310 g HARVARD UNIVERSITY Library of the Museum of Comparative Zoology € GREAT BASIN NATURALIST January 31, 1983 Brigham Young University iviUS. COMP. ZOOL LIBRARY UMiVEJRSiTY GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Chairman, Botany; James R. Barnes, Zoology; Hal L. Black, Zoology; Stanley L. Welsh, Botany; Clayton M. White, Zoology. All are at Brig- ham Young University, Provo, Utah 84602. Ex Officio Editorial Board Members. Bruce N. Smith, Dean, College of Biological and Agricul- tural Sciences; Norman A. Darais, University Editor, University Publications. Subject Area Associate Editors. Dr. Noel H. Holmgren, New York Botanical Garden, Bronx, New York 10458 (Plant Taxonomy). Dr. James A. MacMahon, Utah State University, Department of Biology, UMC 53, Lo- gan, Utah 84322 (Vertebrate Zoology). Dr. G. Wayne Minshall, Department of Biology, Idaho State University, Pocatello, Idaho 83201 (Aquatic Biology). Dr. Ned K. Johnson, Museum of Vertebrate Zoology and Department of Zoology, Uni- versity of California, Berkeley, California 94720 (Ornithology). Dr. E. Philip Pister, Associate Fishery Biologist, California Department of Fish and Game, 407 West Line Street, Bishop, California 93514 (Fish Biology). Dr. Wayne N. Mathis, Chairman, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 (Entomology). Dr. Theodore W. Weaver III, Department of Botany, Montana State University, Boze- man, Montana 59715 (Plant Ecology). The Great Basin Naturalist was founded in 1939 and has been published from one to four times a year since then by Brigham Young University. Previously unpublished manuscripts in English of less than 100 printed pages in length and pertaining to the biological natural his- tory of western North America are accepted. Western North America is considered to be west of the Mississippi River from Alaska to Panama. The Great Basin Naturalist Memoirs was es- tablished in 1976 for scholarly works in biological natural history longer than can be accom- modated in the parent publication. The Memoirs appears irregularly and bears no geographi- cal restriction in subject matter. Manuscripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist for private individuals is $16.00; for institutions, $24.00 (outside the United States, $18.00 and $26.00); and for stu- dent subscriptions, $10.00. The price of single issues is $6.00 each. All back issues are in print and are available for sale. All matters pertaining to subscriptions, back issues, or other busi- ness should be directed to Brigham Young University, Great Basin Naturalist, 290 Life Sci- ence Museum, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same office 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. See Notice to Contributors on the inside back cover. 3-83 650 64094 ISSN 017-3614 TABLE OF CONTENTS Volume 43 Number 1 - January 31, 1983 Life history of the Lahontan cutthroat trout, Salmo clarki henshawi, in Pyramid Lake, Nevada. WilHam F. Sigler, William T. Helm, Paul A. Kucera, Steven Vigg, and Gar W. Workman 1 A review of the genus Soliperla (Plecoptera: Peltoperlidae). Bill P. Stark 30 A bibliography of Colorado vegetation description. William L. Baker 45 Evaluation of a program to control hydatid disease in central Utah. Ferron L. Andersen, John R. Crellin, Craig R. Nichols, and Peter M. Schantz 65 Influence of cryptogamic crusts on moisture relationships of soils in Navajo National Monimient, Arizona. Jack D. Brotherson and Samuel R. Rushforth 73 A vascular flora of the San Rafael Swell, Utah. James G. Harris 79 Pronghom responses to hunting coyotes. Timothy D. Reynolds 88 Floristics of the upper Walker River, California and Nevada. Matt Lavin 93 Agrapyron arizonicum (Gramineae: Triticeae) and a natural hybrid from Arizona. Grant L. Pyrah 131 Species composition, distribution, and phytosociology of Kalsow Prairie, a mesic tallgrass prairie in Iowa. Jack D. Brotherson 137 Deer mouse, Peromyscus maniculatus, and associated rodent fleas (Siphonaptera) in the arctic-alpine life zone of Rocky Mountain National Park, Colorado. R. B. Eads and E. G. Campos 168 Food of larval Tui chubs, Gila bicolor, in Pyramid Lake, Nevada. David L. Galat and Nancy Vucinich 175 Number 2 - April 30, 1983 Utah flora: Compositae (Asteraceae). Stanley L. Welsh 179 Haplopappus crispus and H. zionis (Asteraceae): new species from Utah. Loran C. Anderson 358 A bouquet of daisies {Erigeron, Compositae). Stanley L. Welsh 365 New taxa in Thelesperma and Townsendia (Compositae) from Utah. Stanley L. Welsh .... 369 New Haplopappus variety in Utah (Compositae). Stanley L. Welsh and Frank J. Smith ... 371 New species of Hymenoxys and Perityle (Compositae) from Utah. Stanley L. Welsh and Elizabeth Neese 373 New variety of Stephanomeria tenuifolia (Compositae) from Utah. Sherel Goodrich and Stanley L. Welsh 375 Number 3 - July 31, 1983 A re-evaluation of the postglacial vegetation of the Laramie Basin, Wyoming-Colorado. Deborah L. Elliott-Fisk, Betty S. Adkins, and Jeanine L. Spaulding 377 Comparative life history and floral characteristics of desert and mountain floras in Utah. Patrick D. Collins, Kimball T. Harper, and Burton K. Pendleton 385 Flora of the Lower Cretaceous Cedar Mountain Formation of Utah and Colorado, part I. Paraphyllanthoxylon utahense. G. F. Thayne, W. D. Tidwell, and W. L. Stokes .. 394 Aculeata Hymenoptera of Sand Mountain and Blow Sand Mountains, Nevada. R. W. Rust, L. M. Hanks, and R. C. Bechtel 403 Status and life history notes on the native fishes of the Alvord Basin, Oregon and Nevada. Jack E. Williams and Carl E. Bond 409 Kramer Palouse natural area. Del W. Despain and Grant A. Harris 421 Winter food habits of cougars from northeastern Oregon. Chris Maser and Ronald S. Rohweder 425 A new species of Penstemon (Scrophulariaceae) from the Uinta Basin, Utah. Elizabeth Neese and Stanley L. Welsh 429 A comparative study of coyote food habits on two Utah deer herds. Jordan C. Pederson and R. Cary Tuckfield 432 A small carnivore survey technique. Tim W. Clark and Thomas M. Campbell III 438 Evaluation of Draba oligosperma, D. pectinipila, and D. juniperina complex (Cruciferae). Robert W. Lichvar 441 Presence of maxillary canine teeth in mule deer in Utah. Jordan C. Pederson 445 Comparative successional roles of trembling aspen and lodgepole pine in the Southern Rocky Mountains. Albert J. Parker and Kathleen C. Parker 447 Differential utilization of bat boxes by house wrens {Troglodytes aedon). Hal L. Black 456 Plant and soil relationships in two hydrothermally altered areas of the Great Basin. N. M. Milton and T. L. Purdy 457 Plasticity and polymorphism in seed germination of Mimulus guttatus (Scrophulariaceae). Robert K. Vickery, Jr 470 Predatory behavior of larval Ambystoma tigrinurn nebulosiim on Limnephilus (Trichoptera) larvae. Joseph R. Holomuzki 475 Notes on reproduction of the side-blotched lizard Uta stansburiana stansbiiriana in southwest Idaho. George Bakewell, Joseph M. Chopek, and Gary L. Burkholder .... 477 Observations on alpine vegetation near Schoolroom Glacier, Teton Range, Wyoming. John R. Spence and Richard J. Shaw 483 Winter stomach contents of South Dakota badgers. E. Blake Hart and Michael Trumbo . 492 A list of Utah spiders, with their localities. Dorald M. Allred and B. J. Kaston 494 Number 4 - October 31, 1983 Alpine and subalpine wetland plant communities of the Uinta Mountains, Utah. George M. Briggs and James A. MacMahon 523 Utah flora: Salicaceae. Sherel Goodrich 531 A mammalian humerus from the Upper Jurassic of Colorado. Donald R. Prothero and James A. Jensen 551 Bats of the Colorado oil shale region. Robert B. Finley, Jr., William Caire, and Dallas E. Wilhelm 554 New generic concepts in the Triticeae of the Intermountain Region: keys and comments. Mary E. Barkworth, Douglas R. Dewey, and Riley J. Atkins 561 Reproductive attributes of some Rocky Mountain subalpine herbs in successional context. David J. Schimpf and Robert L. Bayn, Jr 573 Applicability of the universal soil loss equation for southeastern Idaho wildlands. Mark E. Jensen 579 Winter stoneflies (Plecoptera) of New Mexico. Gerald Z. Jacobi and Richard W. Baumann 585 Daily and yearly movement of the Devil's Hole pupfish Cyprinodon diabolis Wales in Devil's Hole, Nevada. Thomas M. Baugh and James E. Deacon 592 A revision of the genus Microrhopala (Coleoptera: Chrysomelidae) in America north of Mexico. Shawn M. Clark 597 Flora of the Stansbury Mountains, Utah. Alan C. Taye 619 New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), part IX. Stephen L. Wood 647 Plant commimity variability on a small area in southeastern Montana. James G. MacCracken, Daniel W. Uresk, and Richard M. Hansen 660 New leafhopper species of Coelidia with a revised key and notes on homonymy and distribution (Homoptera: Cicadellidae, Coelidiinae). Mervin W. Nielson 669 Eye fluke {Diplostomum spathaceum) of fishes from the upper Salmon River near Obsidian, Idaho. Richard Heckmann 675 Evaluation of varieties in Stanleya pinnata (Cruciferae). Robert W. Lichvar 684 Some aspects of the presettlement vegetation of the Piceance Basin, Colorado. William L.Baker 687 New variety of Opuntia hasilaris (Cactaceae) from Utah. Stanley L. Welsh and Elizabeth Neese 700 Vegetative types and endemic plants of the Bryce Canyon Breaks. Robert A. Graybosch and Hayle Buchanan 701 Seasonal growth of the Tui chub, Gila bicolor, in Pyramid Lake, Nevada. Joseph L. Kennedy 713 Bird distributional and breeding records for southeastern Idaho, Utah, and adjacent regions. Clayton M. White, Herbert H. Frost, Dennis L. Shirley, G. Merrill Webb, and Richard D. Porter 717 Birds of southwestern Idaho. Daniel A. Stephens and Timothy D. Reynolds 728 New records for the vascular flora of Wyoming and Montana. Robert W. Lichvar, Robert D. Dom, and Erwin F. Evert 739 Range extensions for two dwarf mistletoes (Arceuthobium spp.) in the southwest. Robert L. Mathiasen and Kenneth H. Jones 741 First nest records for the Plain Titmouse and Blue-gray Gnatcatcher in Wyoming. Scott L. Findholt 747 A new species of Thelesperma (Asteraceae) from Wyoming. Robert D. Dorn 749 Index 751 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 43 January 31, 1983 No. 1 LIFE HISTORY OF THE LAHONTAN CUTTHROAT TROUT, SALMO CLARKI HENSHAWI, IN PYRAMID LAKE, NEVADA William F. Sigler', William T. Helm^ Paul A. Kucera\ Steven Vigg*, and Gar W. Workman' Abstract.— The Pyramid Lake Lahontan cutthroat trout (Salmo clarki henshawi) population was sampled on a monthly basis from November 1975 through December 1977. A subsample of 676 trout, stratified by fish size and lake habitat, provided biological data. The entire population is presently derived from hatchery production, stocked at lengths of approximately 75 to 300 mm. Peak annulus formation occurs in March and April, followed by the peri- od of maximum growth. Scale patterns illustrate a variable growing season. Maximum growth in length is in the first three years of life; after that males begin to grow faster than females. Males attained a greater age in our sample; i.e., the oldest male was seven years old compared to six years for females. The Pyramid Lake Lahontan cutthroat trout exhibit nearly isometric growth. The legal sport fishery removed <20,000 adult fish in 1977 (>380 mm); other decimating factors are poorly un- derstood. No evidence of the following diseases or pathogens was found in the Pyramid Lake population, presuming a carrier incidence of 2 percent at the 95 percent confidence level: infectious pancreatic necrosis, infectious hemato- poietic necrosis, viral hemorrhagic septicema, bacterial kidney disease, enteric redmouth, furunculosis, whirling dis- ease, blood fluke; however, 7 of 235 (=:;3 percent) adults sampled at the Marble Bluff fish way were positive for furunculosis. Small trout feed primarily on zooplankton and benthic invertebrates; cutthroat trout >300 mm are piscivorous, feeding almost exclusively on tui chub {Gila bicolor). The spawning migration of Pyramid Lake cutthroat trout to the Marble Bluff egg taking facility in spring 1976 and 1977 peaked in April and May. Females mature at three or four years (352-484 mm), and males mature at two or three years (299-445 mm). Mean diameter of mature eggs is 4.51 mm; both ovum size and fecundity are a function of fish size. Fecundity ranges from 1241 to 7963 eggs, with a mean of 3815. Lahontan cutthroat trout comprise <2 percent of the numerical relative abundance and <7 percent of the total fish biomass. Distribution patterns vary on a seasonal basis, with maximum activity during late fall and winter. Man- agement objectives are presented and recommendations are discussed. The Lahontan cutthroat trout (Salmo clarki The dechne and ultimate extinction of the henshawi) is unique in its abihty to withstand original strain of cutthroat trout in Pyramid the alkaline-saline waters of remnant Great Lake was caused primarily by degradation of Basin lakes. Coevolution of Pyramid Lake spawning habitat associated with diversion of Lahontan cutthroat trout in a continuous lake water out of the Truckee River-Pyramid environment for 50,000-100,000 years with Lake ecosystem (Trelease 1953). The Pyra- an abundant prey species (tui chub, Gila mid Lake trout fishery has been reestablished bicolor) resulted in a unique predator— the via hatchery propagation of Heenan, Walker, world's largest cutthroat trout (18.6 kg). and Summit lake strains of Lahontan cut- 'W. F. Sigler & Associates Inc., P.O. Box 1350, Logan, Utah 84322. 'Utah State University, Logan, Utah 84322. 'Nez Perce Tribe, Lapwai, Idaho 83540. 'Desert Research Institute, University of Nevada System, Reno, Nevada 89507. 'Utah State UniveRity, Logan, Utah 84322. Great Basin Naturalist Vol. 43, No. 1 Fig. 1. The largest post-1943 Lahontan cutthroat trout (12.7 kg), captured by Ralston Fillmore from Pyra- mid Lake, Nevada, April 1976. Photograph courtesy of Alan Ruger. throat trout, and, in the past, of cutthroat- rainbow trout {Salmo gairdneri) hybrids. The subspecies S. c. henshawi currently has "threatened" status (Deacon et al. 1979). Pyramid Lake presently supports a trophy sport fishery; the average trout retained by fishermen is 500 mm in length and weighs 1 .2 kg. In 1976, a Paiute Indian, Ralston Fill- more, captured a 12.7 kg Lahontan cutthroat trout that represents a record for the post- 1943 fishery (Fig. 1). Recent catches are evi- dence that the environment of Pyramid Lake is capable of supporting at least a limited valuable and unique fishery. However, hu- man demands on limited Truckee River wa- ter and recent droughts have jeopardized the trout in Pyramid Lake. The ethics and prior- ities of our society, as a whole, may ulti- mately decide the fate of the Pyramid Lake Lahontan cutthroat trout. Nine now discrete basins once conjoined to form vast (area 22,300 km^; maximum depth 270 m) Lake Lahontan in the northwestern Great Basin (Hubbs and Miller 1948). Pyra- mid Lake is the deepest remnant of this once great lake system that experienced several cycles of water level fluctuations during the Pleistocene Epoch (Houghton 1976). Great Basin lakes have desiccated to the present state since the last pluvial period some 10-12 thousand years before present (BP). Benson (1978) concludes via sediment analyses that Pyramid Lake was greatly reduced in size 9-5 thousand years BP, but did not become dry and had subsequently been rising until the cultural impacts of the past century. Pyramid is a graben lake approximately 40 km long and 6.5 to 16 km wide, with a north- south axis (Figure 2). At the mean 1976 ele- vation of 1157 m (United States Geological Survey 1977), Pyramid Lake has a surface area of 446.4 km^, a volume of 26.4 km^, a mean depth of 59 m and a maximum depth of 103 m (Harris 1970). Pyramid is the deep- est and most voluminous saline terminal lake in the western hemisphere (Galat et al. 1981). Pyramid Lake, located entirely within the Pyramid Lake Paiute Indian Reservation, is the terminal water body of the endorheic Truckee River system originating 193 river km upstream at oligotrophic Lake Tahoe. The evaporation loss is about 1.2 m annually. Due to transbasin diversion of the Truckee River, the lake level declined 23 m between 1905 and 1979; this amounts to a 30 percent reduction in lake volume. The lake water is highly ionic (Na+ > K+ > Mg2+ > Ca2+; CI- > HCO3 > CO32- > SO42-), with a pH of 9.2. The 1976 total dissolved solids (TDS) concentration was 5235 mg/1 at elevation 1157 m. On a worldwide perspective, 71 per- cent of some 350 saline lakes listed by McCarraher (1972) are more saline than Pyramid, but, compared to USA saline lakes. Pyramid is in the moderate range (Galat et al. 1981). During 1976 and 1977 mean surface tem- perature ranged from 6.1 to 23.1 C. As winds subside and surface water temperature in- creases, a thermocline is formed in June and lasts through December at 16 to 22 m. The lake is monomictic (Hutchinson 1957); turn- over begins in early winter and mixing ex- tends to spring. January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT Fox Valley Needles Cormorant Rock Helk Kitchen Anderson Bay True North PopcortT" Fig. 2. Bathymetric map of Pyramid Lake, Nevada; depth contours are in meters. Physical changes, including out-of-basin and inbasin water diversions, channelization, and destruction of riparian habitat, have ad- versely affected the ecology of the Truckee River-Pyramid Lake ecosystem. Historically the Lahontan cutthroat trout moved out of Pyramid and Winnemucca lakes, Nevada, and spawned in the entire Truckee River and its tributaries, a total length of 525 km. They also moved into Lake Tahoe, Nevada-Califor- nia, and spawned in its tributary streams. Derby Dam, completed in 1905, 62 km Great Basin Naturalist Vol. 43, No. 1 above Pyramid Lake, effected a transbasin diversion of much of the lower Truckee Riv- er flow. This obstacle reduced river spawning to the area below the dam. The dam not only had the direct effect of reducing flows in the lower river, but indirectly caused the buildup of the delta at the mouth. The numbers of trout diminished steadily imtil 1930, which was the last successful spawning year for the original population of Pyramid Lake Lahon- tan cutthroat trout (Sumner 1939). The U.S. Bureau of Fisheries (now U.S. Fish and Wild- life Service) stocked limited numbers in the lake in 1931 and 1932. None was seen in Pyramid Lake after 1943 and very few after 1938. However, before that time millions of eggs had been taken from the Pyramid Lake cutthroat and stocked elsewhere (Townley 1980). In 1976 the Marble Bluff complex, con- sisting of a dam and impoundment, a build- ing, and a 5.6 km fishway ending at the lake, was completed. This facility was built so that spawning fish could migrate upriver when there was not enough water in the river del- ta. The fishway, operating at 0.85 to 1.27 mVsec, allows fish to move upstream via four step-up ladders. At the upper end of the fish- way the fish may be shunted on upstream or into the building. Importance The Pyramid Lake Lahontan cutthroat trout is potentially of substantial economic and social importance to the Pyramid Lake Paiute Indian Tribe. Its adaptation to the highly saline waters of the lake make it a unique natural history entity. In 1977 an estimated 27,241 people spent 276,532 hours fishing for Lahontan cutthroat trout in Pyramid Lake. They landed 43,841 fish, of which 19,930 or 46 percent were le- gal size (381 mm). This is at the rate of 0.16 fish per hour landed and 0.07 kept. It has been estimated that the historic an- nual production of Lahontan cutthroat trout was at least 454,000 kg (Behnke 1974). The Pyramid Lake trout was the mainstay in the diet of the Pyramid Lake Paiutes and many other Indian tribes. They were also shipped to mining camps and other markets as far away as San Francisco. Range and Distrirution An ancestoral cutthroat trout probably in- vaded ancient Lake Lahontan from the Co- lumbia River Basin and developed into what is now known as the Lahontan cutthroat trout (Behnke and Zarn 1976). When the great lake desiccated, two populations of La- hontans evolved, one best adapted to lakes and the other to streams. The major lake populations of Lahontan cutthroat trout were then in Pyramid Lake, Walker Lake, Donner Lake, Independence Lake, and Lake Tahoe (Miller 1951). The trout in some of these lakes, which held an abundance of forage fish, became predatory at an early age, grew fast and large, and were moderately long lived. Currently, the largest population of lake- dwelling Lahontan cutthroat trout is in Pyra- mid Lake. Walker Lake supports a small population that has no opportunity to repro- duce. Summit Lake and Independence Lake may contain the most nearly pure strain of the Lahontan, but both lakes are small and have few forage fish (Behnke and Zarn 1976). A number of western lakes support reproduc- ing populations of Lahontan cutthroat trout. Recently a small stream-dwelling population of Lahontan cutthroat trout, believed to have been transplanted from Pyramid Lake before 1930, was discovered near Pilot's Peak, north of Wendover, Utah-Nevada (Hickman and Duff 1978, Hickman and Behnke 1979). There are a number of stream-dwelling La- hontan cutthroat trout populations in the Great Basin. In 1950, the Nevada Fish and Game De- partment initiated a small-scale stocking pro- gram of Lahontan cutthroat and other trout in Pyramid Lake (Trelease 1969). The pro- gram has now grown to 2.2 million 75-300 mm fish per year, supplied by two hatcheries of the Pyramid Lake Indian Tribal Enter- prises (PLITE) and the Lahontan National Fish Hatchery at Gardnerville, Nevada. Morphology and Genetics Despite the diverse evolutionary histories of western trout (genus Saltno), some species are related closely enough to interbreed free- ly and produce fertile hybrids. It is this po- tential presence of all degrees of hybrids January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT within a habitat that complicates identi- fication and evaluation of pure stock (Behnke and Zam 1976). The original stock of Lahon- tan cutthroat trout was apparently resistant to hybridization due to its long isolation in the Lahontan basin. The present subspecies does not share this characteristic. The isola- tion also encouraged a high degree of adapt- ability for lake habitat. The following are typical meristics of the Lahontan cutthroat trout (Behnke and Zam 1976): Scale counts lateral series two rows above lateral line 150-180 above lateral line (origin of dorsal fin to lateral line) 33-43 Vertebrae 61-63 Gillrakers 21-28 Pyloric caecae 40-75 Basibranchial teeth Numerous and well developed The number of pyloric caecae is higher in the Lahontan cutthroat than in other subspecies of cutthroat. The large, round, rather dull reddish spots that appear on the head as well as on the caudal peduncle and occasionally ventrally are the best field characteristic. The following data were collected as part of a study contracted between W. F. Sigler & Associates Inc. and the United States. This study was to provide an ecological evaluation of Pyramid Lake and its fishery resources and habitat. Procedures Fish life history data were taken by month- ly nettings from November 1975 through De- cember 1977. Fish were sampled by bottom set variable-mesh gill nets, vertical set gill nets, beach seines, fyke nets, and trawls. Fish were measured to the nearest millimeter in fork (FL), standard (SL), and total (TL) lengths and weighed to the nearest gram (Sig- ler and Kennedy 1978). Scale samples were collected from the left side in the region above the lateral line and midway between the posterior edge of the operculum and the origin of the dorsal fin. Five scales per fish were selected and impres- sions made of them on plastic slides with the use of a roller press (Smith 1963). The length-weight relationship is expressed by the formula W = aL'' (Sigler 1951), where W = weight in grams, L = fork length in cm, and a and b are constants. A log transfor- mation of W produces a linear equation. The constants a and b are calculated by the meth- od of least squares. Validity of the scale method was deter- mined by criteria suggested by Van Oosten (1923, 1929, 1944) and Hile (1941). To avoid possible bias, scales were first read without knowledge of the size of the fish. The scales were read at least three times. Further checks for accuracy of age assignment includ- ed comparisons with known age and tagged fish, Peterson's method, and use of year marks on other bony parts. All scales were examined with an Eberbach microprojector at a magnification of SOX. The body-scale relationship was calculated according to Tesch (1971). The condition fac- tor K = WxlOVL^ was calculated according to Carlander (1969), where W = weight in grams and L = fork length in mm. Calcu- lations were accomplished using an age- growth computer program (Nelson 1976). Creel census information was collected from January 1977 to April 1978. Four week- days and six weekend days were randomly se- lected each month for censusing, with holi- days treated as weekend days. On each selected day a check station was manned on the principal highway leading to Pyramid Lake and three aerial counts were made. Check stations were in operation from noon until dark, where all pertinent information was collected from fishermen. Aerial surveys were conducted by dividing a day into three equal time segments and an aerial count was made at the midpoint of each segment (Fig. 3). Inclement weather caused cancellation of 5 percent of the flights (Kennedy 1978). Shore fishermen and boats were counted on each flight, with the number of boat fish- ermen obtained by multiplying the number of boats by the average number of fishermen per boat on the day of the count (Johnson and Wroblewski 1962). When less than 10 boats were checked, the yearly mean number Great Basin Naturalist Vol. 43, No. 1 Needles Hells Kitchen Anderson Bay ''aREAN- Dago Bay EIGHT s^ Depth contours (m] To Reno To Nixon Truckee River Fig. 3. Lake areas used for creel census data collec- tion and location of creel check stations. of fishermen per boat was used. Rate of suc- cess, effort, and harvest were calculated by computer program (David Wheaton, pers. comm. 1977). All calculations were expanded to a 30-day month. Mean lengths and weights of fish caught were also calculated. Food habits were determined by examining the stomachs of five fish per size group from each net catch. Food habit analyses were conducted by percent of frequency of occur- rence and percent of total volume. Fecundities were determined by actual egg counts (Kucera and Kennedy 1977). Criteria described by Nikolsky (1963) were used for determining stage of maturity; only ripe fe- males and fresh ovaries were used for fecun- dity studies. Linear and logio (Y+1) regres- sions between fecundity, fork length, weight, age, ovum diameter, ovary weight, and net weight (body weight minus ovary weight) were used to examine the interrelationships between these variables. Age and Growth Appearance of Scales and Formation of Annuli Fork length at the time of scale formation for hatchery-reared Lahontan cutthroat plan- ted in Pyramid Lake is 25.8 mm. Calhoun (1942) lists 40 mm (FL) as the size of Lahon- tan cutthroat from Blue Lake, California, at time of scale formation. Yellowstone cut- throat develop scales when they are between 40.3 mm and 42.8 mm (FL) (Brown and Bail- ey 1952). Laakso and Cope (1956) report 39.3 mm (FL) as the size of cutthroat trout at the time of scale formation. Cutthroat trout sam- pled in Montana had formed scales at 41.2 to 63.2 mm (FL) (Brown and Bailey 1952). Ir- ving (1953) reports 23.9 mm (FL) for cut- throat trout in Henry's Lake, Idaho. Rob- ertson (1947) found considerable variation in the size of cutthroat trout at the time of scale formation. Nearly all scales examined from Pyramid Lake fish showed early growth patterns char- acteristic of hatchery rearing. Scales also showed crowded circuli beginning in late September 1975 and 1976 and in early No- vember 1977. Nearly all scales aged showed winter bands of thin and closely spaced cir- culi. Summer growth bands appear as thick and widely spaced circuli. The beginning of growth, the first appearance of summer banding, is assumed to correspond with for- mation of the annulus. The period of annulus formation extended from about February through May, peaking in late April 1976 and late March 1977. All annuli were readily visible. Check marks that appeared throughout all fields were prominent during the first year's growth for the majority of fish examined. These stress conditions that resulted in growth interruptions existed during the first (0 age) year for fish from the National Fish Hatchery (Lahontan National Fish Hatchery personnel, pers. comm. 1976). This situation presumably does not occur every year. Seasonal Growth Our analyses of age and growth for Lahon- tan cutthroat trout from Pyramid Lake is based on scale samples from 676 specimens taken almost exclusively with nets from No- vember 1975 through November 1977. The general shapes of the 1976 and 1977 growth curves (Fig. 4) were the same, but during 1977 growth was more rapid and extended over a longer period of time than for com- parably aged fish in 1976. Increments of January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT growth declined with increasing age of fish in 1976, but increased in 1977. Growth in- creased sharply in spring, slowed in late sum- mer, and ceased during fall and winter. Annual Growth Annual growth (back-calculated lengths) values were derived from the body length- scale radius relationship FL = A -I- B(SR); FL = fork length in mm and SR = anterior scale radius (Table 1). Body-scale regression equations, based on data collected over the entire study, were used to calculate the lengths. The results for 676 fish are: for fe- males FL = 155.881 + 3.5364 (SR), for males FL = 79.176 + 4.2599 (SR), for in- determinates FL = 112.872 + 4.0834 (SR), and for combined FL = 132.952 + 3.8079 (SR). Young-of-the-year Lahontan cutthroat trout sampled from Lahontan National Fish Hatchery averaged 152 mm in length at age eight months. By the end of Year I, hatchery trout are approximately 203 mm FL (Lahon- tan National Fish Hatchery personnel, pers. comm. 1976). These data demonstrate that the back-calculated lengths for age I Pyramid Lake trout are accurate. Growth in length is nearly isometric from the end of the first through the seventh years of life. Variation by sex is evident in the growth rates of certain age groups. Annual increments of growth in length for males are greater than for females from age II on. Ac- cording to Irving (1953), male trout from Henry's Lake, Idaho, grow faster than fe- males, and Bulkley (1961) reports male trout outlive females. Pyramid Lake Lahontan cut- throat trout appear to follow these patterns. Others have reported no difference between the sexes in growth rates (Drummond 1966, Snyder and Tanner 1960). The oldest male and female aged from Pyramid Lake were in their seventh and sixth year, respectively. This longevity is some- what less than historical data. Sumner (1939) found the oldest age groups of trout in Pyra- mid Lake were the seven- to nine-year-olds. Studies in smaller high altitude lakes. Upper Blue Lake (Calhoun 1944), and Topaz Lake (Johnson 1958), indicate that few trout live past their sixth year (Table 2). 500 ■ E J. 300 i£ 200 100 ■ AgeV -il91_ Days = 273. Days 1976 1977 AJ AOD FA JAOD Month Fig. 4. Seasonal growth curves, 1976 and 1977, for Lahontan cutthroat trout age groups I through V. Fish were collected from Pyramid Lake, Nevada, from No- vember 1975 through November 1977. The graph ab- scissa is divided into bimonthly intervals beginning with April. Lahontan cutthroat trout greater than 787 mm were not sampled with our nets and thus do not appear in our age and growth studies; however, larger ones were taken by anglers. In the 1977 creel sample, which was about three times that of the net sample, 22.5 per- cent of the 1916 trout exceeded 600 mm in length; the longest one was 990 mm FL. The average size was 505 mm (S.D. = 107.0). Ei- ther these large fish grew faster than the av- erage in our studies, or they were older than the maximum ages of our net-caught fish. In April 1976, an Indian angler, Ralston Fill- more, captured a 12.7 kg Lahontan cutthroat trout, the largest recorded since December 1925, when another Indian, John Skim- merhorn, caught an 18.6 kg trout, the world record cutthroat (Wheeler 1974). There are numerous unconfirmed reports of ones larger than this being marketed around the turn of the century. LeCren (1951) states the length-weight relationship equation, in addition to provid- ing a method of converting length to weight, also indicates taxonomic differences and events in the life history. The value of the constant "B" will equal 3.0 where growth is symmetrical or isometrical (Ricker 1971). Values less than 3.0 demonstrate linear growth is taking place faster than growth in Great Basin Naturalist Vol. 43, No. 1 weight. Values greater than 3.0 demonstrate the reverse; both are allometric growth. The length-weight equations calculated for Lahontan cutthroat trout show growth slightly exceeds the cubic relationship; this represents allometric growth. We combined all years, drawing on data from 561 trout. Sizes ranged from 189 mm (36 g) to 787 mm (6163 g) (Table 3, Fig. 5). Tesch (1971) notes allometric growth in some salmonid stocks. The length-weight curve of the Lahontan cutthroat trout shows the importance of for- age fish. In Pyramid Lake cutthroat trout weight gains tend to exceed the increases in length when the diet shifts from invertebrates to fish. They are in their third year and sim- ilar to cutthroat trout from Independence Lake, California (Lea 1963). Lea states the rate of growth for the Independence Lake cutthroat trout population is only slightly Table 1. Summary of the mean calculated fork lengths and increments of growth for Lahontan cutthroat trout collected from Pyramid Lake, Nevada, from November 1975 through November 1976. Age group Number of fish Calculated fork lengths (mm) at end of each year of life 2 3 4 5 6 (Female) I II III IV V VI 42 68 80 25 23 15 238 236 233 236 236 233 306 305 304 303 306 374 376 374 374 442 442 447 500 508 580 Grand average Increments of growth Number of fish 235 235 253 305 70 211 374 70 143 443 68 63 503 59 38 580 72 15 (Male) I II III IV V VI VII 4 178 14 168 255 26 169 253 341 25 171 254 344 439 24 168 254 339 437 516 16 167 250 341 439 518 4 165 250 345 435 521 588 575 664 Grand average Increments of growth Number of fish 169 169 113 253 84 109 342 89 95 438 96 69 517 79 44 585 66 20 664 88 4 (Indeterminate) I 26 217 II 51 206 286 III 110 203 285 359 IV 71 202 281 353 416 V 40 200 274 343 407 475 VI 10 197 264 334 407 471 VII 2 199 274 335 384 460 (Combined) I 72 225 II 133 219 294 III 216 216 293 365 IV 121 217 290 362 429 V 87 215 287 358 429 494 VI 41 213 286 360 440 506 VII 6 212 286 362 433 508 536 538 574 565 578 Grand average 204 282 353 412 474 537 578 Increments of growth 204 79 72 64 67 68 41 Number of fish 310 284 233 123 52 12 2 629 Grand average Increments of growth Number of fish 217 217 676 291 75 605 362 72 472 431 70 256 499 66 135 573 66 48 629 64 6 January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT curvilinear until age III, at which time the relative weight increase accelerates greatly. Lea also reports that, for Independence Lake cutthroat trout less than 225 mm, forage fish are of minor significance, but for those over 300 mm, fish become the major forage item. Hazzard and Madsen (1933) report cutthroat trout from Jackson Lake, Wyoming, also show a definite change in diet from Crustacea to fish at a length of approximately 300 mm. The condition factor (K = W X 10 VL^) is used as an index of well-being or relative ro- bustness. The average K of 561 Pyramid Lake Lahontan cutthroat trout, sexes and age groups combined, was 1.00. A slight sexual dimorphism is noted for condition factor, with males having a slightly higher K value than females (Table 4). This is in agreement with results from other studies. Fleener (1952) and Madsen (1940) also report higher K values for male over female cutthroat trout. However, the extent of the sexual di- morphism may vary with fish size and season; i.e., gravid females weigh significantly more just before spawning season than after. Thus the K of females is more variable than males on a seasonal basis. A direct relationship between size and condition factor of Pyramid Lake Lahontan cutthroat trout is evident (Table 5). Lea (1963) reports a trend of increasing condition factor with increasing length for Independ- ence Lake Lahontan cutthroat trout. Fleener (1952), however, states condition factor de- creases with length for cutthroat trout from Beaver Creek, a small tributary of the Logan River, Utah. Irving (1953) says size, age, and sex are not related to condition factor for Henry's Lake cutthroat trout. It seems logical that condition factor would be directly re- lated to fish size in lake environments where large fish have a predatory advantage. This situation occurs in Pyramid Lake since, at the critical size of about 300 mm, Lahontan cut- throat trout are able to utilize the huge for- age base of tui chubs. Table 2. Growth of cutthroat trout from 14 Western lakes. Calculated mean total length (mm) and increments at each annulus Location Number I II III IV V VI VII Pyramid Lake, NV^ 676 217 291 362 431 499 573 629 (217) (75) (72) (70) (66) (66) (64) Lower No Name Lake, WY 64 102 145 190 221 231 (Robertson 1947) (102) (43) (45) (31) (10) Priest Lake, ID 90 81 135 211 287 348 371 (Bjomn 1957) (81) (54) (76) (76) (61) (23) Upper Priest Lake, ID 92 94 142 216 292 338 391 (Bjomn 1957) (94) (48) (74) (76) (46) (53) White Rock and Ted's Lake, UT 22 130 185 201 221 (Sigler and Low 1950) (130) (55) (16) (20) Granby Reservoir, CO 109 196 251 290 (Finnell 1966) (109) (87) (55) (39) Yellowstone Lake, WY 5057 46 130 224 312 394 442 486 (Bulkley 1961) (46) (84) (94) (88) (82) (48) (44) Montana Lake, MT 2158 76 163 241 307 384 (Peters 1964) (76) (87) (78) (66) (77) Island Lake, UT 61 157 211 249 300 343 (Sigler and Low 1950) (157) (54) (38) (54) (43) Thompson Lake, MT 41 130 198 262 318 (Echo 1955) (130) (68) (64) (56) Upper No Name Lake, WY 75 112 178 274 381 421 478 (Robertson 1947) (112) (66) (96) (107) (40) (57) Blue Lake, CA 419 66 180 307 378 361 (Calhoun 1942) (66) (114) (127) (71) Heenan Lake, CA 117 97 216 330 445 (Calhoun 1942) (97) (119) (114) (115) Henry's Lake, ID 356 170 325 437 503 551 594 (Irving 1953) (170) (155) (112) (66) (48) "Fork lengths converted to total length by factors of 1.07 (189 mm - 490 mm), 1.05 (500 mm - 590 mm), and 1.03 (> 590 mm). 10 Great Basin Naturalist Vol. 43, No. 1 6163 4938 + 3712 + ? 2487 Graph symbol No. points represented W= .0027L33271* n = 561 • 23456 789 A BC D 1 2 3 4 5 6 7 8 9 10 11 12 13 Age VII . Age VI Age V Age IV 1261 + Age / /..•■ •4 3 » 2 J 2 . ». I X -Age Age I + 2*26S4*3* 189* 2 . .3«'5 I /3. . > ••2*32 •42** • ••7847 •* il 2 I * 2^23489664222 • , I •37AC6943 2 • I 4 5638 37087^ 233 ••• 2 44494S7S732I^^2* • 26528848A94522** • 309" 428 548 677 787 Fork length (mm) Fig. 5. Length-weight relationship of combined data for Lahontan cutthroat trout from Pyramid Lake, Nevada, November 1975 through November 1977, with mean length-weight values by age groups. FL in cm. Conversion Factors Factors for converting TL to FL and SL to FL for cutthroat trout from Pyramid Lake, Nevada are: TL = 1.07 FL (189 - 490 mm) TL = 1.05 FL (500 - 590 mm) TL = 1.03FL(>.590mm) SL = .888 FL (189 - 300 mm) SL = .893 FL (301 - 500 mm) SL = .895FL(>501 mm) FL = .935 TL (202 - 524 mm) FL = .953 TL (525 - 620 mm) FL = .970TL(>608mm) FL = 1.13 SL (168 - 266 mm) FL = 1.12 SL (268 -447 mm) FL = 1.12SL(>447mm) The ratios vary with size, necessitating more than one set of conversion factors. Con- version factors for cutthroat trout have been reviewed by Cope (1953). Hatchery-reared Lahontan Cutthroat Trout, Length-weight Relationships and Condition Factors In 1976, 612 fingerlings were taken from the Lahontan National Fish Hatchery, Gard- nerville, Nevada, to determine length-weight relationships. The fingerlings represent wild Summit Lake stock ranging in fojk length from 57.9 mm to 125 mm (Table 6) and La- hontan fifth-generation domestic stock, origi- nally from Summit Lake, ranging in fork length from 40 mm to 250 mm (Table 7). The length-weight relationships are calculated as: Summit Lake brood W = .00001L2-8749 Lahontan brood W = .000007L30588 The K-factors for domestic stock range from 1.01 to 1.21, the exponent indicating slightly faster growth in weight than length. January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT 11 The K-factors for wild stock show sUghtly de- creasing trends; the exponent is less than 3.0, indicating these fish are getting slimmer as they grow in length. The Lahontan brood attain greater weight per length than do Summit Lake brood. The weight differences can be attributed to brood spawned from wild stock being more active (wild) and domestic stock being more passive (Lahontan National Fish Hatchery Manager, Charles R. Messier, pers. comm. 1976). Mortality and Morbidity Various factors may cause mortality of Pyramid Lake's trout population, including angling, death during stocking of hatchery recruits, predation on juveniles, spawning-re- lated deaths of adults, and disease. When other decimating factors are inoperative or eliminated, senility must ultimately cause death. Among 676 Lahontan cutthroat trout sampled from Pyramid Lake during this study, no females and only four males reached age VII. Sumner's (1939) data in- dicates very few of the original population of Lahontan cutthroat trout lived beyond eight years, although a few may have lived to be 11. Chemical constituents of the aquatic habi- tat are rarely neutral in their effects on the biota. Toxic substances often first express themselves as growth suppressants, reproduc- tive inhibitors, increased vulnerability to dis- ease, or destroyers of the most sensitive link in the food chain. Increased levels of TDS could be detrimental to the Pyramid Lake fishery. Walker Lake's now extinct Sacra- mento perch population reached its limit of "alkalinity" tolerance when it could no long- er reproduce in the early 1950s. At that time, the total alkalinity was approximately 2500 mg/1 as HCO3 (Cooper 1978). In 1952, the TDS of Walker Lake was 6790 mg/1 (Koch et al. 1979). Rainbow trout are stressed by and grow poorly in Pyramid Lake water; they do not survive in more concentrated al- kaline waters such as Walker Lake (Knoll et al. 1979) and Omak Lake (Paul A. Kucera, pers. comm. 1982). Taylor (1972) notes that carbonate and bicarbonate salts are more tox- ic to Lahontan cutthroat trout at elevated TDS levels than sodium chloride alone. Ele- vated temperatures may have a synergistic effect on salinity toxicity and vice versa (Vigg and Koch 1980). Mortalities range widely among fish cap- tured and released. Hooking mortality of lure-caught cutthroat trout in Yellowstone Lake was relatively low (<6.5 percent); however, the combination of natural baits and high water temperature resulted in sig- nificantly higher mortality (Mamell 1969, Marnell and Hunsaker 1970). In Pyramid Lake, where fish <483 mm TL are illegal (as of 1 July 1982), all types of artificial lures are used. The losses from hook and release in Pyramid Lake have not been established. Le- gal sport fishing removed < 20,000 fish in 1977, not a seriously decimating factor for the Pyramid Lake population. Infectious disease is a potential threat to wild and cultured fish alike. Rational man- Table 3. Length-weight relationships (linear and curvilinear) for Lahontan cutthroat trout from Pyramid Lake, Nevada, from November 1975 through November 1977.* Class Log-log transformation (linear) Exponential (curvilinear) (Female) (Male) (Indeterminate) (Combined) logioW = -2.6218 + 3.3754 logioL (i^ = .95, n = 224) F = 602.4 logioW = -2.8052 -I- 3.4778 logioL (1^ = .94, n = 73) F = 1089.9 logioW = -2.3690 -I- 3.2023 logioL (r2 = .92, n = 262) F = 3017.2 logioW = -2.5531 + 3.3271 logjoL (i^ = .94, n = 561) F = 9293.3 W = .0023L3-3754 W = .0016L3'«778 W = .0042L3-2023 W = .0027L3-3271 ^Equations were calculated using fork length in centimeters. 12 Great Basin Naturalist Vol. 43, No. 1 agement and utilization of any fisheries must incorporate a realistic understanding of the serious pathogens extant in the system. A fish population undergoing environmental deg- radation is subjected to environmental factors that may predispose the population to dis- ease. Various interstate and international regulations have been formulated to restrict movement of serious fish pathogens. PLITE has a program to determine the presence of pathogens in Pyramid Lake trout. Particular attention is given to those pathogens included in restrictive lists. A series of inspections begun in 1976 followed proper procedure as specified by the Fish Health Section of the American Fisheries So- ciety and The Colorado River Wildlife Coun- cil-Fish Disease Policy (American Fisheries Society: Fish Health Section 1975), Con- sultation and confirmation of procedure were reviewed by Ron Goede, fish pathologist of the Utah Division of Wildlife Resources, and by Dennis E. Anderson, U.S. Fish and Wild- life Service, Fort Morgan, Colorado. The inspections in 1977 included the fol- lowing diseases and/or the pathogens in- ducing the diseases: Viral: IPN (Infectious pancreatic necrosis) (1976 & 1977) IHN (Infectious hematopoietic necrosis) VHS (Viral hemorrhagic septicemia) Bacterial: Bacterial kidney disease {Renebacteriiirn salmoninarum) Enteric redmouth {Yersinia ruckerii) Funmculosis {Aeromonas sahnonicida) Parasitic: Whirling disease {Myxosoma cerebralis) Blood fluke {Sanguinicola sp.) Pyramid Lake and lower Truckee River fish populations were sampled by hook and line, gill net, and electrofishing. Fish were also collected from the Marble Bluff fishway and the Dunn Hatchery, Sutcliffe, Nevada. The inspection in 1976 was conducted by Biometrics Inc., Tacoma, Washington. In 1977, the Fish Disease Control Center, U.S. Fish and Wildlife Service, Fort Morgan, Col- orado, conducted inspections. Attribute sampling for IPN in 1976 pre- sumed a 5 percent carrier incidence and achieved 95 percent confidence limits. No evidence of any pathogen was detected in 1976 (Ferjancic 1976). Sampling in 1977 pre- sumed a carrier incidence of 2 percent and achieved 95 percent confidence. Inspections conducted in 1977 (Ruger 1977) detected no evidence of IPN, IHN, VHS, bacterial kidney disease, enteric redmouth, furunculosis, whirling disease, or blood fluke, except 7 of 235 adults sampled at the Marble Bluff fish- way were positive for furunculosis. Sample sizes were sufficiently large to ex- tend confidence beyond original required sampling presumption. Regulation and pro- tocol require assumption of a 2 percent car- rier incidence for 95 percent confidence in wild populations. Sample sizes in this study are sufficient at the 1 percent carrier in- cidence to permit 95 percent confidence in detecting IPN and at the 2 percent carrier in- cidence to permit 95 percent confidence in detecting all other listed diseases. Food and Feeding Habits Lahontan cutthroat trout in Pyramid Lake are largely piscivorous after they reach a size of approximately 300 mm. They then feed al- most exclusively on tui chubs, but they may feed opportunistically on other fish and they feed to some extent on aquatic insects. Small trout feed on zooplankton and benthic in- vertebrates. From January through Decem- ber 1976, 192 Lahontan cutthroat trout were examined for food habits; 35 had not recently fed. The highest percentage of the 35 non- feeders occurred during the winter and early spring months when v/ater temperatures and. Table 4. Coefficient of condition for Lahontan cut- Table 5. Coefficient of condition based on increasing throat trout from Pyramid Lake, Nevada, fork length in mm, November 1975 through November 1977. fork length (mm) for Lahontan cutthroat trout from Pyramid Lake, Nevada, November 1975 through No- vember 1977. Male 74 1.08 .634-1.416 Female 225 1.03 .857-1.212 Sexes combined* 562 1.00 .634-1.416 N Mean Range Fork length (mm) 150-350 351-550 551-700 Number of fish 200 273 89 (K) .831 .994 1.160 ^Includes fish in which sex was undetermined. January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT 13 therefore, trout metabolism and feeding ac- tivity were low. The piscivorous nature of Lahontan cut- throat trout was predictable. Fish, the most frequent food item, was eaten by 62.4 per- cent of the trout (Table 8). Fish also account- ed for the largest volume of food (84.5 per- cent). Snyder (1917) found adults in lakes feed largely on minnows, with one fish from Pyramid Lake described as containing three large minnows. In Johnson's (1958) food anal- yses of 20 Pyramid Lake cutthroat trout, fish were dominant. Insects, zooplankton, and amphipods appear in descending order of im- portance. Invertebrates rather than fish are the major source of food for Lahontan cut- throat trout in two Sierra lakes, presumably because the trout occur in different areas of the lake than other species of fish (Calhoun 1942). A diet succession from invertebrates to fish is apparent for Lahontan cutthroat trout as they increase in size (Table 9). Invertebrates make up 51.2 percent and fish 38.3 percent of the volume of food eaten by trout 198-300 mm FL. The volume of invertebrates eaten decreases with increasing trout size. Larger trout, 300-399 mm, consume 32.8 percent in- vertebrates and 60.8 percent fish. This is also true for Utah cutthroat trout (Sigler 1962, Sigler and Miller 1963). Chironomids, the second most important food item, are consumed by 41.4 percent of the trout, but account for only 4.0 percent of the volume. Chironomid pupae are eaten nearly twice as frequently as larvae. This is also true for Lahontan cutthroat in Blue Lake, California (Calhoun 1944) and sea- sonally in Omak Lake, Washington (Paul A. Kucera, unpubl. data 1981). Platts (1959b) re- ports chironomidae pupae are the most im- portant forage item for cutthroat trout in Strawberry Reservoir, Utah. The remaining food items in the cutthroat diet were of relatively minor importance compared to consumption of fish and chi- ronomids. Some of the items, however, can be of significant value seasonally or during certain life stages, such as zooplankton and smaller invertebrates for young-of-the-year trout. Other food items consumed in order of percent of frequency of occurrence were: terrestrial insects (10.2); amphipods, both Hyallela and Gammarus (9); algae (7.6); zoo- plankton (7.6); bottom substrate (4.5); He- mipterans (3.2); odonates (1.9); vascular plants (1.9); coleopterans (.6); and hydracari- nads (.6). Reproduction The Lahontan cutthroat trout spawning migration into the Truckee River and Marble Bluff fishway peaked in April and May of 1976 and 1977 at river water temperatures varying from 8 to 16 C (Fig. 6). Although only 563 fish were captured in 1976, the run was reported to be the largest in recent years Table 6. Expected Lahontan cutthroat trout measurements (FL) based on measurements of 200 Lahontan cut- throat trout, Summit Lake brood. Lahontan National Fish Hatchery, Gardnerville, Nevada, 1976. Length (L) Length (L) Weight (W) Grams per "K" Fish per in inches in millimeters in grams centimeter factor kilogram 2.16 55 1.66 .30 .99 602.94 2.36 60 2.16 .36 .99 463.37 2.55 65 2.64 .40 .96 379.11 2.75 70 3.60 .51 1.04 278.02 2.95 75 4.56 .60 1.08 219.49 3.14 80 4.88 .61 .95 205.10 3.34 85 6.21 .73 1.01 161.16 3.54 90 7.17 .79 .98 139.57 3.74 95 8.50 .89 .99 117.75 3.93 100 10.41 1.04 1.04 96.14 4.13 105 11.74 1.11 1.01 80.84 4.33 110 13.34 1.21 1.00 75.02 4.52 115 14.99 1.30 .98 66.76 4.72 120 16.91 1.40 .97 59.17 4.92 125 18.24 1.45 .93 54.87 5.11 130 20.16 1.55 .91 49.63 14 Great Basin Naturalist Vol. 43, No. 1 (U.S. Fish and Wildlife Service pers. comm. 1976). In comparison with the number of fish in the lake large enough to reproduce, this number is amazingly small. The 1977 creel census (Kennedy 1978) produced an esti- mated sport fishing catch of < 20,000 Lahon- tan cutthroat trout >380 mm TL. This size is a reasonable approximation of the average length at reproductive maturity. It is obvious that the population of mature fish must be considerably greater than 563, the number that were captured. Netting did not indicate an unusual concentration of cutthroat trout in the lake near the Truckee River mouth at this time. However, large