubescence basifixed or shortly malpighian; stems 1-6 cm long, as- cending, the internodes commonly con- cealed by stipules; stipules 2-4 mm long, all distinct; leaves 1.5-8 (11) cm long; leaflets (5) 9-17 (21), 2-9 mm long, 1-4 mm broad, oval to obovate, elliptic, or oblanceolate, strigulose beneath, glabrous or glabrate above; peduncles 1-12 cm long; racemes 3- to 9-flowered, the flowers ascending at an- thesis, the axis 0.5-7 cm long in fruit; bracts 1.5-5 mm long; pedicels 0.8-2.2 mm long; bracteoles 0; calyx 3.6-4.5 mm long, the tube 3.1-3.5 mm long, campanulate, stri- gose, purplish, the teeth 0.5-1 mm long; flowers 8-9 mm long, pink-purple; pods as- cending, sessile or nearly so, narrowly ob- long to lanceolate in outline, 12-21 mm long, 2.3-3 mm wide, straight or curved, triangular in cross-section, the dorsal suture sulcate, strigose, bilocular, often mottled. Rimrock and other slickrock sites in mixed desert shrub and pinyon-juniper commu- nities at 1230 to 1870 m in Garfield and San Juan cos.; endemic. This is a mirrored- image cogener of A. cottamii (q.v.), differing in its smaller floral parts and overall flowers size and in the incipient malpighian pub- escence; 19 (xii). Astragalus musiniensis M. E. Jones. Fer- ron Milkvetch. [Xylophacos musiniensis (M. E. Jones) Rydb.; A. musiniensis var. new- herryoids M. E. Jones, lectotype from the San Rafael Swell.] Perennial, acaulescent, 3-13 cm tall, from a caudex, the branches often clothed with a thatch of persistent leaf bases; pubescence basifixed or in- cipiently malpighian; stipules 3.5-10 mm long, all distinct; leaves 1.5-13 cm long; leaflets (1) 3-5, 5-35 mm long, 2-11 mm wide, elliptic to lanceolate, acute to obtuse, strigose on both surfaces; peduncles 0.5-7 cm long; racemes 1- to 6-flowered, the flowers erect-ascending at anthesis, the axis 0.2-1.4 cm long in fruit; bracts 1.6-4 mm long; pedicels 1.2-4 mm long; bracteoles 0; calyx 12-15.5 mm long, the tube 9.5-12.7 mm long, cylindric, strigose-pilose, the teeth 1.5-4 mm long; flowers 20-28 mm long, pink-purple; pods ascending, sessile, obliquely ovoid, 15-36 mm long, 8-17 mm thick, dorsiventrally compressed, hirsutulous to villous-hirsute, unilocular. Salt desert shrub, mixed desert shrub, and pinyon-juniper communities at 1430 to 2130 m in Carbon, Emery (type from 2 miles south of Ferron), Garfield, Grand, Kane, and Wayne cos.; en- demic. The species was named by Marcus E. Jones for a mountain peak on the Wasatch Plateau many miles remote from the type locality; 30 (vii). Astragalus nelsonianus Barneby. Nelson 292 Great Basin Naturalist Vol. 38, No. 3 Milkvetch. [A. pectinatus var. platyphyllus M. E. Jones.]. Perennial, caulescent, 10-30 cm tall, from a subterranean caudex; pu- bescence basifixed; stems decumbent to as- cending or erect; stipules 4-13 mm long, at least some connate-sheathing; leaves 2.5-9 cm long; leaflets 3-13, 10-45 mm long, 2-5 mm wide, narrowly oblong, obtuse to apicu- late, strigose on both surfaces, the terminal leaflet continuous with the rachis; peduncles 3-12 cm long; racemes 6- to 20-flowered, the flowers ascending at anthesis, the axis 2-12 cm long in fruit; bracts 2.5-7 mm long; pedicels 1.5-4 mm long; bracteoles 1; calyx 10-14.5 mm long, the tube 7-10.2 mm long, cylindric, strigose, the teeth 2-4.5 mm long; flowers 10-14 mm long, white, concolorous; pods deflexed, sessile, oblong- ellipsoid, 13-33 mm long, 6-12 mm wide, finally laterally compressed, glabrous or pubescent, unilocular; ovules 20-28. Saline soils in desert shrub communities at 2070 to 2100 m in Daggett Co.; Wyoming. The Nelson milkvetch differs from A. pectinatus Dougl. ex Don in much the same manner that A. hamiltonii differs from A. loncho- carpus. The leaves and leaflets of both A. nehonii and A. hamiltonii are much broader than those of their counterparts; 3 (0). Astragalus newberryi A. Gray. Newberry Milkvetch. [Xylophacos newberryi (A. Gray) Rydb.; A. newberryi var. castoretis M. E. Jones, type from Beaverdam Mts.]. Pe- rennial, acaulescent, 2-14 cm tall, from a caudex, the branches commonly clothed with a thatch of persistent leaf bases; pu- bescence basifixed; stipules 3-11 mm long, all distinct; leaves 1.5-14 cm long; leaflets 3-15, 3-20 mm long, 2-14 mm wide, obo- vate to elliptic, oblanceolate, or orbicular, acute to obtuse or retuse, villous-tomen- tulous on both surfaces; peduncles 0.5-11 cm long; racemes 2- to 8-flowered, ascend- ing in flower, the axis 0.2-2.7 cm long in frviit; bracts 3.5-10 mm long; pedicels 1.4-5 mm long; bracteoles 0-2; calyx 11.5-20 mm long, the tube 8-17 mm long, cylindric, vilous, the teeth 1.9-6 mm long, subulate; flowers 20-32 mm long, pink-purple; pods ascending to spreading, sessile or subsessile, ovoid, curved, 18-23 mm long, 7-12 mm thick, densely villous-tomentose, unilocular; ovules 20-40. Salt desert shrub, mixed desert shnib, sagebrush, moimtain brush, and pinyon-juniper communities at 830 to 2300 m in Beaver, Box Elder, Garfield, Iron, Juab, Kane, Millard, Sevier, Tooele, and Washington cos.; Oregon, Nevada, Cali- fornia, Arizona, and New Mexico; 65 (xix). Astragalus nidularius Barneby. Bird's- nest Milkvetch. Perennial, caulescent 15-51 cm tall, from a subterranean caudex; pu- bescence basifixed; stems ascending to erect, often branched; stipules 1-6 mm long, at least some connate-sheathing; leaves 1.5-7 cm long; leaflets 5-11, 2-20 mm long, 1.3-2 mm wide, linear to oblong, obtuse to emarginate or acute, the terminal leaflet of upper leaves sometimes continuous with the rachis, pubescent on both sides or glabrous above; peduncles 4-16 cm long; racemes (3) 8- to 33-flowered, the flowers ascending to declined at anthesis, the axis 1.5-28 cm long in fruit; bracts 1.2-2.2 mm long; pedi- cels 1.2-3 mm long; bracteoles 0-2; calyx 3.8-7 mm long, the tube 3.3-5.5 mm long, campanulate, strigose, the teeth 1.5-2.2 mm long, subulate; flowers 10.5-15 mm long, pink-purple; pods pendulous, stipitate, the stipe 3.5-6 mm long, the body narrowly ob- long, 20-32 mm long, 3.5-4.5 mm thick, dorsiventrally compressed, strigose, unilocu- lar; ovules 20-24. Pinyon-juniper and mixed desert shrub communities on Cutler, Moen- kopi, and Navajo formations at 1370 to 1900 m in Garfield, San Juan, and Wayne COS.; endemic. The birds-nest milkvetch is a close ally of A. harrisonii (q.v.), from which it is separated geographically and in techni- cal morphological features; 13 (xii). Astragalus nuttallianus Nutt. Small- flowered Milkvetch. Annual or winter an- nual, caulescent, 2-35 cm long, from a tap- root; pubescence basifixed; stems prostrate to decumbent or erect; stipules 1-5 mm long, all distinct; leaves 1-8.5 cm long; leaflets 7-19, 2-14 mm long, 1-6 mm wide obovate to elliptic or oblong, acute to rounded or retuse to emarginate, strigose to glabrous on both sides; peduncles 1-8 cm long; racemes 1- to 7-flowered, the flowers ascending to declined at anthesis, the axis 0.2-2 cm long in fruit; bracts 0.5-2.5 mm long; pedicels 0.4-1.6 mm long; bracteoles 0; calyx 3.2-5.4 Sept. 1978 Welsh: Utah Flora, Fabaceae 293 mm long, the tube 1.9-2.8 mm long, cam- panulate, strigose, the teeth 1-2.2 mm long, subulate; flowers 4.1-7.6 mm long, pink- purple to whitish or faintly tinged with purplish; pods ascending to declined, sessile or subsessile, narrowly oblong, curved, 12-21 mm long, 1.9-3.3 mm wide, glabrous to strigose, bilocular or nearly so; ovules 12-18. There are two allopatric varieties of this annual milkvetch in Utah. 1. Leaflets 9-15; axis of raceme 0.5-2 cm long in fruit; keel tip blunt; plants of eastern Kane Co. and eastward in Utah A. nuttallianiis var. micranthiformis Leaflets 7-11; axis of raceme 0.1-1 cm long in fruit; keel tip acute to sub- acute, usually beaked; plants from central Kane Co. west to Washington Co., and disjunctly north to Tooele Co A. nuttallianus var. imperfectus Var. imperfectus (Rydb.) Barneby. [Ham- osa imperfecta Rydb.]. Creosote bush, warm desert shrub, and cool desert shrub commu- nities at 670 to 1470 m in Kane and Wash- ington and disjunctly in Tooele (Stansbury Island) COS.; Nevada, California, and Ari- zona; 23 (iv). Var. micranthiformis Barneby. Black- brush, mixed desert shrub, and pinyon- juniper communities at 1130 to 1670 m in Emery, Garfield, Grand, Kane, and San Juan COS.; Arizona, Colorado, and New Mexico; 13 (vii). Astragalus oophorus S. Wats. Egg Milk- vetch. Perennial, caulescent, 15-30 cm tall, from a caudex; pubescence basifixed; stems decimibent to ascending, radiating from the caudex; stipules 1.5-7 mm long, all distinct; leaves 3-21 cm long; leaflets 9-25, 3-20 mm long, 2-11 mm wide, oval to obovate or orbicular, obtuse to retuse or mucronate, glabrous on both surfaces, often ciliate; pe- duncles 4-13 cm long; racemes 4- to 13- flowered, the flowers spreading at anthesis, the axis 1-8 cm long in fruit; bracts 1.5-5 mm long; pedicels 2-6 mm long; bracteoles 0-1; calyx 6-12 mm long, the tube 4-8.5 mm long, cylindric or short-cylindric, glabrous or sparingly strigose, the teeth 2-5 mm long, subulate; flowers 17-24 mm long, ochroleucous, concolorous, or pink-purple and with white wing-tips; pods spreading to pendulous, stipitate the stipe (gynophore) 3.5-12 mm long, the body bladdery-inflated, ellipsoid, 25-55 mm long, 10-30 mm wide (when pressed), glabrous, unilocular, often mottled; ovules 28-54. Two distinctive vari- eties are present in Utah. 1. Flowers pink-purple; calyx tube 7.8-9 mm long; stipe (gynophore) 10-12 mm long; plants of western Iron and Beaver cos A. oophorus var. lonchocalyx Flowers ochroleucous; calyx tube 4-7 mm long; stipe (gynophore) 3.5-8.5 mm long; plants widespread A. oophorus var. caulescens Var. caulescens (M. E. Jones) M. E. Jones. Pallid Egg Milkvetch. [A. megacarpus var. caulescens M. E. Jones, type from Loa Pass; A. artipes A. Gray; Phaca artipes (A. Gray) Rydb.; A. oophorus var. artipes (A. Gray) M. E. Jones]. Sagebrush, pinyon- juniper, and mountain brush communities at 1370 to 2430 m in Beaver, Garfield, Iron, Juab, Kane, Millard, Piute, Sanpete, Sevier, Tooele, Utah, and Washington cos.; Colo- rado, Arizona, and Nevada. Please compare this variety to A. beckwithii var. beckwithii; 66 (vi). Var. lonchocalyx Barneby. Pink Egg Milkvetch. Pinyon-juniper, sagebmsh, and mixed desert shrub communities at 1770 to 2300 m in western Iron and Beaver cos.; Lincoln Co., Nevada; a Great Basin endem- ic; 7 (iv). Astragalus pardalinus (Rydb.) Barneby. Panther Milkvetch. [Phaca pardalina Rydb.] Perennial (short-lived) or functionally an- nual, caulescent, 8-30 (35) cm tall; pub- escence basifixed; stems decumbent to as- cending, forming rounded clumps; stipules 2-5 mm long, all distinct or some shortly 294 Great Basin Naturalist Vol. 38, No. 3 connate-sheathing; leaves 3-7 cm long; leaf- lets 11-17, 3-20 mm long, 1-5 mm wide, oblong to oblanceolate or obovate, truncate to retuse, mucronate, or acute, strigulose on both sides; peduncles 1-4 cm long; racemes 3- to 8-flowered, the flowers ascending at anthesis, the axis 1-4 cm long in fruit, bracts 1-3.5 mm long; pedicels 0.8-3.6 mm long; bracteoles 0-2; calyx 4.8-6.6 mm long, the tube 2.3-2.8 mm long, cam- panulate, villous, the teeth 2.3-2.8 mm long, subulate; flowers 6.3-8.2 mm long, pink-purple (soon fading yellowish); pods declined, sessile or nearly so, obliquely ovoid-ellipsoid, inflated, 13-21 mm long, 8-11 mm thick, straight or only slightly curved, villosulous with spreading-curved hairs, unilocular; ovules 20-28. Mixed desert shrub and pinyon-juniper communities usu- ally in sandy soils at 1270 to 1600 m in Emery (type from Cedar Mt.), Garfield, and Wayne cos.; endemic. The panther milk- vetch shares several salient features with the closely related but largely allopatric taxa A. pubentissimus and A. sabulonum (q.v.). From both of these, A. pardalimis differs in its straight or nearly straight pods, which bear 20-28 ovules (not 9-19). The main area of distribution of the panther milk- vetch is on the sandy eastern foot of the San Rafael Swell which breaks abruptly to the canyons of the Green and Colorado riv- ers; 13 (vii). Astragalus pattersonii A. Gray ex Brand, Patterson Milkvetch. [Tragacantha patter- sonii (A. Gray) Kuntze; Phacopsis pattersonii (A. Gray) Rydb.; Rydbergiella pattersonii (A. Gray) Fedde; Jonesiella pattersonii (A. Gray) Rydlj.]. Perennial, caulescent, 20-45 (50) cm tall, from a branching caudex; stems decum- bent to ascending or erect; stipules 3-8 mm long, all usually distinct; leaves 5-13 cm long; leaflets 7-15 or more, 6-38 mm long, 3-16 mm wide, elliptic to lanceolate, oblan- ceolate, or obovate, obtuse to acute, retuse, or mucronate, strigose to glabrous on both sides; peduncles 3-18 cm long; racemes 6-to 24-flowered, the flowers declined-nodding at anthesis, the axis 2-15 cm long in fruit; bracts 2-8 mm long; pedicels 1-4.5 mm long; bracteoles 2; calyx 8.8-14.2 mm long, the tube 6-8.8 mm long, cylindric, gibbous. pale tan or whitish, colored like the petals, thinly strigulose, the teeth 2.3-6.5 mm long, subulate; flowers 14-22 mm long, white, concolorous or the keel tip faintly purplish; pods erect, sessile, cylindric to ellipsoid or ovoid, 17-35 mm long, 6-10 mm thick, glabrous or puberulent, unilocular; ovules 22-38. Pinyon-juniper and mixed desert shrub communities at 1470 to 2300 m, dis- jimct in Utah, in Carbon, Garfield, Sevier, and Uintah cos.; Colorado and Arizona. The Patterson milkvetch is a primary selenium indicator, usually of clay and silty soils. The Arizona materials are known from the val- ley of Kanab Creek, and this plant might also occur in adjacent Kane Co. The Patter- son milkvetch is a handsome and striking plant, which is similar in many ways with A. praelongus (q.v.), from which it differs in the whitish, concolorous calyces and petals, and in the erect pods; 11 (vi). Astragalus perianus Barneby. Rydberg Milkvetch. Perennial, short-caulescent, 1-6 cm tall, from a shallowly subterranean, branching caudex; pubescence basifixed; stems prostrate, 3-12 cm long; stipules 1-2.5 mm long, at least some connate- sheathing; leaves 1-3 cm long; leaflets 7-19, 1-5 mm long, 1-3 mm wide, oval to ob- ovate, retuse, strigulose on both sides or gla- brate above; peduncles 0.3-2.2 cm long; racemes 2- to 6-flowered, the flowers spreading at anthesis, the axis 0.2-0.8 cm long in fruit; bracts 0.8-1.2 mm long; pedi- cels 1.4-21.5 mm long; bracteoles 0; calyx 3.5-4.2 mm long, the tube 2.3-3 mm long, campanulate, })ilosulous, purplish, the teeth 1-1.4 mm long, subulate; flowers 6.8-8.5 mm long, whitish, faintly suffused \vith pink or purple; pods ascending to declmed, ses- sile, bladdery-inflated, ovoid, 10-23 mm long, 8-14 mm wide (when pressed), stri- gose, purple-mottled, unilocular; ovules 18-20. Tertiary igneous gravels, often on barrens, in alpine sites at 3050 to 3350 m in Garfleld (Sevier Plateau) and Piute (Tushar Mts.) COS.; endemic. This beautiful low plant was first collected by P. A. Rydberg on 23 July 1905. It remained unnamed until its publication by Barneby (1964:973), and was not discovered a second time until 26 June 1975, some 70 years after its original Sept. 1978 Welsh: Utah Flora, Fabaceae 295 collection. The Rydberg milkvelch most closely resembles A. serpens (q.v.), but its relationship purportedly lies elsewhere; 7 (iv). Astragalus pinonis M. E. Jones. Pinyon Milkvetch. [Pisaphaca pinonis (M. E. Jones) Rydb.]. Perennial, caulescent, 10-55 cm tall, from a shallowly subterranean caudex; pu- bescence basifixed; stems erect or reclining, commonly growing through sagebrush cano- py; stipules 1.5-5 mm long, all distinct; leaves 2-11 cm long; leaflets 9-19, 2-18 mm long, 1-4 mm wide, linear to oblong, obtuse to ret\ise, strigose on both surfaces; peduncles 1.5-8 cm long; racemes 5- to 19- flowered, the flowers ascending-spreading at anthesis, the axis 2-7 cm long in fruit; bracts 1-2 mm long; pedicels 1-3 mm long; bracteoles 0-1; calyx 4.3-5.6 mm long, the tube 2.3-3.8 mm long, campanulate, stri- gose, the teeth 1-2 mm long, subulate; flowers 8.2-10.3 mm long, greenish to och- roleucous, suffused with purple; pods spreading-declined, subsessile, oblong- ellipsoid, straight to slightly curved, 20-35 mm long, 5.5-8.5 mm thick, terete or nearly so, strigose, unilocular; ovules 32-42. Sagebnish and sagebRish mixtures in pin- yon-juniper woodlands at 1580 to 2275 m in Beaver and Juab cos.; Nevada; a Great Ba- sin endemic. The pinyon milkvetch is ob- scure and apparently rare in Utah. The type was collected at Frisco, Beaver Co., where M. E. Jones found the plant in 1880 (on June 22). It grows up through low plants of sagebrush and is difficult to see. The plants resemble those of A. convallaritis, but the thicker pods and jointed terminal leaflets of A. pinonis are diagnostic; 2 (0). Astragalus platytropis A. Gray. Broad- keeled Milkvetch. [Tragacantha platytropis (A. Gray) Kuntze; Phaca platytropis (A. Gray) Rydb.; Cystium platytrope (A. Gray) Rydb.]. Perennial, acaulescent or nearly so, 2-7 cm tall, from a branching caudex, the branches often with a thatch of persistent leaf bases; pubescence basifixed; stems 0-2 cm long, prostrate-ascending or erect; stip- ules 1.5-5 mm long, at least some connate- sheathing; leaves 1-7 cm long; leaflets 5-15, 2-11 mm long, 1.5-7 mm wide, elliptic to obovate, oblong, or oval, acute to obtuse or retuse silvery-strigose on both sides; pe- duncles 1.5-6.5 cm long; racemes 2- to 9- flowered, the flowers ascending at anthesis, the axis 0.2-0.6 cm long in fruit; bracts 0.6-2 mm long; pedicels 0.7-1.9 mm long; bracteoles 0-2; calyx 3-5.4 mm long, the tube 2-3.4 mm long, campanulate, strigose, the teeth 0.2-2.1 mm long, subulate; flowers 7-9.5 mm long, pink-purple; pods ascend- ing, sessile, bladdery-inflated, ovoid to sub- globose, 15-33 mm long, 10-22 mm wide (when pressed), purple-mottled, strigulose, semibilocular; ovules 26-34. Ridge tops in shrub communities at 2400 to 3500 m in western Beaver and Tooele cos.; Nevada and California. This milkvetch is known in Utah only from the Mountain Home and Deep Creek ranges, where it occurs on limestone summits in crevices; 2 (i). Astragalus praelongus Sheldon. Stinking Milkvetch. Perennial, caulescent, 10-90 cm tall, from a branching caudex; pubescence basifixed; stems erect or ascending, forming clumps; stipules 2.5-9 mm long, all distinct; leaves 3-22 cm long; leaflets 7-33, 3-50 mm long, 2-24 mm wide, obovate, elliptic, oblong, lanceolate, or oblanceolate, obtuse or retuse to acute, sparingly strigose be- neath, glabrous above; peduncles 4-26 cm long; racemes 10- to 33-flowered, the flow- ers deflexed at anthesis, the axis 3-16 cm long in fruit; bracts 1-7 mm long; pedicels 1-7 mm long; bracteoles 2; calyx 5.8-14 mm long, the tube 4.4-7.5 mm long, cylin- dric, gibbous, glabrous, or thinly strigose, green or yellowish, usually differently col- ored than the petals, the teeth 0.3-6 mm long, subulate; flowers 15-24 mm long, och- roleucous, the keel often faintly purplish- tipped; pods erect to declined, sessile, sub- sessile, or stipitate, inflated, ellipsoid, ovoid, obovoid, or subglobose, 18-42 mm long, 15-25 mm thick, usually straight, glabrous or puberulent, subunilocular, leathery- woody; ovules 40-75. The stinking milk- vetch is represented in Utah by three more or less distinctive varieties. 296 Great Basin Naturalist Vol. 38, No. 3 1. Pods long-stipitate, the stipe 4-8 mm long; plants of San Juan Co. A. praelongiis var. lonchopus - Pods sessile or short-stipitate, the stipe, when present, less than 3 mm long; plants seldom of San Juan Co 2 2(1). Pods narrowly elliptic to oblong, 6-10 mm thick; plants of Beaver, Iron, Millard, and Sevier cos. eastward to east-central Utah ....A. praelongus var. ellisiae Pods broadly oblong to elliptis, 10-15 mm thick; plants of Garfield, Kane, Washington, and Wayne cos., and disjunctly in Sevier Co A. praelongus var. praelongus Var. ellisiae (Rydb.) Barneby in Turner, [Jonesiella ellisiae Rydb.]. Clay soil, com- monly on Mancos Shale, Monekopi, and Chinle formations, but also on alluvial sub- strates containing selenium, at 1330 to 1970 m in Beaver, Carbon, Emery, Grand, Iron, Millard, northern San Juan, Sevier, and Wayne cos.; Colorado, New Mexico, and Texas. The Ellis stinking milkvetch passes by degree into var. praelongus. It seems clear, however, that most of the Great Ba- sin populations are more closely allied with those of the San Rafael Swell region than they are with those of Washington Co. The relationship appears to have resulted from migration of plants of the ellisiae type through passes in the Wasatch Plateau rather than by a north-south exchange of typical phases; 54 (xviii). Var. lonchopus Barneby. Clay, seleni- ferous soils of various formations, in black- brush, mixed desert shrub, and pinyon- juniper communities at 1300 to 1925 m in southern San Juan Co.; Arizona; a Colorado Plateau endemic; 13 (iii). Var. praelongus [A. procerus Gray, not Boiss. & Housskn.; A. pattersonii var. prae- longus (Sheldon) M. E. Jones; Phacopsis praelongus (Sheldon) Rydb.; Rydbergeilla praelonga (Sheldon) Fedde & Sydow; Jone- siella praelonga (Sheldon) Rydb.]. Clay and silt of the Mancos Shale, Tropic Shale, Moenkopi, and Chinle formations, and other seleniferous soils, in salt desert shrub and pinyon-juniper communities at 770 to 2530 m in Garfield, Kane, Sevier, Washington, and Wayne cos.; Nevada, Arizona, and New Mexico. The type variety of stinking milk- vetch demonstrates much variation. The most extreme phase occurs in Washington and portions of Kane cos., and has fistulous tall stems and greatly thickened pods. Still another from Washington Co. has pinkish- lavender flowers. Most of these plants are primary selenophytes, however, a small population from near the mouth of Zion Canyon lacked the characteristic odor of se- lenium and presumably was not seleni- ferous; 81 (xxv). Astragalus preussii A. Gray. Preuss Milk- vetch. Perennial, caulescent, mostly 12-45 cm long, from a woody caudex; pubescence basifixed; stems erect or ascending, forming clumps; stipules 2-7 mm long, all distinct; leaves 3.5-13 cm long; leaflets 7-25, 1-28 mm long, 1-6 mm wide, obovate or obcor- date to oblong, narrowly elliptic, lanceolate, or linear, emarginate to rounded, obtuse, or acute, glabrous; peduncles 2-15 cm long; racemes 3- to 22-flowered, the flowers as- cending, the axis 1-20 cm long in fruit; bracts 1.5-4 mm long; pedicels 1-5.5 mm long; bracteoles 2; calyx 6.4-12.3 mm long, the tube 5.1-9.7 mm long, cylindric, thinly strigose, purple, the teeth 1.3-2.6 mm long, subulate; flowers, 14-24 mm long, pink- purple, or bicolored; pods erect to ascend- ing, stipitate, or subsessile, the stipe 2-7 mm long, the body oblong-ellipsoid, inflat- ed, 12-34 mm long, 6-13 mm thick, glabrous or puberulent, stiffly papery to leathery, imilocular; ovules 20-44. Two rather distinctive varieties are present in Utah. 1. Pods sessile or nearly so; racemes 4-20 cm long in fruit; plants of south- western Washington Co A. preussii var. kixiflorus Pods stipitate; racemes 1-9 cm long in fruit; plants of eastern Kane Co. and northeastward A. preussii var. preussii Sept. 1978 Welsh: Utah Flora, Fabaceae 297 Var. laxiflorus A. Gray. [A. preussii var. laxispicatus Sheldon; Phaca laxiflom (A. Gray) Rydb.]. Creosote bush community at 2200 to 2500 m in the Beaver Dam Moun- tains, Washington Co.; Nevada. This variety is not known from Utah in modern collec- tions, but contemporary specimens have been seen from Nevada; 1 (0). Var. preussii. [Phaca preussii (A. Gray) Rydb.; Tragacantha preussii (A. Gray) Kuntze; Rijdbergiella preussii (A. Gray) Tydb.; A. preussii var. latus M. E. Jones, type from Green River; A. preussii var. sul- catus M. E. Jones, type from Westwater; A. preussii var. arctus Sheldon; Rijdbergiella arcta (Sheldon) Tydb.; A. arctus (Sheldon) Tidestr.; Jonesiella arcta (Sheldon) Rydb.]. Blackbnish, mixed desert shrub, and salt desert shnib communities on seleniferous clays and silts at 1170 to 1570 m in Emery, Garfield, Grand, Kane, San Juan and Wayne COS.; Arizona, Nevada, and California. Our materials belong to var. latus M. E. Jones, sens, str., which differ from the more west- ern var. preussii, sens, str., in having a larger number of leaflets (17-25, not 11-15). Both (in sens, lat.) can be treated conservatively as belonging to a poly- morphic var. preussii; 54 (xxii). Astragalus pubentissimus Torr. & Gray. Green River Milkvetch. [A. multicaulis Nutt. ex Torr. & Gray, not Ledeb.; Trag- acantha pubentissima (Torr. & Gray) Kuntze; Phaca pubentissima (Torr. & Gray) Rydb.]. Perennial, or functionally annual, caulescent, 9-25 cm tall, from a rather weak caudex; pubescence basifixed; stems decumbent to ascending, radiating from the caudex; stipules 1-4.5 mm long, all distinct; leaves 2-8 cm long; leaflets 5-15, 2-14 mm long, 1.5-5 mm wide, oblong to ovate, obo- vate, or elliptic, obtuse to retuse, mucro- nate, or acute, villosulous on both sides; pe- duncles 1-3.5 cm long; racemes 3- to 12- flowered, the flowers spreading at anthesis, the axis 0.4-3.5 cm long in fruit; bracts 1-3 mm long; pedicels 0.5-2 mm long; brac- teoles 0; calyx 4.8-6.3 mm long, the tube 2.8-4.2 mm long, campanulate, villosulous, the teeth 1.8-2.8 mm long, subulate; flowers 8.8-12.2 mm long, pink-purple or ochro- leucous and suffused with purple; pods spreading-declined, sessile, inflated, oblique- ly lance-ellipsoid, 12-20 mm long, 4-8 mm thick, shaggy-pilose, unilocular; ovules 9-18. 1. Plants ascending to erect; flowers monochrome in populations A. pubentissimus var. pubentissimus Plants spreading-decumbent; flowers polychrome in populations A. pubentissimus var. peabodianus Var. peabodianus (M. E. Jones) Welsh comb. nov. based on Astragalus peabod- ianus M. E. Jones Zoe 3:295. 1893. The Peabody phase of the Green River milk- vetch occurs in entrenched channels cut into the escarpments draining the south and west flanks of the Tavaputs Plateaus, in pin- yon-juniper and mixed desert shrub commu- nities at 1300 to 1770 m in Emery and Grand (type from Thompson's Springs) cos.; endemic; 10 (ix). Var. pubentissimus. The Green River milkvetch occurs in pinyon-juniper and mixed desert shrub communities at 1525 to 1925 m in Duchesne and Uintah cos.; Wyoming and Colorado. Reports of live- stock poisoning, mainly of sheep, are at- tributable to this species. In years when it is abundant, the plants constitute the principal forb in much of the lower middle reaches of the Uinta Basin, reminiscent of seeded alfalfa fields among the open juniper-pinyon woodlands; 16 (i). Astragalus purshii Douglas ex Hook. Pursh Milkvetch. Perennial, acaulescent or subacaulescent, 4.5-13 cm tall, from thatch- clothed caudex branches; pubescence basi- fixed; stems 0-2 cm long; leaflets 5-17, 2-14 mm long, 1-7 mm wide, elliptic to oblanceolate, acute densely villous on both sides; pedimcles 1.5-10.5 cm long; racemes 1- to 7-flowered, the flowers ascending at anthesis, the axis 0.3-2 cm long in fruit; bracts 2-7 mm long; pedicels 1-5 mm long; 298 Great Basin Naturalist Vol. 38, No. 3 bracteoles 0; calyx 12-16 mm long, the tube 8.5-12.5 mm long, cylindric, villous-pilose, the teeth 2.2-6 mm long, subulate; flowers 19-26 mm long, whitish to ochroleucous or pink-purple; pods ascending, sessile. obliquely ovoid, usually curved, 13-26 mm long, 5-11 mm wide, shaggy- villous, uni- locular; ovules 20-34. Two varieties are present in Utah. 1. Flowers whitish or ochroleucous, discolorous, the keel purple-tipped; plants of broad distribution in northernmost counties A. purshii var. purshii Flowers pink-purple, concolorous; plants rare in Box Elder and Rich cos. A. purshii var. glareosus Var. glareosus (Douglas) Bameby. [A. glareosus Douglas ex Hook; Tragacantha glareosa (Douglas) Kuntze; A. inflexus var. glareosus (Douglas) M. E. Jones; Phaca gla- reosa (Douglas) Piper; Xylophacos glareosus (Douglas) Rydb.]. Sagebrush community at 1530 to 1830 m in Box Elder and Rich cos.; British Columbia south to Nevada. The var. glareosus simulates A. utahensis (q.v.) both in having pink-purple flowers and in habit. Leaflet shape and the flower structure are important diagnostic features; 2 (0). Var. purshii. [Tragacantha purshii (Douglas) Kuntze; Phaca purshii (Douglas) Piper; Xylophacos purshii (Douglas) Rydb.; A. purshii var. interior M. E. Jones]. Sage- brush, desert shrub, and pinyon-juniper communities at 1530 to 2270 m in Box El- der, Duchesne, Rich, and Uintah cos.; Washington east to Alberta and Saskatche- wan, and south to California, Nevada, Colo- rado, and South Dakota; 15 (ii). Astragalus racemosus Pursh. Alkali Milk- vetch. Perennial, caulescent, 20-53 cm tall, from a branching caudex; pubescence basi- fixed; stems erect or ascending, forming clumps; stipules 3-8 mm long, at least some cormate-sheathing; leaves 4-15 cm long; leaflets 9-19, 3-35 mm long, 1.5-13 mm wide, lance-elliptic to linear-lanceolate, acute to acuminate, glabrous or glabrate on both surfaces, sometimes ciliate; peduncles 3-13 cm long; racemes 9- to 45-flowered, the flowers nodding at anthesis, the axis 5-20 cm long, in fruit; bracts 1.5-7.5 mm long; pedicels 2-8 mm long; bracteoles 0-2; calyx 8.6-12 mm long, the tube 5-6 mm long, short-cylindric, gibbous, glabrous or sparingly strigose, the teeth 3.3-6 mm long, subulate; flowers 14-17 mm long, ochro- leucous, concolorous, or with the keel purple-tipped; pods pendulous, stipitate, the stipe 4-6 mm long, the oblong-ellipsoid body 10-21 mm long, 4-8 mm wide, tri- quetrous, glabrous, unilocular; ovules 12-22. Uinta and Duchesne River formations, rarely otherwise on saline clay and silty soils, at 1570 to 1970 m in Duchesne and Uintah cos; Wyoming. Our plants belong to var. treleasei C. L. Porter. They are pri- mary selenium indicators. Consumption by cattle of this and other selenophyte results in a "alkali disease," or "blind staggers." Plants from along Hill Creek in Uintah Co. have concolorous flowers; those from else- where are discolorous; 26 (v). Astragalus rafaelensis M. E. Jones. San Rafael Milkvetch. [Cnemidophacos rafae- lensis (M. E. Jones) Rydb.]. Perennial, cau- lescent, rush-like, 32-62 cm tall, from a branching caudex; pubescence basifixed; stems erect or ascending, arranged in clumps; stipules 1-5 mm long, at least some connate-sheathing; leaves 2.5-14.8 cm long, all simple, or some with leaflets 3-5, the lateral ones 3-20 mm long, 0.5-1.5 mm wide, linear to oblong, acute, glabrate be- neath, glabrous above, the terminal leaflet longer and confluent with the rachis; pe- duncles 11-27 cm long; racemes loosely 5- to 12-flowered, the flowers ascending to de- clined in anthesis, the axis 2-15 cm long in fruit; bracts 1.2-3.5 mm long; pedicels 2-5.5 mm long; bracteoles 2; calyx 6-9.6 mm long, the tube 5.2-7.5 mm long, short- cylindric, thinly strigose to glabrate, the teeth (0.8) 1.1-2.1 mm long, triangular; flowers 19-26 mm long, pale pink-purple or bicolored; pods deflexed, sessile, oblong- elliptic in outline, 12-25 mm long, 5-7 mm Sept. 1978 Welsh: Utah Flora, Fabaceae 299 wide, laterally compressed, glabrous, leathery-woody, unilocular, ovules 18-20. Seleniferous clays and silts of the Buckhorn Conglomerate, Morrison, Summerville, Chinle, and Moenkopi formations at 1370 to 1625 m in salt desert shrub community in Emery Co.; endemic. There are two main populations; one at the east base of the Ce- dar Mountain and the other in the vicinity of Window-blind Butte in the San Rafael Swell proper. This is a handsome seleno- phyte with close affinities to A. toanus and A. saurinus (q.v.), and lesser affinities to A. woodruffii (q.v.) 17 (vii). Astragalus sabulonum A. Gray. Gravel Milkvetch. [Phaca sabulonum (A. Gray) Rydb.; A. virgineus Sheldon]. Annual, win- ter annual, or biennial, caulescent, 4-30 cm tall, radiating from a taproot; pubescence basifixed; stems decumbent to ascending, rarely erect; stipules 1-4 mm long, all dis- tinct; leaves 1.5-7 cm long; leaflets 9-15, 2-13 mm long, 1-5 mm wide, oblanceolate to oblong or obovate, retuse to truncate or obtuse, loosely villous or both surfaces or glabrate above; peduncles 0.5-4 cm long; racemes 2- to 7-flowered, the flowers ascending-spreading at anthesis, the axis 0.3-2 cm long in fruit; bracts 1-2.5 mm long; pedicels 0.8-2 mm long; bracteoles 0; calyx 3.8-5.5 mm long, the tube 1.8-2.5 mm long, campanulate, hirsutulous, the teeth 1.8-3 mm long, subulate; flowers 6.2-8 mm long, pink-purple or less com- monly ochroleucous and tinged with purple; pods spreading-declined, sessile, obliquely ovoid, inflated, 9-17 mm long, 5-8 mm thick, curved, white-hirsutulous, unilocular; ovules 10-19. Mixed desert shrub, salt desert shrub, and lower pinyon-juniper com- munities, often in sand, at 1130 to 1700 m in Emery, Grand, Kane, San Juan, and Wayne cos.; California, Nevada, Arizona, and New Mexico. The gravel milkvetch is a near ally of A. pardalinus (q.v.), with which it is at least partially sympatric, and of A. pubentissimus whose geographical range is apparently distinct; 21 (vii). Astragalus sabulosus M. E. Jones. Cisco Milkvetch. [Jonesiella sabulosa (M. E. Jones) Rydb.]. Perennial, caulescent, 13-38 cm tall, from a woody caudex; pubescence basifixed; stems decumbent to ascending or erect, forming clumps; stipules 4-9 mm long, all distinct; leaves 3-10.5 cm long; leaflets 5-11, 6-35 (50) mm long, 3-17 mm wide, rhombic-oval to obovate or elliptic, mucro- nate, strigose to glabrous on both sides; pe- duncles 3.5-7 cm long; racemes 4- to 10- flowered, the flowers ascending-spreading at anthesis, the axis 0.5-2 cm long in fruit; bracts 2-6 mm long; pedicels 2-5 mm long; bracteoles 0-2; calyx 15-17.5 mm long, the tube 11.5-14 mm long, cylindric, strigulose, the teeth 3-4 mm long, subulate; flowers 28.5-34 mm long, whitish to ochroleucous, fading yellowish; pods spreading to de- clined, subsessile, inflated, cylindroid, 20-48 mm long, 10-15 mm thick, stiffly papery to leathery, strigose, unilocular; ovules 55-59. Salt desert shrub at 1300 to 1600 m on Mancos Shale east of Thompson in the Grand River Valley in Grand Co.; endemic. The Cisco milkvetch is a primary selenium indicator with close affinities to A. iselyi (q.v.), from which it can be distinguished by its larger yellowish flowers. The Cisco milk- vetch is also allied with A. praelongus which has much smaller flowers and pods. The flowers of A. sabulosus are the largest within Astragalus in Utah, and possibly else- where (though they may not be the long- est); 9 (vi). Astragalus saurinus Barneby. Dinosaur Milkvetch. Perennial, caulescent, rush-like, 25-45 cm tall, from a shallowly sub- terranean caudex; pubescence basifixed; stems erect or ascending, arranged in clumps; stipules 1.2-7 mm long, usually at least some shortly connate-sheathing; leaves 2.5-9 cm long, the uppermost usually simple, the others with leaflets 3-9, mostly 10-28 mm long, 0.5-2 mm wide, linear to linear-elliptic, obtuse to acute, strigose on both surfaces, the terminal leaflet confluent with the rachis; peduncles 7-21 cm long; racemes 3- to 15-flowered, the flowers ascending-spreading at anthesis, the axis 0.5-6 cm long in fruit; bracts 1.2-2 mm long; pedicels 1.5-3 mm long; bracteoles 1-2; calyx 6.4-9.6 mm long, the tube 5.6-6.7 mm long, cylindric, the teeth 0.9-2.9 mm long, triangular-subulate; flow- ers 18-22 mm long, bicolored, pink-purple. 300 Great Basin Naturalist Vol. 38, No. 3 with white wing-tips, rarely all white; pods deflexed, sessile or nearly so, narrowly ob- long in outline, straight or curved, 15-35 mm long, 4.4-6 mm wide, laterally com- pressed, strigose to glabrate, unilocular; ovules 19-29. Duchesne River, Morrison, Carmel, Chinle, and Moenkopi formations in salt desert shrub and pinyon-juniper com- mimities at 1430 to 1700 m in Uintah Co.; endemic. The dinosaur milkvetch is a pri- mary selenophyte with two near gongeners in Utah; A. toaniis of far western Utah, and A. rafaelensis of the San Rafael Swell; 17 (vi). Astragalus scopulorum T. C. Porter. Rocky Mountain Milkvetch. [Tragacantha scopulorum (T. C. Porter) Kuntze; Tium scopulorum (T. C. Porter) Rydb.]. Perennial, caulescent, 15-48 cm tall, from a shallowly subterranean caudex; pubescence basifixed; stems decumbent to ascending, radiating from the caudex; stipules 3-9 mm long, at least some connate-sheathing; leaves 1.5-8.5 cm long; leaflets 15-29, 2-18 mm long, 1-8 mm wide, oblong to elliptic or oblanceo- late, some narrowly so, acute to obtuse or mucronate, thinly strigose to glabrous be- neath, glabrous above, thinly ciliate; pe- duncles 2-14 cm long; racemes 4- to 22- flowered, the flowers declined to nodding at anthesis, the axis 1-7 cm long in fruit; bracts 1.5-7 mm long; pedicels 1-4 mm long; bracteoles 0-2; calyx 9-11.5 mm long; the tube 6.8-8.5 mm long, subcylindric, strigulose, the teeth 1.5-3.5 mm long; flow- ers 18-24 mm long, ochroleucous, con- colorous or the keel faintly purplish; pods pendulous, stipitate, the stipe 4-9 mm long, the body oblong, straight or curved, 18-35 mm long, 3-6.5 mm wide, triquetrous, glabrous, bilocular. Mountain brush, sage- brush, ponderosa pine, pinyon-juniper and aspen-white fir communities at 1670 to 2430 m in Carbon, Grand, eastern Millard, San Juan, Sanpete, Sevier, Summit, Utah, and Wasatch cos.; Colorado, Arizona, and New Mexico. Continued exploration of the plateaus and mountain ranges of central and northern Utah has demonstrated a large area occupied by this distinctive milkvetch of middle elevation brush and woodland; 29 Astragalus serpens M. E. Jones. Plateau Milkvetch. [Phaca serpens (M. E. Jones) Rydb.]. Perennial, caulescent, 3.5-23 cm long, radiating from a weak caudex; pu- bescence basifixed; stems decumbent to ascending-erect; stipules 1.5-3.5 mm long, all distinct; leaves 1.5-4.5 cm long; leaflets 9-15, 2-9 mm long, 1-4 mm wide, obovate to oblanceolate or elliptic, obtuse to emarginate, strigose-pilosulous beneath and above, or glabrate above; peduncles 0.7-2.5 cm long; racemes shortly 2- to 9-flowered, the flowers spreading at anthesis, the axis 0.2-0.8 cm long in fruit; bracts 1-1.5 mm long; pedicels 1-1.8 mm long; bracteoles 0; calyx 4.2-5 mm long, the tube 2.7-3.5 mm long, campanulate, white strigulose (some black hairs sometimes present), the teeth 1.5-2 mm long, subulate; flowers 6.6-8.6 mm long, purplish to pink-purple or whit- ish; pods ascending to declined, stipitate, the stipe (gynophore) 0.7-1.5 mm long, the body bladdery-inflated, ovoid or ellipsoid, 13-29 mm long, 7-17 mm wide (when pressed), red-mottled, strigose, unilocular. Sagebrush, pinyon-juniper, aspen, and aspen-fir communities at 2070 to 2800 m in Garfield, Kane, Iron, Piute, and Wayne cos.; endemic. Barneby (1964) places A. serpens adjacent to A. pubentissimus, a near con- gener of A. sahulonum and A. pardalinus. The similarity to A. perianus was dis- counted, but examination of both species in the field and in the herbarium indicates that the nearest ally of A. serpens is A. perianus, despite the morphological differences noted by Barneby (i.e., the sessile pod, sub- terranean root-crown, and connate stipules) for A. perianus. In some years A. serpens is abundant in the Loa Pass vicinity where it occupies spaces between low sagebrush and igneous gravel; 19 (iii). Astragalus sesquiflorus S. Wats. Sand- stone Milkvetch. [Tragacantha sesquiflora (S. Wats.) Kuntze; Batidophaca sesquiflora (S. Wats.) Rydb.; A. sesquiflorus var. bre- vipes Barneby]. Perennial, caulescent, pros- trate, often mat-forming, 0.5-5 cm tall, radiating from a branching caudex; pu- bescence malpighian; stems 5-28 cm long or more; stipules 1.5-4.5 mm long, all con- nate-sheathing; leaves 1-4 cm long; leaflets Sept. 1978 Welsh: Utah Flora, Fabaceae 301 7-13, 1.5-10 mm long, 0.6-2 mm wide, el- liptic to obovate, acute to obtuse, strigose on both sides (commonly involute); pe- duncles 0.8-4.5 cm long; racemes 1- to 4- flowered, the flowers ascending at anthesis, the axis very short in fruit; bracts 1.2-3 mm long; pedicels 0.7-4 mm long; bracteoles 0-2; calyx 3.7-5.5 mm long, the tube 1.5-2.8 mm long, campanulate, strigulose, the teeth 1.9-3 mm long, subulate; flowers 6-8 mm long, pink-purple; pods spreading- ascending, sessile or subsessile, the body obliquely oblong in outline, 8-10 mm long, 3-4 mm wide, trigonously compressed, strigulose, mixed desert shrub, pinyon-juni- per, and ponderosa pine or aspen commu- nities at 1570 to 2930 m in Kane, San Juan, and disjunctly (accidentally?) in Sanpete COS.; Arizona; a Mohave-San Juan endemic. The sandstone milkevetch is more closely allied to A. humistratus than to other spe- cies of milkvetch in Utah; 21 (iv). Astragalus spatulatus Sheldon. Draba Milkvetch. [Homalohus caespitosus Nutt.; A. caespitosus (Nutt.) A. Gray, not Pallas; Tragacantha caespitosa (Nutt.) Kuntze; A. simplificolius var. spatulatus (Sheldon) M. E. Jones; H. canescens Nutt. ex Torr. & Gray; H. brachycarpus Nutt. ex Torr. & Gray; A. simplex Tidestr., not A. brachij- carpus MB.; A. spatulatus var. simplex Tidestr.; H. uniflorus Rydb.; A. spatulatus var. uniflorus (Rydb.) Barneby]. Perennial, acaulescent, 1.5-9 cm tall, from a branching caudex; pubescence malpighian; stems ob- scured by marcescent leaf bases and stip- ules, tufted, sometimes mat-forming; stipules 2-7 mm long, all connate-sheathing; leaves all or mostly all simple (reduced to phyl- lodia), 0.8-6 cm long, only rarely with leaf- lets 3-5 on some leaves and with the termi- nal one confluent with the rachis, oblanceolate to linear, acute, mucronate, or spinulose, strigose on both surfaces; pe- duncles 0.4-7 cm long; racemes 1- to 11- flowered, the flowers ascending at anthesis, the axis 0.2-3 cm long, in fruit; bracts 0.6-3 mm long; calyx 2.6-5 mm long, the tube 1.9-3.4 mm long, campanulate, strigose, the teeth 0.5-2.5 mm long, subulate; flowers 5.7-9.5 mm long, pink-purplish to ochro- leucous or whitish (in populations); pods erect, sessile, lanceolate to lance- oblong in outline, 4-13 mm long, 1.5-3 mm wide, straight or slightly curved, strigose to rarely glabrous, unilocular; ovules 4-12. Pinyon-juniper, sagebnish, and mountain brush communities, often on exposed ridges, at 1530 to 2630 m in Cache, Carbon, Dag- gett, Duchesne, Emery, Grand, Rich, Uin- tah, and Wasatch cos. (to be expected in Sanpete and Summit); Alberta and Sas- katchewan, south to Idaho, Colorado, and Nebraska. The draba milkvetch is most closely allied to A. chloodes and A. detritalis among our species; 35 (x). Astragalus straturensis M. E. Jones. Sil- ver Reef Milkvetch. [Atelophragma stratu- rense (M. E. Jones) Rydb.; Hamosa atrata Rydb., type from southern Utah; Tium atratiforme (Rydb.) Rydb.]. Perennial, cau- lescent, 13-36 cm tall, from a superficial to shallowly subterranean caudex; pubescence basifixed; stems decumbent to ascending or erect, forming clumps; stipules 1-2.5 mm long, all distinct; leaves 3.5-9.5 cm long; leaflets 9-19, 3-13 mm long, 1-5 mm wide, oblong to linear or oval, obtuse or retuse, strigose beneath, glabrous above; peduncles 1.5-7.5 cm long; racemes 9- to 25-flowered, the flowers ascending to declined at an- thesis, the axis 1.5-14 cm long in fruit; bracts 0.8-1.5 mm long; pedicels 0.8-1.5 mm long; bracteoles 0; calyx 3.5-4.2 mm long, the tube 2.5-3.5 mm long, cam- panulate, strigose, the teeth 0.7-1 mm long, triangular; flowers 6.5-8.5 mm long, pink- purple with white wing-tips; pods pen- dulous, stipitate, the stipe 1.4-2 mm long, the body oblong in outline, curved or straight, 10-15 mm long, 2.2-3 mm wide, strigose, bilocular; ovules 10-13. Sagebrush, pinyon-juniper, and mountain brush commu- nities at 1530 to 2130 m in Beaver, Iron, southeastern Millard, and Washington (type from Silver Reef) cos.; Arizona and Nevada. The Silver Reef milkvetch has no close rela- tives in the milkvetch flora of Utah. In flower size and general habit, it resembles A. wingatanus of southeastern Utah, but the resemblance is only superficial; 10 (iii). Astragalus striatiflorus M. E. Jones. Es- carpment Milkvetch. Perennial, sub- acaulescent to short-caulescent, 1.5-6 cm 302 Great Basin Naturalist Vol. 38, No. 3 tall, radiating from a branching, usually sub- terranean caudex; pubescence basifixed; stems 0-5 cm long, only the tips produced above the sand; stipules 2-4 mm long, all connate-sheathing; leaves 1-4 cm long; leaf- lets 5-13, 1-7 mm long, 0.8-2.5 mm wide, ovate to obovate or oblanceolate, obtuse, mucronate, or emarginate, pilosulous; pe- duncles 1-3 cm long; racemes 2- to 5- flowered, the flowers ascending in anthesis, the axis 0.2-1 cm long; pedicels 1-1.5 mm long; bracteoles 0; calyx 5.5-7 mm long, the tube 3-4 mm long, campanulate, hirsu- tulous, the teeth 1.8-3 mm long, subulate; flowers 9-12 mm long, pink-purple or whit- ish and commonly suffused with purple, the keel-tip purple, long-attenuated, with the stigma protruding; pods spreading, sessile, the body bladdery-inflated, ellipsoid, 12-18 mm long, 8-15 wide (when pressed), mottled, spreading-hairy, bilocular. Interdune valleys, sandy depressions on ledges, and bars and terraces in stream channels at 1530 to 1900 m in Kane (from the Paria River westward) and eastern Washington cos.; Coconino Co., Arizona. The escarpment milkvetch is a singular plant, resembling A. perianus more closely than any other in Utah. The attenu- ate keel-tip, protruding stigma, and inflated bilocular pods are both unusual and diag- nostic features; 12 (iv). Astragalus subcinereus A. Gray. Silver Milkvetch. [Phaca subcinerea (Gray) Rydb.; A. sileranus M. E. Jones, type from Sink Valley (near Alton); Phaca silerana (M. E. Jones) Rydb.; A. sileranus var. cariacus M. E. Jones, type from Elk Head Ranch on the upper Virgin River]. Perennial, caulescent, prostrate, 14-90 cm long, radiating from a subterranean branching caudex; pubescence basifixed; stems prostrate to weakly ascend- ing; stipules 1.5-6.5 mm long, at least some connate-sheathing; leaves 1.5-8.5 cm long; leaflets 9-23, 2-16 mm long, 1-8.5 (10) mm wide, oblong to oblanceolate or obovate, obtuse, emarginate, or retuse, villosulous on both surfaces, or glabrate above; peduncles 1.5-10 cm long; racemes 5- to 37-flowered, the flowers ascending to declined at an- thesis, the axis 1-7 cm long in fruit; bracts 1-3 mm long; pedicels 0.5-2.5 mm long; bracteoles 0-1; calyx 3.4-6.3 mm long, the tube 2.3-3.6 mm long, campanulate, villos- ulous, the teeth 0.9-2.9 mm long, subulate; flowers 6-11 mm long, ochroleucous and commonly suffused with purple; pods spreading to declined, subsessile, inflated, ovoid-ellipsoid to ellipsoid, 12-27 mm long, (3.5) 6-13 mm wide (when pressed), sub- terete to dorsiventrally compressed, thinly villosulous, mottled; ovules 10-20. Barneby (1964: 214) discussed the variation within A. subcinereus sens. lat. noting the morpholog- ical variants in each major segment of the species at large. Much of the material from Kane, Garfield, and Washington counties differs from the typical plants in Mohave County, Arizona in being more leafy (the leaflets 4-10 mm broad), in having longer stems (3-7 dm long), and in having more firmly walled pods 15-28 mm long and 6-10 (13) mm thick. These Utah plants be- long, sens, str., to var. cariacus M. E. Jones, but the features are weak and overlapping at best. There is another variant, not seen by Barneby, that is seemingly worthy of taxonomic recognition. It is separable as fol- lows: Mature pods elliptic-oblong to oblong, 3.5-5.5 mm wide; flowers 8.5-11 mm long; stems 40-90 cm long; plants of volcanic gravels in eastern Sevier and Emery cos ^- subcinereus var. basalticus Mature pods ovoid-ellipsoid, 6-13 mm wide; flowers 6-9 mm long; stems mostly 14-70 cm long; plants commonly of sedimentary gravels, sometimes from igneous gravels, but not in Sevier or Emery cos A. subcinereus var. subcinereus Var. basalticus Welsh var. nov. Astragalo 6-13 mm), floribus magnioribus (8.5-11 subcinereo var. subcinero aemulans, differt mm, nee 6-9 mm), et caulis longioribus leguminibus angustioribus (3.5-5.5 mm, nee (40-90 cm, nee 14-70 cm). Holotype. Utah, Sept. 1978 Welsh: Utah Flora, Fabaceae 303 Sevier Co., open pine woodland adjacent to Utah Highway #72, ca. 10 miles south of junction with Utah Highway #4, Welsh, Isely, Moore 6447, 23 July 1967 (BRY). Par- atypes: Utah, Emery Co., 7 mi S of 1-70, Road to Baker Ranch, Welsh 14788, 18 May 1977 (BRY); Sevier Co., igneous boul- dery slope with juniper, along Utah High- way #72, 5 miles south of junction with Utah Highway #4, Welsh, Isely, Moore 6445, 23 July 1967 (BRY) 12 mi due SSW of Emery, Welsh & Clark 16186, 23 Aug 1977 (BRY). The plants grow at 1830 to 2430 m in pinyon-juniper and sparse ponderosa pine woods in Emery and Sevier COS.; endemic; 4 (iv). Var. subcinereus. Ponderosa pine, pinyon- juniper, and sagebrush communities at 1670 to 2270 m in Garfield, Iron, Kane, and Washington cos.; Lincoln Co., Nevada, and Mohave and Coconino cos.; Arizona; 13 (ii). Astragalus tenellus Pursh. Pulse Milk- vetch. [Erviwi 7nultiflorum Pursh; Homa- lohus multiflorus (Pursh) Nutt. ex Torr. & Gray; A. multiflorus (Pursh) A. Gray; Trag- acantha multiflora (Pursh) Kuntze; H. te- nellus (Pursh) Britton in Britton & Brown; H. strigulosus Rydb.; A. tenellus f. strigu- hsus (Rydb.) Macbr.; A. tenellus var. strigu- losus (Rydb.) F. J. Hermann]. Perennial, caulescent, 10-52 cm tall, from a branching caudex; pubescence basifixed; stems erect, ascending, or less commonly decumbent, forming clumps; stipules 1.5-7 mm long, turning black in drying, at least some connate-sheathing; leaves 2-9 cm long; leaf- lets 11-21, 3-24 mm long, 0.4-6 mm wide, narrowly oblong to elliptic, linear, oblan- ceolate or obovate, acute to obtuse, mucro- nate, or emarginate, thinly strigose beneath, glabrous above; peduncles 0.2-4 cm long, often paired; racemes (1) 3- to 23-flowered, the flowers ascending at anthesis, the axis 0.5-11 cm long in fruit; bracts 0.5-2.7 mm long; pedicels 0.7-3.2 mm long; bracteoles 0-2; calyx 2.6-5.2 mm long, the tube 2-2.7 mm long, campanulate, strigose, the teeth 0.7-2.5 mm long; flowers 6-9 mm long, white to ochroleucous; pods pendulous, stipitate, the stipe 0.6-5.5 mm long, the body elliptic to oblong in outline, straight or curved, 7-16 mm long, 2.5-4.5 mm wide, laterally flattened, glabrous or less commonly strigose, unilocular; ovules 3-9. Mountain brush, sagebrush, pinyon-juniper, ponderosa pine, aspen-fir, and spruce-fir communities at 1670 to 2900 m in Box El- der, Cache, Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Iron, Millard, Piute, Sanpete, Sevier, Summit, Uintah, Utah, Wasatch, and Wayne cos.; Yukon east to northern Manitoba and south to Nevada, New Mexico, Nebraska, and Minnesota. The pulse milkvetch is similar to many salient morphological features to A. wingatanus, but the subterranean caudex, habit of growth, and elongated fruiting racemes of that species indicate affinities elsewhere, even though the fruit and the tiny flowers are similar; 90 (xx). Astragalus tephrodes A. Gray. Ashen Milkvetch. [A. shortianus var. brachylobus A. Gray; Xylophacos brachylobus (A. Gray) Rydb.; A. curtilobus Tidestr., not A. brachy- lobus DC.]. Perennial, acaulescent to short- caulescent, 5-15 cm tall, from a branching caudex; pubescence basifixed; stems 0-8 cm long, the internodes commonly obscured by stipules; stipules 2-11 mm long, all distinct, or rarely some shortly connate-sheathing; leaves 4-16 cm long; leaflets 11-31, 3-17 mm long, 2-11 mm broad, obovate to ob- lanceolate, elliptic, or orbicular, obtuse, acute, or emarginate, strigulose to pilosulous on both sides or glabrous above; peduncles 4-15 cm long; racemes 10- to 25-flowered the flowers ascending at anthesis, the axis 1.5-8 cm long in fruit; bracts 1.5-9 mm long; pedicels 0.6-3.4 mm long; bracteoles 0; calyx 8.8-12.7 mm long, the tube 7.1-10 mm long, cylindric, pilosulous, the teeth 1.7-2.8 mm long, subulate; flowers 15-24 mm long, pink-purple; pods ascending, el- lipsoid to lance-ellipsoid, 17-30 mm long, 6-10 mm thick, strigulose to pilosulous, uni- locular; ovules 24-35. Known in Utah with certainty only on the basis of a collection reported by Barneby (1964) from Washing- ton Co. (Springdale, Jones in 1894, POM); California, Arizona, and New Mexico. Our material reportedly belongs to var. brachy- lobus (A. Gray) Barneby. Astragalus tetrapterus A. Gray. Four- wing Milkvetch. [Pterophacos tetrapterus (A. 304 Great Basin Naturalist Vol. 38, No. 3 Gray) Rydl:).]. Perennial, caulescent, 10-35 cm tall, from a subterranean caudex; pu- bescence basifixed; stems erect or ascending, or finally decumbent; stipules 2-5.5 mm long, all distinct; leaves 1.5-8.5 cm long; leaflets 9-21, 1-33 mm long, 0.3-3.2 mm wide, linear, narrowly oblong, or elliptic, obtuse to acute, strigose to glabrous on both sides, at least some terminal leaflets con- fluent with the rachis in uppermost leaves; peduncles 1-6.5 cm long; racemes 6- to 15- flowered, the flowers ascending at anthesis, the axis 1-4 cm long in fruit; bracts 1.5-3.5 mm long; pedicels 1.4-4.3 mm long; brac- teoles 0-2; calyx 5.5-8.7 mm long, the tube 4.7-7 mm long, cylindric, strigose, the teeth 0.8-2.8 mm long, subulate; flowers 15-19 mm long, white to yellowish tinged faintly with pink, the keel faintly purple-tipped; pods pendulous, sessile, obliquely oblong in outline, curved or coiled, 20-40 mm long, 6-10 mm wide, succulent at first, ultimately (by collapse of fleshy walls), sharply 4- angled, glabrous or strigose, unilocular; ovules 28-38. Pinyon-juniper and sagebrush communities at 1030 to 2130 m in Beaver, Iron, Kane, and Washington (type from 25 miles north of St. George) cos.; Oregon, Ne- vada, and Arizona. The four-wing milkvetch has been reported to produce locoism in livestock, but it is seldom sufficiently abun- dant to produce serious, large-scale losses; 9 (iii). Astragalus toanus M. E. Jones. Toana Milkvetch. [Cnemidophacos toanus (M. E. Jones) Rydb.]. Perennial, caulescent, rush- like, 15-50 cm tall, from shallowly sub- terranean to superficial caudex; pubescence basifixed; stems erect or ascending, in clumps; stipules 1.5-6.5 mm long, at least some connate-sheathing; leaves 2-10 cm long, the uppermost, rarely all, simple, or with the terminal leaflet confluent with the rachis, the lower ones with leaflets 3-9, 3-30 mm long, 1.4-2.5 mm wide, linear- filiform to oblong, obtuse to acute, strigose or glabrous on both sides; peduncles 6-25 cm long; racemes 7- to 35-flowered, the flowers ascending at anthesis, the axis 3-30 cm long in fruit; bracts 1-3 mm long; pedi- cels 0.8-3.5 mm long; bracteoles 0-2; calyx 4.6-8 mm long, the tube 4.1-6.4 mm long, short-cylindric, strigose, the teeth 0.5-2 mm long, subulate; flowers 15-20 mm long, pink-purple with wing-tips white; pods erect, sessile, oblong in outline, 13-25 mm long, 3.7-5.5 mm wide, slightly compressed laterally glabrous to strigose, unilocular; ovules 14-26. Seleniferous clay soils in salt desert shrub communities at 1530 to 1770 m in Box Elder and Millard cos.; Nevada, Oregon, and Idaho. The Toano milkvetch has affinities with both A. saurinus of the Uinta Basin, and A. rafaelensis of the San Rafael Swell. No intermediates are known; 12 (i). Astragalus utahensis (Torr.) Torr. & Gray. Utah Milkvetch. [Phaca millissima var. utahensis Torr. in Stansbury; Trag- acantha utahensis (Torr.) Kuntze; Xylo- pfuicos utahensis (Torr.) Rydb.]