numbers of trout congregated around the Sutcliffe area where hatchery reared cutthroat are planted. As Ball (1955) postulates, since these planted fish were not imprinted on an inflowing stream, they may be milling about the area where they were originally planted. Limited data on the maturation of female Lahontan cutthroat trout in Pyramid Lake suggest that consecutive-year spawning does Table 7. Expected Lahontan cutthroat trout measurements (FL) based on measurements of 412 Lahontan cut- throat trout, Lahontan domestic brood. Lahontan National Fish Hatchery, Gardnerville, Nevada, 1976. Length (L) Length (L) Weight (W) Grams per "K" Fish per in inches in milhmeters in grams centimeter factor kilogram 1.57 40 .65 .16 1.01 1539.84 1.77 45 .93 .20 1.02 1076.23 1.96 50 1.29 .25 1.03 775.87 2.16 55 1.73 .31 1.03 578.54 2.36 60 2.27 .37 1.05 440.92 2.55 65 2.91 .44 1.05 343.94 2.75 70 3.67 .52 1.06 272.71 2.95 75 4.54 .60 1.07 220.46 3.14 80 5.55 .69 1.08 178.57 3.34 85 6.70 .78 1.09 149.39 3.54 90 8.00 .88 1.09 125.11 3.74 95 9.46 .99 1.10 105.80 3.93 100 11.09 1.10 1.10 90.24 4.13 105 12.90 1.22 1.11 77.58 4.33 110 14.90 1.35 Ml 67.15 4.52 115 17.11 1.48 1.12 58.99 4.72 120 19.52 1.62 L12 51.26 4.92 125 22.15 1.77 1.13 45.17 5.11 130 25.02 1.92 1.13 39.99 5.31 135 28.12 2.08 1.14 35.58 5.51 140 31.48 2.24 1.14 31.79 5.70 145 35.10 2.42 1.15 28.51 5.90 150 38.99 2.59 1.15 25.66 6.10 155 43.16 2.78 1.15 23.17 6.29 160 47.62 2.97 1.16 21.01 6.49 165 52.39 3.17 L17 19.09 6.69 170 57.47 3.38 Lie 17.39 6.88 175 62.88 3.59 L17 15.92 7.08 180 68.61 3.81 1.17 14.57 7.28 185 74.70 4.03 1.17 13.38 7.48 190 81.13 4.27 1.18 12.32 7.67 195 87.94 4.50 1.18 11.38 7.87 200 95.12 4.75 1.18 10.52 8.07 205 102.69 5.00 1.19 9.74 8.26 210 110.65 5.26 1.19 9.03 8.46 215 119.03 5.53 1.19 8.40 8.66 220 127.82 5.81 1.20 7.83 8.85 225 137.04 6.09 1.20 7.30 9.05 230 146.70 6.37 1.20 6.81 9.25 235 156.82 6.67 1.20 6.37 9.44 240 167.39 6.97 1.21 5.97 9.64 245 178.44 7.28 1.21 5.60 9.84 250 189.98 7.59 1.21 5.25 January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT 15 not occur. Judging by ovary development, several females of age groups IV and V col- lected prior to the spawning season were not going to reproduce the current year but would the following spring. Similar situations have been reported elsewhere. Some 1-15 percent of the cutthroat in Arnica Creek, Yellowstone National Park, spawn each year after reaching maturity, 10-26 percent are alternate-year spawners, and 46 percent skip two years (Ball and Cope 1961). Seven per- cent of the female Lahontan cutthroat trout in Blue Lake, California, spawned in con- secutive years, and 10.5 percent of the origi- nally marked females were in the run two years later (Calhoun 1942). Female cutthroat trout in Pyramid Lake mature at age III or IV, when they are 352 to 484 mm FL; males mature at ages II or III, when 299 to 445 mm FL. Lahontan cutthroat trout in Independence Lake, California, ma- ture at ages III or IV (Lea 1963). Lea attri- butes the presence of small numbers of ma- ture three-year-olds to a precocial element of the population. Rankel (1976) reports spawn- ing runs of Lahontan cutthroat trout from Summit Lake, Nevada, consist mainly of four-year-old fish. If an alternate-year spawn- ing pattern is typical, then most female La- hontan cutthroat trout in Pyramid Lake will spawn a maximum of twice in their lifetime. If they mature at age IV, they may live to spawn only once. Some of those that mature at age III may spawn again at age V. The sex ratio of Lahontan cutthroat trout in our net catches was 1 male:4.23 females (n = 455). This is not representative of the population in the lake. The ratio of males to females in the spawning runs was variable, i.e., 1.06:1 in 1976 and 1:2.35 in 1977. The ratios of Summit Lake Lahontan cutthroat trout spawning runs, from 1968 to 1975, var- ied from 1:1.3 to 1:2.2 and averaged 1:1.6 males to females (Rankel 1976). Angling is male-selective in Pyramid Lake. This is also true in Omak Lake (Paul A. Kucera, pers. comm. 1981). Female Lahontan cutthroat trout spawn after attaining an average gonadal somatic index (percent gonad weight to total body weight) of 11 percent. The progression in go- nadal somatic indices, observed from October through December 1977, indicates a steady increase in germinal growth through the Table 8. Food of 157 Lahontan cutthroat trout. Pyramid Lake, Nevada, 1976. Percentage of total Percentage Percentage volume frequency Volume of excluding of in total digested Food item Frequency occurrence Rank ml volume Rank matter Fish 98 62.42 1 1225.7 82.89 1 84.50 Chironomidae 22 14.01 3 11.6 0.78 6 0.80 larvae Chironomidae 43 27.39 2 46.8 3.16 3 3.23 pupae Odonata 3 1.91 9 2.7 0.18 9 0.19 Coleoptera 1 0.64 10 0.7 Trace* 13 Trace* Hemiptera 5 3.18 8 1.0 Trace* 12 Trace* Terrestrial 16 10.19 4 19.6 1.33 5 1.35 insects Amphipoda 14 8.92 5 9.2 0.62 8 0.63 Ostracoda 3 1.91 9 1.4 Trace* 11 0.10 Zooplankton 12 7.64 6 10.9 0.74 7 0.75 Algae 12 7.64 6 93.1 6.30 2 6.42 Vascular 3 1.91 9 2.0 0.14 10 0.14 plants Hydracarina 1 0.64 10 Trace* Trace* 14 Trace* Bottom substrate 7 4.46 7 25.8 1.74 4 1.78 Digested matter 18 11.46 28.2 1.91 — Total 1478.7 99.98 99.89 *Trace = less than 0.1 16 Great Basin Naturalist Vol. 43, No. 1 winter. The largest increase involves a shift from an average value of 5.9 percent in No- vember to 8.1 percent in December. The diameter of mature eggs of Pyramid Lake Lahontan cutthroat trout range from 2.76 to 5.09 mm and average 4.51 mm. Monthly progression in egg sizes from Octo- ber through December 1977 indicated a con- sistent, gradual increase in ovum size. The mean egg diameter in December was 4.11 mm, with some eggs as large as 4.60 mm. Some females may be able to spawn in Table 9. Percentage of total volume and frequency of occurrence of food items consumed by Lahontan cutthroat trout from Pyramid Lake, Nevada, in relation to size. Trout were captured from January through December 1976 with bottom-set gill nets. Food item Volume Percent of total volume Frequency by occurrence Frequency of occurrence by percent Size group 99-198 mm (n = = 1) 100.0 1 Fish 1.0 100.0 Fork length = 184 mm Weight = 54 g Size group 198-300 mm (n = 22) Fish Benthic invertebrates Terrestrial insects Zooplankton Digested matter Vascular plants 12.1 38.3 11 50.0 12.2 38.6 12 54.5 4.0 12.7 2 9.5 0.5 1.6 1 4.8 2.0 6.3 2 9.5 0.8 2.5 1 Mean fork length = 251 mm Range = 203-297 mm Mean weight = 136 g Range = 54-240 g Size group 300-399 mm (n = 63) Fish Benthic invertebrates Terrestrial insects Zooplankton Periphyton Vascular plants Bottom substrate Digested matter 67.1 60.8 42 66.7 27.6 25.0 25 39.7 7.3 6.6 7 11.1 1.3 1.2 2 3.2 2.6 2.4 2 3.2 1.0 0.9 1 1.6 Trace* Trace^ 1 1.6 3.4 3.1 3 4.8 Mean fork length = 356 mm Range = 305-398 mm Mean weight = 408 g Range = 218-730 g Size group 399-498 mm (n = 60) Fish Benthic invertebrates Terrestrial insects Zooplankton Periphyton Vascular plants Bottom substrate Digested matter 530.7 77.8 35 58.3 30.0 4.4 25 41.7 8.3 1.2 8 13.3 9.1 1.3 8 13.3 57.5 8.4 9 15.0 0.2 Trace^ 1 1.7 23.5 3.4 6 10.0 22.8 3.3 9 15.0 Mean fork length = 434 mm Range = 400-498 mm Mean weight = 789 g Range = 476-1496 g Size group 500 + mm (n = 11) Fish Benthic invertebrates Periphyton Bottom substrate 615.8 94.1 10 90.9 3.6 0.5 1 9.1 33.0 5.0 1 9.1 2.3 0.4 1 9.1 Mean fork length = 531 mm Range = 500-596 mm Mean weight = 1741 g Range = 1161-2753 g *Trace = less than 0.1 January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TroUT 17 J F 1977 Month and year Fig. 6. Mean monthly Pyramid Lake cutthroat trout catches (15 gill net sets per month) from November 1976 through December 1977, in comparison with the spawning runs at the Marble Bluff fish passage facility. December or January. For example, as early as January, 1981 and 1982, mature cutthroat were running up a small stream south of Sut- cliffe, Nevada, artificially produced by pumping about .057 m^/sec of lake water through rearing ponds and allowing it to run back into the lake. Egg size is positively cor- related with fish length (r = .48; P<0.05) and weight (r = .51; P<0.05), indicating that egg size increases with fish size. The number of eggs produced by a female Lahontan cutthroat trout is significantly re- lated to age, fork length, and weight (P<0.05). The above relationships have sig- nificant linear fits with and without logio transformation of data. Fork length provides the best predictor for fecundity, followed by weight and age. Increases in fecundity corre- spond to increases in length and weight. The log 10 equation for fork length and fecundity is logioF = 2.83 (logioFL) - 4.16, and for weight and fecundity is logioF = .81 (logioWT) + .92. These fish range in fecun- dity from 1241 to 7963 and average 3815 eggs per female. Lea (1963) reports fecun- dities of Lahontan cutthroat trout in Inde- pendence Lake, California, vary from 669 to 2080 eggs and average 1191 eggs per female. HUbitat and Ecology The most characteristic feature of the Pyramid Lake environment is the high level of salts; TDS concentration was about 5350 mg/1 during 1976-1977 (Sigler and Kennedy 1978). Although sodium chloride is the domi- nant salt (over 70 percent), alkalinity may be the most important constituent. The mean pH is 9.2. The historic increase in TDS levels was associated with the decline in lake level (Fig. 7). Since the baseload of salts is relative- ly constant, TDS varies inversely with the volume of the lake. Various studies, although preliminary in nature, have demonstrated that NaCl is relatively irmocuous, but alka- linity (HCO3 + CO3J is toxic to sahnonids 18 Great Basin Naturalist Vol. 43, No. 1 1186 1183 ^ 1180 1176 - 1173 - 1170 - -^ 1167 - a> I 1164 ^ < 1161 - 1158 - 1155 - 1152 _l 1 I 1 1 L. X I \ X \ / \ x-> \" \ • • • • TDS X — X Fragmentary record More complete record (September altitudes are shown 5500 5000 E 4500 H 5 - 4000 - 3500 3000 — I 1 1 1 1 1 1 1 1 1 I 1860 1870 1880 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 Year Fig. 7. Water level and total dissolved solids fluctuations in Pyramid Lake, Nevada, 1867-1979 (From Galat et al. 1981). (Beatty 1959, Mitchum 1960, Taylor 1972, Knoll et al. 1979). The five major species of fish in Pyramid Lake, in order of relative abundance are: tui chub, Tahoe sucker, Lahontan cutthroat trout, cui-ui (Chasmistes cujus), and Sacra- mento perch {Archoplites interruptus). Al- though numerous species have been in- troduced, the current species composition is almost exclusively represented by the original fish species, the only exception being the Sac- ramento perch (Vigg 1981). This fact is prob- ably due to the harsh environmental condi- tions of Pyramid Lake, specifically the TDS levels. In contrast, exotic fish introductions have nearly extirpated the native fish fauna of oligotrophic (low TDS) Lake Tahoe at the upper end of the Truckee River (Miller 1951). The maximum surface (0-1 m) water tem- perature in Pyramid Lake was 21.4 and 23.1 C in July 1976 and August 1977, respec- tively. The lake is thermally stratified from June through December; wind-generated mixing occurs from January through May. The thermocline forms at a depth ranging from 16 to 22 m. The euphotic depth aver- aged 11 m for 1976 and 1977, which resulted in a trophogenic zone of about 4.67 km^ (Ga- lat et al. 1981). Surface-dissolved oxygen (DO) is above 8 mg/1, and thus not limiting to fish. Meta- and hypolimnetic DO depletion occurs beginning in July following stratification and algal de- composition; maximum DO deficits occur in the profundal zone just prior to late fall mix- ing. Hypolimnetic DO deficits in stratified lakes are generally associated with decompo- sition of organic matter, which is generated by primary production in surface water, and gradually sink to the bottom. During Decem- ber, the mean DO level is <4 mg/1 at depths January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT 19 >61 m and <0.2 mg/1 at depths >92 m (Vigg 1980). In contrast to the anoxic condi- tions, which are very hmited on a temporal and spatial basis in Pyramid Lake, Walker Lake exhibits extensive DO depletions that severely restrict fish distribution (Cooper 1978, Koch et al. 1979). Diatoms {Cyclotella sp. and Stephanodiscus spp.) dominate the phytoplankton commu- nity during winter; but the most abundant chlorophyte, Crucigenia sp., attains max- imum abundance in spring (Sigler and Ken- nedy 1978). Blue-green algae are by far the dominant form in Pyramid Lake, comprising >74 percent. Nodularia spumigena is the most abundant species; blooms begin as early as July and may last as late as October. Tem- poral nutrient dynamics inversely relate to phytoplankton abundance. Following vernal increases of algal growth, orthophosphate and nitrate are depleted and remain at low levels during the summer period of maximum primary production. Silica, in addition to ni- trate, probably limits diatom production in Pyramid Lake (Galat et al. 1981). Benthic macroinvertebrates, periphyton, and zooplankton all are important energy sources for juvenile fish in Pyramid Lake. Di- atom domination of the periphyton commu- nity is demonstrated by sampling with glass slides (<99 percent). The chlorophyte, Cladophora glomerata, was the dominant epilithophyton in Pyramid Lake during May and June in 1976 and 1977 (Sigler and Ken- nedy 1978). Chironomids are the lake's most abundant macroinvertebrates (63 percent), followed by Oligochaetes (33 percent), which are especially abundant in the profundal zone (Robertson 1978). Two euryhaline am- phipods, Gammarus lacustris and Hyallela azteca, are associated with tufa and rocks. The zooplankton community is composed of five cladocerans, three copepods, and four rotifers (Lider and Langdon 1978). The cla- doceran, Diaptomus sicilis, is a perennial spe- cies and the most abundant zooplankter throughout the year. The Lahontan cutthroat trout is the third most numerous fish in Pyramid Lake. Com- pared to the more abundant tui chubs and Tahoe suckers, the trout population is numer- ically small, about 1.3 percent (Vigg 1981). However, the relative biomass of the Lahon- tan cutthroat trout population is estimated at 6.4 percent. Theoretically, the biomass of a primary piscivore such as cutthroat trout may be as much as 20 percent of the biomass of the fish forage (McConnell et al. 1978). The trout population at present is far below its theoretical maximum. Activity of the Lahontan cutthroat trout population is at a maximum from December through March. Peak spawning migrations occurred during April and May in 1976 and 1977 (Fig. 6). In 1978 the run was from March 8 to June 13 (Wolcott 1978). The greatest trout activity observed in our study corresponds very closely to the historical spawning period of the winter race of Pyra- mid Lake Lahontan cutthroat trout (Snyder 1917). Snyder observed that the spawning migrations of Lahontan cutthroat occurred in two distinct periods. The larger winter run of trout out of Pyramid and Winnemucca lakes began following the rise in river flows Octo- ber-December; the spawning migration ex- tended through March. As the winter run waned, the spring nm of the smaller, darker, and more heavily spotted trout commenced. This migration peaked in April and extended to May. The sport fishing catch was highest in win- ter, corresponding to the high catches in the 1975-1977 net sampling program (Figure 8). The proportion of large trout was greatest during winter. The high level of winter activ- ity of the Pyramid Lake population of La- hontan cutthroat trout is apparently a mani- festation of innate spawning-related behavior. The larger and older spawners in the cutthroat trout population in Yellowstone Lake are predominant in the early part of each spawning run, with the smaller spawn- ers comprising the latter part of the runs (Bulkley and Benson 1962). The differences between the 1976-1977 Marble Bluff spawning runs and the activity patterns of the lake population of cutthroat trout may be explained by three factors: (1) few Pyramid Lake Lahontan cutthroat trout were apparently imprinted on the Truckee River; (2) the spawning runs were composed of a disproportionately large number of cut- throat-rainbow hybrids that were raised (thus imprinted) in the Truckee River watershed; 20 Great Basin Naturalist Vol. 43, No. 1 -S 38- > -Q 36 ■ o ■S 34 ■ g- 32 o T 30- "I 28 -i 26 24 - 22- 20 u> 16 3 14 I 12- i 10 ■ o I 8 -5 6 o i 4^ S 2 '^ 0 78 92 i i ^ 1 ^;^ 19 i 32 152 i ^ 14 30 35 m 96 i I 1 ^ 76 i I 1 Fig. 8. The Pyramid Lake, DJ FMAMJ JASONDJ FMAMJ JASON 1976 1977 Month and year proportion of large (>550 mm) cutthroat trout taken in the monthly bottom-set gill net catches in Nevada, November 1975 through 1977. and (3) Lahontan cutthroat trout are prob- ably genetically programmed for winter as well as spring spawning, but early winter peak flows are now diminished or eliminated by diversions. For example, during 1976 flows in the lower river peaked in March and steadily decreased to the lowest annual level in December (U.S. Geological Survey 1977). Spawning cutthroat trout instinctively re- turn to the stream in which they were born (Ball 1955, Platts 1959a, McCleave 1967, Jahn 1969). Their olfactory development pat- terns indicate cutthroat trout are capable of imprinting on a home stream odor at a very early age (Jahn 1972). Moreover, mature cut- throat are able to return to home streams even when deprived of their sense of vision or smell, indicating an inborn "compass" homing mechanism. Ball (1955) says, since predetermination of spawning site is established by early life stream association, streams can be lost as nur- sery areas if no natural reproduction occurs, even though mature adults are present in the lake. Ball postulates that nonimprinted fish, such as gravid adults, may randomly move throughout a lake or mill about in the area where they were planted. On an annual basis the majority of Lahon- tan cutthroat trout in Pyramid Lake occur at depths <61 m (Fig. 9). Compared to other species, its depth preference is intermediate, and most closely associated with that of tui chub. There is a differential seasonal depth distribution of cutthroat trout (predator) and tui chub (prey) in Pyramid Lake, i.e., inshore versus 46 m (Fig. 10). Cutthroat trout appar- ently prefer inshore areas during all seasons except summer, when shallow water temper- atures are high. Tui chub are generally in- shore during spring and summer, inter- mediate during autumn, and offshore during January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TroUT 21 Percent species composition " -^ Tui chub Mean catch ■■ ■■ Cutthroat trout ^ -A Tui chub • • Cui-ui o o Tahoe sucker Depth (m) Fig. 9. Percent species composition of tui chub and percent of the mean catch of cutthroat trout, tui chub, cui-ui, and Tahoe sucker by depth in Pyramid Lake, Nevada. Data are derived from the total catch of 108 bottom gill net sets on a quarterly basis (September, December, March, and June) during 1976-1977 (Vigg 1980). winter. Maximum overlap of the two popu- lations occurs during spring, the period of maximum cutthroat trout growth. The two species are opposite with respect to depth distribution during winter, when trout meta- bolism and feeding are low. Changes in net catch/effort and benthic depth distribution of cutthroat trout occurred on a seasonal basis in 1976-1977 (Vigg 1978). During late fall and winter, when the total catch rate of cutthroat trout was about 1.5 times that of other seasons, they inhabited predominantly inshore areas. As surface wa- ter temperatures increase from 10 to 16 C during late spring, the trout population moves into cooler, deeper waters (Fig. 11). This temperature relationship clearly illus- trates the habitat preference of Lahontan cutthroat trout for cooler waters than their prey the tui chub. The lowest summer den- sity of cutthroat trout occurs in littoral ben- thic and inshore surface waters, and the high- est density in benthic waters in or below the thermocline. During the summer months, cutthroat are well represented at depths of 20-60 m in benthic areas, while avoiding the surface waters of the offshore limnetic zone. From June to October the majority of the limnetic trout are at depths of 15-28 m with negligible ntmibers at greater depths (Vigg 22 Great Basin Naturalist Vol. 43, No. 1 100 90 80 70 60 50 40 30 20 10 0 n Cutthroat trout ^ Tuichub n J i i i i t w i i Fall Winter Spring Summer Fall Winter Spring Sumnner Fall (N-D) (J-M) (A-J) (J-S) (O-D) (J-M) (A-J) (JS) (OD) 1975 1976 1977 Season, month and year Fig. 10. Percent of the total seasonal variable mesh bottom-set gill net catches of cutthroat trout and tui chub taken at inshore (versus 46 m) sampling stations in Pyramid Lake, Nevada, from November 1975 through December 1977. 1980). As surface temperatures cool below 16 C in the fall, trout return to surface waters and inshore areas. The profundal zone of Pyramid Lake (>61 m), which constitutes about half of the bot- tom area and 20 percent of the volume, is nearly devoid of cutthroat trout in summer. Temperatures at these depths are less than 7 C during all seasons, and oxygen is low dur- ing the fall and early winter. Cutthroat trout densities are slightly higher in the profundal zone during winter, but this deep area is not an important habitat for trout (Sigler and Kennedy 1978). Management Management of any fishery should ensure that biological, social, economic and political values are given appropriate consideration so as to produce maximum benefits to society from a given stock of fish. Although the pri- mary beneficiary of the Pyramid Lake fishery is the Pyramid Lake Paiute Indian Tribe, so- ciety, the secondary beneficiary, must also be satisfied if the goal of the tribe is to be reached. This goal, as articulated by the U.S. Justice Department, is to produce a viable fishery in Pyramid Lake. The above state- ment may be assumed to be synonymous with or an extended definition of viable fishery. As far as can be determined, there is vir- tually no natural reproduction of Lahontan cutthroat trout in the Truckee River at pres- ent. An important aspect of the management program should be to reestablish successful spawning runs in the Truckee River (Innis et al. 1981). This will require rehabilitation to stabilize stream banks and provide shade to reduce water temperatures, installation of fish ladders to permit spawning adults to mi- grate upriver, fish screens to keep down- stream migrants from entering irrigation canals, and augmented stream flow during critical seasons. Sufficient water will be re- quired for adults to migrate in winter and early spring; to keep temperatures below 13.3 C through the fry stage; and below January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT 23 Fig. 11. Percent of the catches of cutthroat trout and tui chub taken on the surface from surface and bottom gill nets (adjusted to unit of net area) at the 23 m depth in Pyramid Lake, Nevada, from February through November 1977. 21.8 C (Vigg and Koch 1980) during juvenile residence. Since it will take years to restore river habitat, the Lahontan cutthroat trout population must be sustained by stocking. The catch rate of legal size (>380 mm TL) Lahontan cutthroat trout in 1977 was ap- proximately one fish per 14 hours of effort. Under the minimum legal size of 457 mm the catch rate was one fish per 18.9 hours (Alan Ruger, pers. comm. 1982). Catch rates should be increased five- to tenfold to fall within ac- ceptable limits. This will require substantially increased recruitment rates and the reduction of incidental causes of mortality. Since ma- ture fish appear to congregate in the vicinity of stocking sites in late fall, winter, and early spring, during the time when sport fishing is best, some of the stocking effort should be di- rected to the vicinity of popular fishing areas and access points (Table 10). Unless Lahontan cutthroat trout have been imprinted on Truckee River water, they are disinclined to attempt to ascend the river to spawn. Some hatchery fish should be stocked in the lower river to initiate a spawning run. The trout reared in the PLITE Numana Hatchery may be imprinted on the Truckee River because the hatchery outfall runs into the river. The reasons for the very substantial spawning runs of Lahontan cutthroat trout up the Sutcliffe outflow (>9000 in 1982) ap- pears to contradict some long-held beliefs and raises more questions than answers. Alan Ruger (pers. comm. 9 June 1982) thinks the fish are returning to hatchery odors, such as fish feed and juvenile fish. The modified an- nual temperature regime in the Truckee Riv- er, due to the regulation in flow that will be necessary to provide for all uses, must be considered in planning for the reestablish- 24 Great Basin Naturalist Vol. 43, No. 1 ment of natural reproduction in the river. Temperatures acceptable for spawning and incubation of eggs exist during winter and early spring; thus successful natural repro- duction in the river will depend in part on an early spawning migration. Behnke (1979) sug- gests selecting maximum-size fish at first maturity for breeding stock. He also recom- mends using the genetic diversity in remnant stocks to produce the best-adapted strain to Pyramid Lake conditions. As pointed out by Snyder (1917), Pyramid Lake Paiute Indians early in the century recognized two spawn- ing runs of Lahontan cutthroat trout: one in- volved large fish in late November, Decem- ber, and January; the other occurred in the spring when the smaller fish spawned. The spring-spawning fish always faced the haz- ards of high water temperatures, but not at the level that exists today. Competition for river flows is much more intense in spring than in winter. It is generally agreed that some remnant of the original gene pool of Lahontan cutthroat trout persists. Therefore, some of the present population should and apparently do tend to spawn in winter. When hatchery brood stock are to be used as a source of eggs, a program of selective breeding utilizing early-spawning fish should be initiated. Stocking in the river and lake should be limited to Lahontan cutthroat trout; hybrids should not be utilized. Benthic invertebrates are the major food source of Lahontan cutthroat trout until they exceed 300 mm FL. Survival of smaller stocked fish may be limited by the avail- ability of benthic invertebrates. The feasibil- ity of stocking fish as large as 300-330 mm, and their survival and costs in comparison to the size conventionally stocked, should be evaluated. In view of the larger number of trout that must be stocked, introducing larger fish could eliminate benthic invertebrate abundance as a limiting factor and thus in- crease survival rates of stocked fish. Currently all trout <483 mm TL that are landed must be released. The present catch rate of undersized fish is much greater than Table 10. Comparison of gill netting catch rate, surface water temperature, and trout fishing success during 1977 at Pyramid Lake, Nevada. Gill net data Fish per i net Percent (inshore 0 (inshore) Monthly Two-month X Monthly Two-month X January 35 33.5 52 53 February 32 31 54 59 March 30 20 63 54 April 10 19.5 39 54 May 29 22.5 64 54 June 16 13 42 30 July 10 10.5 21 25 August 11 14 28 33 September 17 21.5 37 50 October 26 24.5 65 57 November 23 44.5 50 76 December 66 92 January February March January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TroUT 25 those fish longer than 483 mm. The reason for releasing a fish is the assumption that it will survive to spawn and/ or be caught later. This assumption should be tested, and the size limit implemented accordingly. The Lahontan cutthroat trout now in Pyra- mid Lake probably is physiologically capable of hybridizing with rainbow trout. A popu- lation of mature rainbow trout in the middle and upper Truckee River where Lahontan cutthroat trout spawn would, therefore, po- tentially threaten the maintenance of the lake strain. In addition, brown trout residents in the upper Truckee River will compete with and prey on young Lahontan cutthroat trout. Since a large part of the Truckee River is managed by the Fish and Game Depart- ment of California and the Nevada Wildlife Department, the decision is theirs to imple- ment reduction in the nonnative resident populations of the river fish. Nonfishing recreation on Pyramid Lake represents approximately 500,000 hours of use annually. This use is concentrated during the summer months, but lasts from May through November. Nonfishing recreationists currently represent a significant segment of the lake users, almost twice the use of fishing effort (Fig. 12). Summary The lake form of Lahontan cutthroat trout is the largest of all cutthroat. Its ancestors in- vaded ancient Lake Lahontan from the Co- lumbia River drainage about 70,000 years BP. Before the coming of white men, the La- hontan cutthroat trout was a staple in the diet and an item of trade for the Paiute In- dians of Pyramid Lake. Later, both white men and Indians commercialized the trout fishery in markets as far away as San Fran- cisco. At one time the annual production may have been as much as 454,000 kg. In 1943 the last of the Lahontan cutthroat trout disappeared from Pyramid Lake. Very few had been seen after 1938. Lahontan cutthroat and other trout were stocked in the lake Table 10 continued. Total catch Fish kept Percent fish kept Trout per hour Water temperature Total catch rate Catch rate of fish kept X size surface keeper 6.81 4381 3180 73 .0837 .0608 523 6.60 4993 2495 50 .0956 .0471 538 6.60 7266 3193 44 .1558 .0685 521 8.73 1926 820 43 .0946 .0403 529 10.28 301 101 34 .0458 .0154 478 16.32 439 164 63 .0849 .0317 437 21.28 60 60 .0073 .0073 23.12 26 26 .0130 .0130 21.21 2234 750 34 .3471 .1165 459 16.51 5846 1992 34 .3246 .1106 455 14.50 9164 3495 38 .3452 .1317 498 11.50 7.80 7.10 10.00 7205 4970 2997 4941 3654 2050 969 1924 51 41 32 38 .2250 .1211 .0755 .1513 .1141 .0499 .0244 .0589 488 556 565 482 26 Great Basin Naturalist Vol. 43, No. 1 130, 120 110 100 90 Nonfishing recreation Ibtol hsnermen Fig. 12. J F 1977 Month and year Hours per month of fishing and nonfishing recreation at Pyramid Lake, Nevada, in 1977-1978. Starting in 1950. Today, there is virtually no natural reproduction. Lahontan cutthroat trout in Pyramid Lake live six to seven years. They start maturing at age three to four, some as early as December or as late as April, May, or June. There may be what amoimts to two potential spawning runs. An average-size female produces about 3815 eggs and is mature when the gonadal somatic index reaches 11 percent. None of the eight potential disease organisms ex- plored in 1976-1977 were considered a haz- ard. The fish are most active in the lake from December through March, a time of most fishing effort. Cutthroat <300 mm feed pri- marily on invertebrates; after that size they feed heavily on fish. Five species of fish con- stitute >99 percent of the population. They are, in order of abundance, tui chub, Tahoe sucker, Lahontan cutthroat trout, cui-ui, and Sacramento perch. Pyramid Lake, entirely within the Pyramid Lake Paiute Indian Reservation, is the termi- nus of the Truckee River, which is its only source of water except for a few desert show- ers. The average annual loss to evaporation is 1,2 m. The lake is 40 km long, 6.5 to 16 km wide, covers an area of 446.4 km^, and has a mean depth of 59 m and a maximum of 103 m. Derby Dam, completed in 1905, effects a transbasin diversion of part of the Truckee River flow. The TDS of Pyramid Lake in 1977, at an altitude of 1157 m, was 5235 mg/1. The base load of TDS is reasonably stable. Pyramid Lake stratifies into three well-defined layers in June-July. It destra- tifies in December-January. It is a midlevel productivity lake. Pyramid Lake, a remnant of Lake Lahontan, has a pH of 9.2 and is high in carbonates and bicarbonates. Summer sur- face temperatures are 21-23 C. There is ample dissolved oxygen in the epilimnion and thermocline at all times. Nodularia, a blue- green alga, dominates much of the lake from late summer to early fall. Diatoms dominate the periphyton communities. Chironomids are the most abundant macroinvertebrates. Conclusions The Lahontan cutthroat trout fishery in Pyramid Lake is currently not a viable one. The annual catch was < 20,000; the rate, one fish for > 14 hours effort, when the minimum January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT 27 legal size was 381 mm TL. Fishing success should be increased in the magnitude of five-ten times. Any adverse changes in the lake ecology may stress the fish that will in turn make them more susceptible to disease. The 1905 diversion of the Truckee River, which in dry years may take most of the flow, reduced available stream spawning area for the cutthroat from >500 km to <62 km of substandard stream. Derby Dam, over a period of 25 years, doomed the historical cut- throat fishery. Successful reproduction in the lower river demands stable riparian habitat and water temperatures < 13.3 C, until after spawning-hatching-fry emergence and <21.8 C thereafter. Brood stock or wild egg-pro- ducing fish for hatcheries should be selected for large size at first maturity and for winter or early spring maturing. Part of the matur- ing fish in the lake should be imprinted on the Truckee River. Since the effluent from Numana Hatchery flows into the Truckee River, it may be these trout will be im- printed. The biological implications of the large lam of cutthroat into the Sutcliffe flow should be explored in depth. Since the base load of TDS is constant in Pyramid Lake, the concentration varies in- versely to lake volume. Any significant in- crease in TDS may prove harmful to key or- ganisms in the food chain and to the trout. The median level of productivity that Pyra- mid Lake currently enjoys is considered more desirable than a higher level for Pyramid Lake Lahontan cutthroat trout. Acknowledgments This work was performed under Bureau of Indian Affairs contract H50C 14209487. As- sistance and cooperation was provided by employees of W. F. Sigler & Associates Inc. (WFSAI), members of the Pyramid Lake Paiute Indian Tribe, and the U.S. Fish and Wildlife Service, Fisheries Assistance Office, Reno, Nevada. Denise Robertson and Roy Whaley, formerly of WFSAI, were respon- sible for the two sections on age and growth, and food habits, respectively. The manuscript was reviewed by Alan Ruger, fisheries direc- tor. Pyramid Lake Indian Tribal Enterprises, Sutcliffe, Nevada. Literature Cited American Fisheries Society: Fish Health Section. 1975. Suggested procedures for detection and identification of certain infectious diseases of fishes. Amer. Fish. Soc, Washington, D.C. Ball, O. P. 1955. Some aspects of homing in cutthroat trout. Proc. Utah Acad. Sci., Arts, Lett. 32:75-80. Ball, O. P., and O. B. Cope. 1961. Mortality studies on cutthroat trout in Yellowstone Lake. U.S. Dept. Inter., Fish Wildl. Serv. Res. Rep. 55. 62 pp. Beatty, D. B. 1959. An experimental study of the tox- icity of sodium bicarbonate, sodium chloride, and sodium sulfate to rainbow trout. Unpublished thesis. Univ. of Wyoming. Behnke, R. J. 1974. The effects of the Newlands project on the Pyramid Lake fishery. File Rep. 15 pp. 1979. Monograph of the native trouts of the genus Salmo of western North America. U.S. For. Serv., U.S. Fish Wildl. Serv., U.S. Bur. Land Mgt. Washington, D.C. Behnke, R. J., and M. Zarn. 1976. Biology and manage- ment of threatened and endangered western trout. USDA, For. Serv. Gen. Tech. Rep., Fort Collins, Colorado. 45 pp. Benson, L. V. 1978. Fluctuations in the level of pluvial Lake Lahontan for the past 40,000 years. Quar- temary Res. 9:300-318. BjoRNN, T. C. 1957. A survey of the fishery resources of Priest and Upper Priest lakes and their tribu- taries. Dingell-Johnson Completion Rep. Idaho Fish Game Dep., Boise, Idaho F-24-R. 176 pp. Brown, C. J. D., and J. E. Bailey. 1952. Time and pat- tern of scale formation in Yellowstone cutthroat trout {Salmo clarkii lewisii). Trans. Amer. Mi- crosc. Soc. 71(2): 120-124. BuLKLEY, R. V. 1961. Fluctuations in age composition and growth rate of cutthroat trout in Yellowstone Lake. U.S. Fish Wildl. Serv. Res. Rep. 54. 31 pp. BuLKLEY, R. v., and N. G. Benson. 1962. Predicting year-class abundance of Yellowstone Lake cut- throat trout. U.S. Fish Wildl. Serv. Res. Rep. 50:1-20. Calhoun, A. J. 1942. The biology of the black-spotted trout {Salmo clarkii henshawi [Gill and Jordan]) in two Sierra Lakes. Unpublished dissertation. Stanford University. 218 pp. 1944. The food of the black-spotted trout {Salmo clarkii henshawi [Gill and Jordan]) in two Sierra Nevada lakes. California Fish Game Dep. 30(2): 80-85. Carlander, K. D. 1969. Handbook of freshwater fish- eries biology, Iowa State Univ. Press, Ames, Iowa. 1:1-752. Cooper, J. J. 1978. Contributions to the life history of the Lahontan tui chub, Gila bicolor obesa (Gi- rard), in Walker Lake, Nevada. Unpublished thesis. Univ. of Nevada, Reno, Nevada. Cope, O. B. 1953. Length measurements of Yellowstone Lake trout. U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish. 102. 17 pp. Deacon, J. £., G. Kobetich, J. D. Willl\ms, and S. Contreras. 1979. Fishes of North America en- dangered, threatened, or of special concern. Fish. Bui. Amer. Fish. Soc. 4(2):29-44. 28 Great Basin Naturalist Vol. 43, No. 1 Drummond, R. a. 1966. Techniques in the collection and mounting of trout scales. Prog. Fish Cult. 28(2):113-116. Echo, J. B. 1955. Some ecological relationships between yellow perch and cutthroat trout in Thompson Lakes, Montana. Trans. Amer. Fish. Soc. 84:239-249. Ferjancic, K. p. 1976. The results of the 1976 I.P.N. (In- fectious pancreatic necrosis) survey conducted on Lahontan cutthroat trout (Salmo clarki henshawi) at Pyramid Lake, Nevada. Pyramid Lake Indian Tribal Enterprises, Sutcliffe, Nevada Rept. 4 pp. FiNNELL, L. M. 1966. Granby Reservoir Studies. Colo- rado Div. Wildl., Fisheries Res. No. 3:4-6. Fleener, G. G. 1952. Life history of the cutthroat trout {Salmo clarkii Richardson), in Logan River, Utah. Trans. Amer. Fish. Soc. 81:235-248. Galat, D. L., E. L. Lider, S. Vigg, and S. R. Robertson. 1981. Limnology of a large, deep North American terminal lake. Pyramid Lake, Nevada. USA. Hydrobiologia. 82:281-317. Harris, E. E. 1970. Reconnaissance bathymetry of Pyra- mid Lake, Washoe County, Nevada. Hydrologic Investigation Atlas HA-379, U.S. Geol. Surv. Hazzard, a. S., and M. J. Madsen. 1933. Studies of the food of the cutthroat trout. Trans. Amer. Fish. Soc. 63:198-207. Hickman, T. J., and D. A. Duff. 1978. Status of cut- throat subspecies in the western Bonneville Ba- sin. Great Basin Nat. 38(2): 193-202. Hickman, T. J., and R. J. Behnke. 1979. Probable dis- covery of the original Pyramid Lake cutthroat trout. Prog. Fish Cult. 41:135-137. Hile, R. 1941. Age and growth of rock bass (Ambloplites rupestris Rafinesque) in Nebesh Lake, Wisconsin. Trans. Wisconsin Acad. Sci., Arts, Lett. 33:189-337. HuBBS, C. L., and R. R. Miller. 1948. The Great Basin with emphasis on glacial and postglacial times. n. The zoological evidence. Univ. of Utah. Bull. 38(20): 17-166. Houghton, S. G. 1976. A trace of desert waters. The Great Basin story. Arthur H. Clark Co., Glendale, California. 287 pp. Hutchinson, G. E. 1957. Treatise on limnology: geogra- phy, physics, and chemistry. Vol. 1. John Wiley and Sons, New York. 1015 pp. Innis, G. S., D. F. Hanson, and J. W. Haefner. 1981. A simulation model of management alternatives in a freshwater fishery. Elsevier Scientific Publ. Co. Amsterdam, The Netherlands. 12:267-280. Irving, R. B. 1953. Ecology of the cutthroat trout {Salmo clarkii Richardson) in Henry's Lake, Idaho. Un- published thesis. Utah State Agricultural College. 101 pp. Jahn, L. a. 1969. Movements and homing of cutthroat trout {Salmo clarki) from open-water areas of Yel- lowstone Lake. J. Fish. Res. Bd. Can. 26:1243-1261. 1972. Development of the olfactory apparatus of the cutthroat trout. Trans. Amer. Fish. Soc. 101(2):284-289. Johnson, M. W., and L. Wroblewski. 1962. Errors as- sociated with a systematic sampling creel census. Trans. Amer. Fish. Soc. 91(2):201-207. Johnson, V. K. 1958. Fisheries management report. Lakes— Pyramid, Walker, and Tahoe. Dingell- Johnson Completion Rep. Nevada Fish and Game Dept. FAF-4-R. 47 pp. Kennedy, J. L. 1978. A creel survey of Pyramid Lake, Nevada. Pages 414-437 in W. F. Sigler and J. L. Kennedy, eds.. Pyramid Lake ecological study. W. F. Sigler & Associates Inc., Logan, Utah. Knoll, J., D. L. Koch, R. Knoll, J. Sommer, L. Hoffman, and S. Lintz. 1979. Physiological ad- aptations of salmonid fishes {Salmo clarki hen- shawi, Salmo gairdneri and Oncorhynchus kisutch) to alkaline saline waters and their toxic effects. Report to Office of Water Research and Technology. Desert Res. Inst. Bioresources Cen- ter Piibl. 50009. Reno, Nevada. 82 pp. Koch, D. L., J. J. Cooper, E. L. Lider, R. L. Jacobson, and R. J. Spencer. 1979. Investigations of Walker Lake, Nevada: Dynamic ecological relationships. Desert Res. Inst. Bioresources Center Publ. Reno, Nv. No. 50010. 191 pp. Kucera, p. a., and J. L. Kennedy. 1977. Evaluation of a sphere volume method for estimating fish fecun- dity. Prog. Fish Cult. 39(3):115-117. Laakso, M., and O. B. Cope. 1956. Age determination in Yellowstone cutthroat trout by the scale meth- od. J. Wildl. Mgt. 20(2): 138-153. Lea, R. N. 1963. Ecology of the Lahontan cutthroat trout {Salmo clarki henshawi) in Independence Lake, California. Unpublished thesis. Univ. of California, Berkeley. 95 pp. LeCren, E. D. 1951. The length-weight relationship and seasonal cycle in gonad weight and condition in perch {Perca fluviatilis). J. Anim. Ecol. 20(2):201-219. Lider, E. L., and R. W. Langdon. 1978. Plankton ecolo- gy. Pages 241-293 in W. F. Sigler and J. L. Ken- nedy, eds., Pyramid Lake ecological study. W. F. Sigler & Associates Inc., Logan, Utah. Madsen, M. J. 1940. Report on age and growth of cut- throat trout {Salmo lewisii) of Yellowstone Lake, Wyoming. U.S. Fish Wildl. Serv. Res. Rept. 12 pp. Marnell, L. F. 1969. Hooking mortality of cutthroat trout. Unpublished dissertation, Colorado State Univ. 99 pp. Marnell, N. E., and D. Hunsaker 11. 1970. Hooking mortality of lure-caught cutthroat trout {Salmo clarki) in relation to water temperature, fatigue and reproductive maturity of released fish. Trans. Amer. Fish. Soc. 99(4): 684-688. McCarraher, D. B. 1972. A preliminary bibliography and lake index of the inland mineral waters of the world. FAO. Fish. Cir. 146. McCleave, J. D. 1967. Homing and orientation of cut- throat trout {Salmo clarki) in Yellowstone Lake with special reference to olfaction and vision. J. Fish. Res. Bd. Can. 24(10):201 1-2044. McCoNNELL, W. J., D. L. Galat, and K. Hamilton- Galat. 1978. Potential fish production of Pyra- mid Lake based on organic matter contributions. Colorado Coop. Fish. Res. Unit. Fort Collins, Colorado. 84 pp. January 1983 SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT 29 Miller, R. G. 1951. The natural history of Lake Tahoe fishes. Unpubhshed dissertation. Stanford Univ. 160 pp. MiTCHUM, D. L. 1960. An experimental study of the tox- icity of calcium carbonate, calcium sulphate, magnesium carbonate, and magnesium sulphate to rainbow trout. Unpublished thesis. Univ. of Wyoming. Nelson, L. 1976. SHAD II. A model for analysis of fish- eries age and growth data. Wildl. Sci. Dep., Utah State Univ., Logan, Utah. 15 pp. NiKOLSKY, C. B. 1963. The ecology of fishes. Acad. Press, New York. ,393 pp. Peters, J. C, ed. 1964. Summary of calculated growth data on Montana fishes, 1948-1961. Dingell-John- son Completion Rep., Montana Fish and Game Dept., Helena, Montana. F-23-R. 76 pp. Platts, W. S. 1959a. Homing, movements, and mor- tality of wild cutthroat trout {Salmo clarki) spawned artificially. Prog. Fish Cult. 1959 (January):36-38. 1959b. Food habits of the cutthroat trout in Strawberry Reservoir, Utah. Proc. Utah Acad. Sci., Arts, Lett. 36:119-121. Rankel, G. L. 1976. Fishery management program, 1976, Summit Lake Indian Reservation, Hum- boldt County, Nevada. Manuscript, U.S Fish Wildl. Serv., Reno, Nevada. 35 pp. RiCKER, W. E., ed. 1971. Methods for assessment of fish production in freshwaters. Blackwell Sci. Publ., Oxford. 348 pp. Robertson, O. H. 1947. An ecological study of two high mountain trout lakes in the Wind River Range, Wyoming. Ecology 28(2):87-112. Robertson, S. R. 1978. The distribution and relative abundance of benthic macroinvertebrates in Pyramid Lake, Nevada. Unpublished thesis, Univ. of Nevada. RuGER, A. W. 1977. The results of the 1977 disease sur- vey conducted on Lahontan cutthroat trout (Sal- mo clarki henshawi) at Pyramid Lake, Nevada. Manuscript, Pyramid Lake Indian Tribal Enter- prises, Sutcliffe, Nevada. SiGLER, W. F. 1951. The life history and management of the mountain whitefish (Prosopium williamsoni (Girard)) in Logan River, Utah. Utah State Agri- cultural College Bull. 347. 36 pp. 1962. Bear Lake and its future, 26th Fac. Assn. Honor Lecture. Utah State Univ., Logan, Utah. 23 pp. SiGLER, W. F., AND J. L. KENNEDY, eds. 1978. Pyramid Lake ecological study. W. F. Sigler & Associates Inc., Logan, Utah. 545 pp. SiGLER, W. F., AND J. B. Low. 1950. Age composition and growth of fish and fishermen success in Utah's high Uinta lakes. Utah Acad. Sci., Arts, Lett. 27:32-36. Sigler, W. F., and R. R. Miller. 1963. Fishes of Utah. Utah Dept. Fish and Game, Salt Lake City, Utah. 203 pp. Smith, S. H. 1963. Making plastic impressions of fish scales with a roller press. Prog. Fish Cult. 16(2): 75-78. Snyder, G. R., and H. A. Tanner. 1960. Cutthroat trout reproduction in the inlets to Trappers Lake. Col- orado Div. Wildl. Tech. Bull. 7:1-85. Snyder, J. O. 1917. The fishes of the Lahontan system of Nevada and northeastern California. U.S. Bur. Fish. Bui. 1915-16(35:31-86). Sumner, F. H. 1939. The decline of Pyramid Lake fish- ery. Trans. Amer. Fish. Soc. 69:216-224. Taylor, R. E. L. 1972. The effects of increasing salinity on the Pyramid Lake fishery. Agric. Expt. Sta., University of Nevada. 8 pp. Tesch, F. W. 1971. Age and growth. Pages 98-131 in W. E. Ricker, ed., Methods of assessment of fish production in freshwaters. Blackwell Sci. Publ., Oxford. TowNLEY, J. M. 1980. The Truckee Basin fishery, 1844-1944. Nevada Historical Society in coopera- tion with Desert Research Institute, Univ. of Ne- vada, Reno. Trelease, T. J. 1953. The death of a lake. Field and Stream. February. 1969. The rebirth of a lake. Nevada Outdoors and Wildlife Review. Nevada Dept. Fish and Game 4:3. United States Geological Survey. 1977. Water re- sources data for Nevada, water year 1976. U.S. Dept. Inter., Carson City, Nevada. 344 pp. Van Oosten, J. 1923. The whitefishes [Coregonus clupeaformis). A study of the scales of whitefishes of known ages. Zoologica 11(17):380-412. 1929. Life history of the lake herring (Coregonus artedii LeSueur) of Lake Huron as revealed by its scales, with a critique of the scale method. U.S. Bur. Fish. 44(1928):265-428. 1944. Factors affecting the growth of fish. Trans. Ninth N. Amer. Wildl. Conf. 9:177-183. Vigg, S. 1978. Vertical distribution of adult fish in Pyra- mid Lake, Nevada. Great Basin Nat. 38(4):417-428. 1980. Seasonal benthic distribution of adult fish in Pyramid Lake, Nevada. California Fish and Game J. 66(l):49-58. 1981. Species composition and relative abun- dance of adult fish in Pyramid Lake, Nevada. Great Basin Nat. 41(4):395-408. Vigg, S., and D. L. Koch. 1980. Upper lethal temper- ature range of Lahontan cutthroat trout in waters of different ionic concentration. Trans. Amer. Fish. Soc. 109:336-339. Wheeler, S. S. 1974. The Desert Lake: the story of Ne- vada's Pyramid Lake. Caxton Printers, Ltd., Caldwell, Idaho. 133 pp. Wolcott, Roger S. C, Jr. 1978. Evaluation of the 1978 Lahontan cutthroat trout run to the Pyramid Lake fishway and comparison with past year. U.S. Fish Wildl. Serv., Fish. Asst. Office, Reno, Nevada. 30 pp. A REVIEW OF THE GENUS SOLIPERLA (PLECOPTERA: PELTOPERLIDAE) Bill P. Stark' Abstract.— The western Nearctic stonefly genus Soliperla is reviewed and six species are recognized. Soliperla sierra (Calif.) and S. tillamook (Ore.) are described as new to science and illustrations of diagnostic features are pre- sented for all species. Males and nymphs are keyed and a phylogeny for the group is proposed. Soliperla was proposed by Ricker (1952) as a monotypic subgenus of Peltoperla to con- tain P. thyra Needham & Smith. At that time the species was known only from the male holotype but Jewett (1954) described the fe- male along with males and females of two additional species, P. campanula and P. quadrispinula; the nymph of P. campanula was also described at this time. Jewett (1955) described the fourth member of the group, P. fenderi, from a single male. These species have remained poorly known since their dis- covery, with only synoptic notes and regional keys (Jewett 1959, 1960) appearing until Stark and Stewart (1981) gave additional characters that supported Illies's (1966) ele- vation of the group to generic status. During recent field work with colleagues in Washington, Oregon, and California, Soli perla nymphs were common in splash zones of small streams and springs. Through this work, nymphs were associated for the four known species, and the additional material collected along with specimens obtained from museums permits the first com- prehensive treatment of Soliperla. Methods were given by Stark and Stewart (1981). Soliperla Ricker Peltoperla (Soliperla) Ricker 1952: 157. Type-species of subgenus: Peltoperla thyra Needham & Smith. Monotypic. Soliperla lilies 1966:26. Adults and nymphs of Soliperla are unusual among Nearctic Peltoperlidae in displaying distinctive pigmentation patterns. Adults are typified by a dark mesal pronotal stripe that contrasts sharply with the light yellow back- ground (Figs. 23, 31), and nymphs have con- spicuous white areas on the abdominal terga that contrast with the dark background (Figs. 4, 15). Monophyly for the group is asserted on the basis of the distinctive epiproct, with recurved crenulate apex and the membranous pair of lobes associated with the epiproct (Figs. 5, 13). The genus is currently known from the western Nearctic region from cen- tral California to Washington (Fig. 1). Keys to Soliperla males 1. Meso ventral area of aedeagus with two irregular longitudinal rows of short, thick setae (Figs. 3, 30) 2 — Mesoventral area of aedeagus with short, thick setae, if present, not in longitudinal rows 3 2(1). Lateral aedeagal lobes terminating in sclerotized spine with 2-4 small subapical setae (Fig. 29) sierra 'Department of Biology, Mississippi College, Clinton, Mississippi 39058. 30 January 1983 Stark: Review of Soliperla 31 Fig. 1. Distribution of Soliperla species. S. campanula = open circles, S. fenderi = closed circles, S. quad- rispinula = open squares, S. sierra = triangles; S. thyra = closed squares, S. tillamook = x. 32 Great Basin Naturalist Vol. 43, No. 1 — Lateral aedeagal lobes membranous apically with 2-4 small subapical setae (Fig. 2) campanula 3(1). Ventral aedeagal lobes with large sclerotized spine or a single long, thick seta (Figs. 19,35) 4 — Ventral aedeagal lobes membranous with numerous scattered setae (Figs. 12,45) 5 4(3). Ventral aedeagal lobes with sclerotized bilobed process (Fig. 35) thyra — Ventral aedeagal lobes with a terminal long, thick seta (Fig. 19) quadrispinula 5(3). Ventral aedeagal lobes with irregular row of long, slender setae along apical margin (Fig. 12) fenderi — Ventral aedeagal lobes with scattered short setae along apical margins (Fig. 45) tillamook Preliminary key to nymphs (sierra and tillamook unknown) 1. Some long setae in abdominal tergum 9 posterior fringe bent (Fig. 39); abdominal tergum 5 typically with lateral pale spots (Fig. 22) 2 — Long setae in abdominal tergum 9 posterior fringe straight (Fig. 6); abdominal tergum 5 typically without pale spots (Fig. 4) 3 2(1). Mesal area of abdominal tergum 8 posterior fringe with ca 20 clavate setae be- tween long setae (Fig. 40); mesal pale spots on abdominal terga 5 and 6 rounded (Fig. 38) thyra — Mesal area of abdominal tergum 8 posterior fringe with ca 5-7 clavate setae between long setae; mesal pale spots on abdominal terga 5 and 6 irregularly linear to triangular (Fig. 22) quadrispinula 3(1). Lateral pale spots on abdominal tergum 4 conspicuously larger than mesal spot (Fig. 15); known from Mt. Rainier, Washington fenderi — Lateral pale spots on abdominal tergum 4 subequal to mesal spot in size (Fig. 4); widely distributed in northern Oregon campanula Soliperla campanula (Jewett) Females cannot be distinguished with cer- tainty from several related species, and the Peltaperla {Soliperla) campanula Jewett 1954: 167. Holo- ,^. - r.. , , i features type $ (CAS), Oxbow Springs, Hood River Co., ^§8^ ^^^g^' /' ^> ^^^° f ^"^ /° *^^ , teatures Oregon th^t would distinguish them from other mem- Jewett (1954) detailed the major diagnostic bers of the genus. The species is currently features of this species. Males are distin- known only from northern Oregon (Fig. 1). guished from other Soliperla by the structure „ Material EXAMiNED.