. Perennial, subacaulescent, mostly 2-12 cm tall, radi- ating from a branching caudex; pubescence basifixed; stems 0-10 cm long, the inter- nodes usually concealed by stipules, pros- trate; stipules 3-10 mm long, all distinct; leaves 1.5-12 cm long; leaflets 9-19, 2-15 mm long, 2-12 mm wide, obovate or sub- orbicular to ovate, obtuse to emarginate, rarely acute, densely villous-tomentose on both surfaces; peduncles 1-14 cm long; racemes 2- to 8-flowered, the flowers as- cending at anthesis, the axis 0.4-2.6 cm long in fniit; bracts 4-9 mm long; pedicels 2-4.3 mm long; bracteoles 0-2; calyx 12-14 mm long, the tube 8.5-13 mm long, cylin- dric, villous-tomentose, the teeth 2-4.5 mm long, lance-subulate; flowers 23-31 mm long, pink-purple (rarely white); pods as- cending, sessile or stipitate, the stipe (gyno- phore) 1-2.5 mm long, the body 17-30 mm long, 5.5-7.5 mm wide, obscured by long shaggy-villous hairs, unilocular; ovules 22-31. Sagebrush, pinyon-juniper, mountain brush, and grassland communities at 1250 to 2130 m in all Utah counties except Daggett, Emery, Grand, Iron (to be expected), Kane, Rich, San Juan, and Uintah; Idaho and Ne- vada. The Utah milkvetch is known locally as "ladyslipper," because of a fancied re- semblance of the large flowers to softly cushiony house-slippers. The plants are abundant along the Wasatch front, where they flower in April and May, much to the Sept. 1978 Welsh: Utah Flora, Fabaceae 305 delight of beginning students in taxonomy, each of whom feels compelled to collect at least one plant and deposit it in a herba- rium; 163 (xii). Astragalus wardii A. Gray. Ward Milk- vetch. [Phaca wardii (A. Gray) Rydb.]. Per- ennial, caulescent, 9-50 cm tall, from a branching, superficial (rarely subterranean) caudex; pubescence basifixed; stems decum- bent to erect; stipules 1-3 mm long, all dis- tinct; leaves 3-10 cm long; leaflets 17-23, 3-11 mm long, 0.8-6 mm wide, oblanceo- late to elliptic or narrowly oblong to linear, obtuse to retuse or emarginate, strigose only on the midrib beneath and on margins; pe- duncles 1.5-5 cm long; racemes loosely 5-to 15-flowered, the flowers ascending to de- clined at anthesis, the axis 1-5 cm long in fruit; bracts 1-2.2 mm long; pedicels 1-3.4 mm long; bracteoles 0-2; calyx 2.9-4.6 mm long, the tube 1.7-2.3 mm long, cam- panulate, strigose with black, less commonly black mixed with white, hairs, the teeth 1-2.4 mm long, subulate; flowers 5-8 mm long, whitish or ochroleucous; pods pen- dulous to spreading, sessile or on a gyno- phore about as broad as long, bladdery- inflated, 15-28 mm long, 9-17 mm wide (when pressed), mottled or not, glabrous, unilocular; ovules 12-17. Sagebnish, cotton- wood, pinyon-juniper, ponderosa pine and spruce-fir communities, or less commonly in grassland and salt desert shrub communities, at 1530 to 2730 m in Garfield, Kane, Piute, and Sevier cos.; endemic. The Ward milk- vetch is similar in its inflated, unilocular pods, distinct stipules, and small flowers with the closely contiguous, if not sympa- tric, A. serpens, from which it can be distin- guished by the more numerous leaflets, pre- dominately black strigose calyces, and glabrous pods. The presence of an incipient gynophore in A. wardii strengthens the sim- ilaritv between these distinctive species; 24 (ii). ' Astragalus wetherillii M. E. Jones. Weth- erill Milkvetch. [Phaca wetherillii (M. E. Jones) Rydb.]. Perennial, caulescent, 4-26 cm tall, from a branching caudex; pu- bescence basifixed; stems decumbent to as- cending, in clumps; stipules 1.2-3.5 mm long, all distinct; leaves 2-10 cm long; leaf- lets 7-15, 3-14 mm long, 2-9 mm wide, obovate to oval, obtuse to emarginate or mucronate, thinly strigose beneath, glabrous above; peduncles 1.5-4.5 cm long; racemes 2- to 9-flowered, the flowers ascending to declined at anthesis, the axis 0.3-2.3 mm long in fruit; bracts 1-2.5 mm long; pedi- cels 1-2.5 mm long; bracteoles 0; calyx 4.6-6.2 mm long, the tube 2.8-3.8 mm long, campanulate, strigose, the teeth 1.8-2.4 mm long, subulate; flowers 7.5-11 mm long, whitish or tinged lavender; pods spreading to declined or shortly stipitate, the stipe (gynophore) 1-2.5 mm long, the body inflat- ed, ovoid-ellipsoid, slightly curved, 14-22 mm long, strigulose, unilocular; ovules 9-13. Mountain brush and pinyon-juniper commu- nities at 1430 to 1770 m in Grand Co. (col- lected by Eastwood according to Jones 1923); central western Colorado. The Utah collection is known from the canyon of the Colorado River east of Moab. It has not been recollected in many years, but likely it persists along the canyon slopes on the shaded side. It is a peculiar species, not quite like any other known from Utah. The pods of early produced flowers are often mature and brown while progressively younger pods and flowers continue to be produced as the plants elongate; 2 (ii, both from Colorado). Astragalus wingatanus S. Wats. Fort Wingate Milkvetch. [Homalobus wingatanus (S. Wats.) Rydb.; A. dodgeanus M. E. Jones, type from Thompson's Springs; A. wing- atanus var. dodgeanus (M. E. Jones) M. E. Jones]. Perennial, caulescent, 15-45 (60) cm tall, from a subterranean caudex; pu- bescence basifixed; stems spreading- ascending, forming diffuse clumps; stipules 1.5-5 mm long, at least some connate- sheathing; leaves 1.5-6.5 cm long; leaflets 7-15 (17) 3-18 mm long, 0.4-3.6 mm wide, linear to narrowly oblong, elliptic or oblan- ceolate, acute, obtuse, or retuse, strigose to glabrous beneath, glabrous above, often ci- liate; peduncles 2-14 cm long; racemes very loosely 7- to 35-flowered, the flowers as- cending at anthesis, the axis 3-18 cm long in fruit; bracts 0.5-2 mm long; pedicels 0.8-3 mm long; bracteoles 0-2; calyx 2.5-3.7 mm long, the tube 1.5-2.6 mm long, 306 Great Basin Naturalist Vol. 38, No. 3 campanulate, strigose, the teeth 0.4-1.4 mm long, triangular-subulate; flowers 5.5-8 mm long, pink-purple, the wing-tips white or pale; pods deflexed, subsessile or short-stipi- tate, the stipe to 1.7 mm long, the body el- liptic to oblong in outline, straight or slightly curved, 9-15 mm long, 3-4.5 mm wide, compressed, glabrous, unilocular; ovules 4-8. Pinyon-juniper, mixed desert shrub, salt desert shrub, and less commonly in mountain brush communities at 1530 to 2130 m in Carbon, Duchesne, Emery, Grand, and San Juan cos.; Colorado, New Mexico, and Arizona. The Fort Wingate milkvetch simulates A. tenellus (q.v.), in its small flowers and laterally flattened pods. The buried caudex and elongated fruiting racemes are diagnostic for A. wingataniis, and indicate relationships elsewhere (i.e., with scytocarpous taxa such as A. flexuosus var. diehlii); 29 (xii). Astragalus woodruffii M. E. Jones. Woodruff Milkvetch. [Homalobus wood- ruffii (M. E. Jones) Rydb.]. Perennial, cau- lescent, rush-like, 25-55 (65) cm tall, from a deeply subterranean caudex; pubescence basifixed or incipiently malpighian; stems erect or ascending in broom-like clumps; stipules 10-25 mm long, at least some usu- ally connate-sheathing; leaves 1.5-6.5 (8) cm long, at least the upper ones simple, the others with decurrent leaflets 2-9, 2-17 mm long, 0.5-2 mm wide, acute to obtuse, silvery-strigose on both surfaces, the termi- nal leaflet decurrent also; peduncles 3.5-16 cm long; racemes 8- to 45-flowered, the flowers ascending at anthesis, the axis 2-25 cm long in fruit; bracts 2.5-7 mm long; pedicels 1-5 mm long; bracteoles 0-2; calyx 7.2-10.9 mm long, the tube 4.2-6.6 mm long, short-cylindric, pilosulous, the teeth 2.7-6 mm long, lance-subulate; flowers 12-19 mm long, pink-purple with pale or white wing-tips; pods erect, sessile, oblong in outline, straight to slightly curved, 14-20 mm long, 3.5-4.8 mm wide, laterally com- pressed, strigose, unilocular. Seleniferous, sandy or sandy-silts with sandloving desert shrubs at 1330 to 1670 m in southeastern Emery (type from San Rafael Swell), eastern Garfield and Wayne cos.; endemic. The Woodruff milkvetch has affinities with A. rafaelensis, A. saurinus, and A. toanus, but has been placed in a section by itself by Bameby (1964: 436). The species was de- scribed by Jones (1923: 78) as being "the most beautiful species of the genus when the whole mass is ablaze with pink-purple bloom." The plant is a primary selenium in- dicator; 21 (vii). Astragalus zionis M. E. Jones, Zion Milk- vetch. [Xylophacos zionis (M. E. Jones) Rydb.]. Perennial, subacaulescent or short- caulescent, 3-23 cm tall, from a branching caudex, this sometimes clothed with a per- sistent thatch of leaf bases; pubescence basi- fixed; stems 0-11 cm long, prostrate to as- cending, the internodes often concealed by stipules; stipules 1.5-5.5 mm long, all dis- tinct or some shortly connate-sheathing; leaves mostly 2-15 cm long; leaflets 13-25, 2-16 mm long, 1-6 mm wide, elliptic or ovate, acute or less commonly obtuse, sil- very villous on both sides; peduncles 0.5-15 cm long; racemes 1- to 11-flowered, the flowers ascending at anthesis, the axis 0.3-6 cm long in fruit; bracts 2-5 mm long; pedi- cels 1-3 mm long; bracteoles 0-2; calyx 8.3-18 mm long, the tube 6.5-12.7 mm long, cylindric, villous, the teeth 1.5-5.7 mm long, subulate; flowers 18-26 mm long, pink-purple or sometimes pale; pods ascend- ing, sessile, obliquely ovoid-oblong in out- line, 15-30 mm long, 5.5-9 mm wide, usu- ally curved, dorsiventrally compressed, strigose or villosulous, brightly mottled, usu- ally unilocular; ovules 24-30. On sandstone and in sandy and gravelly soils in mixed desert shrub, mountain brush, and riparian communities at 1130 to 2430 m in Garfield, Kane, San Juan, and Washington cos.; Ari- zona. The Zion milkvetch is allied to A. ar- gophyllus from which it differs in the con- nate stipules and brightly mottled pods. The habitat of A. argophyllus is seldom situated in sand, while that of A. zionis is almost ex- clusively restricted to it; 31 (vi). Caesalpinia L. Unarmed shrubs, glandular-punctate in part; leaves alternate, bipinnate, petiolate, the pinnae bearing several leaflets; stipules small, persistent; flowers several to many, borne in racemes, the axis of the raceme Sept. 1978 Welsh: Utah Flora, Fabaceae 307 densely glandular; calyx 5-lobed; corolla ir- regular; petals 5, conspicuous, the up- permost not enclosing the others in bud; stamens 10, long-exserted, brightly colored and showy, distinct; pistils sessile; pods flat- tened, dehiscent. Caesalpinia gilliesii (Wallich) Dietrich. Poinciana, Bird-of-paradise. [Poinciana gil- liesii Wallich ex Hook.; Erijthrosetmon gil- liesii (Wallich) Klotzsch]. Shrubs, commonly (1) 1.5-2.5 m tall; leaves 15-28 cm long, with 8-12 pairs of pinnae, each with 7-11 pairs of elliptic to oblong leaflets 3-8 mm long, glabrous; petioles 2-4 cm long, glabrous; racemes 10-40 cm long, terminal, the peduncle and rachis conspicuously viscid-glandular; pedicels 1.5-3 cm long; flowers very showy; sepals distinct, 15-20 mm long, stipitate glandular and pu- berulent; petals yellow with orange mark- ings, 20-35 mm long; staminal filaments red, 6-9 cm long; pods ascending, sessile, oblong in outline, 55-120 mm long, 14-20 mm wide, glandular-dotted, dehiscent. Cul- tivated ornamental of startling beauty at low elevations in Washington Co.; Texas to California. Plantings in St. George and in Beaver Dam Wash persist and spread, final- ly becoming established. This is a remark- able addition to the flora of the state; 3 (ii). Caragana Lam. Shrubs; leaves alternate, even-pinnate, the rachis extended as a bristle or spine; stipules small and deciduous or persistent as spines; flowers solitary, yellow, showy; calyx cam- panulate or turbinate, obscurely to con- spicuously 5-toothed; corolla papilio- naceous; stamens 10, diadelphous; ovary sessile; pods subcylindric, linear-oblong, straight, glabrous, elastically dehiscent, the valves coiling upon dehiscence. Caragana arborescens Lam. Pea Tree. [Robinia caragana L.; C. sibirica Medicus; C. inermis Moench; C. caragana (L.) Kar- stin; Aspalathus caragana (L.) Kuntze]. Shrubs to 4 m tall or more; leaves 4-10 cm long; leaflets 8-12, 12-25 mm long, 5-15 mm wide, lance-oblong to elliptic or oval, cuspidate, villous above and below, becom- ing glabrate in age; stipules slender, occa- sionally persisting as spines; bracts reduced to rudiments at juncture of peduncle and pedicel; flowers 17-23 mm long, borne sing- ly on peduncles 12-35 mm long, few to sev- eral from each bud; pedicels 5-15 mm long; calyx turbinate, pubescent, the tube 4.5-7.5 mm long, the teeth small or obsolete, the margin villous; pods ascending to declined, sessile, linear-oblong, straight 35-55 mm long, 4-7 mm thick, glabrous, the valves drying brown. Cultivated ornamental and erosion-control plant in Kane, Salt Lake, Se- vier, Sanpete, Summit, and Utah cos., and probably throughout Utah (not adequately collected); introduced from central Siberia; 12 (ii). There are several other species of Caragana growing in North America, all in- troduced from the Old World. It is ex- pected that several of these should find use in ornamental, hedge, windbreak, and ero- sion control plantings in the future. Cercis L. Small trees or shrubs; leaves alternate, simple, palmately veined, cordate-obvate or orbicular; stipules deciduous; flowers clus- tered, appearing before leaves from spurs on old branches or cauliflorus, pink, showy; calyx turbinate-campanulate, shallowly 5- lobed; corolla irregular, the keel larger than the banner; stamens 10, distinct; ovary sub- sessile; pods short-stipitate laterally flat- tened, the ventral suture somewhat winged, indehiscent or tardily dehiscent. 1. Leaves ovate- or cordate-acuminate; flowers 9-12 mm long; plants cultivate, introduced C. canadensis Leaves cordate-reniform or broader than long, rounded or emarginate, not acuminate apically; flowers 12-14 mm long or more C. occidentalis Cercis canadensis L. American Redbud; Judas Tree. Shrub or small tree, mostly 1.5-3 m tall, rarely more; leaves commonly ovate-cordate, truncate to cordate basally, acuminate to acute apically, 3.5-10 cm long, longer than broad to somewhat broad- 308 Great Basin Naturalist Vol. 38, No. 3 er than long, glabrate to pubenilent be- neath; flowers appearing before the leaves, clustered on spurs, cauliflorus; pedicels 6-10 mm long; calyx asymmetric, 2.5-3.5 mm long, 4.3-6.3 mm wide; corolla pink (pink- purple), 9-12 mm long; keel petals 3.8-5 mm wide; pods pendulous-spreading, later- ally flattened, 40-80 mm long, 8-18 mm wide, winged, the wings 0.8-2.1 mm wide, glabrous. Cultivated ornamental at 1370 to 1570 m in Salt Lake and Utah cos., and un- doubtedly elsewhere; introduced from the eastern U.S., where indigenous from Nebras- ka south to Texas and east to the Atlantic. Our materials seem to belong to var. cana- densis; 9 (i). Cercis occidentalis Torr. ex A. Gray. Western Redbud. Small tree or less com- monly a shrub, mostly 1.5-3.5 m tall, rarely more; leaves cordate-reniform, cordate ba- sally, rounded to emarginate apically, 2-7 cm long, commonly broader than long, glabrous or puberulent along vein axils be- neath; flowers appearing before the leaves, clustered on spurs, cauliflorvis; pedicels 8-12 mm long; calyx asymmetric, 3-4.5 mm long, 5.5-8 mm wide; corolla pink to pink- purple, 12-15 mm long; keel petals 5.5-8 mm wide; pods short-stipitate, 40-100 mm long, 13-20 mm wide, winged, the wings 1.5-2.5 mm wide, glabrous. Indigenous, or rarely cultivated, at 770 to 1230 m in sand- stone canyons and alcoves in Garfield, Kane, Salt Lake (in cultivation), San Juan, and Washington cos.; California, Nevada, and Arizona. Variation in Cercis has been summarized by Isely (1975). Our material was recognized by that worker as being dis- tinctive as a population when compared to typical California specimens. Plants from Utah, Arizona, and Nevada clearly belong to var. orbiculata (Greene) Tidestr. in Tidestr. & Kittell (C. orbiculata Greene, type from Diamond Valley, Washington Co.); 32 (v). Cladrastis Raf. Unarmed trees; leaves alternate, odd- pinnately compound, petiolate; stipules lacking; axillary buds covered by the leaf base; flowers in panicles, white, showy; ca- lyx turbinate-campanulate, 5-toothed, sta- mens 10, distinct or nearly so, included; ovary stipitate; pods pendulous with the panicles, oblong in outline, laterally flat- tened, often irregularly constricted (by abor- tion of the seeds), dehiscent. Cladrastis kentukea (Dum.-Cour.) Rudd. Yellow-wood. [Sophora kentukea Dum.- Cour.; Virginia lutea Raf.; C. lutea (Michx.) K. Koch; C. tinctoria Raf.]. Tree to 10 m tall, rarely more; bark smooth; wood yel- low; leaves 15-32 cm long; leaflets 7-9 (11), 4-11 cm long, 2.3-7 cm wide, elliptic to ovate or obovate, acute to obtuse basally, acuminate to acute or obtuse apically, borne on petiolules, glabrous; panicles 20-40 cm long; flowers numerous, white, 15-25 mm long; calyx 8.5-9.5 mm long, vil- lous, the rounded lobes 1.2-1.8 mm long, densely villous; pods stipitate, the stipe 4-6 mm long, usually shorter than the calyx, the body 55-80 mm long, 8-10 mm wide, flat- tened, glabrous. Cultivated ornamental flowering tree of lower elevations in Cache, Salt Lake, and Utah cos., and likely else- where; introduced from eastern U.S. This is a beautiful tree, worthy of being cultivated more widely than at present; 3 (0). Colutea L. Unarmed shrubs; leaves alternate, odd- pinnately compound, petiolate; stipules more less persistent, papery; flowers in racemes, yellow, showy; calyx turbinate- campanulate, 5-toothed; petals papilio- naceous; stamens diadelphous, 9 and 1; ovary stipitate, the style hirsute along the ventral margin; pods pendulous, bladdery- inflated, strigulose, the incurved style per- sistent. Colutea arborescens L. Bladder-senna. Shrubs, 1.2-3.5 m tall, sometimes dying to tlie ground; herbage strigose; leaves 6-12.5 cm long; leaflets 9-13, 14-23 (30) mm long, 7.5-12.5 mm wide, obovate to oblanceolate or elliptic, rounded to emarginate, mucro- nate, strigose beneath, glabrous above, the veins apparent; racemes 4- to 11-flowered; pedicels 3.5-9 mm long; bracts 1.2-4.2 mm long; bracteoles 2; calyx 4.5-7.5 mm long, the tube 4.2-5.2 mm long, the teeth 1.2-2.2 mm long; flowers 16-22 mm long, yellow, the banner abruptly reflexed, the wings Sept. 1978 Welsh: Utah Flora, Fab ace ae 309 strongly angled upwards, surpassed by the keel; pods stipitate, the stipe 6-10 mm long, the body bladdery-inflated, ellipsoid, 45-60 mm long (or more), 16-24 mm wide (when pressed), usually mottled, indehiscent. Bladder-senna is a cultivated shrub, mainly as an ornamental in Salt Lake and Utah COS., but recently it has been used in ero- sion control plantings, especially on clay soils along roadways in Juab and Sanpete COS., at elevations of 1670 to 2500 m; in- troduced from southern Europe and Africa; 4(i). CORONILLA L. Perennial herbs, caulescent, from a tap- root and caudex; leaves alternate, pinnately compound; stipules herbaceous, becoming chartaceus; flowers papilionaceous, in ax- illary pedunculate umbels; bracts minute, scarious; calyx 5-toothed; petals 5, pink or white, the keel long-attenuate; stamens 10, diadelphous; ovary enclosed in the staminal sheath, the style glabrous; fruit a loment, 4- angled. Coronilla varia L. Crown Vetch. Per- ennial, 30-105 cm long, with spreading stems, glabrous to scabrous; stipules small; leaflets 9-23, 5-21 mm long, 1.5-11 mm wide, oblong to obovate or oblanceolate, acute to retuse apically; peduncles 3-11 cm long; bracts minute; flowers 11-20, 9-12 mm long; calyx 1.5-2 mm long, the teeth much shorter than the tube; loments 35-50 mm long. Cultivated erosion control plant, escaping and persisting, Sevier and Utah COS.; widespread in Western states; in- troduced from Europe; 2 (i). Cytisus L. Deciduous or evergreen shrubs; leaves al- ternate, 3-foliolate or simple; stipules min- ute, thickened; flowers papilionaceous, ax- illary, solitary or sometimes 2 or 3; bracts minute; calyx bilabiate, the upper lip 2- lobed, the lower 3-lobed; petals 5, yellow, the keel not especially alternate; stamen 10, monadelphous, with 4 longer than the oth- er, the style curved, broadened near the concave top; pods flattened, several-seeded. Cytisus scoparius (L.) Link. Scots Broom. Shrubs to 2 m tall or more, the branchlets green, strongly angled; leaves 3-foliolate, becoming simple upwards; flowers solitary, rarely 2 or 3, 16-22 mm long, yellow or tinged purple; pods villous along the mar- gin. Escaped cultivated plants in Weber Co., widespread in the Paciflc Coast states; introduced from Europe; 9 (0). Dalea L. Perennial herbs, unarmed; leaves alter- nate, odd-pinnate with 3 or more leaflets, glandular-dotted; stipules linear to subulate; peduncles opposite the leaves; flowers in dense spikes; calyx campanulate, 5- to 10- ribbed and 5-lobed; corolla papilionaceous, the banner petal attached near the rim of the floral cup, the other four variously in- serted or all near the rim of the staminal tube; stamens 5, monadelphous; petals white, yellowish, pink, or pink-purple to in- digo; pods 1- to 2-seeded, indehiscent, in- cluded in the calyx or slightly exceeding it. Note: Included herein are the herbaceous species traditionally placed in Petalostemon, and excluded are those shrubby species now treated as belonging to Psorothamnus (q.v.). References Wemple, D. K. 1970. Revision of the genus Petalostemon (Leguminosae). Iowa State Jour. Sci. 45:1-102. 2(1). Stems prostrate; spikes slender, commonly less than 8 mm thick; leaflets 5-17; plants of Kane, San Juan, and Washington cos D. lanata Stems decumbent to ascending or erect; spikes thicker, commonly 10 mm or more; leaflets 7 (9) or fewer; distribution various 2 Herbage glabrous 3 Herbage pilose, pilosulous, or hirsute with lustrous hairs 4 310 Great Basin Naturalist Vol. 38, No. 3 3(2). Petals pink to pink-purple; calyx oblique, the teeth long-pilose; plants west of the Colorado River D. searlsiae - Petals white (fading cream); calyx symmetric, the teeth ciliate; plants mostly east of the Colorado River (rarely to the west) D. oligophylla 4(2). Spikes dense, the longest seldom more than 5 cm long; calyx teeth 1.5-2.5 mm long; plants wide-spread in Kane, San Juan, Grand, Emery, Wayne, and Garfield cos D. flavescens Spikes rather lax, the longest 5-14 cm long; calyx teeth 2.7-4 mm long; plants of San Juan Co D. epica Dalea epica Welsh. Hole-in-the-Rock Prairie-clover. Stems ascending to erect, 20-42 cm tall, from a superficial to sub- terranean woody caudex, strigulose to pi- lose; stipules 2-4 mm long, lance-subulate to linear, persistent; leaves 1.5-3.8 cm long; leaflets 3-5, 5-19 mm long, 2-9 mm wide, flat or folded, oblanceolate to elliptic, acute, lustrous-strigulose or -pilose, glandu- lar beneath; terminal leaflet petiolulate on a continuation of the rachis, the rachis pro- longation very short; peduncles 1.5-9 (12) cm long, sparingly villous-hirsute; spikes (2) 2.5-9 (14) cm long, 12-18 mm wide (when pressed), the rachis spreading-hairy; bracts 4-7 mm long, lance-aristate, villous; calyx 5-7 mm long, obscurely 10-ribbed, the tube not translucent, the teeth 2.7-4 mm long; flowers 7.5-11 mm long, the petals white; pistils 11.5-13 mm long, the style 8.5-9.5 mm long; pods villous. Sandstone bedrock and dimes at 1530 m in San Juan Co. (west of Clay Hills Divide); endemic. The Hole- in-the-Rock prairie-clover is a near ally of D. flavescens, within whose range it occurs, but with whom it is apparently allopatric; 5 (V). Dalea flavescens (S. Wats.) Welsh. Kanab Prairie-clover. [Petalostemon flavescens S. Wats.; Kunistera flavescens (S. Wats.) Kuntze.]. Stems 23-52 cm tall, from a su- perficial caudex, glabrous to hairy (espe- cially above); stipules 1-4 mm long, sub- ulate, persistent; leaves 1.5-4.7 cm long; leaflets 3-7, 6-20 mm long, 1-6 mm wide, folded or flat, oblong to oblanceolate or linear, lustrous-strigulose on both surfaces, glandu- lar beneath; terminal leaflet stalked to sub- sessile; peduncles 3-20 cm long, glabrous or sparsely to densely pilose-hirsute; spikes 1.5-6.5 cm long, 10-16 mm wide (when pressed), and sometimes glandular; calyx 10- ribbed, the tube not translucent, the teeth 1.5-2.5 mm long; flowers 6.2-9.5 mm long, the petals white, fading cream; pistils 7.5-10.3 mm long, the style 6-8 mm long; pods villous. Grasslands, mixed desert shrub, blackbrush, and pinyon-juniper commu- nities, commonly in sandy soils, at 970 to 1870 m in Emery, Garfield, Grand, Kane (type from Kanab), San Juan, and Wayne COS.; Arizona, a Navajo Basin endemic. The Kanab prairie-clover is remarkably variable in several features, e.g., in number and posi- tion of glands, in vesture of the peduncle, and in width of the spike. In one specimen (Holmgren & Goddard 9990, frag. BRY) from the bottom of Glen Canyon near the mouth of Bridge (Forbidding) Canyon (now beneath Lake Powell), the calyx teeth are glandular. This seems to represent an exten- sion of the glandular condition from bracts, where they exist or don't exist, to the calyx teeth, where they usually do not exist. Spike width varies with maturity of flowers and fruit, but there is a tendency, especially along the canyons of the Colorado River, towards very broad flowering spikes (12-16 mm), and tliese might warrant taxonomic recognition when more material becomes available. Plants from East Clark Bench and eastward to Last Chance Canyon in Kane Co., mainly occupy stabilized dunes domi- nated by Vanclevea-Ephedm-Oryzopsis mix- tures, and have moderately to densely hairy peduncles which vary from 3-10 cm in length at anthesis. Most of the remainder of the plants in Utah have glabrous to spar- ingly hairy peduncles, which are 9-20 cm long at anthesis. When more specimens are Sept. 1978 Welsh: Utah Flora, Fabaceae 311 at hand, it might become pnident to treat these variants at some taxonomic level; 33 (xiv). Dalea lanata Sprang. Woolly Dalea. [ParoselUi lanata (Spreng.) Britton; D. tertni- nalis (M. E. Jones) Heller]. Stems prostrate, 15-60 cm long or more, from a sub- terranean to superficial caudex, pilosulous to glabrous; stipules 1-2 mm long, subulate, more or less persistent; leaves 0.9-3 cm long; leaflets 5-17, 1.5-10 mm long, 1-5.5 mm wide, obovate to cuneate, truncate to emarginate, commonly folded, pilosulous or glabrous; peduncles 0.5-2.5 cm long; spikes 1.8-7.5 cm long, lax in mid- to late-anthesis and in fruit; bracts ovate-acuminate, pilo- sulous and with one or more large glands; calyx 3.2-4.6 mm long, the tube 2.1-2.7 mm long, glabrous, the teeth 0.9-1.9 mm long, pilosulous dorsally and ciliate; flowers 6-7 mm long, the petals indigo to rose- pink; pistils 5-6 mm long, the style 3.5-4.5 mm long; pods villous or glabrous. Stabi- lized dimes and other sandy sites at 970 to 1370 m in Kane, San Juan, and Washington COS.; Arizona, Colorado, and Kansas south to Mexico. Our material belongs to var. termi- nalis (M. E. Jones) Barneby. This variety differs from D. lanata var. lanata inter alia in the glabrous, shining, membranous calyx tube. Some plants from San Juan Co. (Har- rison 12194, 12194a, 12199 BRY) are glabrous throughout, but seem not to differ otherwise; 8 (0). Dalea oligophylla (Torr.) Shinners. West- em Prairie-clover. [Petalostemon gracilis var. oligophyllus Torr. in Emory; P. gracilis A. Gray, not D. gracilis Kunth; Kuhnistera occidentalis A. Gray ex Heller; P. occiden- tale (A. Gray) Fern.; K. Candida var. occi- dentalis (A. Gray) Rydb.; P. truncatus Rydb.; K. oligophylla (Torr.) Heller; P. oligophyllus (Torr.) Rydb.; P. truncatus Rydb.; P. sonorae Rydb.; P. candidus var. oligophyllus (Torr.) F. J. Hermann.; D. Can- dida var. oligophylla (Torr.) Shinners]. Stems decumbent to erect, 40-90 cm tall, from a superficial caudex, glabrous; stipules 1-4.5 mm long, fragile, often coiled; leaves 1.5-5.2 cm long; leaflets 4-9, 5-27 mm long, 1-7 mm wide, oblanceolate to elliptic or oblong, tnmcate to emarginate, com- monly folded, glabrous, glandular beneath; peduncles 1.5-15 cm long, glabrous; spikes 1-6.8 cm long, 8-12 mm thick the rachis commonly glabrous; bracts lance-acuminate, caducous; calyx strongly 10-ribbed, usually pubescent between the ribs, the tube 2.3-3 mm long, the lobes 1-1.3 mm long; pistils 9-11 mm long, the style 8-10 mm long; pods glabrous or sparingly hairy apically. Sandy drainages and crevices in rim rock in mixed desert shrub, blackbush, pinyon- juniper, and hanging garden communities at 1070 to 1830 feet in Emery, Garfield, Grand, Kane, San Juan, and Wayne cos.; Al- berta east to Saskatchewan and south to Arizona, New Mexico, Texas, Mexico, and Iowa. Utah specimens of the western prairie-clover are remarkably uniform; 35 (vi). Dalea searlsiae (A. Gray) Barneby. Searls Prairie-clover. [Petalostemon searlsiae A. Gray; Kuhnistera searlsiae (A. Gray) Kuntze]. Stems decumbent to erect, 23-65 cm tall, from a superficial caudex, glabrous; stipules 1-3 mm long, deciduous; leaves 1.4-5.2 cm long; leaflets 5-11, 3-18 mm long, 0.8-6 mm wide, oblanceolate to ellip- tic or almost linear, truncate to emarginate or acute, commonly involute or folded, glabrous, glandular beneath; peduncles 3.5-22 cm long, glabrous; spikes 1.6-9 (13) cm long, 8-12 mm wide, the rachis glabrous to hairy; bracts obovate to lanceolate, acuminate-aristate, deciduous; calyx 10- ribbed, the tube glabrous to moderately pi- lose, 2.4-2.9 mm long, the lobes 1-1.4 mm long, usually long-pilose; pistils 5-7 mm long, the style 4-5 mm long; pods villous. Sagebrush, pinyon-juniper, warm-desert shrub, or less commonly in spruce-fir com- munities at 1230 to 2800 m in Box Elder, Garfield, Iron, Juab, Kane, Tooele, and Washington cos.; Arizona, Nevada, and Cal- ifornia. The Searls prairie-clover is closely allied to the ornate prairie-clover {Petaloste- mon ornatwn Douglas in Hook.), whose range is contiguous to the northwest. In a broad sense, possibly D. searlsiae would best be included as a variety of that taxon, but its features differ in about the same order of magnitude as those used to segregate other taxa at specific levels. No attempt at com- 312 Great Basin Naturalist Vol. 38, No. 3 bination is intended or implied herein; 26 (i)- Galega L. Perennial, caulescent, from a caudex and stout taproot; leaves altenate, odd-pinnate; stipules sagittate, distinct; flowers papilio- naceous, borne in axillary racemes, each subtended by a bract; bracteoles 0; calyx 5- toothed; petals 5, purplish-blue, the keel subequal to the wings, attenuate apically; stamens monadelphous; ovarv enclosed in the staminal sheath, the style glabrous; pods sessile, narrowly oblong in outline, more or less constricted between the seeds, unilocu- lar. Galega officinalis L. Goatsrue. Perennial, caulescent, 50-100 cm tall, from a branch- ing caudex; pubescence basifixed; stems erect or ascending; stipules 7-16 mm long, sagittate, the basal lobes again once or twice lobed; leaves 3-22 cm long; leaflets 9-15, 7-52 mm long, 3-17 mm wide, lan- ceolate to elliptic, cuspidate-aristate api- cally, glabrous above, thinly pilose along veins beneath; peduncles 3-9 cm long; ra- cemes 20- to 36-flowered, the flowers de- clined at anthesis, the axis 2.5-7 cm long in fruit; bracts 3.5-6 mm long, some com- monlv semi-sagittate; calvx 4-5.5 mm long, the tube 2-2.5 mm long, campanulate, glabrous, the teeth 2-3.2 mm long, subulate; flowers 9.5-12.5 mm long, blue-purple (fad- ing cream); pods ascending, subcylindric, 18-38 mm long, 2-3 mm thick, longitudi- nallv nerved, glabrous, unilocular. In- troduced, established weedy plant, known in Utah only from Cache Co., where it has grown continuously since at least 1909; ad- ventive from Europe; 4 (0). Gleditsia L. Trees, often armed with simple or branched thorns; leaves alternate, deciduous, pinnately once to twice compound (with both kinds of leaves often on the same branch, or some intermediate); stipules mi- nute, caducous; leaflets 14-36 on once- pinnate leaves and on the 3-5 pinnae; flow- ers in spike-like axillary racemes, polyg- amous, almost regular; sepals equal or nearly so; petals 3-5, very narrow, yellow- ish, the uppermost internal in bud; stamens 3-10, distinct, the anthers in pistillate flow- ers abortive; pods flattened, strap-like, in- dehiscent. Gleditsia triacanthos L. Honey Locust. Trees to 20 m tall or more; bark smooth; leaflets 12-35 cm long, the pinnae 3-5 (8) when bipinnately compound, the leaflets 14-36 on once pinnately compound leaves and on pinnae, 10-42 mm long, 3-14 mm wide, lanceolate to oblong, crenate, obtuse to cuspidate, glabrous above, puberulent along veins beneath; racemes many-flowered, 3-7 cm long, short pedunculate or sub- sessile; sepals separate; petals 4-5 mm long, greenish; pods sessile, oblong in outline, lat- erally flattened, 70-350 mm long, 15-30 mm wide, curved, indehiscent, the seeds im- bedded in tissue. Cultivated shade tree, rarely escaping, in Grand, Millard, Salt Lake, Utah, and Wayne cos., and likely throughout the state at lower elevations; in- troduced from the eastern United States. Many horticultural forms, mainly thornless, are cultivated. The trees are handsome or- namentals, even with thorns; 15 (iii). Glycyrrhiza L. Perennial, caulescent, from stout sweet roots; leaves alternate, odd-pinnate, glandular- punctate; stipules subulate, distinct; flowers papilionaceous, in axillary racemes, each subtended by a lanceolate, deciduous bract; bracteoles 0; calyx 5-toothed; petals 5, white to cream, the keel shorter than the wings; stamens 10, diadelphous; ovary en- closed in the staminal sheath, the style glabrous; pods sessile, elliptic to oblong in outline, bur-like, armed with micinate ap- pendages, indehiscent, few-seeded. Glycyrrhiza lepidota Pursh. Licorice. Plants 40-120 cm tall, from a deep-seated root; stipules 2-7 mm long, subulate; leaves 8-19 cm long; leaflets 13-19, 8-53 mm long, 3-15 mm wide, lanceolate to oblong, mucronate, glabrous above, glandular-dotted and puberulent beneath; peduncles often paired, 3-8 cm long; racemes 20- to 50- flowered, the flowers ascending at anthesis, the rachis, 2.5-9 cm long in fruit; bracts 5-8 mm long, caducous; calyx 4.8-6.9 mm Sept. 1978 Welsh: Utah Flora, Fabaceae 313 long, the tiibe 2.5-4.9 mm long, canip- anulate to short-cylindric, stipitate- glandular, the teeth 1.5-3.6 mm long; flow- ers 9.1-13 mm long, white to cream; pods spreading, laterally compressed, oblong, 13-20 mm long, the body 5-7 mm wide, beset with hooked prickles, simulating cock- leburs. Terraces, streamsides, seeps, and oth- er semi-moist sites in streamside, grease- wood, mixed desert shrub, and pinyon- juniper communities at 670 to 2470 m in Beaver, Cache, Daggett, Garfield, Grand, Kane, Millard, Piute, Rich, Salt Lake, San Juan, Sevier, Summit, Uintah, Washington, Wayne , and Weber cos.; widespread in the United States, except for the southeast; 53 (xii). Gymnocladus L. Unarmed trees; leaves alternate, de- ciduous, bipinnately compound; stipules small, deciduous; leaflets 9-15 on each of the 3-7 pinnae, or the lowermost pinnae represented by leaflets; flowers in terminal panicles, dioecious or polygamous, regular or nearly so; calyx 5-lobed; petals 5, dis- tinct; pods broad-oblong, hard, thick, flat- tened, pulpy. Gymnocladus dioica (L.) K. Koch. Ken- tucky Coffee-tree. [Guilandina dioica L.; Gymnocladus canadensis Lam.]. Tree to 20 m or more; leaves 20-60 (90) cm long; pin- nae 3-7 pair, each with 9-15 leaflets, these 17-70 mm long, 7-30 mm wide, ovate to lanceolate, entire, acuminate, pilose (espe- cially along the veins) beneath, glabrous to glabrate above; panicles many-flowered; pis- tillate panicles to 25 cm long, the staminate much smaller and more dense; flowers 10-13 mm long, greenish white; fruit 100 to 200 mm long; 30-50 mm wide, persistent, tardily dehiscent, the large seeds imbedded in tissue. Cultivated ornamental and bo- tanical curiosity in Salt Lake and Utah cos., and probably elsewhere; introduced from the eastern United States. The Kentucky coffee-tree is so named because the pulp surrounding the seeds is rich in a poly- saccharide which, when roasted, has been used as a coffee substitute; 4 (i). Hedysarum L. Perennial herbs, caulescent, from a cau- dex and taproot; leaves alternate, odd- pinnate, stipules adnate to the petiole base, at least the lowermost connate-sheathing; flowers papilionaceous, in axillary racemes, each subtended by a bract; bracteoles 2; ca- lyx 5-toothed; petals 5, red-purple to pink or pink-purple, the keel much longer than the wings, abruptly bow-shaped; stamens 10, diadelphous; ovary enclosed in the sta- minal sheath, the style glabrous; loments with 2-8 segments, prominently reticulate. References NoRTHSTROM, T. E. 1974. The genus He- dysarum in North America. Ph.D. Dis- sertation. Brigham Young University. 187 pp. NoRTHSTROM, T. E. AND S. L. Welsh. 1970. Revision of the Hedysarum boreale complex. Great Basin Nat. 30:109-130. Rollins, R. C. 1940. Studies in the genus Hedysarum in North America. Rho- dora 42:217-238. Leaflets thin, the veins readily apparent; fruit segments winged; calyx lobes imequal, shorter than the tube; plants rare in eastern Utah H. occidentale Leaflets thick the veins not apparent; fruit segments not winged; calyx lobes subequal, longer than the tube; plants common, widespread H. boreale Hedysarum boreale Nutt. Northern Sweetvetch. Perennial, caulescent, 17-70 cm tall, from branching subterranean to su- perficial caudex; pubescence basifixed; stems decumbent to erect; stipules 2-10 mm long, at least some connate-sheathing; leaves 3-12 cm long; leaflets 5-15, 7-35 mm long, 2-19 mm wide, oblong to elliptic, lance-oblong, or ovate (rarely linear), strigose on both sides or glabrate to glabrous above; pe- dimcles 2.8-15 cm long; racemes 5- to 45- flowered, the flowers ascending at anthesis, 314 Great Basin Naturalist Vol. 38, No. 3 the axis 5-28.5 cm long in fruit; bracts 2-5 less commonly white; loments stipitate, mm long; pedicels 0.8-4.5 mm long; brae- pendulous to spreading, with 2-8 segments, teoles 2; calyx 4.5-8 mm long, the tube not winged, prominently reticulate. Our 2.5-3.5 mm long, campanulate, strigose, the material belongs to ssp. boreale and is sepa- teeth 2-6 mm long, subulate; flowers 10-19 rable into two varieties. mm long, red-purple to pink or pink-purple, 1. Segments of the loment bearing spines on the lateral surfaces; plants of Uintah Co H. boreale var. gremiale — Segments of the loment lacking spines; plants widespread H. boreale var. boreale Var. boreale. [H. carnulosum Greene; H. pabtiUire A. Nels; H. utahense Rydb.; H. pabulare var. rivulare L. O. Williams; H. mackenziei var. pabulare (A. Nels.) Kearney & Peebles; H. utahense Rydb., type from Salt Lake City; H. boreale var. utahense (Rydb.) Rollins; H. canescens Nutt. ex Torr. & Gray; H. cinerascens Rydb.; H. mack- enziei var. canescens (Nutt.) Fedtsch.; H. boreale var. cinerascens (Rydb.) Rollins; H. boreale var. obovatum Rollins]. Mixed desert shrub, pinyon-juniper, mountain brush, pon- derosa pine, and aspen communities at 1175-2500 m in Box Elder, Cache, Carbon, Davis, Duchesne, Emery, Garfield, Grand, Juab, Millard, Salt Lake, San Juan, Sanpete, Sevier, Tooele, Uintah, Utah, Wasatch, Washington, Wayne, and Weber cos.; Al- berta east to Manitoba and south to Ne- vada, Arizona, New Mexico, and Texas. The use of leaf pubescence, or lack thereof, to segregate the var. boreale into fiuther taxa is a function in frustration, leading to two essentially sympatric phases which might re- flect ecology more than genetics; 142 (xxiv). Var. gremiale (Rollins) Northstrom & Welsh. [H. gremiale Rollins]. Pinyon- juniper and mountain brush communities at 1470 to 1670 m in Uintah Co.; endemic. The spines on the loment segments vary from few (or none) to numerous, indicating complete transition with var. boreale. In- cipient spines are found in specimens of var. boreale from outside of Utah, but no- where are the spines so well or so con- sistently developed as in Uintah Co.; 3 (0). Hedysarum occidentale Greene. Western Sweetvetch. [H. lancifolium Rydb.; H. mar- ginatum Greene; H. uintahense A. Nels.]. Perennial, caulescent, 30-90 cm tall, from a branching, superficial caudex; pubescence basifixed; stems ascending to erect; stipules 10-17 mm long, at least some connate- sheathing; leaves 8-20 cm long; leaflets 11-19, 9-37 mm long, 4-16 mm wide, ovate to lance-ovate or elliptic, apiculate to emarginate, strigose on both sides or glabrous above; peduncles 3.7-15 cm long; racemes 10- to 50-flowered, the flowers spreading to declined at anthesis, the axis 6-14 cm long in fruit; bracts 2-8 mm long; bracteoles 2; calyx 3.5-11 mm long, the tube 2.3-6 mm long, campanulate, glabrous to strigose, the teeth 0.5-2 mm long, triangular; flowers 16-23 mm long, pink to red-purple; loments stipitate, pendulous, with 1-5 segments, winged. Mountain brush, sagebrush, and lower spruce-fir-aspen commimities at 1770 to 2430 m in Carbon, Duchesne, Emery, and Summit cos.; British Columbia south of Washington, Montana, Idaho, Wyoming, and Colorado. Utah plants have been collected in flower and fruit in mid-to late summer; 8 (vi). Material from Carbon and Emery counties differs from the main body of the species in leaflet features and is separable as var. canone Welsh var. nov. Hedijsaro occidentali var. occidentali aemulans sed foliolis ovatis ellipticis vel late lanceolatis et apice retusis truncatis vel apiculatis. Holotype: Utah; Carbon Co. 14 mi due ENE of Helper, Soldier Creek, 7408 feet, Welsh & Taylor 15256, 30 June 1977 (BRY). Paratypes: Carbon Co., Soldier Can- yon, Welsh & Christensen 6614, 11 August 1967; 2.5 mi due N. of Sunnyside, Welsh & Taylor 15075, 21 June 1977; east of Sunny- side, Welsh & Murdock 9146, 19 July 1969, do, Welsh & Christensen 6598, 11 Aug 1967. Emery Co., 9 mi due NNE of Wood- Sept. 1978 Welsh: Utah Flora, Fab ace ae 315 side, Welsh 14923, 9 June 1977 (all BRY). HoFFMANSEGGiA CavaniUes Perennial herbs, rhizomatous (?) and with a subterranean caudex; leaves alternate, bi- pinnately compound, odd-pinnate as to pin- nae and even-pinnate as to leaflets, the leaf- lets not glandular punctate; stipules distinct, persistent; flowers in terminal racemes, per- fect, irregular; calyx 5-lobed; petals 5, yel- low, the uppermost internal in bud; stamens 10, the filaments distinct; pods oblong or falcate in outline, laterally compressed, in- dehiscent. Hoffmanse^ia repens (Eastwood) Cock- erell. Creeping Rush-pea. [Caesalpinia re- pens Eastwood; Moparia repens (Eastwood) Britton]. Subacaulescent or shortly caules- cent, 5-12.5 cm tall (above ground), from a deeply subterranean caudex, the branches below ground 3-15 cm long, pale; pu- bescence basifixed; leaves 2.5-9.5 cm long; pinnae 3-7 (9); leaflets 4-14, 3-12 mm long, 1-6 mm wide, asymmetrically ob- ovate-elliptic to oblong, crowded, entire, villosulous; peduncles 1.2-6 cm long; ra- cemes 7- to 26-flowered, the flowers spread- ing at anthesis, the axis 3-8 cm long in fruit; bracts 3-7 mm long, caducous; bract- eoles 0; pedicels 2-7 mm long; calyx 8-10.5 mm long, the tube 2-4.5 mm long, campanulate, retrorsely villosulous, the teeth 6-8.5 mm long, oblong-lanceolate, vil- losulous; flowers opening flat or nearly so, the petals yellow, 10-12 mm long, red- spotted near the base, the whole fading pink-orange; pods pendulous, oblong, 20-50 mm long, 10-20 mm wide, membranous, pilosulous. Sandy deserts with Ephedra, Ory- zopsis, and other arenophilus plants at 1430 to 1670 m in Emery, Garfield, Grand (type from Court House Wash), and Wayne cos.; endemic. This is a striking plant in the light of early morning along the sandy stretches near Hanksville; 20 (iv). Laburnum Medic. Trees, unarmed; leaves alternate, pal- mately trifoliolate; stipules lacking; flowers in terminal, pendulous racemes, perfect; ca- lyx 2-lipped, the upper lip 2-toothed, the lower lip with 3 coalescent teeth; petals all distinct; stamens 10, monadelphous; ovary stipitate; pods pendulous with the raceme, narrowly oblong, laterally compressed, more or less constricted between the few to sev- eral seeds. Laburnum anagyroides Medic. Golden- chain, Bean-tree. [L. vulgare Bercht. & Presl.; Cytissus laburnum L.]. Slender trees to 6 m tall; leaves (including petioles) 1.7 cm long 2.5-15 cm long; leaflets 3, pal- mate, 1.4-7.5 cm long, 0.7-3.5 cm wide, lanceolate to elliptic or oblanceolate, acute to obtuse or rounded, strigulose to glabrate beneath, glabrous above, often ciliate; pe- duncles 1.2-3.8 cm long; racemes (7) 15- to 50-flowered, the flowers inverted and spreading at anthesis, the axis 15-30 cm long in fruit; bracts lacking; bracteoles 0; pedicels 8-14 mm long; calyx oblique, 4.5-5.5 mm long, the tube 3-4 mm long, glabrous, the lobes about 1.5 mm long, tuft- ed hairy; flowers 14-17 mm long, yellow; pods stipitate, the stipe 2-5 mm long, the body oblong in outline, 30-50 mm long, 5-8 mm wide, strigose, tardily dehiscent. Cultivated ornamental trees of great beauty, in population centers, in much of Utah; in- troduced from southern Europe; 4 (0). Lathyrus L. Annual or perennial herbs, clambering, trailing, or climbing; leaves alternate, even- pinnately compound, the rachis terminating in a bristle or prehensile tendril; stipules herbaceous, semi-hastate or semi-sagittate; leaflets 2-12, very variable; flowers in ax- illary racemes, papilionaceous; calyx 5- toothed, obliquely campanulate; petals 5, white or cream to pink, purplish, or other- wise (in cultivated types), the wings not ad- nate to the keel, but fltted together in a groove; stamens 10, diadelphous; style later- ally compressed, bearded along the ventral (upper) edge; pods oblong, several-seeded, the valves coiling upon dehiscence. References Hitchcock, C. L. 1952. A revision of the North American species of Lathyrus. Univ. Wash. Publ. Biol. 15:1-104. Welsh, S. L. 1965. Legumes of Utah III: Lathyrus L. Proc. Utah Acad. Sci. 42:214-221. 316 Great Basin Naturalist Vol. 38, No. 3 1. Leaflets 2; stems winged; plants introduced, annual or perennial 2 — Leaflets 4 or more; stems angled but not winged; plants indigenous, perennial 4 2(1). Plants annual, pubescent; flowers 25-30 mm long L. odoratus — Plants perennial, glabrous; flowers 12-25 mm long 3 3(1). Leaflets narrowly lanceolate to elliptic; flowers 15-18 mm long L. sylvestris — Leaflets lance-elliptic to oblong or ovate; flowers 20-25 mm long L. latifolius 4(1). Keel conspicuously shorter than the wings; calyx glabrous or the teeth merely ciliate, the lower tooth usually longer than the tube; stipules large, foliaceous; petals pink-purple, rarely white L. pauciflorus — Keel commonly subequal to the wings; calyx often hairy, the lower tooth shorter than the tube; stipules not foliaceous; flowers pink-purple, pale lavender, pinkish-violet, cream, or white 5 5(4). Flowers 8-16 mm long; petals pale lavender-tinged to pinkish-violet, cream, or white, often polychrome in populations; plants common at middle eleva- tions, especially in aspen, flowering in summer L. lanzwertii — Flowers 15-30 mm long; petals bright pink- to blue-purple; plants wide- spread at lower elevations, flowering mainly in springtime L. bmchycalyx Lathyrus brachycalyx Rydb. Rcdberg long; racemes 2- to 5-flowered, the flowers Sweetpea. Perennial clambering herbs, de- spreading at anthesis; calyx tube 3.5-7 mm cumbent to erect, 10-50 cm long, the her- long, campanulate, the teeth 1.5-6 mm bage villous to glabrous; stipules 6-15 mm long, triangular to lanceolate; flowers 15-30 long, semi-sagittate; leaves 2-9 cm long (ex- mm long, pink to pink-purple; pods 30-70 eluding tendrils); leaflets 6-12, 5-50 (70) mm long, 5-10 mm wide. There are three mm long, 2-15 mm wide, linear to elliptic, more or less distinctive and allopatric varie- oblong lanceolate, or oblanceolate; tendrils ties in Utah, simple or branched; peduncles 4-10 cm 1. Plants villous; leaflets commonly 10-25 cm long; flowers 18-25 mm long; banner not deeply cordate apically, the blade as long as broad; calyx tube 3.5-5.5 mm long, the teeth 2.2-3.8 mm long; Great Basin L. brachycalyx var. brachycalyx — Plants glabrous or sparingly villous; leaflets commonly 25-70 mm long; flowers, banner and calyx various; mainly not of the Great Basin 2 2(1). Flowers 15-25 mm long; banner often deeply cordate, the blade commonly broader than long; calyx tube 4-5 mm long, the teeth 1.5-2.3 mm long ' L. brachycalyx var. zioniy — Flowers 20-30 mm long; banner but shallowly retuse, the blade much longer than broad; calyx tube 5-7 mm long, the teeth 2-5 mm long L. brachycalyx var. eucosmus Var. brachycalyx. [L. brachycalyx Rydb., Beaver, Box Elder, Juab, Millard, Salt Lake, type from City Creek Canyon]. Mixed Sanpete, Tooele, and Utah cos.; Nevada (?), desert shrub, pinyon-juniper, and mountain a Great Basin endemic; 51 (vi). brush communities at 1575 to 2600 m in Var. eucosmus (Butters & St. John) Sept. 1978 Welsh: Utah Flora, Fabaceae 317 Welsh Stat. nov. based on Lathynis eu- to villous; stipules 7-20 mm long, semi- cosmus Butters & St. John Rhodora 19:160. sagittate; leaves 2-14 cm long (excluding 1917. [L. brachycalyx ssp. eucosmus (Butters tendrils) leaflets 4-12, 7-75 mm long, 3-18 & St. John) Welsh.] Clay soil in washes in (26) mm wide, elliptic to lanceolate, oblan- salt desert shrub communities at 1450 to ceolate, or oval; tendrils short and simple to 1700 m in Emery, Grand, and Sanpete cos.; more commonly branched and prehensile; Colorado, New Mexico, and Arizona; 5 (0). peduncles 2-8.5 cm long; racemes 2- to 5- Var. zionis (C. L. Hitchc.) Welsh stat. flowered, the flowers spreading at anthesis; nov. based on Lathynis zionis C. L. Hitchc, calyx tube 3.5-6 mm long, campanulate, the Univ. Wash. Publ. Biol. 15:36. 1952. [L. lower lateral teeth 1.8-4.2 mm long, brachycalyx ssp. zionis (C. L. Hitchc.) triangular to lanceolate; flowers 12-22 mm Welsh.] Sandy soils in pinyon-juniper, mixed long, pink-purple to white or cream and desert shrub, and riparian communities at commonly suffused or veined with pink or 1200 to 2500 m in Garfield, Grand, Kane, purple; pods 30-60 mm long, 3-7 mm San Juan, and Washington cos.; Arizona; 42 wide. Three rather poorly defined and ap- (vi). parently intergrading varieties are present Lathyrus lanzwertii Kellogg. Lanzwert in Utah. The following arbitrary key will al- Sweetpea. Plants clambering, decumbent to low segregation of most of the specimens, erect, 20-60 cm tall, the herbage glabrous 1. Tendrils reduced to a simple filiform stalk, rarely coiled; leaflets commonly 6 only; plants rare in southern Utah L. lanzwertii var. arizonicus — Tendrils commonly branched and/ or coiled; leaflets often more than 6; plants widespread 2 2(1). Flowers white, less commonly suffused or veined with pink or purple, mostly 15-22 mm long; plants more abundant southward in Utah L. lanzwertii var. laetivirens — Flowers pink-purple or suffused with purple, commonly 12-17 mm long; plants more abundant northward in Utah L. lanzwertii var. lanzwertii Var. arizonicus (Britten) Welsh, comb. and Wasatch cos.; Washington, Oregon, nov. based on L. arizonicus Britton Trans. California, Idaho, and Nevada; 54 (xxii). N. Y. Aca. Sci. 8:65. 1899. Aspen and Lathyrus latifolius L. Perennial Sweet- mountain brush commimities at 2470 to pea. Perennial, climbing vines, 80-200 cm 2770 m in San Juan Co.; Arizona; 4 (0). tall, the stems broadly winged, glabrous; Var. laetivirens (Green) Welsh stat. nov. stipules 9-40 mm long, semi-hastate to based on Lathyrus laetivirens Greene ex semi-sagittate; leaves 6-12 cm long (exclud- Rydb. Fl. Colorado 2.7. 1906. [L. leu- ing tendrils; leaflets 2, 35-80 (150) mm canthus Rydb.; L. leucanthus var. laetivi- long, 5-23 (50) mm wide, lance-elliptic to rens (Greene) C. L. Hitchc] Riparian, as- oblong or ovate; tendrils branched, coiled; pen, mountain baish, coniferous woods, and peduncles 7-15 cm long; racemes 5- to 15- other montane communities, at 1830 to flowered, the flowers spreading at anthesis; 3130 m in Beaver, Carbon, Emery, Garfield, calyx tube 5.8-6.2 mm long, campanulate, Grand, Millard, Salt Lake, San Juan, San- the lower lateral calyx teeth 3-6 mm long, pete, Sevier, and Washington cos.; Colo- lanceolate; flowers 20-25 mm long, pink- rado. 32 (x). purple, pink or white; pods 60-80 mm long, Var. lanzwertii. [L. coriaceus White, type 7-10 mm wide, glabrous. Cultivated, escap- from Wasatch Mts.]. Aspen, Douglas fir, ing and now established, mainly along canal spruce fir, and less commonly in mountain banks in Carbon, Grand, and Utah cos., and brush commimities at 1650 to 2400 m in probably widespread; introduced from Eu- Davis, Duchesne, Garfield, Rich, Tooele, rope; 4 (ii). Salt Lake, Sanpete, Sevier, Summit, Utah, Lathyrus odoratus L. Sweetpea. Annual, 318 Great Basin Naturalist Vol. 38, No. 3 climbing vines, 80-300 cm tall, the stems broadly winged, pubescent; stipules 10-30 mm long, semi-hastate; leaves 6-15 cm long (excluding tendrils); leaflets 2, 25-85 mm long, 8-40 wide, elliptic to ovate or oblan- ceolate; tendrils well-developed, prehensile; pedimcles 3-28 cm long; racemes 2- to 5- flowered, the flowers spreading at anthesis; calyx tubes 5.5-7.5 mm long, campanulate, spreading-hairy, the lower lateral teeth 5-8 mm long, lanceolate; flowers 25-37 mm long, varicolored; pods 30-60 mm long, 5-8 mm wide, pubescent. Cultivated ornamental or greenhouse a'nd outside plantings, grow- ing best in cool middle elevations of the state; introduced from Europe; 2 (ii). Lathy rus pauciflorus Fern. Utah Sweet- pea. [L. utahensis M. E. Jones; L. pauci- florus var. utahensis (M. E. Jones) Peck; L. bradfieldianus A. Nels.j. Perennial, 20-100 cm tall or more, climbing vines, glabrous; stems merely angled; stipules 8-32 mm long, the larger ones, at least, foliose and toothed; leaves 2-12.5 cm long (excluding tendrils); leaflets 8-12, 14-50 mm long, 8-32 mm wide, ovate to elliptic; tendrils well-developed, prehensile; peduncles 3.5-24 cm long; racemes 3- to 10-flowered, the flowers spreading at anthesis; calyx tube 5-7.3 mm long, obliquely campanulate, more or less gibbous, the lower lateral teeth 2.5-7 mm long, often curved and spreading; flowers (13) 15-23 (27) mm long, pink to pink-purple, with keel usually pale or white; pods 40-75 mm long, 7-11 mm wide. Oak-sagebrush, mountain brush, as- pen, lodgepole pine, mixed conifer, and meadow communities at 1370 to 2900 m in Box Elder, Cache, Davis, Millard, Salt Lake, Sanpete, Sevier, Summit, Tooele, Utah, Wasatch, and Weber cos.; Washington, Ore- gon, Idaho, Colorado, and Arizona. The Utah sweetpea forms apparent hybrids with L. lanzwertii. Leaf and flower features are intermediate in such plants (e.g., Higgins 1048; Clark 2008; Welsh & Lsely 6378, all BRY); 71 (x). Lathyrus sylvestris L. Scots Sweetpea. Perennial, 60-200 cm tall, clambering, the stems broadly winged, glabrous; stipules 20-34 mm long, semi-sagittate; leaves 3-12 (15) cm long (excluding tendrils); leaflets 2, 30-120 mm long, 5-40 mm wide, linear- lanceolate to lanceolate or elliptic; pe- duncles 8-22 cm long; racemes 4- to 9- flowered, the flowers ascending-spreading at anthesis; calyx tube 4-5 mm long, cam- panulate, the lower lateral teeth 1.7-4 mm long, lanceolate; flowers 15-18 mm long, red or red-purple; pods 40-60 mm long, 5-8 mm wide, glabrous. Cultivated orna- mental, persisting, escaping, and established in Utah Co., and probably elsewhere; in- troduced from Europe. 2 (0). Lespedeza Michx. Perennial herbs (or woody at the base), from a caudex; leaves alternate, pinnately trifoliolate; stipules inconspicuous; flowers papilionaceous, in axillary racemes or sub- paniculate, each subtended by a bract; brac- teoles 2, attached at base of calyx; calyx 5- toothed; petals 5, pink-purple; stamens 10, diadelphous; ovary 1-ovuled, the style in- curved and beardless, the stigma small and terminal; pod short, partially included in the calyx. Lespedeza thunbergii (DC.) Nakai. Thun- bery Bush Clover. [Desmodiwn thunbergii DC.]. Perennial, 80-100 cm tall or more and 150-200 cm wide, clump-forming; pu- bescence basifixed; stems striate; petioles 0.3-0.8 cm long; leaflets 3, lacking stipules, commonly 20-50 mm long, elliptic to ob- long, glabrous above, strigose beneath; pe- duncles 0.5-1.2 cm long; racemes many- flowered; flowers 10-12 mm long (or more?), pink-purple; bracteoles lanceolate; pods obovate to oblong, to 10 mm long. Cultivated ornamental plant in Utah Co.; introduced from China. Lotus L. Plants annual or perennial herbs or suf- fnitescent, caulescent, from a taproot and caudex; leaves alternate, pinnately (or ap- pearing palmately) compound; stipules fo- liaceous, scarious, or gland-like; flowers pa- pilionaceous, in axillary pedunculate umbels or solitary; bracts leaf-like; calyx 5-toothed; petals 5, yellow or white, sometimes suf- fused with red, the keel long-attenuate; sta- Sept. 1978 Welsh: Utah Flora, Fabaceae 319 mens 10, diadelphous; ovary enclosed in the Reference staminal sheath, the style glabrous; pods Ottley, A. M. 1944. The American Loti flattened or subterete, straight, one to several- with special consideration of a pro- seeded, dehiscent. posed new section, Simpeteria. Brit- onia 5:81-123. 