-Oregon: Cl^ckarna. Co Mt. °c 1 ■ 1 1 4.1 1 Hood, near Timberline Lodge, 31-V-77, K. W. Stewart, of the epiproct and aedeagus. The anterodor- ^ ^ Szczytko, 2 s (reared) (NTSU); same location, 20- sal face of the epiproct is about twice as wide vn-67, S. G. Jewett, l ? (USNM); Mt. Hood, Still Crk. as the stalk, and the lateral margins are Cmp. Gnd., 12-VII-79, B. Stark, K. W. Stewart, 2 s curved inward near the crenulate antero-ven- (^P^); trib. Still Crk, Mt. Hood, 17-VI-67, S. G. Jewett, 1 .. 1 _r /IT- tr\ A .. 1 in 1^ ■ S (USNM); Mt. Hood, 1.2 mi N Hwy 26, 20- vn-67, J. tral surface (Fig. 5). Approximately 12-14 ir- ^^\j ^^^j^^y^ ^^.^^ S^,^^^ ^.^ ^^ ^^^^ 15-vn-54, S. regular teeth are present along this surface. g. Jewett 1 S (USNM). Hood River Co., Oxbow Springs, The ventral aspect of the aedeagus includes a 26-V-40, S. G. Jewett, 15,1? (OSU); Iron Crk, W of large mesal lobe and two small lateral lobes. Bennetts Pass, ll-VII-68, E. Evans, 13,2? (USNM). The mesal lobe has two irregular rows of ^^ ?"' ^f f f'o^nJn^M'^Trf ?t ?V h'^"" , ~ ^r, 1. 1-1 111 78, B. Frost, 1^,1? (OSU); Mack Crk, H. J. Andrews about 5-10 short peglike setae, and the later- g^p p^r., 25-VI-74, N. H. Anderson, l s (OSU); 12.5 al lobes have 2-4 subapical peglike setae mi NE Blue Riv, H. J. Andrews Exp. For., 19-VII-78, B. (Fig. 3). Frost (OSU). Linn Co., Ice Cap Crk, 7-VII-66, J. Bedea, January 1983 Stark: Review of Soliperla 33 Figs. 2-6. S. campanula. Fig. 2. Aedeagus, lateral. Fig. 3. Aedeagus, A = dorsal, B = ventral. Fig. 4 Nymphal ab- domen, dorsal. Fig. 5. Epiproct, anterodorsal. Fig. 6. Nymphal abdominal tergum 8, posterior fringe. 34 Great Basin Naturalist Vol. 43, No. 1 Figs. 7-10. Soliperla eggs. Fig. 7. S. campanula, 280X. Fig. 8. S. campanula, lOOOX. Fig. 9. S. fenderi, 300X. Fig. 10. S. fenderi, lOOOX. 2 3 (OSU); 14 mi NE Blue Riv, H. J. Andrews Expt. For., 6-VII-78, B. Frost 1 wl wl - - 6 _- 5 4 O. D. D- w col O ■Q n -Aponorphic O -Plesionorphic 47 Fig. 47. Proposed phylogeny for Soliperla species. See text for explanation. Acknowledgments Several individuals and museums cooper- ated in providing material for study. These include R. W. Baumann (Monte L. Bean Mu- seum, Brigham Young University), O. S. Flint (United States National Museum), B. Kon- dratieff, J. Lattin (Oregon State University Museum), B. Mather, L. L. Pechuman (Cor- nell University Museum), K. W. Stewart (North Texas State University Museum) and G. Wiggins (Royal Ontario Museujn). K. W. Stewart and D. Ziegler gave valuable assis- tance in collecting and rearing efforts. S. W. Szczytko (University of Wisconsin, Stevens Point) and S. Faison (University of Mississippi Dental School) assisted in preparing SEM mi- crographs. This study was supported, in part, by NSF grant DEB 78-12565. Literature Cited Illies, J. 1966. Katalog der rezenten Plecoptera. Das Tierreich, 82. Walter de Gruyter and Co. 632 pp. 44 Great Basin Naturalist Vol. 43, No. 1 Jewett, S. G. 1954. New stoneflies from California and Oregon (Plecoptera). Pan-Pac. Entomol. 30:167-179. 1955. Notes and descriptions concerning west- em North American stoneflies (Plecoptera). Was- mann Jour. Biol. 13:145-155. 1959. The stoneflies (Plecoptera) of the Pacific Northwest. Oregon State Monographs, Studies Entomol. 3:1-95. 1960. The stoneflies (Plecoptera) of California. Bull. Calif. Insect Surv. 6:125-177. Needham, J. C, AND L. W. Smith. 1916. The stoneflies of the genus Peltoperla. Canadian Entomol. 48:80-88. RicKER, W. E. 1952. Systematic studies in Plecoptera. Indiana Univ. Pub. Sci. Ser. 18:1-200. Stark, B. P., and K. W. Stewart. 1981. The Nearctic genera of Peltoperlidae (Plecoptera). J. Kans. En- tomol. Soc. 54:285-311. Watrous, L. E., and Q. D. Wheeler. 1981. The out- group comparison method of character analysis. Syst. Zool. 30:1-11. A BIBLIOGRAPHY OF COLORADO VEGETATION DESCRIPTION William L. Baker' Abstract.— A list of 658 references to Colorado vegetation description is presented, along with county and sub- ject indexes to the list. This bibliography includes published and selected unpublished references through 1981. Included are references to vegetation description, including studies that contain only qualitative description, along with more detailed quantitative studies. Also included are selected references to age /size structure, fire, succession, floristics, vegetation history (primarily palynology), phenology, plant ge- ography, vegetation zonation, and the tim- berline. An index is included to subjects other than vegetation description. Coverage of these tangential subjects may not be com- prehensive. Autecological and environmental studies are generally excluded, as are studies pertaining only to nonvascular vegetation. References are included here if at least part of the study area is in Colorado. Ex- cluded are the many works from adjoining states that may have relevance, particularly to the margins of Colorado, but have no part of their study areas inside Colorado. Re- searchers should consult bibliographies from adjoining states for these references. Included below is an index to the bibliog- raphy by county. A single reference may per- tain to more than one county. Also included are a list of studies pertaining to the whole state, and a list of "regional studies" that per- tain to an undefined, or poorly demarcated part of the state. Researchers seeking com- plete coverage of a particular county should also check references in these lists. Subject Index Age/Size Structure: 286, 291, 302, 330, 373, 488, 489, 547, 558, 559, 624, 628, 630 Bibliographies: 3, 4, 7, 214, 658 Fire: 2, 3, 4, 5, 13, 19, 24, 25, 37, 38, 49, 70, 92, 93, 98, 113, 114, 150, 152, 153, 171, 211, 228, 234, 235, 249, 306, 395, 396, 417, 490, 556, 557, 655, 656, 658, 659 Floristics: 10, 23, 30, 45, 60, 101, 102, 126, 164, 203, 215, 216, 298, 335, 372, 397, 402, 403, 404, 493, 496, 497, 500, 531, 561, 599, 616 Palynology/Vegetation History: 11, 307, 336, 337, 338, 340, 425, 426, 427, 428, 617 Phenology: 30, 42, 155, 217, 218, 219, 220, 221, 222, 240, 246, 249, 368, 369, 443, 506, 564, 619, 620, 621 Plant Geography: 179, 215, 216, 249, 287, 288, 298, 326, 372, 375, 419, 421, 424, 443, 454, 493, 497, 500, 611, 613, 614, 615, 619, 620, 621 Succession: 5, 8, 13, 15, 21, 24, 25, 26, 32, 37, 38, 49, 69, 70, 73, 83, 84, 92, 93, 98, 105, 112, 113, 114, 125, 128, 131, 140, 141, 146, 150, 152, 153, 161, 171, 178, 182, 183, 184, 185, 188, 211, 228, 229, 234, 239, 242, 255, 258, 261, 263, 286, 299, 301, 311, 312, 318, 319, 321, 327, 343, 345, 346, 347, 352, 354, 360, 375, 384, 395, 396, 400, 401, 407, 415, 417, 418, 420, 429, 448, 466, 472, 478, 483, 490, 504, 507, 510, 517, 518, 519, 520, 524, 525, 537, 542, 545, 546, 547, 548, 556, 557, 558, 559, 580, 611, 624, 625, 626, 627, 630, 632, 633, 635, 636, 637, 638, 639, 640, 641, 646, 653, 656, 658, 659 Timberline: 49, 62, 167, 192, 227, 466, 492, 547, 606 Vegetation Maps: 5, 6, 12, 17, 31, 48, 68, 70, 82, 86, 94, 95, 103, 108, 137, 139, 140, 169, 173, 181, 186, 225, 239, 243, 262, 265, 266, 267, 269, 271b, 290, 291, 292, 293, 294, 299, 301, 322, 323, 329, 336, 337, 350, 351, 353, 361, 369, 376, 388, 400, 401, 403, 404, 409, 461, 468, 479, 486, 487, 512, 521, 522, 533, 538, 544, 547, 562, 563, 568, 582, 586, 588, 589, 597, 623, 631, 647, 660 Vegetation Zonation: 23, 76, 78, 79, 81, 86, 88, 107, 126, 128, 129, 135, 138, 177, 181, 193, 230, 233, 239, 298, 326, 336, 337, 380, 421, 436, 438, 439, 440, 449, 482, 495, 498 County Index Adams: 351, 447, 592, 593 Alamosa: 138, 193, 450, 451, 457, 458 Arapahoe: 351 Archuleta: 508, 542, 648 Baca: 10, 125, 263, 486, 487, 504, 511, 573 Bent: 309, 310, 311, 312, 316 'Colorado Natural Heritage Inventory, 1550 Lincoln Street, Suite 110, Denver, Colorado 80203. 45 46 Great Basin Naturalist Vol. 43, No. 1 Boulder: 8, 14, 20, ,33, 36, 37, 38, 45, 48, 49, 51, 74, 75, 84, 94, 95, 104, 126, 140, 141, 144, 148, 155, 165, 169, 189, 209, 216, 228, 229, 230, 231, 232, 239, 240, 246, 250, 251, 252, 265, 275, 276, 277, 278, 286, 287, 288, 289, 290, 291, 302, 307, 325, 332, .333, 334, 338, 341, 342, ,343, 344, 345, .346, 347, 348, .351, 367, 368, 369, 370, 377, .381, 382, 391, .392, 393, 397, 403, 406, 407, 409, 417, 418, 419, 420, 421, 422, 425, 447, 448, 459, 464, 466, 478, 479, 484, 513, 526, 5.35, .545, 546, 554, 574, 582, 592, 593, 594, 595, 596, 606, 611, 612, 616, 656 Chaffee: 52, 53, 56, 234, .321, 522, 560 Cheyenne: no references Clear Creek: 49, 52, 53, 56, 68, 74, 75, 112, 206, 209, 215, 216, 2.39, 291, 421, 446, 448, 594, 595, 596, 616 Conejos: 138, 139, 193, 450, 457, 458 Costilla: 138, 139, 193, 450, 457, 458, 650, 651, 652 Crowley: no references Custer: 101, 421 Delta: 70, 191, 467, 468, 542, 557, 577, 578 Denver: 351 Dolores: 86, 136, 336, 337, 355, 533, 542 Douglas: 40, 234, 351, ,361, 452, 575, 594, 595, ,596, 616 Eagle: 49, 133, 210, 2,39, 303, 350 Elbert: 161, 351, 361, 619, 620, 621 El Paso: 46, 52, 53, 56, 99, 100, 161, 171, 215, 216, 234, 263, 279, 280, 318, 319, 320, 361, 421, 504, 509, 510, 514, 516, 517, 524, 525, 560, 561, 594, 595, 596, 616, 631, 643, 644, 645, 647 Fremont: 61, 160, 576, 650, 651, 652 Garfield: 58, 59, 70, 82, 133, 157, 158, 177, 204, 208, 213, 239, 253, 266, 267, 269, 270, 271a, 352, 353, 355, 373, 374, 375, 480, 507, 556, ,557, 563, 568/569, 570, 571, 597 Gilpin: 72, 112, 168, 209, 215, 216, 239, 308, 354, 441, 442, 443, 444, 447, 448, 460, 469, 594, 595, 596, 616 Grand: 47, 74, 75, 77, 78, 84, 112, 173, 202, 206, 207, 209, 210, 213, 228, 229, 230, 231, 232, 239, 253, 254, 287, 288, 289, 354, 397, 398, 421, 448, 470, 535, 537, 545, 546, 552, 564, 624, 625, 626, 627, 628, 629, 630, 646 Gunnison: 9, 21, 22, 23, .30, 50, 70, 71, 73, HI, 127, 143, 163, 164, 239, 243, 244, 245, 272, 295, 296, 297, 298, 299, ,300, 301, 321, 327, 328, 329, 340, 385, 413, 414, 415, 506, 537, 542, 557, 572, 598, 600, 649 Hinsdale: 11, 23, 52, 53, 56, 69, 139, 148, 249, 534, 611 Huerfano: 291, 421, 550, 594, 595, 596, 650, 651, 652 Jackson: 2, 74, 75, 77, 78, 209, 213, 228, 229, 2,30, 231, 232, 239, 253, 254, 355, 485, 532, 537, 562, 6,32 Jefferson: 63, 64, 65, 94, 95, 189, 234, 239, 264, 273, 351, 404, 452, 594, 595, 596, 610 Kiowa: 125 Kit Carson: 100, 399, 573 Lake: 215, 216, 359 La Plata: 18, 136, 201, 242, 336, 337, 379, 542, 558, 559, 648 Larimer: 2, 13, 24, 25, 31, 34, 35, 36, 37, 38, 41, 42, 43, 44, 52, 53, 56, 57, 68, 83, 84, 92, 93, 98, 104, 127, 145, 146, 168, 172, 175, 182, 183, 184, 185, 187, 188, 194, 195, 196, 197, 198, 199, 200, 202, 206, 207, 209, 212, 215, 216, 218, 219, 220, 221, 222, 228, 229, 230, 231, 232, 239, 275, 276, 277, 278, 304, 305, 306, 322, 323, 335, 356, 362, 395, 396, 397, 417, 418, 419, 420, 421, 422, 429, 437, 447, 461, 463, 465, 473, 488, 489, 507, 529, 535, 537, 538, 539, 541, 545, 546, 547, 581, 594, 595, ,596, 601, 602, 603, 616, 618, 6,33, 634, 635, 636, 637, 638, 639, 640, 641, 650, 651, 652, 654, 657, 660 Las Animas: 311, 312, 486, 487, 594, 595, 596, 619, 620, 621 Lincoln: 125 Logan: .36, 89, 90, 91, 116, 117, 118, 119, 120, ,364, 505 Mesa: 66, 70, 86, 147, 191, 208, 210, 266, 324, 507, 537, 542, 557, 577, 578, 588, 589 Mineral: 136, 1,39, 542, 548 Moffat: 60, 103, 121, 1,32, 177, 208, 213, 239, 352, ,353, 357, ,358, 659 Montezuma: 16, 19, 134, 150, 151, 152, 153, 355, 426, 427, 428, 542, 617 Montrose: 50, 70, 71, 86, 163, 164, 208, 295, 355, 413, 414, 415, 467, 468, 542, 572, 600, 649 Morgan: 36, 89, 90, 91, 125, 162, 236, 237, 238, 312, 313, 364, 505 Otero:309, 310, 311,312, 316 Ouray: 70 Park: 234, 291, 400, 401, 402, 474, 515, 522, 523, 537, 560, 583, 584, 585, 599, 616 Phillips: no references Pitkin: 52, 53, 56, 70, 210, 239, 247, 557 Prowers: 309, 310, 311, 312, 316 Pueblo: 160, ,309, 310, 311, 312, 316, 594, 595, 596 Rio Blanco: 58, 59, 82, 158, 177, 204, 208, 210, 213, 239, 253, 254, 268, 271b, 281, 282, 352, ,353, 355, 373, 374, 375, 480, 507, 556, 563, 570, 571, 597, 604, 605, 623 Rio Grande: 52, 53, 56, 138, 139, 193, 450, 457, 458 Routt: 77, 78, 79, 80, 213, 239, 248, 355, 553, 556, 632 Saguache: 23, 32, 127, 138, 139, 193, 291, 339, 450, 457, 458, 522, 650, 651, 652 San Juan: 11, 148, 3,36, ,337, 372, 534, 537, 611 San Miguel: 86, 208, 355, 371, 507, 542 Sedgewick: 89, 90, 91, 312, 313, 364, 399, 505, 573 Summit: 49, 52, 53, 56, 123, 167, 173, 174, 209, 234, 239, 285, 359, 360, 400, 401, 402, 537 Teller: 171, 234, 255, 256, 257, 258, 259, 260, 261, 361, 435, 509, 510, 530, 560, 616, 631 Washington: 124, 154, 180, 491, 503, 504, 519, 527 Weld: 15, 36, 54, 55, 84, 89, 90, 91, 105, 162, 166, 176, 223, 224, 236, 237, 238, 241, 283, 284, 312, 313, 314, 330, 331, 364, 366, 378, 383, 384, 399, 405, 447, 454, 455, 456, 471, 472, 477, 505, 521, 527, 555, 592, 593, 594, 653 Yuma: 36, 125, 154, 389, 390, 399 Statewide Studies: 1, 3, 4, 7, 12, 88, 102, 106, 107, 109, 128, 129, 135, 142, 149, 170, 178, 179, 181, 214, 262, 294, 326, 365, 376, 380, 387, 388, 424, 438, 439, 445, 447, 449, 481, 482, 492, 493, 494, 495, 496, 497, 498, 499, 500, 501, 502, 512, 520, 531, 540, 565, 566, 586, 587, 607, 613, 614, 615 Regional Studies: Plains: 39, 87, 110, 115, 205, 235, 274, 433, 454, 549, 608, 609 Mountains: 5, 6, 17, 19, 26, 28, 29, 76, 96, 108, 122, 139, 211, 227, 349, 386, 412, 416, 433, 580 Western Colorado Plateaus: 225, 226, 416, 480, 622, 658 Bibliography 1. Aldous, A. E., and H. L. Shantz. 1924. Types of vegetation in the semiarid portion of the United States and their economic significance. J. Agr. Res. 28: 99-128. January 1983 Baker: Colorado Bibliography 47 2. Alexander, M. E. 1980. Four fire scar records on lodgepole pine (Pinus contorta Dougl.) in north-central Colorado. S. W. Natur. 25:432-434. 3. Alexander, M. E., and D. V. Sandberg. 1976. Fire ecology and historical fire occurrence in the forest and range ecosystems of Colorado: a bibliography. Colo. St. Univ., Dept. of For. and Wood Sci., Occas. Report. 17 pp. 4. Alexander, M. E., and D. V. Sandberg. 1979. Fire ecology and historical fire occurrence in the forest and range ecosystems of Colorado: a bibliography, supple- ment No. 1. Colo. St. Univ., Dept. of For. and Wood Sci., Occas. Report. 4 pp. 5. Alexander, R. R. 1974a. Silviculture of central and southern Rocky Mountain forests: a summary of the sta- tus of our knowledge by timber types. USDA For. Serv. Res. Paper RM-120, Rocky Mt. For. and Range Expt. Stn., Fort Collins, Colo. 36 pp. 6. Alexander, R. R. 1974b. Silviculture of subalpine forests in the central and southern Rocky Mountains: the status of our knowledge. USDA For. Serv. Res. Paper RM-121, Rocky Mt. For. and Range Expt. Stn., Fort Collins, Colo. 88 pp. 7. Allison, E. M. 1908. Bibliography and history of Colorado botany. Univ. of Colo. Studies 6: 51-76. 8. Amundsen, C. C. 1967. The subalpine forest of Wild Basin, Front Range, Colorado. Unpubl. dis- sertation, Univ. of Colo., Boulder. 129 pp. 9. Andersen, D. C, R. S. Hoffman, and K. B. Armi- tage. 1979. Aboveground productivity and floristic struc- ture of a high subalpine herbaceous meadow. Arct. and Alp. Res. 11:467-476. 10. Anderson, J. M., Jr. 1950. A flora of Baca County, Colorado. Unpubl. thesis, Univ. of Colo., Boulder. 39 pp. 11. Andrews, J. T., P. E. Carrara, F. B. King, and R. Stuckenrath. 1975. Holocene environmental changes in the alpine zone, northern San Juan Mountains, Colo- rado: evidence from bog stratigraphy and palynology. Quat. Res. 5:173-197. 12. Armstrong, D. M. 1972. Distribution of Mammals in Colorado. Mus. Nat. Hist., Univ. of Kansas, Monogr. No. 3. [Vegetation: p. 26-31]. 13. Ashton, R. E. 1930. Preliminary observations on revegetation of the Twinsisters Bum in Rocky Mountain National Park. J. Colo.-Wyom. Acad. Sci. 1(2): 13 (Abstract). 14. Bachrach, J. H. 1951. The plant communities of a valley in the montane forest of the Colorado Front Range. Unpubl. thesis, Univ. of Colo., Boulder. 68 pp. 15. Badaracco, R. J. 1971. An interpretive resource analysis of Pawnee Buttes, Colorado. Unpubl. dis- sertation, Colo. St. Univ., Fort Collins. 341 pp. 16. Bader, E. H. 1932. The vegetation of the Mesa Verde National Park, Colorado. Unpubl. thesis, Univ. of Colo., Boulder. 64 pp. 17. Bailey, D. K. 1970. Phytogeography and taxonomy of Pinus subsection Balfourianae. Ann. Missouri Bot. Card. 57:210-249. 18. Baker, C. F. 1898. Botanizing in the La Plata Mountains. Plant World 2:29-32. 19. Baker, F. S. 1925. Aspen in the central Rocky Mountain region. USDA Bull. 1291. 46 pp. 20. Barber, R. A. 1935. An ecological study of a small, montane, spring-fed, stream area in the University of Colorado camp, on Mount Niwot. Unpubl. thesis, Univ. of Colo., Boulder. 21. Barclay, H. G. 1941a. Plant communities of the Gothic area and their successional relations. Bull. Ecol. Soc. Am. 22:34 (Abstract). 22. Barclay, H. G. 1941b. A subalpine climax grass- land in west central Colorado. Bull. Ecol. Soc. Am. 22:35 (Abstract). 23. Barren, J. 1969. Flora of the Gunnison Basin: Gunnison, Saguache, and Hinsdale counties. Natur. Land Inst., Rockford, 111. 24. Barrows, J. S., E. W. Mogren, K. Rowdabaugh, and R. Yancik. 1977. The role of fire in ponderosa pine and mixed conifer ecosystems. Final report, Co-op Agreem. with U.S. Nat. Park Serv. (CX-1200-7-8011) and Rocky Mt. For. and Range Exp. Stn. (16-542-CA), on file at Rocky Mt. For. and Range Expt. Stn., Fort Collins, Colo. 101 pp. 25. Barth, R. C. 1970. Revegetation after a subalpine wildfire. Unpubl. thesis, Colo. St. Univ., Fort Collins. 142 pp. 26. Bates, C. G. 1917. Forest succession in the central Rocky Mountains. J. For. 15:587-592. 27. Bates, C. G. 1923. The transect of a mountain val- ley. Ecology 4: 54-62. 28. Bates, C. G. 1924. Forest types in the central Rocky Mountains as affected by climate and soil. USDA Bull. No. 1233. 152 pp. 29. Bates, C. G., F. B. Notestein, and P. Keplinger. 1914. Climatic characteristics of forest types in the cen- tral Rocky Mountains. Proc. Soc. Am. For. 9:78-94. 30. Bathke, D. M. 1968. The flora of the subalpine and alpine zones in Robinson Basin, Colorado. Unpubl. thesis. Western St. College, Gunnison, Colo. 92 pp. 31. Baumann, T. G. 1978. Winter ecology of bighorn sheep in the Mummy Range, Colorado. Unpubl. thesis, Colo. St. Univ., Fort Collins. 32. Beck, R. F. 1965. Pocket gopher habitat relations in mountain rangelands in Colorado. Unpubl. thesis, Colo. St. Univ., Fort Collins. 61 pp. 33. Beidleman, R. G. 1948. The vertebrate ecology of a Colorado plains cottonwood river bottom. Unpubl. thesis, Univ. of Colo., Boulder. 34. Beidleman, R. G. 1952. The ponderosa-grassland ecotone in north central Colorado. J. Colo.-Wyom. Acad. Sci. 4(4):70 (Abstract). 35. Beidleman, R. G. 1953. The island of pines. Liv- ing Wilderness 46:7-10. 36. Beidleman, R. G. 1954. The cottonwood river-bot- tom community as a vertebrate habitat. Unpubl. dis- sertation, Univ. of Colo., Boulder. 358 pp. 37. Beidleman, R. G. 1957. Fire bum succession stud- ies in the northern Colorado Rockies. J. Colo.-Wyom. Acad. Sci. 4(9):39 (Abstract). 38. Beidleman, R. G. 1967. Twenty-year successional studies in northern Colorado. J. Colo.-Wyom. Acad. Sci. 5(8):63-64 (Abstract). 39. Beidleman, R. G. 1978. The cottonwood-willow riparian ecosystem as a vertebrate habitat, with particu- lar reference to birds. Pages 192-195 in W. D. Graul and S. J. Bissell, eds.. Lowland river and stream habitat in Colorado: a symposium. (Greeley, Colo. 4-5 Oct. 1978). Colo. Chap. Wildl. Soc. & Colo. Audubon Council. 40. Beidleman, R. G., and W. Fischer. 1979. Environ- mental reconnaissance study of Perry Park Ranch, Douglas County, Colorado. Wright-Ingram Institute, Colorado Springs, Colo. 48 Great Basin Naturalist Vol. 43, No. 1 41. Bell, K. L. 1974a. Autecology of Kobresia bel- lardii: why winter snow accumulation patterns affect lo- cal distribution. Unpubl. dissertation, Univ. of Alberta, Edmonton. 167 pp. 42. Bell, K. L. 1974b. Autumn, winter, and spring phenology of some Colorado alpine plants. Am. Midi. Natur. 91:460-464. 43. Bell, K. L. 1977. Autecology of Kobresia beUardii: why winter snow accumulation affects local distribution. Bull. Ecol. Soc. Am. 58(2):41 (Abstract). 44. Bell, K. L., and L. C. Bliss. 1979. Autecology of Kobresia bellardii: why winter snow accumulation limits local distribution. Ecol. Monogr. 49: 377-402. 45. Benedict, J. B. 1977. A master plan for the Bunker Hill Placer Preserve, Boulder County, Colorado. Un- publ. report on file at The Nature Conservancy, Denver, Colo. 61 pp. 46. Bichel, M. A. 1959. Investigations of a Nebraska and a Colorado prairie and their impact on the relict concept. Unpubl. dissertation, Univ. of Nebraska, Lin- coln. 180 pp. 47. Bierly, K. F. 1972. Meadow and fen vegetation in Big Meadows, Rocky Mountain National Park. Unpubl. thesis, Colo. St. Univ., Fort Collins, Colo. 102 pp. 48. Biggins, J. and M. H. Dodson. 1970. Marshall Mesa natural area study. Unpubl. report, Dept. Geogr., Univ. of Colo., Boulder. 54 pp. 49. Bollinger, W. H. 1973. The vegetation patterns af- ter fire at the alpine forest-tundra ecotone in the Colo- rado Front Range. Unpubl. dissertation, Univ. of Colo., Boulder. 2 vols. 50. Bond, H. L. 1959. Revegetation and disintegration of pocket gopher mounds on Black Mesa, Colorado. Un- publ. thesis, Colo. St. Univ., Fort Collins. 51. Bonde, E. K. 1967. Pinus aristata (Pinaceae) in Boulder County, Colorado: northward extension of its known Colorado range. S.W. Natur. 12:474-475. 52. Bonham, C. D. 1966. An ordination of alpine hair- grass (Deschampsia caespitosa Beauv.) meadows. Un- publ. dissertation, Colo. St. Univ., Fort Collins. 124 pp. 53. Bonham, C. D. 1972. Vegetation analysis of grazed and ungrazed alpine hairgrass meadows. J. Range Mgmt. 25: 276-279. 54. Bonham, C. D., and J. S. Hannan. 1978. Blue grama and buffalograss patterns in and near a prairie dog town. J. Range Mgmt. 31:63-65. 55. Bonham, C. D., and A. Lerwick. 1976. Vegetation changes induced by prairie dogs on shortgrass range. J. Range Mgmt. 29: 221-225. 56. Bonham, C. D., and R. T. Ward. 1970. Phytoso- ciological relationships in alpine tufted hairgrass {Des- chanvpsia caespitosa (L.) Beauv.) meadows. Arct. and Alp. Res. 2:267-275. 57. Bowes, A. L. 1958. Ecological analysis of mule deer winter range. Cache la Poudre canyon, Colorado. Unpubl. thesis, Colo. St. Univ., Fort Collins. 58. Boyce, D. A. 1976. A preliminary description of the ecosystems adjacent to the South Fork of the White River, Rio Blanco, and Garfield counties, Colorado. J. Colo.-Wyom. Acad. Sci. 8(1):45 (Abstract). 59. Boyce, D. A. 1977. Vegetation of the South Fork of the White River Valley, Colorado. Unpubl. dis- sertation, Univ. of Colo., Boulder. 60. Bradley, R. A. W. 1950. The vascular flora of Mof- fat County, Colorado. Unpubl. thesis, Univ. of Colo., Boulder. 71 pp. 61. Brandegee, T. S. 1878. Notes from Colorado. Bot. Gaz. 3:166. 62. Brandegee, T. S. 1880. Timber line in the Sawatch Range. Bot. Gaz. 5:125-126. 63. Branson, F. A., R. F. Miller, and I. S. McQueen. 1961. Soil-water availability and use by grasslands on ad- jacent stony and shale-derived soils in Colorado. Short papers in the geologic and hydrologic sciences. USGS Prof. Paper 424C:251-253. 64. Branson, F. A., R. F. Miller, and I. S. McQueen. 1964. Effects of two kinds of geologic materials on plant communities and soil moisture. Am. Soc. Agron. Spec. Publ. No. 5:165-175. 65. Branson, F. A., R. F. Miller, and I. S. McQueen. 1965. Plant communities and soil moisture relationships near Denver, Colorado. Ecology 46:311-319. 66. Branson, F. A., R. F. Miller, and L S. McQueen. 1976. Moisture relationships in twelve northern desert shrub communities near Grand Junction, Colorado. Ecology 57:1104-1124. 67. Branson, F. A., and J. B. Owen. 1970. Plant cover, runoff, and sediment yield relationships on Mancos shale in western Colorado. Water Resources Res. 6:783-790. 68. Braun, C. E. 1969. Population dynamics, habitat, and movements of white-tailed ptarmigan in Colorado. Unpubl. dissertation, Colo. St. Univ., Fort Collins. 69. Brewster, J. F. 1968. Ecological factors in the revegetation of SlumguUion Mudflow near Lake City, Colorado. Unpubl. thesis, Univ. of Illinois, Urbana. 94 pp. 70. Brown, H. E. 1958. Gambel oak in west-central Colorado. Ecology 39:317-327. 71. Brown, H. E., and J. R. Thompson. 1965. Summer water use by aspen, spruce, and grassland in western Colorado. J. For. 63:756-760. 72. Bruderlin, K. 1911. A study of the lodgepole-pine forests of Boulder Park (Tolland, Colorado). Univ. of Colo. Studies 8:265-275. 73. Buck, P. 1960. Vegetational succession in sub- alpine ponds in the Rockies. Proc. Okla. Acad. Sci. 40:2-6. 74. Buckner, D. L. 1976. Occurrence and mainte- nance of ribbon forest in the Colorado Rocky Moun- tains. J. Colo.-Wyom. Acad. Sci. 8(1):46 (Abstract). 75. Buckner, D. L. 1977. Ribbon forest development and maintenance in the central Rocky Mountains of Colorado. Unpubl. dissertation, Univ. of Colo., Boulder. 224 pp. 76. Bujakiewicz, A. 1975. Vegetational zonation in the Front Range of the Rocky Mountains, Colorado. USA [in Polish] Frag. Florist, et Geobot. 21:99-142. 77. Bunin, J. E. 1974. The vegetation of the west slope of the Park Range, Colorado. J. Colo.-Wyom. Acad. Sci. 7(5):30 (Abstract). 78. Bunin, J. E. 1975a. The vegetation of the west slope of the Park Range, Colorado. Unpubl. dissertation, Univ. of Colo., Boulder. 2 pp. 79. Bunin, J. E. 1975b. Aspen forests of the west slope of the Park Range, north-central Colorado. Paper pre- sented at the 1975 AIBS/ESA Meetings, 17-22 Aug. 1975, Oregon St. Univ., Corvallis. 22 pp. 80. Bunin, J. E. 1975c. Aspen forests of the west slope of the Park Range, northcentral Colorado. Bull. Ecol. Soc. Am. 56(2):48 (Abstract). January 1983 Baker: Colorado Bibliography 49 81. Bums, S. F. 1980. A comparison of mountain life zones of the Swiss Alps with the Colorado Front Range. J. Colo.-Wyom. Acad. Sci. 12(1):31 (Abstract). 82. Butler, J. R. and J. L. England. 1979. 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Ecological characteristics of pin- yon-juniper woodlands on the Colorado Plateau— a liter- ature survey. USDI, Bur. of Land Mgmt., Tech. Note No. 310. 183 pp. 659. Zimmerman, T. 1978. Skull Creek Study Area Forestry. Unpubl. report. Bureau of Land Management, Craig District Office, Craig, Colo. 62 pp. 660. Zuck, R. H. 1974. An ecological versus manage- rial classification of forest communities. Unpubl. thesis, Colo. St. Univ., Fort Collins. 110 pp. 'Acknowledgments I am grateful to the numerous people who provided access to unpublished reports, and to J. Scott Peterson, who provided support and criticism. The author would appreciate being informed if there are omissions. EVALUATION OF A PROGRAM TO CONTROL HYDATID DISEASE IN CENTRAL UTAH^ Ferron L. Andersen^, John R. Crellin^-^, Craig R. Nichols*, and Peter M. Schantz^ Abstract.— A program to control hydatid disease in central Utah was evaluated by: (1) surveillance of infection in dogs and sheep, (2) questioning adult residents of two Sanpete County communities (Fountain Green and Spring City) about their knowledge of hydatid disease and their attitudes toward preventive measures, (3) interviewing local officials to evaluate the proposed implementation of proper community-wide measures, (4) analyzing records of prophylactic treatment of dogs with praziquantel, and (5) comparing scores of tests given to third and fourth grade students before and after they colored an educational pamphlet about hydatid disease. Infection rates of Echinococcus granulosus in dogs brought to volunteer diagnostic clinics dropped from 28.3 per- cent in 1972 to 1.0 percent in 1979, but increased to 9.8 percent in 1981. This last rise was due mainly to the fact that some dogs were examined that belonged to range sheepmen who had either not attended a field clinic recently or had never attended at all. Prevalence of the parasite in slaughtered sheep decreased steadily from 13.2 percent in 1972 to 2.8 percent in 1981. With regard to the questionnaire phase of the project, 87.3 percent and 84.3 percent of the respondents understood the role of dogs in the life cycle of E. granulosus in Fountain Green and Spring City, re- spectively. Over 50 percent of the respondents of these two communities had worked directly with sheep sometime in their life. In general, residents were more willing to practice preventive measures involving sheep than they were to implement measures involving dogs alone. However, many of the recommended community-wide preventive mea- sures were not implemented. With the coloring book, students answered an average of 62.5 percent of the questions correctly before they colored the pamphlet and 83.3 percent afterward. Overall, our results suggest that residents of Sanpete County are knowledgeable about hydatid disease and its mode of transmission, and that, in general, progress has been made in control of hydatid disease in central Utah. Utah has had the most autochthonous cases of hydatidosis (39) in man reported from the contiguous United States (Crellin et al., 1982), with the first reported case diagnosed in 1944 (Carlquist and Dowell 1951). Studies on dogs, foxes, and coyotes were undertaken in the early 1950s to identify the definitive host, but these were unsuccessful (Butler and Grundmarm 1951, Grundmann et al. 1953, Butler and Grundmann 1954). The normal hosts of Echinococcus granulosus in Utah (dogs and sheep) were revealed in 1969 as part of an investigation into the death of a nine-year old boy in the community of Herri- man (Kahn et al. 1972), near Salt Lake City. Concurrently, parasitologists at Brigham Young University began surveillance of the parasite in dogs and sheep in central Utah (Fox et al. 1970, Andersen et al. 1973, Love- less et al. 1978). As more work was done, it became apparent that the main foci of in- fection were in that area of Utah, especially Sanpete County. As a result, a cooperative program to study and control hydatid disease in central Utah was begun in 1971 by per- sonnel from Brigham Young University (Pro- vo, Utah), the Utah Department of Health (Salt Lake City, Utah), and the Centers for Disease Control (Atlanta, Georgia) (Andersen et al. 1974). Materials and Methods Description of Study Area Sanpete County is in the center of Utah and has a total area of 4,136 sq km. A valley (1700 m elevation) bordered by mountains (3400 m elevation) runs the length of the county. There are 14,615 people in the coun- ty (U.S. Bureau of the Census 1980a), 90 per- cent of which are Mormons (The Church of Jesus Christ of Latter-day Saints) (Stinner et al. 1978). Unlike most farming regions in the 'This project was supported in part by U.S. Public Health Service Grant AI-10588. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 'Present address: Trinity College, Deerfield, Illinois 60015. •Bureau of Communicable Disease Control, Utah Department of Health, Salt Lake City, Utah 84103. 'Division of Parasitic Diseases, Center for Infectious Diseases, Centers for Disease Control, Atlanta, Georgia 30333. 65 66 Great Basin Naturalist Vol. 43, No. 1 United States, nearly all residents of Sanpete County, regardless of occupation, live in one of 13 communities which vary in population from 153 to 2807 (U.S. Bureau of the Census 1980a). In 1978 there were approximately 1950 dogs (Loveless et al. 1978) and 90,400 sheep in the county (U.S. Bureau of the Cen- sus 1980b). Agriculture is the principal in- dustry, with an emphasis on turkeys and sheep. Range sheep herds are grazed in the mountains during summer, on the valley floor during spring and fall, and on the desert (100-200 km to the west) during winter. In addition to the range herds, there are also semirange herds that are confined during winter but grazed in the mountains during summer. There are also many small flocks confined on a year-round basis, which are usually made up of ewes obtained from range sheepmen (Crellin et al., 1982). Description of Hydatid Disease Control Program The main goals of the program have been reported previously (Andersen et al. 1974) and are summarized on Table 1. The educa- tional portion of the program was accom- plished through press releases, an article in the National Woolgrower (Andersen and Wal- lentine 1976), pamphlets distributed county- wide, talks to civic and church groups, and displays and personal coimseling done during community screening clinics. A filmstrip and coloring book were developed as tools to educate children. Copies of the filmstrip were given to the audiovisual departments of Table 1. Main goals of a program to control hydatid disease in Sanpete County, Utah. Community objectives 1. Covering or fencing animal disposal pits at waste dis- posal sites 2. Eliminating stray or roving dogs 3. Conducting educational programs in local schools Individual objectives 1. Fostering a widespread understanding of the para- site's life cycle 2. Advocating proper preventive measures such as not feeding sheep viscera to dogs, burning or burying sheep that die on the range, and not allowing dogs to roam 3. Encouraging individuals to have themselves and their dogs checked for £. granulosus at yearly screening clinics the two school districts in Sanpete County, and in 1980 coloring books were distributed to all third and fourth grade students in eight elementary schools in the county. The overall goal of this portion of the program was to educate the residents about the life cycle of E. granulosus, and the proper measures that could be taken to prevent the disease. Local governments were advised as to proper community-wide control measures, and it was their choice whether or not such measures were implemented. Included within this phase of the program was prophylactic treatment by a local veterinarian (Dr. M. John Ramsay) of dogs in the northern half of Sanpete County with bunamidine hydro- chloride (1974-1978) and praziquantel (1979-1981). Surveillance of dogs, sheep, and human beings was conducted in several areas of cen- tral Utah. Starting in 1971, clinics to examine dogs were held in communities in Sanpete County, and, in 1980 and 1981, on nearby mountain range lands (Table 2). In other areas (Summit and Utah counties), clinics were usually held at the sheep camp location. Dogs were dosed with arecoline hydro- bromide, and the resulting purge was exam- ined on site. Sheep were checked at slaughter for the presence of cysts by state meat in- spectors, and the identification of cysts sus- pected was confirmed at the parasitology laboratory at Brigham Young University. For- ty-nine coyotes (Andersen et al. 1973, Conder and Loveless 1978) and 74 deer (Jensen et al., 1982b) were also examined to assess their possible role as sylvatic reservoirs of in- fection. Immunodiagnostic clinics for human beings were conducted usually in conjunction with clinics for dogs (Klock et al. 1973, Bar- bour et al. 1978). Clinics were advertised in local newspapers, by posters placed in com- mimities, and by letters and phone calls to sheepmen by individuals from the Utah De- partment of Health, BYU, and a local veter- inarian's office. Evaluation of Control Program The effectiveness of the control measures was evaluated by monitoring trends in in- fection rates in dogs and sheep and the num- bers of new cases diagnosed in human beings. January 1983 Andersen et al.: Hydatid Disease 67 The success of holding screening clinics was evaluated partially by comparing the list of all those who brought dogs for examination with a complete list of all sheepmen in the region (Crellin et al., 1982). Further eval- uation was obtained by questioning adult residents of two Sanpete Coimty commu- nities (Fountain Green and Spring City) con- cerning their knowledge of hydatid disease and attitudes towards proper preventive measures. The procedures employed in devis- ing, distributing, and analyzing these ques- tionnaires were described earlier (Condie et al., 1981). Implementation of community- wide measures was evaluated through inter- views with city officials and visits to the vari- ous community waste disposal sites. The effectiveness of the coloring books in increasing knowledge about hydatid disease and proper preventive measures was ana- lyzed by testing the students before they re- ceived the coloring book and again two weeks afterward. Pre- and posttest scores were compared using a paired t-tesi. Results Of 15,775 sheep slaughtered in five central Utah abattoirs since 1971, 1116 (7.1 percent) were infected with E. granulosus. The range in yearly prevalence was 13.2 percent in 1972 to 2.8 percent in 1980 and 1981 (Fig. 1). During the study period, 83 screening clinics for dogs were held; 109 of 1120 (9.7 percent) dogs purged were infected with the adult cestode. Infection rates of the parasite in dogs declined from 28.3 percent in 1972 (Loveless et al. 1978) to 1.0 percent in 1979, but rose to 8.9 percent in 1980 and to 9.8 percent in 1981 (Fig. 1) (Jensen et al. 1982a). This rise, however, was probably due to the fact that in 1980 7 of the 8 dogs found in- fected were owned by sheepmen who had never attended a clinic, and that in 1981 4 of the 5 dogs infected were owned by sheepmen who had not attended a clinic in several years. Sixty of 593 (10.1 percent) owners who attended a clinic had at least one infected dog (Fig. 2 and Table 3). Generally, the pro- portion of owners with at least one infected dog was higher in communities from the northern half of the county (Table 3). Twenty-seven of the 28 (96.4 percent) range sheepmen presently in the county have taken some of their dogs to at least one clinic, but no more than 19 (67.8 percent) have attended in any one year. Seven clinics were held in Summit County where 6 of 69 (8.7 percent) dogs harbored E. granulosus, and one clinic was held in Utah County where 1 of 21 (4.8 percent) dogs was infected. No echinococ- cosis infections were found among 49 coyotes (Andersen et al. 1973, Conder and Loveless 1978) and 74 deer examined (Jensen et al. 1982b). Table 2. Dogs infected with Echinococcus granulosus in Sanpete County, 1971-1981. Use of dog Community North Sanpete Fairview Fountain Green Mt. Pleasant Spring City Other communities Area total South Sanpete Ephraim Manti Mayfield Gunnison Area total County total Tending sheep House pet Dogs Dogs Dogs Dogs purged infected (%) purged infected (%) 70 14(20.0) 91 4(4.4) 153 32(21.7) 67 3(4.5) 92 5(5.4) 31 2(6.5) 81 16(19.8) 161 12(7.5) 55 4(7.3) 33 0(0.0) 451 72(16.0) 383 21(5.5) 56 8(14.3) 21 0(0.0) 43 4(9.3) 96 1(1.0) 2 0(0.0) 37 0(0.0) 11 3(27.3) 20 0(0.0) 112 15(13.4) 174 1(0.6) 563 87(15.4) 557 22(3.9) 68 Table 3. Dog owners in Great Basin Naturalist Sanpete County who have had dogs infected with Echinococcus grant Vol. 43, No. 1 ilosus, 1971-1981. Rang e sheepmen Semirange sheepmen Not sheepmen Region No. of owners No. with infected dogs (%) No. of owners No. with infected dogs (%) No. of owners No. with infected dogs (%) North South 25 19 16(64.0) 7(36.8) 67 12 15(22.4) 1(8.3) 285 180 21(7.4) 1(0.6) Total 44 23(52.3) 79 16(20.3) 465 22(4.7) Fourteen autochthonous cases of hydatid disease in man have been diagnosed in San- pete County— an average annual incidence of 3.7 per 100,000 (Table 4). Twelve of these 14 cases were from the northern half of the county. The 7 cases in Fountain Green since 1952 translate to a prevalence of 50 per 100,000. Six of 2747 individuals in Sanpete County tested serologically for the presence of hydatid cysts were diagnosed to be asymp- tomatic carriers, and subsequently 4 of these 6 have had surgery (Barbour et al. 1978). In the door-to-door surveys, responses were obtained from 140 of 156 (89.7 percent) and 147 of 176 (83.5 percent) households in Fountain Green and Spring City, respective- ly. In Fountain Green 256 individuals (1980 population of 578), and in Spring City 228 (1980 population of 675) returned completed questionnaires. Twenty-nine percent of the respondents from Fountain Green indicated that they were aware of hydatid disease be- fore 1971, but only 9.0 percent from Spring City were aware of this disease prior to that year. In Fountain Green, 87.3 percent of the respondents currently understood the role of dogs in the life cycle of E. granulosus, and 70.0 percent knew the role of sheep. In Spring City 84.3 percent imderstood the role of dogs and 69.0 percent knew the role of sheep. There were 62.0 percent and 54.0 per- cent of the adult residents of Fountain Green and Spring City, respectively, who had worked with sheep some time in their life. In Fountain Green 83.0 percent of the adult residents and in Spring City 77.5 percent were willing to pay to have a proper animal disposal pit built and maintained. Spring City residents were asked the reasons why they did or did not bring their dogs to clinics. The 35 r Q 30 LU 1- o 2b LU LL Z 20 1- z 15 LU O cc 10 LU CL DOGS SHEEP — • \ / 1971 1973 1975 1977 1979 1981 YEAR Fig. 1. Prevalence oi Echinococcus granulosus in dogs and sheep from central Utah, 1971-1981. January 1983 Andersen et al. : Hydatid Disease 69 Table 4 . Fourteen autochthonous cases of hydatid disease in human beings from Sanpete County, Utah. . Year of City of Connection with Year of Location Name Sex birth residence sheep raising surgery of cyst N.P. F 1944 Fountain Green Direct" 1952 Lung J.B. M 1916 Fountain Green Direct 1964 Lung & Liver P.A. M 1946 Fountain Green Direct 1967 Liver PJ. M 1931 Fountain Green Direct 1971 Liver L.C. M 1949 Fountain Green Direct 1971 Liver M.C. M 1908 Mt. Pleasant Direct 1972 Lung R.L. M 1908 Mt. Pleasant Direct 1973 Lung D.J. M 1960 Fountain Green Indirect^ 1974 Liver R.C. M 1913 Spring City Direct 1975 Liver D.A. M 1921 Fountain Green Direct 1975 Liver B.S. F 1967 Manti Indirect 1976 Lung D.D. F 1955 Gunnison Direct None= Liver C.W. F 1926 Spring City Direct None^ Liver E.G. M 1951 Fairview Indirect 1979 Lung "Patient or family members are sheepmen. "Patient lives near sheepmen, tase diagnosed in 1976. desire to know if their dog was infected (93.9 percent) and their desire to eUminate hydatid disease (90.3 percent) were the reasons most commonly given for attending. Being un- aware of the cHnic (55.9 percent) and a lack of personal time (35.2 percent) were the rea- sons most frequently listed for not attending. Data on sources of information (Table 5), at- titudes and practices of dog and sheep own- ers (Table 6), and dog and sheep populations (Table 7) are also presented herein. Implementation of community-wide pre- ventive measures and participation in the praziquantel treatment program were gener- • RANGE SHEEPMEN A SEMI-RANGE SHEEPMEN □ NOT SHEEPMEN 1971 Fig. 2. Percent of individuals in Sanpete County, Utah, having dogs infected with Echinococcus granubsus, 1971-1981. 70 Great Basin Naturalist Vol. 43, No. 1 ally poor. Only one community (Spring City) of the six that had an animal disposal pit, had a pit cover. Two commimities (Moroni and Manti) did not allow dead animals to be left at their waste disposal sites. Laws making it illegal to leave dead animals on the range were passed recently in Sanpete and five ad- joining counties, but, at this time, enforce- ment is nonexistent. All communities have leash laws, but only one (Manti) has made an effort to eliminate stray dogs. Fairview, Fountain Green, Moroni, and Mt. Pleasant have enacted laws requiring treatment of dogs with praziquantel as a stipulation for obtaining or renewing a dog license; how- ever, few sheepdogs in Sanpete County are licensed and compliance is largely voluntary. In the praziquantel treatment program, only 7 of 63 (11.1 percent) owners participating had ever had dogs known to be infected. Only 22 of the 63 (34.9 percent) owners had ever brought their dogs to a screening clinic. Discussion Among the human population at risk in Sanpete County, cystic hydatid disease has been diagnosed at a rate (3.7 cases per 100,000 population per year) comparable to those in other regions regarded as highly en- demic for this infection. Data for comparison include rates from the Australian state of Tasmania prior to the initiation of a state- wide control program (15 per 100,000 per year; McConnell and Green 1979); from Cyprus (12.9 per 100,000 per year; Anony- mous 1981); from Chile (7.8 per 100,000 per Table 5. Helpfulness of various sources of informa- tion about hydatid disease in Fountain Green and Spring City, Utah. year; Anonymous 1981); and from Yugoslavia (3.7 per 100,000 per year; Anonymous 1981). Also, infection rates in dogs and sheep at the beginning of the study period herein re- viewed were similar to those in other endem- ic regions (Anonymous 1981). Most successful programs to control E. granulosus have had both a formal organiza- tional structure and a mandatory in- volvement of the local population (Gemmell 1979). In contrast, the control effort in cen- tral Utah has been a cooperative project in- volving university, state, and federal agencies, and participation of the community has been completely voluntary. The volun- tary aspect of the program has imposed cer- tain limitations on the rate of progress of sug- gested preventive and control measures. Although the majority of dog owners and sheep ranchers have responded positively to recommended measures, and also availed themselves of the diagnostic services pro- vided, our questionnaire survey indicated that some individuals have not. The data in- dicate that the health educational aspects of the program have succeeded in making most adult residents of the two commimities thor- oughly aware of hydatid disease, but this may not be adequate to sustain interest and to motivate necessary changes. We had pre- viously observed that most dog owners in Sanpete County were able to respond cor- rectly to questions about the life cycle of Table 6. Attitudes and practices of dog and/or sheep owners in Fountain Green and Spring City, Utah. Attitude or practice Percentage of respondents Fountain Green Spring City (%) (%) Attitude— willing to: take dog to clinic" 83.5 78.2 Percentage of respondents who perceived source as helpful" pay tor propnyiacnc treatment of do^ control do^ 86.7 48.8 95.7 65.0 Source of information Fountain Green (%) Spring City (%) use animal pit Practice; allow dog to have access to area 88.0 87.8 Screening clinics Friends 70.8 60.2 56.3 47.8 Pamphlets Veterinarian 56.5 51.6 50.7 32.7 where sheep are killed^ 21.6 11.0 Newspapers Filmstrip Physician 23.5 16.0 12.9 36.7 14.4 13.5 butcher sheep at home or on range*" 63.3 90.6 'Individuals who marked a "4" or "5" on a 1 (no help) to 5 (very helpful) scale. *Dog owners: Fountain Green, 64, and Spring City, ( "Sheep owners: Fountain Green, 34, and Spring City, 27. January 1983 Andersen et al.: Hydatid Disease 71 hydatid disease and how to prevent it; how- ever, many continue to manage their dogs in ways that permit them ready access to sources of infection (Schantz and Andersen 1980). Another problem associated with the vol- untary nature of our program involves the in- terpretation of data collected at diagnostic field clinics for dogs. Generally, rates of in- fection measured annually suggested a down- ward trend, although rates in most recent years have increased somewhat (Fig. 1). This apparent increase most likely reflects differ- ences in the yearly samples of the canine population, rather than actual increases in rates of transmission. Some effort was made in the later years to encourage dog owners who had not previously attended to bring their animals to the voluntary clinics. Also, more diagnostic clinics were held in the sum- mer sheep range land, rather than in local communities as was done previously. Thus, higher rates of infection were found in dogs not previously examined and in those sheep dogs that were from high-risk canine popu- lations associated with herds of sheep. In contrast to these fluctuating changes noted in the infection rates of dogs, a more steady decline was noted for the slaughtered sheep, and yet no changes in sheep marketing practices were known to occur during the study period. It now seems likely to conclude that, after 10 years, E. granulosus has largely been elim- inated from "house" dogs (those dogs not working with sheep), but is still found in small numbers in dogs owned by the county's sheepmen. This partial control has probably been achieved through educational efforts that resulted in a reduction in availability of sheep viscera to dogs. However, recom- mended preventive measures such as control Table 7. Dog and sheep populations in Fountain Green and Spring City, Utah. Fountain Green Spring City Households with dogs 64/140 66/147 Number of dogs 109 101 Dogs per dog-owning household 1.7 1.5 Households with sheep 34/140 27/147 Number of sheep 24,317 3,655 Sheep per sheep- owning household 714 135 of dogs and proper disposal of dead animals have not been effectively implemented. The number of sheep in Sanpete County has dropped 38.5 percent since 1969 (U.S. Bu- reau of Census 1980b), and this could be a factor in the reduction in prevalence of E. granulosus. A drop in total sheep numbers was considered a factor in the elimination of hydatid disease from Iceland (Beard 1973). In spite of the progress noted in our con- trol program to date, there is still a continued potential for transmission of E. granulosus between dogs and human beings in Sanpete County. Basically, this is due to: (1) the lack of a county-wide control program for dogs, (2) the lack of adequately maintained animal disposal pits and covers in most of the com- munities, (3) the persistence of home-slaugh- tering of sheep, and (4) concentration of most county residents within communities. Our re- sults, and those of an earlier study (Condie et al. 1981), suggest that individuals in central Utah are generally unwilling to control their dogs, but would be supportive of community efforts to build and maintain proper disposal pits for dead sheep. In the future we plan to continue the dis- tribution of all educational aids available, and also to encourage the community offi- cials in those areas where hydatid disease has occurred to build and maintain proper ani- mal disposal pits and covers. In addition, most clinics will be held closer to summer grazing areas, and sheepmen with a poor at- tendance record at clinics will be encouraged to have their dogs checked on a regular basis. Table 8. Efficacy of coloring books as educational aids for third and fourth grade students in Sanpete County, Utah. Average Average pretest posttest No. score score School students (%) (%) Fairview Fountain Green Moroni Mt. Pleasant/ Spring City Ephraim Gunnison Manti 47 57.3 79.6° 6 51.7 83.3° 50 66.6 80.8° 107 60.7 78.4° 63 64.4 88.1° 84 66.8 87.5° 56 59.5 86.6° Total 413 62.5 83.3° 'Posttest score significantly greater than pretest score at a = 0.05 using paired t-test. 72 Great Basin Naturalist Vol. 43, No. 1 Finally, since praziquantel (Droncit®) is now an approved drug in the United States, and since dog owners within the two commu- nities surveyed indicated that they are will- ing to pay for treatment, we will encourage continued prophylactic treatment of dogs with this drug. Finally, surveillance of hyda- tid disease will be maintained by monitoring infection rates in slaughtered sheep and by registering diagnosed human cases. Literature Cited Andersen, F. L., P. D. Wright, and C. Mortenson. 1973. Prevalence of Echinococcus granulosus in- fection in dogs and sheep in central Utah. J. Am. Vet. Med. Assoc. 163:1168-1171. Andersen, F. L., J. R. Everett, A. G. Barbour, and F. J. Schoenfeld. 1974. Current studies on hydatid disease in Utah. Proc. 78th Ann. Meeting U.S. Animal Hlth. Assoc. 370-384. Andersen, F. L., and M. V. Wallentine. 1976. Hydatid disease. Nat. Woolgrower 66:16-18. Anonymous. 1981. FAO/UNEP/WHO guidelines on echinococcosis/hydatidosis surveillance, pre- vention and control. Wld. Hlth. Org., V.P.H. 80:23, Geneva. Barbour, A. G., J. R. Everett, F. L. Andersen, C. R. Nichols, T. Fukushima, and I. G. Kagan. 1978. Hydatid disease screening: Sanpete County, Utah, 1971-1976. Am. J. Trop. Med. Hyg. 27:94-100. Beard, T. C. 1973. The elimination of echinococcosis from Iceland. Bull. Wld. Hlth. Org. 48:653-660. Butler, J. M., and A. W. Grundmann. 1951. Fecal ex- amination of 200 dogs from Salt Lake City, Utah, for intestinal helminths. J. Am. Vet. Med. Assoc. 118:396-398. Butler, J. M., and A. W. Grundmann. 1954. The in- testinal helminths of the coyote Canis latrans Say, in Utah. J. Parasitol. 40:440-443. Carlquist, J. H., AND R. J. DowELL. 1951. Echino- coccus disease. Report of four cases contracted in the United States. Rocky Mtn. Med. J. 48:773-776. CoNDER, G. A., AND R. M. LovELESs. 1978. Parasites of the coyote {Canis latrans) in central Utah. J. Wildl. Dis. 14:247-249. CoNDiE, S. J., J. R. Crellin, F. L. Andersen, and P. M. Schantz. 1981. Participation in a community program to prevent hydatid disease. Publ. Hlth. Lond. 95:28-35. Crellin, J. R., F. L. Andersen, P. M. Schantz, and S. J. CoNDiE. 1982. Possible factors influencing dis- tribution and prevalence of Echinococcus gran- ulosus in Utah. Am. J. Epidemiol. 116:463-474. Fox, J. C, F. L. Andersen, and K. H. Hoopes. 1970. A survey of the helminth parasites of cattle and sheep in Utah valley. Great Basin Nat. 30: 131-145. Gemmell, M. a. 1979. Hydatidosis control— a global view. Aust. Vet. J. 55:118-125. Grundmann, A. W., J. M. Butler, and M. J. Selin. 1953. Examination of commercial silver fox as a possible host for Echinococcus granulosus (Batsch 1786) Rudolphi 1805 in Salt Lake County, Utah. J. Am. Vet. Med. Assoc. 123:235-236. Jensen, L. A., F. L. Andersen, and P. M. Schantz. 1982a. The prevalence of Echinococcus gran- ulosus and other taeniid cestodes in sheep dogs of central Utah. Great Basin Nat. 42:65-66. Jensen, L. A., J. A. Short, and F. L. Andersen. 1982b. The internal parasites of Odocoileus hemionus of central Utah. Proc. Helminthol. Soc. Washington 49:317-319. Kahn, J. B., S. Spruance, J. Harbottle, P. Cannon, and M. G. Schultz. 1972. Echinococcosis in Utah. Am. J. Trop. Med. Hyg. 21:185-188. Klock, L. E., S. L. Spruance, F. L. Andersen, D. D. JuRANEK, AND I. G. Kagan. 1973. Detection of asymptomatic hydatid disease by a community screening program. Am. J. Epidemiol. 97:16-21. Loveless, R. M., F. L. Andersen, M. J. Ramsay, and R. K. Hedelius. 1978. Echinococcus granulosus in dogs and sheep in central Utah, 1971-76. Am. J. Vet. Res. 39:499-502. McCoNNELL, J. D., AND R. J. Green. 1979. The control of hydatid disease in Tasmania. Aust. Vet. J. 55:140-145. Schantz, P. M., and F. L. Andersen. 1980. Dogs, own- ers and hydatid disease in Sanpete County. Gr. Bas. Nat. 40:216-220. Stinner, W. F., M. B. Toney, and S. Kan. 1978. Popu- lation changes in rural Utah. Utah Sci. 39:107-109. U.S. Bureau of the Census. 1980a. 7980 Census. Utah preliminary report. U.S. Bureau of the Census, Washington, D.C. 1980b. 1978 Census of Agriculture. Utah prelim- inary report. U.S. Bureau of the Census, Wash- ington, D.C. INFLUENCE OF CRYPTOGAMIC CRUSTS ON MOISTURE RELATIONSHIPS OF SOILS IN NAVAJO NATIONAL MONUMENT, ARIZONA Jack D. Brotherson' and Samuel R. Rushforth' Abstract.— Cryptogamic soil crusts of Betatakin Canyon in Navajo National Monument were investigated to un- derstand the influence of such crusts on soil moisture relationships and potential sediment production. Crusts sam- pled were part of the pinyon-juniper community and were studied in paired units. The presence of crusts on soils significantly increased the depth of water penetration and decreased runoff. Soils showed reduced infiltration of wa- ter where lichen and algal crusts were present and enhanced infiltration rates where mosses were present. Crusts ap- pear to cause surface sealing and therefore likely reduce surface evaporation rates as well. Cryptogamic crusts are nonvascular plant communities that grow on or immediately beneath the soil surface. Such communities are components of most desert ecosystems. They have been described in several ecosys- tems in western North America (Anderson and Rushforth 1976, Anderson et al. 1982a) as well as in the deserts of the Middle East (Evenari et al. 1971). Until recently scant at- tention had been given them and little was known concerning their role in native ecosys- tems. Studies of the past decade indicate that they exert a significant impact on reducing soil erosion (Evenari et al. 1971, Loope and Gifford 1972, Kleiner and Harper 1972, Klei- ner and Harper 1977, Anderson et al. 1982a, Anderson et al. 1982b). Fletcher and Martin (1948) found that fungal and algal crusts in- crease the tensile strength of soil. The algae appear to be the most effective in binding the surface soil particles (Durrell and Shields 1961) because of the thick gelatinous sheaths that enclose the trichomes of several algal species (Anderson and Rushforth 1976). Such gelatinous sheaths add strength and aggregat- ing qualities to the I or 2 mm of surface soil upon which they grow (Anantani and Ma- rathe 1974). Research on the biology of cryptogamic crusts has also been done in several other areas. These studies include taxonomy (Ali and Sandhu 1972, Anderson and Rushforth 1976); nitrogen fixation (MacGregor and Johnson 1971, Reddy and Gibbons 1975); land reclamation (Singh 1950); soil fertility (Shields and Durrell 1964); reproduction, growth and habitat relations (Evenari et al. 1971, Anderson et al. 1982b); and moisture (Booth 1941, Loope and Gifford 1972). The objective of this study was to in- vestigate the influence of cryptogamic crusts in the pinyon-juniper woodlands of north- eastern Arizona on depth of water pene- tration, infiltration, runoff, and potential sediment production. Study Area Navajo National Monument is located in northeastern Arizona (Fig. 1) and is the site of three large Anasazi Indian cliff dwellings. Betatakin Canyon, the site of the present study, is a side canyon of the larger Tsegi Canyon complex and has been described by Hack (1945). The major geological formation comprising the canyon is Navajo Sandstone, which forms sheer towering cliffs 200 m or more in height. The canyon floor consists of deep alluvial deposits of sandy Quaternary fill. Kayenta sandstone outcrops in the lower reaches of the canyon. The annual temperatures recorded at the park headquarters weather station at Betata- kin canyon ranges from -23 to 38 C with a mean of 10 C. The number of frost-free days in the area varies from 107 to 213, with an average of 155 days. Total annual precipi- tation ranges from 17 to 48 cm with a yearly mean of 29 cm. There is a single wet season lasting from late summer through fall. 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 73 74 Great Basin Naturalist Vol. 43, No. 1 G A R F I ELD SAN JUAN KANE ■\\^i « -.^^F , UTAH i^ POWELL ^^ i;ARIZONA I \ KEET SIEl tUIN COCONINO HOUSE I /BiTATAKIN RUIN^,.-— ^^,3 y /, Shontj,' N A V A J 0 APACHE NAVAJO INDIAN RESE RVATI LEGEND = STATE LINE I COUNTY LINE I ' MAIN HIGHWAY O N DIRT aOAD I /-'^ RIVEt I O TOWN I ■ *UINS SITE o (J X UJ LU z Fig. 1. Reference map of study area. Methods Cryptogamic crusts were sampled in the pinyon-juniper {Pinus edulis-Juniperus os- teosperma) community that borders Betatakin Canyon in Navajo National Monument, Ari- zona. Cryptogam crusts were studied in paired units so that varying conditions in habitat (slope, exposure, soil texture, etc.) could be kept to a minimum. Pairs consisted of five sites where crusts were intact and un- disturbed and five adjacent sites where the crusts had been heavily disturbed or de- stroyed. A total of 10 sites were considered for each measurement. Pairs were always lo- cated within 2 m of each other. Water infiltration rates were measured by using a thin-walled aluminum cylinder 12 cm tall and 65 mm in diameter. The cylinder was gently turned into the crust or soil to a depth of 2 cm and then 50 ml of water was ponded above the core inside of the cylinder. Infil- tration into the core was measured as the number of seconds needed for the ponded water to disappear into the core. Depth of water penetration and runoff were assessed by raining 1.5 liters of water January 1983 Brotherson, Rushforth: Cryptogamic Crusts 75 onto the crust or adjacent soil surface through a perforated 80 mm diameter disk. The perforations were evenly spaced on a 0.5 cm grid. The disk was placed at a distance of 1.2 m above the ground surface. Total deliv- ery time for the water to be dispensed onto the crust or soil surface was 60 seconds. These rates were designed to approximate or exceed precipitation at cloudburst propor- tions (i.e., 10 cm/hr). High intensities of pre- cipitation, such as those exceeding infiltration capacities of the soil, are significant because of their effects on runoff and erosion. Once the water had disappeared into the crust or soil surface, depth of penetration was mea- sured immediately. Five depth measurements were taken for each watering at each of the 10 areas and then averaged to give a single value for each site. Runoff was measured by recording the across slope and downslope spread of water rained onto study sites. The area of spread was computed from these measurements us- ing the formula for the area of an ellipse. Soil movement was assessed by estimating the amount of soil moved during a measured rain. The following index was used: 1 = no appreciable movement; 2 = moderate move- ment—up to 10 percent of soil being dis- placed; and 3 = heavy movement— between 10 and 20 percent of soil being displaced. All runoff and soil movement measure- ments were taken during the third week of August 1980. Sampling intensity was deter- mined following the estimation procedures described by Avery 1975. Significant differ- ences in the paired measurements were as- sessed through the use of Students-t statistic. Results and Discussion The influence of cryptogamic crusts on six soil moisture characteristics was assessed. Av- erage values for all measurements taken dur- ing this study are given in Table I. All but one of the measured characteristics showed significant differences between crusted and uncrusted soils. Infiltration measurements on the paired study sites indicated that well-developed cryptogamic crusts (Fig. 2) significantly in- creased the depth of water penetration. This was also found by Loope and Gifford (1972). Downslope movement of water was signifi- cantly greater on the sites that exhibited no crust development. Likewise, the differences in total area of surface spread was signifi- cantly greater on uncrusted soils. These dif- ferences are probably best explained by the micro-topographic changes that develop at the soil surface under the influence of crypto- gamic crust growth. Well-developed crusts form pedestals so that the ground surface looks something like a convoluted brain coral (Fig. 2-4). Hills and valleys a few centimeters in relief develop across broad crusted areas. The small valleys run in all directions and cause pooling of the water as it hits the soil surface (Fig. 5). This pooling holds the water in place for extended periods, thus increasing the time for infiltration to occur and simulta- neously decreasing runoff and movement across the soil surface. With reduced surface movement, deeper penetration of water oc- curs. The net effect is to slow the movement of surface-flowing water, providing longer periods for infiltration, less opportunity for Table 1. Relationships of cryptogamic crusts growing on the soil in Navajo National Monument to measured moisture parameters. Figures represent means and standard deviations (sd). Characteristic Crust Noncrust Significance measured Mean sd Mean sd level Water penetration depth (cm) 5.46 1.35 3.23 0.69 .05 Downslope spread (cm) 67.62 13.74 95.50 4.24 .001 Across slope spread (cm) 47.24 10.87 45.72 6.48 NS Area of spread (sq cm) 10434.11 3041.18 13738.50 2185.26 .001 Soil movement" 1.00 0.00 2.60 0.89 .01 Infiltration (seconds) Moss cover 15.40 3.90 238.00 87.90 .001 Lichen and algae cover 48.00 14.50 31.00 8.10 .001 'Soil movement was assessed as follows: 1 = no movement, 2 between 10-20 percent of soil being displaced. moderate movement— up to 10 percent of soil being displaced, 3 = heavy movement— 76 Great Basin Naturalist Vol. 43, No. 1 Figs. 2-5. Cryptogamic crusts. Left top, moving clockwise: 2. Crusts beneath Utah juniper trees. 3. Well-devel- oped cryptogamic soil crusts. 4. Close-up of cryptogamic crusts showing typical pinnacle development. 5. Close-up of cryptogamic crust after experimental rain showing water ponding. concentration in rills, and decreased power to cause erosion. In other words, cryptogam crusting fosters more infiltration and less run- off of surface water. Well-developed crust areas also showed significantly less soil movement (Table 1). These data support the findings of several other studies (Fletcher and Martin 1948, Loope and Gifford 1972, Kleiner and Harper 1977, Anderson et al. 1982b). Cryptogamic crusts appear to have a protective influence on the soil in four major ways. First, they bind the soil surface particles with the inter- twining growth of algal and fungal filaments (Durrell and Shields 1961). Second, the moss and lichen constituents of cryptogam crusts aid in stabilizing the soil by covering the sur- face with thalli and penetrating the soil sur- face with rhizoids (Anderson et al. 1982b). Third, the irregularities of a well-developed cryptogamic crust surface tend to break up microwind patterns and thus reduce wind- born soil movement (Brady 1974). And fourth, with less water movement there is also significantly less soil movement. Well-developed crusts also influenced wa- ter movement into the soil. Where moss cov- er was high, infiltration rates were greatly enhanced over areas where moss cover had been removed. The enhancement of infil- January 1983 Brotherson, Rushforth: Cryptogamic Crusts 77 tration appeared to be due to the moss thalli acting as a sponge. On the other hand, where they had been removed, a .05 to 1 cm thick layer of silt beneath them acted to retard in- filtration. Infiltration rates were significantly reduced or impeded by lichen and algal crust cover. The highest infiltration rates (most rapid penetration by water) occurred on soils with no cryptogamic cover (Table 1), In gen- eral, where cryptogamic cover was high, in- creased resistance to infiltration occurred. Loope and Gifford (1972) noted this pattern and also found that, when crusts were wetted previous to infiltration trials, infiltration rates on crusted soils were retarded by a factor of two. Fritsch (1922) first suggested that the highly mucilaginous sheaths of blue-green al- gae, which are the major components of cryptogamic crusts in arid environments, might form a layer at the soil surface that would both impede water infiltration into the soil and impede evaporation of soil moisture caught beneath the algal layer. This would provide more water to the plants growing in such areas. Booth (1941) later tested this hy- pothesis and showed that more moisture was to be found in the upper layers of soil (i.e., the upper 2.5 cm) where cryptogamic crusts were prominent than in adjacent soils with no crusts (i.e., 8.9 percent vs. 1.3 percent, respectively). Data from several studies indicate that high cryptogamic crust cover is associated with high silt in the soil surface (Evenari et al. 1971, Loope and Gifford 1972, Kleiner and Harper 1977, Anderson et al. 1982b). Textural observations on our sites showed similar patterns. Kleiner and Harper (1977) also argue that once established the crusts tend to trap silt at the soil surface. Evenari et al. (1971) and Blackburn and Skou (1974) present data that indicate that soils high in silt often have low permeability rates and high runoff. They suggested that soils with high levels of silt in the upper layers often show high initial infiltration rates, but, as more wetting occurs, the percolation rates decrease rapidly and eventually an almost impenetrable layer can be formed. Beneath such a sealed surface, air caught in the voids of the lower layers may have a difficult time escaping and may therefore further retard in- filtration (Evenari et al. 1971). It appears then that at least three factors tend to reduce water infiltration rates in soils with cryptogam crusts: (1) the effect of high levels of silt in the soil and its resultant swell- ing and sealing action when mixed with wa- ter (Evenari et al. 1971); (2) the wetting ac- tion of the water on the gelatinous sheaths of the algal filaments, causing the filament to swell and tightly bind the surface soil par- ticles (Anantani and Marathe 1974, Durrell and Shields 1961, Fritsch 1922); and (3) air trapped beneath the sealed surface to further impede water penetration. Evenari et al. (1971) also indicated from their research on micro-watershed irrigation projects that, as the farm areas receive runoff water ladened with silt from the watersheds and as the silt is deposited on the soil surface, evaporation from the irrigated fields was re- duced to as little as 7.4 mm over a seven month period. This kind of reduction in evaporation in a desert with annual evapo- ration values from 1700 to 2700 mm would be highly important relative to moisture re- tention in the subsurface layers of the soil. Since cryptogamic crusts tend to seal the soil surface and since crusts also increase the depth of water penetration, the effects they have on reducing moisture stress in desert ecosystems could prove to be extremely valu- able. Furthermore, since crust communities tend to grow in association with high silt lev- els at the soil surface, these elevated silt lev- els undoubtedly further reduce water losses by evaporation. This being the case, crypto- gamic crusts may be as important in their role in water conservation in desert systems as they are in preventing soil erosion. Literature Cited Ali, S., and G. R. Sandhu. 1972. Blue-green algae of the saline soils of the Punjab. Oikos 22:268-272. Anantani, Y. S., and D. V. Marathe. 1974. Soil aggre- gating effects of some algae occurring in the soils of Kutch and Rajasthan. Univ. of Bombay 41:94-100. Anderson, D. C, K. T. Harper, and S. R. Rushforth. 1982a. Recovery of cryptogamic crusts from graz- ing on Utah winter ranges. J. Range Manage. 35:355-359. Anderson, D. C, K. T. Harper, and R. C. Holmgren. 1982b. Factors influencing the development of cryptogamic soil crusts in Utah deserts. J. Range Manage. 35:180-185. 78 Great Basin Naturalist Vol. 43, No. 1 Anderson, D. C, and S. R. Rushforth. 1976. The cryptogamic flora of desert soil crusts in southern Utah. Nova Hedwigia 29:691-729. Avery, T. E. 1975. Natural resources measurements. 2d Ed. McGraw-Hill Book Company, New York. 339 pp. Blackburn, W. H., and C. M. Skou. 1974. Infiltration rates and sediment production of selected plant communities in Nevada. J. Range Manage. 27:476-479. Booth, W. E. 1941. Algae as pioneers in plant succes- sion and their importance in erosion control. Ecology 22:38-46. Brady, N. C. 1974. The nature and properties of soils. 8th ed. Macmillan Publishing Co., Inc., New York. 639 pp. DuRRELL, L. W., AND L. M. SHIELDS. 1961. Character- istics of soil algae relating to crust formation. Trans. Amer. Micro. Soc. 80:73-79. Evenari, M., L. Shanan, and N. Tadmor. 1971. The Negev— the challenge of a desert. Harvard Univ. Press, Cambridge, Massachusetts. 345 pp. Fletcher, J. E., and W. P. Martin. 1948. Some effects of algae and molds in the rain-crust of desert soils. Ecology 29:95-100. Fritsch, F. E. 1922. The terrestrial algae. J. Ecology 10:220-236. Hack, J. T. 1945. Recent geology of the Tsegi Canyon. Appendix I in R. L. Reals, G. W. Brainerd, and W. Smith, eds. Archaeological studies in North- east Arizona. Univ. of California Press, Berkeley and Los Angeles. Kleiner, E. F., and K. T. Harper. 1972. Environment and community organization in grasslands of Canyonlands National Park. Ecology 53:229-309. 1977. Soil properties in relation to cryptogamic ground cover in Canyonlands National Park. J. Range Manage. 30:202-205. LooPE, W. L., AND G. F. GiFFORD. 1972. Influence of a soil microfloral crust on select properties of soils under pinyon-juniper in southeastern Utah. J. Soil and Water Conser. 27:164-167. MacGregor, A. N., AND D. E. Johnson. 1971. Capacity of desert algal crusts to fix atmospheric nitrogen. Soil Sci. Soc. of Amer. Proc. 35:843-844. Reddy, G. B., and J. Gibbons. 1975. Nitrogen fixation by algae on fescuegrass soil crusts. Soil Sci. of Sco. Amer. Proc. 39:654-656. Shields, L. M., and L. W. Durrell. 1964. Algae in rela- tion to soil fertility. Bot. Rev. 30:92-128. Singh, R. N. 1950. Reclamation of "Usar" lands in India through bluegreen algae. Nature 165:325-326. A VASCULAR FLORA OF THE SAN RAFAEL SWELL, UTAH' James G. Harris^ Abstract.— The vegetation of the San Rafael Swell in southeastern Utah is examined based on personal field col- lections and previously collected herbarium specimens in the Brigham Young University Herbarium (BRY). An anno- tated checklist includes information on frequency of occurrence and habitat preference for each entity. Treated are 491 vascular plant taxa from 59 families. The San Rafael Swell is the eroded rem- nant of a massive domal anticline, oval in shape, stretching along a northeasterly axis from Capitol Reef National Park in northern Wayne County to the foot of the Tavaputs Plateau in Carbon County. Folding of the earth's crust, which formed the domal anti- cline, and the subsequent erosion of the cen- tral dome left a huge area of concentric stone rings, each progressively younger from the innermost to the outermost. One of these rings, the Ferron Sandstone Member of the Mancos Shale Formation, encloses approx- imately L4 million acres of land and forms the boundary of this study. In some cases, where the Ferron Sandstone has been com- pletely eroded from the structural confines of the swell, artificial boundaries have been fol- lowed (Fig. 1). Throughout the field seasons of 1979 and 1980, I attempted to collect representatives of all vascular plant species growing in the San Rafael Swell. A voucher specimen from each collection has been placed in the Brig- ham Young University Herbarium (BRY). In- cluded in the checklist are species not found by me, but collected by others and vouched for by specimens in the Brigham Young Uni- versity Herbarium. The checklist is not defin- itive; additional taxa will undoubtedly be dis- covered in the swell. The following sources were used for identi- fication and nomenclature: lower vascular plants and gymnosperms, Cronquist et al. (1972); dicotyledons, Neese and Welsh (1981), Welsh (1978, 1980a, 1980b), Welsh and Atwood (1981), Welsh and Moore (1973), Welsh and Reveal (1977), Welsh et al. (1981); monocotyledons, Cronquist et al. (1977). The checklist includes 478 vascular plant taxa found in the study area and an addition- al 13 taxa known to occur on lands immedi- ately adjacent to the study area and to be ex- pected in the San Rafael Swell. These 13 taxa are listed below: Allionia incarnata L. Cordylanthus wrightii Gray Descurainia pinnata (Walter) Britt. var. filipes (Gray) Peck Eriogonum smithii Reveal Euphorbia micromera Boiss. Haplopappus drummondii (T. & G.) Blake Helianthus petiolaris Nutt. Hymenopappus filifolius Hook. var. lugens (Greene) Jeps. Lupinus pusillus Pursh var. rubens (Rydb.) Welsh Phacelia demissa Gray var. heterotricha Howell Physocarpus alternans (Jones) Howell Sphaeralcea munroana (Dougl.) Spach Sporobolus giganteus Nash The flora of the San Rafael Swell is not particularly rich in numbers of species. This is due, at least in part, to the relatively nar- row elevational range in the area. Although not rich in numbers of species, the flora is rather rich in numbers of unique and geo- graphically restricted species. The following eight species are strictly endemic to the San Rafael Swell: Astragalus rafaelensis Jones Cryptantha johnstonii Higgins Cryptantha pnesiana (Payson) Payson Erigeron maguirei Cronquist 'Part of a master's thesis, originally submitted to the Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 'Department of Botany and Range Science, Brigham Young University, Prove, Utah 84602. Present address: Department of Botany, University of Alberta, Edmonton, Alberta T6G 2E9. 79 80 Great Basin Naturalist Vol. 43, No. 1 Emery County Fig. 1. Map of the San Rafael Swell study area. Pediocactus despainii Welsh & Goodrich Sclerocactus wrightii Benson Sphaeralcea psoraloides Welsh Thelypodiopsis harnebyi Welsh & Atwood An additional seven species are endemic to the San Rafael Swell and the adjacent lands in close proximity to the swell. They are list- ed as follows: Astragalus pardalinus (Rydb.) Barneby Astragalus woodruffii Jones Hoffmanseggia repens (Eastw.) Cockerell Hymenoxys depressa (T. & G.) Welsh & Reveal Lomatium junceum Barneby & Holmgren Psorothamnus polyadenius (Torr.) Rydb. var. jonesii Barneby Townsendia apnea Welsh & Reveal Vegetative Zones The vegetation of the San Rafael Swell can be divided into five major zones: the Sand Desert Shrub Zone, the Salt Desert Shrub Zone, the Mixed Desert Shrub Zone, the Pin- yon-Juniper Zone, and the Ponderosa Pine- Mountain Brush Zone. These zones are con- trolled principally by elevation, precipi- tation, and local edaphic factors. Although in some areas the zones are clear and well de- fined, in most areas they interfuse consid- erably and it becomes difficult to draw a sharp boundary line between them. January 1983 Harris: San Rafael Swell Flora 81 Sand Desert Shrub Zone.— The San Ra- fael Desert, in the southeastern region of the San Rafael Swell, ranges from an elevation of about 4200 ft (1280 m) at the lowest point to about 5000 ft (1524 m) at the highest point. It is a sandy desert with moving dunes in some areas. Common shrubs in this zone in- clude: Artemisia filifolia. Ephedra torreyana. Yucca hatrimaniae, Poliomintha incana, Ti- quilia latior, and Coleogyne ramosissima. Abronia elliptica, Oenothera pallida, Hoff- manseggia repens, and Oryzopsis hymenoides are characteristic herbaceous plants of this zone. Salt Desert Shrub Zone.— In the San Rafael Swell the Salt Desert Shrub Zone is widespread, occurring mainly on alkaline clay soils between 4500 and 5800 ft (1372-1766 m) in elevation. Communities in this zone are characteristically dominated by Atriplex confertifolia interspersed with any of the following subdominant species: Hilaria jamesii, Oryzopsis hymenoides, Enceliopsis nudicaulis, Aristida purpurea, Eriogonum in- flatum, Salsola iherica or Ceratoides lanata. Highly alkaline flood plains in this zone are often covered by nearly pure stands of Sarcobatus vermiculatus, which may at times be accompanied by thick clumps of Tamarix ramosissima. Heavy clay soils of the Mancos Shale For- mation are often dominated by Atriplex cu- neata and Atriplex corrugata. During wet springs these areas may be blanketed by an- nual species such as Cleome lutea, Lupinus pusillus, Malcolmia africana, Camissonia sca- poidea, and Camissonia eastwoodiae. Mixed Desert Shrub Zone.— Deep, well- drained soils between 4500 and 5800 ft (1372-1766 m) in elevation are often domi- nated by one or several of the following shrubby species: Artemisia tridentata, Arte- misia bigelovii, Artemisia spinescens, Xantho- cephalum sarothrae or Chrysothamnus vis- cidiflorus. These may be intermingled with an understory of herbaceous plants including: Hilaria jamesii, Stipa comata, Boutehua gra- cilis. Astragalus spp., Sphaeralcea spp., and Leucelene ericoides. Pinyon-Juniper Zone.— Pinus edulis and Juniperus osteosperma are conspicuously dominant at elevations between 5800 and 7800 ft (1766-2377 m). Within this altitudin- al range Juniperus osteosperma is dominant at the lower elevations and Pinus edulis is domi- nant at the higher elevations. Intermediate elevations yield a heterogeneous mixture of both species. Dense stands have almost no understory, but more open or scattered stands will in- clude almost all the species found in the Mixed Desert Shrub Zone. PoNDEROSA Pine-Mountain Brush Zone.— The highest point in the San Rafael Swell is the San Rafael Knob at 7921 ft (2414 m) in elevation. The area surrounding the knob and the summit of Cedar Mountain rep- resent the highest vegetative zone in the swell. Pinus edulis continues to remain domi- nant even at the highest elevations, but above 7500 ft (2286 m) plants characteristic of the Ponderosa Pine-Mountain Brush Zone be- come increasingly common. Some common species in this zone are: Pinus ponderosa, Ar- temisia nova, Cowania mexicana, Mahonia fremontii, Populus tremuloides, Amelanchier utahensis, Cercocarpus montanus, Ribes lep- tanthum, Juniperus scopulorum, Hymenoxys richardsonii, Symphoricarpos longiflorus and Philadelphus microphyllus. In addition to these five major vegetative zones, smaller communities dependent on unique local habitats are scattered through- out the area. These include plants distributed along seeps and streams, along washes and roadsides where runoff collects, in hanging gardens, on rocky outcrops, and on disturbed sites. Annotated Checklist Following each species entry in the check- list (with the exception of the 13 taxa from lands adjacent to the study area) a designa- tion is given concerning the frequency of oc- currence and habitat preference of the taxon. Plants of broad distribution that are not lim- ited to a specific zone or community type are listed as widespread. The following abbrevia- tions are used: C— common O— occasional U— uncommon R— rare SnD— Sand Desert Shrub Zone SID— Salt Desert Shrub Zone MD— Mixed Desert Shrub Zone PJ— Pinyon-Juniper Zone 82 Great Basin Naturalist Vol. 43, No. 1 PM— Ponderosa Pine-Mountain Brush Zone SS— Seeps and streams WR— Washes and roadsides HG— Hanging gardens RO— Rocky outcrops DS— Disturbed sites W— Widespread ACERACEAE Acer glabrum Torr. (R;PM) Anacardiaceae Rhus trilobata Nutt. var. simplicifolia (Greene) Barkl. (C;WR) Rhus trilobata Nutt. var. trilobata (C;WR) Toxicodendron rydbergii (Small) Greene (C;SS,HG) Apiaceae Cymopteris acaulis (Pursh) Raf. (0;SnD,SlD) Cyrnopteris bulbosus Nels. (C;W) Cymopteris fendleri Gray (C;W) Cymopteris purpurascens (Gray) Jones (0;S1D) Cymopteris purpureus Wats. (0;W) Lomatium junceum Barneby & Holmgren (R;S1D,MD,PJ) Stum suave Walt. (R;WR) Apocynaceae Amsonia eastwoodiana Rydb. (C;SnD) Ainsonia jonesii Woodson (0;MD,PJ) Apocynum cannabintim L. (C;SS,HG) Cycladenia humilis Benth. var. jonesii (Eastw.) Welsh & Atwood (R;MD) ASCLEPIADACEAE Asclepias cryptoceras Wats. (C;W) Asclepias labriformis Jones (C;WR,SD,MD) Asclepias macrosperma Eastw. (C;SnD,MD) Asclepias ruthiae Maguire & Woodson (0;W) Asclepias speciosa Torr. (C;WR) ASTERACEAE Ambrosia acanthicarpa Hook. (0;S1D,MD,PJ) Ambrosia artemisiifolia L. (C;S1D,MD,PJ) Artemisia biennis Willd. (U;WR) Artemisia bigelovii Gray (C;MD,PJ) Artemisia campestris L. (U;SnD,MD) Artemisia dracunculus L. (C;WR) Artemisia filifolia Torr. (C;SnD) Artemisia frigida Willd. (R; WR) Artemisia ludoviciana Nutt. (0;WR) Artemisia nova Nels. (C;PJ,PM) Artemisia pygmaea Gray (0;S1D,MD,PJ) Artemisia spinescens Eaton (0;S1D, MD) Artemisia tridentata Nutt. (C;MD,PJ,WR) Aster glaucodes Blake (U;PM) Baccharis emoryi Gray (U;WR) Brickellia longifolia Wats. (U;WR) Brickellia microphylla (Nutt.) Gray (U;WR) Brickellia oblongifolia var. linifolia (Eaton) Robins (0;PJ,WR) Brickellia scabra (Gray) Nels. (R;HG,WR) Chaenactis stevioides Hook. & Arn. (C;S1D,MD) Chamaechaenactis scaposa (Eastw.) Rydb. (0;PJ) Chrysothamnus greenei (Gray) Greene (0;S1D) Chrysothamnus linifolius Greene (0;WR) Chrysothamnus nauseosus (Pall.) Britt. var. consimilis (Greene) Hall (C;WR) Chrysothamnus nauseosus (Pall.) Britt. var. gnapha- loides (Greene) Hall (U;WR) Chrysothamnus nauseosus (Pall.) Britt. var. junceus (Greene) Hall (U;WR) Chrysothamnus nauseosus (Pall.) Britt. var. leio- spermus (Gray) Hall (R;WR) Chrysothamnus pulchellus (Gray) Greene (U;WR) Chrysothamnus viscidiflorus (Hook.) Nutt. var. steno- phyllus (Gray) Hall (U;WR) Chrysothamnus viscidiflorus (Hook.) Nutt. var. vis- cidiflorus (C;WR) Cirsium pulchellum (Greene) Woot. & Standi. (U;WR) Cirsium undulatum (Nutt.) Spreng. (C;WR) Dicoria canescens Gray (U;SnD) Encelia frutescens Gray (U;WR) Enceliopsis nudicaulis (Gray) Nels. (C;S1D, MD) Enceliopsis nutans (Eastw.) Nels. (C;S1D,MD) Erigeron argentatus Greene (0;S1D,MD,PJ) Erigeron compactus Blake var. consimilis Cronquist (0;MD,PJ) Erigeron divergens T. & G. (U;PJ) Erigeron eatonii Gray (U;PJ,PM) Erigeron maguirei Cronquist (R;WR) Erigeron pulcherrimus Heller var. pulcherrimus (C;S1D,MD,PJ) Erigeron pumilus Nutt. ssp. concinnoides Cronq. var. condensatus (Eaton) Cronq. (C;S1D,MD,PJ) Erigeron utahensis Gray (C;WR,RO) Gaillardia pinnatifida Torr. (C;S1D,MD) Gaillardia spathulata Gray (C;S1D,MD,PJ) Grindelia squarrosa (Pursh) Dunal. (0;WR) Haplopappus armerioides (Nutt.) Gray (C;MD,PJ,RO) Haplopappus drummondii (T. & G.) Blake Helianthella microcephala (Gray) Gray (C;PJ,PM) Helianthus annutts L. (C;WR) Helianthus anomalus Blake (C;SnD) Helianthus petiolaris Nutt. Heterotheca villosa (Pursh) Shinners (C;S1D,MD,PJ) Hymenopappus filifolius Hook. var. lugens (Greene) Jeps. Hymenopappus filifolius Hook. var. megacephalus Turner (C;S1D,MD) Hymenopappus filifolius Hook. var. pauciflorus (Johnst.) Turner (0;S1D) Hymenoxys acaulis (Pursh) Parker var. arizonica (Greene) Parker (C;MD,PJ) Hymenoxys depressa (T. & G.) Welsh & Reveal (0;R0) Hymenoxys richardsonii (Hook.) Cockerell (C;PJ,PM) Iva axillaris Pursh (0;SS,WR) Iva xanthifolia Nutt. (0;WR) Leucelene ericoides (Torr.) Greene (C;W) Lygodesmia arizonica Tomb (C;S1D,MD,WR) Lygodesmia grandiflora (Nutt.) T. & G. (C;S1D) Lygodesmia juncea (Pursh) D. Don (R;SnD) Machaeranthera canescens (Pursh) Gray (R;SnD) Machaeranthera grindelioides (Nutt.) Shinners (C;W) Machaeranthera linearis Greene (U;WR) Machaeranthera tanacetifolia (H.B.K.) Nees (C;WR) Malacothrix sonchoides (Nutt.) T. & G. (C;S1D,MD) Oxytena acerosa Nutt. (C;SS,WR) Parthenium ligulatum (Jones) Barneby (U;S1D,MD,PJ) Petradoria pumila (Nutt.) Greene (C;PJ,WR,RO) Platyschkuhria integrifolia (Gray) Rydb. var. oblongi- folia (Gray) Ellison (C;WR) Prenanthella exigua (Gray) Rydb. (C;WR,RO) Senecio multicapitatus Greenm. (U;PJ,PM,WR) January 1983 Harris: San Rafael Swell Flora 83 Senecio multilobatus T. & G. (0;PJ,WR) Stephanomeria exigua Nutt. (0;WR) Stephanomeria runcinata Nutt. (R;WR) Stephanomeria spinosa (Nutt.) Tomb (U;WR) Stephanomeria tenuifolia (Torr.) Hall (C;WR,RO) Taraxacum officinale Weber (C;SS) Tetradymia canescens DC. (0;PJ,PM) Tetradymia glabrata Gray (C;MD,PJ) Tetradymia spinosa Hook. & Am. (0;WR) Thelesperma subnudum Gray (C;W) Townsendia annua Beaman (U;MD) Townsendia aprica Welsh & Reveal (U;S1D,MD,PJ) Townsendia incana Nutt. (C;S1D,MD,PJ) Tragopogon dubius Scop. (C;W) Vanclevea stylosa (Eastw.) Greene (C;SnD) Wyethia scabra Hook. (C;WR) Xanthium strumarium L. (0;SS) Xanthocephalum microcephalum (DC.) Shinners (R;PJ) Xanthocephalum sarothrae (Pursh) Shinners (C;S1D,MD,PJ) Xylorhiza venusta (Jones) Heller (C;RO) Berberidaceae Mahonia fremontii (Torr.) Fedde (C;MD,PJ) Betulaceae Betula occidentalis Hook. (R;SS) Boraginaceae Cryptantha cinerea Greene (R;PJ,PM) Cryptantha confertifolia (Greene) Payson (R;SnD) Cryptantha crassisepala (T. & G.) Greene var. ela- chantha Johnst. (C;W) Cryptantha flava (Nels.) Payson (C;W) Cryptantha flavoculata (Nels.) Payson (C;PJ) Cryptantha fulvocanescens (Wats.) Payson var. ech- inoides (Jones) Higgins (0;S1D) Cryptantha fulvocanescens (Wats.) Payson var. ful- vocanescens (C;MD,PJ) Cryptantha gracilis Osterh. (U;PJ) Cryptantha humilis (Gray) Payson var. nana (Eastw.) Higgins (0;PJ) Cryptantha jamesii (Torr.) Payson var. disticha (Eastw.) Payson (U;MD,WR) Cryptantha jamesii (Torr.) Payson var. setosa (Jones) Johnst. (C;PJ) Cryptantha johnstonii Higgins (U;S1D) Cryptantha jonesiana (Payson) Payson (U;S1D) Cryptantha mensana (Jones) Payson (0;S1D) Cryptantha paradoxa (Nels.) Payson (0;PJ) Cryptantha recurvata Coville (0;MD,PJ,WR) Cryptantha rollinsii Johnst. (0;MD,PJ,WR) Cryptantha tenuis (Eastw.) Payson (0;WR) Cryptantha wetherillii (Eastw.) Payson (0;S1D) Heliotropium convulvulaceum (Nutt.) Gray (U;SnD) Lappula occidentalis (Wats.) Greene var. cupulata (Gray) Higgins (R;MD,PJ) Lappula occidentalis (Wats.) Greene var. occidentalis (C;W) Lithospermum incisum Lehm. (C;PJ) Tiquilia latior (Johnst.) Richardson (0;S1D,MD) Brassicaceae Arabis demissa Greene var. lanugida Rollins (R;PJ,PM) Arabis perennans Wats. (0;PJ) Arabis pulchra Jones var. pallens Jones (0;PJ) Arabis selbyi Rydb. (C;PJ) Caulanthus crassicaulis (Torr.) Wats. (C;PJ) Chorispora tenella (Pall.) DC. (U;DS) Descurainia pinnata (Walt.) Britt. var. filipes (Gray) Peck Descurainia pinnata (Walt.) Britt. var. intermedia (Rydb.) Hitchc. (U;MD) Descurainia pinnata (Walt.) Britt. var. osmarium (Cockerell) Shinners (C;W) Descurainia sophia (L.) Webb. (0;WR) Dithyrea wislizenii Engelm. in Wisliz. (0;SnD) Lepidium densiflorum Schrad. var. pubicarpum (Nels.) Thell. (R;PJ) Lepidium densiflorum Schrad. var. ramosum (Nels.) Thell. (0;MD,PJ) Lepidium montanum Nutt. var. jonesii (Rydb.) Hitchc. (C;W) Lesquerella alpina (Nutt.) Wats. var. alpina (0;PJ,PM) Lesquerella intermedia (Wats.) Heller (0;PJ,PM) Lesquerella kingii Wats. (R;PJ) Lesquerella rectipes Woot. & Standi. (C;W) Lesquerella subumbellata Rollins (U;PJ,PM) Malcolmia africana (L.) R. Br. in Ait. (C;S1D,DS) Physaria acutifolia Rydb. (C;PJ) Physaria chambersii Rollins (U;PJ) Schoencrambe linifolia (Nutt.) Greene (0;PJ,PM) Sisymbrium altissimum L. (C;PJ,PM) Stanleya pinnata (Pursh) Britt. (C;S1D) Stanleya viridiflora Nutt. in T. & G. (U;MD,PJ) Streptanthella longirostris (Wats.) Rydb. (C;W) Streptanthus cordatus Nutt. ex T. & G. (C;PJ) Thelypodiopsis barnebyi Welsh & Atwood (R;MD) Thelypodiopsis divaricata (Rollins) Welsh & Reveal (C;S1D,MD,PJ) Cactaceae Coryphantha vivipara (Nutt.) Britt. & Rose (0;PJ) Echinocereus triglochidiatus Engelm. var. melana- canthus (Engelm.) Benson (C;S1D,MD,PJ) Opuntia basilaris Engelm. & Bigel. (0;SnD,MD,PJ) Opuntia erinacea Engelm. (0;SnD) Opuntia polyacantha Haw. (C;W) Pediocactus despainii Welsh & Goodrich (R;PJ) Pediocactus simpsonii (Engelm.) Britt. & Rose (R;PJ) Sclerocactus parviflorus Clover & Jotter var. inter^ medius (Peebles) Woodruff & Benson (C;W) Sclerocactus wrightiae Benson (U;S1D,MD,PJ) Capparidaceae Cleome lutea Hook. (C;S1D) Cleome serrulata Pursh (0;PJ) Cleomella palmerana Jones (0;PJ) Caprifoliaceae Sambucus coerulea Raf. (U;PJ,PM) Symphoricarpos longiflorus Gray (0;PJ,PM) Caryophyllaceae Arenaria eastwoodiae Rydb. (C;PJ) Arenaria fendleri Gray (C;S1D) Arenaria hookeri Nutt. var. desertorum Maguire (R;PJ) Paronychia sessiliflora Nutt. (0;PJ,WR) Celastraceae Forsellesia meionandra (Koehne) Heller (0;PJ,RO) Chenopodiaceae Allenrolfea occidentalis (Wats.) Kuntze (0;S1D) Atriplex argentea Nutt. (U;S1D) Atriplex canescens (Pursh) Nutt. (C;S1D,MD) Atriplex confertifolia (Torr. & Frem.) Wats. (C;S1D) Atriplex corrugata Wats. (C;S1D) 84 Great Basin Naturalist Vol. 43, No. 1 Atriplex cuneata Nels. (C;S1D) Atriplex graciliflora Jones (U;S1D) Atriplex powellii Wats. {C;S1D) Atriplex saccaria Wats. (U;MD) Bassia hysopifolia Pallas (U;WR) Ceratoides lanata (Pursh) Howell (C;S1D) Chenopodium fremontii Wats. (C;W) Chenopodium glauciim L. (0;S1D) Chenopodium pratericola Rydb. (C;SS) Grayia brandegei Gray (U;MD) Halogeton glomeratus (Bieb.) Meyer (C;DS) Kochia iranica Bomm. (C;WR) Kochia americana Wats. (R;S1D,WR) Monolepis nuttalliana (Schult.) Greene (0;WR) Sahola iberica Sennen & Pan (C;W) Sarcobatus vermiculatus (Hook.) Torr. (C;S1D,SS,WR) Suaeda torreyana Wats. (C;SS,WR) CONVOLVULACEAE Convoltmlus arvensis L. (C;WR) Cuscuta cuspidata Engelm. (C;MD) CuPRESSACEAE Juniperus osteosperma (Torr.) Little (C;PJ,PM) Juniperus scopulorum Sarg. (U;PM) Cyperaceae Carex filifolia Nutt. (C;SS) Carex parryana Dewey (R;SS) Eleocharis palustris (L.) Roemer & Schult. (0;SS) Eleocharis rostellata (Torr.) Torr. (U;SS) Scirpus maritimus L. (0;SS) Scirpus pungens Vahl. (C;SS) Scirpus validus Vahl. (0;SS) Elaeagnaceae Shepherdia rotundifolia Parry (U;PJ) Shepherdia argentea (Pursh) Nutt. (U;WR) Ephedraceae Ephedra torreyana Wats. (C;W) Ephedra viridis Coville (C;W) Equisetaceae Equisetum laevigatum A. Br. (C;SS) Euphorbiaceae Euphorbia albomarginata T. & G. (U;PJ) Euphorbia fendleri T. & G. (C;S1D,MD,PJ) Euphorbia micromera Boiss. Euphorbia nephradenia Bameby (R;S1D,WR) Euphorbia palmeri Engelm. (R;S1D) Euphorbia parryi Engelm. (G;SnD) Euphorbia robusta (Engelm.) Small (0;PJ,PM) Euphorbia serphyllifolia Pers. (C;PJ,PM) Fabaceae Astragalus amphioxys Gray var. amphioxys (C;S1D,MD,PJ) Astragalus amphioxys Gray var. vespertinus (Sheld.) Jones (C;W) Astragalus argophyllus Nutt. var. martinii Jones (U;PJ,PM) Astragalus asclepiadoides Jones (C;S1D) Astragalus brandegei Porter (U;S1D,MD,PJ) Astragalus calycosus Torr. (0;S1D,MD,PJ) Astragalus ceramicus Sheldon (U;WR) Astragalus coltonii Jones var. coltonii (C;S1D,MD,PJ) Astragalus convallarius Greene var. convallarius (U;PJ,PM) Astragalus cymboides Jones (C;S1D,MD,PJ) Astragalus desperatus Jones var. desperatus (C;SnD,MD,PJ) Astragalus desperatus Jones var. petrophilus Jones (C;MD,SnD,PJ) Astragalus episcopus Wats. (C;MD,PJ) Astragalus flavus Nutt. var. argillosus (Jones) Bameby (C;S1D) Astragalus flavus Nutt. var. flavus (C;S1D,MD,PJ) Astragalus flexuosus (Hook.) Don. var. diehlii (Jones). Barneby (0;S1D,MD,PJ) Astragalus gey eri Gray (0;MD) Astragalus kentrophyta Gray var. cohradoensis Jones (U;WR,RO) Astragalus kentrophyta Gray var. elatus Wats. (R;S1D) Astragalus lentiginosus Dougl. var. araneosus (Sheld.) Barneby (G;MD,PJ) Astragalus lentiginosus Dougl. var. palans (Jones) Jones (C;MD,PJ) Astragalus lonchocarpus Torr. (C;S1D,MD,PJ) Astragalus moencoppensis Jones (G;S1D,MD,PJ) Astragalus mollissimus Torr. var. thompsonae (Wats.) Barneby (C;W) Astragalus musiniensis Jones (C;S1D,MD,PJ) Astragalus pardalinus (Rydb.) Barneby (0;SnD,MD,PJ) Astragalus praelongus Sheld. var. ellisiae (Rydb.) Barneby (C;S1D) Astragalus praelongus Sheld. var. praelongus (U;S1D) Astragalus preussii Gray var. preussii (C;S1D,MD,PJ) Astragalus rafaelensis Jones (0;S1D) Astragalus sabulonum Gray (0;PJ,PM) Astragalus spatulatus Sheld. (0;PJ,PM) Astragalus subcinereus Gray var. basalticus Welsh (R;PJ) Astragalus tenellus Pursh (0;MD,PJ) Astragalus wingatanus Wats. (R;S1D,MD,PJ) Astragalus woodruffii Jones (U;SnD,MD) Dalea flavescens (Wats.) Welsh (0;SnD,MD,PJ) Dalea oligophylla (Torr.) Shinners (0;WR,RO) Glycyrrhiza lepidota Pursh (C;WR,SS) Hedysarum boreale Nutt. (C;MD,PJ,PM) Hoffmanseggia repens (Eastw.) Cockerell (C;SnD) Lathyrus brachycalyx Rydb. var. eucosmus (Butters & St. John) Welsh (U;WR) Lupinus argenteus Pursh var. argenteus (R;WR) Lupinus argenteus Pursh var. tenellus Dougl. (U;MD) Lupinus brevicaulis Wats. (U;S1D,MD) Lupinus pusillus Pursh var. pusillus (C;S1D,MD,PJ) Lupinus pusillus Pursh var. rubens (Rydb.) Welsh Medicago sativa L. (C;WR) Melilotus alba Medicus (0;SS,WR) Melilotus officinalis (L.) Lam. (0;SS,WR) Oxytropis lambertii Pursh var. bigelovii Gray (R;PJ,PM) Oxytropis sericea Nutt. (0;MD,PJ) Psoralea lanceolata Pursh var. lanceolata (U;MD,PJ) Psoralea lanceolata Pursh var. stenophylla (Rydb.) Toft & Welsh (0;WR) Psorothamnus polyadenius (Torr.) Rydb. var. jonesii Barneby (R;S1D,MD) Psorothamnus thompsonae (Vail) Welsh & Atwood (R;MD) Robinia pseudoacacia L. (R;WR) Sophora stenophylla Gray (0;MD) Fumariaceae Corydalis aurea Willd. (R;PJ) January 1983 Harris: San Rafael Swell Flora 85 GEI>rrtANACEAE Swertia albomarginata (Wats.) Kuntze (R;PM) Swertih utahensis (Jones) St. John (C;SnD,MD) Hydrophyllaceae Phacelia constancei Atwood (R;S1D) Phacelia corrugata Nels. (C;W) Phacelia demissa Gray var. demissa (0;S1D,PJ) Phacelia demissa Gray var. heterotricha Howell Phacelia indecora Howell (R;S1D) Phacelia ivesiana Torr. (C;S1D,MD,PJ) Phacelia rafaelensis Atwood (C;MD,PJ) JUNCACEAE Juncus arcticus Willd. (C;SS) Juncus balticus Willd. (C;SS) Juncus nodosus L. (U;SS) Juncus torreyi Coville (0;SS) JUNCAGINACEAE Triglochin maritima L. (U;SS) Lamiaceae Poliomintha incana (Torr.) Gray (C;SnD) Liliaceae Allium geyeri Wats. (C;PJ,PM) Allium macropetalum Rydb. (U;S1D) Allium textile Nels. & Macbr. (C;SnD) Androstephium breviflorum Wats. (C;S1D,MD,PJ) Asparagus officinalis L. (0;WR,SS) Cahchortus nuttallii T. & G. (C;S1D,MD) Eremocrinum albomarginatum (Jones) Jones (C;SnD) Smilacina stellata (L.) Desf. (C;HG) Yucca harrimaniae Trel. (C;SnD,MD,PJ) LiNACEAE Linum aristatum Engelm. (U;WR) Linum perenne L. (C;WR) Linum puberulum (Engelm.) Heller (0;S1D) Linum subteres Winkler (0;WR,PJ) LOASACEAE Mentzelia albicaulis Dougl. (C;S1D,MD,PJ) Mentzelia humilis (Gray) DarL (U;WR) Mentzelia multiflora (Nutt.) Gray (C;SnD) Mentzelia pterosperma Eastw. (U;WR) Mentzelia pumila (Nutt.) T. & G. (0;WR) Malvaceae Sphaeralcea coccinea (Nutt.) Rydb. (C;W) Sphaeralcea grossulariifolia (Hook. & Arn.) Rydb. (0;SnD,MD) Sphaeralcea leptophylla (Gray) Rydb. (R;WR) Sphaeralcea munroana (Dougl.) Spach Sphaeralcea parvifolia Nels. (C;W) Sphaeralcea psoraloides Welsh (R;S1D,MD) Nyctaginaceae Abronia elliptica Nels. (C;SnD,MD,WR) Abronia nana Wats. (U;S1D,PJ) Allionia incarnata L. Oxybaphus linearis (Pursh) Robins. (C;W) Tripterocalyx micranthus (Torr.) Hook. (0;SnD) Oleaceae Fraxinus anomala Torr. in Wats. (C;WR) Onagraceae C^lylOphus lavandulaefolia (T. & G.) Raven (0;PT,RO) Cafnissonia eastwoodiae (Munz) Raven (C;SnD,SlD) CdviU^hhia scapoidea (T. & G.) Raven (C;SnD,SlD) Cafnissonia walkeri (Nels.) Raven (U;S1D,MD) Oenothera brachycarpa (Gray) Britt. (U;MD,PJ,WR) Oenothera caespitosa Nutt. (C;SnD,MD,WR) Oenothera pallida Lindl. ssp. pallida (C;SnD,MD,WR) Oenothera pallida Lindl. ssp. trichocalyx (Nutt.) Munz & Klein (C;S1D,MD) Orchidaceae Epipactis gigantea Dougl. ex Hook. (C;HG,SS) Orobanchaceae Orobanche fasciculata Nutt. (0;W) Papaveraceae Argemone corymbosa Greene (C;SnD) Argemone munita Dur. & Hilg. (C;SnD) PiNACEAE Pinus edulis Engelm. (C;PJ,PM) Pinus ponderosa Lawson (C;PM) Pseudotsuga menziesii (Mirb.) Franc. (C;PM) Plantaginaceae Plantago patagonica Jacq. (C;S1D,MD) POACEAE Agropyron caninum (L.) Beauv. (U;MD,RO) Agropyron cristatum (L.) Gaertn. (C;MD,PJ) Agropyron spicatum (Pursh) Scribn. & Smith (U;PJ) Agropyron trachycaulum (Link) Malte (U;WR) Andropogon hallii Hackel (R;WR) Aristida purpurea Nutt. (C;W) Bouteloua curtipendula (Michx.) Gray (R;MD) Bouteloua gracilis (H.B.K.) Lag. ex Steud. (C;S1D,MD,PJ) Bromus japonicus Thunb. (R;PJ) Bromus tectorum L. (C;WR,DS) Calamagrostis scopuhrum Jones (U;WR) Dichanthelium lanuginosum (Elliott) Gould (U;SS) Distichlis spicata (L.) Greene var. striata (Torr.) Scribn. (C;S1D,WR) Elymus canadensis L. (U;WR) Elymus cinereus Scribn. & Merr. (0;MD,PJ) Elymus junceus Fischer (R;S1D,MD) Elymus salina Jones (C;PJ) Erioneuron pilosum (Buckley) Nash (C;MD,RO,WR) Erioneuron pulchellum (H.B.K.) Tateoka (R;PJ) Festuca pratensis Huds. (U;SS) Hilaria jamesii (Torr.) Benth. (C;W) Hordeum jubatum L. (C;WR) Muhlenbergia asperifolia (Nees & Mey.) Parodi (0;WR) Muhlenbergia pungens Thurber (C;SnD) Munroa squarrosa (Nutt.) Torr. (0;PJ) Oryzopsis hymenoides (Roem. & Schult.) Ricker (C;W) Oryzopsis micrantha (Trin. & Rupr.) Thurber (U;PJ) Panicum virgatum L. (0;SS,WR) Phragmites australis (Car.) Trin. ex Steud. (C;SS) Poa fendleriana (Steud.) Vasey (0;PJ,PM) Poa pratensis L. (U;SS,WR) Poa sandbergii Vasey (U;PJ) Polypogon monspeliensis (L.) Desf. (U;SS,WR) Schedonnardus paniculatus (Nutt.) Trel. (R;S1D) Schizachyrium scoparium (Michx.) Nash (0;PJ,WR) Sitanion hystrix (Nutt.) Smith (C;W) Spartina gracilis Trin. (U;SS) Sporobolus airoides (Torr.) Torr. (C;S1D,MD) Sporobolus contractus Hitchc. (0;MD,PJ,WR) Sporobolus cryptandrus (Torr.) Gray (G;S1D,MD,PJ) Sporobolus flexuosus (Thurb.) Rydb. (U;SnD,WR) Sporobolus giganteus Nash Stipa arida Jones (U;WR) Stipa comata Trin. & Rupr. (C;MD,PJ) Stipa Columbiana Macoun. (U;PJ) 86 Great Basin Naturalist Vol. 43, No. 1 SHpa speciosa Trin. & Rupr. (U;PJ) ' Vulpia octoflora (Walt.) Rydb. (0;PJ,WR) . t'oLEMONIACEAE Gilia aggregata (Pursh) Spreng. (C;W) Gilia congesta Hook. (0;S1D) Gilia gunnisonii T. & G. (C;MD,WR) Gilia inconspicua (Smith) Sweet (C;PJ) Gilia latifolia Wats. (R;RO,WR) Gilia leptamena Gray (C;SnD,MD,PJ) Gilia polycladon Torr. in Emory (0;S1D) Gilia pumila Nutt. (0;S1D) Gilia roseata Rydb. (C;WR,RO) Gilia stenothyrsa Gray (0;PJ) Leptodactylon caespitosum Nutt. (U;WR) Leptodactylon pungens (Torr.) Nutt. (0;MD,WR,RO) Leptodactylon watsonii (Gray) Rydb. (U;WR,RO) Phlox austromontana Gov. (0;PJ) Phlox hoodii Rich. (C;MD,PJ) Phlox hngifolia Nutt. (0;S1D,MD) POLYGALACEAE Polygala acanthoclada Gray (R;MD) Polygala subspinosa Wats. (G;SlD,PJ,RO) POLYGONACEAE Eriogonum alatum Torr. in Sitgr. (C;PJ) Eriogonum hatemanii Jones (C;PJ) Eriogonum. bicolor Jones (C;MD,PJ) Eriogonum cernuum Nutt. (C;S1D) Eriogonum corymbosum Benth. var. corymbosum (G;S1D,MD) Eriogonum fasciculatum Benth. var. polifolium (Benth. in DG.) T. & G. (R;MD) Eriogonum flexum Jones (R;S1D) Eriogonum gordonii Benth. (U;S1D,MD) Eriogonum hookeri Wats. (0;MD,PJ) Eriogonum inflatum Torr. & Frem. var. fusiforme (Small) Reveal (0;S1D,MD,PJ) Eriogonum inflatum Torr. & Frem. var. inflatum (G;S1D,MD) Eriogonum jamesii Benth. var. flavescens Wats. (G;MD,PJ,WR) Eriogonum leptocladon T. & G. var. leptocladon (C;S1D) Eriogonum microthecum Nutt. var. foliosum (T. & G.) Reveal (0;S1D) Eriogonum ovalifolium Nutt. var. ovalifolium (C;W,RO) Eriogonum salsuginosum (Nutt.) Hook. (0;MD,PJ) Eriogonum shockleyi Wats. var. longilobum (Jones) Reveal (C;S1D,MD) Eriogonum smithii Reveal Eriogonum tumulosum (Bameby) Reveal (U;PJ) Eriogonum wetherillii Eastw. (0;W) Polygonum aviculare L. (0;WR,DS) POLYPODIACEAE Adiantum capillus-veneris L. (U;HG) Cheilanthes feei Moore (0;HG) Pellaea glabella Mett. ex Kuhn (R;HG) PORTULACEAE Portulaca oleracea L. (R;PJ,PM) Talinum validulum Greene (0;PJ) Ranunculaceae Aquilegia micrantha Eastw. (C;HG) Clematis ligusticifolia Nutt. (G;WR) Delphinium nuttallianum Pritz. ex Walp. (0;W) Delphinium scaposum Greene (0;PJ) Ranunculus cymbalaria Pursh (U;SS) Ranunculus testiculatus Grantz (0;DS) Rosaceae Amelanchier utahensis Koehne (C;WR,SS) Cercocarpus intricatus Wats. (G;PJ,RO) Cercocarpus montanus Raf. (G;PJ,PM,WR) Coleogyne ramosissima Torr. (G;SnD) Cowania mexicana D. Don (C;PJ) Fallugia paradoxa (D. Don) Endl. (0;WR) Physocarpus alternans (Jones) Howell Physocarpus monogynus (Torr.) Gou. (U;PJ) Purshia tridentata (Pursh) DC. (0;WR) Rosa woodsii Lindl. (R;PM) Rubiaceae Galium multiflorum Kellogg (0;WR) Salicaceae Populus x acuminata Rydb. (U;WR,SS) Popidus angustifolia James ex Torr. (0;WR) Populus fremontii Wats. (G;WR,SS) Populus tremuloides Michx. (0;PM) Salix exigua Nutt. (G;SS) Santalaceae Comandra umbellata (L.) Nutt. var. pallida (DG.) Jones (G;W,WR) Saxifragaceae Philadelphus microphyllus Gray (G;PJ,PM) Ribes cereum Dougl. (R;PJ,PM) Ribes leptanthum Gray (0;PM) Scrophulariaceae Castilleja chromosa Nels. (0;W) Castilleja linariaefolia Benth. (0;SS) Castilleja scabrida Eastw. (G;W) Cordylanthus kingii Wats. (0;PJ) Cordylanthus wrightii Gray Mimulus rubellus Gray (R;PM) Penstemon carnosus Pennell (G;PJ) Penstemon cyanocaulis Payson (U;WR) Penstemon eatonii Gray (0;PJ,PM) Penstemon lentus Pennell (U;WR) Penstemon utahensis Eastw. (G;PJ) Selaginellaceae Selaginella mutica Eaton ex Underw. (U;HG) Tamaricaceae Tamarix ramosissima Ledeb. (G;SS,WR) Typhaceae Typha latifolia L. (G;SS) Ulmaceae Celtis reticulata Torr. (U;WR) Ulmus pumila L. (R;WR) Literature Cited Cronquist, a., a. Holmgren, N. Holmgren, and J. Reveal. 1972. Intermountain flora: vascular plants of the Intermountain West. vol. I. Hafner Publishing Gompany, Inc., New York. Gronquist, a., a. Holmgren, N. Holmgren, J. Reveal, AND P. Holmgren. 1977. Intermountain flora: vascular plants of the Intermountain West. vol. VI. Golumbia University Press, New York. Neese, E., and S. L. Welsh. 1981. Astragalus section Desperati (Leguminosae) and a new species from the Uinta Basin of Utah. Rhodora 83:455-460. Welsh, S. L. 1978. Utah flora: Fabaceae (Leguminosae). Great Basin Nat. 38:225-367. January 1983 Harris: San Rafael Swell Flora 87 1980a. Utah flora: Malvaceae. Great Basin Nat. 40:27-37. 1980b. Utah flora: miscellaneous families. Great Basin Nat. 40:38-58. 1981. New taxa of western plants— In tribute. Brittonia 33:294-303. Welsh, S. L., N. D. Atwood, S. Goodrich, E. Neese, K. H. Thorne, and B. Albee. 1981. Preliminary index of Utah vascular plant names. Great Basin Nat. 41:1-108. Welsh, S. L., and G. Moore. 1973. Utah plants: Tracheophyta. 3d ed. Brigham Young University Press, Provo, Utah. Welsh, S. L., and J. L. Reveal. 1977. Utah flora: Brassi- caceae (Cruciferae). Great Basin Nat. 37:297-364. PRONGHORN RESPONSES TO HUNTING COYOTES Timothy D. Reynolds' Abstract.