1. Plants annual, prostrate to ascending; flowers sessile in leaf axils; plants of Washington Co 2 — Plants perennial, sometimes flowering the first year, prostrate to erect 3 2(1). Plants subglabrous or merely strigose; calyx teeth subequal to the tube; pods 10-15 mm long L. subpinnatus — Plants villous; calyx teeth much longer than the tube; pods 5-10 mm long .... L. humistratus 3(1). Flowers sessile, solitary in leaf axils, or on peduncles to 2.6 cm long; plants indigenous in San Juan Co L. wrightii — Flowers pedunculate, solitary or 2 to several; plants variously distributed, not or seldom of San Juan Co 4 4(3). Plants suffruticose, rigid, commonly erect, internodes greatly exceeding leaf length; bracts of inflorescence 1 or 0; plants of Washington Co L. rigidus — Plants herbaceous, prostrate to ascending or erect; internode and leaf length various; bracts of inflorescence 1 or more; distribution various 5 5(4). Stipules leaflet-like; plants introduced, cultivated and escaping 6 — Stipules reduced to glands; plants indigenous 7 6(5). Flowers 3 or 4, 7-9; leaflets of main leaves lance-linear to narrowly elliptic, acute L. tenuis — Flowers 5-12, 8-12 mm long; leaflets of main leaves obovate, rounded L. corniculatus 7(5). Leaves sessile, the leaflets strictly palmate, usually drying a lead-green color; plants widely distributed in central to southwestern Utah L. utahensis — Leaves short-petiolate, the rachis elongate and at least one leaflet pinnately disposed; plants of western Kane, Washington, and Iron cos L. longebracteatus Lotus corniculatus L. Bird's-foot Trefoil, pected elsewhere; introduced from Europe; Perennial, 10-50 cm long, with ascending 2 (0). or procumbent stems, glabrous or strigose; Lotus humistratus Greene. Low Trefoil, stipules foliar, almost or quite as large as Annual, 6-30 cm long, with prostrate to as- the leaflets; leaflets 3, 5-15 mm long, 2-8 cending stems; stipules reduced to glands; mm wide, obovate, rounded apically; pe- leaves pinnate, the rachis flattened; leaflets duncles 0.5-7.5 cm long; bracts 1- to 3- 3-5, 3-15 mm long, 1-8 mm wide, obovate foliolate; flowers (1 or 2) mostly 5-12, 8-12 to oblanceolate, obtuse apically; peduncles mm long, yellow; calyx 3-4 mm long, the 0, the flowers solitary, axillary, 4-6 mm teeth subequal to the tube; pods linear long, yellow or tinged red; calyx 3-4 mm 20-35 mm long, 2-3.5 mm thick, subterete, long, the teeth much longer than the tube; straight, glabrous. Cultivated forage plant of pods 5-10 mm long, 2.5-3.5 mm wide, lat- moist pastures, persisting in Cache, Millard, erally compressed, strigulose-villous. Sandy Utah, and Washington cos., and to be ex- or gravelly sites in creosote bush and warm 320 Great Basin Naturalist Vol. 38, No. 3 desert shnib communities at 670 to 800 m in Washington Co.; Arizona, New Mexico, and California; Mexico; 4 (0). Lotus longebracteatus Rydb. Long- bracted Trefoil. [Anisolotiis longebracteatus (Rydb.) Rydb.; Hosackia rigida var. numu- laria M. E. Jones; Anisolotus numularius (M. E. Jones) Woot. & Standi.; L. numu- larius (M. E. Jones) Tidestr., not Reichb. ex Steud.; L. nwnulus Drayton; L. rigidus x utahensis Ottley]. Perennial, 7-38 cm long, with prostrate to decumbent stems radiating from a herbaceous, superficial caudex; stip- ules reduced to glands; leaves petiolate, pin- nate, with at least one leaf commonly placed along the short rachis; leaflets 3 or 4, 2-22 mm long, 1-8 mm wide, oblanceo- late to elliptic or oval (on lowermost leaves), obtuse to acute; peduncles 0.5-6.5 cm long; bracts 1- to 3-foliolate; flowers 1 or 2, 12-17 mm long, yellow, suffused with red; calyx 5-8 mm long, the tube 2.8-5.1 mm long, strigose, the teeth 2.2-3.9 mm long, shorter than the tube; pods narrowly oblong, 15-28 mm long, 3-4 mm wide, straight, strigose. Sandy and gravelly sites in desert shrub, riparian, and pinyon-juniper commimities at 670 to 1600 m in Iron, Kane (west of the Paria River), and Wash- ington COS.; Nevada and Arizona. Plants of long-bracted trefoil were considered by Ot- tley (1944: 109-113) to be hybrids between L. rigidus and L. utahensis. Specimens from Utah, even those annotated by Ottley, do not support that interpretation. They are not intermediate between the putative par- ents. Rather, the plants have features not shared by either L. utahensis or L. rigidus. Both of these taxa have erect or ascending stems from ligneous to subligneous super- ficial to subterranean caudices. The stems of L. longebracteatus are prostrate-decumbent or procumbent from a herbaceous caudex, a feature evidently overlooked by Ottley, and not shared by either of the purported par- ents. Other more subtle characteristics differ also (see descriptions); 42 (vi). Lotus rigidus (Benth.) Greene. Bush Tre- foil. [Hosackia rigida Benth.; Anisolotus rig- idus Rydb.; L. argensis Coville]. Perennial, 25-70 cm tall, with erect or ascending stems commonly woody at the base, from a ligneous or subligneous, superficial caudex; stipules reduced to glands; leaves petiolate, the rachis flattened, pinnate, with at least one leaflet along the rachis; leaflets 3-5, 2-20 mm long, 0.8-4 mm wide, oblanceo- late to oblong, obtuse to emarginate; pe- duncles 2.3-14 cm long; bracts 1- to 3- foliolate; flowers 1-3, 12-23 mm long, yel- low suffused with red; calyx 6.2-10 mm long, the tube 4.2-6.5 mm long, cylindro- campanulate, strigose, the teeth 2-4 mm long, shorter than the tube; pods narrowly oblong, 32-45 mm long, 3.7-4.2 mm wide, straight, glabrous. Sandstone outcrops and sandy to clay banks and terraces at 800 to 1070 m in Washington Co.; Nevada, Ari- zona, California, and Mexico. The bush tre- foil is a rem.arkable plant, forming rounded clumps of very brittle stems, making diffi- cult the task of representing it well on her- barium mounts; 11 (iii). Lotus subpinnatus Lag. Mohave Trefoil. [Hosackia subpinnata (Lag.) Torr. & Gray; L. wrangelianus Fischer & Meter]. Annual, 3-30 cm long, with prostrate to ascending stems, glabrous to strigose; stipules reduced to glands; leaves pinnate, the rachis flat- tened; leaflets 3-5, 1.5-15 mm long, 0.8-7 (10) mm wide, obovate, obtuse to truncate apically; peduncles 0, the flowers solitary, axillary, 4.5-7 mm long, yellow or suffused red; calyx 2.8-5 mm long, the teeth sub- equal to the tube; pods 10-15 mm long, 2.8-3.1 mm wide, compressed, sparingly strigose. Salt desert shrub and warm desert shrub communities at 870 to 1100 m in Washington Co.; Nevada and California; 4 (0). Lotus tenuis Kit. in Willd. Slender Tre- foil. [L. corniculatus ^ tenuifolius L.; L. tenuifolius (L.) Reichb.]. Perennial, 20-60 cm long, with weak, ascending stems, glabrous; stipules foliar, almost or quite as large as the leaflets; leaflets 3, 5-15 mm long, 2-4 mm wide, lanceolate to narrowly oblanceolate or lance-linear, acute apically; peduncles 2-7.5 cm long; bracts 1- to 3- foliolate; flowers 2-4, 7-9 mm long, yellow (often drying blue); calyx 4-5 mm long, the teeth subequal to the tube; pods linear, 25-30 mm long, 2-3 mm thick, subterete, straight, glabrous. Cultivated forage plant of Sept. 1978 Welsh: Utah Flora, Fab ace ae 321 moist meadows, persisting in Utah Co., and probably elsewhere; introduced from Eu- rope; 1 (0). Lotus utahensis Ottley. Utah Trefoil. Perennial, 15-43 cm tall, with erect- ascending stems from a shallowly sub- terranean, subligneous caudex; stipules re- duced to glands; leaves sessile, palmate; leaflets 3-5, 2-23 mm long, 1.5-7 mm wide, spatulate to oblanceolate or oblong, obtuse to acute; peduncles 1.4-7.5 cm long; bracts 1- to 3-foliolate; flowers 1-5, 12-16 mm long, yellow, suffused with red; calyx 4.5-8.7 mm long, the tube 3.3-4.5 mm long, shorter than the tube; pods narrowly ob- long, 22-35 mm long, 2.5-3.5 mm wide, minutely strigulose to glabrate, shining, straight. Sagebrush, pinyon-juniper, moun- tain brush, aspen, and spruce-fir commu- nities at 1470 to 2730 m in Beaver, Gar- field, Iron, Kane, Millard, Piute, Sevier, Utah, Washington, and Wayne cos.; Nevada and Arizona; 79 (xvi). Lotus wrightii (A. Gray) Greene. Wright Trefoil. [Hosackia wrightii A. Gray; Ani- solotus ivrightii (A. Gray) Rydb.]. Perennial, 12-60 cm tall, with erect-ascending stems from a commonly superficial caudex; stip- ules reduced to glands; leaves petiolate (sometimes shortly so), palmate; leaflets 3-5, 3-22 mm long, 1-5 mm wide, spatulate to oblanceolate, oblong, linear, obtuse to acute; peduncles 0-2.6 cm long; bracts 1-to 5-foliolate; flowers commonly solitary, rarely 2, 14-18 mm long, yellow, suffused with red; calyx 7.5-9.2 mm long, longer than or subequal to the tube; pods narrowly oblong, 25-34 mm long, 2-2.6 mm wide, strigulose to villosulous, straight. Ponderosa pine woods at 1830 to 2130 m in San Juan Co.; Colorado, Arizona, and New Mexico; 4 (0). LupiNus L. Plants annual or perennial herbs; leaves alternate, palmately compound; stipules slender, persistent; flowers borne in terminal racemes, perfect; calyx bilabiate, the lips entire or toothed, commonly with brac- teoles; petals usually blue or blue-purple, less commonly whitish, yellowish, or red- dish, the banner variously reflexed, glabrous or variously hairy dorsally, the wings mostly glabrous, the keel glabrous or ciliate on up- per (less commonly on lower) edges; sta- mens 10, monadelphous, with 5 long fila- ments alternating with 5 short ones; pods laterally compressed, 2- to several-seeded. Note: The genus is notoriously difficult be- cause of lack of clear diagnostic features. Taxa tend to grade morphologically into each other, probably due to hybridization. Wide ranging taxa tend to intergrade with all other taxa which they contact. Because of these problems, and because of the likeli- hood of cleistogamy in some taxa, some specimens can be assigned to described en- tities only arbitrarily. The following key represents an attempt to provide mean- ingful determination of most of the mate- rials from Utah. It is tentative at best. References Cox, B. J. 1970. A monograph of the Lu- pinus ornatus complex. M. S. Thesis, University of Missouri, 133 pp. Dunn, D. B. 1956. Leguminosae of Nevada, Par. II. Lupinus. Contr. Flora Nevada 39:1-64. 1957. A revision of the Lupinus ar- bustus complex of the Laxiflori. Madrono 14:54-73. 1959. Lupinus pusillus and its rela- tionship. Amer. Midi. Naturalist 62:500-510. 1964. Lupinus. pp. 140-43. In: Welsh, S. L., et al. Common Utah Plants, Brigham Young University Press, Provo, Utah 312 pp. Fleak, L. S. and D. B. Dunn. 1971. No- menclature of the Lupinus sericeus complex (Papilionaceae). Trans. Mis- souri Acad. 5:85-88. Hess, L. W. 1969. The biosystematics of the Lupinus argenteus complex and al- lies. Ph.D. Dissertation, University of Missouri. 230 pp. 1. Plants annual, the cotyledons commonly persistent Key I Plants perennial, the cotelydons not present at flowering Key II 322 Great Basin Naturalist Vol. 38, No. 3 Key I. Plants Annual 1. Leaflets long-pilose on both surfaces; cotyledons petiolate; ovules 4-6; plants of Washington Co L. concinnus — Leaflets variously hairy beneath, glabrous or glabrate above; cotyledons sessile; ovules 2-6; plants of various distribution 2 2(1). Flowers borne in subcapitate racemes, the rachis commonly 2 cm long or shorter 3 — Flowers borne in elongate racemes, the rachis commonly 2 cm long or longer 4 3(2). Plants subacaulescent or acaulescent, the internodes seldom to 1 cm long; upper calyx lip 2 mm long or less, entire L. brevicaulis — Plants caulescent, with several developed internodes, at least some more than 2 cm long; upper calyx lip 3-6 mm long, bilobed L. kingii 4(2). Plants caulescent; keel petals ciliate, at least on lower margin near the base; ovules 4-6; plants of Washington Co L. sparsiflorus — Plants acaulescent, subacaulescent, or short caulescent; keel petals ciliate on upper edges towards the apex; ovules 2; plants of various distribution 5 5(4). Peduncles L5-7.5 cm long; pods not constricted between the seeds; plants subacaulescent to short caulescent, known from Washington Co L. flavoculatus — Peduncles 0-3.5 cm long; pods constricted between the seeds; plants mostly short caulescent, widespread in Utah L. pusillus Key n. Plants Perennial L Leaflets glabrous above 2 — Leaflets pubescent above 6 2(1). Banner reflexed at or below the midpoint, glabrous or pubescent distally along the crest; plants of Washington Co L. laxiflorus — Banner reflexed above (beyond) the midpoint, hairy if at all beneath the upper calyx lip, or as if as above then not of Washington Co 3 3(2). Leaves mainly basal, the petioles 8-13 cm long or more, coarsely hirsute; stems from a rhizome; plants of San Juan Co L. anunophilus — Leaves mainly well distributed along the stem, the petioles commonly less than 8 cm long, stri-gose or silvery hairy; stems from a caudex; plants of various distribution 4 4(3). Flowers 5-7 mm long; keel densely ciliate; plants of Salt Lake, Summit, and San Juan cos L. argenteus — Flowers 8-12 mm long; keel thinly ciliate, or the margin glabrous 5 5(4). Leaflets abruptly obtuse to rounded; racemes more than 15 cm long L. argenteus — Leaflets obtuse to acute (rarely rounded); racemes mostly less than 15 cm long L. inaculatus 6(1). Banner glabrous dorsally 7 — Banner pubescent on the back (look beneath upper lip of calyx) 11 Sept. 1978 Welsh: Utah Flora, Fabaceae 323 7(6). Flowers 10-13.5 mm long; calyx saccate-gibbous or shortly spurred; leaves mainly basal; plants of Uintah and Kane cos L. wyethii — Flowers 7-13 mm long; calyx not especially gibbous; leaves distributed along the stem, or all leaves basal; plants of various distribution 8 8(7). Plants acaulescent or short-caulescent; leaves essentially all basal 9 — Plants caulescent; leaves distributed along the stem 10 9(8). Plants low, 3-12 cm tall; racemes sessile, surpassed by the foliage; distribu- tion montane L. caespitosus — Plants 15-40 cm tall; racemes shortly pedunculate, much surpassing the foliage; distributed in western Iron and Washington cos L. volutans 10(8). Stems appressed and commonly silvery-hairy; leaflets densely silvery-hairy; plants of Iron and Washington cos L. hillii — Stems merely strigose, appearing green; leaflets thinly strigose; plants mainly of northern Utah L. argenteus 11(6). Calyx with a gibbous-saccate spur at base of upper lip; wings pubescent, or keel ciliate below the claws (rarely glabrous) 12 — Calyx at most gibbous at base of upper lip; wings and lower edge of keel glabrous 13 12(11). Wings not ciliate; flowers yellow or varicolored in populations; plants of far western Utah L. arbustus — Wings or keel (or both) ciliate below the claws (rarely glabrous); flowers commonly blue-purple; plants widespread L. caudatus 13(11). Banner reflexed at or below the midpoint, strigose to thinly strigose near the tip, or rarely hairy along the crest 14 — Banner reflexed beyond the midpoint, strigose on the back beneath the calyx lobe or over much of the back t 15 14(13). Leaves mainly basal, the plants commonly less than 50 cm tall; known from Juab, Millard, Tooele, and Utah cos L. prunophilus — Leaves mainly well distributed along the stem, the plants commonly more than 50 cm tall; distribution broad L. serviceus 15(13). Stems velvety or woolly hairy; plants commonly of meadows and stream ter- races in Duchesne, Salt Lake, Summit, Wasatch, Uintah, and Utah cos L. leucophyllus — Stems with appressed, ascending, spreading, or retrose hairs, but not velvety or woolly; distribu-tion and habitat various 16 16(15). Stems with spreading or retrose hairs; plants of Washington Co 17 — Stems with suppressed or ascending hairs; plants variously distributed 18 17(16). Flowers 7-9 mm long L. palmeri — Flowers 12-16 mm long L. jonesii 18(16). Flowers semi-orbicular viewed laterally; leaflets usually oblanceolate L. alpestris — Flowers narrow viewed laterally; leaflets linear-oblanceolate L. argenteus 324 Great Basin Naturalist Vol. 38, No. 3 Lupinus alpestris A. Nels. Mountain Lu- pine. Perennial, 25-65 cm tall, from a woody caudex, the basal leaves commonly deciduous by anthesis; pubescence of stems mainly strigose to strigulose or pilosulous; petioles 1.5-7 cm long; leaves mainly cau- line; leaflets 7-9, 8-65 mm long, 3-14 mm vsdde, oblanceolate to elliptic, strigulose to strigose on both sides, sometimes sparingly so above; pedimcles 0.5-4 cm long; racemes 3- 59 35-flowered, 4-15 cm long in an- thesis, 4-23 cm long in fruit; pedicels 2-5 mm long; flowers 8-12 mm long, blue- purple or rarely white; upper lip of calyx somewhat gibbous; banner with a central white or yellow spot, pubescent on the back near the calyx lip, reflexed above the middle; wings glabrous or rarely pubescent near the claw; keel sparingly ciliate near the apex; ovules 3-5. Sagebrush, mountain brush, pinyon-juniper, ponderosa pine, and aspen commimities 1530 to 2800 m in Bea- ver, Box Elder, Carbon, Duchesne, Emery, Garfield, Grand, Iron, Kane, Juab, Millard, Piute, Salt Lake, San Juan, Sanpete, Sevier, Summit, Tooele, Utah, Wasatch, and Wash- ington COS.; Saskatchewan and Montana south to Nevada, Arizona, and New Mexico. The status of mountain lupine, often cited as L. X alpestris, is in doubt, but is represen- tative of the nature of taxa within the genus as a whole. Specimens occupy a morpholo- gical gradient which overlaps both L. ar- genteus sens. lat. and L. caudatus sens. lat. Its distribution and range of variation ex- ceeds that of both of those taxa, however, indicating that more is involved than mere hybridization and introgression. The taxon, as described herein, serves as a kind of bo- tanical wasteland into which those speci- mens are dumped which seem beyond the limits of L. argenteiis and L. caudatus. The enormous range of variation is indicated in the treatment by Hess (1969), in which some 24 synonyms, mainly based on mate- rials from Nevada, are listed for L. alpestris; 51 (X). Lupinus ammophilus Greene. Sand Lu- pine. Plants perennial, 13-70 cm tall, from a rhizome-like subterranean caudex; pu- bescence of stems hirsute-pilose leaves mainly basal; petioles of lowermost leaves (4) 8-19 cm long, those of cauline leaves 3-14 cm long; leaflets 8-11, 12-75 mm long, 4-18 mm wide, oblanceolate to obo- vate, pilose beneath, glabrous above; pe- duncles 4-13 cm long; racemes 19- to 48- flowered, 8-25 cm long in anthesis, 8-36 cm long in fruit; flowers 8-14 mm long, blue-purple or rarely white; pedicels 4.5-9 mm long; calyx gibbous at base of upper lip; banner with a central yellow or white spot, glabrous dorsally, reflexed above the midpoint; wings glabrous, keel sparingly ci- liate near the apex; ovules 3-7. Sagebrush, pinyon-juniper, mountain brush, ponderosa pine, and aspen- Douglas fir woods, com- monly in sandy soils at 1830 to 2730 m in San Juan Co.; Colorado and New Mexico; 10 (iii). Lupinus arbustus Douglas ex Lindl. Spurred Lupine. Perennial, 26-60 cm tall, from a woody caudex; pubescence of stems minutely strigulose; leaves scattered along the stem, but with much shorter petioles upwards; petioles 2.5-16 cm long; leaflets 7-13, 24-50 mm long, 3-10 mm wide, ob- lanceolate, pilose on both surfaces; pe- dimcles 2-5 cm long; racemes 12- to 46- flowered, the axis 2.5-18 cm long in an- thesis, 4-23 cm long in fruit; flowers 10-14 mm long, yellow or white or blue-purple, or all of these in populations, or in the same raceme; pedicels 1.5-6 mm long; calyx with a gibbous-saccate spur at base of up- per lip, the spur 1.5-2.5 mm long; banner with a central white, yellowish, or brownish spot, pubescent dorsally, reflexed near the midpoint, wings ciliate along the upper edge near the apex; keel ciliate along the upper margin; ovules 5 or 6. Sagebrush and pinyon-juniper communities at 2135 to 2440 m in the Deep Creek Mountains, Juab and Tooele cos.; Washington east to Montana and south to California and Nevada. Our material belongs to var. calcaratus (Kel- logg) Welsh Stat. nov. based on Lupinus calcaratus Kellogg Proc. Calif. Acad. Sci. 2:195. 1862. [L. laxiflorus var. calcaratus (Kellogg) Dunn]. This is one of our most beautiful species of lupine; 11 (ii). Lupinus argenteus Pursh, Silvery Lupine. Plants perennial, 18-90 cm tall, from a su- perficial caudex, puberulent to strigose on Sept. 1978 Welsh: Utah Flora, Fabaceae 325 stems and petioles; leaves mainly cauline; above the midpoint, the wings and keel petioles 1.5-8 cm long: leaflets 6-9, 7-95 glabrous or variously sparingly ciliate; mm long, 2-22 mm wide, oblanceolate to ovules 3-6. The silvery lupine is represent- spatulate or almost linear, flat or folded, ed in Utah by several more or less dis- strigulose to strigose on both surfaces or al- tinctive but integrading varieties. Further- most to quite glabrous above; peduncles more, at least some of the phases grade 1.5-14.5 cm long; racemes 15- to 92-flower- further into other taxa, especially into L. al- ed, 5-24 cm long in anthesis, the axis 6-29 pestris, L. caiidatiis, and also into L. se- em long in fruit; flowers (5-7) 8.5-16 mm riceus. Silvery lupine, along with those spe- long, blue-purple, blue, white or rarely oth- cies, constitutes the most common and most er hues; pedicels 1.5-6 mm long; calyx gib- widespread of the perennial lupines in the bous or rounded at base of upper lip; ban- state. The large proportion of the specimens ner with a central yellow or white spot, encountered can be segregated by use of pubsecent or glabrous dorsally, reflexed the following, admittedly arbitrary, key. 1. Leaflets more or less evenly pubescent above, generally folded, the upper surface thus obscured 2 — Leaflets glabrous above, or with hairs scattered or merely with a few near the margin, flat or folded 3 2(1). Flowers (8) 9-11.5 mm long, narrow when viewed laterally; plants of broad distribution L. argenteus var. tenelliis — Flowers 11-14 mm long, orbicular when viewed laterally; plants of low ele- vations in east central Utah L. argenteus var. moahensis 3(1). Leaflets commonly folded, narrowly oblanceolate; banner usually pubescent beneath the upper lip of calyx; keel ciliate near the apex; plants of low to moderate elevations 4 — Leaflets commonly flat, oblanceolate to broadly so; banner usually glabrous dorsally, the keel margin glabrous or hairy only near the apex; plants of moderate to high elevations 5 4(3). Wings 4.5-6 mm wide; flowers 10-15 mm long, orbicular when viewed laterally; plants of central to eastern Utah L. argenteus var. argenteus — Wings 3-4.5 mm wide; flowers 8-11.5 mm long, narrow when viewed laterally; plants mainly of central to western Utah L. argenteus var. tenellus 5(3). Keel densely ciliate near the apex; flowers 5-7 mm long; plants uncommon in Utah, passing into the following varieties L. argenteus var. parviflorus — Keel moderately to sparingly ciliate near the apex; flowers commonly. 8 mm long or more; plants common at high elevations 6 6(5). Flowers spreading at anthesis; stems ^slender, often suffused with red near the base L,- argenteus var. rubricaulis — Flowers pendant at anthesis; stems stout, seldom red at the base L. argenteus var. boreus Var argenteus [L. garrettianus C. P. . Sevier, Uintah, and Wayne cos.; Alberta Smith, type from 1 mile west of Duchesne]. and Saskatchewan south to Oregon, Califor- Grasslands, river terraces, sagebrush, and nia, Nevada, New Mexico, and the Dakotas. pinyon-juniper communities at 1575 to 3050 In Utah this variety passes mainly into var. m in Carbon, Duchesne, Garfield, San Juan, tenellus and var. moahensis. From the latter 326 Great Basin Naturalist Vol. 38, No. 3 it differs inter alia in pubescence position on the leaves; 27 (xi). Var. boreus (C. P. Smith) Welsh comb, nov. based on Lupinus spathulatus var. boreus C. P. Smith, Species Lupinorum 746. 1952. [L. spathulatus Rydb., type from Wasatch Mts.; L. alsophilus Greene.] Spruce-fir woods, meadows, aspen, mixed conifer, and upper mountain brush commu- nities at 2370 to 3350 m in Beaver, Carbon, Davis, Emery, Garfield, Iron, Piute, Salt Lake, Sanpete, Sevier, Summit, Utah, and Wasatch cos.; Montana and South Dakota, south to Wyoming and Colorado. The spathulate lupine forms almost a continuum with var. rubricaulis. They are approx- imately sympatric in about the same kinds of habitats through much of their ranges; 20 (viii). Var. moabensis (Dunn & Harmon) Welsh Stat. nov. based on Lupinus argenteus ssp. moabensis Dunn & Harmon. Mixed desert shrub and pinyon-juniper communities, of- ten in washes, at 1400 to 1630 m in Grand and Wayne cos.; Colorado. The Moab lu- pine differs from var. argenteus, with which it is closely allied, in the features empha- sized in the key. Further differential charac- teristics are to be found in the very early flowering time of the Moab lupine. How- ever, this is at least partly a response to the earlier season of the lower elevation habi- tats, and might not be genetic; 7 (iii). Var. parviflorus (Nutt.) C. L. Hitchc. [L. parviflorus Nutt. ex Hook. & Am.; L. ar- genteus ssp. parviflorus (Nutt.) Phillips]. Spruce-fir, lodgepole pine, and aspen com- munities at 2270 to 2730 m in Duchesne, Salt Lake, Summit, and San Juan cos.; Idaho, Wyoming, and Colorado. Our mate- rial is mostly intermediate towards one or more of the varieties of L. argenteus, espe- cially with those that occur at higher eleva- tions; 4 (0). Var. rubricaulis (Greene) Welsh comb, nov. based on Lupinus rubricaulis Greene, PI. Baker. 3:35. 1901. [L. caudatus var. rubricaulis (Greene) C. P. Smith]. Spruce- fir, meadow, mixed conifer, aspen, and ripa- rian commimities at 2470 to 3370 m in Bea- ver, Carbon, Daggett, Duchesne, Garfield, Iron, Piute, Sanpete, Sevier, Summit, Uin- tah, Utah, and Wasatch cos.; Idaho, Mon- tana, and South Dakota, south to Nevada and New Mexico. The red-stemmed lupine differs in degree only from var. boreus, and the two might best be treated as variants within a single variety. If they are so treated, then the name becomes var. rubr- icaulis as that name has priority in rank. Specimens from the north slope of the Uinta Mts. (Welsh 1584 BRY; Goodman 527, 169 RM) are apparently intermediate towards the more northern L. monticola Rydb.; 38 (xvi). Var. tenellus (Douglas) Dunn. [L. tenellus Douglas ex G. Don; L. laxiflorus var. te- nellus (Douglas) T. & G.; L. foliosus var. stenophyllus Nutt.; L. stenophyllus (Nutt.) Rydb.; L. argenteus var. stenophyllus (Nutt.) Davis]. Sagebrush, pinyon-juniper, mountain brush, aspen, grassland, and mixed conifer communities at 1570 to 3130 m in Cache, Carbon, Davis, Duschesne, Garfield, Grand, Juab, Millard, Piute, Salt Lake, San Juan, Sevier, Uintah, Utah, Wasatch, and Wash- ington COS.; Alberta and Saskatchewan south to California, Arizona, New Mexico, and the Dakotas. The slender lupine is most similar to var. argenteus, from which it differs mainly in size of flowers that aver- age smaller (9-11.5 mm, not 10-16). It is with this phase of L. argenteus that L. al- pestris is most easily mistaken. Indeed, there is no clear way to distinguish all specimens as belonging to one or to the other; 28 (xii). Lupinus brevicaulis S. Wats. Short- stemmed Lupine. Annual, 4-11 cm tall, from a taproot; cotyledons sessile; stems 0-2 cm long, when developed at all usually ob- scured by the leaf bases; leaves in a basal tuft; petioles 0.8-6.5 cm long; leaflets 3-9, 5-18 mm long, 1.5-9 mm wide, oblanceol- ate, flat or folded, pilose beneath, glabrous above (except marginally in some); pe- duncles 0.6-6.5 cm long; racemes 4- to 12- flowered, 1-2.5 cm long in anthesis, the axis 1.5-3 cm long in fruit; flowers 5.2-7 mm long, blue-purple or white; pedicels 0.3-0.8 mm long; calyx tapering to the pedicel, the upper lip very short; banner with a central yellow spot, glabrous dorsally, reflexed near the midpoint; ovules 2 or 3. Salt desert shrub, pinyon-juniper, sagebrush, blackbrush. Sept. 1978 Welsh: Utah Flora, Fabaceae 327 and creosote bush communities at 830 to Wasatch, and Wayne cos.; Oregon to Mon- 1970 m in Beaver, Carbon, Daggett, Gar- tana and south to Cahfornia, Nevada, and field, Millard, Salt Lake, Uintah, Utah, and Colorado. This distinctive taxon has been Washington cos.; Oregon east to Colorado, treated as belonging to an expanded L. le- and south to Arizona and New Mexico. The pidiis Dougl. ex Lindl., but seems best rec- short-stemmed lupine forms apparent hybr- ognized at specific rank, in our flora at ids with L. flavoculatiis; 18 (i). least; 33 (xi). Lupinus caespitosus Nutt. ex Torr. & Lupinus caudatus Kellogg. Spurred Lu- Gray. Stemless Lupine. [L. aridus var. iita- pine. Perennial, 21-80 cm tall, from a hensis S. Wats., type from Parley's Park; L. woody caudex; leaves mainly cauline; pe- watsonii Heller; L. lepidus var. utahensis (S. tioles 1-12 cm long, commonly 2-8 cm Wats.) C. L. Hitchc; L. lepidus ssp. caespi- long; leaflets 5-90, 10-45 (60) mm long, tosus (Nutt.) Detling]. Perennial, caespitose, 2-14 cm wide, oblanceolate to elliptic or 2.5-11.5 cm tall, acaulescent or essentially narrowly oblanceolate, pilose on both sur- so, from a superficial caudex; leaves mainly faces or glabrate above; peduncles 1-6.5 cm basal; petioles 1-9 cm long; leaflets 5-9, long; racemes 10- to 57-flowered, 3-16 cm 3-25 mm long, 1.5-6 mm wide, oblanceol- long in anthesis, the axis 4.5-17 cm long in ate, mucronate, pilose on both sides; pe- fruit; flowers 8-12.5 (13.5) mm long, blue- duncles 0-2 cm long; racemes 12- to 40- purple or less commonly white; pedicels flowered, 1-4 cm long in anthesis, the axis 1-3 (5) mm long; calyx with a gibbous- 2-6 cm long in fruit; flowers 7-8.5 mm saccate spur 0.2-1.5 (2) mm long at the long, blue-purple or white; pedicels 2.5-4 base of the upper lip; banner pubescent mm long; calyx tapering to the pedicel, the dorsally, reflexed at or beyond the mid- upper lip well developed; banner with a point; wings commonly ciliate above and central yellow spot, glabrous dorsally, re- near the claws, the keel commonly ciliate flexed below the midpoint; ovules 2-4. above and near the claws; ovules 3-5. Meadows, open deciduous woodland, mixed Three rather weak varieties are known from conifer, and sagebrush commimities at 2130 Utah. They are separable only arbitrarily, to 3350 m in Beaver, Cache, Carbon, but seem to represent at least trends within Emery, Grand, Kane, Garfield, Iron, Piute, the variation. Sanpete, Sevier, Summit, Uintah, Utah, 1. Banner reflexed at the midpoint; leaflets rather broadly oblanceolate; plants rare in Kane Co L. caudatus var. cutleri — Banner reflexed beyond the midpoint; leaflets only rarely broadly oblanceo- late; plants of various distribution 2 2(1). Leaflets bicolored, green to yellow-green above, dull green to grayish beneath; plants uncommon, of southern Utah L. caudatus var. argophyllus — Leaflets more uniformly colored, either green or gray on both surfaces, or only somewhat bi-colored; plants common in much of Utah L. caudatus var. caudatus Var. argophyllus (A. Gray) Welsh stat. nov. based on Lupinus decumbens var. argo- phyllus A. Gray Mem. Amer. Acad. 4:37. 1849. [L. caudatus ssp. argophyllus (A. Gray) Phillips; L. argophyllus (A. Gray) Cockerell; L. laxifhrus var. argophyllus (A. Gray) M. E. Jones; L. helleri Greene; L. aduncus Greene]. Pinyon-juniper, mountain brush, ponderosa pine, and grassland com- munities at 1570 to 2430 feet in Beaver, Garfield, Kane, and San Juan cos.; Wyom- ing south to New Mexico; 7 (iii). Var. caudatus. [L. holosericeus var. utah- ensis S. Wats., type from Wasatch Mts.; L. argentinus Rydb.; L. lupinus Rydb., type from Bears Ears, Elk Mt.; L. utahensis Moldenke]. Sagebrush, pinyon-juniper, mountain brush, ponderosa pine, aspen, mixed 328 Great Basin Naturalist Vol. 38, No. 3 conifer, and grassland communities at 1470 to 2730 m in Beaver, Box Elder, Cache, Davis, Duchesne, Garfield, Iron, Juab, Kane, Millard, Morgan, Piute, Rich, Salt Lake, San Juan, Tooele, Uintah, Utah, Wasatch, Wayne, and Weber cos.; Oregon, Idaho, and Wyoming south to California, Nevada, Arizona, and Colorado. This variety forms intermediates with L. argentetis (especially with var. tenellus), L. hillii, L. sericeus, and the other varieties of L. caudatus; 67 (vii). Var. cutleri (Eastw.) Welsh stat. nov. based on Lupinus cutleri Eastw., Leafl. West. Bot. 4:192. 1945. [L. caudatus ssp. cutleri (Eastw.) Hess]. Pinyon-juniper wood- land at 1570 m along the Cockscomb in Kane Co.; Arizona, Colorado, and New Mexico; 1 (i). Lupinus concinnus Agardh. Elegant Lu- pine. [L. micensis M. E. Jones; L. orcuttii S. Wats.; L. concinnus ssp. orcuttii (S. Wats.) Dunn.] Annual, caulescent, 6.5-20 cm tall, from a taproot; cotyledons petiolate; stems with apparent internodes; leaves mainly cauline, the lowermost the smallest; petioles 0.6-8 cm long; leaflets 5-8, 4-24 mm long, 1.5-7.5 mm wide, obovate to oblanceolate, pilose on both sides; peduncles 0.4-2 cm long; racemes 6- to 17-flowered, 2-4 cm long in anthesis, the axis 4-7 cm long in fruit; flowers 7-8.8 mm long, blue-purple (rarely white); pedicels 0.5-1.2 mm long; calyx tapering to the pedicel; upper lip well developed; banner with a central yel- low spot, glabrous dorsally, reflexed at or below the midpoint; ovules 2-4. Warm desert shrublands at 770 to 1070 m in Washington Co.; New Mexico, Arizona, and Nevada. Our material belongs to var. or- cuttii (S. Wats.) C. P. Smith; 10 (ii). Lupinus flavoculatus Heller. Yellow-eyed Lupine. [L. rubens var. flavoculatus (Heller) C. P. Smith; L. odoratus var. flavoculatus (Heller) Jepson]. Annual, 3-11 cm tall, from a taproot; cotyledons sessile; stems 0-2 cm long, often with at least one apparent inter- node; leaves mainly basal; petioles 0.5-8.5 cm long; leaflets 6-10, 5-20 mm long, 2-9 mm wide, obovate to oblanceolate, flat or folded, pilose to glabrate beneath, glabrous above, long ciliate; peduncles 1.5-7.5 cm long; racemes 9- to 16-flowered, 1.2-3 cm long in anthesis, the axis 2-4 cm long in fruit; flowers 5.1-7.2 mm long, blue-purple or less commonly white; pedicels 0.5-2 mm long; calyx tapering to the pedicel, the up- per lip short; banner with a central yellow spot, glabrous dorsally, reflexed near the midpoint; ovules 2-4. Warm desert shrub and pinyon-juniper communities at 770 to 2330 m in Washington Co.; Nevada and California. This plant apparently hybridizes with L. brevicaulis, from which it cannot be separated in all cases. Possibly, the yellow- eyed lupine might best be treated at some infraspecific level within that taxon; 5(0). Lupinus hillii Greene. Hill Lupine. [L. pulcher Eastw., type from 18 miles south of Cedar City]. Perennial, 24-65 cm tall, from a woody caudex; leaves mainly cauline; pe- tioles 0.6-8 cm long; leaflets 5-9, 7-46 mm long, 2-8 mm wide, oblanceolate to narrow- ly so, often folded, pilose on both surfaces though less densely so above; peduncles 0.5-18 cm long; racemes 24- to 73-flower- ed, 5-20 cm long in anthesis, the axis 6-23 cm long in fruit; flowers 6-8.5 mm long, blue-purple; pedicels 2-4.5 mm long; calyx gibbous at base of upper lip; banner with a central yellow spot, glabrous dorsally, re- flexed near the midpoint; ovules 2 or 3. Sagebrush and pinyon-juniper to ponderosa pine communities at 2130 to 2730 m in Pine Valley Mts., Washington and Iron cos.; Arizona. The Hill lupine approaches, if not passes into, L. palmeri on the one hand, and into L. caudatus on the other. Much more material is necessary for an adequate inter- pretation of our somewhat meager speci- mens; 6 (ii). Lupinus jonesii Rydb. Jones Lupine. Pe- rennial, 50-120 cm tall, from a woody cau- dex; pubescence of stems spreading-hirsute; leaves mainly cauline; petioles 1-8 cm long; leaflets 6-9, 9-60 (70) mm long, 3-13 mm wide, oblanceolate to oblong-oblanceolate, folded or flat, pilose on both surfaces; pe- duncles 4.5-12 cm long; racemes 18- to 65- flowered, 8-28 cm long in anthesis, the axis 9-30 cm long in fruit; flowers 12-15 (16) mm long, pallid, or blue-purple; pedicels 2-5 mm long; calyx strongly gibbous at base of upper lip; banner with a central yellow or brown spot, more or less strigose dor- Sept. 1978 Welsh: Utah Flora, Fabaceae 329 sally, reflexed at or beyond the midpoint; ovule number not known. Pinyon- juniper and mountain brush communities at 1170 to 2330 m in Washington Co. (type from Sil- ver Reef); endemic. Affinities of L. jonesii appear to lie with L. sericeus through var. barbiger; 3(0). Lupinus kingii S. Wats, King Lupine. [L. sileri S. Wats., type from southern Utah; L. capitatus Greene]. Annual, 6-24 cm tall, from a taproot; cotyledons sessile; stems with two or more apparent internodes; leaves mainly cauline; petioles 0.8-3.5 cm long; leaflet 5-7, 7-23 mm long, 2-6.5 mm wide, oblanceolate, flat or folded, long- pilose on both surfaces or glabrous medially above or overall; peduncles 0-9.5 cm long; racemes 5- to 12-flowered, 1-2.3 cm long at anthesis, the axis 1.8-3.7 cm long in fruit; flowers 7.8-9.2 mm long, blue purple or less commonly pallid; calyx somewhat gib- bous at base of upper lip; banner with a central yellow spot, glabrous dorsally, re- flexed below the midpoint; ovules usually 2. Two varieties are present. 1. Racemes sessile, the peduncles obsolete, contained within the foliage L. kingii var. argillaceus Racemes pedunculate, the peduncles 1-9.5 cm long, commonly produced beyond the foliate L. kingii var. kingii Var. argillaceus (Woot. & Standi.) C. P. Smith. (L. argillaceus Woot. & Standi.) Pinyon-juniper woodland at 2400 m in the Henry Mts., Garfield Co.; New Mexico. This is a striking phase, which is evidently rare in Utah; 1 (0). Var. kingii. Sagebrush, pinyon-jvmiper, mountain brush, and ponderosa pine com- munitfes at 1830 to 2440 m in Garfield, Iron, Kane, Piute, San Juan, Sevier, Wasatch (type from Heber Valley), and Washington cos.; Nevada, Arizona, New Mexico, and Colorado. 24 (iv). Lupinus latifolius Agardh. Broad-leaved Lupine. Perennial, 30-120 cm tall, from a branching caudex; pubescence appressed strigose or almost lacking; leaves mainly cauline; petioles 4-20 cm long, leaflets 5-11, 25-90 mm long, 4-20 mm wide, ob- long to elliptic or oblanceolate, flat, glabrous above, thinly appressed-strigose be- neath; racemes 10- to 35-flowered, 8-25 cm long in anthesis, 10-40 cm long in fruit; flowers 10-14 cm long, blue to pinkish, fad- ing brown; pedicels 6-12 mm only; calyx not gibbous at the base; banner with a cen- tral yellow spot; glabrous dorsally, reflexed below the midpoint; ovules 7-10. Oakbrush and streamside communities at ca. 1230 m in Washington (Zion National Park) Co.; California, Oregon, Washington. Our mate- rials appear to belong to var. columbianus (Heller) C. P. Smith. [L. columbianus Hel- ler] 2(0). Lupinus leucophyllus Dougl. ex Lindl. White-leaved Lupine. [L. eatonanus C. P. Smith, type from Hailstone]. Perennial, 40-90 cm tall or more, from a woody cau- dex; pubescence of stems dense, appressed to spreading; leaves mainly cauline; petioles 1.2-16 cm long; leaflets 7-10, 9-70 mm long, 3-13 mm wide, oblanceolate, flat or folded, villous-pilose on both surfaces; pe- duncles 2.5-5.5 cm long; racemes 22- to 70- flowered, 7-18 cm long in anthesis, the axis 8-19 cm long in fruit; pedicels 2.5-5 mm long; flowers 10-15 mm long, blue-purple or less commonly pallid; calyx gibbous to saccate-gibbous at base of upper lip; banner with a central yellow spot, densely hairy dorsally, reflexed above the midpoint; ovules 4-6. Terraces at 1450 to 2300 m in Duchesne, Summit, Uintah, Utah, and Wasatch cos.; Washington south to Nevada and east to Wyoming. The white-leaved lu- pine forms apparent hybrids with L. cau- datus, among others; 9 (iv). Lupinus maculatus Rydb. Spotted Lu- pine. Perennial, 40-85 cm tall, from a woody caudex; pubescence of stems ap- pressed or subappressed or lacking; leaves mainly cauline; petioles 1.8-4 cm long; leaf- lets 5-8, 14-75 mm long, 2.5-18 mm wide, oblanceolate, mainly flat, sparingly pilose 330 Great Basin Naturalist Vol. 38, No. 3 beneath, glabrous above, ciliate, obtuse to acute apically; peduncles 0.8-9 cm long; racemes 20- to 39 (50) -flowered, 5-15 (23) cm long in anthesis, the axis 8-25 cm long in fruit; pedicels 2-5 mm long; flowers 8.5-14 mm long, blue-purple or white, fad- ing tan; calyx more or less gibbous at base of upper lip; banner with a central brown (or fading brown) spot, glabrous dorsally, re- flexed above the midpoint; ovules 4 or 5. Aspen-conifer, meadow, and open conifer communities at 2230 to 2730 m in Davis, Salt Lake (?), Sevier, Utah [type from P. V. (Pleasant Valley) Junction], and Weber cos.; Idaho. Spotted lupine is a near ally of L. argenteus var. boreus and var. rubricaulis, with which it forms apparent hybrids. Per- haps it would best be treated at in- fraspecific rank in that species; 10 (i). Lupinus palmeri S. Wats. Palmer Lupine. Perennial, 24-60 cm tall, from a woody caudex; pubescence of stems more or less spreading-ascending (at least some); leaves mainly cauline; petioles 0.8-10 cm long; leaflets 6-10, 10-48 mm long, 3-8 (10) mm wide, oblanceolate, flat or folded, pilose on both surfaces; peduncles 0.5-5.5 (7) cm long; racemes 14- to 55-flowered, 4-15 cm long in anthesis, the axis 12-20 cm long in fruit; flowers 1.5-11 mm long, blue-purple; pedicels 1.2-4 mm long; calyx more or less gibbous at base of upper lip; banner with a central yellow spot, usually pubescent dor- sally, reflexed at or beyond the midpoint; ovules 2-6. Pinyon-juniper and sagebrush communities at 1830 to 2130 m in Wash- ington Co.; Nevada, Arizona, and New Mexico. The Palmer lupine, as it is known in Utah, is closely related to L. hillii, from which it is distinguished with some diffi- culty. Additional materials might dictate a change in status for this poorly known en- tity; 5(0). Lupinus prunophilus M. E. Jones. Robin- son Lupine. [L. wyethii var. prunophilus (M. E. Jones) C. P. Smith; L. arcticus var. prunophilus (M. E. Jones) C. P. Smith; L. polyphyllus ssp. polyphyllus var. pruno- philus (M. E. Jones) Phillips; L. tooelensis C. P. Smith, type from Johnson Canyon, Deep Creek Mts.] Perennial, 23-65 cm tall, from a woody caudex; pubescence of stems spreading-hirsute; leaves mainly basal; pet- ioles of lowermost leaves 7-30 cm long; leaflets 8-13, 15-75 above; peduncles 4-10 cm long; racemes 25- to 68-flowered, 6-23 cm long, the axis 8-35 cm long in fruit; flowers 10-16 mm long, blue-purple; pedi- cels 3.5-8 mm long; calyx gibbous at base of upper lip; banner with a yellow spot, glabrous dorsally, reflexed near the mid- pont; wings glabrous; keel sparingly ciliate near the apex; ovules 3-6. Sagebrush, pinyon-juniper, and mountain brush commu- nities at 1530 to 2130 m in Juab (type from Robinson), Millard, Tooele, and Utah cos.; Washington south to Nevada and east to Montana, Wyoming, and Colorado. The Robinson lupine has been regarded by Hess (1969) as L. x prunophilus, but that designa- tion seems inconsistent with the material at hand. The populations in Utah are better defined than some of the other taxa treated as species; 13 (ii). Lupinus pusillus Pursh. Dwarf Lupine. Annual, 3-24 cm tall, from a taproot; co- tyledons sessile; pubescence of stems and petioles spreading long-hairy; leaves mainly cauline; petioles 1-9 cm long; leaflets 3-9 (14), 11-48 mm long, 2-10 mm wide, oblan- ceolate, flat or folded, long-pilose beneath, glabrous above; peduncles 0.5-3.5 cm long; racemes 4- to 38-flowered, 1-17 cm long in anthesis, the axis 4-21 cm long in fruit; flowers 8.5-12 mm long, blue or variously pink or white; pedicels 1-3.5 mm long; ca- lyx tapering to the pedicel; banner with a central yellow spot, glabrous dorsally, re- flexed near the midpoint; ovules 2; pods constricted between the seeds. Three poorly differentiated and intergrading varieties are present in Utah. The following arbitrary key will serve to segregate most materials. 1. Pedimcles seldom more than 1 cm long; inflorescence shorter than the leaves; banner 5 mm wide or less L. pusilhts var. intermontanus Peduncles commonly 1-3.5 cm long; inflorescence usually longer than the leaves; banner 6-10 mm wide 2 Sept. 1978 Welsh: Utah Flora, Fabaceae 331 2(1). Calyx tube and pedicel glabrous; corolla commonly blue; plants completely transitional with the next L. pusillus var. rubens Calyx tube and pedicel pilose; corolla blue or pale to white L. pusillus var. pusillus Var. intermontanus (Heller) C. P. Smith. [L. intermontanus Heller; L. pusillus ssp. in- termontanus (Heller) Dunn]. Dunes and oth- er sandy sites in mixed desert shrub, pinyon- juniper, and mountain brush communities at 1130 to 1770 m in Duchesne, Emery, Gar- field, Kane, Millard, Salt Lake, San Juan, Tooele, and Wayne cos.; Arizona, Califor- nia, Idaho, Montana, Nevada, Oregon, Washington, and Wyoming. The long-hairy raceme axis and pedicels, along with the narrow banner and the short racemes, are diagnostic; 23 (iv). Var. pusillus. Salt desert shrub, mixed desert shrub, and pinyon-juniper commu- nities, commonly in sand, at 12.30 to 1970 m in Emery, Garfield, Grand, Kane, San Juan, Uintah, and Wayne cos.; Alberta and Saskatchewan south to Arizona, New Mexi- co, and Kansas; 39 (ix). Var. rubens (Rydb.) Welsh stat. nov. based on Lupinus rubens Rydb., Bull. Tor- rey Rot. Club 34:45. 1907. [L. odoratus var. rubens (Rydb.) Jepson; L. pusillus ssp. ru- bens (Rydb.) Dunn]. Mixed desert shrub and blackbrush communities, commonly in sand, at 800 to 1650 m in Garfield, San Juan, Washington (type from St. George), and Wayne cos.; Nevada, Arizona, and Colo- rado; 7 (ii). The maintenance of var. rubens in any taxonomic rank is dubious, since the main diagnostic features, i.e., the glabrous pedicel and calyx tube, do not fall into an either/or situation. Never-the-less, the phase of low elevations, mainly in Washington Co., do represent a trend, which, although based on this tenuous pubescence feature alone, seems worthy of taxonomic recogni- tion. Lupinus sericeus Pursh. Silky Lupine. Perennial, 30-120 cm tall, from a branching caudex; pubescence of stems short- villous to pilose or strigose, sometimes spreading; petioles 1.2-9 cm long; leaflets 5-9, 7-78 mm long, 2-15 mm wide, oblanceolate, commonly flat (at least some), pilose to puberulent on both surfaces or glabrous to glabra te above; peduncles 1.3-9 cm long; racemes 14- to 70-flowered, 6-28 cm long in anthesis, the axis 8-37 cm long in fruit; flowers 10-16 mm long, blue, blue- purple, pale, or white; pedicels 2-7 mm long; calyx more or less gibbous at the base of upper lip; banner with yellow or brown eyespot, strigose along the dorsal crest or more widely; ovules 5-7. Widely distributed in Utah, and constituting one of the three important species complexes in the state, L. sericeus, along with L. argenteus and L. caudatus, occupy most of the range avail- able to perennial lupines. Three main popu- lation types are segregated in the following key. 1. Flowers white in populations, the eyespot and veins commonly dark brown or fading dark brown; plants mostly from Sevier Co. southward L. sericeus var. barbiger - Flowers commonly blue or blue-purple in populations, intergrading with the preceding when in contact, the eyespot yellow or brown 2 2(1). Leaflets sparingly pubescent to glabrous above; plants mostly of central southern Utah; intergrading with the following L. sericeus var. marianus Leaflets uniformly puberulent to pilose above; plants of broader distribution L. sericeus var. sericeus Var. barbiger (S. Wats.) Welsh comb, nov. based on Lupinus barbiger S. Wats., Proc. Amer. Acad. 8:528. 1873. [L. leu- canthus Rydb., type from Springdale]. Sage- brush and mountain brush communities at 1930 to 2730 m in Garfield, Kane (type 332 Great Basin Naturalist Vol. 38, No. 3 from Kane Co.), Piute, Sevier, Washington, and Wayne cos.; endemic. This is the com- mon white-flowered variety of south-central Utah; it forms intermediates at points of contact with both var. sericeus and var. ma- rianus; 29 (xiii). Var. marianus (Rydb.) Welsh stat. nov. based on Lupinus marianus Rydb. Bull. Torrey Bot. Club 34:41. 1907. [L. sericeus ssp. marianus (Rydb.) Fleak & Dunn]. Sage- brush, pinyon-juniper, mountain brush, pon- derosa pine, aspen, and spruce-fir commu- nities at 1770 to 2870 m in Garfield, Piute (type from Bullion Creek), and Sevier cos.; endemic; 8 (iv). Var. sericeus. [L. aegra-ovium C. P. Smith, type from Salina Canyon; L. huff- manii C. P. Smith, type from Salina Can- yon; L. sericeus ssp. huffmanii (C. P. Smith) Fleak & Dimn; L. larsonanus C. P. Smith, type from Salina Canyon; L. puroviridis C. P. Smith, type from Salina Canyon; L. quercus-jugi C. P. Smith, type from Salina Canyon vicinity; L. rickeri C. P. Smith, type from Salina Canyon; L. sahnensis C. P. Smith, type from Salina Canyon; L. eg- glestonianus C. P. Smith; L. flexuosus Lindl.; L. sericeus var. flexuosus (Lindl.) C. P. Smith]. Sagebrush, mountain brush, pin- yon-juniper, ponderosa pine, aspen, spruce- fir and meadow communities at 1770 to 3130 m in Beaver, Carbon, Emery, Garfield, Grand, Juab, Kane, Piute, Salt Lake, Sevier, Summit, Tooele, Utah, Wasatch, Wayne, and Weber cos.; British Columbia east to Alberta and south to California, Arizona, and New Mexico; 60 (xxv). Lupinus sparsiflorus Benth. Mohave Lu- pine. Annual, 9-38 cm tall, from a taproot; cotyledons sessile; pubescence of stems ap- pressed or ascending, of two lengths; leaves mainly cauline; petioles 1.4-8 cm long; leaf- lets 5-9, 5-35 mm long, 1-4 mm wide, elliptic-oblong to oblanceolate, flat or fold- ed, long-pilose on both surfaces, less densely hairy above; peduncles 1.3-5 cm long; ra- cemes 8- to 46-flowered, 4-23 cm long in anthesis, the axis 6-29 cm long in fruit; flowers 8.5-11 mm long, blue-purple or rarely white; pedicels 2-5 mm long; calyx gibbous at base or upper lip; banner with a central yellow spot, glabrous dorsally, re- flexed beyond the midpoint; ovules 4-6. Joshua tree, cresote bush, and mixed warm desert shrub communities at 670 to 1100 m in Washington Co.; Arizona, Nevada, Cali- fornia, and Mexico. Our material belongs to var. mohavensis (Dziekanowski & Dunn) Welsh stat. nov. based on Lupinus spar- siflorus ssp. mohavensis Dziekanowski & Dunn Aliso 6:48. 1966; 10 (ii). Lupinus volutans Greene. Rolled Lupine. Perennial, 15-40 cm tall, caespitose from a branching caudex, short-caulescent; pu- bescence of stems long pilose; leaves mainly basal; petioles 4-19 cm long; leaflets 5-7, 6-40 mm long, 2-9 mm wide, oblanceolate to elliptic, flat or folded, long pilose on both surfaces, less densely so above; pe- duncles 0.8-3 cm long; racemes 28- to 85- flowered, 6-23 cm long in anthesis, the axis 10-30 cm long in fruit; flowers 12-14 mm long, blue-purple; pedicels 1-3 mm long; calyx only somewhat gibbous at base of up- per lip; banner with a central yellow spot, glabrous dorsally, reflexed at or below the midpoint; ovules 3 or 4. Sagebrush and pin- yon-juniper communities at 1670 to 1850 m in southwestern Iron and northwestern Washington cos.; Nevada. The rolled lupine is allied to the Lupinus lepidus complex, and has been included by some authors as a synonym of phases of that group. In Utah it is a distinctive plant, with its nearest ally being L. caespitosus, from which it differs both morphologically and ecologically; 2 (i). Lupinus wyethii S. Wats. Wyeth Lupine. Perennial, 19-45 cm tall, from a branching superficial or scarcely subterranean caudex; pubescence of stem appressed-ascending, of two lengths; leaves basal and cauline; pet- ioles of lowermost leaves 3.5-13 cm long; leaflets 6-9, SViS mm long, 1.2-7 mm wide, narrowly oblanceolate, commonly folded, pilose on both surfaces; peduncles 3.5-11 cm long; racemes 13- to 49-flowered, 9-30 cm long in anthesis, the axis 10-35 cm long in fruit; flowers 10-13.5 mm long, blue- purple; pedicels 3-8 mm long; calyx gib- bous to saccate-spurred at base of upper lip; banner with a yellow eyespot, glabrous dor- sally, reflexed near the midpoint; wings glabrous dorsally, reflexed near the mid- point; wings glabrous; keel ciliate along the Sept. 1978 Welsh: Utah Flora, Fabaceae 333 upper margin near the apex; ovules 5 or 6. Wash bottoms and terraces or rimrock in pinyon-juniper and riparian communities at 1470 to 1900 m in Duchesne, Kane, and Uintah cos.; British Columbia east to Sas- katchewan and south to Washington, Idaho, and Wyoming. Our material differs from the main body of specimens considered as belonging within var. wyethii, whose range is mostly north of the Utah outliers. Speci- mens from Kane Co. differ from the north- ern materials in having fewer leaflets (6-9, not 8-13), in possessing petioles which aver- age shorter, and in more attenuate racemes. There are additional more subtle differ- ences. Possibly this population is worthy of taxonomic recognition; 12 (iv). Medicago L. Plants annual or perennial herbs, caules- cent, from a taproot or a caudex; leaves al- ternate, pinnately trifoliolate, the leaflets serrate in the distal half or less; stipules her- baceous, often toothed; flowers papilio- naceous, borne in axillary, pedunculate racemes or heads; bracts subulate; calyx 5- toothed; petals 5, yellow, white, blue, pink, lavender, or purple; stamens 10, dia- delphous; ovary enfolded by the staminal sheath, the style subulate, irritable; pods curved to spirally coiled, 1- to several-seeded, indehiscent, reticulate or spiny. 1. Flowers 2-3 mm long; inflorescence less than 10 mm long in anthesis; pods coiled through a single spiral, 1-seeded, unarmed; plants annual, prostrate to decumbent or rarely erect M. lupulina - Flowers 4-10 mm long; inflorescence longer than 10 mm long (including flower length), or pods differing from above; plants various 2 2(1). Flowers 4-5 mm long, yellow, 2-5 per raceme; racemes less than 10 mm long; pods armed with prickles, several-seeded; plants annual M. polymorpha - Flowers 6-10 mm long, yellow or blue, lavender, pink, purple, or white, 6 to many on at least some racemes; racemes longer than 10 mm; pods unarmed, several-seeded; plants usually perennial 3 3(2). Flowers yellow (sometimes tinged violet); pods merely curved; plants uncommon M. falcata Flowers blue, pink, lavender, purple, or white; pods spirally coiled; plants abundant to common M. sativa Medicago falcata L. Yellow Alfalfa. Pe- rennial (rarely fimctionally annual), 40-100 cm tall or more, the stems erect or ascend- ing, strigulose; stipules 4-12 mm long, per- sistent, conspicuously veined; leaves short- petiolate, the leaflets linear, oblong, oblan- ceolate, or elliptic, 6-20 mm long, 1-6 (10) mm wide, few-toothed, tridentate or merely apiculate apically, strigulose beneath; pe- duncles subequal to the subtending leaves or longer; racemes 6- to 20-flowered, mostly 10-20 mm long; flowers 6-9 mm long, yel- low, sometimes suffused with violet; calyx campanulate, the tube 1-2 mm long, the teeth 1.5-3 mm long, lance-subulate; pods 6-10 mm long, merely curved, unarmed, several-seeded. Sparingly cultivated forage plant, escaping and persisting; introduced from the Old World. The yellow alfalfa forms hybrids with M. sativa, and is some- times considered as a phase of that species; 2(i). Medicago lupulina L. Black Medick, Hop Clover. Annual, the stems prostrate to de- cumbent or sometimes erect, 10-40 cm long; stipules entire or nearly so, 3-6 mm long, persistent; leaves short-petiolate, the leaflets cimeate to obcordate, 4-15 mm long, 2-12 mm wide, toothed in the apical one-third (rarely more), pubescent to glabrous; peduncles mostly equaling or sur- passing the subtending leaves; racemes 6-to 25-flowered, less than 10 mm long at an- thesis (to 25 mm long in fruit); flowers 2-3 334 Great Basin Naturalist Vol. 38, No. 3 mm long, yellow; calyx campanulate, about 1 mm long; pods spiral through about 1 coil, unarmed, 1-seeded. Introduced weedy species of lawns, fields, and other sites in much of Utah (Cache, Garfield, Juab, Kane, Piute, Rich, Salt Lake, San Juan, Summit, Utah, Wasatch, Washington, and Weber COS.), where it should be considered as cos- mopolitan; introduced from Europe; 35 (v). Medicago polymorpha L. Bur Clover. (M. hispida Gaertn.). Annual, the stems pro- strate to erect, 10-40 cm long; stipules deeply divided into long teeth, mostly 3-7 mm long; leaves short-petiolate, the leaflets cimeate to obovate or obcordate, 10-25 mm long, 6-18 mm wide, toothed in the apical one-third or more, pubescent to glabrous; peduncles mostly shorter than the sub- tending leaves; racemes 2- to 5-flowered, less than 10 mm long; flowers 4-5 mm long, yellow; calyx campanulate, the tube 1-1.5 mm long, the lance-subulate teeth 1-2 mm long; pods spirally coiled, armed with spines, several-seeded. Rare weedy species of cultivated land in Utah, but to be ex- pected almost anywhere; introduced from Europe; 1(0). Medicago sativa L. Alfalfa, Lucern. Pe- rennial, or functionally annual, the stems 40-100 cm long or more, ascending to erect, finally sprawling, strigulose, stipules entire or toothed, 4-12 mm long, persistent; leaves short-petiolate, the leaflets elliptic to oblanceolate, 8-40 mm long, 2-15 mm wide, apically few-toothed, pubescent; pe- duncles often surpassing the subtending leaves; racemes 6- to 25-flowered, 10-35 mm long or more; flowers 6-10 mm long, blue, lavender, pink, purple, or white; calyx campanulate to short-cylindric, the tube 1.5-2.5 mm long, the lance-subulate teeth 2-4 mm long; pods spirally coiled, unarmed, several-seeded. Introduced forage plant, escaping and persisting, now almost or quite cosmopolitan at moderate and lower elevations (Cache, Carbon, Garfield, Kane, Salt Lake, San Juan, Sanpete, Sevier, Tooele, Utah, Washington, and Weber cos.); introduced from Europe. This is one of the most important forage plants grown in the state; 35 (iv). Melilotus L. Plants annual or biennual herbs, caules- cent, from a stout taproot; leaves alternate, pinnately trifoliolate, the leaflets dentate- serrate in the distal half or more; stipules herbaceous, distinct, subulate, entire or has- tately lobed; flowers papilionaceous, borne in axillary, pedunculate racemes; bracts sub- ulate; calyx 5-toothed; petals 5, white or yellow, the keel obtuse; stamens 10, dia- delphous; ovary enfolded by the staminal sheath, the style subulate, not irritable; pods straight, ovoid, reticulately veined or cross- ribbed, unarmed, glabrous, 1- to 2-seeded, indehiscent. 1. Flowers 2-3 mm long, yellow; pedicels less than 1 mm long; plants known from Washington Co M. indicus Flowers 3-7 mm long, white or yellow; pedicels 1-2 mm long; plants wide- spread 2 2(1). Flowers white; pods reticulately veined M- alhus Flowers yellow; pods cross-ribbed ^- officinalis Melilotus alhus Descr. ex Lam. White Sweet Clover. Annual or biennial, the stems commonly 50-150 cm tall or more, erect, strigulose; stipules entire or hastately lobed, mostly 5-10 mm long, persistent; leave short-petiolate, the leaflets obovate to ellip- tic or oblanceolate, 8-35 mm long, 1-15 mm wide, pubescent or glabrous; peduncles commonly surpassing the subtending leaves; racemes 38- to 115-flowered, 28-125 cm long or more; flowers 4-5.5 mm long, white; calyx campanulate, the tube 1.2-1.8 mm long, the teeth 1-1.5 mm long, acumi- nate; pods 2.5-6 mm long, reticulately veined, 1- to 2-seeded. Introduced forage plant, now widely established in Beaver, Sept. 1978 Welsh: Utah Flora, Fab ace ae 335 Box Elder, Cache, Garfield, Grand, Juab, Kane, Salt Lake, San Juan, Sanpete, Sevier, Tooele, Utah, Washington, and Wayne cos., and likely cosmopolitan; introduced from Europe. This plant is an excellent source of honey for domestic bees. Plants of sweet clover contain coumarin, and are respon- sible for production of bloat in livestock; 49 (xii). Melilotus indicus (L.) All. India Sweet Clover. Annual or winter annual, the stems 20-80 cm tall, erect, glabrous or strigose; stipules lance-subulate, 4-6 mm long, per- sistant; leaves short-petiolate, the leaflets obovate to oblanceolate or elliptic, 7-30 mm long, 3-14 mm wide, glabrous; pe- duncles shorter to longer than the sub- tending leaves; racemes 21- to 43-flowered, 8-30 mm long; flowers 2-3 mm long, yel- low (fading white); calyx campanulate, the tube 0.6-0.8 mm long, the teeth 0.5-0.7 mm long, triangular; pods 1.5-2 mm long, reticulately veined, 1-seeded. Adventive weedy plants of lower elevations in Wash- ington Co.; introduced from Eurasia; 3 (0). Melilotus officinalis (L.) Lam. Yellow Sweet Clover. (Trifolium melilotus offici- nalis L.). Annual, winter annual, or biennial, the stems 50-150 cm tall or more, erect, strigulose; stipules entire or with 1-3 basal teeth, 3-10 mm long, persistent; leaves shortly petiolate, the leaflets cuneate to el- liptic or oblanceolate, 8-38 mm long, 3-16 mm wide, pubescent or glabrous; peduncles shorter to longer than the subtending leaves; racemes 20- to 65-flowered, 1.8-11 (14) cm long; flowers 4.5-6 (7) mm long, yellow, fading cream; calyx campanulate, the tube 1-1.8 mm long, the teeth 1-1.5 mm long, acuminate; pods 3-5 mm long, cross-ribbed, 1- to 2-seeded. Common ruder- al weed of almost cosmopolitan distribution in Box Elder, Cache, Carbon, Daggett, Gar- field, Grand, Kane, Millard, Piute, Salt Lake, San Juan, Sanpete, Sevier, Tooele, Utah, Wasatch, Washington, Wayne, and Weber cos., and likely anywhere; in- troduced from Europe. This plant has about the same attributes and shortcomings as does M. albus, with which it often occurs; 58 (xi). Onobrychis Adans. Perennial herbs, caulescent, from a cau- dex and taproot; leaves alternate, odd- pinnate; stipules adnate to the petiole base, the lowermost amplexicaul but not connate; flowers papilionaceous, in axillary racemes, each subtended by a bract; bracteoles 2; ca- lyx 5-toothed; petals 5, red-purple to laven- der or pink, the keel much longer than the wings, abruptly bow-shaped; stamens 10, es- sentially diadelphous; ovary enclosed in the staminal sheath, the style glabrous; loment reduced to 1 segment, this armed with prickles. Onobrychis viciifolia Scop. Sainfoin, Holy Clover. [Hedysarum onobrychis L.; O. onobrychis (L.) Rydb.; O. sativa Lam.] Pe- rennial, caulescent, 20-45 cm tall, from a branching, superficial caudex; stems ascend- ing to erect; stipules 3-12 mm long, all more or less amplexicaul; leaves 3-12 mm long; leaflets 11-21 (27), 8-25 mm long, 2-7 mm wide, oblong to elliptic or oblan- ceolate, pilose mainly along veins beneath, glabrous above; peduncles 8-19 cm long; racemes 14- to 39 (50) -flowered, the flow- ers ascending-spreading at anthesis, the axis 4-15 cm long in fruit; bracts 2.5-4.5 mm long; pedicels 0.2-1.5 mm long; bracteoles 1; calyx 5.5-6.5 mm long, the tube 2.3-3 mm long, campanulate, the teeth 2.2-4 mm long, subulate; flowers 10-13 mm long, red- purple, lavender, or pink; loment sessile, as- cending, armed with prickles. Introduced forage plant, escaping and persisting in Juab, Salt Lake, Sanpete, and Utah cos., and likely elsewhere; native to Europe. The genus is closely allied to Hedysarum (q.v.); 4(i). OXYTROPIS DC. Perennial, caulescent, or acaulescent herbs, from a taproot and caudex; leaves al- ternate or basal, odd-pinnate; stipules ad- nate to the petiole base, often connate- sheathing, flowers papilionaceous, scapose or borne in axillary racemes, each sub- tended by a single bract; bracteoles 0 (rarely 2); calyx 5-toothed; petals 5, pink, pink-purple, or white, the keel shorter than the wings, the keel-tip produced into a por- 336 Great Basin Naturalist Vol. 38, No. 3 rect beak; stamens 10, diadelphous; ovary References enfolded in the staminal sheath, the style glabrous; pods sessile or stipitate, straight, Barneby, R. C. 1952. A revision of the erect, ascending, or spreading-declined. Mo- ^orth American species of Oxytwpis culed, 2-lociiled, or partially 2-loculed by ^^- ^^^^- ^^^'^- ^^'a^- ^^i. IV. intrusion of the ventral (upper) suture, de- ^i.i-ii oiz,. hiscent apically or throughout. 1. Plants shortly caulescent; stipules only somewhat adnate to the petioles; flowers 5-9 mm long; pods spreading-declined O. deflexa — Plants acaulescent, scapose; stipules adnate to petioles through half their length or more; flowers commonly more than 9 mm long; pods erect or ascending 2 2(1). Bracts, calyx teeth, pods, and sometimes other plant parts glandular-viscid .... O. viscida — Bracts, calyx teeth, pods, and other plant parts pilose to villosulous, but not at all glandular-viscid 3 3(2). Racemes 1- to 5-flowered, subcapitate 4 — Racemes 6- to many-flowered, the axis elongate, only rarely subcapitate 7 4(3). Calyx swollen at full anthesis, becoming inflated and finally enclosing the pod; pods rarely longer than 10 mm; plants of Daggett Co O. multiceps — Calyx campanulate, not turgid, not becoming inflated or enclosing the fruit, at length rupturing along one side; pods often over 10 mm long 5 5(4). Pods oblong-ellipsoid, not inflated, leathery in texture; flowers 1-4; plants of mountain summits in north central to southern Utah O. parryi — Pods ovoid, inflated, papery in texture; flower number various; plants of various distribution 6 6(5). Flowers 6-12 mm long; leaflets 7-17; plants of limestone, gravel, and basalt in southern Utah O. oreophila — Flowers (11) 14-17 mm long; leaflets 3-7; plants of shale and limestone in Uintah, Emery, and Garfield cos O. jonesii 7(6) Plants dwarf, seldom over 10 cm tall; flowers 6-12 mm long; growing on limestone, gravel, and basalt in southern Utah O. oreophila — Plants mainly over 10 cm tall; flowers 14-27 mm long; growing on various substrates and with distribution various but seldom as above 8 8(7). Petals bright pink-purple 9 — Petals white or ochroleucous to yellowish 10 9(8). Pubescence of basifixed hairs; calyx somewhat swollen and accrescent; plants of Daggett Co O. besseyi — Pubescence of malphighian hairs; calyx not swollen or enlarging; plants of wide distribution O. lambertii 10(8). Wing petals dilated apically, at least 5 mm wide; plants of broad distribution O. sericea — Wing petals not especially dilated, less than 5 mm wide, plants known from the north slope of the Uinta Mts O. campestris Sept. 1978 Welsh: Utah Flora, Fab ace ae 337 Oxytropis besseyi (Rydb.) Blankinship. Bessey locoweed. [Aragalhis besseyi Rydb.] Caespitose, acaulescent, 8-28 cm tall; pu- bescence basifixed, silky-pilose to sub- tomentose; stipules pilose-tomentose; leaves 6-20 cm long; leaflets 11-13 (25), 6-25 mm long, 2-6 mm wide, lanceolate to lance- oblong or elliptic, silky-pilose; scapes 6-19 cm long, subtomentose; racemes 8- to 22- flowered, the flowers spreading-ascending at maturity, the axis 2.5-8 cm long in fruit; bracts silky-pilose; flowers 18-25 mm long, brilliant pink-purple; calyx 10-13 mm long, the tube swollen at anthesis, 7.5-9.5 mm long, the teeth 2.5-4 mm long, lance sub- ulate; pods sessile or nearly so, strongly in- flated, the body ovoid, 5-8 mm thick, semi- bilocular, densely villous. Pinyon-juniper community at 1830 to 2130 m in Daggett Co.; Montana, Idaho, Wyoming, and Colo- rado. Our material is assignable to var. oh- napiformis (C. L. Porter) Welsh comb, no v. based on Oxytropis obnapiformis C. L. Por- ter, Madrono 9:133. 1947. Maintenance of O. obnapiformis at specific rank was dis- cussed by Barneby (1952), who noted that the features used as diagnostic ones "are not strong characters." That information along with the apparent intermediacy of Utah specimens with those of var. fallax Barneby, from nearby in Wyoming, dictates that the reduction of O. obnapiformis is necessary. In specimens from Utah, the leaflet number is fewer than in more typical plants from northwestern Colorado, but the pubescence and pod features seem to substantiate the recognition of the taxon at varietal rank; 5 (0). Oxytropis campestris (L.) DC. Yellow Locoweed. [Astragalus campestris L.]. Caespitose, acaulescent, 4-18 cm tall; pu- bescence basifixed, mainly pilose; stipules glabrous or sparingly pilose; leaves 1.5-12 cm long; leaflets 7-17, 3-23 mm long, 1-6 mm wide, oblong to lanceolate or obovate; scapes 2.5-15 cm long, pilose to glabrate; flowers 14-20 mm long, ochroleucous; calyx 7-10 mm long, the tube cylindric, the teeth 1.5-3 mm long, triangular-subulate, pods 8-18 mm long, erect, sessile or subsessile, pilose, partially bilocular by intrusion of the ventral suture. Meadows and open woods at 2270 to 2600 m in Summit Co. (Goodmans Ranch, Bear River, E. & L. Payson 4868 POM); Oregon, Washington, Idaho, Mon- tana, and Colorado. The range as given is for our materials only, which belong to var. cusickii (Greenm.) Barneby (O. cusickii Greenm.). The species proper is circum- boreal; 1 (0). Oxytropis deflexa (Pallas) DC. Stemmed Oxytrope. Shortly caulescent to sub- acaulescent, (5) 7-48 cm tall; pubescence basifixed, villous-pilose; leaves 2-20 cm long; leaflets 23-41, 3-24 mm long, 1-7 mm wide, lance-oblong to lanceolate, pilose on both surfaces, quite sessile; peduncles 3.5-32 cm long, villous-pilose; racemes 3-to 25-flowered, the flowers ascending to de- clined at anthesis, the axis 3.5-10 cm long in fruit; bracts pilose; flowers 5-10.5 mm long, whitish, lilac, or blue-purple; calyx 4-8 mm long, the tube 2-3.5 (4.2) mm long, campanulate, the teeth 1.5-5 mm long, lance-subulate; pods spreading-declined, sub- sessile to shortly stipitate, the body oblong to ellipsoid, 8-18 mm long, 3-4.5 mm wide, subunilocular, pilosulous. Two rather dis- tinctive varieties occur in Utah. 1. Plants subacaulescent; flowers 9-10.5 mm long, blue-purple; racemes 15-22 mm broad at anthesis; of high elevations in Uinta Mts O. deflexa var. deflexa - Plants commonly short caulescent; flowers 5-9 mm long, whitish, lilac, or blue-purple; racemes commonly less than 17 mm wide at anthesis; of moderate elevations, widespread O. deflexa var. sencea Var. deflexa. [Astragalus deflexus Pallas], rado and north to Yukon; Asia. The North Meadows at 2750 to 3350 m in Duchesne American material here assigned to var. def- and Summit (Goodrich 824, BRY) cos; Colo- lexa might well represent a distinct taxon. 338 Great Basin Naturalist Vol. 38, No. 3 worthy of recognition at varietal level; 2 (0). Var. sericea Torr. & Gray. Moist mead- ows, streambanks, and gravel bars in aspen, mixed conifer, and sagebrush communities at 2430 to 3050 m in Emery, Garfield, Grand, Iron, Summit, Uintah (?), and Wayne cos; Alaska and Yukon south to Cal- ifornia, New Mexico, and North Dakota; 21 VI Oxytropsis jonesii Bameby. Jones Oxyt- rope. Acaulescent, densely pulvinate-caespi- tose; pubescence basifixed, villous-pilose; leaves 0.7-3 cm long; leaflets 1-7, 2-11 mm long, 1-3.5 mm wide, lanceolate to lance- oblong, pilose to strigose on both surfaces, quite sessile; scapes 0-3.5 cm long, villous- pilose; racemes 1- to 5-flowered, the flowers ascending at anthesis, the axis not elongat- ing in fruit; bracts pilose; flowers (11) 14-16.5 mm long, pink-purple; calyx 7.5-9.5 mm long, the tube shortly cylindric, 4.5-7 mm long, the teeth 1.5-3 mm long, triangular to subulate; pods sessile or sub- sessile, erect, bladdery-inflated, 14-25 mm long 8-13 mm wide (when pressed), semi- bilocular, villous-pilose. Ponderosa pine, western bristlecone pine, and mixed desert shrub communities, on Flagstaff Limestone, Pink Limestone member of Wasatch Forma- tion, and on Green River Shale Formation, at 1930 to 2430 m in Emery, Garfield (type from Red Canyon), and Uintah cos.; endem- ic. The Jones oxytrope is closely allied to O. oreophila with which it is sympatric at the type locality; 16 (v). Oxytropis lamhertii Pursh. Lambert Lo- coweed. [O. lamhertii var. bigelovii Gray; Aragallus bigelovii (Gray) Greene; Astra- galus lamhertii var. higelovii (Gray) Tidestr.; Aragallus metcalfei Greene; Aragallus knowltonii Greene; Aragallus patens Rydb.; O. patens (Rydb.) A. Nels.; O. hilocularis A. Nels.]. Caespitose, acaulescent, (10) 14-50 cm tall; pubescence malpighian, strigose; stipules pilose; leaves 3-24 cm long; leaflets 7-13, 7-45 mm long, 2-8 mm wide, lan- ceolate to oblong or linear; scapes 4-28 cm long, strigose; racemes 8- to 40-flowered, the flowers ascending at anthesis, the axis 3-23 cm long in fruit; bracts strigose; flow- ers 17-25 mm long, pink-purple; calyx 6.5-10 mm long, the cylindric tube 4.5-7.5 mm long, the teeth 1.5-4.5 mm long, sub- ulate; pods sessile or shortly stipitate, erect or ascending at maturity, cylindroid to lance-acuminate in outline, the body 15-27 mm long, 2.5-6 mm thick, bilocular, stri- gose to strigulose. Mixed desert shrub, pinyon-juniper, sagebrush, and grass com- munities at 1230 to 2730 m in Carbon, Duchesne, Emery, Garfield, Kane, San Juan, Sanpete, Sevier, Wasatch, and Wayne cos.; the species from Saskatchewan and Mani- toba south to Arizona, New Mexico, and Texas. Our material belongs to var. hige- lovii A. Gray, which occurs from Wyoming south to Arizona and New Mexico. In other places where O. lamhertii contacts O. se- ricea, hybrid swarms of great variability and beauty occur. No such extensive hybrid populations are known for Utah, possibly becauuse the points of contact between these species are few or non-existent; 51 (xii). Oxytropis multiceps Torr. & Gray. Rocky Mountain Oxytrope. [Spiesia multiceps (Torr. & Gray) Kuntze; Aragallus multiceps (Torr. & Gray) Heller; Astragalus hisontum Tidestr.; O. multiceps var. minor A. Gray; Aragallus multiceps var. minor (A. Gray) A. Nels.; Aragallus minor (A. Gray) Cockerell ex Daniels; O. minor (A. Gray) Cockerell; Astragalus hisontum var. minor (A. Gray) Tidestr.]. Caespitose, acaulescent, pulvinate; pubescence basifixed, silky-pilose; stipules amplexicaul but distinct, silky-pilose; leaves 1-5 cm long; leaflets 5-9, 3-13 mm long, 1-4 mm wide, lanceolate to elliptic, oblong, or oblanceolate, silky-pilose; scape 1-4 cm long, long-villous; racemes 1- to 4- flowered, the flowers ascending-spreading at anthesis, the axis to 0.5 cm long in fruit; bracts thinly pilose; flowers 17-24 mm long, bright pink-purple; calyx 7-13 (20) mm long, the tube swollen at anthesis, becoming bladdery-inflated and investing the fruit at maturity, 5.5-10 mm long (8-18 mm in fniit), the teeth 2-3 mm long, triangular- subulate; pods included within the swollen calyx, stipitate, the stipe 0.5-1.5 mm long, the ovoid-ellipsoid body 6-10 mm long, 3-5 mm wide, subunilocular, short-villous. Pinyon-juniper and ponderosa pine commu- Sept. 1978 Welsh: Utah Flora, Fabaceae 339 nities at 1830 to 2270 m in Daggett Co.; Wyoming, Colorado, and Nebraska; 2 (0). Oxytropis oreophila A. Gray. Mountain Oxytrope. Caespitose, acaulescent, often densely pulvinate, 1-14 (23) cm tall; pu- bescence basifixed, silky-pilose; stipules silky-pilose; leaves 0.5-8.5 cm long; leaflets (5) 7-17, 1-15 mm long, 0.5-4 mm wide, lanceolate, elliptic, oval, or ovate, pilose; scapes 0-12 (21) cm long, pilose to hirsute; racemes (1) 2- to 12-flowered, the flowers ascending-spreading at anthesis, the axis to 1 cm long in fruit; bracts pilose; flowers 6-12.5 mm long, pink to pink-purple or white; calyx 4.5-8 mm long, the tube 3.2-5.5 mm long, campanulate to short- cylindric, the teeth 1.3-2.5 mm long, sub- ulate; pods sessile, bladdery-inflated, (7) 9-17 mm long, 5-14 mm wide (when pressed), subunilocular, hirsutulous to vil- lous. Two more or less distinctive, but only arbitrarily separable varieties are present in Utah. 1. Plants densely pulvinate-caespitose; leaves 0.5-2 cm long, the leaflets mostly 2-5 mm long; pods 10 mm long or less, and less than 6 mm wide; flowers 9.5 mm long or less O. orephila var. iuniperina Plants densely to loosely pulvinate-caespitose; leaves often more than 2 cm long, the leaflets commonly more than 5 mm long; pods 9-17 mm long, more than 6 mm wide; flowers commonly 10-12.5 mm long O. oreophila var. oreophila Var. juniperina Welsh var. nov. Oxytropis oreophila var. oreophila aemulans sed differt dense caespitosi-pulvinatis, foliis brevissimis, leguminibus brevissimis et angustis, et flo- ribus brevissimis. Holotype: Utah, Wayne Co., ca 1 mile east of Bicknell, at 2200 m on Carmel formation, in pinyon-juniper woodland, Welsh & Moore 13828, 30 June 1976 (BRY). Paratypes; Utah, Wayne Co., do, Welsh & Moore 13831, 30 June 1976 (BRY); do, Atwood 1863, 18 June 1969 (BRY). Material of the Juniperina phase has long been recognized. Gray (Proc. Amer. Acad. 20:3, 1884) alluded to a specimen by Ward (574 GH, US), from Rabbit Valley (Loa-Bicknell vicinity), Utah, as a possible variety of O. oreophila or a related species "with flowers immersed in tufts of foliage." It is a xeric phase of O. oreophila, which passes by degree in to var. oreophila. Other xeric phases fail to demonstrate the mor- phological features of these striking plants, and the trend seems worth recognition from a taxonomic standpoint; 5 (iii). Var. oreophila. [Spiesia oreophila (Gray) Kuntze; Aragallus oreophiliis (Gray) A. Nels.; Astragalus oreophilus (Gray) Tidestr.; Astragalus munzii Wheeler.] Alpine tundra, ridge tops, meadows, spruce-fir, ponderosa pine, pinyon-juniper, and less commonly in mixed desert shrub communities, on lime- stone, volcanic gravels, beach gravels, and sands, at 1700 to 3350 m in Beaver, Gar- field, Iron, Kane, Piute, Sanpete, and Wayne cos.; California, Nevada, Arizona, and New Mexico; 67 (xiv). Oxytropis parryi A. Gray. Parry Oxyt- rope. [Spiesia parnji (A. Gray) Kuntze; Ara- gallus parryi (A. Gray) Greene; Astragalus parryanus Tidestr., not A. parryi A. Gray.] Caespitose, acaulescent, 2-11 cm tall; pu- bescence basifixed, silky-pilose; leavse 1.5-7 cm long; leaflet 7-17, 2-9 (12) mm long, 0.8-3 mm wide, oblong to elliptic or lan- ceolate, pilose; scapes 1.2-8 (10) cm long, pilose; racemes 1- to 3 (4) -flowered, the flowers erect or ascending, the axis 0.5-1 cm long in fruit; bracts pilose; flowers 7.5-12 mm long, pink-purple; calyx 5-8 mm long, the tube 3-5.5 mm long, cam- panulate to short-cylindric, the teeth 1.5-2.5 mm long, triangular-subulate; pods erect, sessile, oblong to lance-ovoid, 13-22 mm long, 4-8 mm thick, bilocular or nearly so, pilosulous. Alpine timdra, ridge tops, and meadows at 2700 to 3770 m in Beaver, Car- bon, Garfield, Grand, Juab, Piute, San Juan, and Wasatch cos.; Idaho, Wyoming, Califor- nia, Colorado, and New Mexico. The Parry oxytrope is difficult to distinguish in flower from O. oreophila, especially from speci- mens of that species with fewer than five 340 Great Basin Naturalist Vol. 38, No. 3 flowers. The fruit is definitive; 11 (v). Oxytropis sericea Nutt. in Terr. & Gray. Silky or White Locoweed. [O. lamhertii var. sericea (Nutt.) A. Gray; Spiesia lambertii var. sericea (Nutt.) Rydb.; AragaUiis lam- bertii var. senceus (Nutt.) A. Nels.; Ara- galltis majusculus Greene, type from the Mt. Ellen Henry Mts.] Caespitose, acaules- cent, 13-32 cm tall, pubescence basifixed, silky-pilose; stipules pilose to subtomentose; leaves 3.5-21 cm long; leaflets 9-23, 4-32 (40) mm long, 1.5-10 mm wide, lanceolate to oblong elliptic, or ovate, pilose; scapes 7-26 cm long, pilose; racemes 6- to 27- flowered, the flowers ascending to spread- ing, the axis 1.5-12 cm long in fruit; bracts pilose; flowers 15-26 mm long, white or tinged with purple; calyx 8-12 mm long, the tube cylindric, the teeth triangular to subulate; pods erect, sessile, the body sub- cylindric to ovoid-oblong, 10-25 mm long, 4-7.5 mm thick, bilocular or nearly so, stri- gose or pilosulous. Sagebrush, pinyon- juniper, and grassland (rarely mixed desert shrub) communities at 1670 to 3350 m in Box Elder, Carbon, Daggett, Duchesne, Emery, Garfield, Kane, Piute, Summit, and Wayne cos.; the species from the Yukon southward to Nevada, New Mexico, and Oklahoma. Our materials belong to var. se- ricea, which occurs from Montana, Idaho, and Souta Dakota, south to Nevada, New Mexico, and Oklahoma; 38 (vii). Oxytropis viscida Nutt. Viscid Locoweed. [O. campestris var. viscida (Nutt.) S. Wats.; Spiesia viscida (Nutt.) Tidestr.] Caespitose, acaulescent, 4-18 cm tall; pubescence basi- fixed, spreading-hairy; stipules glabrous dor- sally, commonly prominently glandular; leaves 2-17 cm long; leaflets 19-39 or more, 1.5-20 mm long, 1-5 mm wide, ob- long to lanceolate or elliptic, glabrate to glabrous on both sides, sometimes glandular; scapes 2-12.5 cm long, spreading-hairy; racemes 3- to 20-flowered, the flowers spreading-ascending, the axis 2-7 cm long in fruit; bracts glandular; flowers 11-19 mm long, whitish, lilac, or pink-purple; calyx 5-11 mm long, the shortly cylindric tube 4-7 mm long, the teeth 1.5-3.5 mm long, triangular-subulate, commonly glandular; pods erect, sessile, ovoid to subcylindric. 8-20 mm long, 4-5 mm thick, bilocular, glandular. Montane meadows, shniblands, and open woods at 2430 to 3050 m in Emery, Salt Lake, Sanpete, Sevier, Utah, and Wayne cos.; Alaska east to Gaspe and south to California, Nevada, Colorado, and Minnesota. Our material belongs to var. vis- cida. This is a portion of a circumboreal complex with great variation. Utah mate- rials demonstrate two of the extreme phases of that variety. Naming of the phases within the species would lead to an endless en- tanglement. Thus, the whitish flowered plants from Emery, Sanpete, and Sevier COS., are considered as a minor element in this evidently polygenetic and certainly polymorphic series; 14 (iv). Parryella Torr. & Gray ex A. Gray Unarmed shrubs; leaves alternate, pinnate, with numerous leaflets, glandular-dotted; stipules subulate, caducous; peduncles oppo- site the leaves; flowers in loose spicate ra- cemes or panicles; calyx turbinate-camp- anulate, 10-ribbed near the base, 5-lobed; petals lacking; stamens 10, the filaments dis- tinct, inserted on the hypanthium; pods 1- seeded (2-ovuled), indehiscent, obliquely obovoid, glandular-dotted, exserted from the calyx. References Rydberg, p. a. 1919. Parryella. pp. 25-26. In: Rydberg, P. A. Fabaceae-Psoraleae. N. Amer. Fl. 24:1-64. Parryella filifolia Torr. & Gray ex A. Gray. Narrow-leaf Dunebroom. Shrubs to 15 cm tall or more, often partially buried in sand, the branchlets strigose and with mam- miform glandular protuberances; stipules 1-2 mm long, chestnut-brown, fragile; leaves 3.5-13 cm long; leaflets 8-40, 1-21 mm long, linear, all involute, strigose to glabrate and glandular on the visible sur- face; pedimcles 0-1 cm long; racemes 4-10 cm long, the main ones often branched near the base, forming panicles, 35- to 90- flowered, the axis not much elongating in fruit; bracts reduced to gland-tipped ves- Sept. 1978 Welsh: Utah Flora, Fab ace ae 341 tiges; calyx 2.6-3.1 mm long, 10-ribbed near the base, the tube opaque, the teeth 0.2-0.5 mm long, ciliate; flowers 5-6.5 mm long in anthesis, the stamens long-exserted; pods short-stipitate, the body 5-6.5 mm long, 2.5-3 mm wide, the pilose style base per- sistent, glabrous, glandular-punctate. Stabi- lized dime sands at 1470 to 1570 m astride the Grand-San Juan co. line, and previously known from along the San Juan River near the present upper reaches of Lake Powell; New Mexico and Arizona; 5 (iv). Peteria a. Gray Perennial, caulescent, from a subterranean caudex arising from deep-seated tuberous roots; leaves alternate, odd-pinnate; stipules only slightly adnate to the petiole base, spinescent; flowers papilionaceous, borne in terminal racemes, each subtended by a bract, or the lowermost flower sometimes axillary; bracteoles 0; calyx 5-toothed; petals 5, whitish to ochroleucous, rarely pinkish; stamens 10, diadelphous; style hairy at apex; pods narrowly oblong, straight, few- to several-seeded, laterally compressed, dehis- cent, the valves coiling upon dehiscence. Peteria thompsonae S. Wats. Thompson Peteria. [P. nevadensis Tidestr.]. Perennial, 11-48 cm tall; pubescence basifixed, of flat- tened appressed hairs; stems erect to sprawling; stipular spines 1.8-6.5 mm long; leaves 4-18 cm long; leaflets 9-27, 6-17 mm long, 3-11 mm wide, elliptic to ovate, oblong to oval, strigose on both sides or gla- brate above; peduncles 0-4 cm long; ra- cemes 5- to 44-flowered, the flowers as- cending at anthesis, the axis 6-18 cm long in fruit; bracts 5-9 mm long, stipitate- glandular; pedicels 1-5 mm long, strigulose and stipitate-glandular; calyx 11-16.2 mm long, the tube 6.5-8 mm long, short-cylin- dric, stipitate-glandular, the teeth 3.8-9 mm long, lance-subulate; flowers 18-22 mm long, whitish to ochroleucous, rarely pink- ish; pods descending, stipitate, the stipe to 11 mm long, the body 48-55 (70) mm long, 5-7 mm wide, glabrous, more or less con- stricted between the seeds. Pinyon-juniper and mixed desert shrub communities at 1230 to 1870 m in Emery, Grand, Juab, Kane, San Juan, and Washington co.; Arizona, Nevada, and Idaho. The type specimen was collect- ed by Mrs. Ellen Thompson, sister of John Wesley Powell, at Kanab, where she lived in 1872. This is a singular and striking plant; 16 (v). Phaseolus L. Annual herbs, from a taproot; leaves al- ternate, pinnately trifoliolate; stipules her- baceous, distinct; flowers papilionaceous, ax- illary or in axillary racemes, each subtended by a bract; bracteoles 2, attached at base of calyx; calyx 5-toothed; petals 5, pink to purplish or white; stamens 10, diadelphous; ovary few- to several-ovuled, the style twisted or coiled in the keel, bearded to- wards the apex, the stigma oblique; pods linear to oblong, laterally flattened to sub- terete, the valves coiling upon dehiscence. Phaseolus vulgaris L. Kidney Bean. An- nual, 30-200 cm tall or more, clump- forming or twining and vine-like; pu- bescence basifixed, villosulous or finally gla- brate; petioles 4-20 cm long; leaflets 3, stipelate, commonly 40-100 mm long and 20-80 mm wide, ovate to lanceolate, pilo- sulous to villosulous on both sides, mainly along the veins; peduncles 0-2 cm long; racemes few-flowered; flowers 12-16 mm long; bracteoles ovate-lanceolate, prominen- tly veined; pods slender, subterete to flat- tened, most 6-15 cm long. This is the table bean of commerce. It is cultivated widely, escaping commonly, but hardly persisting; native of the New World (?). Additional species are present in the cultivated flora of the state, but the extent is unknown, and they are here excluded. Among them are the scarlet runner bean {P. coccineus L.) and the lima bean (P. limensis Macf.); 2 (i). PiSUM L. Plants herbaceous, annual, from a tap- root; stipules prominent, larger than the leaflets, semi-sagittate to ovate or reniform; stems clambering, not winged; leaves alter- nate, even-pinnate, the terminal extension of the rachis forming prehensile tendrils; ra- cemes axillary, pedunculate, 1- to few- 342 Great Basin Naturalist Vol. 38, No. 3 flowered; stamens 10, diadelphous; style lat- erally compressed, bearded along the ven- tral edge; pods 1-loculed, oblong in outline, several-seeded, the valves coiling upon de- hiscence. Pisum sativum L. Garden Pea. Plants 20-200 cm long or more, sprawling or clambering, the stems merely angled, glabrous; stipules foliaceous, larger than the leaflets; leaflets 4-6, elliptic to ovate to oblong-lanceolate, 9-60 mm long, 6-40 mm wide, glabrous; tendrils with 1-3 pairs of lateral branches, prehensile; flowers 1-3 per raceme, white, red, or bicolored, 18-30 mm long; calyx 12-18 mm long, the teeth long- er than the tube; pods commonly 4-10 cm long, glabrous. Cultivated pea of commerce, widely grown, escaping but not persisting; introduced from Eurasia. Several horticul- tural forms are grown, mostly with white flowers. 2 (i). Prosopis L. Armed shrubs or small trees; leaves alter- nate, bipinnate, with an obscure gland be- tween the lower pair of pinnae; leaflets sev- eral to numerous; stipules small, or modified as spines; flowers perfect, borne in spikelike racemes, yellowish to ochroleucous; calyx 5- toothed; corolla regular, the 5 petals dis- tinct or nearly so; stamens 10, distinct, ex- serted, the anthers terminally glandular; ovary pubescent, sessile or stipitate, the stigma concave; pods indehiscent, narrowly oblong and more or less constricted be- tween the seeds or spirally coiled. References Johnston, M. D. 1962. The North Ameri- can Mesquites, Prosopis Sect. Alga- robia (Leguminosae). Brittonia 4:72-90. Leaflets 5-8 pairs, mostly less than 10 mm long; pods coiled spirally into a woody cylinder P. pubescens Leaflets 10-18 pairs or more, often exceeding 10 mm long; pods narrowly oblong, straight or nearly so P. glandulosa Prosopis glandulosa Torr. Mesquite. [P. juliflora authors, not (Sw.) DC.; P. chilensis authors, not (Mol.) Stuntz]. Armed shrubs or broad-crowned trees, commonly 3-5 m tall; leaves petiolate, the rachis produced as a spine; pinnae 2; leaflets 7-17 pairs, oblong to narrowly oblong, 7-38 mm long, 1-4 mm wide, glabrous on both sides, sometimes cil- iate; spines nodal but not stipular, single or paired, 3-35 mm long or more; stipules in- conspicuous, subulate; flowers in yellowish to greenish spikes, clustered from spur branches, ascending to declined; ovary pi- lose; pods stipitate, the stipe 5-8 mm long, the body narrowly oblong to linear, often curved, subterete to somewhat flattened, mostly 100 to 200 mm long, a bony endo- carp around each seed. Terraces and bars at 670 to 1170 m in Washington Co.; the spe- cies from California, Nevada, Arizona, New Mexico, Kansas, Oklahoma, Texas, and southward. Our specimens belong to var. torreyana (Benson) M. C. Johnston (P. juli- flora var. torreyana Benson), with distribu- tion from California, Arizona, New Mexico, and Texas southward. In the herbarium of the University of Utah (UT) there is a speci- men of mesquite labeled as collected in a vacant lot in North Salt Lake; whether ac- cidental or cultivated, is not known; 17 (i). Prosopis pubescens Benth. Screwbean. [P. odorata Torr. & Frem. in Frem.; Strombo- carpa pubescens (Benth.) A. Gray; S. odo- rata A. Gray.] Armed shrubs or small trees with slender branches, commonly 2.5-3.5 m tall; leaves shortly petiolate, the rachis pro- duced as a spine; pinnae 2 (rarely 4); leaf- lets 5-8 pairs, elliptic to oblong, puberulent to glabrate; spines paired at nodes, appar- ently stipular, mostly 5-25 mm long; flow- ers in clusters or solitary, yellowish spikes; ovary villous; pods tightly coiled into a woody cylinder, 30-50 mm long, 4-6 mm thick. Benchlands, slopes, and terraces along drainages at 730 to 900 m in Washington Co.; California, Nevada, Arizona, New Mex- ico, Texas, and Mexico; 9 (i). Sept. 1978 Welsh: Utah Flora, Fabaceae 343 PsoRALEA L. References Perennial herbs, unanned, from rhizomes Matthews, E. H. 1969. A preliminary revi- or tuberous roots; leaves alternate, pal- sion of the genus Psoralea in Utah, mately compound, the leaflets 3-5, glandu- Unpublished Mss. Brigham Young Uni- lar-dotted; stipules triangular to subulate; versity. 42 pp. flowers in axillary racemes or spike-like ra- Ockendon, D. J. 1965. A taxonomic study cemes; calyx subcylindric to campanulate, of Psoralea subgenus Pediomelum (Le- 5-lobed; corolla papilionaceous; stamens 10, guminosae). Southwestern Nat. diadelphous (rarely all connate); petals 10:81-124. bluish or purplish to lavender or white; Toft, C. A. and S. L. Welsh. 1972. A re- pods 1-seeded, indehiscent or irregularly so, vision of the Psoralea lanceolata com- included in the calyx or slightly exceeding plex (Leguminosae). Great Basin Nat. it. 32:76-87. 1. Plants strongly caulescent, commonly 25-100 cm tall, arising from rhizomes; leaves mainly 3-foliolate (except in P. tenui flora); leaflets narrowly obovate to oblanceolate or linear; bracts of inflorescence inconspicuous 2 — Plants subacaulescent to short-caulescent, mostly shorter than 25 cm, arising from tuberous roots or thickened rhizomes; leaves mainly 5-foliolate (except in P. pariensis); leaflets obovate to sub-orbicular; bracts of inflorescence conspicuous 4 2(1). Peduncles 15-40 cm long or more; leaves few, mostly deciduous by an thesis, the leaflets sharply acuminate; plants of southeastern Utah P. jiincea — Peduncles mainly shorter than 15 cm; leaves numerous, persisting through the season, the leaflets obtuse to rounded or cuspidate; distribution various 3 3(2). Plants commonly 40-100 cm tall, at least some leaves 4- to 5-foliolate; flowers mainly indigo; plants of western Kane, Washington, and Iron cos P. teniiiflora — Plants often less than 40 cm tall, all leaves commonly 3-foliolate; flowers white to purple; plants widespread in Utah P. lanceolata 4(1). Plants definitely caulescent, or if acaulescent (as occasionally in P. epipsila), then leaflets glabrous or glabrate above 5 — Plants acaulescent to short-caulescent; leaflets definitely pubescent above 6 5(4). Leaflets conspicuously bicolored, cinerous beneath, bright green and glabrous to glabrate above (except along some veins); plants known only from southern Kane Co. and adjacent Arizona P- epipsila — Leaflets dull-green beneath, green above, pubescent as above; plants of central eastern Utah P. arornatica 6(4). Leaflets commonly 3, strongly white strigose along the veins above; plants of the Paunsagaunt and Paria regions P. pariensis — Leaflets commonly 5, uniformly strigose above, or glabrous, or thinly strigose along the veins; plants of various distribution 7 7(6). Petioles and peduncles appressed or ascending pubescent; plants rather wide- spread in eastern Utah P- megalantha — Petioles and peduncles retrorsely hairy; plants of Washington Co P. mephitica 344 Great Basin Naturalist Vol. 38, No. 3 Psoralea aromatica Payson. Paradox Breadroot. [P. rafaelensis M. E. Jones, type from LaSal Mts.; P. rafaelensis var. magna M. E. Jones, type from the San Rafael Swell.] Caulescent, 8-15 (20) cm tall, from slender subterranean caudex branches aris- ing from deep-seated tuberous roots; stems with 2-4 (5) elongated internodes, strigose to strigulose; leaves mainly 5-foliolate; pet- ioles 1.2-8 cm long, pubescent like the stems; leaflets 6-26 mm long, 3-16 mm wide, cimeate-obovate, gray-green, strigu- lose, and punctate beneath, yellow-green, punctate, and strigose overall or only along the veins above; stipules scarious, 2-9 mm long; peduncles 0.2-0.5 cm long; racemes 3- to 7- flowered, 1-2 cm long; pedicels 1-2.5 mm long; bracts lanceolate, 4-7 mm long; flowers 10-13 mm long, the banner, wings, and keel more or less suffused with purple; calyx 10-11 mm long, the tube 3-4 mm long, strongly gibbous, the lower tooth 5-7 mm long, about twice as broad as the lower lateral ones; pods to 15 mm long. Pinyon- juniper woodland and mixed desert shrub communities at about 1530 m in Emery and Grand cos.; Paradox Basin, Colorado, a middle Navajo Basin endemic; 8 (2). Psoralea epipsila Bameby. Kane Bread- root. Subacaulescent or short-caulescent, 3.5-10.5 cm tall, from slender subterranean caudex branches arising from deep-seated tuberous roots; stems with 2 or 3 elongated internodes, strigose to ascending hairy; leaves mainly 5-foliolate; petioles 0.8-5 cm long, pubescent like the stems; leaflets 5, 8-25 mm long, 3-15 mm wide, obovate, grayish, strigulose, and punctate beneath, bright yellow-green, punctate and glabrous to thinly strigose above (especially along the veins); peduncles to 5 cm long; racemes 2-4 cm long; pedicels about 3 mm long; bracts broadly lanceolate, 10-13 mm long; flowers 11-14 mm long, the banner, wings, and keel pale violet; calyx 11-14 mm long, the tube 5-6 mm long, strongly gibbous, the lower tooth about 8 mm long, the upper ones shorter and narrower; pods 1-seeded. Pinyon-jimiper woodland at about 1670 m on Chinle and Moenkopi formations in Kane Co., and in adjacent Arizona; 2 (0). The Kane breadroot is still imperfectly un- derstood, even though the first collection was taken by M. E. Jones in 1890. Many more collections are necessary before ade- quate descriptions and relationships can be drawn. This is a Mohave Strip endemic; 2 (0). Psoralea juncea Eastw. Rush Scurfpea. [Psoralidium funceum (Eastw.) Rydb.] Cau- lescent, 48-90 cm tall, from a rhizome; stems with 5 or more elongated internodes, strigose; leaves commonly 3-foliolate, often deciduous by anthesis; petioles 1.4-7 cm long, pubescent like the stems; leaflets 3, 19-44 mm long, 3-7 mm wide, oblanceolate to eliptic, acuminate apically, strigose and glandular on both surfaces, greenish; stipules acuminate, strigose; peduncles (8) 11-48 cm long; racemes 7- to 20-flowered, 5-11 cm long; bracts lance-acuminate, glabrous dor- sally, 1.5-2.5 mm long, deciduous; flowers 4.2-5.8 mm long, the petals indigo; calyx 2.7-3.4 mm long, the tube 1.3-2.5 mm long, campanulate, not especially gibbous, the lower tooth 0.7-1.4 mm long, longer than the others; pods 1-seeded, densely silky- villous. Stabilized dunes and other sandy sites at 1200 to 1370 m in Garfield, Kane, and San Juan cos.; Coconino Co., Arizona; a Glen Canyon-San Juan endemic; 21 (iv). Psoralea lanceolata Pursh. Dune Scurfpea. Caulescent, 15-68 cm tall, from a rhizome, clump-forming; stems with 5 or more elongated internodes, glabrous or stri- gose; leaves commonly 3-foliolate, persistent at flowering; petioles 0.8-3 cm long, stri- gose to glabrate; leaflets 3, 14-50 mm long, 0.5-9 mm wide, oblanceolate below becom- ing linear upwards, obtuse to acute or cus- pidate, sparingly strigose on both sides or glabrous above, yellow-green; stipules lance- attenuate, 3-16 mm long; peduncles 3.3-24 cm long; racemes 5- to 41-flowered, 1.2-17 cm long; bracts ovate to elliptic or lanceo- late, glabrous dorsally, 1.3-2.8 mm long, persistent, flowers 4.8-6.3 mm long, blue, white, or bicolored; calyx 1.6-2.8 mm long, the tube 1.3-2 mm long, campanulate, not especially gibbous, the lower tooth 0.2-0.8 mm long, not much larger than the others; pods 1-seeded, conspicuously glandular. Three rather distinctive but apparently in- tergrading varieties are present in Utah. Sept. 1978 Welsh: Utah Flora, Fab ace ae 345 1. Racemes lax, 25-170 mm long at anthesis, the flower nodes widely separated, often with 2 or 3 flowers per node; blue flower color predominating; plants of southeastern Utah P. lanceolata var. stenophylla Racemes compact to moderately lax, 12-55 mm long at anthesis, the flower nodes seldom widely separated or mainly with more than 3 flowers per node; blue and white flowers variably abundant; plants of various distribution 2 2(1). Racemes (12) 27-55 mm long, with (17) 27-41 flowers; plants of the Great Basin P. lanceolata var. stenostachys Racemes 10-25 (28) mm long, with 5-24 flowers; plants of the Colorado Basin P. lanceolata var. lanceolata Var. lanceolata. Plains Scurf pea. [Psorali- ditim lanceolatum (Pursh) Rydb.; P. elliptica Pursh; Lotodes ellipticum (Pursh) Kuntze; P. laxiflora Nutt. ex Torr. & Gray; P. micran- tJia A. Gray ex Torr.; Psoralidiwn niicran- thwn (A. Gray) Rydb.] Sand dunes and oth- er sandy sites at 1230 to 1770 m in Daggett, Garfield, Grand, Kane, Uintah, Washington, and Wayne cos.; Washington and Saskatchewan south to California, Ari- zona, New Mexico, and Oklahoma; 36 (viii). Var. stenophylla (Rydb.) Toft & Welsh. Canyon Scurfpea. [P. stenophylla Rydb.; Psoralidium stenophyllum (Rydb.) Rydb.] Sandy sites at 1270 to 1830 m in Emery, Garfield, Grand (type from Wilson Mesa), Kane, San Juan, and Wayne cos.; endemic; 26 (xiii). Var. stenostachys (Rydb.) Welsh comb, nov. based on Psoralea stenostachys Rydb., Bull. Torrey Bot. Club 42:46. 1913. Basin Scurfpea. [Psoralidium stenostachys (Rydb.) Rydb.] Dunes and other sandy sites in Juab, Kane, Millard, Salt Lake, Tooele (type from Government Well) and Weber cos.; endem- ic. Specimens from Salt Lake and Weber COS. are placed here tentatively because of the nature of the existing materials. The plants have not been collected in those counties in recent times, and they might well be extinct. The question of their varie- tal status might therefore be moot; 27 (ii). Psoralea megalantha Woot. & Standi. Large-flowered Breadroot. [Pediomelum megalanthtim (Woot. & Standi.) Rydb.] Sub- acaulescent to short-caulescent, 4-15 cm tall, from slender subterranean caudex branches arising from deep-seated tuberous roots; stems with 0-3 elongated internodes. incurved-strigose or with a few (rarely most) hairs spreading-ascending; leaves mainly 5 (8) -foliolate; petioles 1.2-9.5 cm long, pu- bescent like the stems or more commonly mainly spreading-hairy; leaflets 9-34 mm long, 4-23 mm wide, cuneate-obovate to subrhombic, gray-green, strigulose, and pimctate beneath, yellow-green, punctate, and strigose overall above; stipules scarious, 2-15 mm long; peduncles 1.4-5 cm long; racemes mainly 6- to 24-flowered, 2-5 cm long; pedicles 1.5-5 mm long, bracts com- monly bidentate, lance-ovate, 3-12 mm long; flowers 12.5-21 mm long, the banner, wings, and keel commonly purple or suf- fused with purple; calyx 12.5-18.5 mm long, the tube 5.5-8 (9) mm long, strongly gibbous, the lower tooth 4-9 (10) mm long, only somewhat broader than the lateral ones; pods included in the calyx. Mixed desert shrub, juniper-pinyon woodland, and blackbnish communities at 1430 to 1830 m in Duchesne, Grand, San Juan, Uintah, and Wayne cos.; Colorado and New Mexico. The plants of large-flowered breadroot are quite uniform, with two exceptions. Those from San Juan Co. tend to produce stems with elongate internodes, some of them with dense ascending-spreading pubescence. A collection from eastern Wayne Co. has flowers only 12.5 mm long. Much more ma- terial is necessary before decisions as to status of these variations will be possible; 16 (viii). Psoralea mephitica S. Wats. Skunk Bread- root. [Pediomelum mephiticum S. Wats.) Rydb.; Pediomelum retrorsum Rydb.; P. ret- rorsa (Rydb.) Tidestr. in Tidestr. & Kittell; P. mephitica var. retrorsa (Rydb.) Kearney 346 Great Basin Naturalist Vol. 38, No. 3 & Peebles.] Acaulescent to subacaulescent, 4.5-15 cm tall, from slender subterranean caudex branches arising from deep-seated tuberous roots; stems lacking or with very short intemodes above ground, retrorsely hairy; leaves mainly 5-foliolate; petioles 3.2-12 cm long, retrorsely hairy; leaflets 11-35 mm long, 4-28 mm wide, obovate to broadly so, gray-green, strigose overall, and punctate beneath, green to yellow-green, strigose overall (more densely along veins), and punctate above; stipules scarious, 4-12 mm long; peduncles 2-6 cm long; racemes mainly 12- to 35-flowered, 1.5-4.5 cm long; pedicels 1.5-3.5 mm long; bracts mainly not bidentate, elliptic-obovate, 5-12 mm long; flowers 10.5-12.8 mm long, the banner whitish or yellowish, the wings and keel purple or suffused with purple; calyx 9-12.5 mm long, the tube 3.5-4.5 mm long, strong- ly gibbous at the base, the lower tooth 4.5-9 mm long, about twice as broad as the others; pods included in the calyx. Black- brush and pinyon-juniper communities at 1470 to 1700 m in southeastern Washington Co.; Arizona, Nevada, and California. This species was described by Sereno Watson si- multaneously with P. castorea (Proc. Amer. Acad. 14:291. 1879). The type locality of both of these taxa is cited in the original descriptions as "near Beaver City, S. Utah." The type collections were both by Dr. E. Palmer (No. 96 for P. castorea, and No. 97 for P. mephitica). Palmer (Amer. Nat. 12:601. 1878) in describing the use of these plants by Indians (as food) noted that P. castorea S. Wats, "grows in exposed sandy localities between Beaver Dams, Arizona, and St. Thomas, Nevada." [sic]. Of P. meph- itica. Palmer (I.e.) notes that "it is abundant on the low places between the hills south- east from St. George, Southern Utah, and the Pah-Utes resort there to collect its roots" (sic). Thus, the type localities can be inferred as being in northwestern Arizona or south-eastern Nevada for P. castorea, and in the hills to the southeast of St. George for P. mephitica. There is no evidence to support the idea that P. castorea was ever collected in Utah, and it is herein excluded. Finally, no specimen of any of the bread- root psoraleas is known from the Great Ba- sin portion of Utah in modern collections, and it seems likely that none was taken there in the past; 6 (0). Psoralea pariensis Welsh & Atwood in Welsh, Atwood, & Reveal. Paria Breadroot. Acaulescent or subacaulescent, 2-9 cm tall, from slender subterranean caudex branches arising from deep-seated tuberous roots; stems lacking or with very short internodes above ground, strigose; leaves mainly 3- foliolate; petioles 1.3-7 cm long, strigose; leaflets 9-25 mm long, 7-22 mm wide, ob- ovate or orbicular, cuneate, rounded to truncate or emarginate apically, gray-green, glandular, and strigose beneath, yellow- green, glandular and strongly strigose along veins above; stipules scarious, 4-10 mm long; peduncles 0.5-2.8 cm long, the hairs appressed to ascending; racemes mainly 6-to 15-flowered, 1-2.5 cm long; pedicels 1-3.8 mm long; bracts abruptly acuminate, 4-8 mm long; flowers 8.8-12.5 mm long, the banner cream to ochroleucous, the wings and keel purple or suffused with purple; ca- lyx 8.6-11.4 mm long, the tube 3.3-4.6 mm long, more or less gibbous at the base, the lower calyx teeth 5.3-6.8 mm long, about twice broader than the others; pods includ- ed in the calyx, about 9 mm long. Ponde- rosa pine and juniper-piny on woods at 1700 to 2430 m in Garfield (type from Bryce Canyon) and Kane cos.; endemic; 8 (ii). Psoralea tenuiflora Pursh. Prairie Scurfpea. [Psoralidium tenuiflorum (Pursh) Rydb.; P. obtusiloba Torr. & Gray; Psorali- dium bigelovii Rydb.; P. bigelovii (Rydb.) Tidestr.; P. tenuiflora var. bigelovii (Rydb.) Macbr.; P. floribunda Torr. & Gray; Psorali- dium floribundum (Nutt.) Rydb.] Caules- cent, 40-100 cm tall or more, forming large clumps; stems with 8 or more elongated in- ternodes, strigose; leaves 3- to 5-folioIate, persistent at flowering; petioles 0.1-2.2 cm long, strigose to strigulose; leaflets 3-5, 6-40 mm long, 2-16 mm wide, oblanceolate throughout, mainly rounded to a cuspidate apex, gray-green and strigose beneath, yellow-green and glabrous or pubescent only along veins above; stipules scarious, 2-13 mm long, strigose; peduncles 0.5-7 cm long, or lacking, sometimes bracteate or some flowers axillary; racemes mainly 7- to Sept. 1978 Welsh: Utah Flora, Fabaceae 347 21 -flowered, 9-5.9 cm long; bracts ovate- acuminate, glabrous dorsally, 1.5-2.5 mm long, persistent; flowers 4.5-6 cm long, the petals indigo; calyx 2.5-3.2 mm long, the tube 1.5-2 mm long, campanulate, not gib- bous, the lower tooth 0.8-1.7 mm long, no- ticeably larger than the others; pods 1 -seeded, glabrous, conspicuously glandular. Pinyon- juniper, sagebrush, and mountain brush communities at 1700 to 2200 m in Garfleld, Iron, Kane, and Washington cos.; North Da- kota and Montana southward to Arizona, Texas, and Mexico. The segregation of vari- eties from among our specimens seems un- warranted; 14 (iii). PsoROTHAMNUS Rydb. Shrubs, armed; leaves alternate, odd-pin- nate, with 5 or more leaflets, glandular- dotted; stipules subulate or vestigial; pe- duncles opposite the leaves; flowers in lax racemes; calyx campanulate, 10-ribbed, 5- lobed; corolla papilionaceous, the petals all inserted on the hypanthium; stamens 9 or 10, monadelphous; petals mainly indigo; pods 1- to 2-seeded, indehiscent, exserted from the calyx. Note: Traditional treatments have placed the species herein recognized as Psorothamnus within an expanded Dalea (q.v.). References Rydberg, p. a. 1919. Psorodendron Rydb., and Psorothamnus Rydb. N. Amer. Fl. 24:41-48. 1. Branchlets densely reflexed-puberulent, bearing conspicuous yellow- to red- orange resinous glands 2 Branchlets merely appressed-strigose, lacking glands or only obscurely glandular 3 2(1). Calyx lobes ovate to oval, obtuse to rounded, or the lowermost ovate-acute .. P. thompsonae - Calyx lobes all lance-attenuate P. polyadenius 3(1). Calyx villous with contorted spreading hairs, the lateral teeth linear- lanceolate, quite as long as the tube; plants rare in Kane Co P. arborescens Calyx strigose with appressed hairs or glabrate, the lateral teeth lance- attenuate, mostly shorter than the tube; plants locally common in Garfield, Kane, San Juan, and Washington cos P. fremontii Psorothamnus arborescens (Torr.) Barn- eby. Beauty Indigo-bush. [Dalea arborescens Torr. in A. Gray; Parosela arborescens (Torr.) Heller; Psorodendron arborescens (Torr.) Rydb.; Dalea amoena S. Wats.; Paro- sela amoena (S. Wats.) Vail; Psorodendron amoenum (S. Wats.) Rydb.; Parosela john- sonii var. pubescens Parish; Psorodendron pubescens (Parish) Rydb.; Dalea fremontii var. pubescens (Parish) L. Benson; D. amoena var. pubescens (Parish) Peebles.] Armed shrubs, 4-10 dm tall or more; branchlets strigose to strigulose, sparingly glandular; stipules 1-2 mm long, subulate; leaves 1.4-3.8 cm long; leaflets 7-15, 1-10 (12) mm long, 0.7-1.5 mm wide, glandular beneath, strigose on both sides, linear to narrowly oblong, obtuse to rounded, the up- permost lateral leaflet often confluent with the terminal; peduncles 0.2-0.6 cm long, or the lowermost flower axillary; racemes 11- to 21-flowered, 1.8- to 4.5 cm long, the rachis pilosulous; bracts 1.5-3.5 mm long, lance-aristate, villosulous; calyx 8-10 mm long, the tube 3.8-4.8 mm long, definitely 10-ribbed, villous, the teeth 3.6-5.2 mm long, villous, not markedly differing in width; flowers 8.1-10.6 mm long, indigo; pods conspicuously glandular-dotted. Mixed desert shrub at 1230 to 1530 m in Kane Co.; Arizona and Nevada. Our material be- longs to var. pubescens (Parish) Barneby; 3 (i)- Psorothamnus fremontii (Torr.) Barneby. 348 Great Basin Naturalist Vol. 38, No. 3 Fremont Indigo-bush. [Dalea fremontii Torr. in A. Gray; Parosela fremontii (Torr.) Vail; Psorodendron fremontii (Torr.) Rydb.; Dalea johnsonii S. Wats., type from near St. George; Parosela johnsonii (S. Wats.) Vail; Psorodendron johnsonii (S. Wats.) Rydb.; Parosela johnsonii var. minutifolia Parish; D. fremontii var. minutifolia (Parish) L. Benson]. Armed shrubs, 5-15 dm tall or more; branchlets strigose, sparingly if at all glandular; stipules 0.3-1.5 mm long, fragile; leaves 1.8-6.5 cm long; leaflets (1) 3-9, 3-14 mm long, 0.8-6 mm wide, glandular beneath, strigose on both sides, linear to ob- long or elliptic, obtuse to rounded, the up- permost often confluent with the rachis; pe- duncles 0-2.2 cm long, or some flowers axillary; racemes 7- to 41-flowered, 3.5-14.3 cm long, the rachis strigose to strigulose; bracts 0.8-1.8 mm long, lanceolate, glabrous or else foliose and with 1-5 leaflets; calyx 4.5-6.5 mm long, the tube 2.5-3.3 mm long, obscurely if at all 10-ribbed, ap- pressed strigose to glabrate, the teeth 1.8-3.2 mm long, strigose, the upper markedly wider than the others; flowers 8.5-12 mm long, indigo; pods glandular- dotted. Blackbrush, creosote bush, mixed desert shrub, and (less commonly) pinyon- juniper communities at 730 to 2270 m in Garfield, Kane, San Juan, and Washington COS.; Nevada, Arizona, and California. The plants are strikingly beautiful, contrasting indigo-colored flowrs against grayish foliage; 44 (vii). Psorothamnus polyadenius (Torr.) Rydb. Glandular Indigo-bush. Armed shrubs, 1.5-8 dm tall or more; branchlets velvety with retrorse short hairs, conspicuously glandular with yellow- or red-orange resinous glands; stipules vestigial or to 0.5 mm long; leaves 0.4-2.8 cm long; leaflets 5-13, 1.2-6.5 mm long, 1-5 mm wide, oval to obovate, or ob- cordate, glandular beneath, strigose on both sides, obtuse, rounded, or emarginate, the uppermost often confluent to each other and to the rachis; peduncles 0.8-2.7 cm long ; racemes mainly 6- to 18-flowered, 2-5 cm long, the rachis retrorsely hairy; bracts 1-2.5 mm long, lanceolate, pilose; calyx 5.5-7.2 mm long, the tube 2.5-3.5 mm long, 10-ribbed, villous, the teeth 3.5-4.5 mm long, villous, lance-subulate, all about alike; flowers 7.5-9 mm long, indigo; pods glandular-dotted. Two rather distinctive va- rieties are present in Utah. 1. Branches strongly divaricate; leaflets flat, commonly less than 4 mm long; plants more than 2.5 dm tall, of Washington Co P. polyadenius var. polyadenius Branches merely ascending, or rarely some divaricate; leaflets curved, at least some over 4 mm long; plants commonly less than 2.5 dm tall, of Emery Co P. polyadenius var. jonesii Var. jonesii Barneby. [Dalea nummidaria M. E. Jones, type from Green River.] Salt desert shrub community, on Mancos Shale formation (Blue Gate Member) at ca. 1470 m in Emery Co.; endemic; 3 (0). Var. polyadenius. [Dalea polyadenia Torr. ex S. Wats.; Parosela polyadenia (Torr.) Heller.] Creosote bush-Joshua tree community on pedimental gravels, Beaver Dam Mts., Washington Co.; Nevada and California; 2 (0). Psorothamnus thompsonae (Vail) Welsh & Atwood. Thompson Indigo-bush. Armed shrubs, 2.5-8 dm tall or more; branchlets velvety with retrorse short hairs, con- spicuously glandular with yellow- to orange- red mammiform resinous glands; stipules vestigial or to 0.8 mm long; leaves 0.7-5 cm long; leaflets 7-17, 1-10 mm long, 0.6-4 mm wide, linear to oblong, oval, or obcor- date, glandular and strigose to glabrate be- neath, strigose above, the uppermost jointed to the rachis; peduncles 0.4-1.8 cm long, rarely obsolete and some flowers axillary; racemes mainly 8- to 25-flowered, 2-9 cm long, the rachis retrorsely hairy; bracts 0.5-1.5 mm long, lanceolate, glabrous or hairy, soon deciduous; calyx 3.7-5 mm long, the tube 2.1-3.2 mm long, conspicuously 1- ribbed, glabrous or villous, ovate to oval, Sept. 1978 Welsh: Utah Flora, Fabaceae 349 obtuse or only the lowermost acute; flowers pods glandular-dotted. Two distinctive vari- 7.8-10.8 mm long, indigo or purple-pink; eties occur in Utah. 1. Calyx tube villous; leaflets linear; plants of San Juan Co P. thompsonae var. whitingii Calyx tube glabrous; leaflets oblong to oval or obcordate; plants of various distribution P. thompsonae var. thompsonae Var. thompsonae. [Parosela thompsonae Vail]. Mixed desert shrub and salt desert shrub communities at 1170 to 2270 m in Garfield, Kane (?, the type supposedly from near Kanab, but more likely from Arizona), San Juan, and Wayne cos.; Arizona. There are no modern collections of this taxon from Kane Co.; 20 (iv). Var. whitingii (Kearney & Peebles) Barn- eby. [Dalea whitingii Kearney & Peebles]. Mixed desert shrub community at 1230 to 1530 m in San Juan Co.; Arizona; 1 (0). ROBINIA L. Shrubs or trees, often armed; leaves alter- nate, odd-pinnate, the leaflets petiolulate and stipulate; stipules setaceous and ca- ducous or persistent as spines; flowers in ax- illary racemes, white, pinkish, or pink, very showy, often aromatic; calyx campanulate to turbinate the 5 teeth triangular to triangular-acuminate; corolla papilio- naceous; stamens 10, diadelphous; ovary subsessile or sessile; pods oblong, straight, laterally flattened, tardily dehiscent. 