— Six accounts of pronghorn antelope {Antilocapra americana) chasing or attacking coyotes (Canis lat- rans) are described: three chases by individual pronghorn does, two by herds of antelope, and one joint effort by a pronghorn doe and a Short -eared Owl {Asio flammeus). Modifications of Berger's (1979) ungulate antipredatory de- fense model are proposed. Coyotes {Canis latrans) in the western United States feed on pronghorn antelope {Antilocapra americana). Pubhshed accounts indicate that coyote predation on pronghorn is not a particularly rare event (Thompson 1949, Arrington and Edwards 1951, Udy 1953, Beale and Smith 1973). In fact. Springer and Smith (1981) recorded prong- horn remains in more than 50 percent of the summer coyote scats they examined. Con- trariwise, until recently, published accounts of responses of pronghorn to predators were uncommon, and records of pronghorn chas- ing cr attacking coyotes were lacking. Berger (1979) described a "previously unknown de- fense strategy in pronghorn" in which a group of antelope chased a coyote. From this observation he developed a schematic repre- sentation of antipredatory defenses in un- gulates, and concluded that predator harass- ment is beneficial to the prey by (1) giving naive individuals the opportunity to recog- nize predators in a low risk situation, (2) al- lowing the prey to safely monitor the pred- ator's position, and (3) making the predator reluctant to attack in the future. Lipetz and Bekoff (1980) analyzed 25 antelope-coyote chases and concluded only that such encoun- ters appear to have direct survival value for proiighom fawns. Pescribed here are six observations of pronghorn, either singly or in groups, chasing coyotes. One event, detailed below, included a joint effort between a Short-eared Owl {Asio flammeus) and a pronghorn doe. A re- finement of Berger's (1979) antipredatory de- fense model is proposed for pronghorn. All observations were recorded in the sagebrush {Artemisia tridentata) dominated habitat at the National Environmental Research Park on the U.S. Department of Energy's Idaho National Engineering Laboratory (INEL) Site in southeastern Idaho. Observations Group Response On 14 July 1978 and 18 November 1979, I witnessed groups of pronghorn chase coyotes. The first occasion was similar to Berger's (1979) report. A coyote was observed stalking a small band of antelope (4 does, 1 fawn) that was loafing and feeding about 300 m from a larger group (5 does, 2 bucks, 2 fawns). One feeding doe from the smaller band apparent- ly sighted the coyote at a distance of nearly 100 m, stared toward the coyote for a few seconds, and sounded an alarm call. The re- maining antelope of both groups were then alert and directed their attention toward the vicinity of the coyote. When the stalking coyote approached within 40 m, the group of 5 quickly joined the larger group. The coyote followed, maintaining a distance of 40-50 m from the antelope, and sat down as the groups merged. One doe (thought to be a yearling) took a few steps toward the coyote, then returned to the main group. She re- peated this investigative sequence twice. On the fourth foray she was accompanied by the 13 other pronghorn. All antelope stopped momentarily about 30 m from the coyote, then burst into a full run toward the coyote. 'Department of Biology, Idaho State University, Pocatello, Idaho 83209; and U.S. Department of Energy, Radiological and Environmental Science Labo- ratory, 550 2nd Street, Idaho Falls, Idaho 83401. Present address: Biology Department, Boise State University, Boise, Idaho 83725. 88 January 1983 Reynolds: Pronghorn Responses 89 The coyote fled and was pursued for 300-400 m before disappearing from view. The November 1979 encounter differed from the previous one in that 2 coyotes were observed moving near a large group of about 120 pronghorn. The pronghorn sighted the coyotes at a distance of about 200 m. A large buck left the group and walked directly to- ward the coyotes. He was followed by about 20 animals (both does and bucks). This group began running toward the coyotes when the distance had closed to less than 100 m. The remaining 100 or so pronghorn simply ob- served the chase. The coyotes immediately took flight. The pronghorn stopped chasing after running about 150 m. The coyotes con- tinued their retreat, but at a slow run or trot, while repeatedly looking back at the prong- horn. The coyotes vanished from view at about 600 m. My first example above, and Berger's (1979) report, suggest that pronghorn groups must contain sufficient numbers of animals before a chase will be initiated. This lower limit or threshold concept may be valid un- der certain conditions, but as evidenced by the following accounts it is by no means a universal trend in antelope antipredatory behavior. Doe with Fawns On 20 June 1978, a female pronghorn was observed nursing two fawns, seemingly una- ware of a coyote furtively approaching her from the rear. While licking one fawn, the doe apparently noticed the coyote 30 m away and gave an alarm call. The fawns promptly dropped into the immobility re- sponse (Autenrieth and Fichter 1975) and the doe stared intently at the advancing coyote. The coyote approached to within 20 m, and then made a dash toward the antelope. The doe responded by charging the coyote, caus- ing it to veer away from the fawns. The doe pursued the coyote for nearly 50 m, and then returned to a position about halfway between the fawns and the predator. Twice more the coyote ran toward the fawns, and each time was thwarted by the charging doe, who again positioned herself between her young and the coyote. The coyote slowly moved away from the antelope while the doe intently watched its progress. When the coyote had withdrawn to a distance of 80 m, it abruptly changed its direction, putting itself on a course that would bring it within 20-30 m of the fawns. The doe again charged the coyote and pur- sued it for nearly 400 m before both dis- appeared from view. The doe returned to the area 85 minutes later, called her fawns from seclusion, and resumed nursing them. Another postparturient doe and coyote in- teraction was observed on 30 June 1978. Other than the fact that this doe had only one fawn, this encounter closely followed the sequence described above: the coyote ap- proached to within 30 m before charging, only to be charged by the doe. Second and third attacks followed; each time the coyote was chased a short distance away by the doe. On the fourth attempt, as the coyote veered from its course, the doe actually butted it in the side, rolling it over. The coyote regained its footing without losing momentum, and was vigorously pursued by the doe for about 150 m. The doe stopped, watched the coyote run away, then intermittently fed, or sham- fed (Autenrieth and Fichter 1975), for nearly 30 minutes before returning to the vicinity of the fawn. A third antelope doe was observed defend- ing two neonates, approximately two weeks old, from a pair of coyotes on 8 June 1979. When observations began (0925 hours MST) the doe was feeding and the fawns were ca- vorting nearby. At 0932 hours the doe spotted coyote No. 1 about 50 m to the north and gave an alarm call. The fawns immedi- ately lay down, separated from each other by a distance of 8-10 m. As the doe focused her attention on the now stationary coyote, coy- ote No. 2 appeared behind coyote No. 1 and began moving in an arc toward the east. Coy- ote No. 2 had approached to within 20 m of the fawns when the doe charged it, causing it to move further eastward from the fawns. Al- most simultaneously, coyote No. 1 dashed to- ward the fawns and was within 10 m of them before the doe whirled and charged, forcing it to the west of the secluded young. Coyote No. 2 then advanced and was driven off, again to the east. Coyote No. 1 again at- tacked, this time advancing within 1-2 m of one of the fawns before being repulsed by the doe. Bleating, the fawn burst from its bed, 90 Great Basin Naturalist Vol. 43, No. 1 and ran in a southerly direction accompanied by the doe. At the sound of the bleat, the sec- ond fawn immediately took flight, but was quickly brought down from behind by coyote No. 2. The doe and the surviving fawn ran at full speed for nearly 200 m and abruptly stopped. The fawn lay down and the doe moved in a seemingly leisure manner, roughly in a southwestward direction, fre- quently looking back toward the feeding coyotes. Joint Interspecific Response At 0545 hours on 13 May 1977, an extraor- dinary predator-prey encounter was observed and recorded. A single pronghorn doe was observed feeding about 400 m west of my po- sition and about 80 m west of a Short-eared Owl nest known to contain two yoimg. A Short-eared Owl was noticed flying oddly about 100 m north of the doe. The owl was flying in a southerly direction and repeatedly "dive bombing" from a height of 10-15 m to the top of the sage. The owl continued this undulating flight toward the now alert pronghorn. As the owl closely approached, the doe ran through the sagebrush in the same direction as the owl's flight, alternating a head up and head down posture. The latter was coordinated with short bursts of speed. The animals continued this pattern for about 100 m. As they emerged from the sagebrush into a crested wheatgrass (Agropyron crista- tum) planting, a coyote was seen running ahead of the antelope and below the owl. To- gether, they pursued the coyote for nearly 300 m before the coyote reentered the sage- brush. Both the pronghorn and the owl then abandoned the chase. The doe looked in the direction of the coyote for nearly five min- utes, then resumed feeding. The owl circled to a height of about 50 m and began hunting activities. Carrying a prey item, it visited the nest 15 minutes later. Further investigation indicated that the owl's mate had been on or near the nest throughout the joint anti- predatory defense. It is doubtful that the an- telope participating in the chase was pro- tecting a fawn. The earliest record of pronghoms fawning on the Idaho National Engineering Laboratory Site is 23 May (1980), with the peak of fawning normally occurring the last week of May and the first week of June each year. Conclusions Figure 1 is an adaptation of Berger's (1979) antipredatory defense model for ungulates, and represents my proposed spectrum of pronghorn responses to hunting coyotes. The wide solid arrows indicate the responses most likely to occur in pronghorn coyote encoun- ters. Narrow solid lines represent docu- mented responses that occur less often, and the wavy arrows account for the rare obser- vation of concurrent, interspecific chasing. The dashed arrows indicate some possible re- actions of pronghorn to hunting coyotes that were not recorded in my observations. The actions taken by pronghorn when con- fronted by coyotes appear to be generally re- lated to the size and composition of the pronghorn group. Individuals unaccompanied by fawns, or small groups of pronghorn, tend to retreat from coyote predators, often join- ing other bands of pronghorn. Larger groups of pronghorn exhibit a continuum of respon- ses ranging from mild interest, or curiosity, to actual attack that in the broadest context represents mobbing behavior (Harvey and Greenwood 1978). My observations indicate the postparturient does, with fawns nearby, invariably attack or chase coyotes advancing toward them. The intensity of these attacks, and the context in which they occur, closely resembles the antipredatory response of "snarling" described by Curio (1975). The constancy of this behavior supports the thesis that, in certain situations (i.e., does with fawns nearby), predator harassment has di- rect survival value for pronghorn fawns (Lipetz and Bekoff 1980). The significance of the joint (cooperative?) chase by a pronghorn doe and a Short-eared Owl is unknown. There are two plausible interpretations of this event. First, as several instances of pronghorn chasing Short-eared Owls in an antipredatory context have been observed (Fichter, pers. comm., Copeland, in litt.), it is possible that the doe was responding to both the coyote and the owl as potential pred- ators. However, if the doe was not protecting a fawn (the date of this encounter suggests she was not), the adaptive advantage, or evo January 1983 Reynolds: Pronghorn Responses 91 Individual or small group Interspecific cooperation DOE with fawn(s) Attack Fig. 1. A proposed model of the responses of pronghorn antelope to coyotes. See text for the explanation of arrows. lutionary significance of her actions is not ob- vious. On the other hand, as this paper and that by Lipetz and Bekoff (1980) suggests, pronghorn chasing coyotes is not an excep- tionally rare event. Although published re- cords are few. Short-eared Owls have not in- frequently been observed mobbing predatory species, including coyotes (pers. obs., Trost, pers. comm., Clark 1975). It is likely that the antelope-owl-coyote interaction described here represents a mutual, albeit fortuitous, ef- fort by the antelope and owl to hustle the coyote. The proximity of the chase to the owl nest makes the reasons behind the owl's in- volvement obvious. The factors precipitating the antelope's behavior are less certain. Fich- ter (unpublished data) witnessed a buck band of over a dozen pronghorn chase a coyote in mid- June 1965. These animals pursued the predator for 1-1.5 km, passing in front of and 92 Great Basin Naturalist Vol. 43, No. 1 circling the running coyote at least twice, a sequence frequently associated with moving vehicles in pronghom country. It is possible that in certain low-risk situations, such as when a coyote is already fleeing from harass- ment, antelope may participate in the chase as a playlike exercise. This might represent a learning experience for the prey and/ or predator, lending support to any or all of Berger's (1979) explanations for ungulates at- tacking predators. However, the rarity of birds and mammals jointly mobbing a poten- tial predator precludes a convenient analysis of the role of thjs interspecific behavior in the relationship of predators and prey. More data are required before the evolutionary sig- nificance of this and similar observations can be properly assessed. Acknowledgments This is a contribution from the INEL Site Ecological Studies Program, supported by the Office of Health and Environmental Re- search, U.S. Department of Energy. I thank R. E. Autenrieth for his suggestions and M. W. Barrett, P. T. Bromley, E. Fichter, O. D. Markham, J. M. Peek, F. L. Rose, and G. E. Svendsen for improving the drafts of this manuscript. Literature Cited Arbington, O. N., and A. E. Edwards. 1951. Predator control as a factor in antelope management. Trans. N. Am. Wildl. Conf. 16:179-195. Autenrieth, R. E., and E. Fichter. 1975. On the be- havior and socialization of pronghom fawns. Wildl. Monogr. No. 42. Ill pp. Beale, D. R., and a. D. Smith. 1973. Mortality of pronghorn antelope fawns in western Utah. J. Wildl. Manage. 37:343-352. Berger, J. 1979. "Predator harassment" as a defensive strategy in ungulates. Am. Midi. Nat. 102:197-199. Clark, R. J. 1975. A field study of the Short-eared Owl Asio flammeus (Pontoppidan) in North America. Wildl. Monogr. No. 47. 67 pp. Curio, E. 1975. The functional organization of anti- predator behavior in the Pied Flycatcher, a study of avian visual perception. Anim. Behav. 23:1-115. Harvey, P. H., and P. J. Greenwood. 1978. Anti- predator defense strategies: some evolutionary problems. Pages 129-151 in J. R. Krebs and N. B. Davies, eds.. Behavioral ecology: an evolutionary approach. Blackwell Scientific Pubs., Oxford. 494 pp. Lipetz, V. E., and M. Bekoff. 1980. Possible functions of predator harassment in pronghom antelope. J. Mammal. 61:741-743. Thompson, W. K. 1949. Predation on antelope. J. Wildl. Manage. 13:313-314. Udy, J. R. 1953. Effects of predator control on antelope populations. Utah State Dept. Fish and Game Publ. No. 5. 48 pp. FLORISTICS OF THE UPPER WALKER RIVER, CALIFORNIA AND NEVADA Matt Lavin' Abstract.— A checklist of the vascular flora of the upper Walker River is presented. Listed are 1078 taxa from this 4000 km^ area. The upper Walker River encompasses a portion of the boundary between the Intermountain and Sierra Nevada floristic regions, and hence displays much floristic diversity within a relatively small area. Due to its location along the east slope of the Sierra Nevada, the Walker River drainage is unique in that it contains much elevational variation eastward into the Intermountain Region. This elevational extension is due to the presence of large mountain ranges including the Sweetwater Mountains, the Bodie Hills, and the Wassuk Range. As a result of this elevational variation, there is much overlapping of the two floristic regions. Additionally, the easternmost expo- sures of the Sierran granodiorites occur within the Walker River basin and may enhance the eastward migration of Sierran plants. The 90 percent floristic similarity (Sorenson's) between the Sweetwater Mountains, lying to the east of the Sierra, and the east slope of the Sierra Nevada (within the Walker River drainage) indicates the Sweetwaters to be more affiliated with the Sierran flora instead of the Intermountain flora. The upper Walker River drainage includes an area of approximately 4000 km 2, located at the north end of Mono Coimty, California, the southern ends of Douglas and Lyon coun- ties, Nevada, and the western edge of Miner- al County, Nevada. The western boundary of this drainage generally follows the crest of the Sierra Nevada from the Conway Summit- Virginia Lakes area, northward to the Topaz Lake-Monitor Pass area. The eastern bound- ary is delimited by the Bodie, Masonic, and Pine Grove hills and the southwest slopes of the Pine Nut Range. The Sweetwater Moim- tains and the Wellington hills lie between these hills and the Sierra Nevada. East slope Sierran vegetation present here has been generally described by Billings (1951) and Rundel et al. (1977). The area is dominated by mixed conifer forests composed of Pinus jeffreyi and Abies concolor at the low elevations, and Abies magnifica, Tsuga mertensiana, Pinus monticola, and P. albi- caulis at the higher elevations. The alpine vegetation of the Sierra is uniquely adapted for extreme svmimer drought (Chabot and Billings 1972). To the east of the Sierra, Intermountain vegetation (high elevation sagebrush steppe situated above woodlands composed of Pinus monophylla and Juniperus osteosperma) pre- dominates. This vegetation has also been briefly described by Billings (1951). The Sweetwater Mountains, situated between the Intermountain and Sierran floristic regions, display characteristics of both. The vascular flora of the upper Walker River is modestly represented in literature. Works such as Hinton (1975), Reveal (1968), Cox (1972), Reveal and Ertter (1980), Hard- ham and True (1972), Strother (1974), Munz (1968), Bameby (1964), Dempster and Ehren- dorfer (1965), Halse (1981), and others cite specific collections made within this area. Davis (1979) compiled a plant list and keys to the plants occurring on the east central Sierra (Owens Lake to Lake Tahoe) and asso- ciated desert ranges. This work was based solely on distributions given in existing floras. Sharsmith (1940) includes the Walker Riv- er portion of the Sierra Nevada as the north- ernmost boundary of the Sierra alpine floris- tic region. This area includes Leavitt Peak south to Dunderberg Peak. Thome (1982) de- fines the upper Walker River basin as the northern boundary of the transmontane Cali- fornian floristic region. Major and Taylor (1977) conducted a vege- tation study of the alpine zone of the Sweet- water Mountains. They list 43 species from the area. Taylor (1977) indicates a 40-50 'Department of Biology, University of Nevada, Reno, Nevada 89557. Present address: P.O. Box 13494, Reno, Nevada 89507. 93 94 Great Basin Naturalist Vol. 43, No. 1 percent floristic similarity (Sorenson's) be- tween the Carson Pass area of the Sierra Ne- vada and the Sweetwater Mountains. Since a 50 percent turnover rate in plant species oc- curs every 650 km along the Cascade-Sierran axis (Taylor 1977), the Walker River portion of the Sierra must have a very high degree of similarity with Carson Pass, 60 km to the north. An interpretation of this would mean the Walker River portion of the Sierra has approximately a 50 percent floristic sim- ilarity to the Sweetwaters. Bell (1980) described the alpine flora of the Wassuk Range, located in the lower Walker River drainage just west of Walker Lake (50 km east of the Sweetwater Mountains). She lists 70 vascular plants from the alpine flora of this region. Bell has also recently com- pleted a study of the alpine flora of the Sweetwater Mountains, but this information is unavailable. Messick (1982) completed a flora of the Bodie Hills, and the results of this work are incorporated in Table 1 . The upper Walker River drainage is the region of several type collections. The Sweet- water Mountains are the type locale for Cordylanthus ramosus ssp. setosus, Senecio pattersonensis and Draba lemmonii var. in- crassata. In the Bodie Hills, we have the type locale for Draba quadricostata, Pinus mon- ophylla, Arabis bodiensis (see Rollins 1982), Streptanthus oliganthus, and Phacelia mon- oensis. A. bodiensis, S. pattersonensis, D. lem- monii var. incrassata, and D. quadricostata are endemic to the Walker River drainage. The Wellington Hills are the type locale for Astragalus oophorus var. lavinii, and So- nora Pass for Raillardella argentea, Cymop- terus cinerarius, and Wyethia mollis. "A dry rocky mountain near Sonora Pass" is the type location for Astragalus platytropis, A. lentigi- nosus var. ineptus, and A. whitneyi. Bameby suggests that the type locale for these three Astragali may be the Sweetwater Moujitains. However, during the course of this in- vestigation, all three, including A. platy- tropis, were found on Leavitt and Emma Peak of the Sierra Nevada. Therefore, the type locale given by Gray could have possi- bly referred to the Sierra instead of the Sweetwaters. Methods The checklist (Table 1) was developed from collections made during the course of this study: July 1979 to August 1982. Addi- tional collections were recorded from the herbaria at the University of Nevada, Reno (RENO), The New York Botanical Garden (NY), and the California Academy of Sciences (CAS). Collections from the upper Walker River made by Arnold Tiehm and Margaret Williams, Reno, Nevada; Steve Wharff, Tonopah, Nevada; Frank Smith, Smithfield, Utah; Dennis Breedlove, CAS; Joe Robertson, E. F. Kleiner, Tom Lugaski, Pat and Ham Vreeland, H. N. Mozingo, and Fred Ryser, all from the University of Nevada, Reno, were also recorded in this checklist. Approximately 3850 numbers were recorded. Distributional information contained within the checklist was continually refined while in the field. Various documentary works, mentioned in the introduction, were used to determine those plants that have a probable distribution within the upper Walker River drainage, but were not observed during this study. Almost all of the taxa listed can be found on deposit at the University of Nevada, Reno, herbarium and the Toiyabe National Forest Supervisor's Office, Reno, Nevada. The purposes of the checklist (Table 1) are to both document the flora present in the up- per Walker River and to document the geo- graphical and altitudinal distribution of each taxon within this area. Nomenclature gener- ally follows Kartesz and Kartesz (1980). Results and Discussion Table 1 lists 1078 taxa of vascular plants from the upper Walker River. The upper Walker River drainage is unique in that it ex- tends much elevational variation of the east slope of the Sierra Nevada eastward into the Intermountain Region, due to the presence and close proximity of such large mountain ranges as the Sweetwater Mountains, the Bodie Hills, and the Wassuk Range. Along any floristic boundary there is bound to be some overlap of unique plant species or char- acteristic vegetation of one flora into anoth- er. However, many plants having their center of distribution in the Intermountain Region January 1983 Lavin: Walker River Floristics 95 also have outlying populations in the mon- tane environments of the Sierra Nevada. This phenomenon is discussed by Taylor (1976) for the Carson Pass area of the Sierra. He attri- butes the occurrence of many Intermountain plants on the east slope of the Sierra to Xero- thermic invasion. A list of these plants at the headwaters of the Walker River follow: Agoseris glauca var. monticola Allium biceptrum Allium parvum Arabis bodiensis A. pulchra var. pulchra Artemisia arbuscula Aster ascendens Astragalus platytropis A. iodanthus Amelanchier utahensis Antennaria dimorpha Balsamorhiza sagittata Calyptridium roseum Cercocarpus ledifolius Cirsium utahense Cheilanthes gracillima Chenopodium overi Chorizanthe brevicomu var. spathulata Chrysothamnus nauseosus ssp. albicaulis C. viscidiflorus Crepis acuminata C. modocensis ssp. subacaulis Castilleja linariifolia Cordylanthus ramosus ssp. setosus Cryptantha circumscissa Cryptantha echinella Erigeron aphanactis E. nevadincola E. breweri var. porphyreticus E. eatonii ssp. plantagineus Eriogonum elatum E. microthecum var. ambiguum E. ovalifolium var. nevadense E. ivrightii var. subscaposum Galium multiflorum Gilia leptantha ssp. salticola Grayia spinosa Haplopappus acaulis Heuchera duranii Hydrophyllum capitatum var. alpinum Leptodactylon pungens Linanthus nuttallii Lupinus caudatus ssp. caudatus L. nevadensis Lomatium nevadense var. nevadense Lomatium nevadense var. parishii Melica striata Mentzelia congesta Mimulus densus M. rubellus Navarretia breweri Opuntia polyacantha var. rufispina Poa nevadensis var. juncifolia P. nevadensis var. nevadensis Paeonia brownii Penstemon bridgesii Phoenicaulis cheiranthoides Phacelia humilis Phlox covillei Plagiobothrys hispidus P. kingii var. harknessii Prunus andersonii Purshia tridentata Pinus monophylla Ribes velutinum Rosa woodsii var. ultramontana Scrophularia desertorum Senecio canus S. pattersonensis S. spartioides Sisyrinchium halophilum Sphaeromeria cana Stephanomeria spinosa Streptanthus oliganthus Tetradymia canescens Thelypodium crispum Zigadenus paniculatus These 77 plant taxa represent approx- imately 10 percent of the flora present on the east slope of the Sierra within the Walker River drainage. This is below Taylor's (1976) estimate of 20 percent for the east slope within the Carson River drainage just to the north. However, when distribution records become more complete, it would not be sur- prising to find 20 percent of the Sierran flora within the Walker River basin being com- posed of Intermountain elements. The Xerothermic climate may also have been responsible for the northern migration of Amelancheir pallida var. covillei, Cea- nothus greggii var. vestitus, Cercocarpus ledi- folius var. intricatus, Ivesia purpurascens ssp. congdonis, Cryptantha confertiflora, Phacelia peirsoniana, and Plagiobothrys jonesii. The Walker River drainage may be the north- ernmost location for these plants. {Cryp- tantha confertiflora has been fovmd just to the north in the Carson River drainage.) It is documented that the eastward migra- tion of Sierran plant species into the Inter- mountain Region is small relative to the westward migration of Rocky Mountain plants into this region (Harper et al. 1978). However, the Walker River drainage has many Sierra or Pacific cordilleran plant spe- cies occurring well into the Intermountain Region, some of which occur as far east as Masonic Mountain or the Wassuk Range. A list of these plants follows: Agropyron pringlei Allium campanulatum 96 Great Basin Naturalist Vol. 43, No. 1 Allophyllum gilioides A. violaceum Amelanchier pallida Anelsonia eurycarpa Angelica lineariloba Arabis inyoensis A. platysperma var. howellii Arnica nevadensis Astragalus kentrophyta var. danaus A. lentiginosus var. ineptus A. purshii var. lectulus Calochortus leichtlinii Carex tahoensis Chaenactis nevadensis Chrysothamnus parryi ssp. monocephalus Claytonia nevadensis Cryptantha glomeriflora Cryptantha nubigena Cymopteris cinerarius Draba lemmonii (var. incrassata) D. oligosperma var. subsessilis D. stenoloba var. ramosa Erigeron petiolaris E. pygmaeus Eriogonum rosense Gentianopsis hohpetala Gentiana newberryi Geum canescens Galium hypotrichium ssp. hypotrichium Haplopappus apargioides Hieracium hcrridum Ivesia lycopodioides I. purpurascens ssp. congdonis Juniperus occidentalis ssp. australis Kalmia microphylla Ledum glandulosum var. californica Leucophysalis nana Lupinus andersonii L. caudatus ssp. montigenus L. confertus L. hypolasius L. meionanthus L. caudatus ssp. montigenus L. sellulus var. lobbii L. tegeticulatus (breweri bryoides) Luzula divaricate Mimulus coccineus Penstemon davidsonii var. davidsonii Pinus jeffreyi P. monticola Polygonum douglasii var. latifolia Raillardella argentea Rhamnus rubra ssp. rubra Scirpus dementis Senecio fremontii var. occidentalis S. scorzonella Sisyrinchium idahoense var. occidentale Symphoricarpos parishii Trifolium andersonii ssp. andersonii Additionally, work done by Goodrich (1981) in central Nevada has shown that sev- eral other Pacific cordilleran plants extend their ranges far into the Intermountain Re- gion. These plants, also found extending east- ward within the Walker River basin, include Artemisia rothrockii, Carex helleri, Silene sar- gentii, Astragalus whitneyi (see Reveal 1979) and Astragalus purshii var. tinctus (see Bam- eby 1964). Silene sargentii has been listed as endemic to the Sierra Nevada. However, the type collection, as given by Hitchcock and Maguire (1947), is from Table Mountain of the Monitor Range in central Nevada. The eastward distribution of these Sierran plants may be enhanced by the very eastern exposures of the Sierran granodiorites. These granites can be found as far east as the Was- suk Range (Bateman 1967). However, the most likely factor enhancing the migration of these Sierran plants is the abundance of mon- tane and alpine habitats found eastward into the Intermountain Region. The floristic com- ponents of the Sweetwater Moujitains point to this eastward migration. The Sweetwaters, situated between the Sierra and Intermountain floristic regions, are regarded as belonging to the Intermountain flora (Cronquist et al. 1972). Sierra conifer forests are extensive on the Sweetwaters, es- pecially toward the southern end. The tim- berline vegetation is dominated solely by Pinus albicaulis, a Sierran characteristic. Using the information provided in Table 1, Sorenson's index of similarity (Billings 1978) between the Sierra and Sweetwaters can be determined with regard to floristic elements in both the alpine and the montane conifer forests. These are calculated to be 90 and 93 percent, respectively. This indicates the Sweetwater Mountains to be more affiliated, floristically, with the Sierra than previously thought. However, it might be considered that the east slope of the Sierra, within the Walker River drainage, being influenced by Inter- mountain vegetation, is bound to yield a high index of similarity to the Sweetwaters. In other words, the whole of the upper Walker could be considered "Intermountain" with regard to vegetational composition. This is reinforced by the fact that the Walker River portion of the Sierra does lack typical "un- derstory" brush vegetation that is found just to the north in the Carson River drainage. Arctostaphylos patula, Ceanothus cordulatus January 1983 Lavin: Walker River Floristics 97 and C. prostratus have not been found here (or are at least not abundant) and A. neva- densis was foujid only twice in very small populations. Instead, the dominant brush cover consists of Purshia tridentata, Cea- nothus velutinus, Artemisia tridentata ssp. vaseyana, and Symphoricarpos oreophilus. In comparing all the upper Walker River flora with the flora of central Nevada (Good- rich 1981), both floras being just about equal in number of taxa, approximately a 50 per- cent similarity can be determined. The Sweetwaters could, therefore, be easily con- sidered as part of the east slope Sierran flora. Aside from the high indices of similarity, the Sweetwaters are dominated by many Sierra or Pacific cordillera plant species; Lupinus hypolasius, Chrysothamnus parryi ssp. mon- ocephalus, and Draba oligosperma var. sub- sessilis at the highest elevations, and Pinus jeffreyi and P. contorta var. murrayana at the lower elevations. Acknowledgments This study was funded by the U.S. Forest Service, Toiyabe National Forest. I am very grateful to them for this and for their cooper- ation. Thanks to Sue Sullivan, U.S. Forest Service, the checklist (Table 1) and its trans- fer to Brigham Young University Press was made possible. For their help with identi- fication of plant specimens, I am indebted to Warren Wagner (Botrychium), Lincoln Con- stance (Apiaceae), Ted Barkley (Senecio), Al- mut Jones (Aster), Guy Nesom (Erigeron), Gerald Ownbey (Cirsium and Argemone), Reed Rollins (Brassicaceae), Arthur Cronquist (miscellaneous), Alfred Schuyler (aquatic plants), Gary Wallace (Pyrola), Rupert Barn- eby (Fabaceae), Richard Halse (Phacelia), Duane Atwood (Phacelia), Douglass Hender- son (Sisyrinchium), Paul Fryxell (Malvaceae), David Boufford (Circaea), Alva Day (Polemo- niaceae), Lauramae Dempster (Galium), Har- lan Lewis (Gayophytum), Lawrence Heckard (Castilleja, Cordylanthus, Orthocarpus), Fred- rick Meyer (Valeriana), John T. Howell (Carex and miscellaneous), Margaret Wil- liams (miscellaneous), Arnold Tiehm (mis- cellaneous), and Ken Genz (miscellaneous). I am especially grateful to John Thomas How- ell, Margaret Williams and Arnold Tiehm for sharing with me their intimate knowledge of the Intermountain and Sierran floras, and to the curators at NY and CAS for allowing ac- cess to their herbaria. Literature Cited Barneby, R. C. 1964. Atlas of North American Astra- galus. New York Botanical Gardens. 2 Vols. Bateman, p. C. 1967. The Sierra Nevada Batholith. Sci- ence 158: 1407-1417. Bell, K., and R. Johnson. 1980. Alpine flora of the Wassuk Range, Mineral County, Nevada. Madroiio 27: 25-35. Billings, W. D. 1951. Vegetation zonation in the Great Basin of western North America. Pages 101-122 in Les Bases e regeneration do la des zones arides. International Colloquium. Intemat. Union of Bio. Sci. Ser. B.9. 1978. Alpine phytogeography across the Great Basin. Great Basin Nat. Mem. 2: 105-117. Chabot, B. F., and W. D. Billings. 1972. Origins and ecology of the Sierran alpine flora and vegeta- tion. Ecol. Monogr. 42: 163-199. Cox, B. J. 1972. Biosystematics of Lupinus lepidus-L. caespitosus complex. Unpublished dissertation. Univ. of Missouri, Columbia. 443 pp. Cronquist, A., A. Holmgren, N. Holmgren, and J. Reveal. 1972. Intermountain flora. Vol. 1. Haf- ner. New York. 584 pp. Davis, B. 1979. Draft: Plant list and keys. East Side Cen- tral Sierra and associated desert Ranges. U.S. Forest Service, Toiyabe National Forest, Reno, Nevada. 124 pp. Dempster, L. T., and F. Ehrendorfer. 1965. Evolution of the Galium multiflorum complex in western North America. II. Critical taxonomic revision. Brittonia 17: 289-334. Goodrich, S. 1981. A floristic study of central Nevada. Unpublished thesis, Brigham Young Univ. 400 pp. Halse, R. R. 1981. Taxonomy of Phacelia Sect. Miltitzia (Hydrophyllaceae) Madrono 28: 121-132. Hardham, C. B., and G. H. True. 1972. Malacothrix tor- reyi (Compositae) in California. Madrono 21: 535. Harper, K. T., D. C. Freeman, W. K. Ostler, and L. C. Klikoff. 1978. The flora of the Great Basin mountain ranges: diversity, sources and dispersal ecology. Great Basin Nat. Mem. 2: 81-103. HiNTON, W. F. 1975. Systematics of the Calyptridium umbellatum complex (Portulacaceae). Brittonia 27: 197-208. Hitchcock, C. L., and B. Maguire. 1947. A revision of the North American species of Silene. Univ. of Washington Pub. Biol. 13: 1-73. Kartesz, J., AND R. Kartesz. 1980. A synonymized checklist of the vascular flora of the United States, Canada and Greenland. Univ. of North Carolina Press. Chapel Hill. 500 pp. Lavin, M. 1981. Floristics of the headwaters of the Walker River, California and Nevada. Unpub- lished thesis. Univ. of Nevada, Reno. 141 pp. Major, J., and D. W. Taylor. 1977. Alpine. Pages 601-675 in M. G. Barbour and J. Major, eds.. 98 Great Basin Naturalist Vol. 43, No. 1 Terrestrial vegetation of California. John Wiley Interscience, New York. Messick, T. 1982. The flora and phytogeography of the Bodie Hills of Mono County, CA and Mineral County, NV. Unpublished thesis, Humboldt State Univ. MuNZ, P. A., AND D. Keck. 1968. A California flora. Univ. of California Press. Berkeley. 1681 pp. and supplement. Reveal, J. L. 1968. Notes of Eriogonum— IV. A revision of the Eriogonum deflexum complex. Brittonia 20: 13-33. 1979. Biogeography of the Intermountain Re- gion. Mentzelia 4: 1-92. Reveal, J. L., and B. Ertter. 1980. Noteworthy collec- tions. Madrono 27: 142. Rollins, R. C. 1982. Studies on Arahis (Cruciferae) of Western North America II. Contr. Gray Her- barium of Harvard. No. 212:111-114. RuNDEL, P. W., D. J. Parsons, and D. T. Gordon. 1977. Montane and subalpine vegetation of the Sierra Nevada and Cascade ranges. Pages 559-599 in M. G. Barbour and J. Major, eds., Terrestrial vegeta- tion of California. John Wiley Interscience, New York. Sharsmith, C. W. 1940. A contribution to the history of the alpine flora of the Sierra Nevada. Unpub- lished dissertation. Univ. of California, Berkeley. 274 pp. Strother, J. L. 1974. Taxonomy of Tetradymia (Com- positae: Senecioeae). Brittonia 26: 177-202. Taylor, D. W. 1976. Disjunction of Great Basin plants in the northern Sierra Nevada. Madrono 23: 301-310. 1977. Floristic relationships along the Cascade- Sierran axis. Amer. Midi. Natur. 97: 333-349. Thorne, R. F. 1982. The desert and other transmontane plant communities of southern California. Aliso 10:219-257. Table 1. Checklist of the vascular flora of the upper Walker River. For the columns under the heading of "AREA":N = Sierra Nevada; S= Sweetwater Mountains; W = Wellington Hills; M = Masonic Hills, Bodie Hills, Pine Grove Hills, and the southwest slopes of the Pine Nut Range. For the columns under the heading of "HABITAT": 1 = riparian; 2= low elevation sagebrush-grass zone; 3=pinyon-juniper woodland; 4 = high elevation sagebrush -grass zone; 5 = Jeffrey pine-white fir forests; 6 = red fir forests; 7 = lodgepole pine forests; 8 = whitebark pine forests; 9= al- pine zone (see Billings 1951, Lavin 1981, for a general description of these). An X indicates the plant has been ob- served in the field or by herbarium specimen; an O indicates the plant has a probable distribution within the area due to information obtained in the literature. AREAS HABITATS N S W M 123456789 Adiantaceae - pteridophyta Aspidotis densa (Brack.) Lellinger Cheilanthes gracillima D.C. Eat. Cryptogramma crispa (L.) R. Br. ex Hook. ssp. acrostichoides (R. Br.) Hulten Pellaea breweri D.C. Eat. bridgesii Hook. Aspleniaceae Athyrium distentifolium Tausch ex Opiz var. americanum (Butters) Boivin Cystopteris fragilis (L.) Bemh. Woodsia oregana D.C. Eat. scopulina D.C. Eat. Dennstaedtiaceae Pteridium aquilinum (L.) Kuhn var. pubescens Underwood Equisetaceae Equisetum arvense L. hyemale L. var. affine (Engelm.) A. A. Eat. laevigatum A. Braun X X X X X X X X X X X X X o X X o 0 X X X X X X X X X X X X X X X X X X X X X X XXX X XX X X X o X XX XXXX X XXX January 1983 Lavin: Walker River Floristics 99 Table 1 continued. AREAS HABITATS N S W M 12 3 4 5 7 8 9 ISOETACEAE Isoetes bolanderi Engelm. var. bolanderi Marsileaceae Marsilea vestita Hook. & Grev. Ophioglossaceae Botrychium lunaria (L.) Sw. var. minganese (Victorin) Dole simplex E. Hitchcock Selaginellaceae Selaginella watsonii Underwood CUPRESSACEAE - PINOPHYTA Juniperus communis L. occidentalis Hook. ssp. australis Vasek osteosperma (Torr.) Little Ephedraceae Ephedra nevadensis S. Wats. viridis Coville Pinaceae Abies concoloT (Gord. & Glend.) Hildebr. magnifica A. Murr. Pinus albicaulis Engelm. contorta Dougl. ex Loud. var. murrayana (Grev. & Balf.) Engelm. flexilis James jeffreyi Grev. & Balf. monophylla Torr. & Frem. monticola Dougl. ex D. Don Tsuga mertensiana (Bong.) Carr. AcERACEAE - ANTHOPHYTA Acer glabrum Torr. var. torreyi (Greene) Smiley Alismataceae Sagittaria cuneata Sheldon Amaranthaceae Amaranthus alhus L. blitoides S. Wats. retroflexus L. Apiaceae Angelica breweri Gray lineariloba Gray X X X X XX o XX o X X X X X X X X X X X X X X X X X X X X XX X XXX XXX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XXX X O X X X X X O XX XXX XXX X XX XXX XXX o o o X X X X X X X X X o X X 100 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS N S W M HABITATS 123456789 Berula erecta (Huds.) Coville Cicuta douglasii (DC.) Coult. & Rose Conium maculatum L. Cymopterus cinerarius Gray globosus (S. Wats.) S. Wats. panamintensis Coult. & Rose var. panamintensis Heracleum lanatum Michx. Ligusticum grayii Coult. & Rose Lonuitiiim dissectum (Nutt.) M. & C. var. multifidum (Nutt.) M. & C. foeniculaceum (Nutt.) Coult. & Rose ssp. macdougalii (Coult. & Rose) Theobald nevadense (S. Wats.) Coult. & Rose var. nevadense var. parishii (Coult. & Rose) Jepson plummerae (Coult. & Rose) Coult. & Rose var. sonnei (Coult. & Rose) Jepson Osmorhiza chilensis Hook. & Am. occidentalis (Torr. & Gray) Torr. Perideridia bolanderi (Gray) A. Nels. & J.F. Macbride ssp. bolanderi lemmonii (Coult. & Rose) Chuang & Const. parishii (Coult. & Rose) A. Nels. ssp. latifolia (Gray) Chuang & Const. Podistera nevadensis (Gray) S. Wats. Pteryxia terebinthina (Hook.) Coult. & Rose var. californica (Coult. & Rose) Mathias Sphenosciadium capitellatum Gray Apocynaceae Apocynum androsaemifolium L. ssp. pumilum (Gray) Boivin X medium Greene ASCLEPIADACEAE Asclepias cryptoceras S. Wats. ssp. cryptoceras ssp. davisii (Woods.) Woods. fascicularis Dene. speciosa Torr. ASTERACEAE Achillea millefolium L. var. alpicola (Rydb.) Garrett var. lanulosa (Nutt.) Piper X X X X X X X X X o X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X o o X X o o XXXX XXX X XXXX X xxxxxx XXX O X 0 o XXX X X X X X X XXXX XXX XX X XXXX X X XXX XXX January 1983 Lavin: Walker River Floristics 101 Table 1 continued. AREAS HABITATS N S w M 1 2 3 4 5 6 7 8 9 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X Agoseris aurantiaca (Hook.) Greene elata (Nutt.) Greene glauca (Pursh) Raf. var. nwnticola (Greene) Cronquist var. laciniata (D.C. Eat.) Smiley Ambrosia acanthicarpa Hook. Anisocoma acaulis Torr. & Gray Antennaria alpina (L.) Gaertn. var. media (Greene) Jepson dimorpha (Nutt.) Torr. & Gray micTophylla Rydb. umbrineUa Rydb. Arnica chamissonis Less. ssp. foliosa (Nutt.) Maguire var. andina (Nutt.) Ediger & Barkley ssp. foliosa (Nutt.) Maguire var. incana (Gray) Hulten cordifolia Hook. var. cordifolia diversifolia Greene longifolia D.C. Eat. mollis Hook. nevadensis Gray parry i Gray var. sonnei (Greene) Cronquist sororia Greene Artemisia arbuscula Nutt. cana Pursh dougtasiana Bess. dracunculus L. ludoviciana Nutt. ssp. incompta (Nutt.) Keck ssp. ludoviciana norvegica Fries var. saxitalis (Bess.) Hook. nova A. Nels. rothrockii Gray spinescens D.C. Eat. tridentata Nutt. ssp. vaseyana (Rydb.) Beetle ssp. wyomingensis Beetle & Young ssp. tridentata Aster ascendens Lindl. in Hook. alpigenus (Torr. & Gray) Gray ssp. andersonii (Gray) Onno campestris Nutt. var. bloomeri Gray eatonii (Gray) T.J. Howell integrifolius Nutt. occidentalis (Nutt.) Torr. & Gray var. occidentalis scopuhrum Gray X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o o 0 X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X o o X X X X X X X X X X X X X X X X X X X X X X X X X X X 102 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats N S W M 12 3 4 5 6 7 Bahamorhiza hookeri (Hook.) Nutt. sagittata (Pursh) Nutt. Bidens cemua L. Brickellia grandiflora (Hook.) Nutt. var. petiolaris Gray microphylla (Nutt.) Gray oblongifolia Nutt. var. linifolia (D.C. Eat.) B.L. Robins. Chaenactis alpigerm C.W. Sharsmith douglasii (Hook.) Hook. & Am. var. rubricaulis (Rydb.) Ferris nevadensis (Kellogg) Gray xantiana Gray Chrysothamnus nanseosus (Pall.) Britton ssp. consimilis (Greene) Hall & Clem, ssp. albicaulis (Nutt.) Hall & Clem, ssp. hololeucus (Gray) Hall & Clem. parryi (Gray) Greene ssp. monocephalus (Nels. & Kenn.) Hall & Clem, ssp. nevadensis (Gray) Hall & Clem. viscidiflorus (Hook.) Nutt. ssp. puberulus (D.C. Eat.) Hall & Clem, ssp. viscidiflorus Cichorium intybus L. Cirsium andersonii (Gray) Petrak congdonii Moore & Frankton eatonii (Gray) B.L. Robins. pastoris J.T. Howell tioganum (Congd.) Petrak utahense Petrak vulgare (Savi) Tenore Conyza canadensis (L.) Cronquist var. glabrata (Gray) Cronquist Crepis acuminata Nutt. ssp. acuminata intermedia Gray modocensis Greene ssp. subacaulis (Kell.) Babcock & Stebbins nana Richards. ssp. ramosa Babcock & Stebbins ssp. nana occidentalis Nutt. ssp. pumila (Rydb.) Babcock & Stebbins ssp. occidentalis ssp. conjuncta (Jepson) Babcock & Stebbins runcinata (James) Torr. & Gray ssp. hallii Babcock & Stebbins Dugaldia hoopesii (Gray) Rydb. XXX X X X X X X XXX XX X XX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 0 X o o X X X X X X X 0 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X January 1983 Lavin: Walker River Floristics 103 Table 1 continued. AREAS HABITATS NSWM 123456789 Eatonella nivea (D.C. Eat.) Gray Erigeron aphanactis (Gray) Greene var. aphanactis barbellulatus Greene bloomeri Gray var. bhameri breweri Gray var. breweri var. porphyretictis (M.E. Jones) Cronquist clokeyi Cronquist compositus Pursh var. glabratus Macoun coulteri Porter divergens Torr. & Gray eatonii Gray ssp. plantagineus (Greene) Cronquist lonchophyllus Hook. nevadincola Blake peregrinus (Pursh) Greene ssp. callianthemus (Greene) Cronquist var. angustifolius (Gray) Cronquist ssp. callianthemus (Greene) Cronquist var. hirsutus Cronquist petiolaris Greene pygmaeus (Gray) Greene tener (Gray) Gray vagus Payson Eriophyllum lanatum (Pursh) Forbes var. integrifolium (Hook.) Smiley Eupatorium occidentale Hook. Glyptopleura marginata D.C. Eat. Gnaphalium palustre Nutt. microcephalum Nutt. var. thermale (E. Nels.) Cronquist Grindelia squarrosa (Pursh) Ehinal var. squarrosa Gutierrezia sarothrae (Pursh) Britt. & Rusby Haplopappus acaulis (Nutt.) Gray apargioides Gray bloomeri (Hook.) Gray lanceolatus (Hook.) Torr. & Gray var. lanceolatus macronema Gray racemosus (Nutt.) Torr. ssp. glomeratus (Nutt.) Hall suffruticosus (Nutt.) Gray uniflorus (Hook.) Torr. & Gray ssp. uniflorus X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X o X X X o 0 X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o o X X X X X X X X X X X X X o o X X X X o o X X X X X X X X 104 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats NSW M 1 2 3 4 5 6 7 8 9 X X X X X X X X X X X 0 X X X X X X X X X X X o X X o 0 X X X X X X X X X XXX X X X X XXX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X Helianthus annuus L. Heterotheca breweri AGray) Shinners Hieracium albiflorum Hook. gracile Hook, var. gracile horridum Fries Hulsea algida Gray heterochroma Gray Hymenopappus filifolius Hook. var. nanus (Rydb.) B.L. Turner Hymenoxys coaperi (Gray) Cockerell var. canescens (D.C. Eat.) Parker Iva axillaris Pursh Lactuca serriola L. tartarica (L.) C.A. May ssp. pulchella (Pursh) Stebbins Layia glandulosa (Hook.) Hook. & Am. ssp. glandulosa Leucanthemum vulgare Lam. Machaeranthera canescens (Pursh) Gray shastensis Gray var. montana (Greene) Cronquist & Keck var. gossophylla (Piper) Cronquist & Keck Madia ghmerata Hook. gracilis (Sm.) Keck Malacothrix sonchoides (Nutt.) Torr. & Gray var. torreyi (Gray) E. WiUiams Microseris lindleyi (DC.) Gray Nothocalais alpestris (Gray) Chambers Psihcarphus brevissimus Nutt. var. brevissimus Raillardella argentea (Gray) Gray scaposa (Gray) Gray Senecio canus Hook. cymbalarioides Beuk fremontii Torr. & Gray var. occidentalis Gray hydrophilus Nutt. integerrimus Nutt. var. exaltatus (Nutt.) Cronquist multilobatus Torr. & Gray ex Gray X X o X X o o o X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X January 1983 Lavin: Walker River Floristics 105 Table 1 continued. AREAS HABITATS N s w M 1 2 3 4 5 6 7 8 9 X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X o o X X o X X X X X X X X X X X X X X X X X X X X X X X X X X pattersonensis Hoover scorzonelUi Greene serra Hook, var. serra spartioides Torr. & Gray streptanthifolius Greene triangularis Hook. wemeriifolius (Gray) Gray Solidago canadensis L. var. salebrosa (Piper) M.E. Jones multiradiata Ait. occidentalis (Nutt.) Torr. & Gray spectahilis (D.C. Eat.) Gray Sonchus asper (L.) Hill Sphaeromeria cana (D.C. Eat.) Heller potentilloides (Gray) Heller var. potentilloides Stephanomeria exigua Nutt. ssp. coronaria (Greene) Gottlieb ssp. exigua spinosa (Nutt.) S. Tomb Taraxacum officinale Weber Tetradymia axillaris A. Nels. var. longispina ( M.E. Jones) Strother canescens DC. glahrata Torr. & Gray spinosa Hook. & Am. tetrameres (Blake) Strother Townsendia condensata Parry ex Gray scapigera D.C. Eat. Tragopogon dubius Scop. Wyethia mollis Gray Betulaceae Alnus incana (L.) Moench ssp. tenuifolia (Nutt.) Breittung BORAGINACEAE Amsinckia tessellata Gray Cryptantha affinis (Gray) Greene circumscissa (Hook. & Am.) Johnston var. hispida (J.F. Macbr.) Johnston var. circumscissa confertiflora (Greene) Payson echinella Greene flavoculata (A. Nels.) Payson glomeriflora Greene humilis (Gray) Payson var. humilis X X X X X X X X X o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 106 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats N S W M 123456789 jamesii (Torr.) Payson var. abortiva (Greene) Payson nuhigena (Greene) Payson pterocarya (Torr.) Greene var. pterocarya torreyana (Gray) Greene var. torreyana var. pumila (Heller) Johnston watsonii (Gray) Greene sp. nov. Hackelia floribunda (Lehm.) Johnston micrantha (Eastw.) J.L. Gentry Lappula redowskii (Homem.) Greene var. redowskii Mertensia oblongifolia (Nutt.) G. Don var. nevadensis (A. Nels.) L.O. Williams Pectocarya setosa Gray Plagiobothrys hispidus Gray jonesii Gray kingii (S. Wats.) Gray var. harknessii (Greene) Jepson var. kingii scouleri (Hook. & Am.) Johnston var. scouleri Tiquilia nuttallii (Benth. ex Hook.) Richards. Brassicaceae Anelsonia eurycarpa (Gray) Macbr. & Payson Arabis bodiensis Rollins cobrensis M.E. Jones davidsonii Greene divaricarpa A. Nels. drummondii Gray fernaldiana Rollins var. stylosa (S. Wats.) Rollins glabra (L.) Bemh. hirsuta (L.) Scop. var. glabrata Torr. & Gray holboellii Homem. var. penduhcarpa (A. Nels.) Rollins var. pinetorum (Tidestrom) Rollins var. retrofracta (Grahm.) Rydb. var. holboellii inyoensis Rollins lemmonii S. Wats. var. lemmonii var. depauperata (A. Nels. & Kenn.) Rollins lyallii S. Wats. var. lyallii platysperma Gray var. platysperma var. howellii (S. Wats.) Jepson X X X X X X X X X X X X X o X X X X X X X X X X X X X X X o o X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XXX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X o X X X X X X X X X X XXX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o o X X o X X X X X X X X X X X X X X January 1983 Table 1 continued. Lavin: Walker River Floristics 107 puberula Nutt. pulchra M.E. Jones ex S. Wats, var. pulchra var. gracilis M.E. Jones sparsiflora Nutt. var. subvillosa (S. Wats.) Rollins var. sparsiflora Barbarea orthoceras Ledeb. var. dolichocarpa Fern, var. orthoceras Capsella bursa-pastoris (L.) Medic. Cardamine breweri S. Wats, var. breweri Cardaria pubescens (C.A. Mey) Jarmolenko Caulanthus pilosus S. Wats. Descurainia califomica (Gray) O.K. Schulz pinnata (Walt.) Britt. ssp. filipes (Gray) Detling ssp. halictorum (Cockerell) Detling ssp. menziesii (DC.) Detling richardsonii (Sweet) O.E. Schulz ssp. incisa (Engelm.) Detling ssp. viscosa (Rydb.) Detling sophia (L.) Webb ex Prantl Draba albertina Greene breweri S. Wats. densifolia Nutt. douglasii Gray var. crockeri (Lemmon) C.L. Hitchc. lemmonii S. Wats, var. lemmonii var. incrassata Rollins oligosperma Hook. var. subsessilis (S. Wats.) O.E. Schulz var. oligosperma paysonii J.F. Macbride quadricostata Rollins stenoloba Ledeb. var. ramosa C.L. Hitchcock Erysimum argillosum (Greene) Rydb. capitatum (Dougl.) Greene perenne (S.Wats, ex Coville) Abrams repandum L. Hymenolobus procumbens (L.) Nutt. ex Torr. & Gray Lepidium lasiocarpum Nutt. perfoliatum L. virginicum L. var. pubescens (Greene) C.L. Hitchc. AREAS ] HABITATS ) N S W M 1 2 3 4 5 6 7 8 9 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XXXX X X X X XXXX X X X XXX XXXX XXX XXXX X xxxxxxx o o x x XXXX X XXXX XXXX XX X X X X XXXX X X o o XX O O O X X XXXX xxxxxxxx X X O X XXX X X O XX XXXX XXX XXX X XX X X X X X o XXX X X X X X X X X X X o o o X X X o XXX o XXXX XXX X XXX X XXX XX XX X X XX XXXX XXXX XXXX XXXX 108 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats N S W M 123456789 Lesquerella kingii S. Wats, var. kingii Nasturtium officinale R. Br. Phoenicaulis cheiranthoides Nutt. Polyctenium fremontii (S. Wats.) Greene Rorippa curvisiliqua (Hook.) Bess, ex Britton teres (Michx.) R. Stuckey Sisymbrium altissimum L. Stanleya pinnata (Pursh) Britton var. pinnata Streptanthus cordatus Nutt. ex Torr. & Gray oliganthus Rollins tortuosus Kellogg var. orbiculatus (Greene) Hall Thelypodium crispum Greene ex Payson integrifolium (Nutt.) Endl. ssp. complanatum Al-Shehbaz laciniatum (Hook.) Endl. Cactaceae Opuntia polyacantha Haw. var. rufispina (Engelm. & Bigelow) Benson pulchella Engelm. Callitrichaceae CalUtriche heterophylla Pursh emend. Darby var. bolanderi (Hegelm.) Fassett. verna L. emend. Kuetz. Campanulaceae Nemacladus rigidus Curran Porterella carnosula (Hook. & Am.) Torr. Capparidaceae Cleomella hillmanii A. Nels. parviflora Gray Caprifoliaceae Lonicera involucrata (Rich.) Banks ex Spreng. Sambucus caerulea Raf. racemosa L. ssp. pubens (Michx.) House var. microbotrys (Rydb.) Kearney & Peebles Symphoricarpos longifloTus Gray X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o 0 o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XX X X X X 0 X X X X X X o X X X X X X X X X o o o X X X X X X X X X X X X X X X X X X X X X X o X X January 1983 Lavin: Walker River Floristics 109 Table 1 continued. oreophilus Gray var. oreophilus parishii Rydb. Caryophyllaceae Arenaria aculeate S. Wats. Cerastium alpinum L. vulgatum L. Minuartia nuttallii (Pax) Briq. ssp. fragilis (Maguire & Holmgren) McNeill ssp. gracilis (B.L. Robins.) McNeill obtusiloba (Rydb.) House rossii (R. Br.) Graebn. rubella (Wahlenb.) Hiem Pseudostellaria jamesiana (Torr.) Weber & Hartman Sagina saginoides (L.) Karst. Saponaria officinalis L. Silene bemardina S. Wats, ssp. maguirei Bocquet var. maguirei ssp. bemardina menziesii Hook. ssp. dorrii (Kellogg) C.L. Hitchc.