1. Uppermost pair of calyx teeth connate almost to the apex, forming an emarginate lip; branchlets and peduncles lacking hispid processes R. pseudoacacia - Uppermost pair of calyx teeth free for about two-thirds of their length; branchlets and often the peduncles hispid 2 2(1). Branchlets and peduncles both glandular-hispid; shrubs, cultivated R. hispida - Branchlets lacking processes; peduncles glandular-hispid; trees, cultivated or indigenous (in Washington Co.) R- neomexicana Robinia hispida L. Rose- Acacia. Shrubs, arising from root sprouts, spreading, com- monly 1-2 m tall, except when grafted; branchlets both hispid and glandular-hispid; leaves 9.5-27 cm long; leaflets 7-13, 11-60 mm long, 7-40 mm wide, ovate or lance- ovate, elliptic, or oblong, obtuse, cuspidate, sparingly villous beneath, glabrous above; petioles hispid near the base; stipules obso- lete; peduncles 1-4 cm long, hispid; ra- cemes 3- to 10-flowered, 3-8 cm long; flow- ers 18-30 mm long, rose-pink; calyx turbinate-campanulate, the tube 4.5-6 mm long, the teeth 3.5-7 mm long, triangular- acuminate; pods hispid, seldom produced. Cultivated ornamental in Utah Co., and likely elsewhere; indigenous from Virginia and Kentucky to Georgia and Alabama. Evidence seems to indicate that rose-acacia is a sterile triploid, at least in part. Lack of fruit in plants from Utah seems to support this contention; 5 (ii). Robinia neomexicana A. Gray. New Mexico locust. [R. neomexicana var. luxuri- ans Dieck.; R. hixurians (Dieck.) Schneid ex Taroua & Schneid; R. breviloba Rydb.; R. subvelutina Rydb.] Small trees or shrubs, mainly 1-8 m tall; branchlets villosulous, rarely glandular; leaves 8-20 (28) cm long; leaflets 9-19, 12-40 mm long, 7-20 mm wide, lance-oblong to oblong, obtuse, cuspi- date, sparingly pubescent above and below, finally glabrate on one or both sides; pet- ioles villosulous near the base; stipules 3-10 mm long or more, often spiny; peduncles 1-4 cm long, glandular-pubescent to hispid throughout; racemes 3- to 14-flowered, 3-8 cm long; flowers 16-24 mm long, pink; ca- 350 Great Basin Naturalist Vol. 38, No. 3 lyx campanulate, the tube 5-8 mm long, the teeth 3-5 mm long, triangular-acuminate pods 40-80 mm long, glandular-pubescent to hispid or glabrous. Talus slopes and ter- races in Zion Canyon, Zion National Park, Washington Co.; Colorado, Nevada, Ari- zona, New Mexico, Texas, and Mexico. The plant is also known from cultivation in Cache and Utah cos., and is probably pres- ent elsewhere. The pink-flowered tree lo- custs of our region are all more or less in- volved through hybridization with the New Mexico locust and the black locust; 4 (i). Robinia pseudoacacia L. Black Locust. Trees to 25 m tall or more; branchlets pu- berulent to villosulous, lacking glands; leaves (6) 8.5-26 cm long; leaflets (5) 11-25, 11-60 mm long, 6-30 (38) mm wide, lance- oblong to oblong, obtuse to retuse or cuspi- date, puberulent to glabrate on both sides; petioles villosulous to pilose near the base; stipules minute, or represented by spines; peduncles 1.4-4.3 cm long, puberulent to villosulous; racemes 11- to 27-flowered, 4-13 cm long; flowers 12-20 (23) mm long, white (fading cream), or pale pink; calyx broadly campanulate to turbinate, the tube 3.5-5.5 mm long, the teeth 1.5-2 mm long, the upper pair connate, the sinus shallow; pods 40-120 mm long, glabrous. Cultivated ornamental, street, and shade trees, long- persisting and escaping; now known from Carbon, Cache, Grand, Kane, Millard, Salt Lake, Sanpete, Utah, Washington, and Wayne cos., and likely more widely culti- vated; indigenous in the eastern United States. This is a handsome shade tree, with very hard wood. The flowers yield nectar, and the wood has been used for fence posts; 34 (x). SOPHORA L. Trees or herbs, unarmed; leaves alternate, odd-pinnately compound; stipules obsolete or herbaceous; flowers in terminal racemes or panicles, perfect, papilionaceous; calyx 5- toothed; petals all distinct; stamens 10, dis- tinct or essentially so; ovary stipitate; pods spreading to pendulous, subterete, con- stricted between the seeds, indehiscent or tardily so. 1. Plants trees, cultivated; flowers borne in panicles in midsummer S. japonica Plants herbaceous, indigenous; flowers in racemes, opening in spring 2 2(1). Flowers blue-purple to blue; leaflets more than 5 times longer than broad, silvery-hairy S. stenophylla Flowers white to cream; leaflets less than 5 times longer than broad, gray- QYQen ^- nuttalliana Sophora japonica L. Japanese Pagoda- tree. Trees to 12 m tall, the bark green and smooth for some years; branchlets sparingly villosulous to glabrate, leaf bases expanded, obscuring the axillary buds; leaves 12-23 cm long; leaflets 7-17, 14-60 mm long, 8-29 mm wide, lance-oblong to lanceolate, acute and apiculate, strigose beneath, gla- brate or glabrous above; stipules obsolete; panicles many-flowered, 15-40 cm long; flowers 11-14 mm long, white to cream; ca- lyx broadly campanulate 5.5-6.5 mm long, the tube 4.5-5.1 mm long, the teeth 0.8-1.2 mm long, broadly triangular; pods mainly 50-100 mm long, fleshy, constricted be- tween the seeds. Cultivated ornamental, widely planted, but our records only from Salt Lake and Utah cos.; introduced from China; 5 (i). Sophora nuttalliana Turner. Silky Soph- ora. [S. sericea Nutt.] Perennial, caulescent, 12-27 (30) cm tall, from a rhizome; pu- bescence essentially basifixed; stems ascend- ing to erect; stipules 1.5-4 mm long, dis- tinct, caducous; leaves 2.5-6 (7) cm long; leaflets 13-23, 2-11 mm long, 0.8-6 mm wide, oblong-obovate to obovate, rounded to retuse, often folded, strigose beneath, glabrous above; peduncles 0.8-3 cm long; racemes mainly 8- to 52-flowered, 3-9 cm long at an thesis, the axis 3-13 cm long in fruit; bracts 3-7 mm long; pedicels 2-4 mm Sept. 1978 Welsh: Utah Flora, Fabaceae 351 long; bracteoles 1; calyx 8-10.5 mm long, gibbous, the tube 6.4-8.5 mm long, oblique- ly short-cylindric, the teeth 2-2.5 mm long, triangular; flowers 14-19 mm long, white to cream (fading cream); pods erect-ascending, stipitate, the stipe 6-12 mm long, the body 12-40 mm long, 3-4.5 mm thick, con- stricted tightly between the usually 1-3 seeds, strigose. River bottoms and roadsides at 1070 to 1670 m in San Juan and Wash- ington COS.; Wyoming and South Dakota, south to Arizona, New Mexico, and Texas; 2 (0). Sophora stenophylla A. Gray. Silvery Sophora. Perennial, caulescent, 13-41 cm tall (above ground), from deeply seated rhi- zomes; pubescence basifixed; stems ascend- ing to erect; stipules 3-12 mm long, dis- tinct, caducous, or obsolete; leaves 1.7-5.6 cm long; leaflets 9-15, 7-27 mm long, 0.5-4 mm wide, linear to narrowly oblong, acute to attenuate, silvery pilosulous, the pu- bescence fading yellowish in time; pe- duncles 1.7-5 cm long; racemes mainly 12- to 39-flowered, 5-17 cm long at anthesis, 6-23.5 cm long in fruit; bracts 3-7 mm long; pedicels 1-8 mm long; bracteoles 0; calyx 6.5-10.8 mm long, gibbous, the tube 4.8-7.2 mm long, obliquely short-cylindric, the teeth 1.7-3.6 mm long, ovate-triangular; flowers 15-27 mm long, blue-purple to blue; pods spreading declined, stipitate, the stipe 8-16 mm long, the body 15-60 mm long, 6-8 mm wide, strongly constricted be- tween the usually 1-5 seeds, strigose. Sand dunes and other sandy sites, mainly in mix- ed desert shrub communities, at 900 to 2270 m in Emery, Garfield (at Red Canyon, with ponderosa pine). Grand, San Juan, Uintah, Washington, and Wayne cos.; New Mexico and Arizona. This is one of the most beauti- ful of the legume species in Utah; 41 (vi). Spartium L. Shrubs, imarmed; leaves alternate to sub- opposite, simple; stipules obsolete; flowers in terminal, erect racemes, papilionaceous, perfect; calyx 1-lipped, the 5 teeth margin- al, all on the lower side of the calyx, the three central ones approximate, the lateral ones somewhat removed; petals yellow, the keel pubescent along the lower edge, and with a porrect beak; stamens 10, mon- adelphous; ovary sessile; pods spreading-as- cending, laterally compressed, many-seeded. Spartium junceum L. Spanish Broom. [Genista juncea (L.) Lam.] Shrubs, 15-25 dm tall; leaves simple, 8-27 mm long, 0.5-4 mm wide, linear to narrowly oblanceolate, acute to obtuse, strigose on both sides; pe- duncles 3-22 cm long (from last leaf to first flower); racemes 3- to 16-flowered, 3-16 cm long; bracts minute, caducous; bracteoles 0; pedicels 1-4 mm long; calyx oblique, 7-8 mm long, glabrous, the teeth minute; flow- ers 21-26 mm long, yellow; pods sessile, 50-80 mm long, 5-7 mm wide, strigose, di- hiscent. Cultivated ornamental in Washing- ton Co.; widely cultivated in the South; in- troduced from southern Europe; 1 (i). Sphaerophysa DC. Perennial, caulescent, from rhizomes; leaves alternate, odd-pinate; stipules adnate to the petiole base, all distinct; flowers pa- pilionaceous, borne in axillary racemes, each subtended by a single bract; bracteoles 1; calyx 5-toothed; petals 5, dull red, drying lavender to brown; stamens 10, diadelphous; ovary enfolded by the staminal sheath; style glabrous except for a tuft of hair below the stigma; pods stipitate, bladdery-inflated, subunilocular. Sphaerophysa salsula (Pallas) DC. [Phaca salsula Pallas; Swainsona salsula (Pallas) Taub. in Engl. & Prantl.] Perennial, caules- cent, 40-70 cm tall, from a deeply placed rhizome; pubescence basifixed; stipules 1-4 mm long, all distinct; leaves 3-10 cm long; leaflets 15-25, 3-18 mm long, 1-7 mm wide, oblong-obovate to elliptic, retuse to obtuse and apiculate, strigose beneath, glabrous above; peduncels 2.5-9 cm long; racemes 5- to 17-flowered, the flowers as- cending in anthesis, finally nodding, the axis 2.5-9 cm long in fruit; bracts 1-2 mm long; pedicels 2.5-8 mm long; bracteoles 2; calyx 5-6 mm long, the tube 3.8-4.6 mm long, campanulate, the teeth 1.2-2 mm long, triangular; flowers 12-14 mm long, dull-red, fading lavender to brown; pods ascending to declined, stipitate, the stipe 4-7 mm long, 352 Great Basin Naturalist Vol. 38, No. 3 the body bladdery-inflated, ovoid, 13-24 mm long, 9-20 mm wide (when pressed), unilocular, strigulose. Introduced weedy species known from the Uinta Basin and to be expected anywhere; widespread in the western United States; adventive from Asia. This plant resembles an Astragalus species, and has been mistaken twice as belonging to previously imdescribed and imnamed in- digenous species (A. violaceus St. John; A. iochrous Barneby). Generic concepts revolv- ing aroimd this genus are unresolved and it seems likely that the plants will ultimately be placed with some earlier named genvis; 1 (0). Thermopsis R. Br. Perennial herbs, caulescent, from rhi- zomes; leaves alternate, palmately trifoliol- ate; stipules foliaceous; flowers papilio- naceous, borne in terminal racemes; bracts herbaceous, persistent; calyx 5-toothed; pet- als 5, yellow or suffused with purple, the keel rounded; stamens 10, distinct; ovary stipitate, the style glabrous; pods narrowly oblong, flattened, many-seeded. Pods straight not especially loment-like, erect or ascending; plants mostly 20-70 cm tall or more, common through most of Utah L. montana Pods curved, loment-like, spreading to recurved; plants mostly 14-40 cm tall, known from Uintah Co L. rhombifolia Thermopsis montana Nutt. in T. & G. Golden Pea, Yellow Pea. Caulescent, 20-75 (100) cm tall, the stems erect, pilosulous to glabrate; stipules foliar, lanceolate to ovate; 13-60 mm long, 3-30 mm wide; petioles 0.8-3.7 cm long; leaflets 3, 21-92 mm long, 5-36 mm wide, elliptic to lanceolate or ob- lanceolate, acute to rounded, pilosulous be- neath, glabrous to glabrate above; peduncles 2.2-13 cm long; bracts 5-11 mm long; ra- cemes mainly 2- to 23-flowered, 6-25 cm long in anthesis, the axis 9-28 cm long in fruit; flowers 20-26 mm long, yellow; calyx 10.2-12.3 mm long, the tube 7-8.3 mm long, obliquely campanulate, the teeth 3.1-4 mm long, ovate-triangular; pods erect or as- cending, stipitate, the stipe 2.5-6 mm long, the body 40-54 mm long, 5-7 mm wide, pi- lose, stramineus or turning black. Moist sites along streams, in meadows, around seeps and springs at 1251 to 3416 feet in Box El- der, Daggett, Duchesne, Garfield, Juab, Kane, Millard, Morgan, Piute, San Juan, Sanpete, Sevier, Summit, Uintah, Utah, Wasatch, Washington, Wayne, and Weber COS.; British Columbia east to Montana and south to California, Arizona, and New Mex- ico. The abundant materials present in Utah herbaria demonstrate a wide range of varia- tion in leaflet and stipule size and shape. There appears to be no basis for segregation of subordinate taxa, even those from the southern tier of counties known previously as T. pinetorum Greene or as T. ovata (Rob- ins.) Rydb. [T. montana ssp. ovata Robins, ex Piper; T. montana var. ovata (Robins.) St. John]. The leaves of plants from Wash- ington Co. average larger, but that is hardly a basis for segregation; 106 (xv). Thermopsis rhombifolia Nutt. ex Rich- ards. Caulescent, 15-40 cm tall, the stems erect, glabrate; stipules foliose, ovate to ob- lanceolate, 6-30 mm long, 2-22 mm wide; petioles 0.3-2.5 cm long; leaflets 3, 15-47 mm long, 7-25 mm wide, obovate to oblan- ceolate, obtuse to rounded, glabrous beneath and above; peduncles 0.5-5.8 cm long; bracts simple to foliose; racemes mainly 4- to 30-flowered, 2-10 cm long in anthesis, the axis 2-12 cm long in fruit; flowers 18-22 mm long; calyx 7.5-10 mm long, the tube 4.5-4 mm long, triangular-ovate; pods divaricate, finally recurved, loment-like, stipitate, the stipe 1.5-4 mm long, the body 25-70 mm long, 5-7 mm wide, pilose to glabrate. Sandy and clay soils, mainly where moist, at 1500 to 1800 m in Daggett (Neese 4532 BRY) and Uintah Co. (Welsh 158; Cronquist 11499 BRY); Alberta and Sas- katchewan south to Colorado and Nebraska. Trifolium L. Perennial or short-lived perennial or an- Sept. 1978 Welsh: Utah Flora, Fabaceae 353 nual, caulescent or acaulescent, from a tap- tion Lupinaster (Leguminosae). Brit- root and caudex or rhizome; leaves alter- tonia 17:107-112. nate, palmately to pinnately 3-foliolate, or 1969. Taxonomy of Trifolitim rarely 4- to 7-foliolate, commonly serrate (Leguminosae). II. The T. longipes throughout, rarely entire; stipules mem- complex in North America. Canad. braneous to foliaceous, often connate; flow- Jour. Bot. 47: 93-113. ers papilionaceous, borne in terminal or ax- 1971. Taxonomy of Trifolium illary, pedunculate to sessile, subcapitate (Leguminosae). III. T. eriocephalum. heads or racemes; calyx 5-toothed; petals 5, Canad. Jour. Bot. 49:395-405. pink, white, or red-purple, withering and 1972. Taxonomy of Trifolium persistent, finally investing the pod; stamens (Leguminosae). IV. The American spe- 10, diadelphous; pods usually shorter than cies of section Lupinaster (Adanson) the calyx, indehiscent, 1- to several-seeded. Seringe. Canad. Jour. Bot. 50:1975-2007. References 1965. Taxo Hum: Five American species of Sec- folium. San Francisco. 325 pp. McDermott, L. F. 1910. An illustrated key Gillett, J. M. 1965. Taxonomy of Trifo- to the North American species of Tri- 1. Plants acaulescent or subacaulescent, mainly 1.5-10 cm tall 2 - Plants caulescent, with 1 or more elongated internodes (see also T. parryi); mainly 10-60 cm tall 7 2(1). Heads 1- to 4-flowered, essentially sessile, the flowers 15-23 mm long; plants of high elevations in Uinta and LaSal mountains T. nanum - Heads several-flowered, sessile or pedunculate, the flowers either shorter or heads pedunculate; plants of various distribution 3 3(2). Plants densely pulvinate-caespitose, matted 4 Plants loosely caespitose, not especially mat-forming 5 4(3). Calyces densely woolly-villous; herbage silvery-hairy; plants known from San Francisco Mts., Beaver Co T. andersonii Calyces strigose to strigulous; herbage green, merely strigose; plants of Daggett, Summit, and San Juan cos T. andinum 5(3). Leaflets toothed; calyces villosulous to pilosulous; flowers less than 9 mm long; plants of moderate elevation, broadly distributed T. gijmnocarpon Leaflets entire, or if toothed then the calyces glabrous; flowers more than 9 mm long; plants of high elevations 6 6(5). Leaflets entire; calyces strigose; plants of the Uinta, LaSal, Abajo, and Henry mountains T. dasyphyllum Leaflets toothed; calyces glabrous; plants of the Uinta, LaSal, and Abajo mountains T. parryi 7(1). Plants stoloniferous, prostrate and rooting at the nodes; flowers white; calyx not bladdery-inflated; introduced, widespread T. repens Plants not stoloniferous (except in T. fragiferum), usually erect or ascending, not rooting at the nodes; flowers mainly pink to red-purple 8 8(7). Calyx soon bladdery-inflated and enclosing the corolla; plants introduced, uncommon T. fragiferum 354 Great Basin Naturalist Vol. 38, No. 3 — Calyx not accrescent or only slightly so, never enclosing the corolla; plants various 9 9(8). Heads sessile or nearly so, commonly immediately subtended by a trifoliolate bract; plants cultivated, escaping, and persisting T. pratense — Heads with well-developed peduncles, not immediately subtended by foliose bracts; plants indigenous or cultivated 10 10(9). Peduncles bent at the apex, the heads reflexed or appearing turned to one side 11 — Peduncles straight, the heads erect 13 11(10). Calyx tube and teeth villous; ovary with hairs near the apex along the ventral side; plants of lower middle elevations in Juab, Sevier, and Beaver (?) cos T. eriocephalum — Calyx tube and teeth sparingly villosulous to glabrous; ovary glabrous, or scaly, but not hairy as above; plants of various distribution 12 12(11). Heads definitely longer than broad; calyx teeth shorter than the tube; basal leaves prominent, long-petioled; plants of Washington and Iron cos T. macilentum — Heads about as broad as long; calyx teeth subequal to the tube; basal leaves more shortly petioled than the subbasal ones; plants of Beaver, Emery, Garfield, Grand, Piute, San Juan, Sanpete, Sevier, Summit, and Utah cos T. kingii 13(10). Heads subtended by a spinose-toothed involucre 14 — Heads lacking an involucre; plants from a caudex, variously distributed, common or uncommon 15 14(13). Plants annual; head 5-20 mm wide; plants of Salt Lake, Weber, and Cache cos T. variegatum — Plants perennial; head 20-30 mm wide; of Tooele and Salt Lake cos T. wormskpldii 15(17). Flowers mainly 7-9 mm long; heads axillary, from the uppermost nodes; plants cultivated, escaping, and persisting T. hybridum — Flowers 10-16 mm long; heads terminal, solitary; plants indigenous 16 16(15). Calyx glabrous; flowers mainly 14-16 mm long; plants rare in Sevier Co T. beckwithii — Calyx villosulous to pilosulous; flowers mainly 10-12 mm long; plants wide- spread in Utah T. longipes Trifolium andersonii A, Gray. Anderson or less obscured by pubescence, villosulous Clover. Acaulescent, densely pulvinate- on both sides with hairs 0.3-0.7 mm long, caespitose, mat-forming, 0.8-3 cm tall, from commonly folded, mucronate; stipules woody caudex branches and a thick taproot, scarious, pilose, 5-9 mm long; heads lacking the stems obscured by imbricated stipules an involucre, borne subsessile or on pe- and persistent leaf bases; petioles 0..3-1.1 duncles 0.2-0.6 cm long, these densely pi- cm long; leaflets 3-3.8 mm long, 1.5-3.4 lose; flowers 4-9, the banner reddish-purple, mm wide, oblanceolate to obovate, entire or the keel and wings pale, 8-9 mm long, on toothed near the apex and the teeth more pedicels ca. 0.5 mm long (obscured by pi- Sept. 1978 Welsh: Utah Flora, Fabaceae 355 lose hairs); calyx 8.5-9.5 mm long, the tube 3-4 mm long, campanulate, obscured by villous hairs to 1.5 mm long, the teeth 5-7 mm long, subulate, villous; pods unknown (in our material). Volcanic gravels in pin- yon-juniper woodland at 2130 m in San Francisco Mts., Beaver Co.; California and Nevada. Our plant differs in several salient features from T. andersonii sens. lat. It is therefore described as var. friscanum Welsh var. nov. Similis Trifoliae andersonii a qua imprimis differt pedunculis brevioribus, flo- ribus parvioribus, et folium pili brevioribus. Holotype: Utah, Beaver Co., Grampian Hill, San Francisco Mts., Peabody, Taylor, & Thome 406, 3 June 1976 (BRY); 3 (0). Trifolium andinum Nutt. in Torr & Gray. Andean Clover. Acaulescent, densely pulvinate-caespitose, mat-forming, 0.3-5 cm tall, from woody caudex branches and a thick taproot, the stems obscured by imbri- cated stipules and persistent leaf bases; petioles 0-2.3 cm long; leaflets, 3, 2-12.5 mm long, 1.2-5 m wide, oblanceolate to obovate, toothed in the apical third, strigose to strigulose on both sides, commonly fold- ed, abruptly acuminate; stipules scarious, yellowish, glabrous except towards the tip, 3-9 mm bug; heads closely subtended by reduced leaf-like bracts, these with or with- out a trifoliolate bract, sessile or appearing pedunculate by elongation of intemodes on floriferous branches, the internodes glabrous; flowers 7-15, in two closely asso- ciated heads, the banner violet-purple, the keel and wings ochroleucous, 9-12 mm long, on pedicels 0.5-1 mm long, these glabrous to strigose; calyx 5-7 (8.5) mm long, the tube 2.2-3.6 (4) mm long, spar- ingly pilose to glabrous, the teeth 2-3.8 (4.5) mm long, lance-subulate, pilose; pods 4-5 mm long, 2.3-2.7 mm wide. Ponderosa pine, sagebrush, or mixed shrub commu- nities at 1970 to 2730 m in Daggett, Sum- mit, and San Juan cos.; Wyoming and Ari- zona. The existence of Andean Clover in two centers in Utah (i.e., on the north slope of the Uintas and on Navajo Mountain) is without precedence among the legumes of Utah. The plants from Navajo Mountain differ in features of the flower, mainly in the length of the calyx teeth and tube. Those from the Uinta Mountain centrum have calyx teeth subequal to or surpassing the tube; those from Navajo Mountain have calyx teeth much shorter than the tube; 6 (i)-' . Trifolium beckwithii Brewer ex S. Wats. Beckwith Clover. Caulescent, 12-40 cm tall, from rhizomes and thick roots; petioles of lower leaves 4-15 cm long, becoming short- er upwards; leaflets 3, 12-45 (50) mm long, 7-20 mm wide, elliptic to lance-elliptic or oblong, toothed from near the base, glabrous, flat, emarginate to apiculate; stipules prominent, herbaceous to scarious, 12-28 mm long; heads not subtended by an involucre, about as broad as long, 25-35 mm wide, terminal or axillary, on peduncles 5-26 cm long, these glabrous; flowers nu- merous, the petals pink, fading brown, 12-16.5 mm long; calyx 6.5-8 mm long, the tube 2.5-3.1 mm long, gibbous, glabrous, 5- veined, the teeth 3.5-4.9 mm long, lance- subulate, glabrous; pods 4.8-5.2 mm long, 1.9-2.1 mm wide. Meadows at 2070 to 2200 m in Sevier and Piute (?) cos.; California, Oregon, Nevada, Idaho, Montana, and South Dakota. Our collections from Utah have flowers and parts which average larger than for the species as a whole. The one lo- cality for the species in Utah is common to T. eriocephalum; 2 (i). Trifolium dasyphyllum Torr. & Gray. Uinta Clover. Acaulescent, loosely mat- forming, 2-14 cm tall, from a caudex and thick taproot, the stems obscured by imbri- cated stipules and leaf bases; petioles 0.3-4 cm long; leaflets 3, 3-28 mm long, 1-5 mm wide, oblanceolate, entire, strigose beneath, strigose to glabrate above, flat or folded, sharply apiculate; stipules scarious, glabrous or strigose, 5-17 mm long; heads closely subtended by a long-lobed involucre, 11-18 mm wide, terminal, sessile or on peduncles 0.5-10 cm long, these strigose; flowers 6-16, erect, the banner violet to ochro- leucous, the keel and wings purple, or all purple, 11-13 mm long; pedicels 0.5-1.5 mm long; calyx 4.7-9.1 mm long, the tube 2.5-2.9 mm long, strigose to glabrate, the teeth 1.9-6.4 mm long, subulate, strigose to glabrate; pods 4-6 mm long. Alpine mead- ows at 2730 to 3800 m in Daggett, Gar- 356 Great Basin Naturalist Vol. 38, No. 3 field. Grand, San Juan, Summit, and Uintah COS.; Montana, Wyoming, Colorado, and New Mexico. Plants from Utah are assign- able to var. uintense (Rydb.) Welsh stat. nov. based on T. uintense Rydb., Bull. Tor- rey Bot. Club 34: 47. 1907, type from Uinta Mts. [T. dasyphijllum f. uintense (Rydb.) McDermott; T. dasyphijllum ssp. uintense (Rydb.) J. M. Gillett]; 117; (0). Trifolium eriocephalum Nutt. in Torr. & Gray. Woolly Clover. Caulescent, 12-36 (42) cm tall, from a caudex and tuberous to slender taproot; petioles 0.8-10 cm long; leaflets 3, 5-53 mm long, 5-12 mm wide, oblong to oblong-lanceolate or elliptic, toothed from near the base, thinly pilose to pilosulous on both sides, flat, obtuse to acute, apiculate; stipules foliaceous, glabrous or pilosulous, 15-55 mm long; heads lacking an involucre, 20-30 mm long, 20-27 mm wide, terminal, on peduncles 4-11 cm long, bent apically, thinly villo- sulous; flowers numerous, curved at base and reflexed, the petals purple to red- purple, rarely white, fading brown, 12-13.5 mm long; pedicels essentially obsolete; calyx 6.5-8 mm long, the tube 2.5-3.3 mm long, villosulous, the teeth 3.5-5.1 mm long, villo- sulous to glabrate, subulate; pods 2.5-7 mm long. Meadows at 15.30 to 2270 m in Bea- ver (?), Juab, and Sevier cos.; Washington, Idaho, Montana, California, and Nevada. Our few specimens are assignable to var. villiferum (House) Martin [T. villiferum House; T. eriocephahun f. villiferum (House) McDermott; T. eriocephalum ssp. villiferum (House) J. M. Gillett]. The type (US) of T. villiferum is labeled as "S. Utah. Dr. E. Pal- mer 91, 1877." Gillett (1971) cites the type locality as "Beaver City, Beaver Co., Utah." The problem of type locality is the same as that for Psoralea mephitica (q.v.) and P. cas- torea, both cited as being from "near Bea- ver City, S. Utah." That neither came from there was alluded to by Palmer (see citation under P. mephitica), and also by Jones (Cal. Acad. Sci. II. 5:632. 1895) who noted that "Dr. Palmer never collected any plants at Beaver City, Utah, during the year in which he collected these species P. mephitica and P. castorea, but he did collect in that year in the Beaverdam Mountains on the north- eastern comer of Arizona at a place called Beaverdam [sic]. Thus, the type locality of T. villiferum is in doubt, but it seems likely that the specimens came from Washington Co., where it is unknown from recent col- lections; 8 (ii). Trifolium fragiferum L. Strawberry-head- ed Clover. Caulescent, 5-30 cm long, rhizo- matous and sometimes stoloniferous, decum- bent to ascending; petioles 0.5-13 cm long; leaflets 3, 6-22 mm long, 4-15 mm wide, obovate, toothed from near the base, trun- cate to retuse and apiculate, flat, glabrous to glabrate on both sides; stipules scarious, 8-20 mm long; heads involucrate, many- flowered, subglobose, 10-22 mm wide, on peduncles 2-17 cm long, these glabrous; flowers 4-6 mm long, purplish, finally in- cluded within the accrescent calyx; calyx fi- nally bladdery-inflated, pilose, reticulately veined around translucent lacimae. Mead- ows, roadsides, and other disturbed sites at 1370 to 2135 m in Cache, Garfield, Juab, Millard, Summit, Piute, Salt Lake, Utah, and Weber cos., and likely elsewhere; in- troduced from Europe; 10 (ii). Trifolium gymnocarpon Nutt. in Torr. & Gray. Nuttall Clover. Acaulescent to short- caulescent, 4-16 cm tall, from a caudex and taproot, the stems mainly obscured by im- bricated stipules and leaf bases; petioles 1-9 cm long; leaflets 3-5, 6-23 mm long, 2-10 mm wide, elliptic to oblong, ovate, or ob- ovate, toothed from near the base, pilo- sulous beneath, glabrous to pilosulous above, flat or folded, rounded to acute and apicu- late; stipules scarious to herbaceous, 6-2.3 mm long; heads without an involucre, he- mispheric, commonly 12-18 mm wide, ter- minal and axillary, on peduncles 1-6.5 cm long, these strigulose, erect to bent apically; flowers 6-15, the lower ones reflexed in age, the petals pink to lavender or purple, 8.5-11 mm long; pedicels 0.5-1 mm long; calyx 4.4-7 mm long, the tube 2.2-3.3 mm long, strigose, the teeth 1.8-3.7 mm long, subulate, strigose; pods 4-8 mm long, 3-4.5 mm wide. Two weak, sympatric varieties occur in Utah. They are hardly worth tax- onomic recognition, but possibly represent trends. Sept. 1978 Welsh: Utah Flora, Fab ace ae 357 Leaflets glabrous above T. gijmnocarfxm var. gymnocarpon Leaflets pilose above T. gyitinocarfwn var. phimmerae Var. gymnocarpon. [T. suhcaulescens A. Gray in Ives; T. gijmnocarpon var. suh- caulescens (A. Gray) A. Nels.; T. nemorale Greene]. Mixed desert shrub, pinyon-juni- per, mountain brush, sagebnish, ponderosa pine, Douglas fir, and spruce-fir commu- nities at 1570 to 2900 m in Carbon, Dag- gett, Duchesne, Garfield, Juab, Kane, Mil- lard, Piute, San Juan, Sanpete, Sevier, Uintah, Utah, and Wayne cos.; Nevada, Wyoming, Colorado, Arizona, and New Mexico; 25 (x). Var. plummerae (S. Wats.) Martin. [T. phimmerae S. Wats.; T. gymnocarpon f. phimmerae (S. Wats.) McDermott; T. gyyn- nocarpon ssp. phimmerae (S. Wats.) Gillett]. Sagebrush, pinyon-juniper, and other com- munities at 1530 to 2430 m in Beaver, Car- bon, Daggett, Duchesne, Millard, San Juan, Sanpete, Summit, Uintah, Wasatch, and Wayne cos.; Oregon, Idaho, Wyoming, Cali- fornia, and Nevada. This phase intergrades completely with the preceding in Utah, and if recognized at any taxonomic level, should best be treated at the rank of forma, as was done by McDermott; 10 (0). Trifolium hybridum L. Alsike Clover. Caulescent, 15-70 cm tall, erect or ascend- ing (rarely decumbent), from a caudex and taproot; petioles 0-16 cm long; leaflets 3, 5-38 mm long, 3-28 mm wide, oval to lance-elliptic, ovate or obovate, flat, toothed from near the base, glabrous on both sides, obtuse to retuse and apiculate; stipules her- baceous, 5-25 mm long; heads without an involucre, 12-25 mm wide, terminal and ax- illary on peduncles 1.5-13 cm long, these glabrous or glabrate, erect; flowers many, the lower reflexed in age, the petals white to pink or reddish, fading red-brown, 5-9 mm long; calyx 2.7-4 mm long, the tube 1.2-1.6 mm long, glabrous, scarious, the teeth 1-2.5 mm long, subulate, glabrous; pods 1- to 3-seeded. Cultivated, short-lived, forage plant, escaping and persisting (?) in Cache, Davis, Garfield, Piute, Salt Lake, San Juan, Sanpete, Svmimit, Utah, Wasatch, and Weber cos.; and perhaps universal; in- troduced from Europe; 21 (ii). Trifolium kingii S. Wats. King Clover. Caulescent, (2) 7-40 cm tall, erect or as- cending, from a caudex and taproot; pet- ioles 0.8-15 cm long, the longest near the base but not basal; leaflets 3, 5-80 mm long, 4-26 mm wide, elliptic to lance-ellip- tic or lanceolate, flat, toothed from near the base, glabrous on both sides, obtuse to acute or attenuate and apiculate; stipules fo- liaceous, 8-30 mm long; heads nodding to suberect, without an involucre, 15-32 mm long, 15-30 mm wide (when pressed), ter- minal, on peduncles 3-13 cm long, these glabrous; flowers many, reflexed, the petals violet to purplish (rarely white), 12-16 mm long; calyx 5.3-9 mm long, glabrous, the tube 2.1-3.5 mm long, the teeth 1.8-6 mm long, subulate; pods 1- to 3-seeded. Mead- ows and open woods at 2270 to 3700 m in Beaver, Emery, Garfield, Grand, Piute, San Juan, Sanpete, Sevier, Summit, and Utah COS.; endemic. With its nodding heads and reflexed flowers, the King clover simulates T. eriocephahim in all salient aspects. The main differential feature involves the pu- bescence of herbage and calyces in T. erio- cephahim-, 35 (v). Trifolium longipes Nutt. in Torr. & Gray. Rydberg Clover. Caulescent (rarely acaulescent), 5-31 (37) cm tall, erect or as- cending from a branching caudex and stout to slender taproot; petioles 1.2-10 cm long; leaflets 3, 5-47 (57) mm long, 3-18 mm wide, narrowly oblong to elliptic, oblanceol- ate, or obovate, flat, toothed from near the base, pilosulous beneath, glabrous above, acute to obtuse and apiculate; stipules fo- liaceous, 8-40 mm long; heads erect, with- out an involucre, 17-31 mm long, 15-33 mm wide (when pressed), terminal, on pe- duncles 0.5-17 cm long, these strigulose; flowers many, finally reflexed, the petals whitish to pink or purple, 11-13 mm long; calyx 4.5-7.8 mm long, the tube 1.6-2.5 mm long, scarious, pilose distallv, the teeth 2.9-5.8 mm long, pilose, subulate; pods 1-to 4-seeded. This is the common clover in mountains in Utah. There are two rather distinctive and largely allopatric varieties. 358 1. Great Basin Naturalist Vol. 38, No. 3 Leaflets of main leaves commonly more than 5 times longer than broad; roots slender, not much enlarged; plants of the Uinta, Deep Creek, LaSal, and Abajo mountains T. longipes var. reflexum Leaflets of main leaves commonly less than 4 times longer than broad; roots tuberous-thickened; plants mainly of southwestern Utah (also in San Juan Co.) T. longipes var. pygmaeum Var. pygmaeum A. Gray in Ives. [T. long- ipes ssp. pygmaeum (A. Gray) J. M. Gillett; T. longipes var. brachypus S. Wats., type from St. George; T. brachypus (S. Wats.) Blankinship; T. rusbyi Greene; T. confusum Rydb., type from southern Utah, i.e., Sheep Range, Cedar City; T. oreganum f. rusbyi (Greene) McDermott; T. oreganum f. bra- chypus (S. Wats.) McDermott; T. longipes var. rysbyi (Greene) Harrington]. Alpine meadows, open woods, stream banks, and grasslands at 1830 to 3500 m in Beaver, Emery, Garfield, Iron, Kane, Piute, San Juan, Sevier, and Washington cos.; Colo- rado, New Mexico, and Arizona; 47 (v). Var. reflexum A. Nels. [T. longipes ssp. reflexum (A. Nels.) J. M. Gillett; T. rydbergii Greene; T. oreganum var. rydbergii (Greene) McDermott]. Meadows, stream banks, woods, and willow communities at 1870 to 3050 m in Cache, Daggett, Duchesne, Grand, Juab (Deep Creek Mts.), Salt Lake, San Juan, Summit, Uintah, Utah, and Wasatch cos.; Oregon, Idaho, Montana, Ne- vada, Wyoming, Colorado, and New Mexi- co; 30 (iv). Trifolium macilentum Greene. Lean Clo- ver. [T. kingii ssp. macilentum (Greene) J. M. Gillett]. Caulescent, 12-35 cm tall, as- cending to decumbent, from a caudex and taproot; petioles 2-16 cm long, the longest definitely basal; leaflets, 3, 14-45 mm long, 4-25 mm wide, those of basal leaves broad- ly ovoid to lance-ovoid, those of cauline leaves narrowly lanceolate, flat, toothed from near the base, glabrous on both sides, retuse to obtuse, acute to attenuate and apiculate; stipules dimorphic, the lower ones scarious and very long, the cauline ones herbaceous and small; heads nodding to suberect, without an involucre, 21-40 mm long, these glabrous; flowers many, re- flexed, violet to purplish, 13.5-17 mm long; calyx 4-5.7 mm long, pilose, the tube 2.2-3 mm long, the teeth 1.5-3.3 mm long, sub- ulate; pods 1- to 3-seeded. Mountain bnish, pinyon-juniper, and ponderosa pine commu- nities at 1370 to 2270 m in western Beaver (Ostler 1276 BRY), Iron, and Washington (type from S. Utah, Palmer 90 US) cos.; Lincoln Co. Nevada. The lean clover was subordinated by Gillett (1972) within T. kingii, on the basis of specimens from Ne- vada which were mistaken for typical T. kingii. The lean clover is a plant of dryish hillsides at lower elevations, differing fur- ther in its strongly dimorphic leaves, flow- ers which average larger, and pilose (though sparingly) calyces which average shorter. Thus, it differs from T. kingii in about the same order of magnitude as does T. erio- cephalum; 12 (ii). Trifolium nanum Torr. Dwarf Clover. Acaulescent, pulvinate-caespitose, 2-4 cm tall, from a caudex and taproot, the stems obscured by imbricated stipules and leaf bases; petioles 0.3-2 mm long; leaflets 3, 3-11 mm long, 1-5 mm wide, oblanceolate to obovate, toothed to entire, glabrous or with some hairs on the lower surface, folded or flat, acute to mucronate; stipules scarious to herbaceous; heads 1- to 4-flowered, with an involucre of distinct to connate bracts, terminal, sessile or on peduncles 0.3-4 cm long, these glabrate to glabrous; flowers 15-23 mm long, pale purplish (fading dark violet), erect; pedicels 1-2 mm long; calyx 5-7 mm long, t 1b tube 3.5-4 mm long, scarious, glabrous, the teeth 2.2-2.8 mm long, trianular-subulate, glabrous; pods 1- to 4-seeded. Alpine meadows at 2900 to 3730 m in Daggett, Grand, San Juan, Summit, and Uintah cos. (probable in Duchesne Co. also); Montana, Wyoming, Colorado, and New Mexico. The dwarf clover is one of our most distinctive, yet poorly collected, clovers. More work is necessary to elucidate its true range in Utah; 7 (0). Trifolium parryi A. Gray. Parry Clover. Acaulescent, or short caulescent and with Sept. 1978 Welsh: Utah Flora, Fabaceae 359 one elongate internode, 4-25 cm tall, from a caudex and taproot; petioles 0.6-13 cm long; leaflets 3, 5-43 mm long, 1.5-13 (16) mm wide, oblanceolate or obovate to ellip- tic or oblong, flat, toothed from near the base, glabrous on both sides, acute to obtuse and a piculate; stipules scarious to her- baceous, 6-18 mm long; heads 5- to 20- flowered, subtended by involucral bracts, terminal, on peduncles 1.8-22 cm long, these glabrous or sparingly hairy near the apex; flowers 12-17 mm long, the petals pale to dark pink-purple (fading dark vio- let), erect; pedicels 0-1 mm long; calyx 4-7.1 mm long, the tube 2-3.9 mm long, scarious, glabrous, the teeth 2-3.2 mm long, lance-subulate; pods 1- to 4-seeded. Alpine meadows, openings in spruce woods, and other coniferous woods, and on talus slopes at 2730 to 3800 m in Daggett, Duchesne, San Juan, Summit, Uintah, and Wasatch cos.; Montana, Wyoming, and Colorado. Our materials belong to var. montanense (Rydb.) Welsh stat. nov. based on T. mon- tanense Rydb., Mem. N. Y. Bot. Card. 1:263. 1900. [T. parryi ssp. montanense (Rydb.) J. M. Gillett; T. inequale Rydb., type from Bear River Canyon, Uinta Mts.]; 26 (iii). Trifolium pratense L. Red Clover. Cau- lescent, short-lived perennial, 18-60 cm tall or more, from a taproot, erect or ascending; petioles 0.8-19 cm long; leaflets 3, 11-54 mm long, 8-28 mm wide, elliptic to lan- ceolate, ovate, or obovate, flat, toothed from near the base (the teeth inconspic- uous), long-pilose beneath, glabrous above, obtuse to retuse; stipules scarious to sub- herbaceous, 10-24 mm long; head closely subtended by one or more foliose bracts, these often 3-foliolate, sessile, or spreading hairy peduncles to 3 cm long, many-flow- ered, 22-36 mm long, 20-34 mm wide, ax- illary, erect; flowers 13-20 mm long, deep red; calyx 7.5-9.7 mm long, the tube 3.2-4.1 mm long, strigose, scarious, the teeth 4.3-5.6 mm long, subulate, pilose; pods 2-seeded. Cultivated forage plant, es- caping but seldom persisting, in Beaver, Box Elder, Daggett, Davis, Garfield, Kane, Mil- lard, Salt Lake, Sevier, Summit, Utah, Wasatch, and Washington cos.; and perhaps universal; introduced from Europe; 26 (i). Trifolium repens L. White Clover. Cau- lescent, 8-35 cm tall, the stems stoloni- ferous, creeping and rooting at the nodes, the petioles and peduncles often arising at right-angles to the stem axis, radiating from a root crown; petioles 1.8-24 cm long; leaf- lets 3, 5-22 mm long, 4-18 mm wide, ob- cordate or obovate to oval or elliptic, flat, toothed from near the base, glabrous on both sides, truncate to emarginate; stipules scarious, 3-10 mm long; heads without an involucre, many-flowered, 10-32 mm long, 15-30 mm wide, axillary, on peduncles 6-33 cm long, these glabrous or sparingly pilose, erect; flowers 5-9 (10) mm long, white or pinkish, fading brown, the lower reflexed in age; calyx 3.2-5.4 mm long, the tube 2.2-2.7 mm long, scarious, glabrous, the teeth 1-2.7 mm long, subulate, glabrous; pedicels 1-6.4 mm long; pods 1- to 3-seed- ed. Cultivated forage and pasture plant, now widely established in Beaver, Box El- der, Cache, Carbon, Garfield, Grand, Kane, Piute, Salt Lake, Sanpete, Sevier, Summit, Utah, Wasatch, and Washington cos. (per- haps cosmopolitan); introduced from Eu- rope; 39 (iv). Trifolium variegatum Nutt. in T. & G. Variegated Clover. Annual, caulescent, with prostrate to erect stems 10-40 (60) cm long; petiole 0.5-4.5 cm long; leaflets 3, 3-27 mm long, 1.5-10 mm wide, obcordate to obovate or oblanceolate, flat, sharply toothed from near the base, glabrous on both sides; stipules herbaceous, ovate, laciniately toothed; heads involucrate, the involucre flaring, sau- cer-shaped, lobed and lacerate, 3- to 40- flowered, 6-20 mm broad, axillary on pe- duncles 0.8-6.5 long, these glabrous; flowers 5-12 (20) mm long, purplish, often white- tipped, fading brown, ascending to erect; calyx 5- to 20-nerved, the teeth subulate-setaceous, glabrous; pedicels very short; pods 1- to 2-seeded. Stream banks and roadsides, at 1300 to 1600 m in Cache, Salt Lake, and Weber cos.; British Colum- bia and Montana south to California and Nevada. Our only annual clover species is not common in Utah. Trifolium wormskjoldii Lehm. Worm- skjold Clover. [T. involucratum Ortega, not Lam.; T. willdenovii Spreng.; T. fimbriatum Lindl.; T. involucratum var. fi7nbnatum 360 Great Basin Naturalist Vol. 38, No. 3 (Lindl.) McDermott; T. willdenovii var. fim- hriatum (Lindl.) Ewan; T. spinulosum Douglas ex Hook.; T. heterodon Torr. & Gray.] Caulescent, 12-35 cm tall or more, from a taproot, or sometimes rooting at de- cumbent stem bases, erect or ascending; petioles 1.2-4 cm long; leaflets 3, 6-30 mm long, 2-14 mm wide, oblanceolate to ellip- tic or obovate, flat, toothed from near the base, glabrous throughout; stipules herbaceous, 8-15 mm long, man-toothed; heads subtended by a toothed involucre, 20-30 mm wide, many-flowered, axillary, erect; flowers 10-18 mm long, reddish to purple; calyx 7-9 mm long, the tube 2.9-3.7 mm long, glabrous, the teeth 4.1-5.3 mm long, subulate, glabrous; pods 1- to 4- seeded. Meadows at lower middle elevations in Juab and Salt Lake cos.; British Colum- bia and Idaho south to California, Mexico, and New Mexico. This plant is rare in col- lections, yet our earliest record, that for Salt Lake Co., was collected in 1880; 2 (0). Trigonella L. Annual or short-lived perennial, caules- cent from a taproot; leaves alternate, pin- nately 3-foliate, serrate in the apical one- third to one-half; stipules herbaceous, dis- tinct, toothed or entire; flowers papilio- naceous, borne in axillary racemes or sub- umbellately disposed; calyx 5-toothed; petals 5, yellow; stamens 10, diadelphous; pods laterally compressed, much surpassing the calyx, several seeded. Trigonella corniculata (L.) L. [Trifolium (Melilotxis) corniculata L.]. Annual, caules- cent, 12-26 cm tall; stipules 3-6 mm long, laciniately toothed; petioles 0.5-4.5 cm long; leaflets 3, the terminal on a short con- tinuation of the rachis, 7-20 mm long, 2.5-16 mm wide obovate to obcordate, toothed in the apical half, sparingly pilose (mainly along veins) beneath, glabrous above, apiculate; peduncles 0.8-2.8 cm long, sparingly pilose; racemes 5- to 16- flowered, 0.8-1.4 cm long; pedicels 1-2 mm long; flowers 5.5-6.5 mm long, yellow; ca- lyx 2.8-3.2 mm long, the tube 1.4-1.8 mm long, sparingly pilose, the teeth 1.2-1.4 mm long, subulate; pods sessile, 12-15 mm long, 2-2.5 mm wide, curved, reticulately veined, the veins leaving the margins at about right-angles, glabrous. Introduced revegeta- tion plant, not known to persist, but to be expected, now known only from Sanpete Co.; introduced from Europe. A second spe- cies of Trigonella is known from Sanpete Co., also, but its identity has not been es- tablished. It differs from T. corniculata in its flowers being 10-12 mm long and sub- umbellately arranged. Further study is in- dicated; 2(0). ViciA L. Annual or perennial herbs, clambering, trailing, or climbing; leaves alternate, even- pinnately compound, the rachis terminating in a usually prehensile tendril; stipules her- baceous, entire to semi-sagittate; leaflets 4-12 or more, very variable; flowers soli- tary, axillary, or in axillary racemes, papilio- naceous; calyx 5-toothed, obliquely cam- panulate to short-cylindric; petals 5, pink to white, the wings adnate to the keel; sta- mens 10, diadelphous; style filiform, beard- ed around the circumference below the stigma; pods oblong, 2- to several-seeded, the valves coiling upon dehiscence. Reference Herman, F. J. 1960. Vetches of the United States— native, naturalized, and culti- vated. U.S.D.A. Agr. Handb. 168: 1-84. 1. Flowers 15 or more in dense secund racemes; introduced plants of cultivated lands and other disturbed sites V. villosa - Flowers 10 or fewer, in secund racemes or otherwise; introduced or indigenous plants 2 2(1). Flowers 5-8 mm long; plants very slender, indigenous in southern Utah V. ludoviciana Sept. 1978 Welsh: Utah Flora, Fabaceae 361 - Flowers 12-25 mm long or more; plants not very slender, indigenous and widespread or cultivated 3 3(2). Flowers 3-10 in pedunculate racemes; plants indigenous, widespread V. americana Flowers 1-3, sessile or very shortly pedunculate in leaf axils; plants uncommon, cultivated V. faba Vicia americana Muhl. ex Willd. Ameri- can Vetch. Perennial, 12-127 cm tall, the stems glabrous or pubescent; stipules 3-10 mm long, semi-sagittate, deeply toothed in the lower portion; leaves (excluding tendrils) 2-3 cm long; leaflets 8-16, 3-44 mm long, 1-19 mm wide, linear, elliptic, oblong, ovate, lanceolate, oblanceolate, or obovate, glabrous to pubescent, acute to truncate, rounded, or retuse and apiculate, less com- monly toothed apically; tendrils branched or simple; peduncles 1.8-6.7 cm long; racemes 3- to 7 (10) -flowered, the flowers spreading in anthesis; calyx 6.2-8.4 mm long, the tube 4.8-6.5 mm long, the lowermost tooth 0.7-1.9 (2.5) mm long, triangular; flowers 13-22 (25) mm long, pink to pink-purple; pods stipitate, the stipe 2.5-4.5 mm long. the body 23-35 mm long, 6-8 mm wide, glabrous. This widespread, indigenous vetch is extremely variable with regard to leaflet length-width ratios and shape. Thickness of the leaflets and pubescence also varies con- siderably. Several subordinate taxa have been based on these variations, but contin- ued recognition seems possible only when diagnostic criteria are arbitrarily applied, and even then with great difficulty, espe- cially in plants with dimorphic leaves. More importantly, much of the variation seems to be ecologically influenced, and further rec- ognition of most of the types seems unwar- ranted. Therefore, it seems best to recog- nize only two infraspecific taxa, the one widespread and common, the other restrict- ed and rare. 1. Leaflets 20-40 mm long, 2-5 mm wide, coriaceous, pubescent with short curved hairs; lateral veins prominent, leaving the midrib at a narrow angle; plants of Uintah Co V. americana var. minor Leaflets 3-44 mm long, 1-19 mm wide, thin or glabrous, or both; lateral veins prominent or not, leaving the midrib at a wide angle; plants through- out Utah V- americana var. americana Var. americana. [V. oregana Nutt. in Torr. & Gray; V. americana var. oregana (Nutt.) A. Nels. in Coult. & Nels.; V. ameri- cana ssp. oregana (Nutt.) Abrams; V. trun- cata Nutt. in Torr. & Gray; V. americana var. truncata (Nutt.) Brew, in Brew. & Wats.] Sagebrush, pinyon-juniper, mountain brush, ponderosa pine, aspen, and spruce-fir communities at 1270 to 3050 m in Beaver, Carbon, Cache, Davis, Duchesne, Emery, Garfield, Iron, Juab, Kane, Millard, Morgan, Rich, Salt Lake, San Juan, Sanpete, Sevier, Summit, Tooele, Utah, Wasatch, Washing- ton, and Weber cos.; British Columbia east to Ontario and south to Mexico, Arizona, New Mexico, Kansas, Missouri, and Vir- ginia; 167 (xxxv). Var. minor Hook. [V. sylvatica Nutt.; V. sparsifolia Nutt. in Torr. & Gray; Lathyrus linearis Nutt. in Torr. & Gray; V. linearis (Nutt.) Greene; Lathyrus dissitifolius Nutt. in Torr. & Gray; V. dissitifolius (Nutt.) Rydb.; V. americana var. angustifolia Nees in Wied.; V. caespitosa A. Nels.; V. trifida Dietr.] Pinyon-juniper and mixed desert shrub communities at 1670 to 1830 m in Uintah Co.; Alberta east to North Dakota and south to Colorado and Texas; 1 (0). Vicia faba L. Broadbean. Annual, 40-100 cm tall or more, the stems glabrous; stipules 4-15 mm long, semi-sagittate, not or only somewhat toothed; leaves (excluding ten- drils) 6-14 cm long; leaflets 2-6, 30-75 (100) mm long, 13-40 mm wide, ovate- 362 Great Basin Naturalist Vol. 38, No. 3 lanceolate to elliptic or obovate, glabrous or with short curved hairs on veins and mar- gin, obtuse to acute and apiculate; tendrils unbranched; peduncles very short or obso- lete; racemes 1- to 4-flowered, the flowers erect-ascending; calyx 12-16 mm long, the short-cylindric tube 7.5-9.5 mm long, the lowermost tooth 3.5-6.5 mm long, lance- subulate; flowers 25-31 mm long, white, spotted with maroon or blackish-violet; pods subsessile to substipitate, the body 70-130 mm long or more, 10-30 mm wide, glabrous. Sparingly cultivated vegetable plant, mainly for the large edible seeds, known from Wasatch Co., but to be ex- pected in most communities in the state; in- troduced from Asia; 1 (1). Vicia ludoviciana Nutt. Louisiana Vetch. [V. exigua Nutt. in Torr. & Gray; V. thur- beri S. Wats.; V. producta Rydb.] Annual or winter annual, 30-85 cm tall, the stems glabrous or puberulent; stipules semi-hastate to linear-oblong, 1-4 mm long; leaves (ex- cluding tendrils) 1.8-5.5 cm long; leaflets 6-10, 7-28 mm long, 0.6-4 mm wide, linear to oblong or oblanceolate, pilosulous to glabrous, obtuse to acute and mucronate; tendrils branched and prehensile; peduncles 0.4-4.8 cm long; racemes 1 (2) -flowered, the flowers ascending to spreading at an- thesis; calyx 2.2-3.3 mm long, the cam- panulate tube 1.4-1.9 mm long, the lower- most tooth 0.8-1.4 mm long, lance-subulate; flowers 6.3-7.4 mm long, lavender; pods stipitate, the stipe 0.8-1.4 mm long, the body 16-28 mm long, 5-6.2 mm wide, glabrous. Blackbrush, creosote bush, and pinyon-juniper communities at 900 to 1730 m in San Juan and Washington cos.; Ore- gon, California, Nevada, Arizona, New Mex- ico, Colorado, and Texas; 8 (i). Vicia villosa Roth. Hairy Vetch. Annual or biennial, 50-200 cm tall, the stems spreading-hairy; stipules toothed or entire, 5-15 mm long; leaves (excluding tendrils) 2.3-8 cm long; leaflets 10-18, 8-30 mm long, 1-6 mm wide, linear to oblong or nar- rowly lanceolate, long-pilose or hirsute on both sides, acute to obtuse and apiculate; peduncles 1.8-7.5 cm long; racemes mainly 15- to 25-flowered, the flowers declined at anthesis; calyx 7-7.8 mm long, the gibbous tube 3.8-^.7 mm long, the teeth 3.1-4.3 mm long, subulate, pilose; flowers 15-17 mm long, pink-purple or reddish-violet; pods 20-30 mm long, 7-10 mm wide, glabrous. Weedy introduction of cultivated lands and other disturbed sites, often along fence-rows, in Salt Lake, Utah, Washington and Weber cos., but to be expected else- where; adventive from Europe; 5 (0). Wisteria Nutt. Twining woody vines, imarmed; leaves al- ternate, odd-pinnate with 7 or more leaflets; stipules ovate to lance-linear, caducous; pe- dimcles opposite the leaves; flowers in elon- gate, pendulous racemes, calyx turbinate- campanulate, 2-lipped, the upper lip shortly 2-toothed, the lower 3-toothed; corolla pa- pilionaceous, the banner strongly reflexed; stamens 10, diadelphous; petals white, pink, lavender, purple, or blue; pods stipitate, lat- erally flattened, constricted between the seeds. Longest pedicels 12-28 mm long or more; calyx lobes very short; pods silky- to velvety-hairy W. florihunda Longest pedicels 6-11 mm long; calyx lobes almost as long as the tube; pods glabrous ^- macrostachya Wisteria floribunda (Willd.) DC. Japa- nese Wisteria. [Glycine floribunda Willd.; Kraunia floribunda (Willd.) Taub.; W. macrobotrys Sieb. & Zucc] Twining vines to several meters long; leaves petiolate, 9-35 cm long; leaflets 9-15, 13-90 mm long, 9-30 mm wide, ovate to lance-oblong. long-attenuate, pilose to glabrate or glabrous on both sides; peduncles 0.5-5 cm long; racemes pendulous, many-flowered, 25-50 cm long; flowers 15-20 mm long, white, pink, purple, or lavender; calyx tube strigulose, 3.5-4.3 mm long, the upper teeth almost completely connate, the lowermost Sept. 1978 Welsh: Utah Flora, Fabaceae 363 tooth 1.3-2.1 mm long, triangular; pods 100-180 mm long, 15-18 mm wide, taper- ing to the base, velvety hairy in lines. Culti- vated ornamental of great beauty in both grafted and free growing forms in Utah Co., and likely in most low elevation population centers in the state; introduced from Japan; 3 (ii). Wisteria macrostachya Nutt. Nuttall Wisteria. Freely twining vines to several meters long; leaves 15-40 cm long; leaflets 9-13, 28-90 mm long, 9-33 mm wide, ovate to lance-oblong, long-acuminate, pi- lose along the veins beneath at maturity, pi- lose to glabrate or glabrous otherwise; pe- duncles 1-5 cm long; racemes pendulous, many-flowered, 22-55 cm long; flowers 18-22 mm long, lilac-purple; calyx tube pilosulous, 4.5-5.5 mm long, the upper teeth cleft, the lowermost tooth 5.5-6.5 mm long, ovate-acuminate; pods 70-120 mm long, 11-15 mm wide, oblong, glabrous. Or- namental of charm and beauty in Utah Co., and likely elsewhere in Utah; introduced from the southeastern United States; 3 (iii). Index of Names and Synonyms Acacia 231 greggii 231 Albizia 231 julibrissin 231 Amorpha 231 canescens 232 fruticosa 232 Anisolotus longebractcatus 320 numularius 320 rigidus 320 wrightii 321 Anthyllis 232 vulneraria 232 Arachis 232 hypogaea 2,32 Aragallus besseyi 337 bigelovii 338 knowltonii ,338 lambertii 340 sericeus 340 viajusculus 340 metcalfei 338 minor 338 multiceps 338 minor .338 oreophilus ,339 parryi 339 patens ,338 viscidus 340 Aspalathus caragana 307 Astragalus 232 aboriginum 264 adanus 261 adsurgens 261 robust ior 262 agrestis 262 polijspermus 262 alpinus 262 amphioxys 262 amphioxys 263 cymbellus 273 melanocalyx 263 vespertinus 263 ampullarius 263 angustus 268 ceramicus 268 pictus 268 araneosus 285 aretioides 264 arcttis 297 arietinus 270 argillosus 278 argophyllus 263 argophyllus 264 cnicensis 264 martinii 264 panguicensis 264 pephragmenoides 264 arrecttis 277 eremiticus 277 artemisiaru77i 265 artipes 293 asclepiadoides 264 australis 264 bamebyi 265 beckwithii 265 beckwithii 265 purpeueus 265 bigelovii 290 thompsonae 290 bisontum 338 minor 338 bisulcatus 266 bisulcatus 266 haydenianus 266 major 266 bodinii 266 yukonis 266 brandegei 267 brevicaulis 268 bryantii 267 caespitosus 301 callithrix 267 calycosus 268 calycosus 268 mancus 268 scaposus 268 campestris 3.37 diversifolius 274 candensis 268 brevidens 268 canadensis 268 candicans 268 canovirens 271 carltonii 290 carolinianus 268 castaneiformis 271 consobrinus 271 ceramicus 268 pictus 268 chamaeleuce 269 panguicensis 264 chloodes 270 cibariiis 270 cicadae 269 laccoliticus 269 cicer 270 coltonii 271 aphyllus 271 coltonii 271 foliosus 271 moabensis 271 confertiflorus 279 flaviflorus 279 consobrinus 271 convallarius 272 convallarius 272 diversifolius 274 finitimus 272 cottamii 272 coulteri 285 fremontii 285 cronquistii 272 curtilobus 303 cymboides 273 dasyglottis 262 debilis 266 decumbens 289 oblongifolius 289 deflexus 337 desereticus 273 desperatus 273 conspectus 265 petrophilus 273 detritalis 274 deihlii 279 diphysus 285 albiflorus 285 diversifolius 274 junceus 272 roborum 272 dodgeanus 305 drummondii 274 duchesnensis 275 eastwoodae 275 elegans 277 emoryanus 275 ensiformis 276 gracilior 276 episcopus 276 eremiticus 277 eucosmus 277 eurekensis 277 falcatus 278 fallax 280 familicus 280 flavus 278 argillosus 278 candicans 278 flavus 279 flexuosus 279 diehlei 279 flexuosus 279 fremontii 285 fucatus 279 garrettii 290 geyeri 279 gilviflonis 280 glareosus 298 goniatus 262 gracHentus 280 fallax 280 hallii 280 364 Great Basin Naturalist Vol. 38, No. 3 fallax 280 hamiltonii 280 harrisonii 281 haydenianiis 266 major 266 henrimontanensis 281 humistratus 281 humivagans 282 hylophilus 289 ublongifolius 289 hypoglottis 262 polyspermus 262 ibapensis 274 impensis 283 inflexus 298 glareosus 298 iodanthus 282 diaphanoides 282 iselyi 282 jejunus 282 jessiae 283 junceus 272 attenuatus 272 diversifolius 274 orthocarpus 274 juncifonnis 272 kaibensis 276 kentrophyta 282 coloradoensis 283 elatus 283 jessiae 283 im plexus 283 rotundus 283 lambertii 338 bigelovii 338 lancearius 283 lentiginosus 284 albiflorus 285 araneosiis 285 chartaceus 285 diphysus 285 fremontii 285 palans 285 platyphyllidius 286 salinus 286 scorpionis 286 stramineus 286 ursinus 286 vitreus 286 wahweapensis 286 limnocharis 287 loanus 287 lonchocarpus 287 lutosus 288 macer 287 malacoides 288 mancus 268 marcus-jonesii 263 marianus 288 megacarpus 289 caulescens 293 parryi 289 prodigus 289 minthorniae 276 gracilior 276 miser 289 oblongifolius 289 tenuifolius 290 missouriensis 290 amphibolus 290 moencoppensis 290 mollissinius 290 montanus 28.3 coloradoensis 283 impensus 283 tegetarius 283 montii 291 monumentalis 291 mortonii 268 brevidens 268 inulticaulis 297 mtdtiflorus 303 munzii .339 musiniensis 291 newberryoides 291 nelsonianus 291 newberryi 292 castoretis 292 wardiantis 287 nidularius 292 nitidiis 261 robustior 261 nuttallianus 292 imperfectus 29.3 micranthiformis 293 oophorus 293 artipes 293 caulescens 293 lonchocalyx 293 oreophihis 339 oroboides 271 americanus 211 palans 285 araneosus 285 panguicense 264 pardalinus 29.3 parryanus 339 pattersonii 294 praelongtis 296 peabodianus 297 pectinatiis 291 platyphyUus 291 perianus 294 pictus 268 angtistus 268 foliolosus 268 magnus 268 pinonis 295 platytropis 295 praelongus 295 ellisiae 296 lonchopus 296 praelongus 296 preussii 296 arctus 297 eastwoodae 275 /afus 297 laxiflonis 297 Uixispicatus 297 preussii 297 sulcatus 275, 297 procerus 296 .pubentissimus 297 peabodianus 297 pubentissimus 297 purshii 297 glareosus 298 interior 298 purshii 298 pygmaetis 269 laccoliticus 268 racemosus 298 treleasei 298 rafaelensis 298 sabinarwn 264 sabulonum 299 sabulosus 299 salinus 286 saurinus 299 scaposus 268 scobatinatulus 266 scopulorum 300 sericoleucous 264 arctioides 264 serotinus 272 campestris 272 serpens 300 sesquiflonis .300 brevipes .300 short ianus 263 brachylobus 303 minor 26.3 sileranus 302 cariacus .302 simplex 301 simplicifolius 301 spatulatus 301 spatulatus 301 simplex 301 uniflorus 301 spectabilis 274 stocksii 281 straturensis .301 striatiflorus 301 striatus 261 subcinereus 279, .302 basalticus 302 subcinereus 303 syrticolus 290 tegetarius 283 e/a .. L. M L. I^rcant Sal Moisture Wklw Addad by Irrigation I , I JAN FEB MAR APR MAY JUN 1976 JUL AUG SEP OCT NOV OEC Fig. 1. Precipitation pattern, irrigation regime, and comparative soil moisture data for irrigated and nonirri- gated pasture plots, Provo, Utah, 1976. December 1978 BuLLiCK, Andersen: Nematode Biology 373 & 20 - - - ' 0 25 50 75 100 125 150 175 Precipitation (mm| 200 225 250 27S Fig. 2. Bioclimatograph of monthly mean maximum temperature vs. precipitation for irrigated and nonirrigated pasture plots, Provo, Utah, 1976. 