& Maguire nuda (S. Wats.) C.L. Hitchc. & Maguire ssp. insectivora (Henders.) C.L. Hitchc. & Maguire sargentii S. Wats. Stellaria crispa Cham. & Schlecht. longipes Goldie umbellata Turcz. ex Kar. & Kir. Chenopodiaceae Atriplex argentea Nutt. ssp. argentea canescens (Pursh) Nutt. ssp. canescens confertifolia (Torr. & Frem.) S. Wats. heterosperma Bunge patula L. ssp. hastata (L.) H. & S. rosea L. Bassia hyssopifolia (Pallas) Kuntze Ceratoides lanata (Pursh) J.T. Howell var. lanata Chenopodium album L. atrovirens Rydb. botrys L. dessiccatum A. Nels. var. leptophylloides (J. Murr) H.A. Wahl. var. dessiccatum AREAS HABITATS ! NSW M 1 2 3 4 5 6 7 8 9 XXX X o X X X o X X X XXX X X X X X X X X X X X X X X X X X X X X XXX o o O X X X X X X X X X X X X o X ox X X X X x o X X o o o X X X o o X X o X X X o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X XXX X O X X X X X X X o o o 110 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats fremontii S. Wats. leptophyllum (Moq.) Nutt. ex S. Wats. overi Aellen Grayia spinosa (Hook.) Moq. Halogeton glomeratus (Stephan ex Bieb) C.A. Mey. Kochia scoparia (L.) Schrad. Monolepis nuttalliana (Roem. & Schult.) Greene spathulata Gray Sakola iberica Sennen & Pau Sarcobatus vermiculatus (Hook.) Torr. var. vermiculatus Sueda occidentalis S. Wats. Clusiaceae Hypericum formosum H.B.K. ssp. scouleri (Hook.) C.L. Hitchc. CONVOLVULACEAE Calystegia polymorpha (Greene) Munz Convolvulus arvensis L. CORNACEAE Cornus sericea L. ssp. sericea Crassulaceae Sedum integrifolium Coult. & A. Nels. lanceolatum (Nutt.) Britton & Rose obtusatum Gray Crossosomataceae Forsellesia nevadensis (Gray) Greene CUSCUTACEAE Cuscuta suksdorfii Yunker var. subpedicellata Yunker Cyperaceae Carex abrupta Mackenzie aquatilis Wahlenb. athrostachya Olney aurea Nutt. brevipes W. Boott. canescens L. capitata L. congdonii Bailey disperma Dewey douglasii Boott N s w M 1 2 3 4 5 6 7 8 9 X X O o o o O o X X X X X X X X X X X X X X X X X X X X X X X X X X X X o o o o O X X X X XXX X X X X X XXX XX o X X X X o X X XXX X XXX X X O X X X XXX X X X X X X X X X 0 X X X X X X o X o X X X X X X X o o o 0 o o o o o o X X o O X X X X X X X X X X X X X X X X X X X X X X X January 1983 Lavin: Walker River Floristics 111 Table 1 continued. exserta Mackenzie festivelhi Mackenzie fissuricola Mackenzie haydeniana Olney helleri Mackenzie heteroneura W. Boott var. epapillosa (Mackenzie) F.J. Herm. var. heteroneura hoodii Boott jepsonii J.T. Howell jonesii Bailey lanuginosa Michx. leporinella Mackenzie luzulifolia W. Boott microptera Mackenzie nebraskensis Dewey nervina Bailey nigricans C.A. Mey. pachystachya Cham, ex Steudel. phaeocephala Piper praegracilis W. Boott rossii Boott ex Hook. rostrata Stokes ex With. scopulorum T. H. Holm. var. bracteosa (Bailey) F.J. Herm. var. scopulorum simulate Mackenzie specifica Bailey spectabilis Dewey straminiformis Bailey subnigricans Stacey tahoensis Smiley vallicola Dewey vemacula Bailey vesicaria L. Eleocharis engelmannii Steud. palustris (L.) Roemer & Schultes pauciflora (Lightf.) Link Eriophorum crinigerum (Gray) Beetle Scirpus acutus Muhl. ex Bigelow americanus Pers. dementis M.E. Jones microcarpus Presl. nevadensis S. Wats. pungens Vahl. Elaeagnaceae Elaeagnus angustifolia L. Shepherdia argentea (Pursh) Nutt. Elatinaceae Elatine rubella Rydb. AREAS HABITATS ! N S w M 1 2 3 4 5 6 7 8 9 X X X X X X X X X X X X X X X o o 0 o 0 X X o X X o o o o o o X X X X X X X X X X X X X X X X X X X o o 0 X X X X X X X X X X o o X X X X X X X 0 o o o o o X X X X X X X X X X X o o o o o X X o X o o X X X X X X X X X o X X X 0 o o o o o o X X o X X X X X X X X X X o X o o X o X X X o o o o o o o o o X X o X X X X X X X X o X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X o o o X X o X o o o X X X o o X X X X X o o X o 112 Table 1 continued. Great Basin Naturalist Vol. 43, No. 1 AREAS habitats NSWM 12 3 456789 Ericaceae Arctostaphylos tievadensis Gray Cassiape mertensiana (Bong.) D. Don Kalmia microphylla (Hook.) Heller var. microphylla Ledum glandulosum Nutt. var. californicum (Kellogg) C.L. Hitchc. Orthilia secunda (L.) House ssp. secunda Phyllodoce breweri (Gray) Heller Pterospora andromedea Nutt. Pyrola californica Krisa dentata Sm. var. dentata minor L. Sarcodes sanguinea Torr. Vaccinium caespitosum Michx. var. paludicola (Camp) Hulten uliginosum L. ssp. occidentale (Gray) Hulten EUPHORBIACEAE Chamaesyce serpyllifolia (Pers.) Small Fabaceae Astragalus andersonii Gray bolanderi Gray calycosus Torr. ex S. Wats. var. calycosus canadensis L. var. brevidens (Gand.) Bameby casei Gray curvicarpus (Heller) J.F. Macbr. var. curvicarpus gibbsii Kellogg iodanthus S. Wats. var. iodanthus johnnis-howellii Bameby kentrophyta Gray var. danaus Bameby lentiginosus Dougl. ex Hook. var. ineptus (Gray) M.E. Jones malacus Gray obscurus S. Wats. oophorus S. Wats. var. lavinii Bameby in ed. platytropis Gray X X X X X X X X o X X XX X XXX X X X X X O O X X X X X o X XXX X X X X X X XXX X X X X X X X X X X X X X X X X X X X XXX X o X o XXX X X X o o o XXX XX X X X o X X X X X X X X X X X X X X X X X X January 1983 Lavin: Walker River Floristics 113 Table 1 continued. AREAS HABITATS N S W M 123456789 purshii Dougl. ex Hook. var. lectulus (S. Wats.) M.E. Jones var. tinctus M.E. Jones whitneyi Gray var. whitneyi Dalea omata (Dougl. ex Hook.) Eat. & Wright Lupinus andersonii S. Wats. arbustus Dougl. ex Lindl. ssp. calcaratus (Kellogg) Dunn brevicaulis S. Wats. caudatus Kellogg ssp. caudatus ssp. montigenus (Heller) Hess & Dunn confertus Kellogg hypolasius Greene lyallii Gray var. danatis (Gray) S. Wats, var. lyallii meionanthus Gray nevadensis Heller polyphyllus Lindl. ssp. superbus (Heller) Munz sellulus Kellogg var. lobbii (S. Wats.) Cox var. sellulus tegeticulatus Eastw X inyoensis Heller Medicago lupulina L. sativa L. Melilotus albus Medic. officinalis (L.) Pallas Oxytropis parryi Gray Robinia pseudoacacia L. Trifolium andersonii Gray ssp. beatleyae Gillett ssp. andersonii cyathiferum Lindl. hybridum L. longipes Nutt. ssp. longipes monanthum Gray var. monanthum pratense L. productum Greene repens L. wormskjoldii Lehm. Vicia americana Muhl. ex Willd. ssp. americana Fagaceae Castanopsis sempervirens (Kellogg) Dudley X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XXX XXX XXX XXX XXX X X X X X X XX XX X X XXX X X X X XXX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X o X X X X X X X o X o X 0 X X X X X o X X X X X X X X X X X X o o X X X X X X X X X X X X X X X X X X X X X X X X 114 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS HABITATS N S W M 123456789 Gentianaceae Frasera speciosa Dougl. ex Griseb. Gentiana calycosa Griseb. newberryi Gray Gentianopsis holopetala (Gray) litis Geraniaceae Erodium cicutarium (L.) L'Her. Geranium richardsonii Fisch. & Trautv. Haloragidaceae Hippuris vulgaris L. Hydrocharitaceae Elodea canadensis Michx. Hydrophyllaceae Hesperochiron califomicus (Benth.) S. Wats. Hydrophyllum capitatum Dougl. ex Benth. var. alpinum S. Wats. Noma aretioides (Hook. & Am.) Brand densum Lemmon rothrockii Gray Nemophila spatulata Coville Phacelia bicolor Torr. ex S. Wats. curvipes Torr. ex S. Wats. frigida Greene hastata Dougl. ex Lehm. ssp. compacta (Brand) Heckard heterophylla Pursh ssp. virgata (Greene) Heckard humilis Torr. & Gray var. humilis hydrophylloides Torr. ex Gray linearis (Pursh) Holz. monoensis Halse peirsoniana J.T. Howell ramosissima Dougl. ex Lehm. var. ramosissima tetramera J.T. Howell Tricardia watsonii Torr. ex S. Wats. Iridaceae Iris missouriensis Nutt. Sisyrinchium fialophilum Greene idahoense Bickn. var. occidentale (Bickn.) Henderson X X X X X X o o o o X X X X X X X o XXX O X X X X X X X X X X X X X XXX O X X XX X X X X X X X XXX XX o o XX X XXX X X X X X XXX O X o X X X X o o X X X X X X X X X XXX X X X X X XXX X X X X X X X X X X X X X X X o X X X X X X X o o X X X X X X X X X X X X XX XXXX XXXXX XX X X X X X X X XXXX X X X XX January 1983 Lavin: Walker River Floristics 115 Table 1 continued. AREAS HABITATS N S W M 123456789 JUNCACEAE Juncus balticus Willd. bryoides F.J. Herm. bufonius L. var. occidentalis F.J. Herm. var. bufonius capillaris F.J. Herm. chlorocephalus Engelm. drumniondii E. Mey. ensifolius Wikstr. var. montanus (Engelm.) C.L. Hitchcock hemiendytus F.J. Herm. kello^ii Engelm. longistylis Torr. macrandrus Coville megaspermus F.J. Herm. mertensianus Bong. mexicanus Willd. nevadensis S. Wats. orthophyllus Coville parry i Engelm. saximontanus A. Nels. Luzula ditxiricata S. Wats. multiflora (Retz.) Lej. ssp. comosa (E. Mey.) Hulten spicata (L.) DC. subcongesta (S. Wats.) Jepson JUNCAGINACEAE Triglochin concinna Burtt-Davy var. debilis (M.E. Jones) J.T. Howell maritima L. Lamiaceae Agastache urticifolia (Benth.) Kuntze Marrubium vulgare L. Mentha arvensis L. ssp. haplocalyx Briq. Monardella odoratissima Benth. ssp. glauca (Greene) Epling Salvia dorrii (Kellogg) Abrams var. dorrii Trichostema austromontana Lewis Lemnaceae Lemna gibba L. minuta H.B.K. trisulca L. Lentibulariaceae Utricularia vulgaris L. X X X X X X X X X X X X X X X X X X X X X o X o X X X X X X X o o o o o o o o o o o o o X X X X o o o o 0 o o X X X X X X X o o o X X X X X X X X X X X X X X X X X X X o X X X o o X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X O X X X XX X X X XXX X XXX o o o X O X X X X XX X X X X XX XXX X X X X X o o o o o o X X X X 0 X 0 X X o X X X X X 116 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS N S W M HABITATS 123456789 LiLIACEAE Allium amplectens Torr. anceps Kellogg atroTubens S. Wats, var. atrorubens bisceptrum S. Wats. campanulatum S. Wats. lemmonii S. Wats. parvum Kellogg validum S. Wats. Cahchortus bruneaunis A. Nels. & J.F. Macbride leichtlinii Hook. f. Camassia leichtlinii (Baker) S. Wats. Fritillaria atropurpurea Nutt. pinetorum A. Davids. Lilium parvum Kellogg Muilla transmontana Greene Smilacina stellate (L.) Desf. var. stellata Triteleia gracilis (S. Wats.) Greene ixioides (Ait. f.) Greene ssp. analina (Greene) Lenz Veratrum califomicum Durand var. californicum Zigadenus paniculatus (Nutt.) S. Wats. venetiosus S. Wats, var. venenosus LiMNANTHACEAE Floerkea proserpinacoides Willd. LiNACEAE Linum lewisii Pursh var. lewisii LOASACEAE Mentzelia albicaulis (Hook.) Torr. & Gray congesta (Nutt.) Torr. & Gray dispersa S. Wats. laevicaulis (Hook.) Torr. & Gray montana (A. Davids.) A. Davids. torreyi Gray var. torreyi veatchiana Kellogg LORANTHACEAE Arceuthobium divaricatum Engelm. Phoradendron juniperinum Engelm. ex Gray ssp. juniperinum o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o o o o o o o X X X X X X X X X X X X X X X X X X X X X X X o o o o X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X O X O X X X O X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X o X X X X X X X X January 1983 Lavin: Walker River Floristics 117 Table 1 continued. AREAS HABITATS NSWM 123456789 Malvaceae Malva neglecta Wallr. Sidalcea glaucescens Greene multifida Greene oregana (Torr. & Gray) Gray ssp. spicata (Kegel) C.L. Hitchc. Sphaeralcea atnbigua Gray ssp. monticola Kearney grossulariifolia (Hook. & Am.) Rydb. MORACEAE Humulus lupulus L. var. lupuhides E. Small Nyctag inaceae Abronia turbinata Torr. ex S. Wats. Mirahilis bigelovii Gray var. retroTsa (Heller) Munz Nymphaeaceae Nuphar luteum (L.) Sibthorp. & Sm. ssp. polysepalum (Engelm.) E.O. Real Oleaceae Menodora spinescens Gray Onagraceae Boisduvalia densiflora (Lindl.) S. Wats. Camissonia claviformis (Torr. & Frem.) Raven ssp. integrior (Raven) Raven nevadensis (Kellogg) Raven pubens (S. Wats.) Raven pusilla Raven subacaulis (Pursh) Raven tanacetifolia (Torr. & Gray) Raven ssp. tanacetifolia Circaea alpina L. ssp. pacifica (Aschers. & Magnus) Raven Epilobium anagallidifolium Lam. angustifolium L. brachycarpum Presl. ciliatum Raf. ssp. ciliatum glaberrimum Barbey lactiflorum Hausskn. latifolium L. obcordatum Gray ssp. obcordatum oregonese Hausskn. X X X X o X X XX X X X X X o o X X X X X X o X X o o o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 0 o X X X X X X X X X X X X X X X X X X X 0 o X 118 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats N s w M 1 2 3 4 5 6 7 8 9 X X X X X X X X X X X X 0 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o o o o o X X X X X X X X X X X o X X o X X o o o 0 Gayophytum decipiens Lewis & Szweykowski diffusum Ton. & Gray ssp. parviflorum Lewis & Szweykowski heterozygum Lewis & Szweykowski ramosissimum Torr. & Gray Oenothera caespitosa Nutt. ssp. marginata (Nutt.) Munz hookeri Ton. & Gray ssp. angustifolia (R.R. Gates) Munz ORCmOACEAE Corallorhiza maculata Raf. Epipactis gigantea Dougl. ex Hook. Listera convallarioides (Sw.) Nutt. Platanthera dilatata (Pursh) Lindl. ex Beck var. leucostachys (Lindl.) Luer sparsiflora (S. Wats.) Schlechter Spiranthes romanzoffiana Cham. Orobanchaceae Orobanche corymbosa (Rydb.) Ferris var. corymbosa fasciculata Nutt. var. lutea (Parry) Ashey var. fasciculata Paeoniaceae Paeonia brownii Dougl. ex Hook. Papaveraceae Argemone munita Dur. & Hilg. ssp. rotundata (Rydb.) G. Ownbey Corydalis aurea Willd. Dicentra uniflora Kellogg Eschscholzia calif omica Cham. Plantag in aceae Plantago lanceolata L. POACEAE Agropyron dasystachum (Hook.) Scribn. & Sm. var. dasystachum desertorum (Link.) Schultes pringlei (Scribn. & Sm.) A.S. Hitchc. scribneri Vasey trachycaulum (Link.) Malte ex H. Lewis var. latiglume (Scribn. & Sm.) Beetle var. trachycaulum X X o X X X X X X X X X X X X X X X X X X o XXX X X X o XX X X X o o XX X XX XX X XXX XX X O X X X X X X X X X X X X o X X January 1983 Lavin: Walker River Floristics 119 Table 1 continued. AREAS HABITATS N S w M 1 2 3 4 5 6 7 8 9 X X X X X X X X X o X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X o X X o X X X X X X X X X X X X X X X X X X X X X X X X X o o o o o X X X X X X X X X X X o X X X X X X X Agrostis capillaris L. exarata Trin. filiculmis M.E. Jones lepida A.S. Hitchc. scabra Willd. Alopecurus aequalis Sobol. geniculatus L. Beckmannia syzigachne (Stendl.) Feam. Bromus anomalus Rupr. ex Foum. carinatus Hook. 6f Am. catharticus Vahl commutatus Schrad. inermis Leyes. rubens L. suksdorfii Vasey tectorum L. CakiTnagrostis breweri Thurb. canadensis (Michx.) Beauv. purpurascens R. Br. Dactylis glomerata L. Danthonia intermedia Vasey Deschampsia cespitosa (L.) Beauv. ssp. cespitosa danthonioides (Trin.) Munro ex Benth. elongata (Hook.) Munro ex Benth. Distichlis spicata (L.) Greene var. stricta (Torr.) Beetle Echinochloa crusgalli (L.) Beauv. Elymus cinereus Scribn. & Merr. glaucus Buckl. ssp. glaucus ssp. virescens (Piper) Gould triticoides Buckl. var. triticoides var. pubescens A.S. Hitchc. Elysitanion X hansenii (Scribn.) Bowden Eragrostis orcuttiana Vasey Festuca brachyphylla Schultes pratensis Huds. rubra L. Glyceria elata (Nash) M.E. Jones Hilaria jamesii (Torr.) Benth. Hordeum brachyantherum Nevski jubatum L. X X o X X X X X X X X X X X X X o X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 120 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats N S w M 1 2 3 4 5 6 7 8 9 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X y X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o 0 o X X X X X X X o o o X X X X X o o o Koeleria nitida Nutt. Leucopoa kingii (S. Wats.) W.A. Weber Melica bulbosa Geyer ex Porter & Coult. stricta Boland. Muhlenbergia asperifolia (Nees & Meyem) Parodi filiformis (Thurb.) Rydb. richardsonis (Ttin.) Rydb. Oryzopsis hymenoides (Roemer & Schultes) Ricker Panicum capillare L. Phleum alpinum L. pratense L. Phragmites australis (Cav.) Trin. ex Steud. Poa ampla Merr. annua L. bolanderi Vasey bulbosa L. compressa L. cusickii Vasey epilis Scribn. fendleriana (Steud.) Vasey gracillima Vasey incurva Scribn. & Williams interior Rydb. leibergii Scribn. leptocoma Trin. nervosa (Hook.) Vasey nevadensis Vasey ex Scribn. var. juncifolia (Scribn.) Beetle var. nevadensis palustris L. pratensis L. rupicola Nash, ex Rydb. scabrella (Thurb.) Benth. ex Vasey secunda Presl. suksdorfii (Beal) Vasey ex Piper Polypogon monspeliensis (L.) Desf. Puccinellia distans (Jacq.) Pari. lemmonii (Vasey) Scribn. nuttalliana (Schultes) A.S. Hitchc. Secale cereale L. Sitanion hystrix (Nutt.) J.G. Smith var. brevifolium (J.G. Smith) C.L. Hitchc. jubatum J.G. Smith Sphenopholis obtusata (Michx.) Scribn var. obtusata Sporobolus cryptandrus (Torr.) Gray X X X X X X X X X X X XXX X o O 0 X X January 1983 Lavin: Walker River Floristics 121 Table 1 continued. AREAS HABITATS N S w M 1 2 3 4 5 6 7 8 9 X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X o X o X o o 0 o 0 X X X X X X X X X X X X X X X X X X X X X X X X X X o o X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Stipa californica Merr. & Burtt-Davy Columbiana Macoun. comata Trin. & Rupr. var. comata nevadensis B.L. Johnson occidentalis Thurb. ex S. Wats. pinetorum M.E. Jones speciosa Trin. & Rupr. thurberiana Piper webberi (Thurb.) B.L. Johnson Trisetum spicatum (L.) Richt. ssp. spicatum triflorum (Bigelow) Love & Love ssp. molle (Michx.) Love & Love Vulpia megalura (Nutt.) Rydb. octoflora (Walt.) Rydb. var. hirtella (Piper) Henr. var. octoflora POLEMONIACEAE Allophyllum gilioides (Benth.) A. & V. Grant integrifolium (Brand) A. & V. Grant violaceum (Heller) A. & V. Grant Collomia grandiflora Dougl. ex Lindl. linearis Nutt. tinctoria Kellogg Eriastrum sparsiflorum (Eastw.) Mason wilcoxii (A. Nels.) Mason Gilia brecciarum M.E. Jones ssp. brecciarum capillaris Kellogg inconspicua (Sm.) Sweet interior (Mason & A. Grant) A. Grant leptalea (Gray) Greene ssp. leptalea leptantha Parish ssp. salticola (Eastw.) A. & V. Grant leptomeria Gray malior Day & V. Grant modocensis Eastw. ophthalmoides Brand sinuata Dougl. ex Benth. Gymnosteris parvula (Rydb.) Heller Ipomopsis aggregata (Pursh) V. Grant ssp. ag^regata ssp. attenuata (Gray) V. & A. Grant congesta (Hook.) V. Grant ssp. montana (A. Nels. & Kennedy) V. Grant ssp. palmifrons (Brand) Day 122 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats N S W M 123456789 Leptodactylon pungens (Torr.) Nutt. ex Rydb. ssp. pulchriflonim (Brand) Mason Linanthus ciliatus (Benth.) Greene var. neglectus (Greene) Jepson nuttallii (Gray) Greene ex Milliken ssp. nuttallii septentrionalis Mason MicTosteris gracilis (Hook.) Greene ssp. gracilis ssp. humilfs (Greene) V. Grant Navarretia breweri (Gray) Greene divaricata (Torr. ex Gray) Greene var. divaricata Phlox caespitosa Nutt. ssp. pulvinata Wherry covillei E. Nels. diffusa Benth. ssp. subcarinata Wherry ssp. diffusa hoodii Richards ssp. canescens (Torr. & Gray) Wherry longifolia Nutt. Polemonium caeruleum L. ssp. amygdalinum (Wherry) Munz califomicum Eastw. chartaceum Mason eximium Greene pulcherrimum Hook. POLYGALACEAE Polygala intermontana Wendt subspinosa S. Wats. POLYGONACEAE Chorizanthe brevicomu Torr. var. spathulata (Rydb.) C.L. Hitchc. watsonii Torr. & Gray Eriogonum baileyi S. Wats. var. baileyi beatlleyae Reveal brachyantherum Coville caespitosum Nutt. var. cemuum elatum Dougl. ex Benth. var. elatum esmeraldense S. Wats. heermannii Dur. & Hilg. var. humilius (Stokes) Reveal hookeri S. Wats. incanum Torr. & Gray X X X X X X X X XXX X X X X o X X X X X X X X X X X X X X X X o o o X X X X X X X X X X X X X X X X X o o X X X X X X X X X o o X X o o 0 X X X X X X X X X X o X X X X X X X X X X X X X X X X X o 0 o o X X X X X X X X X X X X X X X X o X X X X X X X X X o X o X X X X X X X X X X X X X X X X X X X X X o X o X X X o X X 0 X X X X January 1983 Lavin: Walker River Floristics 123 Table 1 continued. \REAS HABITATS N s w M 1 2 3 4 5 6 7 8 9 lobbii Torr. & Gray var. lobbii X X X X X maculatum Heller X X X marifolium Torr. & Gray X X X X X micTothecum Nutt. var. laxiflorum Hook. X X X X X X X X X var. ambiguum (M.E. Jones) Reveal X X X X X X X nidularium Coville X X X X X nudum Dougl. ex Benth. var. deductum (Greene) Jepson X X X X X X nutans Torr. & Gray var. nutans X X o ochrocephalum S. Wats. var. alexandrae Reveal in ed X X X X X ovalifolium Nutt. var. ovalifolium X X X X X X X X X var. nivale (Canby) M.E. Jones X X X X X X X var. nevadense Gandoger X X X X X X X pusillum Torr. & Gray X X X rosense A. Nels. & Kennedy X X X X X X X spergulinum Gray var. reddingianum (M.E. Jones) J.T. Howell X X X X X X X X X X strictum Benth. ssp. proliferum (Torr. & Gray) Stokes X X o umbellatum Torr. var. nevadense Gandoger X X X X var. umbellatum X X X X X X X X X X X vimineum Dougl. ex Benth. X X X X X wrightii Torr. ex Benth. var. subscaposum S. Wats. X X X X X X X Oxyria digyna (L.) Hill X X o X X Oxytheca dendroidea Nutt. ssp. dendroidea X X Polygonum amphibium L. var. stipulaceum Coleman X X X X aviculare L. X X X X X bistortoides Pursh X X X X X X douglasii Greene var. douglasii X X X X X X X var. johnstonii Munz o o o o O 0 var. latifolium (Engelm.) Greene X X X X X X kelloggii Greene X X X X X X X X X X minimum S. Wats. X X X X persicaria L. X X X shastense Brewer ex Gray X X X Rumex acetosella L. X X X X calif omicus Rech. f. X X o X o o o o o crispus L. X X X X X X X lacustris Greene o 0 o o occidentalis S. Wats. o o paucifolius Nutt. ex S. Wats. ssp. paucifolius X X X X X X X X ssp. gracilescens (Rech. f.) Rech. f. X X X salicifolius Weijnm. X X triangulivalvis (Danser) Rech. f. X X X X X X X 124 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats N S W M 123456789 PORTULACACEAE Calyptridium roseum S. Wats. umbellatum (Torr.) Greene Claytonia nevadense S. Wats. rubra (T.J. Howell) Tidestrom umbellata S. Wats. Leivisia nevadensis (Gray) B.L. Robins. pygmaea (Gray) B.L. Robins, ssp. pygmaea rediviva Pursh ssp. rediviva ssp. minor (Rydb.) A. Holmgren sierrae Ferris triphylla (S. Wats.) B.L. Robins. Montia chamissoi (Ledeb. ex Spreng.) Greene POTAMOGETONACEAE Potamogeton gramineus L. richardsonii (Benn.) Rydb. Primulaceae Androsace septentrionalis L. var. subumbellata A. Nels. Dodecatheon alpinum (Gray) Greene ssp. alpinum ssp. majtis H.J. Thompson jeffreyi Van Houtte Primula suffrutescens Gray Ranunculaceae Aconitum columbianum Nutt. Actaea rubra (Ait.) WiUd. ssp. arguta (Nutt.) Hulten Anemone drummondii S. Wats. Aquilegia formosa Fisch. var. paucifhra (Greene) Boothman var. formosa pubescens Goville Caltha leptosepala DC. ssp. howellii (Huth) P.G. Smith Clematis ligusticifolius Nutt. var. brevifolia Nutt. Delphinium andersonii Gray ssp. andersonii glaucum S. Wats. X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X O X o X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X XXX X X X X X X X X X X o o O O 0 o o o ox X X X X X X X o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X O X X January 1983 Table 1 continued. Lavin: Walker River Floristics 125 AREAS HABITATS NSWM 123456789 parishii Gray ssp. parishii polycladon Eastw. Myosurus aristatus Benth. ex Hook. minimus L. ssp. montanus Campbell Ranunculus alismifolius Geyer ex Benth. var. alismellus Gray andersonii Gray aquatilis L. var. capillaceus (Thuill.) DC. cymbalaria Pursh var. saximontanus Fern. eschscholtzii Schlecht. var. eschscholtzii var. oxynotis (Gray) Jepson glaberrimus Hook. var. ellipticus Hook. occidentalis Torr. & Gray var. ultramx)ntantis Greene testiculatus Crantz Thalictrum fendleri Gray var. fendleri sparsiflorum Turcz. ex Fisch. & Mey. var. saximontanum Boivin Rhamnaceae Ceanothus greggii Gray var. vestitus (Greene) McMinn velutinus Dougl. ex Hook. Rham.nus rubra Greene ssp. rubra ROSACEAE Amelanchier pallida Greene pumila Torr. & Gray utahensis Koehne var. covillei (Standi.) Clokey var. utahensis Cercocarpus ledifolius Nutt. ex Torr. & Gray var. ledifolius var. intricatus (S. Wats.) M.E. Jones Chamaebatiaria millefolium (Torr.) Maxim. Fragaria virginiana Duchesne ssp. platypetala (Rydb.) Staudt Geum canescens (Greene) Munz macrophyllum Willd. X X X X XX X o X X X X X X X X X X X X X X X X X X XXX X X X X X X X X X X X X X X X X X X X X o X X 0 o o o o X X X o X X o X o X X X X X X X X X X X X X X X X X X X o X X X X X X 0 0 0 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o 0 O O 0 o X X X XXX X X X X X XXX XX 126 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats N S W M 123456789 Holodiscus dumosa (Nutt.) Heller var. glabrescens (Greenm.) C.L. Hitchc. Horkelia fusca Lindl. ssp. pseudocapitata (Rydb.) Keck Ivesia gordonii (Hook.) Torr. & Gray lycopodioides (Hook.) Torr. & Gray ssp. lycopodioides ssp. megalopetala (Rydb.) Keck muirii Graj^ purpurascens (S. Wats.) Keck ssp. congdonis (Rydb.) Keck shockleyi S. Wats. Potentillc anserina L. ssp. anserina biennis Greene breweri S. Wats. diversifolia Lehm. drummondii Lehm. ssp. bruceae (Rydb.) Keck ssp. drummondii fhbellifolia Hook, ex Torr. & Gray var. fhbellifolia fruticosa L. ssp. floribunda (Pursh) Elkington glandulosa Lindl. ssp. nevadensis (S. Wats.) Keck gracilis Dougl. ex Hook. var. glabrata (Lehm.) C.L. Hitchc. var. flabelliformis (Lehm.) Torr. & Gray newberryi Gray pectinisecta Rydb. pseudosericea Rydb. Prunus andersonii Gray emarginata (Dougl. ex Hook.) Walp. virginiana L. var. demissa (Nutt.) Sarg. var. melanocarpa (A. Nels.) Sarg. Purshia tridentata (Pursh) DC. Rosa woodsii Lindl. var. ultrarrumtana (S. Wats.) Jepson Rubus parviflorus Nutt. var. bifarius Fern. Sibbaldia procumbens L. Sorbus califomica Greene Spiraea densiflora Nutt. ex Torr. & Gray ssp. densiflora X X X X XXX XX X X 0 X X X X X X X X X X X X X X o o o O X o X X X X X X X X X X X X X X X X X X X X X X o o X X X X X o X X X X X X X o o o o 0 X X X o X X X X o X X X X X X X X X X X X X X X o X X X X X X X X X o X o o X o X X X X X X X X X X X X X X X X X X X X X X X X X X X o X o X X X X X X X X X X X XXXX X xxxxxx X XX X X O X XX X O O X X X XXX January 1983 Lavin: Walker River Floristics 127 Table 1 continued. AREAS HABITATS N S W M 123456789 RUBIACEAE Galium aparine L. bifolium S. Wats. hypotrichium Gray ssp. ebbettsense Dempster & Ehrend. ssp. hypotrichium multiflorum Kellogg trifidum L. ssp. pacificum Wieg. ssp. subbiflorum (Wieg.) Puff trifhrum Michx. Kelloggia galioides Torr. Salicaceae Populus balsamifera L. ssp. trichocarpa (Torr. & Gray) Brayshaw fremontii S. Wats. tremuloides Michx. Salix arctica Pallas drummondiana Barratt ex Hook. eastwoodiae Heller exigtia Nutt. geyeriana Anderss. var. geyeriana var. argentea (Bebb) Schneid. lasiandra Benth. var. lasiandra var. caudata (Nutt.) Sudworth lasiolepis Benth. lemmonii Bebb ligulifolia (Ball) Ball ex Schneid. lutea Nutt. melanopsis Nutt. myrtillifolia Anderss. var. myrtillifolia orestera Schneid. planifolia Pursh var. monica (Bebb) Schneid. reticulata L. ssp. nivalis (Hook.) Love, Love & Kapoor scouleriana Barratt ex Hook. Saxifragaceae Heuchera duranii Bacig. rubescens Torr. var. alpicola Jepson Lithophraffna glabrum Nutt. Mitella breweri Gray pentandra Hook. Pamassia palustris L. var. califomica Gray X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X o X X 0 X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X o o o o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 0 X X X X X X X o X X o o o o X X X X X X o o X X X X X X X o X X X o o 0 o X X o X X X X X X X X X X X X X X X X o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 128 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats N S W M 123456789 Ribes aureutn Pursh var. aureum cereum Dougl. inebrians Lindl. inerme Rydb. var. inerme montigenum McClatchie velutinum Greene var. velutinum var. glanduliferum (Heller) Jepson viscosissimum, Pursh var. viscosissim,um. Saxifraga aprica Greene bryophora Gray nidifica Greene odontoUmui Piper oregana T.J. Howell var. sierrae Coville tolmiei Torr. & Gray var. ledifolia (Greene) Engl. & Irmsch. SCROPHULARIACEAE Antirrhinum kingii S. Wats. Castilleja applegatei Fern. var. fragilis (Zeile) N. Homgren var. pallida (Eastwood) N. Holmgren chromosa A. Nels. exilis A. Nels. lemmonii Gray linariifolia Benth. ex DC. miniata Dougl. ex Benth. nana Eastwood peirsonii Eastwood pilosa (S. Wats.) Rydb. Collinsia parviflora Dougl. ex Lindl. Cordylanthus helleri (Ferris) J.F. Macbride ramosus Nutt. ex Benth. ssp. setosus Pennell Keckiella breviflora (Lindl.) Straw ssp. glabrisepala (Keck) Straw Limosella aquatica L. Mimetanthe pilosa (Benth.) Greene Mimulus breweri (Greene) Coville coccineus Congd. densus A.L. Grant floribundus Dougl. ex Lindl. guttatus Fisch. ex DC. lewisii Pursh mephiticus Greene X X X X X X X o o X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X o o o 0 X X X X X X o X X X X X X o X o X X X X X X X X X X o X o X X X X X X X X X X X X X X X X X X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o X X X X X X X o o o X X o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o o o o o o X X X X X X X X X X January 1983 Table 1 continued. Lavin: Walker River Floristics 129 AREAS HABITATS NSWM 1234567 moschatus Dougl. primuloides Benth. var. primuloides rubellus Gray suksdorfii Gray tilingii Kegel Orthocarpus copelandii Eastwood var. cryptanthus (Piper) Keck hispidus Benth. luteus Nutt. Pedicularis attollens Gray ssp. attollens groenlandica Retz. ssp. groenlandica semibarbata Gray var. semibarbata Penstemon bridgesii Gray dovidsonii Greene var. davidsonii deustus Dougl. ex Lindl. ssp. deustus heterodoxus Gray var. heterodoxus humilis Nutt. ex Gray newberryi Gray ssp. newberryi procerus Dougl. ex Grahm. ssp. formosus (A. Nels.) Keck rubicundis Keck rydbergii A. Nels. var. varians (A. Nels.) Cronq. speciosus Dougl. ex Lindl. Scrophularia desertorum (Munz) R.J. Shaw Verbascum thapsus L. Veronica americana (Raf.) Schwein. ex Benth. anagallis-aquatica L. beccabunga L. peregrina L. ssp. xalapensis (H.B.K.) Pennell serpyllifolia L. ssp. humifusa (Dickson) Syme ssp. serpyllifolia wormskjoldii Roemer & Schultes ssp. alterniflora (Fern.) Pennell SOLANACEAE Leucophysalis nana (Gray) Averett Nicotiana attenuata Torr. ex S. Wats. Solanum triflorum Nutt. X X X X X X O X X X X XX X X X X XXXX XXXX XXX XXXX X X X XXX XXX XXX XX O X O X X o o o o X X X X X X X X X X X X XXX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XX X X XXX X XXXX XXX XXX XXXX XXX XXXX XXXXX X XX X X X X X o o o o o X X O X O X X XX X X XX XX XXX X X X X X XX XX XXXX XXXX XX X o o o 130 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. AREAS habitats NSWM 123456789 Typhaceae Typha latifolia L. Urticaceae Urtica dioica L. var. holosericea (Nutt.) C.L. Hitchc. Valebianaceae Plectritis macrocera Torr. & Gray ssp. grayit (Suksdorf) Morey Valeriana californica Heller Verbenaceae Verbena bracteata Lag. & Rodr. ViOLACEAE Viola adunca Sm. var. adunca bakeri Greene ssp. bakeri beckwithii Torr. & Gray ssp. beckwithii macloskeyi Lloyd ssp. macloskeyi nephrophylla Greene var. nephrophylla purpurea Kellogg ssp. atriplicifolia (Greene) Baker 6f Clausen ssp. aurea (Kellogg) Clausen Zygophyllaceae Tribulus terrestris L. X X X X X X X X XX o X X ox XXX X X o o o X X X o 0 o o o X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X AGROPYRON ARIZONICUM (GRAMINEAE: TRITICEAE) AND A NATURAL HYBRID FROM ARIZONA Grant L. Pyrah' Abstract.— The new hybrid X Agrositanion pinalenoensis (Gramineae: Tribe Triticeae) is found in disturbed, for- ested areas of higher elevations in southern Arizona. In the Pinaleno Mountains where logging has been heavy, nu- merous disturbed habitats have permitted frequent hybrid populations to persist. Intermediate phenotype, chromo- some behavior, lack of seed set, and pollen sterility were used to interpret the status of this hybrid derivative. Introgression and/or segregation are not apparent. Of the many natural and experimental in- tergeneric hybrids reported in the Triticeae, none have been reported between Agropyron arizonicum Scribner & Smith and Sitanion hystrix var. brevifolium (J. G. Smith) C. L. Hitchcock. The present paper describes ex- tensive hybridizations between these two taxa. Agropyron arizonicum has flat leaves, dis- tinct flexuous spikes, one spikelet per node, and more or less ascending awns (although they are somewhat divergent at maturity). Si- tanion hystrix has long been recognized as an extremely variable species; however, erect spikes, very long awns on the glumes and lemmas, and 2 spikelets per rachis node are typical. The variation in .Arizona has been treated by Wilson (1963) as S. longifolium. Although Sitanion hystrix has very exten- sive distribution in western North America, it is limited to higher elevations of isolated mountain ranges in the southwestern United States and northern Mexico. On these iso- lated mountain ranges, the range of this spe- cies frequently overlaps that of Agropyron arizonicum, a species of high elevations re- stricted to west Texas, southwestern New Mexico, and southeastern Arizona (Fig. 1). In the Pinaleno Mountains, Graham Co., Ari- zona, and the Santa Catalina Mountains, Pima Co., Arizona (Fig. 2), where extensive areas have been distiubed by logging (Fig. 3), recreation, summer home development, and road building, numerous hybrids between Agropyron arizonicum and Sitanion hystrix var. brevifolium are formed. Eleven hybrid Fig. 1. Cross-hatched area represents sympatric range of Sitanion hystrix var. brevifolium and Agropyron anzonicum. Fig. 2. Cross-hatched areas represent the distribution of X Agrositanion pinalenoensis Pyrah in Arizona. (Larger area is the Pinaleno Mountains, smaller area is the Santa Catalina Mountains). 'Southwest Missouri State University, Springfield, Missouri 65802. 131 132 Great Basin Naturalist Vol. 43, No. 1 Fig. 3. Typical disturbed forest with dense stands oi Agropyron arizonicum and Sitanion hystrix var. brevifolium. populations were found and studied in the Pinaleno Mountains at elevations generally between 7000 and 10,000 ft. Materials and Methods Eleven hybrid populations in the Pinaleno Mountains and two hybrid populations from the Santa Catalina Mountains were studied. Prepared herbarium specimens from each are deposited in the Southwest Missouri State University herbarium. Five additional mature inflorescences were obtained from each hy- brid and four parent specimens from the High Peak population. These were put in en- velopes and used for comparative measure- ments of length of glume, lemma, awn, and rachis joint, and also determination of the number of spikelets per rachis node. A few late-flowering inflorescences were fixed in a solution of one part glacial acetic acid to three parts absolute ethyl alcohol. These were stored in 70 percent ethyl alco- hol and used for cytological studies. Pollen grains were obtained from mature spikes and viewed with the scanning electron microscope to determine viability. Soil pH was determined by sampling five sites each for parental species and the hybrid. The soil sample was carefully obtained by taking soil from the entire soil profile of 0 to 5 inches and mixed. A soil-water slurry was prepared and the pH determined by a standard pH meter. In an attempt to assess the pollen parent and the seed parent and hybrid success, a large population near High Peak was studied in the following way. A circular area 6 m in diameter was marked around each of 25 hybrids. The number of specimens of each parental species as well as other hybrids within this circle were recorded. Results and Discussion Pure stands of Sitanion typically grow in rather open, unshaded, shallow soil, with top- soil and litter depths from 1 to 3 inches and clay with scattered rock constituting the re- maining root zone. Soil pH range is 5.4 to 5.8. Agropyron arizonicum grows in richer soils, with the topsoil and litter occupying January 1983 Pyrah: Agropyron Hybrid 133 the upper 10 inches and only a limited amount of clay and rock toward the bottom of the root zone. Soil pH range is 5.9 to 6.4. Characteristically, this species is more vigor- ous in partially shaded areas, but it also grows in open sun. Nearly all combinations of the above soil conditions and other habitat requirements have been created by logging and road building, resulting in numerous dis- turbed habitats, as well as habitats for each parental species in very close proximity. In many of these situations, plants of both spe- cies either touch each other or are within only a few inches. Since flowering occurs over the same time period, this allows for showers of pollen to accomplish hybridiza- tion. In nearly all these situations hybrids are found. The frequency of hybridization between these species is difficult to assess, since a ma- ture hybrid plant is the only indication that hybrid pollination occurred. A circular area (6 m in diameter) around each of 25 hybrids near High Peak was examined and the num- ber of associated parental and hybrid plants was counted (Table 1). It is suggestive from the columns opposite hybrids 8, 9, 10, 11, and Table 1. Comparison of the number of parental and hybrid plants within an area (6 meters in diameter) around 25 hybrids in a population near High Peak (see text). Hybrid No. Sitanion Agropyron Hybrids 1 8 0 0 2 15 0 0 3 14 0 2 4 15 0 2 5 10 7 0 6 20 4 1 7 20 1 1 8 8 8 5 9 8 8 5 10 8 8 5 11 7 9 5 12 5 2 1 13 6 6 5 14 15 1 . 0 16 15 0 1 17 15 0 1 18 20 0 0 19 9 1 1 20 15 0 1 21 13 0 0 22 1 2 0 23 5 0 1 24 5 0 1 25 8 3 0 13 that hybrid success is dependent upon the density and proximity of both parents. Al- though this may appear to be obvious, there are situations in some dicots in which hy- bridization is abundant but one parent is rare (Stutz 1964, Pyrah 1965). A series of artificial hybridizations would be necessary to deter- mine whether or not Agropyron or Sitanion is more important as the seed or pollen parent of the natural hybrids. Hybrid plants are readily recognized be- cause of their robust size and slightly nodding spikes (Fig. 4), and most are growing in dis- turbed habitats (Fig. 3). Soil pH ranges from 5.7 to 6.5. Of the 11 populations studied, nearly every hybrid specimen fell within an intermediate range in morphological features and fragility of the rachis. A diagramatic il- lustration (Fig. 5) of Sitanion hystrix var. bre- vifolium and Agropyron arizonicum with their putative hybrids shows average lengths of the glume, lemma, awn, and rachis joint Fig. 4. Spikes of Sitanion hystrix var. brevifolium (left), hybrid (center), and Agropyron arizonicum (right). 134 Great Basin Naturalist Vol. 43, No. 1 \ \^ \/ \/ \/ \/ \/ \/ \/ N//r x=io.o \ / \ •X=66.5 X=11.0 =80. 1 / / \ / \/ \/ \/ \/ \/ X=10.7 \ / \ / \ \ \ / X=15.3 / / -X=36.4 \ \//\T::;:::Yfi X=26.6 ■K=13.7 ►-X=14.2 Sitanion Hybrid Agropyron Fig. 5. Schematic representation of some spike and spikelet characters of S. hystrix var. brevifolium, A. arizoni- cum, and their natural hybrid. Measurements used in this figure are means (X) from one population. A = Spike; cen- ter line between oblique lines (spikelets) represent the rachis joint. B = Lemma (triangle) with attached awn. C = Glume. and the number of spikelet pairs per spike. Measurements were derived from a popu- lation near High Peak consisting of 25 hy- brids and 4 parental specimens (Table 2). Field observation and examination of numer- ous herbarium specimens reveal clearly that the 2 parents are rather uniform with regard to the characters used and that where varia- tion exists the range does not overlap that of the hybrid. January 1983 Pyrah: Agropyron Hybrid 135 Table 2. Summary of measurements of spike characters used from one population to distinguish Agropyron ari- zonicum, Sitanion hystrix var. brevifolium, and their natural hybrid. No. nodes No. single No. double Glume Lemma awn Lemma Rachis joint per spike spikelets spikelets length length length length Agropyron X 11.900 11.600 0.300 14.156 26.588 13.739 15.349 N 40.000 40.000 40.000 109.000 119.000 119.000 109.000 # 2.329 2.318 0.648 2.290 4.758 9.122 3.059 XSE 0.368 0.367 0.103 0.219 0.436 0.836 0.293 Sitanion X 10.900 1.900 9.000 80.091 66.505 11.037 10.020 N 40.000 40.000 40.000 99.000 109.000 109.000 99.000 # 1.236 0.709 1.013 11.639 8.759 0.849 2.162 XSE 0.195 0.112 0.160 1.170 0.839 0.081 0.217 Hybrid X 13.360 8.730 4.630 28.071 36.372 11.609 10.737 N 400.000 400.000 400.000 1236.000 1336.000 1336.000 1236.000 S2 2.380 2.888 2.733 4.741 7.441 1.220 1.969 XSE 0.119 0.144 0.137 0.135 0.204 0.033 0.056 The hybrid did not set seed and no viable pollen was produced because the pollen grains were collapsed and empty (compare Figs. 6, 7, and 8). Preliminary cytological ex- amination of one hybrid showed 14 bivalents at metaphase; however, some bivalents show irregular pairing of chromosome segments that could cause cryptic structural hybridity (Fig. 9; arrows indicate conspicuous asynap- tic and synaptic pairing within two biva- lents). Morphology of spike characters was generally intermediate. These evidences strongly suggest that the plants studied are first generation hybrids and that little or no introgression occurs. These hybrid plants conform to the generic description of X Agrositanion as reported by Bowden (1967). Descriptions of the Hybrids X Agrositanion pinalenoensis Pyrah, Hyb. nov. (Agropyron arizonicum Scribner & Smith X Sitanion hystrix var. brevifolium (J. G. Smith) C. L. Hitchcock). Hybrida sterilis, inter Sitanion hystrix var. brevifolium et Agropyron arizonicum proba- biliter sed differt ab utroque spica moderate nutanti et inferioribus sex nodis spicae cum binatis vel binatis singulisque spiculis et supe- rioribus nodis spicae ciun singulus spiculus; differt a prima articulis rhachis et lemma- tibus longioribus et glumis et aristis lemma- tum brevioribus; differt a secunda articulis rhachis et lemmatibus brevioribus et glumis et aristis lemmatum longioribus. Figs. 6, 7, and 8. Electron micrographs (SEM) of pollen from Sitanion hystrix var. brevifolium (Fig. 6), hybrid (Fig. 7), and Agropyron arizonicum (Fig. 8). ca 600X. 136 Great Basin Naturalist Vol. 43, No. 1 Fig. 9. Meiotic metaphase I of hybrid. Arrows in- dicate two pairs of chromosomes with synaptic and asy- naptic regions. Sterile hybrids differing from both parents by having moderately nodding spikes and usually having the lower 6 to 8 spike nodes with paired spikelets or a combination of paired and single spikelets and only single spikelets at the upper nodes; differing from Sitanion by having longer rachis joints and lemmas but shorter glumes and lemma awns; differing from Agropyron by having shorter rachis joints and lemmas but longer glumes and lemma awns. Type.— Open grassy meadows of disturbed forests along State Rt. 366 near mile marker 141 on Mt. Graham, Pinaleno Mountains, Graham Co., Arizona, 13 Aug. 1975, Grant L. Pyrah 3051 (SMS). Discussion The classification of genera, species, and hybrids in the Triticeae is still open to ques- tion. Many taxonomists question the validity of recognizing all the genera now published, although Gould (1947) and Church (1967) are two of only a few who have initiated some consolidation. Widespread hybridization, similar chromosome behavior, and several variable spike characteristics were the pri- mary bases for consolidating Sitanion, Agro- pyron, and Hystrix with Elymus. Hitchcock et al. (1969) defend the retention of estab- lished separate generic names for this agro- nomically important tribe, primarily on the basis of practicality. Agropyron arizonicum is probably closely allied to A. spicatum and there are still conflicts about the variation found in Sitanion hystrix. Wilson (1963) treats the plants of this region as Sitanion longifolium, but Hitchcock et al. (1969) pre- fer S. hystrix var. brevifolium. Confusion is now arising in the literature for numerous named hybrids because of the lack of agreement as to the generic status of each of the parents involved. If a consoli- dation of genera were accepted, hybrids would bear the appropriate specific designa- tion within the genus Elymus. However, Bowden (1967) recognized the classical treat- ment of this tribe and made appropriate no- menclatural designations for intergeneric hybrids in conformity with the International Code of Botanical Nomenclature. Hybrid naming in this case had resulted in such ge- neric combinations as Agroelymus, Agrohor- deum, Agrositanion, Elyhordeum, and Sitor- deum. I have chosen to use the classification of Bowden and to create the binomial X Agrositanion pinalenoensis Pyrah. Literature Cited Bowden, W. 1967. Taxonomy of intergeneric hybrids of the tribe Triticeae from North America. Cana- dian J. Bot. 45:720. Church, G. L. 1967. Taxonomic and genetic relation- ships of eastern North American species of Elymus. Rhodora 69:135. 1967. Pine Hills Elymus. Rhodora 69:330-351. Gould, F. W. 1947. Nomenclatorial changes in Elymus with a key to the California species. Madrono 9:120-128. 1974. Nomenclatural changes in the Poaceae. Brittonia 26:60. Hitchcock, C. L., A. Cronquist, M. Owenby, and J. W. Thompson. 1969. Page 701 in Vascular plants of the Pacific Northwest. Part I. Pyrah, G. 1964. Cytogenetic studies of Cercocarpus in Utah. Unpublished thesis. Brigham Young Univ., Provo, Utah. Stutz, H. C, and L. K. Thomas. 1963. Hybridization and introgression in Cowania and Purshia. Evolu- tion 18:183-195. Wilson, F. D. 1963. Revision of Sitanion (Triticeae, Gramineae). Brittonia. 15:303-323. SPECIES COMPOSITION, DISTRIBUTION, AND PHYTOSOCIOLOGY OF KALSOW PRAIRIE, A MESIC TALL-GRASS PRAIRIE IN IOWA Jack D. Brotherson' Abstract.— Species composition, distribution, and phytosociology of an 8 hectare area of tall grass prairie was in- tensively studied. Elevation and soils data were correlated with species distribution patterns. All species showed a response. Nine general patterns of distribution were observed in relation to elevation and soil types. Ordination and interspecific association analyses were used to identify clusters or groups of species having similar ecological ampli- tudes. Sporobolis heterolepis is the dominant plant of the upland prairie. The vegetation of the prairie is best de- scribed and represented by the continuum concepts of phytosociology. A government survey started in March 1832, when Iowa was still a territory, and completed in August 1859 first documented the original extent of Iowa's prairie. The sur- vey indicated that in the 1850s grassland cov- ered about 85 percent of Iowa (U.S. Govern- ment 1868, Hayden 1945, Hewes 1950, and Dick-Peddie 1955). Today there are only a few tracts of this once vast Iowa prairie remaining. Provision for state-owned prairies was made in 1933 when the Iowa State Con- servation Commission prepared a report known as the Iowa Twenty-five Year Con- servation Plan. This plan led to the purchase of several prairies that are now owned by state agencies. The prairies were purchased and set aside as natural areas with the intent that the various typical landscapes, wild flowers, and wild life of the native tall-grass prairie region be preserved for posterity. It was also intended that these areas would be useful as '^ame and wild life sanctuaries; as examples of the native prairie soil types, where comparisons could be made with culti- vated soils of the same soil association; and as reserves of prairie where scientific in- vestigations could be made on problems con- cerning the native vegetations, floras, and faunas of the various topographic, climatic, and prairie districts throughout Iowa. There- fore, they were meant to serve as a reference point by which future generations could compare the influences of man on Iowa since settlement (Hayden 1946, Moyer 1953, Aii- man 1959, Landers 1966). Kalsow Prairie, 64.8 ha (160 acres) of un- plowed grassland in Pocahontas County, Iowa, is one such area. Criteria for its pur- chase dictated that this area satisfy the re- quirements of a game preserve, contain one or more soil types of an association, and in- clude several regional vegetation types (Hay- den 1946). Since its purchase in 1949 it has been the object of several studies on the na- ture and description of its vegetation, soils, management, insects, response to fire, mam- mals, and nematodes (Moyer 1953, Eh- renreich 1957, Esau 1968, Richards 1969, Brennan 1969, Norton and Ponchillia 1968, Schmitt 1969). The characteristics of Iowa prairie in terms of vegetation types, structure, and general ecology of the dominant species was the sub- ject of several papers during the 1930s and 1940s (Steiger 1930, Rydberg 1931, Weaver and Fitzpatrick 1934, Hayden 1943). These authors recognized the existence of six major types of grassland or vegetative communities and generally concluded that water relations, as affected by climate, soil, and topography, are responsible for local variations in the structure and distribution of Iowa prairie vegetation. Weaver and Fitzpatrick (1934) state: In varying the water relations of soil and air they merely bring about changes in the groupings of the dominant grasses and accompanying segregations and rearrange- ments of the forbs. 'Department of Botany and Range Science, Brigham Young University, Prove, Utah 84602. 137 138 Great Basin Naturalist Vol. 43, No. 1 The major grassland types, as alluded to in the above studies, were labeled "Con- sociations" after Weaver and Clements (1938) and were designated as follows: 1. Big Bluestem type {Andropogon gerardi)— found on the lower moist slopes and well- aerated lowlands. 2. Slough Grass type {Spartina pectinata)— found on poorly aerated and wet soils of sloughs and natural drainage systems. 3. Tall Panic Grass- Wildrye type {Panicum virgatum and Elymus canadensis)— found to occur on soils intermediate between Slough Grass and Big Bluestem types. 4. Little Bluestem type {Schizachyrium sco- parius)— most important upland type (well-drained soils). 5. Needle Grass type {Stipa spartea)— found on the uplands, often occurring as a nar- row zone following the shoulders of the ridges. 6. Prairie Dropseed type {Sporobolus hetero- lepis)—iound locally on the dryest upland sites. Moyer (1953), Aikman and Thome (1956), Ehrenreich (1957), and Kennedy (1969) in re- cent studies present ecological and taxonomic descriptions of four state-owned native prairie tracts. All accounts contain extensive reviews of prairie literature. The vegetation complex as treated in these studies is limited basically to upland prairie. The studies also include information on soils, microclimate, topography, and management. Aikman (1959) reviewed in some detail the state of prairie research in Iowa. Investigations involving the distribution of individual species within the prairie associ- ation began with the work of Shimek (1911, 1915, 1925). Weaver (1930) and Weaver and Fitzpatrick (1932) discuss the role of the ma- jor grasses and forbs within the commimity. Steiger (1930) and Cain and Evans (1952) mapped the spatial distributions of several species. They conclude that the principal fac- tors affecting the local distribution patterns of prairie species are as follows: (1) micro- climatic conditions, (2) edaphic variations, (3) the biology of the species concerned, particu- larly methods of reproduction and dispersal, (4) the relations of the species and other or- ganisms, animal as well as plant, occurring in the community, and (5) the element of chance in the dispersal and establishment of new individuals. Local distribution patterns of species have been of interest to many ecol- ogists (Curtie 1955, Kenshaw 1964, Sanders 1969). Species in general show varying degrees of aggregation or association due to exhibited preferences for or tolerances of certain envi- ronmental conditions. The distributional pat- terns and interactions of the component spe- cies of a community express its phytosociological structure. Studies of grass- land phytosociology have been concerned with either classification or ordination of ba- sic species groups (Crawford and Wishart 1968). This investigation was undertaken to pro- vide information on the phytosociology of an 8 ha tract of Kalsow prairie in relation to edaphic and topographic variation. It in- cludes information on species composition and distribution, factors affecting the distri- butional patterns of these species, community types, and interrelationships within and be- tween these communities. Materials and Methods Study Site Kalsow Prairie is one of several state- owned Iowa prairies. It is 5 miles northwest of Manson, Iowa, and comprises the NE V4 of Section 36, Belleville Township, T 90N, R 32W, Pocahontas County. It occurs in a part of northcentral Iowa that was glaciated dur- ing the most recent advances of the Wiscon- sin Glacier and within the Clarion-Nicollet- Webster soil association area (Ruhe 1969). The area was chosen for study on the basis of its vegetational composition (i.e., floristic richness and the presence of several plant community types). The Vegetation Taxonomy Voucher specimens from the prairie were collected in duplicate throughout the grow- ing seasons. All specimens were identified, and identical sets have been deposited in the herbaria of Iowa State University, Ames, Iowa, and Brigham Young University, Provo, January 1983 Brotherson: Kalsow Prairie 139 Utah. Nomenclature follows Pohl (1966) for the grasses, Gilly (1946) for the sedges, and Gleason (1952) for the forbs. Community Types The 8 ha tract within Kalsow Prairie is ad- jacent to the south boundary of Kalsow Prairie and contains within its borders two vegetation types or subcommunities. These vegetation units were identified and delimit- ed as follows (Fig. 1): 1. Upland prairie— includes a major por- tion of the study area on the ridges and adjacent lower slopes. 2. Potholes and drainage— found in the swales and lowlands of the study site. The vegetation of these community types was analyzed using two separate approaches. The first involved the identification and list- ing of all species foimd within their bound- aries. The second utilized random plots to de- termine percent cover, composition, and -^WEST MAP OF THE KALSOW PRAIRIE A AREAS AFFECTED BY SOIL DRIFT FROM ADJACENT FIELDS ■ 20 ACRES OF. PRAIRIE INVOLVED IN SOIL AND PLANT DISTRIBUTION STUDIES d©^ POTHOLES AND DRAINAGE Fig. 1. Map of Kalsow Prairie. 140 Great Basin Naturalist Vol. 43, No. 1 interspecific relationships of species within these subcommunities. Quadrat Analysis The vegetation of each area was sampled by using 20 X 50 cm (1000 cm 2) quadrats. The quadrats were located on a restricted basis to reduce bias and to keep adjacent quadrats equal distances apart. Sampling was done between August 1 and September 15, when most species had reached their max- imimi growth. Cover estimates were made for each quadrat through use of Dauben- mire's (1959) method. Coverage was determined separately for all species overlapping the plot regardless of where the individuals were rooted. Coverage was projected to include the perimeter of overlap of each species regardless of super- imposed canopies of other species. The ca- nopies of different species are commonly in- terlaced or superimposed over the same area; therefore coverage percents often total great- er than 100 percent. Community Analysis Plant distribution and topographic studies. An 8 ha (20 acres) plot of prairie (Figure 1) containing a large segment of potholes and drainage was selected and staked off in a 9 X 9 m grid. Each 27 square-meter block was then surveyed and a presence list compiled for all plant species found within the area. A total of 968 blocks was thus surveyed, and distribution data were tabulated for 160 spe- cies. Topographic readings were taken at 968 points and recorded in tenths of feet on the same 8 ha grid. Points were located at the comers of the 27 square meter plots. Soil mapping. Soils were mapped on the 8 ha intensive study area. Mapping was done in cooperation with the Iowa State University Soils Survey under the supervision of Dr. Thomas E. Fenton, with Mr. J. Herbert Hud- dleston doing the actual mapping in the field. The mapping criteria for decisions on soil series delineation were as follows: A Clarion— typical well-drained soil on convex ridges. Surface color lOYR 2/2-3/2; subsoil color lOYR 4'3-5'4. A" Clarion-Nicollet— an intergrade with respect to drainage as interpreted from the color profile. Surface is still lOYR 2/2-3/3, but the subsoil colors are duller, not exceeding /3 chroma. Profile is not mottled as in Nicollet. N Nicollet-typical Nicollet, lOYR 21^-2/2 surface color, /2 chromas in the subsoil, which is mottled. Depth to carbonates generally greater than 30 inches. Na Calcareous Nicollet— as above, but calcareous at some depth less than 30 inches. W Webster— typical Webster with black (N2/-10YR 2/1) surface colors and gray (lOYR 4/1-4/2-5/2) sub- soil colors. Depth to gray subsoil ranges from 23/41 inches, but is commonly 30-35 inches. Carbonates occur at some depth below 22 inch- es, but the usual range in depth to carbonates is 22-36 inches. Some soils identified as Webster are non- calcareous in the entire probe depth (42 inches). Wh Heavy Webster— typical colors of Webster but heavier textures, strong- er development in the B and a lack of carbonates in 42 inches. In many places spots of Wh are included in the regular Webster mapping unit. On the other hand, some areas iden- tified as regular Webster but non- calcareous to 36 inches or more might better have been called heavy Webster. The Webster soils, as mapped, include a rather broad range of texture and depths to carbo- nates, which could be more precisely subdivided only with further investigations. N- Webster-Nicollet— an intergrade whose surface color and friability is like Nicollet but whose subsoil is darker or grayer than true Nicollet. The soil is drier than Webster. Na- Calcareous Webster-Nicollet— as above but calcareous somewhere above 22 inches. H Harps— typical Harps, a loamy, weakly developed soil that effer- vesces strongly to violently from the surface downward. Calcium carbo- nate equivalent probably in the range 20-40 percent. January 1983 Brotherson: Kalsow Prairie 141 C Canisteo— this is essentially cal- careous Webster. As mapped, it may be noncalcareous in the surface, but carbonates must be detected some- where in the 0-15 inch layer. It has lower calcium carbonate equivalent, heavier textures, and stronger devel- opment than Harps. H" Harps-Canisteo— an intergrade that has either Harps-like characteristics in the surface and becomes more Ca- nisteo-like with depth or Canisteo- like surface characteristics and a Harps-like subsoil. Wa Webster-Canisteo— an intergrade in which carbonates are first detected in the 15-22 inch layer. All other characteristics of Wa, W, and C are essentially the same. C" Inverted Canisteo-Heavy Webster— this represents a rather peculiar con- dition that tends to occur as a nar- row band around the potholes. The surface is moderately to strongly cal- careous, but carbonates decrease with depth to a noncalcareous, heavy, well-developed subsoil like that of heavy Webster. G Glenco— a poorly drained soil that occupies small potholes, the outer portions of large potholes, or con- necting drainage-ways. It has a black, highly organic surface but a gray, mineral, heavy, well-developed subsoil. In many respects it is similar to heavy Webster except for the or- ganic surface and lack of grit and pebbl-.s. Depth to carbonate is gen- erally greater than 42 inches, but may be up to 36 inches. Ga Calcareous Glenco— Glenco that be- comes calcareous above 36 inches. It usually lacks the heavy textures and good development of regular Glenco as well. O Okoboji— a black, mucky silt loam, very weakly developed soil occur- ring in the deepest areas of the potholes. GO Glenco-Okoboji— an intergrade that may have the heavy textures of Glenco, but is darker, more organic, less well developed, and wetter than Glenco. Seventeen soil series were recognized and mapped in the field, utilizing soil samples ob- tained with a 42-inch hand probe. Data Analysis General descriptive data. Data collected from quadrat studies, mapping studies, soil studies, and topographic studies were used to describe generally the vegetation. Frequency values and average cover values were deter- mined for all species in every stand. Ordination analysis. An ordination tech- nique proposed by Orloci (1966) was em- ployed to ordinate vegetation units within the different subcommunities listed above. Through this technique the entities to be or- dinated (i.e., plant species or stands of vege- tation) are projected as points into n-dimen- sional space. Such points are positioned by attribute scores through the application of the R and Q techniques of factor analysis. Once established, this multidimensional array of points is then reduced to a three dimen- sional system. This is accomplished by select- ing the two most different stands or species and placing one at zero and the other at some distance along the abscissa. All other stands or species under consideration are then positioned linearly in relationship to these two extremes. This action thus estab- lishes the X-axis. The above process is repeat- ed until all points have been established in three dimensional space (i.e., Y and Z axes have been added). Coordinate values for the X, Y, and Z axes are given as output from the computer. Interspecific association analysis. Expres- sions of interspecific association were at- tempted utilizing Cole's Index (1949). Step one in the computation of the index involves the accumulation of 2 X 2 contingency ta- bles. Actual calculation of the index involves the following three sets of formulas: when ad = be: C7± e = ad - be (a + c)(c + d) (a + b)(b + d) n(a + b)(b + c) when be > ad and d = a: C7± c = ad - be (b + d)(c + d) (a + b)(a + c) n(a + b)(a + c) 142 Great Basin Naturalist Vol. 43, No. 1 when be > ad and a > d: f^ , _ ad - be (a + b)(a + c) (b + d)(e + d) n(b + d)(e + d) where C7 = Cole's Index of Interspecific Association c = standard deviation Cole's Index n = total number of samples and a, b, c, and d represent the four cells of the 2x2 contingency table. Tests of statistical significance were per- formed by means of the Chi-square test. The chi-squares were computed by the formula: X2 = (ad - bc)2n (a + b)(a + c)(c + d)(b + d) where X^ = Chi-square value n = number of samples and a, b, c, and d represent the different cells of the 2x2 contingency table. In all cases a single degree of freedom was used. Chi-square values greater than 3.84 were considered to be significant at the 5 percent level, and values greater than 6.63 were considered to be significant at the 1 percent level. Data representation. Graphic representa- tion of data obtained from topographic stud- ies and from ordination analysis was drawn by the computer. Such representation was accomplished through the use of a plotting technique developed and programmed by Mr. Howard Jesperson, Agricultural Experi- mental Station, Iowa State University. Results and Discussion Species composition Information on species sampled in the up- land regions of Kalsow Prairie is presented in Table 1. Cover, composition (i.e., based on cover), and frequency values of Sporobolus heterolepis, Andropogon gerardi, Poa pra- tensis, and Panicum leibergii indicate these are the dominant grasses of the upland sites. Important or subdominant forbs include Sol- idago canadensis, Solidago rigida, Helianthus grosseserratus, Amorpha canescens. Aster eri- coides, Desmodium canadense, Zizia aurea, Helianthus laetiflorus. Aster laevis, Ratibida pinnata, Ceanothus americanus, and Rosa suffulta. Average cover values (Table 1) ranged from a high of 25.4 for Sporobolus heterolepis to a low of 0.01 for several species. Percent- age frequency values, on the other hand, ranged from 73.1 for Andropogon gerardi to 0.1 for many species. No tests of correlation were made between average cover values and percentage frequency, but those species showing the highest cover values generally showed correspondingly higher percentage frequency values. Since Sporobolus heterolepis is the domi- nant plant of the upland sites, Kalsow Prairie is placed within the "Consociation" desig- nated by Weaver and Fitzpatrick (1934) as the Prairie Dropseed type {Sporobolus hetero- lepis). Weaver and Fitzpatrick (1934) de- scribed this particular consociation as being the least extensive and least important tall- grass subcommunity. It was found to occupy drier upland sites and included the two sub- cominants Stipa spartea and Schizachyrium scoparius. Although these two species were present (Table 1), they were not foimd in suf- ficient quantity to be labeled subdominants. The important grass species found with Sporobolus heterolepis in this study (i.e., An- dropogon gerardi, Poa pratensis, and Panicum leibergii) suggest that the present-day upland regions of Kalsow Prairie are vegetatively distinct from the Prairie Dropseed Con- sociation of similar areas described earlier by Weaver. Both the species and their charac- teristics suggest that this difference is due ei- ther to change in the original vegetation, to differences in community characteristics, or to variations in the more recently glaciated land. Poa pratensis, for example, is an in- troduced species whose characteristics are such that it is able to compete well within the environment of prairie protected from fire and, under conditions of grazing, mow- ing, and other disturbance, is known to in- crease in importance (Weaver 1954). An- dropogon gerardi, on the other hand, is a native grass described by Weaver and Fitz- patrick (1934) as the dominant of the most extensive tall-grass consociation that occu- pied the lowlands and lower moist slopes of the tall-grass prairie region. Historical information, as well as evidence obtained in this study, indicates that much of the Kalsow Prairie has been subjected to January 1983 Brotherson: Kalsow Prairie 143 Table 1. Cover, composition, and frequency percentages for species sampled on upland prairie sites. Cover Composition Frequency Frequency* Species (%) (%) (%) (%) Sporobolus heterolepis 25.42 27.81 66.7 85.0 Andropogon gerardi 15.93 17.43 73.1 87.5 Poa pratensis 12.36 13.52 54.5 90.0 Solidago canadensis 4.12 4.51 38.9 15.0 Solidago rigida 2.57 2.81 19.4 5.0 Panicum leibergii 2.56 2.80 34.0 12.5 Helianthus grosseserratus 2.30 2.52 31.6 10.0 Amorpha canescens 2.08 2.28 15.0 27.5 Aster ericoides 1.99 2.19 40.7 37.5 Desmodium canadense 1.66 1.82 23.0 5.0 Zizia aurea 1.62 1.77 36.1 67.5 Helianthus laetiflorus 1.49 1.63 11.4 7.5 Aster laevis 1.30 1.42 16.9 7.5 Ratibida pinnata 1.26 1.39 20.6 10.0 Ceanothus americanus 1.15 1.26 .9 2.5 Rosa suffulta 1.07 1.17 18.4 45.0 Lysimachia hybrida .84 .92 .7 5.0 Convolvulus sepium .76 .83 1.2 2.5 Silphium laciniatum .72 .79 12.4 15.0 Achillea lanulosa .68 .74 11.7 45.0 Galium obtusum .65 .71 29.1 5.0 Spartina pectinate .60 .66 10.3 17.5 Artemisia ludoviciana .59 .65 4.4 2.5 Comandra umbellata .59 .65 9.1 45.0 Schizachyrium scoparius .52 .57 1.7 85.0 Fragaria virginiana .52 .57 20.7 7.5 Physalis heterophylla .47 .51 2.1 Elymus canadensis .45 .49 19.4 12.5 Stipa spartea .45 .49 6.6 20.0 Aster simplex .43 .47 13.1 Muhlenbergia racemosa .37 .40 11.2 Panicum virgatum .36 .39 12.1 75.0 Senecio pauperculus .36 .39 9.3 Lithospermum canescens .35 .38 13.0 17.5 Heliopsis helianthoides .31 .34 5.8 17.5 Psoralea argophylla .29 .32 3.8 5.0 Solidago missouriensis .28 .31 2.7 5.0 Apocynum sibiric^ .i .27 .30 3.3 Asclepias tuberosa .26 .28 3.5 2.5 Setaria viridis .23 .25 .7 Cirsium altissimum .23 .25 4.6 17.5 Sorghastrum nutans .21 .23 5.4 42.5 Liatris pycnostachya .21 .23 9.8 27.5 Petalostemum purpureum .20 .22 29.5 32.5 Pycnanthemum virginianum .20 .22 7.0 5.0 Lythrum alatum .20 .22 .6 Phlox pilosa .16 .18 6.3 10.0 Physalis virginiana .16 .18 3.1 7.5 Viola pedatifida .14 .15 5.5 5.0 Setaria lutescens .13 .14 .5 Viscia americana .12 .13 4.7 7.5 Lathyrus venosus .12 .13 2.5 5.0 Equisetum kansanum .11 .12 4.7 42.5 Eryngium yuccifolium .11 .12 1.0 2.5 Petalostemum candidum .11 .12 3.9 7.5 ^Figures taken from Moyer (1953) for comparison purposes. 144 Great Basin Naturalist Vol. 43, No. 1 Table 1 continued. Species Cover (%) Composition (%) Frequency Frequency* Baptisia leucophaea Asclepias syriaca Ambrosia artemisifolia Baptisia leucantha Carex gravida Oxalis stricta Teucrium canadense Viola sp. Gentiana andrewsii Potentilla arguta Scutellaria leonardii Thalictrum dasycarpum Lespedeza capitata Solidago riddellii Anemone cylindrica Helenium autumnale Pedicularis canadensis Bouteloua curtipendula Chenopodium album Lathyrus palustris Liatris aspera Lycopus americanus Lysimachia chiliata Mentha arvensis Solidago gymnospermoides Vemonia fasciculata Taraxacum officinale Echinacea pallida Agropyron repens Agropyron smithii Anemone canadensis Arabis hirsuta Asclepias sullivantii Asclepias verticillata Astragalus canadensis Cicuta maculata Helianthus maximiliani Juncus tenuis Lactuca scariola Lysimachia quadrifhra Panicum capillare Phleum pratense Rudbeckia hirta Veronicastrum virginicum Allium sp. Aster novae-angliae Cacalia tuberosa Prenanthes racemosa Solidago nemoralis Trifolium pratense .09 .10 .7 15.0 .07 .08 1.4 .06 .07 1.4 .06 .07 .7 .06 .07 2.0 .06 .07 .9 .06 .07 1.7 .06 .07 3.2 .05 .06 1.1 2.5 .05 .06 .5 .05 .06 2.2 .05 .06 1.6 7.5 .04 .04 1.0 2.5 .04 .04 2.4 .03 .03 .5 35.0 .03 .03 .8 .03 .03 1.2 22.5 .03 .03 .9 7.5 .02 .02 .2 .02 .02 1.1 .02 .02 .8 25.0 .02 .02 .7 .02 .02 .6 .02 .02 .4 .02 .02 1.4 5.0 .02 .02 .3 .02 .02 .9 .02 .02 .4 10.0 .01 .01 .5 .01 .01 1.2 2.5 .01 .01 1.7 .01 .01 .1 .01 .01 .2 5.0 .01 .01 .4 2.5 .01 .01 .3 .01 .01 .3 10.0 .01 .01 .6 .01 .01 .1 .01 .01 .4 .01 .01 .8 .01 .01 .1 .01 .01 .6 85.0 .01 .01 .3 .01 .01 .1 .01 .01 .1 .01 .01 .4 .01 .01 .1 .01 .01 .7 .01 .01 2.6 2.5 .01 .01 .1 12.5 ^Figures taken from Moyer (1953) for comparison purposes. mowing, grazing to some extent, and abun- dant pocket gopher activity. Both Mima moimds and pocket gopher {Geomys bur- sanins) activity are widely scattered across the prairie. The Mima mounds are poorly un- derstood areas of disturbance. Other disturb- ance areas are along the south and west boimdaries of the prairie, where dust from adjacent plowed fields has been deposited in depths up to two or three feet. The distribution of soil types in the 8 ha intensive study site is here of interest. Our survey showed that much of the 8 ha is of lowland soil types. In fact, a large part of the upland prairie may occupy lowland soil types. The disturbance will give possible ex- planation to the high incidence of Poa pra- tensis found with Sporobolus heterolepis, and the large tracts of lowland soil types might well explain the abundance of Andropogon January 1983 Brotherson: Kalsow Prairie 145 gerardi. Why Sporobolus heterolepis is found growing in such abundance on the lowland areas is difficult to explain, but it might be due to the high amount of calcareous soil types found within Kalsow Prairie. Early studies (Weaver and Fitzpatrick 1934, Shimek 1925) suggest the distribution of Sporobolus heterolepis as restricted to driest uplands. Because these areas often show a lack of soil profile development or outcroppings of parent material often high in carbonates (Oschwald et al., 1965), it is fea- sible that Sporobolus heterolepis is adapted to grow on soils of high carbonate content and that it might easily be extended to lowland soils high in carbonate content. Moyer (1953), in a study of the Kalsow Prairie vegetation, gave percentage frequen- cy values for many of the species included in Table 1 of this paper. His figures are report- ed in column four of Table 1 for comparison. These figures suggest that there have been some changes in the species composition of the upland prairie since 1953. Some of the species that show increases in percentage fre- quency in the past are Solidago canadensis. Solidago rigida, Panicum leibergii, Helianthus grosseserratus, Desmodium canadense, Ga- lium obtusum, and Fragaria virginiana. Spe- cies that show decreases in percentage fre- quency over this same period are Phleum pratense, Poa pratensis, Zizia aurea, Rosa suffulta, Schizachyrium scoparius, Panicum virgatum, Sorghastrum nutans, Equisetum kansanum. Anemone cylindrica, Liatris as- pera, and Sporobolus heterolepis. Such changes are not easily explained but might be related to general fluctuations of the vegeta- tion over a period of years, to fluctuations in climatic conditions (i.e., time and duration of rainfall, drought, etc.), to interspecific com- petition, to differences in the technique and intensity of sampling, and to the possible in- fluence of slight disturbance upon the prairie due to increased populations of pocket goph- ers, dust accumulation from adjacent fields, public visitors, and management practices. To describe in greater detail the inter- relationships of species in the upland prairie a three-dimensional stand and species ordina- tion treatment was attempted using Orloci's (1966) method. The results are shown in Fig- ures 2, 3, and 4. Data used in the ordination Fig. 2. Three-dimensional ordination of 37 upland prairie sites. 146 Great Basin Naturalist Vol. 43, No. 1 were from 444 samples taken from 37 sites in the upland prairie community. These 37 sites are shown as points in three-dimensional space in Figure 2 and as dots in two-dimen- sional space in Figure 3. Initially, attempts were made to place the individual sites into specific groups. Groups were designated on the basis of where the stands fell when plotted in three-dimensional space (i.e., those sites which fell close together were consid- ered to be the most similar and were placed within the same group). Attempts to under- stand the meaning of such groupings were unsuccessful. Fiuther attempts to understand the ordering pattern led to the conclusion that discrete grouping within these upland prairie regions is not feasible. It appears that the ordering of the stands into three-dimen- sional space was controlled by the response of several of the major species to environ- mental gradients. Of these species, Andropo- gon gerardi (Fig. 5) and Sporobolus hetero- lepis (Fig. 6) were plotted against the X and Y coordinates of the ordination. As can be seen, both species show continuous distribu- tion in relationship to the axes. Stands plotted near the origin and adjacent to the Y- axis were found to be from drier sites, and those found away from the origin were found on wetter, more moist sites. These facts tend to support the hypothesis that the vegetation of the upland prairie is a continuum as ear- lier described by Curtis (1955) and Dix and Butler (1960). Kennedy (1969), in studying an upland prairie in Guthrie County, Iowa, also concluded that prairie vegetation there is best described through the use of the contin- uum-index concept. Attempts at environmental factor correla- tion with the ordination axes were not made since only general information on environ- mental gradients was available. It seems, however, that these axes represent environ- mental gradients and that the ordering of stands or species along these axes is accom- plished through the response of the different stands or species to certain factors such as moisture, texture, soil carbonates, or other soil factors. The species ordination is shown in Figure 4. Spartina pectinata and Ceanothus ameri- canus are the most different entities on the X-axis, and Andropogon gerardi is the most distinct entity on the Y-axis. Other species having distinct distribution patterns are Phys- alis virginiana, Silphium laciniatum, Oxalis stricta, Amorpha canescens, Solidago mis- souriensis, Desmodium canadense, He- lianthus grosseserratus. Aster ericoides, Vicia americana, Pycnanthemum virginianum, Ratibida pinnata. Aster laevis, and He- lianthus laetiflorus. All other species either showed no definite distribution patterns or were too rare to establish a meaningful pat- tern. The circles A, B, and C in Figure 4 rep- resent the points where 76 of the 92 species fell. This ordering of species has not deli- neated associated groups but has pointed out Fig. 3. Two-dimensional ordination of 37 upland prairie sites. Factors responsible for pattern are unknown. Fig. 4. Two-dimensional ordination of species found in upland prairie study sites, factors responsible for pat- terns unknown: A, B, and C. = Clusters of species not showing distinct distribution patterns, d. = Helianthus laetiflorus. e. = Aster laevis. f. = Ratibida pinnata. g. = Pycnanthemum virginianum. h. = Aster ericoides. i. = Vicia americana. j. = Helianthus grosseserratus. k. = Desmodium canadense. 1. = Amorpha canescens. m. = Oxalis stricta. n. = Silphium laciniatum. o. = Physalis virginiana. p. = Andropogon gerardi. q. = Spartina pectinata. January 1983 Brotherson: Kalsow Prairie 147 those species that exhibit pecuHar distribu- tion patterns or that show a definite response to one or several environmental stimuli. Sanders (1969) found that the R-analysis of Orloci (1966) did give him some fairly dis- tinct groups of associated species as well as groups of species that could not be consid- ered associated. Collins (1968) used the tech- nique to identify taxa that were distinct and different and used them as indicator species in his interpretation of the ecological rela- tionships of fossil diatom populations. It is evident from Figure 4 that the method has not provided information on groups of associ- ated species but rather has indicated taxa that are distinct and therefore may have some usefulness as indicator species. Attempts to discover groups of positively associated species within the upland regions of Kalsow Prairie were made using Cole's In- dex (1949). Those species showing positive as- sociation with other taxa are shown in Table 2. A total of 298 significant associations were found. Some species, such as Achillea lanu- losa, Agropyron smithii, Amorpha canescens, Andropogon gerardi, Asclepias tuberosa, Carex gravida, Comandra umbellata, Helen- ium autumnale, Lespedeza capitata, Phleum pratense, and Solidago gymnospermoides, ex- hibit positive association with only a limited number of species. Other species, however, show positive association with a large num- ber of species. Some of these species are As- ter ericoides, Desmodium canadense, Fragaria virginiana, Galium obtusum, Helianthus grosseserratus, Poa pratensis, Solidago cana- densis, Solid go rigida, Sporobolus hetero- lepis, and Zizia aurea. Many species showed no significant association or expressed values of high negative association. Positive values of Cole's Index indicate that species occur to- gether more often than would otherwise be expected due to chance (Hale 1955, Hurlbert 1969). Therefore, through the use of such an index one can deduce groups of species that consistently show positive values of associ- ation with one another. Figures 7, 8, and 9 were constructed from values taken from Table 2 to illustrate the existence of such groups within the upland prairie. In all three cases one species was picked and the corre- sponding figure was then built up around this species. Species Distribution Patterns Eight hectares of the prairie adjacent to its southern boundary (Figure 1) were selected for intensive study of the distribution of plant species in relation to soils and topography. The area was chosen because it included within its boundaries a representation of all vegetation types occurring on Kalsow Prairie. The area was staked on a 9 X 9 m grid that placed 968 points within the 8 ha. From these points all factors included in this study were examined. The presence of all plant species found in the area was recorded in relation to each 27 square-meter section of the grid. From these present figures, distribution maps for 160 species were constructed. Examples of these maps are shown in Figures lOA through lOHH. These figures illustrate examples of distribution patterns often shared by several species. Andropogon gerardi (Fig. lOD) illus- trates a type of pattern typical of many spe- cies commonly found in the upland prairie. Fig. 5. Two-dimensional ordination of upland prairie with percentage cover values of Sporobolus heterolepis for each site shown relating directly to the Y-axis. Fig. 6. Two-dimensional ordination of upland prairie with percentage cover values of Andropogon gerardi for each site shown relating directly to the X-axis. 148 Great Basin Naturalist Vol. 43, No. 1 Table 2. Cole's Index values expressing positive interspecific association on upland prairie. Species Species X2a Ct" o^- Achillea lanulosa Agropyron repens Agropyron smithii Ambrosia artemisifolia Amorpha canescens Andropogon gerardi Schizachyrium scoparius Anemone canadensis Apocynum sibiricum Artemisia ludoviciana Asclepias sullivantii Andropogon gerardi Aster ericoides Poa pratensis Solidago rigida Sporobolus heterolepis Carex gravida Convolvulus septum Physalis heterophylla Andropogon gerardi Aster ericoides Galium obtusum Helianthus grosseserratus Muhlenbergia racemosa Petalostemum purpureum Pycnanthemum virginianum Solidago canadensis Solidago riddellii Asclepias syriaca Helianthus grosseserratus Senecio pauperculus Setaria lutescens Setaria viridis Stipa spartea Taraxacum officinale Andropogon gerardi Panicum leibergii Solidago rigida Sporobolus heterolepis Aster ericoides Poa pratensis Solidago canadensis Sporobolus heterolepis Comandra umbellata Lithospermum canescens Panicum leibergii Petalostemum purpureum Phlox pilosa Solidago rigida Sporobolus heterolepis Zizia aurea Artemisia ludoviciana Petalostemum candidum Poa pratensis Solidago missouriensis Aster simplex Calamagrostis canadensis Carex aquatilis Carex retrorsa Carex lasiocarpa Fragaria virginiana Heliopsis helianthoides Polygonum coccimeum Spartina pectinata Aster ericoides Convolvulus sepium Helianthus laetiflorus Poa pratensis Elymus canadensis 18.77 .73 .17 6.84 .22 .08 5.03 .27 .11 13.41 .21 .05 11.06 .51 .15 4.91 .22 .09 7.04 .44 .16 5.31 .22 .09 5.96 .73 .29 8.97 .45 .15 4.33 .34 .16 5.43 .43 .18 24.25 .27 .05 8.26 .20 .06 5.65 .18 .07 13.22 .53 .14 25.22 .24 .04 83.22 .43 .04 5.12 .23 .10 155.77 .77 .06 58.48 .28 .03 165.69 .57 .04 4.00 .24 .12 172.49 .55 .04 15.62 .51 .13 28.29 .24 .04 19.00 .20 .04 41.54 .77 .11 126.73 .18 .01 176.68 .30 .02 85.47 .15 .01 238.41 .46 .02 10.40 .21 .06 19.30 .35 .08 5.62 .41 .17 7.53 .18 .06 11.12 .19 .05 6.01 .25 .10 8.23 .77 .26 5.23 .31 .13 5.53 .19 .08 6.16 .19 .07 5.57 .80 .33 25.23 .16 .03 14.68 .26 .06 102.95 .70 .06 44.71 .35 .05 70.51 .55 .06 66.57 .39 .04 3.83 .38 .19 4.39 .27 .12 15.26 .18 .04 13.73 .18 .05 4.53 .42 .19 51.32 .22 .03 14.98 .28 .07 15.37 .78 .19 5.94 .67 .27 *Chi -square "Cole's Index ^Standard deviation Cole's Index January 1983 Table 2 continued. Brotherson: Kalsow Prairie 149 Species Species X2a i "^l:"'-' '■^v -.1 w ^iif jp> -IV ^iiii:; 2ii ^ X :::y iu.j^-";:;ij' /I --;^ HH Fig. 10. Distribution patterns in 8 hectare study area of: A. Agropyron smithii. B. Ambrosia artemisifolia. C. Amorpha canescens. D. Andropogon gerardi. E. Solidago nemoralis. F. Calamagrostis canadensis. G. Carex atherodes. H. Scirpus fluviatilis. I. Helenium autumnali. J. Silphium laciniatum. K. Aster iaeuis. L. Aster simplex. M. Apocynum sibericum. N. Artemisia ludoviciana. O. Ceanothus americanus. P. Conuo/uuZus sepium. Q. Desmodium canadense. R. Fragaria virginiana. S. Helianthus grosseserratus. T. Helianthus laetiflorus. U. Liatris aspera. V. Liatris pyonostachya. W. Lisimachia hybrida. X. Lycopus americanus. Y. OxaZis striata. Z. Panicum leibergii. AA. Polygonum coccineum. BB. Phalris arundinacea. CC. Psoralea aryophylla. DD. Senecio aurens. EE. Spartina pectinata. FF. Sporobolus heter- olepis. GG. Teucrium canadense. HH. Vemonia fasciculata. 156 Great Basin Naturalist Vol. 43, No. 1 Fig. 11. Soil series map of 20-acre intensive study area, abbreviations described in Methods section. A = Clarion, A~ = Clarion-Nicollet, N = Nicollet, Na = calcareous Nicollet, N~ = Nicollet-Webster, Na~ = calcareous Nicol- let-Webster, W = Webster, Wh = heavy Webster, Wa = calcareous Webster, C = Canisteo, H = Harps, H" = Harps-Canisteo, C" = inverted Canisteo-heavy Webster, G = Glenco, Ga = calcareous Glenco, GO = Glenco- Okoboji, O = Okoboji. several edaphic factors (soil and water re- gimes, macronutrients, micronutrients, tex- ture, organic matter, etc.) (Curtis 1959, Greig-Smith 1964, Kershaw 1964). From this we can conclude that species showing similar patterns of distribution may be equally well adapted in their response to one or more en- vironmental stimuli and yet differ greatly in their basic ecological amplitudes. The re- sponse of individuals to the environmental complex is measured in a species distribution pattern as well as in its importance within the community. Attempts were made to access the response of the species included in this study to the factors of soil and topography. Soil and ele- vation readings were recorded at all 968 points of the grid. From these readings a soils map (Figure 11) and contour and elevation maps (Figs. 12, 13) were constructed for the Fig. 12. Topographic map of 20-acre intensive study area. WEST-«- January 1983 Brotherson: Kalsow Prairie 157 8 ha plot. This made it possible to group all plant samples according to 0.5 ft changes in elevation or according to soil series. Once grouped, average cover values were com- puted for all participating species and re- corded in Tables 3 (elevation data) and 4 (soils data). As can be seen from these tables, all species showed response to these factors. Several species, Andropogon gerardi, Amorpha canescens. Aster ericoides, Elymus canadensis, Panicum leibergii, Poa pratensis, Solidago canadensis, Sporobolus heterolepis, and Zizia aurea, showed wide tolerance in relation to both soil and elevation, but all ex- hibited peak^ or plateaus of occurrence. These peaks or plateaus are interpreted to represent the optimum conditions under which a particular species can reach its high- est importance within the community in rela- tion to the entire species complex. Other species showed rather narrow ranges of tolerance. Some of these were Schiz- achyrium scoparius, Apocynum sibiricum, Asclepias sullivantii, Calamagrostis cana- densis, Carex atherodes, Eryngium yuccifo- lium, Lysimachia hybrida, Physalis hetero- phylla, Viola pedatifida and Ceanothus americanus. Those exhibiting narrow ranges also showed peaks of occurrence. For species exhibiting narrow tolerances, four basic types of distribution patterns as related to elevation (Table 3) are recognizable: (1) pothole and drainage, (2) lower slopes, (3) mid- and upper slopes, and (4) ridges. For species showing response to the soil factor (Table 4) three basic classes are recog- nizable: (1) Glenco, Glenco-Okoboji, and Okoboji, (2) calcareous, and (3) noncalcareous and ridge. Species indicating preference for class 1 were Calamagrostis canadensis, Carex atherodes, Carex aquatilis, Carex lasiocarpa, Carex retrorsa, Lysimachia hybrida. Poly- gonum coccineum, and Scirpus fluviatilis. Species showing preference for the cal- careous soils (class 2) were Agropyron smithii, Desmodium canadense, Galium obtusum, Helenium autumnale, Petalostemum purpu- reum, Senecio pauperculus, Silphium lacinia- tum, Solidago canadensis, Solidago nemoralis and Solidago riddellii. Examples of species preferring class 3 were Amorpha canescens, Artemisia ludoviciana, Asclepias tuberosa, Baptisia leucophaea, Eryngium yuccifolium, Lathyrus palustris, Panicum leibergii, Poa pratensis, Solidago missouriensis, Vicia ameri- cana, and Ceanothus americanus. These groups of recognizable patterns, each involving several species, suggest the ex- istence of subcommunities within the prairie area. To ascertain the existence of such com- munities, the data from Tables 3 and 4 were treated using Orloci's (1966) method of ordi- nation. When the results from the soils analy- sis were plotted (Fig. 14), four basic groups Fig. 13. Topographic map of 20-acre intensive study area plotted by computer. IQ y . 9 of 9' Fig. 14. Two-dimensional ordination of vegetation found on the different soil types in the 20-acre study area; cluster A indicates vegetation on Nicollet and Ni- collet-Webster soil types; B indicates vegetation on Clarion, Clarion-Nicollet, Webster, heavy Webster, cal- careous Nicollet, calcareous Nicollet-Webster, cal- careous Webster, Canisteo, Harps, Harps-Canisteo, and inverted Canisteo-heavy Webster soil types; C indicates vegetation on Glenco-Okoboji and Okoboji soil types; D indicates vegetation on Glenco and calcareous Glenco. 158 Great Basin Naturalist Vol. 43, No. 1 Table 3. Average cover values for species in relation to elevation in 20-acre intensive study area. 12 3 4 5 6 Species .7-1.2 1.3-1.8 1.9-2.4 2.5-3.0 3.1-3.6 3.7-4.2 Achillea lanulosa Agropyron smithii Ambrosia artemisifolia Amorpha canescens Andropogon gerardi Schizachyrium scoparius Anemone canadensis Anemone cylindrica Apocynum sibiricum Arabis hirsuta Artemisia ludoviciana Asclepias incamata Asclepias sullivantii Asclepias syriaca Asclepias tuberosa Aster ericoides Aster laet>is Aster simplex Baptisia leucantha Baptisia leucophaea Calamagrostis canadensis Carex atherodes Carex aquatilis Carex gravida Carex lasiocarpa Carex retrorsa Chenopodium album Cicuta maculata Cirsium altissimum Comandra umbellata Convolvulus sepium Desmodium canadense Elymus canadensis Equisetum kansanum Eryngium yuccifolium Fragaria virginiana Galium obtusum Gentiana andrewsii Helenium autumnale Helianthus grosseserratus Helianthus laetifhrus Helianthus maximiliani Heliopsis helianthoides Lactuca scariola Lathyrus palustris Lathyrus venosus Lespedeza capitata Liatris pycnostachya Lithospermum canescens Lycopus americanus Lysimachia chiliata Lysimachia hybrida Lysinwchia quadriflora Lythrum alatum Mentha arvensis Muhlenbergia racemosa Oxalis stricta Panicum capillare .01 .23 .23 .28 .02 .12 .09 .01 .01 .55 .20 1.49 4.27 .52 5.57 9.06 9.49 10.10 .25 .92 .75 .02 .55 .03 1.36 1.98 .52 1.15 .65 .08 .04 .01 .06 .02 .19 .02 .13 .07 .28 .02 .02 .44 .71 .41 1.87 2.09 1.72 .10 .77 1.84 .56 2.72 2.55 1.66 .13 .82 1.59 33.65 36.13 9.52 1.09 34.55 16.35 4.28 .23 2.05 3.22 1.33 .19 .01 .02 1.63 1.91 .53 .08 4.51 7.24 .02 2.67 .57 .14 .02 .54 .55 .67 .44 .03 .02 .14 .38 .09 .32 .03 .72 2.37 3.49 2.96 .11 .49 1.05 .85 .04 .14 .21 .05 .24 .91 .89 .92 1.91 1.11 .90 1.91 1.69 1.84 .93 .21 .01 .06 .13 .03 .28 .19 .18 .02 2.95 8.89 8.88 7.46 4.59 .01 .43 .01 1.39 .28 .22 .24 .02 .06 .03 .03 .02 .08 .06 .06 .01 .01 .11 .20 .80 .63 .65 .07 .26 .34 .33 .24 .13 .01 .03 21.50 1.42 .07 .04 .03 .03 .24 .20 .12 .28 .08 .52 .78 .18 .25 .22 January 1983 Brotherson: Kalsow Prairie 159 Table 3 continued. 7 8 9 10 11 12 13 14 15 4.3-4.8 4.9-5.4 5.5-6.0 6.1-6.6 6.7-7.2 7.3-7.8 7.9-8.4 8.5-9.0 9.1-9.6 .71 1.00 1.07 .13 8.00 .05 .71 2.03 4.50 1.79 1.71 7.86 .83 7.50 1.25 11.70 14.67 10.95 26.21 14.29 57.92 40.00 20.00 1.42 .33 .08 .71 2.50 4.29 3.00 7.89 2.50 3.50 1.25 7.37 2.50 5.83 1.00 .71 1.46 .25 .71 3.35 3.08 2.02 1.37 2.42 4.76 .09 .28 .33 1.08 .33 .17 .28 .08 .24 .08 .47 .25 .12 .33 .24 .12 2.36 2.08 2.50 .71 .25 .48 .09 .08 .09 .08 .71 .99 .08 .52 .25 .12 3.82 .83 .24 3.16 2.67 2.50 .50 .71 .05 .71 .05 2.14 .79 2.14 2.92 3.00 7.50 .26 2.14 .26 .71 .42 1.25 .36 .50 .13 .13 .71 8.16 5.71 .14 .79 .12 .39 .24 .53 .42 160 Great Basin Naturalist Vol. 43, , No. 1 Table 3 continued. 1 2 3 4 5 6 Species .7-1.2 1.3-1.8 1.9-2.4 2.5-3.0 3.1-3.6 3.7-4.2 Panicum leihergii .02 .51 1.19 2.31 Panicum virgatum .26 .66 .86 .60 Pedicularis canadensis .47 .09 .13 Petabstemum candidum .02 .02 .02 Petalostemum purptireum .03 .13 .59 .40 .35 Phalaris arundinacea 7.05 5.49 3.98 .49 Phlox pilosa .04 .05 .13 .13 Physalis heterophylla Physalis virginiana .01 .04 .03 Poa pratensis 1.24 2.82 3.57 4.24 Polygonum coccineum 27.27 16.81 4.02 1.52 .01 Potentilh arguta .02 Psoralea argophylla .02 .09 Pycnanthemum virginianum .37 1.83 .77 .35 Ratibida pinnata .21 .30 1.65 1.93 1.50 Rosa suffulta .24 .46 .91 Rudbeckia hirta .10 .01 .03 Scirpus atrovirens .52 Scirpus fluviatilis 2.05 6.22 1.41 .08 Scutellaria leonardii .07 .04 .13 .08 Senecio pauperculus .42 3.15 3.92 .59 .35 Setaria lutescens .13 .01 .09 Setaria viridis .01 .03 .24 Silphium laciniatum 2.09 4.84 2.75 2.10 Solidago canadensis .66 3.98 6.02 5.68 2.12 Solidago gymnospermoides .01 .31 .11 Solidago missouriensis .01 .09 Solidago rigida 1.81 3.20 5.44 Sorghastrum nutans .09 .17 .42 .08 Spartina pectinata 1.36 4.27 3.74 1.79 .84 .30 Sporobolus heterolepis .52 7.76 23.83 40.98 49.78 Stipa spartea .05 Teucrium canadense .03 1.07 .31 .01 Thalictrum dasycarpum .04 .44 .33 Vernonia fasiculata .59 Veronicastrum virginicum .02 Viola pedatifida .04 .08 Viola sp. .07 .12 .12 .08 Vicia americana .05 .05 Zizia aurea .21 1.43 3.28 4.18 2.74 Allium sp. .02 Aster novae-angliae .23 .02 Cacalia tuberosa .02 Ceanothus americana .01 Panicum implicatum .06 .09 Prenanthes racemosa .08 Solidago nemoralis .02 .13 .45 1.69 Solidago riddellii .03 .13 .69 .18 .03 Taraxacum officinale .09 Echinacea pallida .02 were recognizable. These groups are labeled A, B, C, and D, with group A corresponding to the noncalcareous and ridge entity de- scribed previously and made up of plants showing preference for Nicollet and Nicollet- Webster soils. Group B includes all but one of the calcareous soil types plus four non- calcareous types. The noncalcareous types are found at the periphery of the group and include Clarion, Clarion-Nicollet, Webster, and heavy Webster soil types. Group C in- cludes the Glenco-Okoboji and Okoboji soils, and group D includes Glenco and calcareous- Glenco soils. These last two groups corre- spond to class 1 for species showing response to the soil factor described above. January 1983 Brotherson: Kalsow Prairie 161 Table 3 continued. 7 8 9 10 11 12 13 14 15 4.3^.8 4.9-5.4 5.5-6.0 6.1-6.6 6.7-7.2 7.3-7.8 7.9-8.4 8.5-9.0 9.1-9.6 2.64 2.16 2.62 9.74 2.50 2.92 .50 .24 .67 .17 .13 .13 .09 .58 .12 4.87 2.14 5.00 .09 .12 .13 2.50 7.36 9.25 26.55 .71 22.50 .13 27.86 26.67 38.50 61.25 .05 1.00 .83 .13 .71 2.92 .50 .24 .17 1.46 2.75 2.11 2.03 1.50 2.62 1.18 1.25 .09 .36 .57 2.50 .79 2.41 4.88 .13 .61 2.22 3.50 2.38 4.34 1.58 11.43 2.92 2.50 7.50 5.66 3.17 2.74 .79 .09 .26 .05 42.74 43.08 26.90 17.63 33.21 8.75 8.00 .09 .08 .95 .26 .13 .50 1.25 .42 .09 .05 .09 2.36 .08 .25 1.31 .26 .13 .42 .42 .05 .05 .50 1.97 2.14 2.50 24.50 Ordination of elevation data (Fig. 15) showed no recognizable groupings. Instead it separated the different elevation classes (Table 3) along a curve, point 14 represent- ing the ridge tops and point 1 representing the bottom of the potholes. This would tend to support statements made earlier that the vegetation of Kalsow Prairie is best repre- sented by the continuum concept of Curtis and Mcintosh (1951). The definable subcommunities or groups (Fig. 14) as based on soils data represent the response of the different taxa in the vegeta- tion to an environmental stimulus (i.e., carbo- nate soils) that is not distributed along gradients (i.e., at 9 X 9 m sampling levels) 162 Great Basin Naturalist Vol. 43, No. 1 Table 4. Average cover values for species in relation to soil series in 20-acre intensive study area. Species A A" N Na N" Na" W Achillea lanulosa .14 .19 .57 .44 .21 .37 Agropyron smithii .01 Ambrosia artemisifolia Amorpha canescens 3.47 2.87 4.16 4.06 6.57 2.56 .56 3.47 2.87 4.16 4.06 6.57 39.72 19.25 12.26 21.56 10.04 .08 Andrcypogon gerardi 39.72 19.25 12.26 21.56 10.04 10.63 6.70 Schizachyrium scoparius .08 .80 Anemone canadensis .03 Apocynum sibiricum .09 Arabis hirsuta Artemisia ludoviciana 3.19 .56 .38 1.88 .12 Asclepias incamata Asclepias sullivantii Asclepias syriaca Asclepias tuberosa Aster ericoides Aster laevis Aster simplex Baptisia leucophaea Calamagrostis canadensis Carex atherodes Carex aquatilis Carex gravida Carex lasiocarpa Carex retrorsa Chenopodium album Cirsium altissimum Comandra umbellata Convolvulus sepium Desmodium canadense Elymus canadensis Equisetum kansanum Eryngium yuccifolium Fragaria virginiana Galium obtusum Gentiana andrewsii Helenium autumnale Helianthus grosseserratus Helianthus laetiflorus Helianthus maximiliani Heliopsis helianthoides Lactuca scariola Lathyrus palustris Lathyrus venosus Lespedeza capitata Liatris pycnostachya Lithospermum canescens Lycopus americanus Lysimachia chiliata Lysimachia hybrida Lysimachia quadriflora Lythrum alatum Mentha arvensis Muhlenbergia racemosa Oxalis stricta Panicum capillare Panicum leibergii Panicum virgatum Pedicularis canadensis Petalostemum candidum . 19 .09 Petalostemum purpureum .65 .12 .21 .28 .01 .65 .61 1.88 .73 .19 3.89 1.94 3.58 2.19 .42 1.50 3.47 4.44 1.56 9.68 1.29 .08 2.50 .46 2.12 .14 .85 .33 1.88 .24 .48 .03 10.48 .75 1.11 .01 .28 2.07 .01 .60 .14 .19 .19 .08 .21 .05 .97 .28 .04 .65 1.88 2.19 2.71 1.64 .28 .19 .24 2.19 .44 1.04 .60 .14 .05 .61 .31 .44 .65 .42 .12 .38 .14 .19 .24 .16 2.15 .14 .65 .42 1.83 .01 .03 1.65 .31 2.58 1.25 9.26 3.19 6.20 3.87 .28 6.87 2.18 .24 4.79 .18 .03 .01 .08 .56 .09 .65 .24 1.88 .21 .14 .31 1.21 .40 .28 .56 .09 .52 .31 1.88 .56 .04 1.04 .21 .21 .17 .11 .03 .01 .12 .23 .01 6.67 7.50 2.64 4.06 .77 3.33 1.20 1.20 .14 .44 .04 2.71 .49 January 1983 Brotherson: Kalsow Prairie 163 Table 4 continued. Wh Wa H- Ga GO .42 .40 .07 .02 .09 .65 .10 .09 .22 .17 .09 .73 1.93 .33 .33 .04 .02 12.19 9.09 10.87 14.42 11.57 14.25 .05 .13 .63 .77 .46 1.74 .02 1.31 .02 .73 .17 .03 .71 .09 .02 .18 .64 .25 .02 .16 .07 .81 .74 2.06 3.50 2.63 1.88 2.81 1.22 1.70 2.81 1.18 2.59 .31 2.42 .55 1.76 3.54 1.02 .17 1.36 1.63 1.18 3.98 1.80 2.38 11.15 2.46 1.41 .81 13.61 45.26 56.88 1.50 .63 .25 7.73 8.25 32.38 1.88 .20 .11 .64 1.48 3.90 3.00 1.04 .04 .16 .25 .39 2.48 3.88 .25 6.88 .37 .58 1.23 1.38 8.27 10.63 6.00 .10 .77 .50 .67 .68 .19 .35 .45 .64 .16 .30 .09 .10 .48 .14 1.35 4.66 3.53 2.79 4.15 2.69 .14 .31 1.08 1.10 .56 1.10 .37 .07 .10 .31 .26 .15 .02 .11 .17 .52 2.24 1.62 .96 .98 1.57 .07 .13 2.08 1.42 2.43 .18 .31 1.52 .58 2.16 .30 1.85 .09 .14 .13 13.85 5.17 9.96 7.86 6.31 12.41 3.36 4.63 .10 .20 .02 .04 .34 .31 .18 .34 .65 .06 .02 .10 .06 .05 .09 .02 .10 .23 .63 1.02 .72 .67 .42 .28 .10 .48 .33 .22 .21 .09 .10 .03 .02 .02 .18 .04 .09 .05 .13 .10 .31 .31 .03 .28 .02 .11 .04 .20 .42 1.47 .43 2.70 1.57 .05 .68 .80 1.03 .51 .30 .20 .46 .45 .25 .06 .02 .60 .50 .87 .42 .09 .93 .83 .28 .19 .25 .02 1.31 .05 .75 21.50 17.25 164 Great Basin Naturalist Vol. 43, No. 1 Table 4. Average cover values for species in relation to soil series in 20-i acre intensive study area. Species A a- N Na N- Na- w Phalaris arundinacea 1.09 Phlox pilosa .19 .16 .03 Physalis heterophylla 2.31 .28 1.88 Physalis virginiana .14 .09 .28 .08 .03 Poa pratensis 16.53 31.09 5.99 32.81 5.21 3.22 Polygonum coccineum .44 Potentilla arguta .14 .56 .05 Psoralea argophylla 1.81 1.30 .15 .31 .08 .21 Pycnanthemum virginianum .28 .43 Ratibida pinnata 2.92 3.24 .05 .32 1.46 1.29 Rosa suffulta .14 3.06 1.56 2.19 1.67 .29 Rudbeckia hirta .04 Scirpus atrovirens Scirpus fluviatilis .01 Scutellaria leonardii .09 .05 .08 .14 Senecio pauperculus 1.24 Setaria lutescens .56 Setaria viridis .56 .31 Silphium laciniatum .38 1.25 2.15 Solidago canadensis .14 2.41 3.21 .31 2.86 1.25 3.16 Solidago gymnospermoides .09 .69 1.88 Solidago missouriensis 1.67 .56 .28 Solidago rigida .56 3.25 .31 4.27 2.50 3.79 Sorghastrum nutans .31 .20 .11 Spartina pectinata .04 .83 Sporobolus heterolepis 12.08 21.76 62.69 37.19 58.95 27.92 37.41 Stipa spartea .97 .09 .14 .08 .21 .05 Teucrium canadense .05 .21 .23 Tahlictrum dasycarpum .05 Vemonia fasciculata .09 Veronicastrum virginicum .01 Viola pedatifida .09 .12 .05 Viola sp. .03 Vicia americana .28 .09 .05 .31 .04 .03 Zizia aurea .14 .93 4.58 4.06 1.73 3.33 1.52 Allium sp. Aster novae-angliae .09 Cacalia tuberosa .01 Cearwthus americana 10.56 .56 .04 Panicum implicatum .04 .15 Prenanthes racemosa .09 Solidago nemoralis .28 .93 .75 Solidago riddellii .01 Taraxacum officinale but in mappable units with fairly discrete boundaries. This would tend to cause vegeta- tion sensitive to carbonate influence to group accordingly. An ordination of species, utilizing the data from Tables 3 and 4, isolated taxa having dis- tinct distribution patterns. These species are Amorpha canescens, Andropogon gerardi. As- ter ericoides. Aster laevis, Calamagrostis canadensis, Carex athorodes, Carex aquatilis, Desmodium canadense, Helianthus grosseser- ratus, Helianthus laetiflorus, Panicum lei- bergii, Phalaris arundinacea, Poa pratensis. Polygonum coccineum, Ratibida pinnata, Scirpus fluviatilis, Silphium laciniatum, Sol- idago canadensis, Solidago rigida, Spartina pectinata, Sporobolus heterolepis, Zizia aurea, and Ceanothus americanus, all of which show distinct distribution patterns and in many cases high preference for certain soil groups or elevations. The relationships between elevation and soil series are shown in Figure 16. The soil types are positioned along the base line as January 1983 Table 4 continued. Brotherson: Kalsow Prairie 165 Wh Wa H- c- Ga GO 1.56 .63 1.35 .10 .10 .10 .21 3.65 7.40 2.50 .63 .10 1.88 29.38 .73 .21 .31 .10 .20 .06 2.81 .06 .45 1.73 .68 .11 .48 2.95 4.12 .26 5.34 .17 .28 44.40 .28 .09 .09 3.41 .09 .02 3.86 .99 3.05 .42 .09 .04 2.45 3.84 7.30 3.77 .35 .15 25.42 .15 .20 .31 .02 5.31 .13 .07 .05 3.15 2.17 1.94 .20 .16 .09 3.59 4.69 7.39 .05 1.36 .63 .54 24.08 .38 .38 .09 .02 4.64 .02 .05 .17 3.18 .25 1.99 1.91 .25 .09 2.84 .25 4.79 5.30 .09 .34 .85 32.80 .25 .17 .04 .25 5.13 .04 .09 5.74 .83 .09 .09 9.17 3.61 13.43 1.20 .09 .56 9.44 .09 .09 .09 3.52 .56 6.24 .05 8.74 6.63 .75 40.50 36.75 .35 1.87 .14 .28 .05 7.22 .79 .30 .49 13.63 17.50 .13 .75 .13 3.38 .13 .73 2.92 1.02 .09 .04 .31 .50 .47 .45 .17 .03 .19 they appeared in the field. In all cases where the noncalcareous soils had adjacent cal- careous variants the calcareous variants showed higher average elevations. Summary and Conclusions 1. Sporobolus heterolepis is the dominant plant of the upland prairie that places Kal- sow Prairie within the "Consociation" desig- nated by Weaver and Fitzpatrick (1934) as the Prairie-Dropseed type. 2. The vegetation of the upland prairie communities is best described and represent- ed by the continuum concept as described by Curtis (1955). 3. The vegetation of the upland prairie has changed since Moyer's 1953 study. Species showing increased importance in my study are Solidago canadensis, Solidago rigida, Pan- icum leibergii, Helianthus grosseserratus, and Fragaria virginiana. Species decreasing in im- portance were Phleum pratense, Poa pra- tensis, Zizia aurea, Schizachyrium scoparius. 166 Great Basin Naturalist Vol. 43, No. 1 tc Fig. 15. Two-dimensional ordination of vegetation found at different elevations in the 20-acre study area; 1 = .7-1.2 feet elevation, 2 = 1.3-1.8 feet elevation, 3 = 1.9-2.4 feet elevation, 4 = 2.5-3.0 feet elevation, 5 = 3.1-3.6 feet elevation, 6 = 3.7-4.2 feet elevation, 7 = 4.3-4.8 feet elevation, 8 = 4.9-5.4 feet elevation, 9 = 5.5-6.0 feet elevation, 10 = 6.1-6.6 feet elevation, 11 = 6.7-7.2 feet elevation, 12 = 7.3-7.8 feet elevation, 13 = 7.9-8.4 feet elevation, 14 = 8.5-9.0 feet elevation. Panicum virgatum, Sorghastrum nutans, and Sporobolus heterolepis. 4. Soil series, elevations, and species distri- bution patterns were mapped on an 8 ha in- tensive study plot. Elevation and soils data are correlated with species distribution pat- terns. All species show a response. Nine gen- eral patterns of distribution are described, with the following species as examples: a. Andropogon gerardi—species of wide distribution, limited only by conditions peculiar to the drainage areas of the prairie. b. Silphium laciniatum—a. pattern closely resembling that of Andropogon gerardi but showing limited distribution on the higher and drier ridges. c. Ambrosia artemisifolia— species limited to the border weed communities. d. Amorpha canescens—a. pattern common to species limited to the ridges and lower slopes. e. Solidago nemoralis— species limited to mid- and upland slopes of the prairie. f. Ceanothus americanus—a. pattern limit- ed to the ridges and drier sites of the prairie. g. Helenium awfumna/e— limited to growth on soils that are highly cal- careous to the surface. h. Calamagrostis canadensis— limited to growth along the shallower areas of the pothole and drainage system. i. Scirpus fluviatilis— growth corresponds to deeper areas within the drainage system. 5. Species occurring in the intensive study were ordinated using Orloci's (1966) method. The technique did not delineate associated groups of species, yet it pointed out species exhibiting peculiar distribution patterns. Such species are useful as indicator species. 