4 7 14 21 26 35 42 49 56 63 Da,. ■v 100 ^^^^^^^^^ ___^ S '0 .; N .,,'-'' ^'^^^^-^^ 5 01 , -^^_ i 100 0 N S 100 V CN.™«, 0 i '0 :/ ~~~- ----__ ^^TT^^ 1 . . . 2 4 ' 4 21 28 Days 42 49 56 63 100.0 ■\^ 10.0 . \^ -- --\ IBR.GAIE ° 10 • / ^v .-'-' 0.1 ■' 1 ;: •1-,.™ 1 001 1000 r 1O0 A °" ■""'"■ no 10 • Ky, ^^^>\ -"-""^ ai -. _.----^^ \ -J- j_ . ^ ■ 26 35 42 49 56 63 Fig. 3. Recovery of H. contortus third-stage larvae from irrigated and nonirrigated pasture plots. Experiments 1-6. 374 Great Basin Naturalist Vol. 38, No. 4 '\,^-"^ 1 wo ^ — "--=2:, o.., .... 1 -KIXflD n. i..n lo, .»r I ''" .,-.,-—- „.^o. .. ^^^ — 1 »< ■/ 1 II' ; 1 1 Ituirio 14 itM>t'» 1 , \ 1 lao ■,-^ ' . . . " -^ ^"^ ~~^^ 2 4 7 21 28 3 5 <2 49 56 eo Fig. 4. Recovery of H. contortus third-stage larvae from irrigated and nonirrigated pasture plots, Experiments 7-12. I o J too •-N S^ro .m. a. » «)„ .~t. ■/[ ._, /T^^^^^^ ^ ■n-- -^ "^^^^.^ IRRIGATED ..... 1 \ V ^^ .^-....0 ■ *s N, / ^"''^^^^^^^ Fig. 5. Recovery of H. contortus third-stage larvae from irrigated and nonirrigated pasture plots, Experiments 13-18. December 1978 BuLLicK, Andersen: Nematode Biology 375 t test for paired samples, t = 5.58; P<0.01) from the irrigated than from the nonirri- gated section. The highest average recovery from the irrigated section at that time inter- val was from experiments started in Sep- tember (21.9 percent) and lowest from ex- periments started in July (9.1 percent). The highest average recovery from the nonirri- gated section after seven days' exposure was from experiments started in May (8.3 per- cent) and lowest from experiments started during August (1.3 percent). Irrigation also appeared to affect the mi- gratory ability of the infective larvae after the pellets had once been placed on the plots. After approximately seven days, lar- vae recovered from the irrigated plots were generally found in greater percentages from the glass clippings, whereas larvae recov- ered from the nonirrigated plots were gen- erally found in greater percentages in the soil and remaining pellets (Figs. 3-7). Discussion Ecological studies on the free-living stages of trichostrongylid nematodes have been reported by such workers as Stewart and Douglas (1938), Levine (1963), Kates (1965), Andersen et al. (1970), Levine and Andersen (1973), Williams and Bilkovich (1973), Levine et al. (1974), Gibson and Everett (1976), and Todd et al. (1977). These individuals demonstrated that opti- mum conditions for development and/or survival differ between species of nematodes and that each particular species is limited generally to a imique climatic environment. Gordon (1953) reported that H. contortus flourishes in climatic regions where average monthly maximum temperatures exceed 18.3 C and where total monthly precipi- tation is above 50 mm. In our study, the addition of water by irrigation was suf- ficient to provide a total monthly water ac- cumulation well in excess of 50 mm. Also, the resultant increase in soil moisture helped maintain microenvironmental tem- peratures within the optimum limits pre- scribed above. As a result, environmental conditions on the irrigated plots were with- in the range for optimum pasture transmis- sion of H. contortus for five of the six V 100 ■ >=^-:,^,^..,....,^KO;^;».;»^.»,, -. . .: — ° 10 ■ / " " - - Cj '__, .„„.,~~,llr. „ "~ ~^^^^^^> ot ■/ |,,..,;;::;„,„„,J 1000 lao E\ --- ,.0 '- ' \ ~~--~^,_^^ ^^^...-^ -^--.^^^ 01 ^ 2 4 7 14 21 28 35 42 49 56 63 Days \ 100 ■\ 10 ^TT^^^ ''"^^^ " \....:;: .......1 ~^--~-^^ 1000 r 1Q0 \ ,ON-,«R.G TED 1.0 . V\^ 01 ~~"^^^ / . ^«. . ^^^, ^ ~. —^ 2 4 7 14 21 28 35 42 49 56 63 Oays 1000 ^~~^--^-— — -^ 10 /^^'^'^'^"^^^^"^^ ,„.,0«E " 01 ' 1 ...,'" ..,..,. \ ~"""*>^^-:^„^ 100 0 ^^^ too ■ "V-— -. °"-"""" lEO , 10 - .' ', ^"~"~--^ 01 ./ ' .-"-, \ 0 01 /.'.'. -,. \ . 56 63 1000 100 1 1,0 k ^'^^ -..^..^^^.^ ....... i 01 ...,;;-....J 3 100 0 5 100 i 10 "■ 01 ^ ^ X-\ 2 4 7 Days 49 56 63 1 '" /'[ "---- -— ^ . '" 1 01 ; 001 Z looo S 1O0 ...,;::„™..,.l N * 10 -■ 01 \ 2 14 21 28 35 4 Days 2 49 56 1 10 /[ -"^ — -^..^ ' — 1 01 > 0 01 Jj 1000 • lao i '0 ...;;■ :.=..., J \ ->-- x^ ED Fig. 19-24. 6. Recovery of H. contortus third-stage larvae from irrigated and nonirrigated pasture plots. Experiments 376 Great Basin Naturalist Vol. 38, No. 4 months of the study period. Temperatures on the nonirrigated plots, however, were usually too high for optimum larval survival and the moisture accumulation from natural precipitation alone was consistently too low. Andersen et al. (1974) previously com- pared meteorologic measurements for irri- gated and nonirrigated plots in the same area as the current study, and they demon- strated that the average monthly mean max- imum temperatures for May through Octo- ber at soil surface under grass cover for 1970, 1971, and 1972 were 23.7 and 31.1; 22.5 and 27.6; and 26.2 and 34.9 C, respec- tively. They further showed that the aver- age percent soil moisture for those same months and same years from irrigated and nonirrigated plots were 16.1 and 8.4; 16.3 and 5.8; and 8.3 and 3.8 percent, respec- tively. The comparable figures for temper- ature and soil moisture content from irri- gated and nonirrigated plots for the current study were 24.8 C and 19.0 percent, and 42.5 C and 4.6 percent, respectively. Some of the discrepancy from measurements taken at the same site during different years could have resulted from the fact that in the current study sprinkling irrigation was used to minimize lateral movements of the infective larvae, whereas flood irrigation was used in the 1970-72 study. The lowering of environmental temper- atures following irrigation has also been re- ported by such workers as DeVries and Birch (1961) in Australia, who noted a de- crease during the summer of about 10 C 100.0 10.0 1.0 0.1 2 0.01 5 100.0 • 10.0 o -■ 0.1 0.01 100.0 10.0 > $ 1.0 o u £, 0.1 2 0.01 5 100.0 • 10.0 o -^ 0.1 0.01 1 ^>^-»-l-TOTAl LARVAE RECOVERED FROM GRASS, SOIL, AND PELLETS IRRIGATED / ^ ■ 1 > ■- — =^-=-»^ r l— LARVAE RECOVERED FROM GRASS N^ -1 EXP25 \ 1 1 STARTED leOCTOBERITO . X 1 1 1 V NON-IRRIGATED /\ \^___^ - / 1 ■ — —_ ' > \ ,' , \ , , \, 1 1 1 1 2 4 14 21 28 35 42 49 56 63 Days X IRRIGATED /^\ ■ / \\ / \ V -/ EXP 26 / SIART£025OCTOBERiq76 >^»r— -=r^ "^"^^V^~^=^"=-^=r^ NON-IRRIGATED 1 1 L. 1 1 1 2 4 7 14 21 28 35 Days 42 49 56 63 Fig. 7. Recovery of H. contortus third-stage larvae from irrigated and nonirrigated pasture plots, Experiments 25-26. December 1978 BuLLicK, Andersen: Nematode Biology 377 when the temperature was measured 5 cm below grass-covered ground, and by Leon- ard et al. (1971), who observed a similar de- crease in an irrigated pine-covered forest in New York. Fowler and Helvey (1974) re- ported that air temperatures along the Co- lumbia River Basin in Washington were well below established normals now that ir- rigation is so widespread in the region. The lowering of temperatures in irrigated regions is mainly a reflection of the in- creased soil moisture resulting from the ad- dition of irrigation water. Levine and Todd (1975) stated that soil moisture is a more important criterion than precipitation alone in determining optimum conditions for de- velopment and survival of H. contortiis, since soil moisture results from an inter- action of precipitation, soil type, and evapotranspiration. Soil moisture content also is the factor most responsible for the maintenance of vegetative cover. Knapp (1964) showed that different species of for- age plants influenced the survival and also the infectivity of H. contortus in lambs. In our study, the lack of moisture on the non- irrigated section eventually killed the Ken- tucky bluegrass, which was subsequently re- placed by a few plants with deeper tap roots. These new plants did not provide ad- equate cover, and soil temperatures rose drastically during ensuing months. Con- sequently, the pellets quickly dried and the larvae were sealed within. Andersen and Levine (1968) reported that sheep pellets routinely lose 50 percent of their original weight within 12 to 24 hr when stored at 30 C and 65 to 75 percent RH, which con- ditions would approximate those on our study plot. The irrigated section of the pres- ent project site maintained ample grass cov- er for all six months of the study period and pellets placed on the plots remained moist or disintegrated as water was added. The number of days third-stage trichost- rongylid larvae may be recovered from arti- ficially infected pastures is not only depen- dent upon existing meteorologic conditions in the specific geographical location where the study occurs, but also upon the particu- lar species of nematode studied. Gibson and Everett (1976) recovered third-stage larvae of H. contortus for as long as 40 weeks in England, whereas Levine et al. (1974) re- covered 0.1 percent infective larvae of H. contortus for nine weeks in Illinois. In the present study, 0.01 percent of larvae sur- vived on the irrigated plots for a maximum of nine weeks several times during the six- month study period and at least for a min- imum of four weeks even during October, when conditions for pasture transmission were not optimum. On the nonirrigated plot, 0.01 percent of larvae were recovered for a maximum of eight weeks during May, when soil moisture content was still ade- quate. Thereafter, survival time dropped considerably until in October larvae were recovered for only two weeks. In addition to an increased larval survival time on irrigated plots in the present study, an increased percentage of larvae survived on the irrigated section. Furthermore, lar- vae on irrigated plots were recovered al- most exclusively from grass clippings after one week, whereas at that time larvae from nonirrigated plots were recovered mainly from pellets and soil scrapings. Irrigation thus aided migration of larvae from the pel- lets onto the vegetation, where they would be in the optimum position to be ingested by the grazing host under natural field con- ditions. In summary, the present study demon- strated that irrigation enhanced the ability of H. contortus larvae to survive longer and in greater numbers on experimental pasture plots, and also increased their ability to mi- grate from pellets onto adjacent vegetation. Without suitable moisture and temperature, optimum conditions for pasture transmission would not have been achieved. Irrigational practices in Utah undoubtedly contribute to the relatively high incidence of H. contortus in this region where the prevailing climatic conditions indicate the parasite could not otherwise flourish. Literature Cited Andersen, F. L., and N. D. Levine. 1968. Effect of desiccation on survival of the free-living stages of Trichostrongylus colubriforrnis. J. Parasitol. 54: 117-128. Andersen, F. L., N. D. Levine, and P. A. Boatman. 1970. Survival of third-stage Trichostrongylus cohibrifortnis larvae on pasture. J. Parasitol. 56; 209-232. 378 Great Basin Naturalist Vol. 38, No. 4 Andersen, F. L., and P. R. Roper. 1975. Com- puterized reduction of meteorologic measure- ments from irrigated and non-irrigated plots in Central Utah. Great Basin Nat. 35: 20,3-230. Andersen, F. L., P. D. Wright, and J. C. Fox. 1974. A comparison of meteorologic measurements from irrigated and nonirrigated plots, Provo, Utah, 1970-1972. Brigham Young Univ. Sci. Bull., Biol. Ser. 19: 1-37. Devries, D. a., and J. W. Birch. 1961. The modifica- tion of climate near the ground by irrigation for pastures on the Riverine Plain. Aus. J. Agr. Res. 12: 260-272. Fowler, W. B., and J. D. Helvey. 1974. Effects of large scale irrigation on climate in the Colum- bia Basin. Science 184: 121-127. Fox, J. C, F. L. .\ndersen, and K. H. Hoopes. 1970. A survey of the helminth parasites of cattle and sheep in Utah Valley. Great Basin Nat. 30: 131-145. FuRMAN, D. P. 1944. Effects of environment upon the free-living stages of Ostertagia circumcincta (Sta- delmann) Trichostrongylidae: II. Field Experi- ments. Am. J. Vet. Res' 5: 147-153. Gibson, T. E., and G. Everett. 1976. The ecology of the free-living stages of Haemonchus contortus. Br. Vet. J. 132: 50-59. Gordon, H. M. 1953. The epidemiology of helmin- thosis in sheep in winter-rainfall regions of Aus- tralia. I. Preliminary observations. Aust. Vet. J. 24: 17-45. HoNESS, R. F., and R. C. Bergstrom. 1966. Tricho- strongylosis of cattle in Wyoming. Sci. Mon- ograph 2, Agr. Expt. Sta., Univ. Wyoming, Lar- amie, Wyoming. Irrigation Handbook and Directory. 1972. Irrigation Data, Inc., Lincoln, Nebraska. 327 p. Kates, K. C. 1965. Ecological aspects of helminth transmission in domesticated animals. Amer. Zool. 5: 95-130. Knapp, S. E. 1964. Relationship of different species of forage to the survival and infectivity of Hae- monchus contortus in lambs. J. Parasitol. 50: 144-148. Leonard, R. E., A. L. Leaf, J. V. Berglund, and P. J. Craul. 1971. Annual soil moisture-temperature patterns as influenced by irrigation. Soil Sci. 3: 220-227. Levine, N. D. 1963. Weather, climate, and the bio- nomics of ruminant nematode larvae. Adv. Vet. Sci. 8: 215-261. Levine, N. D., and F. L. Andersen. 1973. Devel- opment and survival of Trichostrongylus coluhri- fortnis on pasture. J. Parasitol. 59: 147-165. Levine, N. D., and K. S. Todd, Jr. 1975. Micro- meteorological factors involved in development and survival of free-living stages of sheep nema- todes Haemonchus contortus and Tricho- strongylus cohibriformis. Int. J. Biometeor. 19: 174-183. Levine, N. D., K. S. Todd, Jr., and P. A. Boatman. 1974. Development and survival of Haemonchus contortus on pasture. Am. J. Vet. Res. 35: 1413-1422. Shore, D. A. 1937. A method of separating infective larvae of Haemonchus contortus (Trichostrongy- lidae) from free-living nematodes. Proc. Helm. Soc. Wash. 4: 52. Stewart, M. A., and J. R. Douglas. 1938. Studies on the bionomics of Trichostrongyhis axei (Cob- bold) and its seasonal incidence on irrigated pastures. Parasitology 30: 477-490. Surtees, G. 1970. Large-scale irrigation and arbovirus epidemiology, Kano Plain, Kenya. 1. Description of the area and preliminary studies on the mos- quitoes. J. Med. Ent. 7: 509-517. Todd, K. S., Jr., N. D. Levine, and B. N. Wagher. 1977. Effects of repeated desiccation on survival of infective Cooperia punctata larvae. J. Para- sitol. 63: 956-957. Utah Agricultural Statistics. 1976. Utah State De- partment of Agriculture, Salt Lake City, Utah. 104 p. Williams, J. C, and F. R. Bilkovich. 1973. Distribu- tion of Ostertagia ostertagi infective larvae on pasture herbage. Am. J. Vet. Res. 34: 1337-1.344. World Health Organization. 1950. Joint study group on bilharziasis in Africa. Rep. 1st Sess. Tech. Rep. 17: 16. Wright, P. D., and F. L. Andersen. 1972. Parasitic helminths of sheep and cattle in central Utah. J. Parsitol. 58: 959. MOUNTAIN ANTS OF NEVADA George C. Wheeler' and Jeanette Wheeler' Abstract.- Introductory topics include "The High Altitude Environment," "Ants Recorded from High Alti- tudes," "Adaptations of Ants," "Mountain Ants of North America," and "The Mountains of Nevada." A Nevada mountain ant species is defined as one that inhabits the Coniferous Forest Biome or Alpine Biome or the ecotone between them. A table gives a taxonomic list of the mountain ants and shows the biomes in which they occur; it also indicates whether they occur in lower biomes. This list comprises 50 species, which is 28 percent of the ant fauna we have found in Nevada. Only 30 species (17 percent of the fauna) are exclusively montane; these are in the genera Mymiica, Manica, Stenamma, Leptothorax, Camponottis, Lasiiis, and Formica. The article concludes with "Records for Nevada Mountain Ants." All known records for each species are cited. For each record we give first the county, next the mountain range, then the peak (or other local feature), and finally the elevation. Epigraphs "The first compilation of world ants found at elevations of 2000 m (6560 ft) or more shows that, while many species may be found at the 2000 m level, the numbers decrease rapidly with increase in altitude. Few ants are found at 3000 m (9840 ft), and at 4000 m (13,129 ft) or more only nine species are known. The world altitudinal re- cord is of Formica picea lockmatteri Stiircke at 4800 m. (15,740 ft) in the Himalayas" (Weber 1943: 351). "While other branches of entomology have made great advances in recent years, our knowledge of the insect life of the North American mountains is, however, ex- tremely fragmentary" (Mani 1968: 365). Acknowledgments As soon as we moved to Nevada in 1967 we began a study of the ants of the state. Most areas were readily accessible, thanks to an excellent highway system and a Jeep Wagoneer. But we soon found that we were too old for hiking and backpacking in the rarified air of the mountaintops. To remedy this handicap, we applied for and were awarded two National Science Foundation grants to employ students to collect for us. The first was for the Alpine Biome (BMS74- 13679). For this study we hired Alvin McLane (one of Nevada's most experienced mountaineers) and Jane Ram- burg (a senior botany major at Wellesley College) for two months during the summer of 1975. The second grant (DEB76-11131) was for the upper levels of the Coniferous Forest Biome. For this study we hired two University of Nevada, Reno, students, Gary Nigro (a graduate entomology student) and Wendy Guyer (a senior botany major) for two months during the summer of 1977. The above should not be taken to mean that we have had no personal experience with high-altitude ants. We have taken ad- vantage of the excellent gravel road up to 12,000 ft on Mt. Grant (elevation 12,200 ft) near Hawthorne and a passable road up to 10,000 ft on Mount Jefferson (elevation 11,949 ft). There are also many roads up to 9000 ft, but these one-lane roads without turnouts generally lead to mines. We thank Alvin McLane for reading the manuscript. Definition In his book, Mani (1968:8) preferred the term "hypsobiont" or "high altitude" to "al- pine" and gives this definition: "The high altitude insects may thus be described as an ecologically highly specialized, mountain 'Adjunct Research Associates, Desert Research Institute, Reno, Nevada 379 380 Great Basin Naturalist Vol. 38, No. 4 autochthonous group existing exclusively in the biome above the forest, at elevations above 2000-2500 in." None of our Nevada ants can qualify because of the adverb ex- clusively, so we will use the more modest term alpine. The student of mountaintop faunas must be duly warned (as does Mani, p. 4) that not all insects occurring at high altitudes are hypsobionts: "Incredibly large numbers of insects [even heavy flying insects] are lifted from the lowland by warm updraft air currents to high altitudes, to be chilled dead, blown passively and eventually cast on high mountain slopes." The converse, however, is not true: hypsobionts are rarely encountered on the lowlands. In this study we define a Nevada moun- tain ant species as one that inhabits the Coniferous Forest Biome or the Alpine Biome or the ecotone between them. We cannot use an elevational boundary because the lower limit of the forest is too uneven. High-altitude Environment The high-altitude environment, like all others, is a complex of many interrelated factors, but it differs from all others in one respect: reduced atmospheric pressure, which is itself the result of high altitude. This seems to say that the high-altitude en- vironment is characterized by high altitude, but we shall avoid being so foolish by de- scribing some of the effects of reduced at- mospheric pressure. 1. Deficient oxygen, the most important characteristic. In the Himalaya at the tim- berline the oxygen is 68 percent of what it is at sea level; at 6000 m it is only 45 per- cent. Mountain sheep, ibexes and yaks live up to 5800 m; man without an artificial ox- ygen supply lives up to 8540 m. Certain in- sects, mites, and spiders flourish at 6800 m, because they are only slightly affected by decreased oxygen or by sudden changes in atmospheric pressure (Mani 1968:10). 2. Atmospheric cold. While it is true that cold does slow down the activities of in- sects, high-altitude insects can exist only be- cause of the atmospheric cold: it enables them to withstand the atmospheric aridity (Mani 1968:22-23). 3. Atmospheric aridity. 4. Intense solar radiation. "Regardless of atmospheric temperature, objects exposed to direct sunshine warm up far more rapidly than at sea level" (Mani 1968:21). This is enormously important for insects because of the short days and the short summers. 5. Snow cover. This is absolutely essential for high-altitude insects. It prevents freezing and desiccation and, because the habitat un- der the snow is not frozen, makes possible an earlier start of summer activities. 6. Biotic factors. (After Mani 1968:44) Most biotic factors are ultimately based on the following: (a) Trees are absent. (b) The scant cespitose vegetation has a short growing period. (c) The active feeding period is severely restricted by the short summer. In the northwestern Himalaya on south slopes the average annual feeding pe- riod may last 10 weeks at 3000-4000 m. On north slopes it starts later and is shorter. (d) Sources of food are extremely irregu- lar, relatively scant, and often local- ized. (1) Autochthonous sources are plants and animals normally living at high altitudes. (2) Wind-blown organisms from the lowlands are the predominant source and are most abundant at the melting edges of snow where dead plants and animals (mostly insects) become exposed. The sur- face of the snow is likewise im- portant; it is almost the exclusive source in the Himalaya above 5000 m. (e) Suitable microhabitats are scarce. Ac- tually there are only two of any sig- nificance: (1) cracks in the soil and rocks and (2), of far greater impor- tance, under stones. (f) Crowding and isolation, caused by this scarcity, may result in "a state of . . . armed neutrality rather than peaceful coexistence!" (p. 81) (g) The majority of high-altitude insects are predators, parasitoids, or parasites. "It would seem that almost every December 1978 Wheeler, Wheeler: Nevada Ants 381 member of a high-altitude commimity spends practically all its time devour- ing and predating [sic] on every other member species" (pp. 80-81). (h) The base of the ecological pyramid is CoUembola. (i) The fauna is impoverished in number of species (perhaps to only three of four in a community), but the number of individuals per species may be very large. Ants Recorded from High Altitudes Weber (1943:341-346) has assembled a list of records with locations and elevations in meters and in feet. The following totals include only workers recorded above 2000 m ( = 6560 ft), except in North America where for some unexplained reason he in- cludes only those at or above 10,000 ft: the Himalaya 37 species, other Asiatic records 15, Alps 1, North America 16, South Ameri- ca 12, Africa 62. Gregg (1963) recorded for Colorado 33 species at or above 10,000 ft. In Nevada there are 19 species at or above 10,000 ft. Formica picea lochmatteri, which is found at an elevation of 15,749 ft in the Himalaya (Weber 1943:351), is the world's highest known ant. The Nearctic champion is ap- parently Formica neorufibarbis, which has been taken at 14,260 ft on the summit of Mt. Evans in Colorado (Gregg 1963:533). The Nevada champion is likewise Formica neorufibarbis, taken on Boundary Peak in Esmeralda County at 12,160 ft. (The sum- mit of Boundary Peak is the highest eleva- tion in Nevada: 13,145 ft.) Deserving special mention here is Tapi- noma sessile. Creighton (1950:353) gave its range as "southern Canada and the entire United States with the exception of desert areas in the southwest. The incidence of sessile appears to decrease sharply in the Gulf Coast region but it has been taken in Florida, Alabama, Mississippi and Texas." We collected it three times in Deep Can- yon near Palm Desert, California (Wheeler and Wheeler 1973:106), which is in the Colorado desert. "The ants have been found to nest all the way from sea level to heights of over 10,000 feet" (Smith 1928:319). Gregg (1963:446) reported it up to 10,505 ft in Colorado. So the Nevada record of 11,320 ft on Boundary Peak must be the highest not only in Nevada but also any- where the species is found. Adaptations Because the Arctic-alpine is the harshest terrestrial environment on earth, one may ask what special adaptations permit certain species to live and even thrive in it: 1. Pigmentation. The insects of high alti- tudes have a large amount of melanin in their integument. The black color enables them to warm up faster and earlier in the morning as well as earlier in the season. This ensures them a longer working period during the all-too-short summer. 2. Atrophy of wings. This enables them to stay in their suitable environment in spite of violent winds. (This does not apply to ants.) 3. Prolonged hibernation. Such species hi- bernate most of the year and sometimes for two years. 4. Subsurface life. Few species live on the surface. (This does not apply to ants, whose workers may be very active on the surface when conditions are favorable.) 5. Increased clothing. Hairs, scales, and wax are more abundant. 6. Cold stenothermy. High altitude in- sects are usually active at temperatures near freezing; this prevents dessication. Many de- velop normally at -1.5 to 5 C (31-41 F) during summer. Some will be killed in a few minutes by exposure to the warmth of a human hand. Adaptations of Ants Ants by their very nature are preadapted to life in a wide variety of environments, including some of the harshest on earth. We have discussed elsewhere (Wheeler and Wheeler 1973:7) their preadaptations to an- other harsh environment, the desert: 1. Social life: The cooperation of many individuals is advantageous anywhere in for- aging, nest construction, defense, and care of the young. 382 Great Basin Naturalist Vol. 38, No. 4 2. Nest structure: Since ants' nests are ex- cavated in the soil, they require no biologi- cally expensive building materials; they are completely flexible as to plan; they are ex- tremely efficient in that they afford a wide range of temperature and moisture condi- tions, from which the ants can select an op- timum. 3. Nocturnal activity: Many species are active both day and night and, of course, all ants are able to function in the total dark- ness of their nests. 4. Speed: Many species can nm rapidly. This would be especially useful in high alti- tudes, for there is a lot of work to be done in the short summer. 5. Omnivorous diet: The majority of ants are omnivorous; their food consists of in- sects, honeydew, seeds, and plant exudates. 6. Integument: One of the general adap- tations of insects to life in any terrestrial environment is an integument that is rela- tively resistant to water loss. It also aids in the regulation of body temperature, a very important role in small, cold-blooded ani- mals. Are there, then, any special adaptations of ants for high altitudes? Did evolution need to do any special remodeling before ants could thrive at high altitudes? The an- swer is no. But color is a preadaptation that becomes especially important at high alti- tudes. Black and red are common ant colors in all biomes, but there are many ant col- ors. Among our Alpine Biome ants, how- ever, all species are black or dark brown or a combination of red and black or red and dark brown— colors which absorb heat most rapidly, black being most efficient. Further- more the bicolored species of Formica are polymorphic: the large major workers have the head and thorax (or only the thorax) red, but the gaster is black or dark brown. The smaller workers, however, become pro- gressively more infuscated until the minims are practically black. These small minims warm up and become active earlier in the morning; the larger and redder majors begin work later. If the midday sun becomes too hot for the minims, the majors and medias can keep on working. Mountain Ants of North America Ant faunas are impoverished at high alti- tudes. Van Pelt (1963:205) found this to be tnie of the much lower mountains in the Blue Ridge Province of the southeastern United States, where the highest elevation is 6684 ft: "The number of ant forms, and in most cases the numbers of colonies, de- creased with increasing altitude." W. M. Wheeler (1917:460) made much of slope: "In mountain regions slope exposure in its relation to insolation is a very impor- tant factor in the local distribution of ants. . . . Northern slopes in the northern hemisphere are usually, for very obvious reasons, almost or quite destitude of ants. . . . [Forel] finds that ants prefer the eastern and southern slopes as these are the situations in which they have the longest day for their activities during the breeding season, since they are early awakened by sufficiently high temperatures of the soil and air from the lethargy induced by the chill night hours, and even though the slope may be in the shade during the afternoon the warmth is sufficient to sustain their ac- tivities till sunset. On western slopes, how- ever, the morning hours are too cool and are therefore practically lost to the ants, whereas the afternoon hours are too warm." Wheeler also stressed the importance of steepness (p. 462). In front of a steep slope facing the sun the heated air rises more rapidly to greater heights before it is cooled to the general temperature of the stratum it penetrates. Ants "always greatly prefer the more gradual slopes and alpine meadows, prob- ably because the soil of such places retains a more abundant and more equable supply of moisture and because their surfaces are much less exposed to rapid evaporation both from direct insolation and from air- currents" (W. M. Wheeler 1917:462). If treeless alpine environment is a harsh one, then a forest ought to be a great im- provement. But this is not necessarily so. A dense forest with a solid canopy is a hostile environment. "Ecologically significant are those areas in which no ants are found. All of them are above 6000 feet, and almost all occur within forests of spruce-fir or fir" (Van Pelt 1963:221). December 1978 Wheeler, Wheeler: Nevada Ants 383 If dessication is such a hazard to ants, then a moist environment should be favor- able. Again this is not necessarily true. Said W. M. Wheeler (1917:460): "Even moder- ately low temperatures, when coupled with considerable humidity, a condition which prevails in California during the winter months, is very unfavorable to ants, and when such conditions are most accentuated, the ant-fauna is reduced to a mere remnant, although the vegetation, if the temperature is not too low, may be luxuriant. This is the case in New Zealand where I sometimes searched in vain for an ant-colony in forests whose luxuriance rivalled those of the trop- ics. But we have a striking example of the depressing effects of cold and moisture on ant-life much nearer home. The cool Selkirk Mts. of British Columbia have an abundant supply of moisture and an unusually rich flora, but their ant-fauna is reduced to a few boreal species. The adjacent Canadian Rockies, however, though in the same lati- tude, are less humid and have a poorer flora, but their ant-fauna is decidedly richer in species and colonies." The most favorable environment for mountain ants is an opening in the forest. Here the ants can find insolation or shade, whichever and whenever needed; the cor- rect humidity may be selected; the workers can forage in the opening and/or the forest. They may nest under stones, but additional nesting sites are afforded by fallen dead trees (or branches): under bark, in solid dead wood, in rotten wood, or in the soil under the fallen trunk or branch. In the Al- pine Biome they of necessity usually nest under stones, but occasionally thatching ants construct nests with plant debris. Mountains of Nevada When we were studying geography in the grades, we visualized the Great Basin as a sort of huge washpan, the bottom a flat plain bordered by mountains. Later in phys- iography we learned that the Great Basin was a part of the Basin and Range Pro- vince, which had mountain ranges rising up from the floor. Still later, when we drove across Nevada and saw a few low ranges, we were not properly impressed. It was only after we had begun our study of the ants of the state that we were forced to the realization that Nevada is a mountainous state. We proved it to our satisfaction by exploring all parts of the state. We were es- pecially impressed when we stood on the summit of Grant Peak (12,200 ft) and view- ed in all directions numerous mountain ranges separated by basins. Perhaps the best confirmation is to view the United States Geological Survey's large (1:500,000) relief map of the state (see fig. 1). Nevada's topographical uniqueness lies in the fact that most of its surface consists of numerous (more than 300) short, isolated mountain ranges separated by basins— usual- ly called valleys both by local inhabitants and on maps. The floors of some of these basins are 7000 ft above sea level. It seem- ed strange to us at first to call anything a "valley" if its "floor" is higher than any peak east of the Rocky Mountains (includ- ing the Black Hills as part of the Rockies). Nevada's ranges are subparallel, and their axes generally approach a north-south direc- tion. They are short-50-75 miles ( = 80-120 km) long and nearly straight. Not all of them are high, but 43 ranges have peaks between 7000 and 9000 ft; another 54 are above 9000 ft and attain a maximum of 13,140 ft on Boundary Peak. The higher ranges show biotic zonation, because, with increasing altitude, temper- ature decreases and precipitation increases from 3 to 30 inches annually. The Alpine Biome (Fig. 2) may be found on the sum- mits of a few of the higher ranges above 10,000 ft. Below the Alpine is the Con- iferous Forest Biome (Fig. 3) extending from about 5000 to 11,500 ft. We are inclined to suspect that the mixed bristlecone pine {Pinits longaeva) and limber pine (P. flexilis) constitute an ecotone (Fig. 4) between the Alpine and Coniferous Forest Biomes. This is especially true of the older forests, which are widely open. The Alpine Biome is treeless and the plants consist typically of grasses, sedges and forbs. Some of the summits are devoid of plants, having just bare rocks. They are without ants. Billings (1954) recognized three sub- divisions of the coniferous forests of Ne- 384 Great Basin Naturalist Vol. 38, No. 4 Fig. 1. Nevada is a mountainous state. Courtesy of Nevada Bureau of Mines December 1978 Wheeler, Wheeler: Nevada Ants 385 vada; (1) Sierran, (2) Rocky Mountain, and (3) Great Basin. (1) The Sierran is found only in the Car- son Range a few miles south of Reno. The pine-fir zone occurs between 5000 and 7500 ft and comprises large trees from 75 to 200 ft high; it is an open forest dominated by yellow pine {Pinus ponderosa), Jeffrey pine (P. jeffreyi), and white fir {Abies concolor). There is an extensive understory of shrubs, etc. The next zone (7500-9300 ft) is domi- nated by red fir {Abies magnifica) and may include lodgepole pine (P. murrayana), western white pine (P. monticola), Jeffrey pine (P. jeffreyi), and mountain hemlock {Tsuga mertensiana) . Aspen {Populus tremu- loides) often forms groves in moister places. From about 9300 ft to the timberline (about 10,300 ft) is a semi-open patchy subalpine forest. The principal tree is whitebark pine (P. albicaulis), which diminishes in height from about 40 ft at 9300 ft to 2 or 3 ft at timberline, where it forms the characteristic krummholz. Other trees are limber pine, lodgepole pine, and mountain hemlock. (2) The Rocky Mountain subdivision com- prises the eastern ranges from the Jarbidge Mountains in Elko County to the Spring Mountains in Clark County. The complete Rocky Mountain zonal series is (in ascend- ing order): ponderosa pine; Douglas fir {Pseiidotsuga menziesii), and white fir; sub- alpine fir {Abies lasiocarpa), Engelmann spmce {Picea engelmannii), and often limber pine or bristlecone pine. In most cases one or more zones are missing or zones may be telescoped, producing mixtures. (3) In Billing's third division the ranges have the simplified forest zones of the Great Basin proper. The Pinyon-Juniper Biome reaches up to 7500-8500 ft. Above this is an almost treeless zone of sagebrush, mountain mahogany {Cercocarpus ledifolia), and other shrubs reaching 9500-10,000 ft. Above the treeless zone is an open sub- alpine forest of limber pine and bristlecone pines, which we regard as ecotone. In all zones of all three subdivisions mountain streams are bordered by aspen, Fig. 2. Alpine Biome. Foreground with typical mat vegetation. Esmeralda County: Summit of Boundary Peak in background. Photograph by Gary Nigro. 386 Great Basin Naturalist Vol. 38, No. 4 Fig. 3. Coniferous Forest Biome. Elko County: Snowslide Gulch, Jarbidge Mountains. Photograph by Gary Nigro. December 1978 Wheeler, Wheeler: Nevada Ants 387 chokecherry {Primus virginiana), water birch {Betula occidentalis), willows, and cot- tonwoods. Mountain Ants of Nevada Because our tentative list of Nevada ants totals 180 species, the list (Table 1) of 50 montane species includes 28 percent of the fauna. But only 30 species are exclusively montane— 17 percent of the total, 60 per- cent of the montane. All 50 montane spe- cies occur in the Coniferous Forest Biome. Nineteen of the montane species have been taken in the Ecotone. Five species which have been reported in the Coniferous Forest and in the Alpine have not been taken in the Ecotone, but it is reasonable to assume that they occur here; these would make a hypothetical Ecotone count of 24 species. No species is exclusively Ecotone. Fourteen species have been reported from the Alpine, none of which is exclusive to that biome. We do not find the ant fauna to be in ac- cord with Billings's subdivisions of the con- iferous forests of Nevada. Only the Sierran is at all distinctive; six species occur in the Sierra Nevada which occur no where else in the state, and conversely there are seven species that do not occur in the Sierra, but are found in many other parts of the state. Most of our montane species are too widely distributed to show any pattern. Montane species limited in Nevada to the Carson Range of the Sierra Nevada include Manica bradleyi, Stenamma ivheeleronim, Camptonotus essigi, Formica integroides, F. microphthalma, and F. sibylla. Montane species which have not been re- corded in Nevada from the Carson Range include Mynnica erneryana, M. lobifrons, Manica hunteri, Leptothorax crassipilis, Lasius vestitus, Formica hewitti, and F. sub- nuda. Table 1 shows that our montane ants are a relatively unspecialized lot. Myrmica comes first on everybody's list of Myrmi- cinae and Manica is second. Stenamma and Aphaenogaster are not far above them. But Fig. 4. Ecotone between Alpine and Coniferous Forest biomes. Open stand of bristlecone pines. Clark County: Charleston Peak, Spring Mountains. Photograph bv Gary Nigro. 388 Great Basin Naturalist Vol. 38, No. 4 what is Leptothorax doing here? It is a mod- erately specialized myrmicine. Tapinoma is somewhat specialized, but it is one of the most versatile ants on earth— it deserves its own paragraph (see above). Among the For- micinae Camponotus, Lasius and Formica rate rather low on the scale of special- ization in structure, but they rate much higher in behavior, Formica being perhaps the most plastic of all ant genera. Polyergtis is, by contrast, highly specialized as an ob- ligatory slave-maker. Apparently it can adapt to any environment in which it slaves {Formica spp.) can function, although we do not yet have it from the Alpine Biome. The list is interesting not only for what it contains, but also for what it does not con- tain: the common genera in the lower biomes of Nevada: Crematogaster, Mon- omorium, Solenopsis, Iridomyrmex, and Con- omyrrna; all the harvesters {Pogonomyrmex, Veromessor, and Fheidole); and the honey ants (Myrmecocystus). Mountain Ant Nests in Nevada Many of our records are based on stray workers only; for these we have no nest data. But those with nests are numerous enough to give a good picture of the nest- ing habits of Nevada mountain ants. In the following summary we have lumped togeth- er the data for all species. Table 1. Mountain Ants of Nevada. A = Alpine; E = Ecotone; C = Coniferous Forest Biome; -I- = lower biomes (Pinyon-Juniper and/or Cool Desert). We have only one record of a mountain ant from the Hot Desert; even that was riparian. Myrmicinae Myrmica Stenamma americana E C -1- diecki C brevinodis C healthi C emery ana E C wheelerorum C fracticornis E C Aphaenogaster lobifrons E C occidentalis c -1- tahoensis C Leptothorax sp. nov. A C crassipilis c Manica muscorum A E c bradleyi C nevadensis A E c -1- hunteri E c nitens rugatulus c c + -1- DOLICHODERINAE Liometopum Tapinoma luctuosum c + sessile FORMINCINAE A E c -1- Camponotus integroides c essigi c lasioides A c + laevigatus c microphthalma c modoc c neortifibarbis A E c viciniis c + nevadensis c Lasius obscuripes A c + alienus A E c obscuriventris A c -1- flavus c oreas c -1- neoniger c -1- planipilis c sitkaensis c propinqua c + vestitus c puberida c Formica Sibylla E c argentea A c + subnuda E c dakotensis A E c + subpolita A E c -1- densiventris E c -1- subsericea E c -(- fusca A E c Polyergus hewitti A E c breviceps E c -1- December 1978 Wheeler, Wheeler: Nevada Ants 389 Coniferous Forest Biome: under a stone (mostly) or wood lying on the surface, or usually somewhat buried, 228; in rotten wood, 64; under earthen mounds construct- ed by the ants, 12; in soil with craters, 15; thatch mounds, 9; thatch and wood, 4. Ecotone: Under a stone (mostly), or wood lying on the surface, or usually somewhat buried, 42; in rotten wood, 3; in soil with crater, 1. Alpine: Under a stone lying on the sur- face or more commonly somewhat buried, 41; in soil with crater, 1; thatch mound, 1. Records for Nevada Mountain Ants In the following list the arrangement of genera is that of Creighton (1950); for each genus, subgenus, or species-group, the spe- cies are arranged alphabetically. Under each species the county name is given first in italics; under each county the names of the ranges or mountains are followed by a dash; in each range (or mountain-group) are given the localities (peak or other topographic feature); finally the elevation of the record is given in feet above sea level. See Figure 5 for named localities. Abbreviations and symbols: Co. = Coun- ty; Hwy = Highway; mi = miles; Mt. = Mountain; Mts. = Mountains; nr = near; = feet. Compass directions are represented by the symbols E, N, S, W, and various combinations thereof. They are understood to be followed by the word of; e.g., "5 mi WSW Reno" would be read aloud as "five miles west-southwest of Reno." When locality is given by legal descrip- tion, the first figure is the section, the sec- ond is the township, and the third is the range. Because all Nevada ranges are east we have not so indicated, but township is cited as north or south; if the section is not known, we have given only township and range. Example: 23-7S-60, if fully expanded would read "section 23, Township 7 South, Range 60 East." The word range here has nothing to do with mountain ranges. (If the uninitiated reader is now thoroughly con- fused, we refer him to Wheeler and Whee- ler 1963:76-77.) Dr. Francoeur has made a preliminary ex- amination of our material included under Mynnica spp. nov. He thinks we have at least four new species, which he will de- scribe later. We have, therefore, added to each record our field number to make it possible to associate localities with the new names. Subfamily Myrmicinae Genus Myrmica Latreille Myrmica americana Weber Clark Co.: Spring Mts.— Charleston Peak 10,400'. Mineral Co.: Wassuk Ra.- Grant Peak 8000'. Myrmica brevinodis Emery Washoe Co.: Carson Ra.— Little Valley 6400'; Hwy 27 nr Mt. Rose 8800'. White Pine Co.: Schell Creek Ra.— North Schell Peak 9700'. Myrmica emeryana Forel Clark Co.: Spring Mts.— Charleston Peak 10,400'. White Pine Co.: Schell Creek Ra.- South Schell Peak 9000'. Myrmica fracticornis Emery Clark Co.: Spring Mts.— Charleston Peak 8100', 9800'. Elko Co.: Jarbidge Mts.- 4 mi S Jarbidge 6600', 6700'. Ruby Mts.- La- moille Canyon 8200'. Esmeralda Co.: White Mts.- Boundary Peak 9800'. Humboldt Co.: Pine Forest Ra.— Onion Reservoirs 8000'. Nye Co.: Grant Ra.- Troy Peak 10,800'. Washoe Co.: Carson Ra.— Little Valley 6400'; Whites Canyon on Mt. Rose 6800'; Tahoe Meadows on Mt. Rose 8400'. White Pine Co.: Snake Ra.- Mt. Moriah 10,000'; Mt. Washington 10,400'; Wheeler Peak 8300', 10,000', 10,400'. Myrmica lobifrons Pergande Elko Co.: Pilot Ra.- Pilot Peak 8800'. Nye Co.: Grant Ra.- Troy Peak 8400'. Wliite Pine Co.: Schell Creek Ra.- North Schell Peak 9800', 10,000'. 390 Great Basin Naturalist Vol. 38, No. 4 3 (Sgrays pk. 4^ :^. .♦'pilot pk. S^OIYABE SUMMIT 'ikN. SCHELL PK. WS. SCHELL PK. •^- #iylURRY: 12 I ?? -^i^SUMMIT •;MT. MORIAH ^^i}WH■EELERPK#K I ftPYRAMID PK. I NYE: *i>-- ••■;= -'i x^.-HnccLCR-rrv^: 3K \f? DSi*E|l\,^|;N pftRANT PK-^w^HINffTONf! """^ ' ^ 5-^ :#!MT. .JEFFERSON -^^j "'' ^{i — '•' ''"' ■' T. GRANT f^lNERAL ..•JaBLE MT.-ijitTIMBER •^ J«TROY PK. 7 n5 MT. «b'oundary pk. ^'6 LINCOLN ;^t;1400 mg/1), the lethal temperature range is con- siderably lowered (Vigg and Koch 1978). The periods of maximum surface orienta- tion of the cutthroat trout population corre- sponded to the spawning run at the Marble Bluff fishway (April-May) and the maximum activity of the trout in the lake (December) (Vigg 1978b). The temperature regime of the surface waters functions as a gateway, allowing access from the hypolimnion of the lake through the shoals and into the Truckee River. This "gateway" is open dur- ing spring and again in early winter; inter- acting with river flows and temperatures, the lake's temperature mechanism thus de- limits the potential spawning periods of the cutthroat trout population. Snyder (1917) documented that historical spawning runs of Pyramid Lake Lahontan cutthroat trout originally occurred both in early winter and spring. The response of tui chubs to the temper- ature regime was nearly opposite that of cutthroat trout. The proportion of the tui chub catch taken in the surface waters was directly related to temperature from Febru- ary through June; as temperature increased from 6.6 to 16.3 C, the proportion of the catch /effort per unit of net area taken on the surface increased from less than 10 per- cent to over 75 percent. The mean propor- tion on the surface during July and August was about 51 percent, at surface temper- atures of about 22 C. Temperatures de- clined from 21.2 C in September to 14.5 C in November; during this period the per- cent of tui chubs taken in the surface sam- ples had decreased to about 20 percent. This movement of tui chubs from surface to benthic areas during fall is consistent with previously documented onshore-offshore benthic distribution patterns (Vigg 1978a). For example, during August 1977, 95 per- cent of the tui chub catch was taken in lit- toral (0-15 m) areas; by November, 64 per- cent was taken offshore at 46 m. Similar Gila distribution patterns have been observed by other workers. Snyder (1917) reported an inshore movement of tui chub in Pyramid Lake in late May, associ- ated with warm water temperatures and commencement of spawning. The Eagle Lake tui chub population also shifted in the spring from the deep waters inhabited dur- ing the winter into warming shallow waters (Kimsey 1954). The limnetic form of Lake Tahoe tui chub inhabits upper waters dur- ing the summer and disappears into deep water during the winter; the benthic form likewise inhabits deep water during the winter, but shallow littoral areas during the summer (Miller 1951). Gaufin (1964) ob- served that Utah chubs, Gila atraria, gener- ally concentrated in the warmest areas of Fish Lake. Food consumption and assimila- tion, growth, and metabolic rates of Utah chub are directly related to temperature and are highest at 20 C (Cheng 1975). The Utah chub in Hebgen Lake, Montana, con- centrated in shallow areas in the spring, were widely distributed in the summer and fall, and concentrated in deeper waters in winter (Graham 1961). In Flaming Gorge Reservoir, the Utah chub population moved into the littoral and eulittoral areas about a month prior to the commencement of spawning, which occurred mid-June through mid- July (Varley and Livesay 1976). This in- shore movement corresponded to peak catches (activity) observed during July. December 1978 Vigg: Pyramid Lake Fish 423 The offshore, Umnetic tui chub popu- lation was surface oriented during the warm season. From June through October, about 92 percent of the hmnetic tui chub catch was taken in the upper 23 m (Table 2). Vertical limnetic distribution of tui chub is undoubtedly affected by temperature; however sampling was conducted during the warmest months of the year (June-Octo- ber) when mean surface temperature ranged only 16.3-23.1 C. Therefore, all of the sea- sonal distribution patterns (e.g., downward winter shift) were not qualified by the verti- cal gill net sampling of the limnetic zone. Nonetheless, monthly changes in vertical distribution occurred within this upper tem- perature range. Maximum surface temper- ature (mean = 23.1 C) was associated with maximum occurrence (70.4 percent) of tui chub in the upper 7.5 m. The temperature regimes in June (mean surface temperature was 16.3 and increas- ing) and October (mean surface temperature was 16.5 and decreasing) were similar; like- wise, the vertical distribution patterns of tui chubs were similar. During June and Octo- ber, 94.8 and 90.8 percent, respectively, of the total catch was taken in the upper 15 m. July and September also exhibited similar temperature regimes (mean surface temper- ature was 21.25 C) and similar (dispersed) tui chub distribution patterns. The 7.2-22.5 m depth interval was most abundantly rep- resented, 72.6 percent of the total in July and 72.1 percent in September. The limnetic tui chub population of Pyra- mid Lake is composed almost exclusively of the planktiverous (fine gill rakered) pectini- fer form which feeds almost entirely on zooplankton (Langdon 1978). The vertical distribution of limnetic tui chubs was associ- ated with that of the zooplankton they were selecting in their diet (Fig. 3). In June, Cyclops Moina Organisms/lifer Moinn Moina 0 i 75 15.0 e 1 22.5 Q 30.0 ^ 375 45.0 Tui chub percent! Zooplankton lorganisms/liter 50 10010 50 10010 50 10010 50 10010 50 10010 50 Percent June July August September October Fig. 3. Vertical limnetic distribution (percent by depth interval) of tui chub in Pyramid Lake, Nevada, from June through October 1977 in relation to the vertical distribution (density by depth interval) of the zooplankton taxon selected in the diet of tui chubs. 424 Great Basin Naturalist Vol. 38, No. 4 Ceriodaphnia quadranguh was very abun- dant in the upper 15 m, with decreasing density to 30 m; Hkewise, tui chubs were most abundantly represented in the upper 15 m, with an appreciable proportion to the 22.5 m depth. The selected zooplanker in July, Cyclops vernalis, was less abundant than C. quadrangula of the previous month; however, its absence in the surface waters and predominance in the 7.5-22.5 m depth interval mirrored the vertical distribution of the tui chub. The tremendous abundance of tui chubs in the upper 7.5 m in July may have kept the standing crop of its selected food item, Moina hutchinsoni, at low den- sities; and the vertical distributions of the fish and zooplankton were strikingly similar. During September the tui chub population was less dense (about 14 percent of the Au- gust level), and more evenly dispersed over the entire 0-22.5 m depth interval. Con- currently, M. hutchinsoni was very abun- dant in the same depth strata. Monia hut- chinsoni populations were most dense in the surface waters; however, due to the rela- tively high proportion of zooplankton to fish, it is unlikely that zooplankton were re- stricting the distribution of tui chubs within the upper 22.5 m of the limnetic zone. Tui chub distribution shifted upwards into the 0-15 strata during October. Monia hutchin- soni was less abundant this month and showed a very similar distribution pattern to that of the tui chub. The preliminary hydroacoustic survey conducted during April 1976 provided sup- plemental information on the vertical distri- bution of tui chubs. Apparently the upward vertical migration had already occurred by this early spring month; the limnetic tui chub population was concentrated in the 4-18 m strata, with appreciable densities extending to 48 m (Nunnallee et al. 1976) Concurrent total zooplankton densities ex- hibited a very similar vertical distribution pattern (Lider and Langdon 1978). Upward prevernal migration was substantiated by in- itial sightings of abundant tui chub schools on the surface in March— during weekly ae- rial observations, 1976-1978 (Joseph L. Kennedy, pers. comm.). Thus, seasonal vertical migration of tui chubs are probably concurrent with inshore- offshore distribution trends already docu- mented. In summary, it is likely that tem- perature-related factors are directive in- fluences on the fish and zooplankton which interact within their predator-prey relation- ships to form the observed limnetic distribu- tion patterns. Diel activity patterns and vertical distri- bution was determined during June and Au- gust 1977. Sampling was conducted over continuous 24-hour periods in the vicinity of Cormorant Rock (northwest Pyramid Lake). Surface and bottom gill net samples were taken at 4-hour intervals along the 23 m depth contour; i.e., a total of 24 net sets were made. Tui chubs were predominantly taken on the surface, and cutthroat trout were pre- dominantly taken on the bottom at 23 m. During both June (Fig. 4) and August (Fig. 5), however, fish catches exhibited con- sistent distribution changes with respect to time-of-day. Tui chubs were almost exclusi- vely taken in the surface waters from 2000- 0400 hours; the surface catch was negligible during the twilight and daylight hours. This could be attributed solely to net avoidance during daylight. For example, Lindenberg (1976) attributed the absence of alewives {Alosa pseudoharengus) in the catch of ver- tical gill nets during to day to net avoid- ance, even though the net appeared invis- ible to divers in the water. However, my experience with surface gill net sampling in- dicates that if fish were present in appre- ciable numbers they would be captured. For example, I captured large numbers of suck- ers (Catostomus) and shiners (Notemigonus) in daytime, surface-set, white, multifilament gill nets in littoral areas of Ruth Reservoir. I believe the observed patterns actually illustrate a movement of the tui chub popu- lation out of the surface into midwater. During both diel samples, the early morning movement of tui chubs out of surface wa- ters corresponded to peak bottom catches at 0400-0800 hours. I think this represents an overcompensation in downward movement, which occurred when the bulk of the tui chub population moved to midwater. Sim- ilar over-compensatory vertical migrations of zooplankton after sunrise have been documented (Hutchinson 1967). The diel December 1978 ViGG: Pyramid Lake Fish 425 vertical distribution of zooplankton closely corresponded to that of tui chubs. The prey species of zooplankton were most dense in surface waters from 2000-0400 hours and least dense at 1200; during midday, the highest densities were observed at 5-10 m (Lider and Langdon 1978). Benthic densities of zooplankton were generally low during all time periods. Very few cutthroat trout were captured 110- 100- _Q E _Q 3 75 50- 25- 20- 15- 10- 5- 0 Tui Cutthroat chub trout Depth A- A #--€) Surface A-A (t) (g) Bottom - 4 -3 -2 0) E C 3 o -1 ^^ m m 2400- 0400 0400- 0800 '■,a^,. 0800- 1200 Time AT 0 1200- 1600- 2000- 1600 2000 2400 Fig. 4. Diel activity and vertical distribution patterns of tui chub and cutthroat trout at the 23 m depth in the vicinity of Cormorant Rock, Pyramid Lake, Nevada, during June 1977. 426 Great Basin Naturalist Vol. 38, No. 4 in surface nets. The morning hours (2400- (0400-2000). Because trout are sight feeders, 1200) accounted for the entire surface trout it is logical that their activity would be catch. On the bottom, cutthroat trout catch- greatest during this time period. Although es were greatest during the daylight hours no samples were taken midwater, I hypoth- 100 - Tui chub Cutthroat trout Depth A- A #--# Surface A-A (D (g) Bottom _Q E _Q 0) _Q E z> c o ID 2400- 0400 0400- 0800 0800- 1200 1200- 1600 1600- 2000 2000- 2400 Time Fig. 5. Diel activity and vertical distribution patterns of the tui chub and cutthroat trout at the 23 m depth in the vicinity of Cormorant Rock, Pyramid Lake, Nevada, during August 1977. December 1978 ViGG: Pyramid Lake Fish 427 esize that cutthroat trout moved into the midwater zone during the dayUght hours to feed on the tui chub population which was concurrently at maximum densities there. Acknowledgments The Pyramid Lake Paiute Indian Tribe was instrumental in initiating the ecological research effort on Pyramid Lake. This re- search was funded by the Bureau of Indian Affairs, Contract H50C14209487. Mr. Floyd (Sam) Dunn and Mr. Dale Hebel carried out a great deal of the field work during this study. Mr. Edward Lider provided in- formation on zooplankton distribution, and Mr. Richard Langdon provided information on tui chub diet. Literature Cited BoRGES, M. H. 1950. Fish distribution studies, Niangua Arm of the Lake of the Ozarks, Missouri. J. Wildhfe Management 14(1): 15-33. Cheng, F. C. 1975. Bioenergetics of Utah chub and rainbow trout. Ph.D. Dissertation. Univ. of Idaho. 69 pp. Cooper, J. J. 1978. Contributions to the hfe history of the tui chub {Gila bicolor) in Walker Lake, Ne- vada. Unpubhshed master's thesis. Univ. of Ne- vada, Reno. 98 pp. Dendy, J. S. 1948. Predicting depth distribution of fish in three TVA storage-type reservoirs. Trans. Am. Fish Soc. 75(1945):65-71. DvTi-ER, W. P., AND R. H. Kramer. 1975. The in- fluence of temperature on scope for activity in cutthroat trout, Salmo clarki. Trans. Am. Fish Soc. 104(3):552-554. Fast, A. W. 1973. Effects of artificial hypolimnion ae- ration on rainbow trout {Salmo gairdneri Rich- ardson) depth distribution. Trans. Am. Fish. Soc. 102(4):715-722. Ferguson, R. G. 1958. The preferred temperature of fish and their midsummer distribution in tem- perate lakes and streams. J. Fish. Res. Bd. Can- ada. 15(4):607-627. Gaufin, R. F. 1964. Ecology of the Utah chub in Fish Lake. Unpublished master's thesis. Univ. of Utah. Ill pp. Gebhabt, G. E. and R. C. Summerfelt. 1975. Factors affecting the vertical distribution of white croppie {Pomoxis annuluris) in two Oklahoma reservoirs. Proc. Ann. Conf. Southeast. Assoc. Game Fish Comm. 28:355-366. Graham, R. J. 1961. Biology of the Utah chub in Heb- gen Lake, Montana. Trans. A. Fish. Soc. 90(3):269-276. Harris, E. E. 1970. Reconnaissance bathymetry of Pyramid Lake, Washoe County, Nevada. Hydro- logic Investigation Atlas HA-379, U.S. Geol. Surv. HoRAK, D. L. and H. a. Tanner. 1964. The use of vertical gill nets in studying fish depth distribu- tion, Horsetooth Reservoir, Colorado. Trans. Amer. Fisheries Soc. 93:137-145. Hutchinson, G. E. 1957. Treatise on limnology, Vol. 1: Geography, physics, and chemistry. John Wiley & Sons, Inc., New York. 1015 pp. 1967. A treatise on limnology. Vol. II: In- troduction to lake biology and the limnoplank- ton. John Wiley & Sons, Inc., New York. 1115 pp. Johnson, V. K. 1958. Fisheries management report- Pyramid Lake. Lakes Pyramid, Walker, and Tahoe Investigations. July 1, 1954 to June 30, 1958. Dingle-Johnson Project FAF-4-R. Nevada Fish and Game Department. 47 pp -I- appen- dices. KiMSEY, J. B. 1954. The life history of the tui chub, Siphateles bicolor (Girard), from Eagle Lake, California. California Fish and Game. 40(4):395-410. Koch, D. L. 1972. Life history information on the cui- ui lakesucker {Chasmistes cujus Cope, 1883) en- demic to Pyramid Lake, Washoe County, Ne- vada. Unpublished Ph.D. dissertation. Univ. of Nevada, Reno, 343 pp. 1973. Reproductive characteristics of the cui- ui lakesucker {Chamistes cujus Cope) and its spawning behavior in Pyramid Lake, Nevada. Trans. Am. Fish. Soc. 102((1): 145-149. Langdon, R. W. 1978. Food habits of the tui chub {Gila bicolor) in Pyramid Lake. Unpublished master's thesis. Humboldt State Univ. 43 pp. LaRivers, I. 1962. Fishes and fisheries of Nevada. Ne- vada State Fish and Game Comm. 782 pp. Lider, E. L. and R. W. Langdon. 1978. Plankton ecology. Chapter 9. In: W. F. Sigler and J. L. Kennedy (eds.). Pyramid Lake, Nevada ecologi- cal study— final report. W. F. Sigler & Associ- ates, Inc. Reno, Nevada. LiNDENBERG, J. G. 1976. Seasonal depth distribution of landlocked alewives, Alosa pseudoharengus (Wilson), in a shallow, eutrophic lake. Trans. Am. Fish Soc. 105(3):395-399. Martin, M. 1972. Morphology and variation of the Moduc sucker, Catostomus microps Rutter, with notes on feeding adaptations. Calif. Fish and Game 58(4):277-284. May, B. E. 1973. Seasonal depth distributions of rain- bow trout {Salmo gairdneri) in Lake Powell. Utah Acad. Sci., Arts, and Letters. 50(part 2):65-72. McCoNNELL, W. J., W. J. Clark, and W. F. Sigler. 1957. Bear Lake, its fish and fishing. Utah State Department of Fish and Game. 76 pp. Miller, R. G. 1951. The natural history of Lake Tahoe fishes. Unpublished Ph.D. dissertation. Stanford Univ. 160 pp. NUNNALLEE, E., R. JOHNSON, AND O. MaTHISEN. 1976. A hydrocoustic survey of the limnetic fish popu- lation of Pyramid Lake, Nevada. Final report to W. F. Sigler & Associates, Inc., Reno, Nevada. 13 pp -I- appendices. 428 Great Basin Naturalist Vol. 38, No. 4 OvERHOLTZ, W. J., A. W. Fast, R. A. Tube, and R. Miller. 1977. Hypolimnion oxygenation and its effects on the depth distribution of rainbow trout {Salmo gairdneri) and gizzard shad (Doro- soma cepedianum). Trans, am. Fish Soc. 106(4):371-375. Russell, I. C. 1885. Geological history of Lake Lahon- tan. A Quarternary lake of Northwestern Ne- vada. U.S. Geol. Surv. Monogr. ll:xiv-288. Snyder, J. O. 1917. The fishes of the Lahontan system of Nevada and northeastern California. U.S. Bur. Fisheries Bull. 1915-16(35):31-86. Sumner, F. H. 1938. An investigation of the present status of the Pyramid Lake fishery, Nevada. Stanford University, California. June 30, 1938. 1939. The decline of Pyramid Lake fishery. Trans. Am. Fish. Soc. 69:216-224. United States Geological Survey. 1977. Water re- sources data for Nevada, water year 1976. U.S. Dep. Inter., Carson City, Nevada. 344 pp. Varley, J. D. AND J. C. LivESAY. 1976. Utah ecology and life history of the Utah chub, Gila atraria. in Flaming Gorge Reservoir, Utah-Wyoming. Utah Div. of Wildlife Resources Pub. No. 76-16. 29 pp. Vice, S. 1978a. Seasonal bottom depth distribution of adult fish in Pyramid Lake, Nevada. Manuscript submitted to Calif. Fish and Game Journal, April 1978. 1978b. Fish ecology. Chapter 8. In: W. F. Sig- ler and J. L. Kennedy (eds.). Pyramid Lake, Ne- vada ecological study— final report. W. F. Sigler & Associates, Inc. Reno, Nevada. ViGG, S., AND D. L. Koch. 1978. Thermal tolerance of Lahontan cutthroat trout, Salmo clarki hen- shawi. Manuscript submitted to Desert Re- search Institute. April 1978. VON Geldern, C. E., Jr. 1964. Distribution of white catfish, Ictalurtis catits, and rainbow trout, Sal- rno gairdneri, in Folsom Lake, California, as de- termined by gill netting from February through November 1961. Calif. Fish and Game Depart- ment. Inland Fisheries Admin. Report No. 64- 15. Dingle-Johnsson Project California F-18-R. AN ADDITION TO THE AMPHIBIAN FAUNA OF CALIFORNIA David A. Mullen' and Robert C. Stebbins' Abstract.— a population of salamanders of the genus Ambystoma has been found at Grass Lake, Siskiyou County, in Northern California. A five-year study has established that the salamanders are reproducing success- fully and may represent a relictual population of native amphibians. The Grass Lake area has a sparse human population, which may help to explain how this secretive amphibian could have escaped notice. Attempts to de- termine the taxonomic position of the Grass Lake salamander, through comparisons of body measurements and coloration with those of other western ambystomatids, were inconclusive. These comparisons suggest, however, a close relationship with the geographically most proximate subspecies, A.t. californiense Gray and A.t. melanostic- tum Baird. Grass Lake is near the midpoint of a gap area of approximately 800 km that separates these sub- species. Regardless of the origin of the population, it must now be listed as an established addition to the am- phibian fauna of California. An adult ambystomid salamander was col- lected at Grass Lake, Siskiyou County, Cali- fornia, on 19 October 1969. The specimen was tentatively identified as a California ti- ger salamander, Ambystoma tigrinum cali- forniense Gray, though its coloration and lo- cation more than 500 km north of the northernmost portion of the known range of this subspecies in the San Francisco Bay re- gion did not strongly support this identi- fication. Large numbers of these animals were observed three days earlier by Mr. John Q. Hines, a long-time resident and nat- uralist in the area. This was the first time that he had seen salamanders at the lake. The animals were crossing the road, moving away from the lake during the first heavy rain of the year. The California tiger salamander occurs in central California, west of the crest of the Sierra Nevada from Sonoma County in the north to Santa Barbara County in the south (Stebbins 1966). It is found in foothill and valley grassland habitats from near sea level to approximately 1200 m elevation. The California form is distinguished from other western subspecies by its distinctive color pattern of oval, or bar-shaped, white, cream, or yellow spots on a black back- ground color. It is isolated from the others by great distances, there being no represen- tatives of the species known from the west- em part of the Great Basin south of the Co- lumbia River area or in the Sonoran Desert. The northern Great Basin, however, is oc- cupied by the subspecies A.t. melanosticum Baird whose nearest populations along the Columbia River are found approximately 432 km north of Grass Lake. The coloration of this subspecies differs considerably from that of the Grass Lake animal. The Grass Lake discovery initiated a five- year study of the area from October 1969 to October 1974, during which time many additional individuals were found. This pa- per reports the results of that study and at- tempts to illucidate the circumstances which could have permitted the undetected existence of such a large amphibian in the area. Comparisons of morphological data with those of other western subspecies of A. trigrinum are presented to suggest possible taxonomic relationships. Habitat Grass Lake is at an elevation of 1555 m in a shallow depression between Deer and Goosenest Mountains in the Cascade Range of northern California. U.S. Highway 97 'Department of Biology, University of San Francisco, San Francisco, California 94117. 'Museum of Vertebrate Zoology, University of California, Berkeley, California 94720. 429 430 Great Basin Naturalist Vol. 38, No. 4 crosses the lake approximately 34 km north- east of the town of Weed, California. This two-lane highway, built across the lake on a bed of earth and volcanic rock, was com- pleted in May 1932 (Anonymous 1932a). It is a well-used roadway between central Cal- ifornia and southeastern Oregon and carried 100 to 300 vehicles per day from its earliest days (Anonymous 1932b), a figure greatly exceeded today. A rail line parallels the southern margin of the lake with a spur northward into the lake for a short distance. This spur was probably used to obtain water for steam en- gines, and a small area immediately around its terminus was dredged to a depth of 1 to 2 m, forming a basin which held water in all months of the year since its construction in September 1906. Except for a small cattle ranch and highway maintenance and U.S. Forest Service stations at the western end of the lake, there is no nearby human habitation. The size of the lake varies greatly, depen- ding upon seasonal precipitation and the flow of two small streams. Bear Wallow and Dairy Creeks, which empty into its northern side. When filled by spring runoff, the lake, situated in a closed basin, mea- sures 4.25 km along its east-west axis, 1.75 km at its widest point. Its contour is very regular, except for the dredged area near the rail spur, which has a maximum depth of 1.5 m. The shoreline slopes gently into the lake around most of its perimeter. The level of the lake can be controlled by regulating the flow of water into a vol- canic vent, approximately 6 m in diameter, situated near the south-central shore. This drain has been encircled by an earthen dam to prevent natural drainage. The dam is breeched at two points by ditches which, when opened, can drain the lake completely to permit the grazing of cattle. The pres- ence of this vent indicates that the lake may have formed in the remains of an old volcanic caldera. Attempts to trace the route of the effluent from this vent, using marker dyes, were unsuccessful. Much of the lake bed supports a dense growth of tule bulrush, Scirpus acutus, which attains an average height of about 1 m. When the lake is drained, a mixture of native and introduced grasses and sedges grow under the Bulrush, providing the pri- mary source of forage for cattle. There are small stands of common cat-tail, Typha lati- folia, in the extreme western portion of the lake where somewhat deeper water occurs. A narrow strip of heavily grazed land, grown to grasses and sedges separates the lake from the forested hill sides of mixed western juniper, Juniperus occidentalis, and yellow pine, Pinus ponderosa. The under- story of the forest is primarily rabbit brush, Chrysothaminus nauseosus, and manzanita, Arctostaphylas viscida. The lake is used for breeding and nesting by a wide variety of resident and migratory birds. Large wading birds, including the great blue heron and common egret, fre- quent the lake and probably prey heavily on reptiles and amphibians since there are no fish in the lake. The Pacific tree frog, Hyla regilla Baird & Girard, long-toed salamander, Ambystoma macrodactylum Baird, and western toad, Bufo horeas Baird & Girard, are found in or near the lake, and the common garter snake, Thamnophis sirtalis L., is abundant. The burrows of the meadow vole, Microtus montanus (Peale), the western Mole, Sca- panus latimanus (Bachman), and the pocket gopher, Thomomys umbrinus leucodon Mer- riam, are found around the lake and extend into it when it is drained. The salamanders probably use these mammal burrows as un- derground refuges. Climatological data from the U.S. De- partment of Commerce Weather Station lo- cated at the nearby Mount Hebron ranger station indicates that the area has a mild climate for a mountain region. The weather at Grass Lake during this study was charac- terized by moderately warm summers, with a high mean monthly temperature of 17.3 C in July. The lake basin is subject to inter- mittent freezing and light snow from late October to early March, with a low mean monthly temperature in January of -4.0 C. Though the mean annual temperature of 6.5 C at Grass Lake did not differ greatly from the 9.0 C temperature recorded at Weed, only 34 km to the southwest, the difference in total precipitation at these two sites is striking. The mean annual precipitation at December 1978 Mullen, Stebbins: California Salamander 431 Grass Lake during this five-year study was 35.9 cm compared to the 68.8 cm recorded at Weed during the same interval. This var- iation in precipitation over so short a dis- tance illustrates the tremendous influence exerted over local weather patterns by the 4317 m Mount Shasta. Most of the precipi- tation falls in the form of rain during the winter and spring months of October through April. Summer thunderstorms ac- count for a small portion of the total but can influence the level of Grass Lake signif- icantly during critical summer periods. Methods Because of the great distance from San Francisco to the study area and the diffi- culty experienced in finding specimens of the Grass Lake salamander, collecting trips were conducted mainly in October to coin- cide with the expected fall migration. Table 1 summarizes the conditions at Grass Lake and the collections made during this study. This information is presented to illustrate the difficulty encountered in our attempts to observe and collect specimens from a population we knew existed in the area. These difficulties offer some evidence, we believe, that this population may have exist- ed, undetected in the area for a long time. Preserved Material Preserved material from the Grass Lake population was compared with that loaned by various western university museums. Loan materials were usually of random col- lections from widely scattered locations and often contained representatives of more than one subspecies of A. triginum. For the purpose of this study, unidentified speci- mens were designated members of a par- ticular subspecies if the collection site was within the range of that subspecies accord- ing to the range maps of Stebbins (1966). Figure 1 (modified from Stebbins 1966) shows the areas from which the materials used in this study were collected and the subspecies to which they belonged. Only data from fully transformed, adult specimens were compared. There was a general paucity of adult material available, with most loan collections composed of a few adults among large numbers of larvae. The difficulties encountered by the authors in obtaining adult specimens from the Grass Lake population suggest the reason for the Table 1. Summary of observations and collections at Grass Lake, Siskiyou County, California. 1969 1970 1971 1972 1973 1974 Date April 3-4 Water level October 19 Low December 8 Low December 21 Low High October 21 Low October 22 High October 23-25 High October 13-15 High October 27-29 High October 13-14 Low October 20-22 Low October 27 High Weather conditions Freezing with snow and ice present Snowing, no ice present Clear with freezing temperatures Clear, warm, and sunny Clear and dry Raining and warm Snow with melting snow present Clear and dry, cold Clear and dry, cold Clear and dry, warm Overcast with freezing temperatures Raining with temperatures just above freezing Specimens collected 1 (live) 1 (live) 4 (live)* 1 (dead) 1 (live) 1 (dead) none 1 (live) 1 (live) 1 (dead) none none none none 27 (live) •Collected by Dr. John O. Sullivan, Department of Biology, Southern Oregon College, Ashlund, Oregon. 432 Great Basin Naturalist Vol. 38, No. 4 scarcity of adult material in other collec- anterior opening of the vent; head width at tions. the posterior angle of the mouth; head Data taken from preserved materials in- length, from the tip of the snout to the fold eluded: total body length, snout to tail tip; of skin at the base of the skull; and the po- snout-vent length, from lower jaw to the sition of the eyes, distance between the in- 4 / melanosticfum I ; I A. t. californiense \^- J A.tnebulosum * A.t movortium other subspecies Grass Lake,Siskiyou Co., Calif. Fig. 1. Map of the western United States showing the distribution of the subspecies of Ambystoma tigrinum (adapted and modified from Stebbins 1966.) December 1978 Mullen, Stebbins: California Salamander 433 nermost eyelid margins. All measurements were taken with calipers to the nearest 0.1 mm. Colors and patterns of coloration were also noted for live and preserved specimens. Results Body measurements.— Measurements for western subspecies of A. tigrinum are pres- ented in Table 2. Unfortunately, few adult specimens are available in collections and collecting efforts vary greatly. In southern populations of nebulosum Hallowell, for in- stance, almost all of the adult material available was collected at a single location and time. Allowing for these intrinsic sources of error, however, the Grass Lake salamander seems to more closely resemble the slightly smaller californiense and slightly larger melanostictum than it does the other geographically proximate subspecies nebulo- sum and mavortium Baird. If arranged in ascending order, the various measurements of the Grass Lake sample fall next to those of melanostictum in all comparisons and be- tween this subspecies and califoriense in three of the five comparisons. The ratios of certain body characteristics presented in Figure 2 are considered more useful as indicators of possible taxonomic relationships because they do not rely on sample size or the uniformity of collecting efforts. The subspecies are arranged accord- ing to their distance from Grass Lake. The resemblance of the Grass Lake am- bystomatid with other western subspecies of A. tigrinum is demonstrated. The possession of a slightly wider head with wider set eyes seems to be the only morphologic charac- teristic examined which might be diagnostic of the Grass Lake form. Coloration.— Living specimens of the Grass Lake ambystomatid usually have a black to dark, olive green dorsal, back- ground color which gives way gradually on the lower flanks to a lighter grey-green col- or on the ventrum (Fig. 3). The background coloration is usually uniform from head to tail on both the dorsal and ventral surfaces and is overlaid by cream-colored, diffusely outlined, irregular spots which tend toward a mottled pattern on the flanks. This pat- tern extends onto the flanks, belly and gular areas in larger specimens and from the tip of the tail onto the head on the dorsum of most specimens. Color comparisons of preserved speci- mens with those of other western ambys- tomatids, generally considered unreliable because of the variable effects of fixation, did not show any variations which could be considered unique. The Grass Lake salaman- der resembles californiense in the possession of cream rather than black spots, and mela- Table 2. Comparison of body measurements taken from preserved specimens of the Grass Lake salamander with those of the western subspecies of Ambystoma tigrinum. Mean measurements in millimeters ± S.E. (Sample size) Total Snout-vent Head _ Eye Subspecies body length length Width Length width melanostictum 16L6±n.3 80.8 ±5.4 17.9±1.1 22.9 ±1.6 8.51 ±0.6 (13) (11) (13) (13) (13) Grass Lake 150.0 ±4.2 77.9 ±2.0 19.3 ±0.5 25.0 ±0.6 8.69 ±0.2 (30) (32) (33) (33) (33) californiense 147.1 ±8.0 77.9 ±2.8 17.5 ±0.6 25.3 ±1.0 6.27 ±0.2 (21) (21) (22) (22) (22) nebulosum, southern 140.4 ±2.8 72.4 ±1.0 16.6 ±0.2 22.9 ±0.2 6.58 ±0.1 (10) (10) (10) (10) (10) nebulosum, northern 188.4 ±6.7 86.0 ±2.6 20.3 ±0.6 27.4 ±0.6 9.27 ±0.3 ( 8) ( 9) ( 9) ( 9) ( 9) mavortium 172.6 ±13.4 86.6 ±5.3 20.1 ±1.5 26.3 ±1.6 8.83 ±0.6 (5) ( 5) ( 5) ( 5) ( 5) 434 Great Basin Naturalist Vol. 38, No. 4 nostictum in having mottling on a dark dor- sal background. It is also similar to califor- niense in the extent to which patterns of spots extend over the upper body and onto the ventrum and gular regions. Although the individual variation in this characteristic is also great, the typical Grass Lake sala- mander tends to have a greater number of smaller and more diffuse spots on the dor- sum and a more extensive ventral pattern than does californiense. Discussion The population at Grass Lake is known to have maintained itself for nine years. It appears to be well established and should be listed as an addition to the amphibian fauna of California. Local residents apparently knew nothing of the existence of salamanders at Grass Lake prior to 1969. One amateur naturalist who had lived at the Grass Lake Ranger /l.f. melanostictum A.t GRASS LAKE A.t. californiense A.t.nebulosum^ southern A.t.nebulosum, northern A.t.mavortium B. I 1 r .60 .70 .80 Head Width (rotio to head length) .30 -a^ -^ * ~-| — .40 .50 Distance Between Eyes (ratio to heed width) A.t. melanostictum A.t GRASS LAKE A.t californiense A. t nebulas urn, southern A.tn, northern A. t mavortium D. .40 1 .50 .60 Snout -vent Length (ratio to total length) * -^ ^ ^ -^ + .20 .30 Head Width (ratio to snout-vent length) Fig. 2. Morphologic characteristics of the Grass Lake salamander compared with those of western subspecies of Ambystoma tigrinum. Horizontal line represents range; vertical line represents mean; open rectangle represents one standard deviation on either side of the mean; black rectangle represents twice the standard error of the mean on either side of the mean. December 1978 Mullen, Stebbins: California Salamander 435 Station from 1953 to 1960 stated that she had never seen salamanders in the area, nor had she known of anyone who had. She was aware of the existence of a salamander nearer to Mount Shasta, probably Ambys- toma macrodactylum, known to exist in that area. Other residents reported occasionally seeing "slimy lizards" in creeks which drained into the Klamath River away from Grass Lake. Occasionally caught on baited trout hooks, they were said to resemble mud puppies (Necturus) and were probably larval Dicamptodon. A very large number of adult salamanders were first noticed migrating from Grass Lake in the fall of 1969. Reports of hun- dreds of salamanders crossing the highway from the lake indicate that the population had grown, unnoticed, to a substantial size. The migration is usually completed in a single night during the first heavy annual rainfall and would be observed mainly by motorists. Unless large numbers of salaman- ders were on the road, it is unlikely that disinterested motorists would report their presence. The animals blend with the pave- ment and could easily be overlooked during the hard rains that stimulate migration. Ti- ger salamanders in semi-arid western parts of the range are subterranian much of the year and are seldom encountered except during their brief breeding migrations. It is thus possible that a relatively large popu- lation could have existed in this isolated re- gion for some time without being noticed. We obtained no larvae despite collecting attempts during both summer and winter months; however, our efforts were not in- tensive. We have considered the possibility that the Grass Lake salamander was introduced. "Water dogs" had been used as fish bait in Dwinnel Reservoir, a large impoundment about 32 km southwest of Grass Lake. A re- Fig. 3. Living, anesthetized Ambystoma tigrinum collected from Grass Lake, Siskiyou County, California. A. MVZ 99976, a 148mm female collected 8 December 1969; B. RCS 11766, a 186mm male collected 19 October 1969. 436 Great Basin Naturalist Vol. 38, No. 4 tailer in Weed stated that the best source of his bait had been a dealer from the state of Washington who had supplied good num- bers of gilled larvae described as uniformly dark grey above and lighter grey below. This source was stopped by California au- thorities in 1965 or 1966. It is most prob- able that the subspecies involved in these transactions was mehnostictum. A second source of "water dogs," de- scribed as not having gills and with bright spots on the tail, was from a wholesaler who bought them in Texas or Louisiana. They were not as desirable a bait because they would not swim down into the lake as did the larvae from Washington. This cir- cumstance, plus their much higher price, greatly limited the number imported. Cali- fornia authorities stopped their importation after 1969, though in that year alone the bait dealer sold more than 7,000 of them. It is most probable that the salamanders in- volved in these transactions were A. t. ma- vortium or the newt, Notophthalmus virides- cens. Another bait dealer in Redding, Califor- nia, confirmed that they too had obtained good numbers of what they called "yellow water dogs" prior to 1969. These salaman- ders from a Texas supplier were described as about 10-12 cm in length and a imiform light yellow in color. Some had gills and some did not. It is again probable that these salamanders were A. t. mavortiiim. After the Texas supply was stopped, small numbers of salamanders were obtained oc- casionally from wholesalers in Nevada. These animals were described as lacking gills and uniformly light grey in color. The subspecies involved in the Nevada transac- tions could have been A. t. nebulosum. Bait purchased in the Redding area was usually used in nearby Lake Shasta. Very little bait was probably transported the 130 km northeast to Dwinnel Reservoir. Indeed, the great majority of fishermen would have traveled to this lake from the population- dense Redding area and farther south. Sur- viving baits, not released at the fishing site, would probably have been discarded by fishermen along the road leading from Dwinnel Reservoir to the southwest and away from Grass Lake. Oregon fishermen from the Klamath Falls area, most likely to use Highway 97 across Grass Lake, prob- ably would not fish in California, where an out-of-state fishing license would be re- quired. These factors tend to minimize the possibility of an accidental introduction of ambystomatids into Grass Lake. There re- mains, nonetheless, the possibility that the Grass Lake population of salamanders re- sulted from an accidental introduction of unused fish bait. The Grass Lake population might have resulted from an increase in the numbers of an ambystomatid native to the area due to recent favorable environmental circum- stances. If this is the case, the question re- mains: why were they not observed prior to 1969? Access to the area was only possible by a system of unimproved logging roads prior to 1932. Efforts to study the biology of this remote area were probably few prior to this time, and it is possible that these rarely en- countered animals could have been present, undetected, or unreported. With the paving of the road came travel across the lake on a daily basis by hundreds of vehicles. This in- creased traffic, however, would have passed quickly through the area and few travelers would have stopped, as there were no sid- ings or recreation facilities until the late 1960s. Further, the numbers of salamanders were probably very low and any limited use of the area for recreation would probably not have occurred during the rainy period when migration takes place. Any discussion of the existence of a pre- viously unknown ambystomatid native to the Grass Lake area, must account for their high numbers in 1969. The dredging and maintenance of the railroad water supply pond may have stabilized and even permit- ted a gradual increase in the numbers of a small population of salamanders by provid- ing a stable breeding habitat. Later, agricul- tural practices at the lake, initiated in the 1960s, may have provided the necessary en- vironment for a more rapid growth of the population. When the natural drainage was interrupted, the water level of the basin was significantly altered. This may have dramatically increased the breeding success of the population by providing a large. December 1978 Mullen, Stebbins: California Salamander 437 stable body of water necessary for increased reproduction while at the same time reduc- ing the success of predators and permitting increased escapement of adults. Successful reproduction in this relatively arid, high al- titude location would certainly depend heavily on the conditions at this small, iso- lated water source. The intervention of man may have proved highly beneficial and re- sulted in the resurgence of a relictual popu- lation. Evidence from this study seems com- mensurate with the hypothesis that a relic- tual population of Amhystoma tigrinum has existed, undetected, in the Grass Lake area and that recent manipulation of the lake by man has permitted a resurgence of breeding success and growth of this remnant popu- lation. Additional study will be required to clarify its taxonomic position. Acknowledgments We thank the following persons and in- stitutions for the loan of specimens in their care: I. McT. Cowan, University of British Columbia; John M. Legler, University of Utah; T. Paul Maslin, University of Colo- rado, Boulder; and Richard L. Wallace, University of Idaho. Our appreciation also to Robert M. Winokur, University of Utah, for his generous donation of specimens from southwestern Utah and to John O. Sullivan, Southern Oregon College, for his provision ^ of additional specimens of the grass Lake Ambystomatid and data concerning their ecology at that site. We also appreciate the opinions of Robert W. Reese, University of Colorado, Colorado Springs, and the time he expended to compare specimens of the new salamander with those in his collection. Finally, our sincerest appreciation to John Q. Hines for his many kindnesses during this study and for his assistance in the collection of field data and specimens from the Grass Lake area, and to Gene M. Christman for preparation of the illustrations. Literature Cited Anonymous. 1932a. Final Report Covering Application of Heavy Fuel Oil and Asphaltic Road Oil as a Dust Palliative on Portions of the State High- way Between Weed and the Oregon State Line. State of Calif., Dept. Publ. Works, Div. High- ways Rep., June 30, 1932. 1932b. Preliminary Report Covering Construc- tion of a Portion of Secondary State Highway between Dorris and the State Line in Siskiyou County. State of Calif., Dept. Publ. Works, Div. Highways Rep., Dec. 7, 1932. Dunn, E. R. 1940. The Races of Amhystoma tigrinum. Copeia, 1940: 154-162. Stebbins, R. C. 1966. A field guide to western reptiles and amphibians. Houghton-Mifflin Company, Boston, Massachusetts. RAPTORS OF THE UINTA NATIONAL FOREST, UTAH James A. Mosher', Clayton M. White^ Joseph R. Murphy^ and M. Alan Jenkins^ Abstract.— The past and present use of the Uinta National Forest by birds of prey was determined from his- torical records and surveys conducted during 1973, 1974, and 1975. Data describing the history, present status, and physical characteristics of each nest site were collected, along with the type and intensivity of human activi- ty within a half-mile radius of the nest. In addition, limited data on fall migration of raptors along the Wasatch Front and the food habits of Golden Eagles are presented. Management guidelines, based on the results of this study, are proposed. The Uinta National Forest occupies an area in excess of 3600 km^ in central Utah. It lies southeast of Salt Lake City (Fig. 1) and encompasses much of the Wasatch Mountain range and the northern portion of the San Pitch Mountains. The area is char- acterized by high, rugged mountains (to 3622 m) cut, especially on the western front, by deep canyons which are generally within the 1523 to 2745 m elevation range. These steep-walled canyons provide numer- ous potential nest sites for cliff-nesting birds. Riparian habitat found along the can- yon floors and higher forested slopes and plateaus provide nesting habitat for wood- land species. Habitat type changes with alti- tude, beginning with a zone of sagebrush at the valley floor and extending through scrub oak, aspen, and dense conifer before reach- ing the tree line. Each of these habitat types offer nesting potential for various spe- cies of raptors. Agricultural and urban development are encroaching up the slopes of much of the western front of the Wasatch Range, and recreational development of the canyon areas is increasing. This activity and a gen- erally increasing awareness of the .suscep- tibility of some raptor species to human dis- turbances prompted the management personnel of the Uinta National Forest to initiate this study of its rare and endangered raptors. Field work relating to this study was conducted from January 1972, to June 1975. The goals of this research were to de- termine: (1) what species of raptors were found in the Forest; (2) when and where they used Forest lands; (3) Forest use by raptors, and particularly by Golden {Aquila chrysaetos) and Bald eagles {Haliaeetus leu- cocephalus) and Peregrine Falcons {Falco peregrinus); (4) guidelines for management of raptors. Methods An intensive literature search on Utah birds of prey was conducted, and the field notes of R. G. Bee (Brigham Young Univer- sity Life Science museum) were searched and summarized. Public cooperation to in- crease sightings and nest records was solic- ited via radio and television publicity and distribution of posters. Surveys by vehicles and on foot were conducted over most of the Forest from 1973 to 1975, paying particular attention to cliff areas. All recorded and reported nest sites were checked. Aerial surveys in a Ces- sna 172 were conducted over some exten- sive cliff areas and other areas of known Bald Eagle use during the winter. Nest site data collected included eleva- tion above sea level, height on cliff, height 'Appalachian Environmental Laboratory, GEES, University of Maryland, Frostburg State College, Frostburg, Maryland 21532. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. Present address of Jenkins is: U.S. Fish and Wildlife Service, Denver Wild- life Research Center, Building 16, Denver Federal Center, Denver, Colorado 80225. 438 December 1978 MOSHER ET AL.: UtaH RaPTORS 439 of cliff, directional orientation of nest site, occupancy history, distance to nearest hu- man activity (i.e., roads, campgrounds, pas- tures, etc.), type and intensity of human ac- tivity within a half-mile radius of the nest site, breeding chronology, and prey remains. Estimates of home range sizes of Golden Eagles are based on multiple observations of individual birds, identified by characteristic plumage patterns, around active nest sites. Migration data were collected by direct ob- servations from a point east of Provo, Utah, on the Wasatch Mountain front at about 1980 m above sea level. Results and Discussion Historical Summary Table 1 is a compilation of raptor sight- ings on the Uinta National Forest up to the time of this study, largely from the unpub- Ushed field notes of R. G. Bee and K. Tur- ley. Except as noted, breeding status is not given. More complete data are available for the Golden Eagle and Peregrine Falcon. Table 2 summarizes the history of the Gold- UTAH Fig. 1. Location of the Uinta National Forest (1972) within Utah. en Eagle nest sites. Old nests located during this study were listed as sites with no re- corded activity. Identification of Golden Eagle nests was based on size and location. There is a long history of breeding activi- ty by Golden Eagles in the Uinta National Forest. It is the most conspicuous breeding raptor. Allowing for alternate nest sites, some 21 territories are now known. Based on Bee's field note records of nest sites, the number of eagles appears to have changed little since the 1930s. The breeding season for this species extends from January to late June (Smith and Murphy 1973, and this study). The Peregrine Falcon has occupied three known eyries in the forest. The last known nesting activity was in 1968. According to Porter and White (1973), ". . . the known active eyries of this species in Utah are now only about 10 percent of those known to have been present earlier in the century." The Peregrine was first recorded in Utah in 1871 (Allen 1872) and the first active eyrie reported in 1899 (Johnson 1899). Fossil evi- dence establishes both the Peregrine and the Prairie falcons {Falco mexicanus) in Utah back perhaps as far as 40,000 years (Porter and White 1973). The known range of the Peregrine in Utah includes only a small part of the Uinta National Forest lands; however, three prior eyrie locations are known within the Forest boundary (Porter and White 1973). The his- tory of these eyries is reproduced from Por- ter and White (1973) in Table 3. We have a recent report of repeated sightings of a Per- egrine in the Spring Lake area from April to June or July 1972 (Turly, pers. comm.). An interesting review of the history and status of the Peregrine Falcon with refer- ences to relations with the Prairie Falcon in Utah is to be found in Porter and White (1973). Very little historical information about the Prairie Falcon in central Utah is available. It was, however, reported to be common in the rocky canyons of the Wasatch Mountains prior to the turn of the century (Ridgway 1874). We have records of its nestings in six locations on Forest land from 1930 to 1940, as well as sightings in four other Forest locations up to 1955. These birds may still nest on the Forest, al- 440 Great Basin Naturalist Vol. 38, No. 4 though no individuals were seen during this study. Recent Activity The results of the 1973 survey, which was the most extensive and systematic survey, are summarized in Table 4. These results need careful interpretation because birds are not necessarily sighted in proportion to their actual numbers. Eagles and Turkey Vultures {Cathartes aura) are very con- spicuous because of their size and soaring habits. Kestrels {Falco sparverius) are also very conspicuous in spite of their small size because they hunt from perches (especially power lines) near roadways. Buteos are gen- erally conspicuous, but less so than eagles Table 1. History of raptor sightings on the Uinta National Forest prior to 1973. Species Date (month /year) Location Bald Eagle Golden Eagle (see also Table 2) Peregrine Falcon Prairie Falcon Sharp-shinned Hawk Cooper's Hawk Goshawk Harris Hawk Red-tailed Hawk Rough-legged Hawk Swainson's Hawk Turkey Vulture Boreal Owl Great Homed Owl Long-eared Owl Pygmy Owl Saw-whet Owl Screech Owl Short-eared Owl Spotted Owl 10/69 Mt. Timpanogos 3/69 Y Mountain 4/69 Mt. Nebo W-'72/'73 South fork, Provo 4-6/72 Spring Lake 4/29 ? °3/30 Dry Canyon, Mona "5/30 Provo Canyon •3/31 Alpine (several years). Fort •4/36 Grove Canyon •4/38 Days Canyon, Hobble Creek •4/40 Alpine, Box Elder 5/71 Spring Lake •5-8/72 Spanish Fork, Canyon 10/71 Spring Lake 8/70 Rock Canyon, Provo 6/71 Left fork. White River 2/72 Spring Lake 12/72 Wallsburg, south 8/72 Current Creek guard station 3/69 Fairview 1/70 Current Creek 3/71 Spring Lake 2/72 Spring Lake 1/72 Spring Lake 8/69 Nebo Loop Road 8/72 Nebo Loop Road 3/69 Provo 2/68 Y Mountain 1/71 Spring Lake 1/72 Spring Lake •7/72 Squaw Peak Trail 12/71 Diamond Fork 1972 Dry Canyon, Linden 6/72 Spring Lake 1/71 Spring Lake 7/71 Rock Canyon 'Indicates sighting was at active nest. December 1978 MOSHER ET AL.: UtaH RaPTORS 441 or vultures. Falcons and accipiters are quite inconspicuous. Owls, by their nocturnal or crepuscular habits, are the least conspicuous of all in diurnal surveys. Subjective estimates of population density were made for several resident species by estimating availability of suitable habitat, relative degrees of conspicuousness, and numbers seen on the Uinta National Forest. These were as follows: Golden Eagles, 84 (1/39 km^); Kestrels, 128 (1/26 km-); Red- tailed Hawks (Buteo jamaicensis), 60 (1/54 km^); Harriers {Circus cyaneus), 16 (1/202 km^); and Turkey Vultures, 52 (1/62 km^). The total raptor density for these species was 1 per 9.6 square kilometers. There was insufficient information to permit an esti- mate of population sizes for the other spe- cies. Some are, of course, wintering birds or migrant visitors. Fall Migration The use of north-south-oriented mountain chains by migrating raptors is a well known phenomenon, and the observation of fall mi- Table 2. Nest site history of Golden Eagles on the Uinta National Forest. Nest Site 1973 1972 1971 1970 1969 1968 1967 1966 Records prior to 1966° Alpine American Fork Heber Levan Nephi Pleasant Grove Provo Spanish Fork Springville Strawberry Thistle Wales -Dry Canyon -Fort Canyon -Canyon Head -Mt. Timpanogos— N -Pittsburgh Lake -East of 40 and 189° -Chicken Creek -Deep Creek— N -Deep Creek— S -Pigeon Creek Road -Pigeon Creek Trail -Footes Canyon -Red Cliffs -Hop Creek° -Salt Creek— S -Grove Canyon -Cascade -Rock Canyon -Y Mountain -Wallsburg-W -Wildwood -Lower Diamond Fork -Upper Diamond Fork -Little Rock Canyon -Balsam Campsite -Days Canyon -Kirkman Divide -Power House -Hobble/Diamond -Sulphur Campite -Current Creek" -Current Creek and 6° -Current Creek and 10 -Current Creek and 12 -Current Creek- W° -Big Jane Canyon -South* -UNF boundary A = Active nests ® = Unsuccessful nests (i.e., no young fledged) • = Not inside UNF boundary °* = Records from the field notes of R. G. Bee ® ® A A A '32, '34, '40, '50, '51 '33, '51 '34 '42 '37 '37, '38, '39, '46 '37, '40, '41, '44 '33 '39 '38, '39, '40, '41 '44, '46 '31, '45, '46, '55 '38 '55 '45 '37 '41 442 Great Basin Naturalist Vol. 38, No. 4 grations from look-out points provides much useful population data (e.g., Hawk Moun- tain Sanctuary, Heintzelman 1975). The Table 3.— History of Peregrine Falcon eyries on the Uinta National Forest.' Site no. First located and subsequent history Last known to be active 17 1930s, 1940s, 1950s, 1967 1968 18 1930s, 1939-46 1969 19 1930-32 1932 Wasatch Front, which forms the eastern boundary of the Great Basin, would appear to provide a similar corridor in the inter- mountain west. Data collected on the Uinta National Forest near Provo, Utah, from 1972 to 1974 suggest that considerable rap- tor movement occurs along this Front (Table 5) and more continuous observation might prove valuable. The greatest amount of movement apparently occurred about mid-September, when 35 and 60.5 birds per hour of observation were recorded. 'Taken from Table 1 of Porter and White (1973) Table 4. Summary of 1973 raptor sightings on the Uinta National Forest. Number of Number of Season Number of Number Number Species individuals sightings present nest sites active fledged Bald Eagle 27 8 FWS Golden Eagle 42 35 SFWS 27 4 4 Osprey Sharp-shinned Hawk 2 2 3 2 t' SEW Cooper's Hawk Goshawk 11 2 8 2 SFWS SFWS 5 4 Ferruginous Hawk Red-tailed Hawk 1 15 1 13 SFS SFWS Rough-legged Hawk Swainson's Hawk 2 4 3 2 FWS SFS Harrier 8 6 SFWS Kestrel 32 20 SFWS 4 4 Merlin 1 1 t Peregrine Falcon Prairie Falcon 1 2 1 2 SFWS SFWS Turkey Vulture Great Homed Owl 39 3 16 3 SFS SFWS Snowy Owl 1 1 WS Total of 18 species 195 127 36 12 4 't means transient Table 5. Observations of fall movements of raptors along the Wasatch Front at Provo, Utah' Date 1 (Hours of observat ion) August 18 25 September October Totals 2 4 9 15 20 22 27 29 4 6 7 13 19 (2.25) (2) (1) (0.5) (5.5) (2) (2) (2) (2) (1) (2) (1) (2) (1) (2) Sharp-shinned 0 2 1 0 16 32 1 0 13 2 7 3 2 0 0 79 Cooper's 0 0 0 0 3 1 1 0 4 2 1 1 0 0 1 14 Red-tailed 0 2 0 1 8 1 2 0 10 0 0 0 1 0 5 30 Ferruginous 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Golden Eagle 1 1 0 1 2 1 0 7 1 5 0 0 0 0 . 1 20 Harrier 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 American Kestrel 3 2 0 0 8 30 112 0 11 1 5 0 1 0 0 73 Unidentified 0 3 0 3 4 5 0 7 0 0 0 1 0 0 0 23 Total 4 10 1 5 41 70 116 14 44 10 13 5 4 0 7 341 Totals/hour of observation 1.8 5 1 10 7.5 35 60.5 7 21 11 6.5 5 2 - 3.5 'Combined for 1972, 1973 and 1974 December 1978 MOSHER ET AL.: UtAH RaPTORS 443 Wintering Bald Eagles Field observations from January 1973 through May 1974 point out two aspects of the wintering Bald Eagle population. First, eagles are dispersed over the Forest during the arrival and departure periods. Second, a midwinter concentration occurs along the lower 14 km of one stream drainage in the southern sector. During February 1974, aerial and ground surveys were conducted in an effort to lo- cate a communal roost site for the dozen or more eagles observed in the area. The aerial survey was conducted at sunrise and the ground survey from late afternoon to sun- down. Individual eagles were observed, but no communal roost was located. Unlike the desert population of Bald Eagles described by Piatt (1976), where roosting is limited to relatively few groves of suitable trees, the Forest population has an abundance of roosting sites. Golden Eagle Nesting and Human Disturbance The Golden Eagle is a year-round resi- dent and cliff nester on the Uinta National Forest. In 1973 and 1974, 18 pairs laid a to- tal of at least 11 eggs and successfully fledged 10 young between 3 June and 29 June (Table 6). Nest sites were situated on cliff ledges exclusively and averaged 1891 m in elevation (1495 to 2562 m). There was a significant tendency (a = 0.01, n = 37) for nests to be situated in a northward direction (Mosher and White 1976). Prey remains collected at three nest sites are summarized in Table 7. These data con- trast sharply with comparable data from desert nesting eagles, in that the mountain birds have a broader prey base and appear to rely less heavily on the Black-tailed jack- rabbit {Lepus californicus) (Smith 1971). Prey which inhabit the higher elevations, such as Blue Grouse {Dendragapus ob- scurus). Mule Deer fawns (Odocoileus he- mionus), and Snowshoe Hare {Lepus ameri- canus), are of particular interest. These data may reflect a low rabbit density which pre- vailed during this study. An estimate of minimum territory size was provided by multiple observations of two nesting pairs of eagles. The observa- tions were plotted on 7^2 minute USGS top- ographic maps and the perimeter points connected. The area contained within the polygon thus formed was estimated using a 1/10 in^ grid. The territory sizes were 8.8 and 17.9 km^ averaging 13.3 km^ The effect of human activity on nesting raptors is an area of debate. Although few "hard" data exist, this is an important as- pect of raptor management. In this study, we recorded several types of human activi- ties and their estimated intensity within 0.8 km of active and inactive Golden Eagle nests (Table 8). The most common activity Table 6. Golden Eagle nesting, 1974. Nest Pair in Eggs Young Fledging site attendance laid fledged date 1973 1974 1973 1974 1973 ] 1974 1973 1974 Heber yes yes yes no 1 - 6/3-10 - Alpine— Dry Canyon yes yes — yes — 2 — 6/27 Pleasant Grove-Chris Flat — yes — ? ? — — Provo— Cascade yes yes - ? no - - Springville- Hobble Creek (Sulphur) yes yes — yes — 1 — by 6/29 Diamond Fork— lower yes — 2 — 1 — 6/19-20 — Diamond Fork— 7 mi. cliff - yes - yes - 1 - by 6/29 Thistle— Big Jane Canyon yes yes yes no 1 — by 6/28 - Spring Lake - yes yes (nest destroyed) Payson Canyon no yes - yes — 1 — v Levan— Chicken Creek no yes — ? — ? — — Levan— Deep Creek yes no yes - 1 - 6/10-16 — Wales— UNF boundary yes no yes - 1 - by 6/28 — Totals 8 10 6 5 5 5 6/3-28 6/27-29 444 Great Basin Naturalist Vol. 38, No. 4 Table 7. Golden Eagle food habits on the Uinta National Forest. Prey species Hobble Creek 1974 Nest site and year Diamond Fork 1969-79° 1973 Alpine 1974 Total Mule Deer Black-tailed Jackrabbit Cottontail Rabbit Snowshoe Hare Uinta Ground Squirrel Rock Squirrel Muskrat Domestic Sheep American Coot Blue Grouse Black-billed Magpie Common Flicker Steller's Jay Unidentified 12 6 14 6 3 5 5 2 1 1 2 Blackbirds 2 2 Nest site totals 6 34 5 3 48 •From W. B. Amell, 1971 Table 8. Human activity at active and inactive Golden Eagle nest sites. Active sites Observed activity (intensity)' 1— Alpine— Dry Canyon 2— Alpine— Fort Canyon 6-Heber-40 and 189 8— Levan— Deep Creek 13— Pleasant Grove— Chris Flat 14— Provo— Cascade 18— Spanish Fork— Upper Diamond 19— Spanish Fork— Lower Diamond 23— Springville— Sulphur Creek' 24— Strawberry— Current Creek 1 27-Thistle-Big Jane 28-Wales-UNF boundary 30— Spring Lake Hiking, vehicle traffic (1) None Vehicle traffic. Shooting (2) Vehicle traffic (1) Vehicle traffic, grazing (1) Vehicle traffic (1) Vehicle traffic, hiking, climbing, shooting (3) Climbing, grazing, shooting (1) Vehicle traffic, camping, hiking, climbing, shooting (3) Vehicle traffic (2) Vehicle traffic, grazing, hiking, shooting (2) Vehicle traffic (1) Climbing (3) Inactive sites 9— Levan— Pigeon Creek 10-Nephi-Red Cliffs 11— Nephi— Hop Creek 12— Pleasant Grove— Grove Canyon 15— Provo— Rock Canyon 20— Springville— Power House 21— Springville— Kirkman Divide None Vehicle traffic (1) Grazing (1) Riding, hiking, shooting (3) Vehicle traffic, riding, hiking, shooting (3) None None 'Intensity classes are (1) little or none, (2) moderate, (3) severe. Classifications were assigned on basis of frequency, proximity, and number of types of disturbance present within a half mile of the nest site. December 1978 MOSHER ET AL.: UtAH RaPTORS 445 was vehicular traffic. The relationships be- tween intensity and type of activity near active and inactive nests are shown in Fig- ure 2. Sample sizes are small, but the re- sults suggest that there may be a great deal of individual variability in tolerance by eagles to one or more types of activity. Of 12 active nests in 1973 and 1974, five were visited on three or more occasions to collect data. No difference in fledging success be- tween nests visited and not visited was found (Table 9). The failure of one nest was attributed directly to removal of the egg by rock climbers. The following management guidelines were recommended to the Uinta National Forest as a result of the study: 1. All known Peregrine Falcon eyries should be preserved from further man- induced alteration until decisions are made about reintroduction, via captive breeding, of this species to central Utah. Table 9. Fledging success of disturbed and undis- turbed Golden Eagle nests. Year Fledging Nest site active Disturbed' success 1— Alpine- Dry Canyon 1974 Yes 2 2— Alpine- Fort Canyon 1973 No 6-Heber-40 and 189 1973 No 8— Levan— Deep Creek 1973 No 14— Provo— Cascade 1973 No 18— Diamond Fork- Upper 1974 No 19— Diamond Fork- Lower 1973 Yes Springville— 23— Sulphur Camp 1974 Yes 23— Sulphur Camp 1975 No (1) 28-Wales 1973 No 29— Payson Canyon 1974 Yes 30— Spring Lake 1974 Yes 0 Disturbed sites— 5, Mean fledging success— 1 Undisturbed sites— 7, Mean fledging success— 10 :::: 4 J::: I I ACTIVE SITES ::::: inactive sites L::::i O) o ACTIVITY TYPE Fig. 2. Comparison of intensity of disturbance by type of activity at active and inactive Golden Eagle nest sites. (Numbers above bars refer to the number of nest sites in the sample.) 446 Great Basin Naturalist Vol. 38, No. 4 2. Human activity compatible with rap- tor nesting be permitted, provided the following criteria are met: (a) no major alteration affecting the actual nest site or prey base oc- curs. (b) disturbance is minimized prior to hatching. (c) the nest cliff itself is protected from climbing. 3. Raptor surveys should be conducted annually to monitor population status. (a) Golden Eagle nest sites should be checked during late May or early June to determine breeding activi- ty and productivity. (b) Road count of Bald Eagles should be conducted during January or February. Acknowledgments This study was funded by the U.S. Forest Service, Uinta National Forest, under Con- tract 50-905. We are grateful for the support provided by the Uinta National Forest personnel, es- pecially R. Wiesert, F. Savage, and K. Tur- ley. Critical reviews of earlier drafts of this paper by Dr. C. P. Stone and Dr. G. A. Feldhamer were particularly helpful. This is Contribution 848, Appalachian Environmental Laboratory, Center for Envi- ronmental and Estuarine Studies, University of Maryland. Literature Cited Allen, J. A. 1872. Notes on an ornithological recon- naissance of portions of Kansas, Colorado, Wyoming, and Utah. Bull. Mus. Comp. Zool. 3(6): 113-183. Heintzelman, D. S. 1975. Autumn hawk flights. Rut- gers Univ. Press, New Brunswick, New Jersey. 397 pp. Johnson, H. C. 1899. A successful day with the Duck Hawks. Condor l(3);45-46. MosHER, J. A. AND CM. White. 1976. Directional ex- posure of Golden Eagle nests. Canad. Field Natural. 90(3):356-.359. Platt, J. B. 1976. Bald Eagles wintering in a Utah desert. Amer. Birds 30(4):783-788. Porter, R. D., and C. M. White. 1973. The Peregrine Falcon in Utah, emphasizing ecology and com- petition with the Prairie Falcon. Brigham Young Univ. Sci. Bull., Biol. Ser. 18(1): 1-74. RiDGWAY, R. 1874. Notes on the bird fauna of the Salt Lake Valley and adjacent portions of the Wasatch Mountains. Bull. Essex Inst. 5:168-173. Smith, D. G. 1971. Population dynamics, habitat selec- tion and partitioning of breeding raptors in the eastern Great Basin of Utah. Ph.D. dissertation. Brigham Young University. 260 pp. Smith, D. G. and J. R. Murphy. 1973. Breeding ecolo- gy of raptors in the eastern Great Basin of Utah. Brigham Young Univ. Sci. Bull, Biol. Ser. 18(3): 1-76. TWO NEW SPECIES OF FLEAS OF THE GENUS MERINGIS (SIPHONAPTERA: HYSTRICHOPSYLLIDAE) Richard B. Eads' Abstract.— Two new species of Meringis are described. Meringis disparalis, n. sp., has been most commonly taken from Dipodomys merriami but has also been taken from D. ordii and Onychomys leucogaster. Collection lo- calities include the counties of Dona Ana, Eddy, and Luna in southern New Mexico. Meringis facilis, n. sp., has been taken from D. ordii, several other rodents, and Sylvilagus audubonii. Collection localities include Crowley County, Colorado, and Bernalillo, Chaves, and Valencia counties in New Mexico. A key to the species of $ Me- ringis is given. Numerous fleas in the collection of the New Mexico Health and Social Services De- partment and this facility labeled Meringis dipodomys from New Mexico and Colorado were studied. It is believed that two pre- viously undescribed species are present. True M. dipodomys have not been seen east of Arizona. Holotypes and allotypes of the new species are in the National Museum of Natural History, Washington, D.C. Para- types are in the collections of this labora- tory and the New Mexico Health and Social Services Department, Santa Fe. In addition to these 2 Meringis species, the other 15 species in this genus which I consider valid are: M. agilis Eads 1960, M. altipectin Traub and Hoff 1951, M. arachis Jordan 1929, M. hilsingi Eads and Menzies 1949, M. deserti Augustson 1953, M. dipo- domys Kohls 1938, M. nidi Williams and Hoff 1951, M. rectus Moreland 1953, M. vitabilis Eads 1960, M. californicus August- son 1953, M. cummingi Fox 1926, M. hiib- bardi Kohls 1938, M. jamesoni Hubbard 1943, M. parkeri Jordon 1937, and M. shan- noni Jordan 1929. Meringis disparalis, n. sp. Figs. 1-5 Diagnosis.— In the male an apical sinus divides the immovable process of the clas- per into two equal lobes in disparalis; in dipodomys these lobes are markedly un- equal. The arac/its-group of Meringis is unique in that the posterior margin of the caudal process of sternum 9 is truncate, forming almost 90 degree angles at the juncture with the dorsal and ventral mar- gins. The caudal process is short in dis- paralis, not reaching subapical spiniform bristles of distal arm of sternum 9; in dipo- domys, the caudal process usually extends beyond the subapical spinform bristles of sternum 9. Meringis disparalis females have a sinus on the caudal margin of sternum 7 that is shallow relative to that of dipo- domys, but a definite dorsal lobe is usually present. Male.— Head: Frontoclypeal margin evenly rounded. Micropunctations or pores scattered from first row of bristles to mar- gin of head. Preantennal region with three large dermal pores along cephalic margin and three smaller pores along ventral mar- gin. First preantennal row of three thin bristles, caudal or ocular row of four larger bristles; a few microsetae interspersed be- tween bristles in each row; ventrad of re- duced, lightly pigmented eye an irregular row of about five microsetae. Genal process bluntly acuminate, apex barely visible beyond genal comb which arises ventrad of eye; outer spine superimposed on basal 'Vector-Borne Diseases Division, Center for Disease Control, Public Health Service, U.S. Department of Health, Education, and Welfare, P.O. Box 2087, Fort Collins, Colorado 80522. 447 448 Great Basin Naturalist Vol. 38, No. 4 three-fourths of narrower but longer inner spine; inner spine more acuminate at apex than outer spine. Three short, stout setae at oral angle of head at base of 4-segmented maxillary palp. Acuminate maxillary lobe extending almost to apex of segment 3 of 5- segmented labial palp. Labial palp longer than maxillary palp, reaching about three- fourths length of fore coxae. Scape of an- tenna with three tiny bristles at base and four somewhat larger median ones; second antennal segment with apical fringe of short bristles not extending beyond segment 1 of club. A row of fine hairs along the dorsal AE.P. FULLL Figs. 1-3. Meringis disparalis: (1) $ head, pro- and mesothorax; (2) ,5 modified abdominal segments; (3) ae- deagus. December 1978 Eads: Two New Fleas 449 margin of antennal fossa, caudal two long- est. Three rows of postantennal bristles ar- ranged about 3:5:5, ventrad bristle longest in each row. Thorax: Pronotal comb of seven spines per side, preceded by a row of seven large bristles with fine intercalary bristles. Ante- rior and posterior margins of pronotum al- most straight, about same width throughout. Mesonotal flange with four pseudosetae per side. Mesepisternum with one median bristle along pleural rod preceded by a smaller one; mesepimeron with four cephalic bristles in an irregular row and a caudal row of two larger ones. Metanotum with a short row of three or four bristles, followed by a row of seven or eight medium-sized bristles and a row of about six longer bristles alternating with small ones. Lateral metanotal area well defined, with an oblique row of four bristles (dorsal bristle large, next one medium size, next one large, ventrad bristle medium sized). Pleural arch well developed, strongly convex. Legs: Procoxa with numerous bristles scattered over entire segment, ventromar- ginal bristles extending beyond trochanter. Mesocoxa with bristles limited to anterior marginal area; metacoxa with bristles on an- terior margin and about 13 submarginal bristles on distal half; metacoxa with an oblique row of five or six spiniforms begin- ning at anterior margin on apical one-fifth. Profemora each with 8 to 10 thin lateral and submarginal bristles. All femora with two ventromarginal subapical bristles, both long in femur 1, markedly unequal in length in femora 2 and 3. Femora with two long, unequal, ventromarginal, subapical bristles strongly arched toward tibia. Hind tibia with about 16 lateral and sublateral bristles; six lateral notches of caudal margin with unequal spines in ratio 2:2:2:3:3:3 (base to apex); distal margin of hind tibia with a row of 3 long, 2 small, and 3 long bristles; hind tibial tooth well developed. Abdomen: Tergum 1 with three or four scattered small bristles followed by a row of five or six larger bristles and a second row of a similar number of still larger bristles with fine intercalary bristles. Terga 2-6 typ- ically with two rows of about eight bristles each, caudal row of large bristles and five or six small intercalaries; tergum 7 with both rows of bristles reduced in number. Sternum of abdominal segment 2 with one bristle on ventral margin; sterna 3-6 each with a row of three bristles frequently pre- Figs. 4-5. Meringis disparalis: (4) apical third of aedeagus; (5) $ sternum 7 and spermatheca. 450 Great Basin Naturalist Vol. 38, No. 4 ceded by a single bristle; sternum 7 with a row of about four bristles preceded by two bristles. Three antepygidial bristles present; middle bristle over twice as long as upper one; ventral bristle longer than dorsal one. Modified abdominal segments: Tergum 8 reduced in size, narrow dorsally between antepygidial bristles and sensillum, expand- ing ventrad to below apex of proximal arm of sternum 4; with a mesal row of eight or nine thin bristles on dorsal half. Sternum 8 large, extending dorsad to near apex of pro- ximal arm of sternum 9 and caudad about half length of distal arm; with an irregular row of five or six median small bristles fol- lowed by a row of about five larger bristles; ventrocaudal margins of sternum 8 pro- duced into a process which is truncate at apex. Distal arm of sternum 9 tapering to a narrow tip, but not as narrow as bilsingi; subapical spiniform bristle always present; usually a second distal spiniform present which is smaller and less darkly pigmented. Immovable process of clasper com- paratively slender, apical portion not as wide as movable finger; divided into two lobes of approximately same height at apex by a shallow sinus; single acetabular bristle inserted near ventrocaudal margin (bristle over twice as long as this distance). Mov- able process of clasper roughly rectangular in shape, with broadly rounded (almost truncate) dorsal margin; over one-third of its height extends distad of the immovable process. Cephalic margin almost straight and caudal margin gently concave; a row of 10-12 microsetae along or near cephalic margin; several thin bristles scattered over movable process; about 15 marginal and four submarginal bristles of approximately the same size along caudal margin. Manu- brium about eight times as long as basal width; beyond basal third sides parallel; tip directed dorsad. Aedeagus (Fig. 3) comparatively long and slender; median lamina over three times as long as wide, about same width throughout except for a slight constriction immediately anterior to fulcrum. Fulcrum with a curved fulcral latero-ventral lobe and a thicker, straight fulcrual medial lobe. Two strong proximally directed lateral shafts of capsule pass under medial lobes of fulcrum. Cres- cent sclerite well sclerotized. Armature of inner tube prominent, longer than crescent sclerite, lying above sclerotized inner tube. Median dorsal lobe, bifid, narrowing api- cally to ventrad-directed, subacute apex. Lateral lobes long, narrow projections of pouch wall, broadest at base, tapering gent- ly to bluntly rounded ventrad-directed apex. Crochet well developed, much wider than lateral lobes. Female.— Outline of sternum 7 variable. Sinus shallow, but usually with a small, dor- sal lobe bluntly produced; anterior row of about six bristles, posterior row of about six longer bristles present. Anal stylet about 3.5 times as long as broad, with one long apical bristle and two very small, subapical ones. Bulga of spermatheca cylindrical, both ends of approximately equal width, ventral mar- gin almost straight and dorsal margin con- cave; hilla appreciably longer than bulga, about one-half as wide, sharply upturned and caudad-directed; sternum 8 reduced, three to five bristles at tapered apex. Ter- gum 8 large, two or three bristles near dor- sal margin just posterior to three antepygi- dial bristles; two mesal rows of unequal bristles present, posterior row complete from dorsal to ventral margins; anterior row only on lower half; caudal margin with an irregular row of heavy bristles. Type specimens were all collected by field personnel of the Plague Branch in New Mexico. HoLOTYPE $ .— Ex Dipodomys merriami. Dona Ana Co., 13-XII-1951. Allotype ? .— Ex D. merriami. Dona Ana Co., 12-XII-1951. Paratypes.— 3 $ S , 4 $ ? ex D. mer- riami. Dona Ana Co., 12-XII-1951; 2 $ $ ex Onychomys leucogaster. Dona Ana Co., 12-XII-1951; 1 ? ex D. merriami, Eddy Co., 6-III-1952; 1 $ ex D. merriami, Luna Co., 4-V-1939; 1 $ ex D. merriami, Luna Co., 19-XII-1950; I S ex O. leucogaster, Luna Co., 5- V- 1939; 1 ,5 ex O. leucogaster. Dona Ana Co., 14-IV-1939; 2 S $ ex D. merriami Dona Ana Co., ll-XII-1951; 1 S ex D. merriami, Luna Co., 19-XII-1950; 2 $ $ ex D. merriami. Dona Ana Co., 13- XII-1951; 1 <5 ex D. ordii. Dona Ana Co., 13-IV-1939; 1 $ ex D. merriami. Dona Ana Co., 15-1-1952. December 1978 Eads: Two New Fleas 451 Comment.— This species has been most commonly taken from the kangaroo rat, Dipodomys merriami, in the New Mexico , counties of Dona Ana, Eddy, and Luna. It has also been taken from D. ordii and Ony- chomys leucogaster. Meringis facilis, n. sp. Figs. 6-9 Diagnosis.— In the male, apical sinus di- vides the immovable process of the clasper into two markedly unequal lobes as in M. dipodomys and M. bilsingi. The short, thick (at midpoint) movable finger of the clasper (height only slightly more than twice its width) tapers to an evenly rounded, sub- acute apex, not at all tending to be rec- tangular in shape as with M. dipodomys and M. disparalis. The process of the ventro- caudal margin of sternum 8 much wider distally than in related species, wider than movable process of clasper. The distal arm of male sternum 9 extends a very short di- tance beyond the insertions of the two spin- iform bristles, the second and smaller of the two is on the truncate, caudal margin. In M. bilsingi the distal arm of sternum 9 ta- pers to a very narrow tip. Females are diffi- cult to identify in the absence of males. The sinus of sternum 7 is shallow but has a defi- nite dorsal lobe which tends to be sharply produced. Male.— Head: Frontoclypeal margin evenly rounded, with micropunctations scat- tered from first row of bristles to margin of head. Preantennal region with row of three thin bristles, followed by four larger bristles (dorsal bristle extends beyond inner tooth of genal comb) and an irregular row of about three microsetae. Postantennal region with micropunctations scattered anteriorly to first row of bristles; three large dermal pores and two smaller pores present; three rows of bristles arranged about 4:5:5, with ventral bristle in each row much longer; a row of about 10 fine hairs along dorsal mar- gin of antennal fossa. Thorax: Pronotal comb of seven large spines per side, preceded by row of six large bristles with fine intercalary bristles. Mesonotum with two rows of bristles, larger second row with fine intercalary bristles, preceded by numerous scattered small bristles; mesonotal flange with 2-4 pseudo- setae per side. Mesepisternum with three unequal bristles in irregular row along pleural rod. Mesepimeron with two rows of bristles, three in first and two larger ones in second. Lateral metanotal area with irregu- Figs. 6-7. Meringis facilis: (6) P>.5). Comparisons of difference between P. m. artemisiae measurements from my study to the expected ratio mean of 80 percent (X- = 2.75, 5df., .8>P>.5) and my P. m. sonoriensis data with its ex- pected ratio mean of 73 percent (X- = 3.80, Idf., .1>P>.05) also proved to be non- significant. With the range of P. m. serratus thus es- tablished a number of pertinent questions arise. Foremost of these is what parameters are serving to maintain the subspecies P. m. serratus? Cover and overstory vegetation were measured and recorded using the line intercept method (Smith 1973) at each trapping site. Where P. m. serratus and P. m. sonoriensis ranges meet an apparent boundary is provided because P. m. sono- riensis is found primarily in sagebrush and bunchgrass associations and P. m. serratus remains on the forested hillsides and creek- bottoms. No gross vegetational factors were found to deliniate a range separation where P. m. Table 1. Body and cranial measurements of P. maniculatus specimens. Assignment of specimens to subspecies was based primarily on tail to head and body ratios as well as body markings. Measurements in P. m artemisiae P. m. serratus P. m. sonoriensis Intergrades' millimeters X C.L.^ X C.L. X C.L. X C.L. Total length 170.5 (167.8-173.2) 183.4 (181.0-185.8) 170.1 (166.2-174.0) 167.8 (164.2-171.2) Tail length 76.1 (74.3-77.9) 87.4 (86.0-88.8) 72.8 (70.9-74.7) 74.3 (72.3-76.3) Hindfoot length 21.0 (20.7-21.3) 23.4 (23.2-23.6) 20.6 (20.3-20.9) 21.6 (21.1-22.1) Ear length 18.9 (18.4-19.4) 22.8 (22.5-23.1) 17.8 (17.2-18.4) 20.5 (19.9-22.1) Ratio, taihhead and body 80.3 (78.6-83.0) 91.0 (89.5-92.5) 74.9 (73.3-76.5) 79.5 (76.7-82.3) Zygomatic breadth 13.1 (12.9-13.3) 13.4 (13.2-13.6) 13.4 (13.2-13.6) 13.3 (13.1-13.5) Occipital-nasal length 26.4 (26.1-26.7) 27.0 (26.7-27.3) 26.6 (26.2-27.0) 26.7 (26.3-27.1) Cranium breadth 11.2 (11.0-11.4) 10.9 (10.7-11.1) 11.0 (10.9-11.1) 11.0 (10.9-11.1) Maxillary tooth row length 3.9 (3.7-4.1) 3.9 (3.8-4.0) 3.8 (3.7-3.9) 3.9 (3.8-4.0) Incisor-maxillary diastema 7.4 (7.2-7.6) 7.1 (7.0-7.2) 6.9 (6.7-7.1) 7.2 (7.0-7.4) Palate length 4.2 (4.0-4.4) 4.0 (3.9-4.1) 4.3 (4.2-4.4) 4.3 (4.1-4.5) Incisive foramen length 5.9 (5.8-6.0) 5.9 (5.8-6.0) 5.5 (5.3-5.7) 5.8 (5.6-6.0) Post-palital length 9.7 (9.5-9.9) 9.6 (9.4-9.8) 9.2 (9.0-9.4) 9.4 (9.2-9.6) Interorbital constriction 4.0 (3.9-4.1) 4.2 (4.1-4.3) 3.9 (3.8-4.0) 4.1 (4.0-4.2) Nasal length 11.5 (11.3-11.7) 11.0 (10.8-11.2) 10.8 (10.6-11.0) 11.0 (10.8-11.2) Baculum length 8.7 (8.5-8.7) 10.9 (10.6-11.2) 8.4 (7.8-9.0) 9.0 (8.0-10.0) 'Assumed intergrades P. m. serratus 'Confidence limits at the 95 percent X P. m. sonoriensis taken at trapping Site 22 (Fig. 1). confidence level. December 1978 Thurow: Peromyscus Ecology 471 artemisiae and P. m. serratus ranges meet. Further investigation of such factors as ge- ographic barriers, chmatic patterns, and soil types offered no solution. One environmen- tal factor was discovered to have a high correlation with the range of P. m. serratus, that being an isobar illustrating the average date (30 June) of the last killing frost (Year- book of Agriculture 1941). An apparent discrepancy between this frost date isobar and the range of P. m. ser- ratus appears in the eastern portion of the range approaching Challis, Idaho. However, since no weather data of this nature are available from that area the frost line could in fact include this drainage in question. Another one of many possibilities is that P. m. serratus is a glacial remnant, having sur- vived that geologic time period through iso- lation in the rugged terrain of the Sawtooth Mountains. Physical, physiological, and be- havioral barriers to breeding between P. m. artemisiae and P. m. serratus should be in- vestigated. Until the isolating mechanisms between these gene pools is identified, the subspecific ranking of P. rn. serratus will re- main in doubt. Further research, in- vestigating these and other aspects, would certainly be a warranted step to gaining a fuller knowledge of this restricted member of North American fauna. I thank Drs. Jerran T. Flinders and Er- nest D. Abies for their appreciated guidance throughout various stages of this project and Table 2. Average ratios (expressed as percent) of tail to head and body measurements of all adult P. maniculatus specimens from 22 trapping sites in central Idaho. Site number Sample area Sample size Mean Ratio (percent) of tail to head and body 7 8 9 10 11 12 13 14 15 16 17 18 19 30 mi. E. of Burgdorf 20 mi. E. of McCall 9 mi. E. of Warm Lake 5 mi. W. of Cape Horn 14 mi. W. of Challis Alturus Lake Total Peromyscus maniculatus serratus Peromyscus maniculatus artemisiae 10 mi. E. of Riggins Chamberlain airstrip Cold Meadows airstrip Big Creek airstrip Taylor Ranch-Cliff Creek Taylor Ranch-Rush Creek Flying "B" airstrip 18 mi. W. of Donnelly 5 mi. W. of Donnelly 8 mi. N.E. of Donnelly Indian Creek airstrip Alder Creek— 25 mi. N. of Challis 5 mi. W. of Lowman Total 20 9 mi. W. of Challis 21 18 mi. N.E. of Sun Valley Total P. 22 3 mi. N. of Pine Peromyscus maniculatus sonoriensis Secondary intergradation of m. serratus with P. m. sonoriensis 5 89.6 5 94.0 12 93.5 8 88.2 11 90.5 5 89.0 46 91.0 10 82.3 11 80.0 10 83.9 3 84.6 21 78.5 19 77.5 1 79.1 15 79.5 10 81.8 4 84.7 8 83.1 12 79.2 15 80.1 139 80.3 8 73.8 14 75.5 22 19 74.9 79.5 472 Great Basin Naturalist Vol. 38, No. 4 for critically reading the manuscript. I also wish to thank Drs. Donald Johnson and Ste- ven Peterson for their helpful comments and Kenneth Sowles for his continual proj- ect aid and expert flying into various back- coimtry airstrips. This work was supported through an honorarium from the Wilderness Research Center of the University of Idaho. Literature Cited Davis, W. B. 1939. The recent mammals of Idaho, pp. 288-293. Caldwell, Idaho: Caxton Printers, Inc. Elliott, C. L. 1976. The effect of altitude upon vari- ous physical and reproductive factors in a popu- lation of Peromyscus maniculatiis. Under- graduate research report. Student Wilderness Study Program. 13 pp. Unpublished manuscript. Hall, E. R., and K. R. Kelson. 1959. The mammals of North America II: 612-624. New York: Ron- ald Press Co. Mayr, E., E. G. Linsley, and R. L. Usinger. 1953. Methods and principles of systematic zoology. New York: McGraw-Hill Book Co. ,328pp. Osgood, W. H. 1909. Revision of the mice of the American genus Peromyscus. N. Amer. Fauna 28:1-285. Smith, R. L. 1974. Ecology and field biology, pp. 695-696. New York: Harper and Row Pub. U.S. Department of Agriculture. 1941. Yearbook of agriculture: climate and man. pp. 829-840. Washington D.C.: United States Government Printing Office. THE EYE FLUKE DISEASE (DIPLOSTOMATOSIS) IN FISHES FROM UTAH Richard A. Heckmann' and James R. Palmieri' Abstract.— During 1976 and 1977, 798 fish representing 11 species from eight collection sites in Utah were examined for metacercariae of Diplostomtnn rfpathaceum, which causes the fish eye fluke disease, diplostomatosis. Eight species were infected. The infection rate ranged from 7 percent of 46 Salmo gairdneri from Soldiers Creek Reservoir to 100 percent in 7 species of fish from four collection sites. Summary charts for four years of data are given for collection sites, piscine hosts in Utah, and fish hosts for Strawberry Reservoir, Utah. During the past four years we have been studying the incidence, hfe history, mor- phology, host-parasite relationships, and control of Diplostomum spathaceum in Utah fishes. Palmieri et al. (1977) reported on the life history and habitat analysis of the eye fluke. The incidence of infection for 1974 and 1975 was reported in a previous pub- lication (Evans et al. 1976). Diplostomwn spothaceum (Rudolfi 1819), or the fish eye fluke which causes the dis- ease diplostomatosis, was reported in Straw- berry Reservoir, Utah, by the Division of Wildlife Resources in 1973. Fish are the most common second intermediate hosts; however, infections in amphibians, reptiles, and mammals have also been reported (Fer- guson 1943). Once the cercariae have pene- trated the second intermediate host, they lose their forked tails and migrate to the lens tissue, where the metacercariae develop in 50-60 days (Erasmus 1958). Diplostoma- tosis causes cataracts of the lens tissue, due to the presence of the metacercarial stage of this parasite. Visual acuity for infected fish can be slightly hampered or lost, de- pending on the number of worms present. In addition to visual loss, fish show retarded growth and a change in food habits. In- crease in the incidence of the disease in the state has generated public and academic concern. The purpose of this paper is to report on the last two years of survey data (1976-1977) for the prevalence of the eye fluke in fish, indicate new hosts for Utah and sites of infection, and summarize four years of data for Strawberry Reservoir, Utah. Materials and Methods A survey of piscine hosts was accom- plished with the cooperation of the Utah Division of Wildlife Resources. Regional fisheries biologists provided heads of fish sampled during routine and annual in- vestigations of fish populations throughout the state. Arrangements were made to ob- tain the samples following the inventories accomplished by the state fisheries biolo- gists. Fish were collected by gill net, seine, and hook and line, then placed in ice chests until the senior author obtained the samples for study at Brigham Young University. The species was determined and recorded. In the laboratory, fish were examined for metacercariae of D. spathaceum. Each indi- vidual eye was extracted and carefully ex- amined, records of fish and lens condition determined, and individual numbers of worms found infecting the right and left eye recorded. Results A total of 798 fish were examined from eight collection sites in Utah during 1976 'Department of Zoology, Brigham Young University, Prove, Utah 84602. 473 474 Great Basin Naturalist Vol. 38, No. 4 and 1977 (Table 1). Of the 11 species of fish, 8 were infected with the metacercariae of D. spathaceum (Table 1). There are 13 known piscine hosts for the eye fluke in Utah (Table 2), which have been found at 14 sites (Table 3). From pre- Table 1. Location, fish host, and parasite infection data of Diphstomum spathaceum for Utah (1976 and 1977). Reservoir Number Percent Average number collection Hosts of infection metacercariae/host Collection date locality collected fish of fish (Range in parenthesis) 27-VI-1976 East Canyon Salmo gairdneri 19 95 7.4 (0-30) Salmo clarki 3 100 2.0(1-3) Richardsonius balteatus 6 100 9.7 (1-16) Catostomus ardens 7 100 66.7 (7-105) 27-1 V- 1976 Echo Salmo gairdneri 8 13 1 (0-1) Prosopium williamsoni 6 0 0 Cypriniis carpio 1 0 0 Catostomus ardens 2 0 0 Catostomus platyrhynchus 4 0 0 27-1 V- 1976 Rockport Salmo trutta 3 0 0 Salmo gairdneri 14 14 1.5 (0-2) Catostomus platyrhynchus 1 0 0 Gila atraria 12 25 1 (0-1) 30-IV-1976 Lost Creek Salmo clarki 13 0 0 Salmo gairdneri 6 0 0 Catostomus ardens 14 72 18.0 (0-131) Gila atraria 27 48 3.1 (0-6) 14- V- 1976 Hyrum Dam Salmo gairdneri 55 100 16.4 (3-42) Salmo trutta 5 80 4.3 (0-7) Cyprinus carpio 8 50 6.2 (0-10) 13- V- 1976 Pineview Ictalurus melas 9 78 2.6 (0-5) Salmo gairdneri 16 86 2.7 (0-7) Cyprinus carpio 2 50 4.0 (0-4) Gila atraria 12 100 3.1(1-7) Lepomis macrochirus 12 100 3.6 (1-10) 19-V-1976 Soldiers Salmo gairdneri 46 7 1.3 (0-2) Creek Salvelinus fontinalis 1 0 0 20- V- 1976 Strawberry Salmo clarki 19 84 3.0(0-11) Salmo gairdneri 82 100 30.1(1-200-1-) Catostomus platyrhynchus 15 100 151.5 (58-200 -(-) Richardsonius balteatus 18 100 6.3 (1-23) 28- V- 1976 Strawberry Richardsonius balteatus 25 100 5.9 (1-72) 28-XI-1976 Strawberry Salmo clarki 30 80 2.6 (0-18) Gila atraria 34 97 3.6(0-10) Salmo gairdneri 52 100 85.4(3-200-1-) Richardsonius balteatus 9 89 2.8 (0-6) 3- VI- 1977 Strawberry Salmo gairdneri 27 100 34.2 ( 1-200 -^) Salmo clarki 15 93 3.4 (0-12) Richardsonius balteatus 6 100 4.1(1-15) Gila atraria 20 90 3.6 (0-9) Catostomus platyrhynchus 10 100 189.1(62-200-1-) 15- VI- 1977 Strawberry Salmo gairdneri 31 100 35.2(1-200-1-) Salmo clarki 14 86 2.8 (0-7) Gila atraria 16 100 3.4 (1-14) Catostomus platyrhynchus 10 100 176.2(58-200-1-) Richardsonius balteatus 10 90 3.5 (0-9) Catostomus ardens 12 100 12.3 (3-17) Salvelinus fontinalis 1 100 168(168) Rhinichthyes cataractae 10 0 0 December 1978 Heckmann, Palmieri: Fish Eye Fluke 475 vious published data (Palmieri et al. 1976b, 1977) we have added 3 additional hosts and included 4 more sites for diplostomatosis in Utah. The rate of infection varied from 7 percent for a Salmo gairdneri from Soldiers Creek Reservoir to 100 percent found in seven species of fish {Salmo gairdneri, Rich- ardsoniiis balteatus, Catostomus ardens, Gila atraria, Lepomis macrochirus, Catostomus platyrhynchus, Salvelinus fontinalis) from 4 of the 8 collection sites (Table 1). To date, a total of 1637 fish, including 22 species from 21 collection sites throughout Utah, have been checked for diplostoma- tosis. Although the number of metacercariae in the right and left lenses of individual hosts seldom was identical, no significant lens preference was noted. Examination of data related to host-sex susceptibility to in- fection by metacercariae of D. spathaceum revealed no significant correlation. The rate of infection of D. spathaceum across Utah is high, as exemplified by sam- ples from Strawberry Reservoir (Table 4). For this body of water, three species of fish, Salmo gairdneri, Salvelinus fontinalis, and Catostomus platyrhynchus, have large num- bers of metacercariae which probably im- pair vision. Discussion Samples of fish from eight collection sites in Utah were examined for Diplostomum spathaceum metacercariae. These data svib- stantiated previous results (Palmieri et al. 1977) that the rate of infection across Utah is high and widespread. Three additional piscine hosts and four reservoirs are includ- ed on the lists for intermediate hosts and habitats respectively. There are sites in Utah where the fluke has not been found. The reason for this is indicated by the lack of the needed shoreline and bottom vegeta- tion, so important for the development, growth, and reproduction of snails. The lack of vegetation, snail, and gull hosts, as well as the presence of low water temperatures, probably accounts for the low infection rate in high alpine lakes. The lack of shoreline vegetation is another reason for the absence of the disease in reservoirs and lakes at lower elevations. There has been an increase in the in- cidence of metacercariae for fish in Straw- berry Reservoir (Table 4). Salmo gairdneri has gone through an increase in prevalence and numbers during the last four years. The last samples of this piscine host suggest a stabilization in number of worms per eye (range 1 to 200 -I- ). The Utah chub, Gila atraria, has become prominent in recent gill net samples from Strawberry Reservoir. All Table 2. Summary of fish hosts positive for metacer- cariae of Diplostomum spathaceum in Utah. Range of infection with Number metacercariae Fish host examined (percent) Catostomus ardens 42 72-100 Catostomus discobolus 26 0-5.8 Catostomus platyrhynchus 117 60.5-100 Cyprinus carpio 8 0-50 Gila atraria 210 0-100 Ictalurus melas 9 0-78 Lepomis macrochirus 12 0-100 Micropterus salmoides 61 0-49 Salmo clarki 179 0-100 Salmo gairdneri 792 0-100 Salmo trutta 21 0-100 Salvelinus fontinalis 14 0-100 Richardsonius balteatus 91 0-100 Table 3. Collecting sites for intermediate hosts (fish) of Diplostomum spathaceum in Utah. Collection site County Ash and LaVerkin creeks Washington Deer Creek Reservoir Wasatch Echo Reservoir" Morgan East Canyon Reservoir" Davis Fish Lake" Sevier Flaming Gorge Reservoir" Daggett Hyrum Reservoir" Cache Kolob Reservoir Washington Lake Powell Kane and San Juan Lost Creek Reservoir" Summit Mantua Reservoir" Box Elder Mirror and Lost lakes Summit Nine-Mile Reservoir" Sanpete Otter Creek Reservoir" Piute Palisade Lake" Sanpete Pineview Reservoir" Weber Rockport Reservoir* Summit Scofield Reservoir Carbon Soldiers Creek Reservoir" Wasatch Strawberry Reservoir" Wasatch Utah Lake Utah 'Contain infected fish 476 Great Basin Naturalist Vol. 38, No. 4 the chub from the last sample were infected with limited mimbers (1 to 14 per host) of metacercariae. There has been no signifi- cant change in the infection rate for Salmo clarki, Catostomus platyrhynchus, and Salve- lintis fontinalis during the four-year sam- pling period for Strawberry Reservoir (Table 4). The pathological effects of Diplostomum spathaceum upon the fish host are many. Examination of those fish blinded with cata- ract and containing a heavy burden of lar- val metacercariae revealed stunted growth (length, girth, and weight), abnormal feed- ing behavior (lack of response to visual stimuli), and decreased vital acuity (Palmieri et al. 1977). Ashton et al. (1969) reported that larvae migrate to the eye via vascular- venous channels and showed that the lens, vitreous, or cortex of the eye may be pro- Table 4. Fish sampled from Strawberry Reservoir, four years. Average number of Host Date of Number of Percent metacercariae species sample fish infection (Range in parenthesis) Salmo clarki 20-IV-74 19 48 3.4 (0-5) (Cutthroat trout) 6-IX-74 9 56 4.2 (0-6) 5-VI-75 75 88 7.3 (0-23) 30-X-75 10 90 2.1 (0.5) 20-V-76 19 84 3.0 ((0-11) 28-XI-76 30 80 2.6(0-18) 3-VI-77 15 93 3.4 (0-12) 15-VI-77 14 86 2.8 (0-7) Salmo gairdneri 20-VI-74 4 75 8.6 (0-20) (Rainbow trout) 9-VIII-74 49 98 13.4 (0-22) 6-XI-74 71 98 11.9(0-27) 5-VI-75 35 97 40.0(0-200-1-) 8-VII-75 2 100 33.5 (1-200-1-) lO-VII-75 2 100 2.0(1-13) 30-X-75 53 93 14.5 (0-54) 20-V-76 82 100 30.1 ( 1-200 -h) 18-XI-76 52 100 85.4(1-2004-) 3-VI-77 27 100 34.2(1-200-1-) 15-VI-77 31 100 35.2(1-200-1-) Salvelinus fontinalis 30-X-75 3 100 32.6 (5-31) (Brook trout) 5-VI-75 4 100 140.9(25-200-1-) 15-VI-77 1 100 168(168) Catostomus platyrhynchus 20-VI-74 8 100 102.9(35-200-1-) (Mountain sucker) 6-XI-74 15 100 81.1 (6-200 -h) 5-VI-75 21 100 112.0(25-200-1-) 30-X-75 9 100 159.6 (.39-200 -h) 20-V-76 15 100 151.5(25-200-)-) 3-VI-77 10 100 189.1(62-200-1-) 15-VI-77 10 100 176.2 (58-200 -^) Catostomus ardens 5-VI-75 11 100 8.8 (2-14) (Utah sucker) 15-VI-77 12 100 12.3 (3-17) Richardsonius balteatus 5-VI-75 5 80 30.6 (0-57) (Redside shiner) 30-X-75 22 95 7.2 (0-16) 20-V-76 18 100 6.3(1-23) 28-V-76 25 100 5.9 (1-22) 28-XI-76 9 89 2.8 (0-6) 3-VI-77 6 100 4.1 (1-15) 15-VI-77 10 90 3.5 (0-9) Gila atraria 28-XI-76 34 80 2.6(0-18) (Utah chub) 3-VI-77 20 90 3.6 (0-9) 15-VI-77 16 100 3.4 (1-14) Rhinichthyes cataractae 15-VI-77 10 0 00 December 1978 Heckmann, Palmieri: Fish Eye Fluke 477 liferated with metacercariae. In older fish, chronic infections produced subacuate in- flammatory reactions in the vitreous in- volving heterophils and eosinophils, and macrophages with ingested lens material oc- curred. There are many possible techniques to in- vestigate concerning the control of diplo- stomatosis. One that shows promise is bio- logical control by the use of a protozoan hyperparasite, Nosema strigeoidea (Protozoa: Microsporida). Hussey reported in 1971 the above species of microsporidia to be host specific for hyperparasitizing sporocysts of Diplostonium spathaceu7n. Palmieri et al. (1976a, 1976c) substantiated Hussey 's work for the eye fluke in Utah. Spores of N. strigeoideae were introduced to laboratory-reared snails (Lymanaea au- ricularia) containing sporocysts of D. spa- thaceum. The spores selectively attack the mother and daughter sporocysts as well as the developing cercarial embryos and re- tard, disfigure, and disrupt normal cercarial development. The microsporidian spores have no outward pathological effect upon the molluscan host. Further work is needed to determine the potential use of this con- trol method for diplostomatosis and in- vestigations pertaining to resistant fish hosts, stimulus for cercarial migration, immune re- sponses, and the potential human health hazard. Acknowledgments The authors wish to thank the Utah Divi- sion of Wildlife Resources for specimen col- lection and the National Science Founda- tion (Grant EPP 75-04396) for financial support. Literature Cited AsHTON, N., N. Brown, and D. Easty. 1969. Trema- tode cataract in freshwater fish. J. Small Anim. Pract. 10:471-478. Erasmus, D. A. 1958. Studies on the morphology, biol- ogy, and development of a strigeid cercariae (cercaria X Baylis). Parasitology 48;.312-335. Evans, R. S., R. A. Heckmann, and J. R. Palmieri. 1976. Diplostomatosis in Utah. Proc. Utah Acad. Sci. 53(l):20-25. Ferguson, M. 1943. Development of eye flukes of fish- es in the lenses of frogs, turtles, birds and mam- mals. J. Parasitol. 29:136-142. Hussey, K. L. 1971. A microsporidean hyperparasite of strigeoid trematodes Nosema strigeoidea sp. n. J. Parasitol. 18(4):676-679. Palmieri, J. R., R. A. Heckmann, and R. S. Evans. 1977. Life history and habitat analysis of the eye fluke Diplostonium spathaceum (Trematoda; Diplostomatidae) in Utah. J. Parasitol. 63:427-429. 1976b. Life cycle and incidence of Diplosto- miim spathaceum Rudolphi (1819) (Trematoda: Diplostomatidae) in Utah. Great Basin Nat. 36(l):86-96. Palmieri, J. R. and R. A. Heckmann. 1976a. Potential biological control of diplostomatosis {Diplosto- i7ium spathaceum) in fishes by hyperparasitism. Proc. Utah Acad. Sci. 53(1):17-19.' Palmieri, J. R., R. A. Heckmann, and A. Call 1976c. Experimental biological control of the eye fluke Diplostomum spathaceum by a hyperparasite, Nosema strigeoidea (Protozoa: Microsporida). J. Parasitol. 62(2):325-326. BITE OF WOLF SPIDER Dorald M. Allred' Abstract.— A man bitten on the finger by a wolf spider, Lycosa sp., experienced temporary bleeding and tin- gling of the finger, but no other symptoms occurred. On September 11, 1978, a male construc- tion worker about thirty years of age was admitted to the emergency ward of the Utah Valley Hospital, Provo, Utah, for treatment of a spider bite. He had grasped the underside of a broken concrete block and felt a pain like a thorn prick on the back of his middle finger. A wolf .spider was found on the underside of the block. The worker had apparently imprisoned it by pressing one of its legs against the block. The paired cheliceral wounds were situat- ed transversely on the finger next to the fin- gernail. They bled freely for a few minutes, and the victim experienced a temporary tingling of his finger which lasted only for a few minutes. He was discharged without treatment other than application of a dis- infectant. No further symptoms were expe- rienced. Unfortunately a co-worker struck the spi- der with a hammer. The remains brought in were identified as a Lycosa female, prob- ably carolinensis based on size and color pattern of the legs and cephalothorax. '290 Life Science Museum, Brigham Young University, Provo, Utah 84602. 478 INDEX TO VOLUME 38 The genera and species described as new to science in this volume appear in bold type in this index. A comparative analysis of the diatom flora on the snail Ampullaria cuprina from the Goshen Ponds, Utah, p. 211. A new subspecies of the snake Lichnnura trivirgata from Cedros Island, Mexico, p. 411. Allelopathic effects of Bur Buttercup tissue on germination and growth of various grasses and forbs in vitro and in soil, p. 90. AUred, Dorald M., article by, p. 478. An addition to the amphibian fauna of Cali- fornia, p. 429. An annotated list of the aquatic insects of southeastern Idaho, Part III. Ephem- eroptera, p. 55. An ecological and taxonomic investigation of Peromysciis maniculatus serratiis in Idaho, p. 469. Andersen, Perron L. and Graham R. Bullick, article by, p. 369. Astragalus lentiginosus var. wahweapensis, p. 286. Astragalus montii, p. 11. Astragalus subcinereus var. basalticus, p. 302. Atwood, N. Duane, Stanley L. Welsh, and Joseph R. Murdock, article by, p. 125. Axler, R. P., R. M. Gersberg, and L. J. Paul- son, article by, p. 187. Basidiomycetes that decay junipers in Ari- zona II, p. 42. Baumann, Richard W., and Bill P. Stark, ar- ticle by, p. 97. Bite of wolf spider, p. 478. Bright, Donald E., article by, p. 71. Brotherson, J. D., G. Nebeker, M. Skougard, and J. Fairchild, article by, p. 19. Brotherson, Jack D., Mark C. Whiting, and Samuel R. Rushforth, article by, p. 31. Buchanan, Bruce A., K. T. Harper, and Neil C. Frischknecht, article by, p. 90. Bullick, Graham R., and Perron L. Ander- sen, article by, p. 369. Campbell, W. P., and E. A. Rechel, article by, p. 456. Conophthorus echinatae, p. 398. Contributions to the biology of the wound- fin, Plagopterus argentissimus (Pisces: Cynrinidae), an endangered species, p. 463. Correction: Rhamnus cathartica L. is not Primus nigra Ait. from Alberta, p. 123. Cross, Jeffrey N., article by, p. 463. Current status of cutthroat trout subspecies in the western Bonneville Basin, p. 193. Differential parasitism of Peromysciis ma- niculatus and Peromyscus truei, by Cut- erebra larvae, p. 51. Distributional records for the Colorado flora, p. 208. Duff, Donald A., and Terry J. Hickman, ar- ticle by, p. 193. Eads, Richard B., article by, p. 447. Effect of irrigation on survival of third-stage Haemonchus contortus larvae (Nematoda: Trichostrongylidae), p. 369. Endangered and threatened plants of Utah: A reevaluation, p. 1. Engstrom, Mark D., and Eugene D. Ple- harty, article by, p. 180. Environmental interaction in summer algal communities of Utah Lake, p. 31. Eridiphas sleveni macrosensis, p. 407. Evans, Howard E., and William L. Rubink, article by, p. 59. Pairchild, J., J. D. Brotherson, G. Nebeker, and M. Skougard, article by, p. 19. Peddema, C, D. Wilken, W. Harmon, and H. Harrington, article by, p. 208. Pleharty, Eugene D., and Mark D. Eng- strom, article by, p. 180. Fluctuation of a marsh habitat and the re- productive strategy of the Yellowheaded Blackbird, p. 85. Frischknecht, Neil C, Bnice A. Buchanan, and K. T. Harper, article by, p. 90. Gersberg, R. M., R. P. Axler, and L. J. Paul- 479 480 Great Basin Naturalist Vol. 38, No. 4 son, article by, p. 187. Gilbertson, R. L., and J. P. Lindsey, article by, p. 42. Growth of Dipodomys ordii (Rodentia: Het- eromyidae), p. 215. Halford, Douglas K., and Jere B. Millard, article by, p. 64. Harmon, W., D. Wilken, C. Feddema, and H. Harrington, article by, p. 208. Harper, K. T., Bruce A. Buchanan, and Neil C. Frischknecht, article by, p. 90. Harrington, H., D. Wilken, W. Harmon, and C. Feddema, article by, p. 208. Havvksworth, Frank G., and Robert L. Mathiasen, article by, p. 89. Heckmann, Richard A., and James R. Pal- mieri, article by, p. 473. Hedijsarum occidentale var. canone, p. 314. Hickman, Terry J., and Donald A. Duff, ar- ticle by, p. 193. Hosts of juniper mistletoe at Walnut Can- yon National Monument, Arizona, p. 89. Jenkins, M. Alan, James A. Mosher, Glayton M. White, and Joseph R. Murphy, article by, p. 438. Johnson, Mark K., and James G. Mac- Cracken, article by, p. 222. Jorgensen, Clive D., H. Duane Smith, and Gary H. Richens, article by, p. 215. Kaiparowits flora, p. 125. Kay, P. A., and C. G. Oviatt, article by, p. 49. Kennedy, Joseph L., and Paul A. Kucera, article by, p. 181. Kucera, Paul A., and Joseph L. Kennedy, article by, p. 181. Kucera, Paul A., article by, p. 203. Lederer, Roger J., article by, p. 85. Leptosporomyces juniperinus, p. 45. Lichanura trivirgato bostici, p. 411. Lindsey, J. P., and R. L. Gilbertson, article by, p. 42. Llewellyn, Jeffrey B., article by, p. 51. Lyniantor alaskanus, p. 399. MacCracken, James G., and Mark K. John- son, article by, p. 222. Mathiasen, Robert L., and Frank G. Hawks- worth, article by, p. 89. Maughan, O. Eugene, article by, p. 115. Meringis disparalis, p. 447. Meringis facilis, p. 451. Millard, Jere B., and Douglas K. Halford, article by, p. 64. Minshall, G. Wayne, and Robert L. Newell, article by, p. 55. Morphometry of sculpins (Cottus) in the Clearwater drainage, Idaho, p. 115. Mosher, James A., Clayton M. White, Jo- seph R. Murphy, and M. Alan Jenkins, article by, p. 438. Mountain ants of Nevada, p. 379. Mullen, David A., and Robert C. Stebbins, article by, p. 429. Murdock, Joseph R., Stanley L. Welsh, and N. Duane Atwood, article by, p. 125. Murphy, Joseph R., James A. Mosher, Clay- ton M. White, and M. Alan Jenkins, ar- ticle by, p. 438. Nebeker, G., J. D. Brotherson, M. Skougard, and J. Fairchild, article by, p. 19. Neoperlo carlsoni, p. 97. Neoperla catharae, p. 98. Neoperla choctaw, p. 99. Neoperla freytagi, p. 101. Neoperla gaufini, p. 101. Neoperla stewarti, p. 104. Newell, Robert L., and G. Wayne Minshall, article by, p. 55. New range and a new subspecies for the snake Eridiphas sleveni, p. 406. New species of nearctic Neoperla (Plecop- tera: Perlidae), with notes on the genus, p. 97. New synonymy, new species, and taxonomic notes of North American Pityophthorus (Coleoptera: Scolytidae), Part III, p. 71. New synonymy and new species of Ameri- can bark beetles (Coleoptera: Scolytidae), Part VII, p. 397. Observations on the prey and nests of seven species of Cerceris (Hymenoptera: Sphe- cidae), p. 59. Ottley, John R., article by, p. 411. Ottley, John R., and Wilmer W. Tanner, ar- ticle by, p. 406. Oviatt, C. G., and P. A. Kay, article by, p. 49. Oxytropis oreophila var. juniperina, p. 339. Palmieri, James R., and Richard A. Heck- mann, article by, p. 473. Paulson, L. J., R. P. Axler, and R. M. Gers- berg, article by, p. 187. Piniis longaeva in the Stansbury Mountains, Utah, p. 49. Pityophthorus carinatus, p. 73. Pityophthorus cavatus, p. 74. December 1978 Index 481 Pityophthonis carinatus monticolae, p. 73. Pitijophthoriis delicatus, p. 399. Pityophthonis euterpes, p. 75. Pityophthonis hesperius, p 76. Pityophthonis imperus, p. 76. Pityophthonis indigus, p. 398. Pityophthonis intentus, p. 77. Pityophthonis laticeps, p. 78. Pityophthonis malleatus, p. 79. Pityophthonis mesembria, p. 80. Pityophthonis montezumae, p. 81. Pityophthonis rubidus, p. 400. Pityophthonis scalptus, p. 82. Pityophthonis trepidus, p. 83. Pityophthonis vesculus, p. 401. Pityophthonis vespertinus, p. 83 Plants of Navajo National Monument, p. 19. Primary productivity in meromictic Big Soda Lake, Nevada, p. 187. Range extension of Onychomys torridtis longicaudiis (Rodentia: Cricetidae) in northwestern Nevada, p. 180. Raptors of the Uinta National Forest, Utah, p. 438. Rechel, E. A., and W. F. Campbell, article by, p. 456. Reproductive biology of the tui chub, Gila bicolor, in Pyramid Lake, Nevada, p. 203. Richins, Gary H., H. Duane Smith, and Clive D. Jorgensen, article by, p. 215. Rvibink, William L., and Howard E. Evans, article by, p. 59. Rushforth, Samuel R., Larry L. St. Clair, and Lorin E. Squires, article by, p. 211. Rushforth, Samuel R., Mark C. Whiting, and Jack D. Brotherson, article by, p. 31. St.Clair, Larry L., Lorin E. Squires, and Samuel R. Rushforth, article by, p. 211. Scolytodes comitabilus, p. 401. Scolytodes crinalis, p. 402. Scolytodes crinitus, p. 402. Scolytodes decorus, p. 402. Scolytodes genialis, p. 403. Scolytodes habilis, p. 403. Scolytodes libidus, p. 404. Scolytodes perpusillus, p. 404. Scolytodes semipunctatus, p. 405. Shaw, R. Keith, article by, p. 123. Similarity between pronghorn and mule deer fecal pellets, p. 222. Skougard, M., J. D. Brotherson, G. Nebeker, and J. Fairchild, article by, p. 19. Smith, H. Duane, Gary H. Richins, and Clive D. Jorgensen, article by, p. 215. Squires, Lorin E., Larry L. St.Clair, and Samuel R. Rushforth, article by, p. 211. Stark, Bill P., and Richard W. Baumann, ar- ticle by, p. 97. Stebbins, Robert C, and David A. Mullen, article by, p. 429. Tanner, Wilmer W., and John R. Ottley, ar- ticle by, p. 406. The effects of radiation from uranium mill tailings on Tradescajitia, p. 456. The eye fluke disease (Diplostomatosis) in fishes from Utah, p. 473. The reproductive ecology of the Tahoe sucker, Catostomus tahoensis, in Pyramid Lake, Nevada, p. 181. Thurow, Thomas L., article by, p. 469. Two new species of fleas of the genus Me- ringis (Siphonaptera: Hystrichopsyllidae), p. 447. Utah flora: Fabaceae (Leguminosae), p. 225. Vertebrate fauna of a radioactive leaching pond complex in southeastern Idaho, p. 64. Vertical distribution of adult fish in Pyra- mid Lake, Nevada, p. 417. Vigg, Steven, article by, p. 417. Welsh, Stanley L., articles by, p. 1, 225. Welsh, Stanley L., N. Duane Atwood, and Joseph R. Murdock, article by, p. 125. Wheeler, George C, and Jeanette Wheeler, article by, p. 379. Wheeler, Jeanette, and George C. Wheeler, article by, p. 379. White, Clayton M., James A. Mosher, Jo- seph R. Murphy, and M. Alan Jenkins, article by, p. 438. Whiting, Mark C, Jack D. Brotherson, and Samuel R. Rushforth, article by, p. 31. Wilken, D., W. Harmon, C. Feddema, and H. Harrington, article by, p. 208. Wood, Stephen L., article by, p. 397. Xyleborus devexulus, p. 398. The Great Basin Naturalist VOLUME 38, 1978 Editor: Stephen L. Wood Published at Brigham Young University, by Brigham Young University NOTICE TO CONTRIBUTORS Original nianiiscripts in English pertaining to the biological natnral history of western North America and intended for publication in the Clrcat Basin Xatiiralist should be di- rected to Brigham Young University, Stephen L. Wood, Editor, Great Basin Xafiiralist, Provo, Utah 84602. Those intended for the C^reat Basin Xattiralist Metnoirs should be sim- ilarly directed, but tiiese manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. Thev should be typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. Use a recent issue of either journal as a format, and the Council of Biolo^ii Edi- tors Sti/lc Manual, Third Edition (AIBS 1972) in preparing the manuscript. An abstract, about 3 percent as long as the text, but not exceeding 200 words, written in accordance with Biological Abstracts guidelines, should precede the introductory paragraph of each ar- ticle. Authors mav recommend one or two reviewers for their article. All manuscripts re- ceive a critical peer review by specialists in the subject area of the manuscript imder con- sideration. Manu.scripts that are accepted and that are less than 100 printed pages in length will automatically be assigned to the Great Basin Xaturalist. Those manuscripts larger than 100 printed pages in length will be considered for the Memoirs series. Illustrations and Tables. All illustrations and tables should be made with a view toward having them appear within the limits of the printed page. Illustrations that form part of an article should accompany the manuscript. Illustrations should be prepared for reduction by the printer to either a single-column (2 5/8 inches) or double-column (5 1/2 inches) width, with the length not exceeding 7 1/2 inches. Costs Borne hi/ Contributor. Contributors to the Great Basin Xaturalist should be pre- pared to donate from $10 to $30 per printed page toward publication of their article (in addition to reprint costs outlined in the schedule below). Authors publishing in the Great Basin Xaturalist Memoirs mav be expected to contribute $35 per printed page in addition to the cost of printed copies thev purchase. No printed copies are furnished free of charge. A price list for reprints and an order form are sent with the gallev proof to contributors. Reprint Schedule of the Great Basin Xaturalist 100 copies 200 copies 300 copies Great Basin Xaturalist Memoirs No. 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodbury, H. H. Frost. $10. No. 2 lutermountain biogeographv: A symposium. By K. T. Harper, J. L. Reveal, et al. $15. 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each $20 $24 $28 $32 $36 $40 additional 28 32 36 40 44 48 2 pp. 36 40 44 48 52 56 $4 TABLE OF CONTENTS Effect of irrigation on survival of third-stage Haemonchus contortus larvae (Nema- toda: Trichostrongylidae). Graham R. Bullick and Ferron L. Andersen 369 Mountain ants of Nevada. George C. Wheeler and Jeanette Wheeler 379 New synonymy and new species of American bark beetles (Coleoptera: Scoly- tidae). Part VII. Stephen L. Wood 397 New range and a new subspecies for the snake Eridiphas slevini. John R. Ottley and Wilmer W. Tanner 406 A new subspecies of the snake Lichanura trivirgata from Cedros Island, Mexico. John R. Ottley 411 Vertical distribution of adult fish in Pyramid Lake, Nevada. Steven Vigg 417 An addition to the amphibian fauna of California. David A. Mullen and Robert C. Stebbins 429 Raptors of the Uinta National Forest, Utah. James A. Mosher, Clayton M. White, Joseph R. Murphy, and M. Alan Jenkins 438 Two new species of fleas of the genus Meringis (Siphonaptera: Hystrichopsyllidae). Richard B. Eads 447 The effects of radiation from uranium mill tailings on Tradescantia. E. A. Rechel and W. F. Campbell 456 Contributions to the biology of the woundfin, Plagopterus argentissimus (Pisces: Cyprinidae), an endangered species 463 An ecological and taxonomic investigation of Peromyscus maniculatus serratus in Idaho. Thomas L. Thurow 469 The eye fluke disease (Diplostomatosis) in fishes from Utah. Richard A. Heckmann and James R. Palmieri 473 Bite of wolf spider. Dorald M. Allred 478 * cincme Bookbinding Co.. Inc. 100 Cambridge St. Charlestcwn. MA 02129 3 2044 072 231 020