6. Indices of interspecific association were computed for all participating species (Cole 1949) and found to be extremely useful in identifying clusters or groups of species hav- ing similar ecological amplitudes. Literature Cited AiKMAN, J. M. 1959. Prairie research in Iowa. The Amer- ican Biology Teacher 21:7-8. Aikman, J. M., AND R. F. Thorne. 1956. The Cayler Prairie: an ecologic and taxonomic study of a northwest Iowa prairie. Iowa Acad. Sci. 63:177-200. Brennan, K. M. 1969. Vertebrate fauna of Kalsow Prairie. Unpublished thesis. Iowa State Univ. H G„ GO O A A- N N„ N- 1^ W V^t, \% C SOIL SERIES Fig. 16. Relationships between elevation and soil series as found in 20-acre intensive study area; mean value in- dicated for each soil by short horizontal line. January 1983 Brotherson: Kalsow Prairie 167 Cain, S. A., and F. C. Evans. 1952. The distribution patterns of three plant species in an old-field community in southeastern Michigan. Contribu- tions from the Laboratory of Vertebrate Biology No. 52. Cole, L. C. 1949. The measurement of interspecific as- sociation. Ecology 30:411-424. Collins, G. B. 1968. Implications of diatom succession in postglacial sediments from two sites in north- em Iowa. Unpublished dissertation. Iowa State Univ. Crawford, R. M. M., and D. Wishart. 1968. A rapid classification and ordination method and its ap- plication to vegetation mapping. Ecology 56:385-404. Curtis, J. T. 1955. A prairie continuum in Wisconsin. Ecology 36:558-566. Daubenmire, R. 1959. A canopy-coverage method of vegetational analysis. Northwest Science 33:43-66. Dick-Peddie, W. a. 1955. Presettlement forest types in Iowa. Unpublished thesis. Iowa State Univ. Due, R. L., AND J. E. Butler. 1960. A phytosociological study of a small prairie in Wisconsin. Ecology 41:316-327. Ehrenreich, J. H. 1957. Management practices for maintenance of native prairies in Iowa. Unpub- lished dissertation. Iowa State Univ. Esau, K. L. 1968. Carabidae (Coleoptera) and other ar- thropods collected in pitfall traps in Iowa corn- fields, fencerows, and prairies. Unpublished dis- sertation. Iowa State Univ. GiLLY, C. L. 1946. The Cyperaceae of Iowa. Iowa State Coll. J. Sci. 21:55-151. Gleason, H. a. 1952. The new Britton and Brown illus- trated flora of the northeastern United States and adjacent Canada. Lancaster Press, Lancaster, Pennsylvania. 3 vols. Greig-Smith, p. 1964. Quantitative plant ecology. But- terworth. Washington, D.C. Hale, M. E., Jr. 1955. Phytosociology of corticolous cryptograms in the upland forests of southern Wisconsin. Ecology 36:45-62. Hayden, Ada. 1943. A botanical survey in the Iowa lake region of Clay and Palo Alto counties. Iowa State Coll. J. Sci. 17:277-416. 1945. The selection of prairie areas in Iowa which should be preserved. Proc. Iowa Acad. Sci. 52:127-148. 1946. A progress report on the preservation of prairie. Proc. Iowa Acad, of Sci. 53:45-82. Hewes, L. 1950. Some features of early woodland and prairie settlement in a central Iowa county. An- nals Assoc. Amer. Geog. 40:40-57. HuRLBERT, S. H. 1969. A coefficient of interspecific asso- ciation. Ecology 50:1-9. Kennedy, R. K. 1969. An analysis of tall-grass prairie vegetation relative to slope position, Sheeder Prairie, Iowa. Unpublished thesis. Iowa State Univ. Kershaw, K. A. 1964. Quantitative and dynamic ecolo- gy. Edward Arnold, Ltd., London, England. Landers, R. Q. 1966. Visit the virgin prairie. Iowa Farm Science 21:418-419. MoYER, John F. 1953. Ecology of native prairie in Iowa. Unpublished dissertation. Iowa State Univ. Norton, D. C, and P. E. Ponchillia. 1968. Stylet-bear- ing nematodes associated with plants in Iowa prairies. J. Iowa Acad. Sci. 75:32-35. Orloci, L. 1966. Geometric models in ecology. I. The theory and application of some ordination meth- ods. J. Ecology 54:193-215. Oschwald, W. R. et al. 1965. Principal soils of Iowa. Special Report No. 42, Department of Agronomy, Iowa State Univ. Pohl, R. W. 1966. The grasses of Iowa. Iowa. Iowa State J. Sci. 40:341-566. Richards, M. S. 1966. Observations on responses of prairie vegetation to an April fire in central Iowa. Unpublished thesis. Iowa State Univ. RuHE, R. V. 1969. Quaternary landscapes in Iowa. Iowa State Univ. Press, Ames Iowa. 249 pp. Rydberg, p. A. 1931. A short phytogeography of the prairies and great plains of central North Ameri- ca. Brittonia 1:57-66. Sanders, D. R. 1969. Structure and pattern of the her- baceous understory of deciduous forests in cen- tral Iowa. Unpublished dissertation. Iowa State Univ. ScHMiTT, D. P. 1969. Plant parasitic nematodes and nematode populations in the Kalsow Prairie. Un- published thesis. Iowa State Univ. Shimek, B. 1911. The prairies. Bull. Lab. Nat. Hist., State Univ. Iowa 6:169-240. Shimek, B. 1915. The plant geography of the Lake Oko- boji region. Bull. Lab. Nat. Hist., State Univ. Iowa 7:1-90. Shimek, B. 1925. The persistence of the prairie. Univ. of Iowa Studies 11:3-24. Steiger, T. L. 1930. Structure of prairie vegetation. Ecology 11:170-217. U.S. Government. 1868. First survey of the State of Iowa. Plats deposited in the State House, Des Moines. Weaver, J. E. 1930. Underground plant development in its relation to grazing. Ecology 11:543-557. Weaver, J. E. 1954. North American prairie. Lincoln, Nebraska, Johnsen Publishing Company. 348 pp. Weaver, J. E., and F. E. Clements. 1938. Plant Ecolo- gy. 2d ed. McGraw-Hill, New York. Weaver, J. E., and T. J. Fitzpatrick. 1932. Ecology and relative importance of the dominants of the tall- grass prairie. Bot. Gaz. 93:113-50. Weaver, J. E., and T. J. Fitzpatrick. 1934. The prairie. Ecol. Monogr. 4:109-295. DEER MOUSE, PEROMYSCUS MANICULATUS, AND ASSOCIATED RODENT FLEAS (SIPHONAPTERA) IN THE ARCTIC-ALPINE LIFE ZONE OF ROCKY MOUNTAIN NATIONAL PARK, COLORADO R. B. Eads' and E. G. Campos' Abstract.— Peromyscus maniculatus and related small rodents have been examined for ectoparasites in the tun- dra region of the Rocky Mountain National Park 1974-1979. One hundred and ninety-four P. maniculatus were ex- amined from two tundra sites. Flea infestation rates were 1.9 fleas per mouse examined and 4.1 fleas per infested mouse. Species taken in significant numbers were Monopsyllus thambus (51 percent), Peromyscopsylla hesperomys (34 percent), Malaraeus euphorbi (9 percent), and Catallagia calisheri (4 percent). Peromyscus maniculatus host/flea relationships in the tundra are compared with those in other life zones in the park. Personnel of the Vector-Borne Diseases Di- vision have studied ectoparasite/host rela- tionships in the Rocky Mountain National Park (RMNP) for a number of years. Of prin- cipal interest have been vectors and reser- voirs of Yersinia pestis and Colorado tick fe- ver virus. All life zones in the Park have been investigated. Because of a dearth of published information concerning fleas and other ecto- parasites of rodents in the arctic-alpine life zone of the Rocky Mountains, data obtained during the period 1974-1979 in the zone are presented here. Emphasis has been placed on the deer mouse, Peromyscus maniculatus, as it is by far the most abundant rodent in the zone. Perom.yscus maniculatus fleas in the tundra are compared with those in other life zones. Study Area The RMNP is located in Larimer and Boul- der counties in north central Colorado. It covers 1046 km^ of mountainous terrain, with elevations ranging from 2400 to over 4300 m. Regional ecosystems of north central Colo- rado, with approximate elevations as defined by Marr (1961), include the grassland-lower montane ecotone region (1707-1829 m). lower montane forest climax region (1829-2347 m), lower montane-upper mon- tane ecotone region (2347-2438 m), upper montane forest climax region (2438-2743 m), upper montane-subalpine ecotone region (2743-2835 m), subalpine forest climax re- gion (2835-3353 m), subalpine-alpine eco- tone region (3353-3475 m), and alpine tun- dra climax region (3475 m-mountain tops). Collections in the tundra were concen- trated at sites 1 and 2, located as shown in Figure 1. Site 1 is 2.5 km west of Rainbow Curve on Trail Ridge Road at an elevation of around 3475 m. Site 2 is on Fall River Road, some 1.5 km below the Alpine Visitors Cen- ter at an elevation of approximately 3523 m. Sites 1 and 2 are in areas with limited human activity. Trapping was confined to an area of 5 or 6 ha in both locations. In the tundra (Fig. 2) short grasses, sedges, and forbes pre- dominate in the exposed meadows. Some- what taller grasses and dwarf shrubs are found in low-lying areas partially protected from the violent winds that sweep the tundra. Methods Mammals were captured in the tundra, primarily in Sherman live traps (7.62 X 7.62 'Veclor-Bome Diseases Division, Center for Infectious Diseases, Centers for Disease Control, Public Health Service, U.S. Department of Health and Hu- man Services, Post Office Box 2087, Fort Collins, Colorado 80522-2087. 168 January 1983 Eads, Campos: Colorado Fleas 169 V "■■■ f ^////////////////////////////////i//////////////////////////A Fig. 1. Flea collection sites in the Rocky Mountain National Park, Larimer County, Colorado. X 22.86 cm), with National live traps (12.7 X 12.7 X 40.64 cm) also used on occasion. No attempts were made to collect larger mammals. Traps baited with rolled oats were set in parallel lines of 20 stations at approx- imately 20 m intervals. Traps were placed adjacent to rocky outcroppings, when avail- able, to provide trapped rodents some pro- tection from the elements. The number of lines varied, but there were usually 4 or 5, 25 m apart. Traps were set in the morning and inspected the following day. Trapping in the tundra was limited to the summer months. Trail Ridge and Fall River roads were only open to vehicular traffic June-September, with some closures at ir- regular intervals during this period due to hazardous driving conditions. As the Sherman traps were picked up, those containing rodents were placed in plas- tic bags to prevent loss of fleas during trans- port to a field laboratory. Each mammal was lightly anesthetized witii ether and brushed in a white enamel pan to remove ectopara- 170 Great Basin Naturalist Vol. 43, No. 1 Fig. 2. Tundra region, Rocky Mountain National Park. sites. Certain animals were bled, ear tagged, and released, and serological tests for plague and Colorado tick fever were performed. Others were held for more extensive ecto- parasite recovery efforts, including exam- ination of the mice under a dissecting micro- scope. Fleas collected were held in 2 percent saline solution for plague testing or in 70 per- cent alcohol for definitive taxonomic studies. Results and Discussion Trapping over a six-year period (1974-79) has provided considerable information on deer mouse host/ flea relationships in the Col- orado tundra region. As shown in Table 1, 156 P. maniculatus were captured in this pe- riod during the summer months at Site 1. An average of 8 P. maniculatus were taken per 100 trap nights. About 47 percent were in- fested with fleas, with a mean number of 2.1 fleas per animal examined and 4.4 per in- fested animal. Site 2 was trapped less fre- quently than Site 1. As shown in Table 2, the flea burden was 1.1 fleas per mouse and 2.9 fleas per infested mouse. Ectoparasites other than fleas were in- frequently encountered on deer mice in the timdra. Fewer than 10 percent were infested, usually lightly, with the sucking louse, Hopl- opleura hesperomydis. Larval chigger mites were somewhat more prevalent. Neo- trombicula harperi, N. microti, and Eu- schoengastia guntheri were taken. A single larval argasid tick was recovered from a P. maniculatus above the tree line, probably an Argas sp. The second most prevalent rodent taken in the tundra was the heather vole, Phenacomys intermedius. Twenty-three were examined, and 65 percent were infested with three spe- cies of fleas at the rate of 2.4 fleas per animal and 3.6 per infested animal. Species included Peromyscopsylla selenis (67 percent), Mega- bothris abantis (19 percent), and Monopsyllus thambus (15 percent). Two species of chigger mites, N. harperi and E. guntheri, were recov- ered from heather voles. One vole was heavi- ly infested and several lightly infested with the dermanyssid mite Hirstionyssus isabellinus. January 1983 Eads, Campos: Colorado Fleas 171 Table 1. Peromyscus maniculatus examined for fleas in the tundra region of the Rocky Mountain National Park (Site 1). Percent Mean traps with Number Number with Total Mean number number fleas/ Date P. manic. P. manic. fleas fleas fleas/animal infested animal 7-16-74 9 11 6 19 1.73 3.17 7-24-74 8 6 3 31 5.17 10.33 8-14-74 10 10 4 11 1.1 2.75 8-15-74 8 6 3 16 2.67 5.33 9-26-74 15 18 4 6 .33 1.5 8-22-75 14 17 11 19 1.12 1.73 8-18-76 6 7 3 7 1.0 2.33 6-29-77 7 2 2 10 5.0 5.0 8-17-77 5 10 4 22 2.2 5.5 9-02-77 5 4 3 6 1.5 2.0 9-09-77 10 4 1 4 1.0 4.0 6-27-78 5 6 4 26 4.33 6.5 7-07-78 3 3 1 2 .67 2.0 7-25-78 5 4 2 12 3.0 6.0 8-03-78 3 4 4 6 1.5 1.5 9-07-78 16 17 7 22 1.29 3.14 7-10-79 7 7 3 16 2.29 5.33 7-18-79 6 6 3 54 9.0 18.0 8-24-79 12 14 6 36 2.57 6.0 8.11 156 74 325 2.08 4.39 Lesser numbers of the least chipmunk, Eu- tamias minimus (7); golden-mantled ground squirrel, Spermophilus lateralis (3); yellow- bellied marmot, Marmota flaviventris (2); and pika, Ochotona princeps (3) were captured and examined. Eleven fleas, 10 Monopsyllus eumolpi, and one M. thambus were removed from 4 of the 7 chipmunks. The 2 marmots were infested with 7 Thrassis stanfordi and 2 Oropsylla ru- pestris. Three Oropsylla idahoensis were re- covered from 2 of the 3 golden-mantled ground squirrels. The 3 pikas examined were infested with the following fleas: 46 Am- phalius necopinus, 46 Ctenophyllus terribilis, and 1 M. thambus. The chigger mite, N. mi- croti, was also taken from all 3 pikas. Peromyscus maniculatus host/flea relation- ships in the tundra have proven to be dis- tinctive relative to the other life zones in the Table 2. Peromyscus maniculatus examined for fleas in the tundra region of the Rocky Mountain National Park (Site 2). Number of mice Number with Number and species Date trapped fleas of fleas 7-17-74 3 1 2 Monopsyllus thambus 7-24-74 4 2 3 M. thambus 1 Malaraeus euphorbi 8-17-77 10 4 9 M. thambus 7 M. euphorbi 4 Peromyscopsylla hesperomys 1 Megabothris abantis 1 Catallagia calisheri 9-02-77 4 3 5 P. hesperomys 2 M. thambus 1 C. calisheri 9-09-77 15 4 3 M. thambus 2 P. hesperomys 1 M. euphorbi 9-24-77 2 1 1 Peromyscopsylla selenis Totals 38 15 43 172 Great Basin Naturalist Vol. 43, No. 1 RMNP. As shown in Table 3, 21 species were recovered in 1974 and 20 in 1975 in all life zones. This is well over twice the number of species found to be parasitizing deer mice in the tundra. The 2 X ratio remains virtually unchanged when the recoveries are limited to normal or true parasites of deer mice, ex- cluding species that have strayed from envi- rormiental associates. Monopsylhis wagneri was the principal flea below the treeline (68 percent); over 60 percent of the deer mice were infested in 1974-1975. The only other prevalent species were Malaraeus telchinum (10 percent of to- tal fleas) and Opisodasys keeni (8 percent of total fleas). Each of the other 11 species nor- mally parasitic on deer mice made up 1-5 percent of the total fleas. As shown in Table 4, mean infestation rates for the 2,090 P. ma- niculatus examined in all life zones in the RMNP in 1974-76 were 1.2 fleas per mouse and 2.5 fleas per infested mouse. Higher flea infestation rates per deer mouse were obtained in the tundra than in the other life zones. This may be a reflection of more intensive examination of the mice in the tundra. Also, tundra mice were usually held for several examinations, and in the other life zones normal procedure was to in- spect them once, tag and release them. A to- tal of 194 deer mice were examined from the two tundra sites, and 368 fleas were recov- ered from 89 of them. This is a rate of 1.9 fleas per mouse examined and 4.1 fleas per infested mouse. In the tundra, fewer species were involved in parasitizing deer mice than in the other life zones. Only four were taken in significant numbers. Monopsyllus wagneri disappears completely from the mice in the tundra and is replaced by M. thambus (51 percent of to- tal fleas at Site 1 and 44 percent at Site 2). Overlapping of the two species occurred at Rainbow Curve, elevation approximately 3290 m. This scenic overlook consists of an exposed rocky slope with large boulders bor- dered by subalpine forest. Table 3. Species of fleas from Peromyscus maniculatus in all life zones in Rocky Mountain National Park in 1974-1975. 1974 1975 Number offleas-1,018 Number of fleas- 1,357 Percent of Percent of animals Percent of Percent of animals Species total fleas with species total fleas with 5 ipecies Ceratophyllidae Malaraeus euphorbi M. telchinum 5.9 1.3 7.3 1.5 14.5 1.3 17.5 Megabothris abantis Monopsyllus eumolpi M. tliambus < < 4.1 < < 1 1 3.6 < < < 1 1 1 < < < M. wagneri M. vison ( < 84.8 I < 60.8 1 < 70.3 1 1 < 60 Opisocrostis Labis Opisodasys keeni Orchopeas leucopus Oropsylla idahoensis < < 9.9 < < 1 9.5 1 1.7 < < 1 5.8 1 < 7.3 1.6 Hystrichopsyllidae Callistopsyllus deuterus Catallagia calisheri < — < 1 < < 1 1 < < C. decipiens C. neweyi Epitedia wenmanni < 4.4 2.4 < 3.9 1 4.1 < 1.8 1 2 < 3.6 Hystrichopsylla occidentalis Megarthroglossus sp. M. divisus < < < < 1 1 < < 1 1 < < - Rhadinopsylla sectilis < < 1 < 1 < Leptopsyllidae Amphipsylla sibirica Peromyscopsylla hesperomys P. selenis < < 4.2 < < 1 4.5 1 < < 1 1 < 1.9 January 1983 Eads, Campos: Colorado Fleas 173 Table 4. Summary of flea collections from Peromyscus maniculatus from all life zones in Rocky Mountain Nation- al Park, 1974-1976. Number of mice Number of Percent Total Number of Number of examined mice of mice number of fleas/ fleas/ Year for fleas with fleas infested fleas mouse infested mouse 1974 955 431 45.1 1,018 1.1 2.4 1975 957 507 53.0 1.357 1.4 2.7 1976 178 80 44.9 183 1.0 2.3 Totals 2,090 1,018 48.7 2,558 1.2 2.5 Peromyscopsylla hesperomys was the sec- ond most prevalent P. maniculatus flea in the tundra (35 percent of the total fleas at Site 1 and 26 percent at Site 2). This flea is found on deer mice in all life zones of the RMNP but in much smaller numbers at the lower elevations, making up <5 percent of the to- tal fleas collected in the park. The third most common species on deer mice in the tundra, Malaeraeus euphorbi (7 percent of the total fleas at Site 1 and 21 per- cent at Site 2) was also more prevalent than at lower altitudes. Less than 2 percent of to- tal fleas collected in the park were this species. A nest flea, Catallagia calisheri, was a poor fourth on tundra deer mice. However, the fact that 4 percent of the fleas at Site 1 and 5 percent at Site 2 were this species is in- dicative of much greater numbers in the nests. Catallagia calisheri was not taken be- low the tree line. At lower elevations in RMNP, the common deer mouse nest Catal- lagia was decipiens. Single specimens of the nest fleas, Callistopsyllus deuterus and Me- garthroglossus sp. (female) were taken from deer mice in the timdra. Both genera were also rarely encountered in other RMNP life zones. Orchopeas leucopus, a common Per- omyscus parasite at lower elevations, was rare in the park. It was taken but once in the timdra and once in other RMNP life zones. The only other fleas taken from deer mice in the tundra were a few Megabothris abantis and Peromyscopsylla selenis, strays from heather voles. Conclusions Our data concerning the fleas of P. ma- niculatus support the conclusions of Wenzel and Tipton (1966) that the altitudinal ranges of many ectoparasites do not necessarily coincide with those of the hosts. Peromyscus maniculatus is prevalent in the RMNP at all elevations. However, vertical stratification is evident in the species of fleas parasitizing them, especially at the upper levels. In the tundra, around 3475 m, the number of flea species on P. maniculatus was greatly re- duced from the lower elevations, but the mean number per animal was somewhat higher. Only four species were encountered in significant numbers on P. maniculatus in the tvmdra. Monopsyllus thambus and Catal- lagia calisheri, found only on the tundra, re- placed M. wagneri and C. decipiens of lower elevations, respectively. Peromyscopsylla hes- peromys and Malaraeus euphorbia, found at all elevations studied in RMNP, were signifi- cantly more prevalent on P. maniculatus in the tundra than at lower elevations. Acknowledgments Excellent cooperation has been received from naturalists of the National Park Service, U.S. Department of Interior, during these studies. Ray Bailey, Vector-Borne Diseases Division statistician, was most helpful in sup- plying computerized data. Gary Maupin took the tundra photograph. Numerous persons, presently and formerly with VBDD, contrib- uted to the field investigations. Included were Dr. R. G. McLean, Dr. A. M. Barnes, Ronald Shriner, Karen Pokorny, Gary Mau- pin, Leon Carter, Edwin Heidig, William Ar- chibald, and David Pegg. Taxonomic assistance was kindly provided by the following authorities: Dr. J. E. Keirans (ticks). Dr. R. B. Loomis (chigger mites), and Dr. K. C. Emerson (sucking lice). 174 Great Basin Naturalist Vol. 43, No. 1 Literature Cited Wenzel, R. L., and v. J. Tipton. 1966. Some relation- ships between mammal hosts and their ectopara- Marr, J. W. 1961. Ecosystems of the east slope of the sites. Pages 677-723 in Ectoparasites of Panama, front range in Colorado. Univ. of Colorado Stud- Field Mus. Nat. Hist. Chicago, Illinois, ies (Series in Biology) No. 8. Univ. of Colorado Press. Boulder. FOOD OF LARVAL TUI CHUBS, GILA BICOLOR, IN PYRAMID LAKE, NEVADA' David L. Galat* and Nancy Vucinich^ Abstract.— Four near-surface locations in Pyramid Lake, Nevada, were sampled for larval tui chubs {Gila bicolor) during summer and early fall 1979. Numbers of larvae collected were highest in mid-July. Zooplankton was the only food eaten throughout the survey; the cladoceran Moina hutchinsoni was the major species eaten at all locations. An- other cladoceran, Diaphanosoma leuchtenbergianum, was also important to the diet of pelagic larvae, and the cope- pod Cyclops vemalis was eaten in significant quantities by nearshore fish. Changes in diet composition of larval tui chubs during summer corresponded to seasonal succession of zooplankton species in Pyramid Lake. The tui chub, Gila bicolor, is a polytypic cyprinid native to drainage systems in Ore- gon, California, and Nevada (Moyle 1976). It is the most abundant fish in Pyramid Lake, Nevada, where adults constitute over 90 per- cent of gill net catches (Vigg 1978, 1981). A major role of this species in the economy of Pyramid Lake is as forage for the primary sport fish, the Lahontan cutthroat trout (Sal- mo clarki henshawi), which commonly attains trophy weights of over 3 kg (Snyder 1917, Kucera 1978, Galat et al. 1981). LaRivers (1962), Langdon (1979), and Vu- cinich et al. (1981) provided information on food of adult and juvenile tui chubs in Pyra- mid Lake; Miller (1951), Kimsey (1954), and Cooper (1978) presented similar information for this species in other waters; and Williams and Williams (1980) described the food of re- lated Gila species. We present data on abun- dance, distribution, and food of the pre- viously uninvestigated larval phase of the tui chub in Pyramid Lake. Methods Two shallow littoral stations in Pyramid Lake (40° 00' N, 119° 35' W), one northeast and the other southeast of Sutcliffe, Nevada, were quantitatively sampled for larval tui chubs between 18 June and 9 August 1979. A metered 1 mm-mesh net, 0.5 m in diameter, was hand-towed along the surface at about the 1 m depth contour and parallel to shore. However, only 2 of 10 attempts at collecting larval tui chubs in the shallow littoral region were successful and only three larvae were captured. Because so few fish were recovered from these locations they are not discussed further. Two deeper littoral locations were also quantitatively sampled for larval fishes: a sur- face pelagic station, northeast of Sutcliffe, Nevada, at the 72 m depth contour, and a surface nearshore station, southeast of Sut- cliffe at the 5-10 m depth contour, were sampled every two weeks from 26 June through 26 October 1979. Larvae were col- lected with the 0.5 m net described above, towed 1-3 m below the surface behind a boat. Fish collected were immediately killed in MS-222 to minimize regurgitation and preserved in 10 percent formalin. After identifying larvae as tui chubs and measuring their fork lengths (FL), the entire digestive tract was removed. Contents from a maximum of 10 nonempty larval tracts were pooled for each date and station where more than two fish were collected. All zooplank- ters recovered from larval fish were identi- fied to species and enumerated under a com- pound microscope at 40-200X. Algae and unidentifiable matter (e.g., detritus and di- gested material) were never observed in sub- stantial amounts and were not quantified. Numbers of organisms recovered from guts were converted to carbon equivalents based on the average carbon content of whole 'The Colorado Cooperative Fishery Research Unit is supported by the U.S. Fish and WildUfe Service, Colorado Division of Wildlife, and Colorado State University. "Cooperative Fishery Research Unit, Colorado State University, Fort Collins, Colorado 80523. 'Pyramid Lake Indian Tribal Enterprises, Star Route, Sutcliffe, Nevada 89510. 175 176 Great Basin Naturalist Vol. 43, No. 1 60 55 ■ CO O '>< 25 fO E « 20 H o t ^ 15 H o Z 10 - 5 - • Pebgic station o- — o Near- shore station ,.--- c-. o Jun Jul Aug Sep Oct Fig. 1. Numbers of tui chub larvae collected near the surface at two locations in Pyramid Lake, Nevada, 1979. organisms as measured with a Hewlett-Pack- ard model 185B CHN analyzer, following Sharp's (1974) procedures. Because carbon values were not obtained for Daphnia schod- leri or copepod nauplii, these organisms were assigned approximate carbon equivalents rel- ative to their intact size. We selected this ap- proach over volumetric or weight techniques because it more accurately reflected the bioenergetic significance of a food item to the fish's diet. Results and Discussion The length range of larval tui chubs cap- tured was 6-15 mm. Following Snyder's (1976) terminology, only mesolarvae (<12 mm FL) and metalarvae (>12 mm) were identified from our collections. Yolk sacs were not apparent in any larvae. The alimen- tary tract was straight and tubelike in larvae up to 13 mm long; in progressively larger fish it began to loop and swell anteriorly. The transition of fish from metalarva to juvenile occurred at a length of about 15 mm. Estimated larval abundance at the pelagic and nearshore stations peaked in mid- July at 0.06 and 0.03 larvae/m^ respectively (Fig. 1), when water temperature from the surface to a depth of 5 m was 21 C. This period coin- cides with the reported time of peak tui chub spawning in Pyramid Lake (Kucera 1978). By September few larvae were collected at the surface nearshore station, and none at the surface pelagic site. Digestive tracts from tui chubs taken on four pelagic and three nearshore sampling dates, totaling 25 and 14 larvae, respectively, were examined. Guts were usually one-half to three-quarters full and contained only zoo- plankton (Tables 1 and 2). Moina hutchinsoni was the dominant zoo- plankter recovered from all pelagic larvae sampled and was also of greatest significance to larvae in two of three nearshore collec- tions. Diaphanosoma leuchtenbergianum ranked second in importance among food items for pelagic larvae but was of minor im- portance among nearshore larvae. In con- trast, Cyclops vernalis appeared in all three nearshore samples but was insignificant in pelagic larval stomachs. Alona costata was observed in larvae from two nearshore sam- ples but was not recovered from pelagic lar- vae. Food items eaten in small amounts were Eucypris sp., Branchionus spp., Daphnia schodleri, and copepod nauplii. We anticipated finding more rotifers and copepod nauplii in larval tui chub stomachs, particularly since these groups were abun- dant in littoral zooplankton samples (Vucin- ich et al. 1981). Kimsey (1954) reported that January 1983 Galat, Vucinich: Food of Tui Chubs 177 Table 1. Pooled stomach contents of pelagic larval tui chubs from Pyramid Lake, Nevada. Carbon values are ap- proximate and represent reconstructed organisms. Numbers in parentheses are subtotals for the various categories. June 26 July 17 July 31 August 15 Total C Percent Total C Percent Total C Percent Total C Percent Food item No. (Mg) Total C No. (Mg) Total C No. (Mg) Total C No. (Mg) Total C Cladocera (8) (22) (98.7) (32) (88) (99.7) (55) (150) (98.4) (50) (139) (96.8) Moina 8 22 98.7 24 67 75.7 34 95 62.5 45 126 87.7 Diapharwsoma 7 18 20.5 21 55 35.9 5 13 9.1 Alona Daphnia 1 2.8 3.2 Copepoda (2) (1.0) (0.7) (4) (4.5) (3.2) Cyclops 1 3.0 2.1 Nauplii 2 1.0 0.7 3 5 1.1 Rotatoria Brachionus 1 0.3 1.3 2 0.5 0.6 5 1.4 0.9 Ostracoda Eucypris Grand total 9 22 34 88 62 152 54 144 No. tracts examined 3 14 7 6 No. tracts with food 2 10 7 6 Mean fish length (mm) 10.2 10.2 10.9 11.9 newly hatched tui chubs in Eagle Lake, Cali- fornia, fed on rotifers, diatoms, desmids, and other microscopic material. Perhaps tui chub larvae smaller than those captured in the present study fed on these organisms. Digestive tracts from larvae captured at the surface nearshore station contained more C. vernalis, A. costata, and Eucypris sp. than were recovered from larvae collected at the surface pelagic station. Conversely, D. leuch- tenbergianum was more abundant in guts from pelagic tui chubs. A probable explana- tion for these differences is that the first three zooplankton taxa named prefer a ben- thic habitat and hence would be more avail- able than D. leuchtenbergianum, a limnetic species, to nearshore tui chubs (Pennak 1978). Shifts in larval tui chub diet composition Table 2. Pooled stomach contents of near-shore larval tui chubs from Pyramid Lake, Nevada. Carbon values are approximate and represent reconstructed organisms. Numbers in parentheses are subtotals for the various categories. June 26 July 17 July 31 Total C Percent Total C Percent Total C Percent Food item No. ^g) Total C No. (Mg) Total C No. H) Total C Cladocera (7) (18) (33.2) (66) (183) (80.2) (31) (87) (96.7) Moina 2 5.6 10.9 60 168 73.6 31 87 96.7 Diaphanosoma 4 10 4.6 Alona 5 12 22.3 2 4.6 2.0 Daphnia Copepoda (23) (35) (66.8) (8) (19) (8.3) (1) (3.0) (3.3) Cyclops 23 35 66.8 6 18 7.9 1 3.0 3.3 Nauplii 2 1.0 0.4 Rotatoria Brachionus 21 5.7 2.5 Ostracoda Eucypris 3 21 9.0 Grand total 30 53 98 228 32 90 No. tracts examined 3 10 2 No. tracts with food 2 10 2 Mean fish length (mm) 10.3 12.8 12.8 178 Great Basin Naturalist Vol. 43, No. 1 from C. vemalis to D. leuchtenbergianum and M . hutchinsoni as the summer progressed paralleled seasonal changes in the relative abundance of these zooplankton species in Pyramid Lake (Galat et al. 1981), suggesting that larval tui chubs, like adults (Langdon 1979), are opportunistic feeders. ACKNOW^LEDGMENTS We thank the Pyramid Lake Paiute Indian Tribe for granting us permission to conduct this study. We also thank E. P. Bergersen, K. Hamilton-Galat, E. P. Eschmeyer, and D. Snyder for their critical review of the manu- script. Mark Coleman assisted with field col- lections and zooplankton identification. Study funding was provided by the Bureau of Indian Affairs under Contract 14-16-0008- 974 to the U.S. Fish and Wildlife Service and Colorado State University. Literature Cited Cooper, J. J. 1978. Contributions to the life history of the Lahontan tui chub, Gila hicolor obesa (Gi- rard), in Walker Lake, Nevada. Unpublished thesis. Univ. of Nevada, Reno. 98 pp. Galat, D. L., E. L. Lider, S. Vigg, and S. R. Robertson. 1981. Limnology of a large, deep. North American terminal lake, Pyramid Lake, Nevada, USA. Hydrobiologia 82:281-317. KiMSEV, J. B. 1954. The life history of the tui chub, Siph- ateles bicolor (Girard), from Eagle Lake, Califor- nia. California Fish Game 40:395-410. Kucera, p. a. 1978. Reproductive biology of the tui chub, Gila bicolor, in Pyramid Lake, Nevada. Great Basin Nat. 38:203-207. Langdon, R. W. 1979. Food habits of the tui chub, Gila bicolor, in Pyramid Lake, Nevada. Unpublished thesis. Humboldt State Univ., Areata, California. 45 pp. La Rivers, I. 1962. Fishes and fisheries of Nevada. Ne- vada State Fish and Game Comm. 782 pp. Miller, R. G. 1951. The natural history of Lake Tahoe fishes. Unpublished dissertation. Stanford Univer- sity. 160 pp. MoYLE, P. B. 1976. Inland fishes of California. Univ. of California Press, Berkeley. 405 pp. Pennak, R. W. 1978. Freshwater invertebrates of the United States. Wiley-lnterscience, New York. 803 pp. Sharp, J. H. 1974. Improved analysis for "particulate" organic carbon and nitrogen from seawater. Lim- nol. Oceanogr. 19:984-989. Snyder, D. E. 1976. Terminologies for intervals of larval fish development. Pages 41-60 in J. Boreman, ed.. Great Lakes fish egg and larval identi- fication: proceedings of a workshop. National Power Plant Team, U.S. Fish and Wildl. Serv. FWS/OBS-76-23. Snyder, J. O. 1917. The fishes of the Lahontan system of Nevada and northwestern California. U.S. Bur. Fish. Bull. 35(1915- 16):33-86. Vice, S. 1978. Vertical distribution of adult fish in Pyra- mid Lake, Nevada. Great Basin Nat. 38:417-428. 1981. Species composition and relative abun- dance of adult fish in Pyramid Lake. Great Basin Nat. 41:395-408. VuciNiCH, N., D. L. Galat, and M. C. Coleman. 1981. Size-class structure and food habits of young-of- year tui chubs (Gila bicolor, Girard), in Pyramid Lake, Nevada. Cooperative Fishery Research Unit, Colorado State University, Fort Collins. 64 pp. Williams, J. E., and C. D. Williams. 1980. Feeding ecology of Gila boraxobius (Osteichthyes: Cyprin- idae) endemic to a thermal lake in southeastern Oregon. Great Basin Nat. 40:101-114. NOTICE TO CONTRIBUTORS Manuscripts intended for publication in the Great Basin Naturalist or Great Basin Natural- ist Memoirs must meet the criteria outlined in paragraph one on the inside front cover. They should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Natu- ralist, 290 Life Science Museum, Provo, Utah 84602. Three copies of the manuscript are re- quired. 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 Council of Biology Editors Style Manual, Fourth Edition (AIBS 1978) 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 article. All manuscripts receive a critical peer review by specialists in the subject area of the manu- script under consideration. 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TABLE OF CONTENTS Life history of the Lahontan cutthroat trout, Salmo clarki henshawi, in Pyramid Lake, Nevada. William F. Sigler, WilHam T. Helm, Paul A. Kucera, Steven Vigg, and Gar W. Workman 1 A review of the genus Soliperla (Plecoptera: Peltoperlidae). Bill P. Stark 30 A bibhography of Colorado vegetation description. William L. Baker 45 Evaluation of a program to control hydatid disease in central Utah. Ferron L. Ander- sen, John R. Crellin, Craig R. Nichols, and Peter M. Schantz 65 Influence of cryptogamic crusts on moisture relationships of soils in Navajo National Monument, Arizona. Jack D. Brotherson and Samuel R. Rushforth 73 A vascular flora of the San Rafael Swell, Utah. James G. Harris 79 Pronghom responses to hunting coyotes. Timothy D. Reynolds 88 Floristics of the upper Walker River, California and Nevada.. Matt Lavin 93 Agropyron arizonicum (Gramineae: Triticeae) and a natural hybrid from Arizona. Grant L. Pyrah 131 Species composition, distribution, and phytosociology of Kalsow Prairie, a mesic tall- grass prairie in Iowa. Jack D. Brotherson 137 Deer mouse, Peromyscus maniculatus, and associated rodent fleas (Siphonaptera) in the arctic-alpine life zone of Rocky Mountain National Park, Colorado. R. B. Eads and E. G. Campos 168 Food of larval Tui chubs, Gila bicolor, in Pyramid Lake, Nevada. David L. Galat and Nancy Vucinich 1'^ i HE GREAT BASIN NATURALIST ume 43 No. 2 April 30, 1983 Brigham Young University BRARY 1P« GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Youing University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Chairman, Botany; James R. Barnes, Zoology; Hal L. Black, Zoology; Stanley L. Welsh, Botany; Clayton M. White, Zoology. All are at Brig- ham Young University, Provo, Utah 84602. Ex Officio Editorial Board Members. Bruce N. Smith, Dean, College of Biological and Agricul- tural Sciences; Norman A. Darais, University Editor, University Publications. Subject Area Associate Editors. Dr. Noel H. Holmgren, New York Botanical Garden, Bronx, New York 10458 (Plant Taxonomy). Dr. James A. MacMahon, Utah State University, Department of Biology, UMC 53, Lo- gan, Utah 84322 (Vertebrate Zoology). Dr. G. Wayne Minshall, Department of Biology, Idaho State University, Pocatello, Idaho 83201 (Aquatic Biology). Dr. Ned K. Johnson, Museum of Vertebrate Zoology and Department of Zoology, Uni- versity of California, Berkeley, California 94720 (Ornithology). Dr. E. Philip Pister, Associate Fishery Biologist, California Department of Fish and Game, 407 West Line Street, Bishop, California 93514 (Fish Biology). Dr. Wayne N. Mathis, Chairman, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 (Entomology). Dr. Theodore W. Weaver III, Department of Botany, Montana State University, Boze- man, Montana 59715 (Plant Ecology). The Great Basin Naturalist was founded in 1939 and has been published from one to four times a year since then by Brigham Young University. Previously unpublished manuscripts in Enghsh of less than 100 printed pages in length and pertaining to the biological natural his- tory of western North America are accepted. Western North America is considered to be west of the Mississippi River from Alaska to Panama. The Great Basin Naturalist Memoirs was es- tablished in 1976 for scholarly works in biological natural history longer than can be accom- modated in the parent publication. The Memoirs appears irregularly and bears no geographi- cal restriction in subject matter. Manuscripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist for private individuals is $16.00; for institutions, $24.00 (outside the United States, $18.00 and $26.00); and for stu- dent subscriptions, $10.00. The price of single issues is $6.00 each. All back issues are in print and are available for sale. All matters pertaining to subscriptions, back issues, or other busi- ness should be directed to Brigham Young University, Great Basin Naturalist, 290 Life Sci- ence Museum, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same office 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. See Notice to Contributors on the inside back cover. 6-83 65066305 ISSN 017-3614 Hie Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 43 April 30, 1983 No. 2 UTAH FLORA: COMPOSITAE (ASTERACEAE) Stanley L. Welsh' Abstract.— A revision of the sunflower family, Compositae (Asteraceae), is presented for the state of Utah. In- cluded are 613 taxa in 100 genera. Keys to genera, species, and infraspecific taxa are provided, along with detailed descriptions, distributional data, and pertinent discussion. Many nomenclatural changes have been necessary to bring Utah composites into line with contemporary interpretations of the family. They include: Artemisia tridentata Nutt. var. wyomingensis (Beetle & Young) Welsh; Aster kingii D.C. Eaton var. barnebyana (Welsh & Goodrich) Welsh; Brickellia microphylla (Nutt.) Gray var. ivatsonii (Robins.) Welsh; Chrysothamntis nauseosiis (Pallas) Britt. — var. abbreviatiis (Jones) Welsh, var. arenarius (L.C. Anderson) Welsh, var. glareosus (Jones) Welsh, var. iridis (L.C. Anderson) Welsh, and var. nitidus (L.C. Anderson) Welsh; Cirsium arizonicum (Gray) Petrak var. nidulum (Jones) Welsh; Cirsium calcareum (Jones) Woot. & Standi. — var. bipinnatum (Eastw.) Welsh and var. pulchellum (Greene) Welsh; Cirsium neomexicanum Gray var. utahense (Petrak) Welsh; Cirsium undulatum (Nutt.) Spreng. var. tracyi (Rydb.) Welsh; Crepis runcinata (James) T. & G. var. ghiuca (Nutt.) Welsh; Erigeron speciosus (Lindl.) DC. — var. mollis (Gray) Welsh and var. uintahensis (Cronq.) Welsh; Gutierrezia petradoria (Welsh & Goodrich) Welsh; Gutier- rezia pomariensis (Welsh) Welsh; Haplopappiis racemosus (Nutt.) Torr. — var. sessiliflortts (Greene) Welsh and var. prionophyllus (Greene) Welsh; Haplopappus watsonii Gray var. rydbergii (Blake) Welsh; Lygodesmia grandiflora (Nutt.) T. & G. — var. arizonica (Tomb) Welsh and var. dianthopsis (D.C. Eaton) Welsh; Machaeranthera canescens (Pursh) Gray — var. commixta (Greene) Welsh, var. leucanthemifolia (Greene) Welsh, and var. vacans (A. Nels.) Welsh; Petradoria pumila (Nutt.) Greene var. graminea (Woot. & Standi.) Welsh; and Senecio spartioides T. & G. var. multicapitatus (Greenm. in Rydb.) Welsh. This paper is one of a series of works lead- ornamentals. Few actual crop or food plants ing to a definitive treatment of the flora of are derived from members of this family. Utah. Previous papers have dealt with the Only lettuce and sunflower are grown as Brassicaceae, Fabaceae, Rosaceae, and mis- crops or as garden plants from the vast array cellaneous smaller families. of species in this great family. Despite the The sunflower family has long been recog- paucity of food plants, there are many oma- nized for its great size and complexity both mental species. These have been included in in Utah and elsewhere; it is probably the the present treatment only when they have largest flowering plant family on earth. In escaped, or when they have been planted Utah it consists of 100 genera and 613 taxa, routinely for many years, and when speci- of which some 40, or about 7 percent, are in- mens have been preserved in the herbarium, troduced. The 573 indigenous taxa comprise The cultivated flora requires a separate in- about 20 percent of the flora native to the tensive effort not herein attempted, state. This large family is apparently unique The importance of members of this family in total numbers, but is also unique in having to wildlife, both as cover and as food, is well such a small proportion of adventive taxa. known by range managers. Despite the pres- The figures are misleading, to an extent, be- ence of chemical substances produced by the cause of the omission of numerous cultivated plants, which impart unpleasant flavors or 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 179 180 Great Basin Naturalist Vol. 43, No. 2 even poisonous substances, many of them are eaten by both wildlife and by livestock. Sage- brush and rabbitbrush species are well known for their value in reclamation of sites requir- ing rehabilitation. Negative values are reflected in the weedy species that occupy cultivated lands, in the mechanically injurious species (such as thistles and their relatives), and in the numer- ous poisonous kinds. Livestock losses have been reported in literature from utilization of species of the family, i.e., Baileya, Hyme- noxys (especially richardsonii), Oxytenia ace- rosa, Psilostrophe, Senecio, and Tetradymia. Other plants are rendered unpalatable by their complex biochemical compounds, and they tend to increase on range lands where other more palatable plants are eaten selec- tively. Gutierrezia and Chrysothamnus spe- cies fit this latter category, although selected phases of the same Chrysothamnus species might be palatable or differentially palatable. Certainly there is much room for in- vestigation of members of this huge assemblage. The largest genus, and one of the most complex taxonomically, is Erigeron, with more than 60 taxa. Without the able mono- graph by Cronquist, and lacking his coopera- tive help, the species of Erigeron would have been much more difficult to interpret. Be- cause of his knowledge, his dedication, and his helpful cooperation, this treatment is ded- icated to him. The same kinds of problems, made even more complicated by extensive hybridization, is true for Senecio, which Barkley has treated so competently. Fortu- nately, monographs or revisions are available for most of the larger genera. However, no modem work is definitive for Aster, which requires interpretation on a cosmopolitan, rather than provincial, basis. Also, Artemisia has not been treated in its entirety for several decades. Much research on the nature of the biochemical constituents of Artemisia has shed light on the taxonomy of the group. Difficulties in the composite genera and species, aside from those involving the great number of taxa, include those due to hybridi- zation, ploidy level, and apomixis. Blending of morphological forms due to hybridization, subtle changes due to variations in chromo- some sets, and the problems of interpretation of apomictic races each lead to problems not easily soluble. And the end results are subject to a variety of preliminary conclusions, each subject to change as additional information becomes available. Further, generic lines in the family are not definitive, with aggrega- tions of species sometimes representing con- venience rather than actual affinities. Recent workers have tended to segregate traditional genera into finer subunits or additional gen- era. This seems to represent a continuing trend, and it seems probable that some of the genera treated herein will be broken up in the future. The weight of evidence for doing so lies with the future workers. Those who find it modern to give "new" generic inter- pretations are often merely opting previous workers, whose interpretations were flawed. Another later generation will opt for a differ- ent set of names based on what they consider to be "modern." Some of the species are edaphic specialists, occurring on definite substrates of restricted aerial or elevational distribution. The woody asters (Xylorhiza) are selenophytes and are re- stricted to shales and silt- and mudstones of fine-textured geological formations in eastern to southwestern Utah. The present work should be considered tentative at best. Despite the large number of specimens available for study (13,700), many of the taxa are imperfectly known and distri- butions are yet to be understood. Mono- graphic work is required for practically all groups, whether completed in the recent past or not. It is hoped that this work will provide a useful summary for those who work with Utah composites. To the extent possible the work presented below contains the most modern names for the entities involved, based on application of contemporary International Rules of Bo- tanical Nomenclature. Following the name of the entities are the synonyms that apply to Utah plants specifically. No attempt has been made to list synonymy exhaustively. The basionym has been included more or less uni- formly, and an attempt has been made to cite all synonyms based on Utah plants, including a brief notation of type locality. The discussion following the description of each species, or the name of the infraspecific taxon, includes the plant communities in April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 181 which the taxon occurs. They are arranged in ascending order from dry low elevation com- munities to mesic high elevation commu- nities. Community data is followed by eleva- tional range (given in meters), counties of known occurrence (in alphabetical order), and the distribution outside Utah. At the end of the discussion are two numbers, the first, in Arabic niunerals, indicates the number of Utah specimens examined for the taxon and the second, in Roman numerals, is the num- ber collected in Utah by the author. The numbers are given to provide the reader with the basis of understanding of the taxon by the writer and his familiarity with the plants in the field. The plants are not equally well known by this writer, and the user should be able to make judgements when the taxonomy presented herein does not adequately ap- proach the conditions as noted in the field. The approach to reality is always an approx- imation, and much improvement will take place in the future, as more information is derived from specimens not now available for the present study. COMPOSITAE (ASTERACEAE) Sunflower Family Annual, biennial, or perennial herbs, or shrubs; leaves alternate, opposite, or whorled, simple, pinnatifid, or compound; in- florescence of involucrate heads, these soli- tary or several in corymbose, racemose, pan- iculate, or cymose clusters; flowers few to numerous on a common receptacle, sur- rounded by green bracts forming a cup- shaped, cylindrical, or urn-shaped involucre enclosing the flowers in bud; heads entirely of tubular (disk) corollas, entirely of ligulate (ray) corollas, or with tubular corollas form- ing a central disk and an outer radiating row of ligulate corollas; receptacle flat, convex, conic, or cylindric, naked or bearing chaffy bracts, scales, or hairs; calyx lacking, or crowning the summit of the ovary and modi- fied as a pappus of capillary bristles, scales, or awns; stamens alternate with corolla lobes; filaments free (rarely connate); the anthers united and forming a tube (rarely separate); ovary inferior, of 2 carpels, 1-loculed and with a single ovule; styles 1, 2-cleft, exserted through the anther tube; fruit an achene. Note: All involucral measurements are from dried pressed herbarium specimens. The width measurements are sometimes broader than in fresh material. Arnow, L., B. Albee, and A. Wyckoff. 1980. Flora of the central Wasatch Front, Utah. Univ. of Utah Printing Service, Salt Lake City. 663 pp. Meyer, S. E. 1976. Annotated checklist of the vascular plants of Washington County, Utah. Unpublished thesis, Univ. of Nevada, Las Vegas. 276 pp. 1. Corollas all raylike; plants usually with milky juice KEY 1 — Corollas not all raylike, some or all of them tubular; juice seldom if ever milky 2 2(1). Corollas all tubular; no ray flowers present, or the rays vestigial and minute KEY II — Corollas not all tubular; ray flowers present 3 3(2). Pappus of capillary bristles, at least in part KEY III — Pappus of awns or scales, or lacking 4 4(3). Pappus lacking KEY IV — Pappus present, of awns or scales KEY V KEY I. Corollas all ray like; plants usually with milky juice. Pappus lacking 2 Pappus present 3 182 Great Basin Naturalist Vol. 43, No. 2 2(1). Rays 10-20 mm long; plants glabrous, with leaves in basal rosette Atrichoseris — Rays 5-7 mm long; plants pubescent, with well-developed cauline leaves .... Lapsana 3(1). Pappus, at least in part, of plumose bristles 4 — Pappus of simple bristles, of awns, or of scales 7 4(3). Plants acaulescent, with merely bracteate stems Hypochaeris — Plants caulescent 5 5(3). Achenes not beaked, truncate at apex; involucres usually less than 15 mm long Stephanomeria — Achenes tapering or beaked at apex; involucres usually more than 15 mm long 6 6(5). Leaves pinnatifid; corollas white or pinkish; involucre with an outer series of short bractlets; southern Utah Rafinesquia — Leaves not pinnatifid, entire; corollas yellow or purplish; involucre lacking short outer bractlets; widespread Tragopogon 7(3). Pappus of 1-3 series of unawned or awned scales 8 — Pappus of capillary bristles 9 8(7). Pappus of 2 or 3 series of unawned scales; corollas blue, closing by mid- morning Cichorium — Pappus scales in a single series, awned; corollas yellow, not closing by mid- morning Microseris 9(7). Achenes more or less flattened; stems leafy; heads in panicles or in umbellate clusters 10 — Achenes not flattened; stems leafy or scapose; heads solitary or variously dis- posed 11 10(9). Involucres cylindric or ovoid-cylindric; achenes beaked; flowers yellow or blue Lactuca — Involucres broadly campanulate to hemispheric; achenes not beaked; flowers yellow Sonchus 11(9). Corollas pink or purplish 12 — Corollas yellow or yellowish, or white or cream colored 14 12(11). Plants annual; heads mainly 5-7 mm long (from base of involucre to tip of pappus) Prenanthella — Plants perennial; heads mainly 8-20 mm long or more 13 13(12). Plants with rigid spine-tipped branches Stephanomeria — Plants unarmed, the branches soft Lygodesmia 14(11). Leaves all basal; heads solitary on scapose peduncles 15 — Leaves not all basal, the stems leafy; heads not on scapose peduncles 17 15(14). Achenes not beaked, truncate; pappus bristles barbellate Microseris — Achenes beaked or tapering to apex; pappus not of barbellate bristles 16 16(15). Achenes 10-ribbed or 10-nerved, not spinulose; involucral bracts usually imbri- cated in several series Agoseris — Achenes 4- to 5-ribbed, spinulose, especially near apex; principal bracts in a single series, the outer much shorter Taraxacum April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 183 17(14). Achenes ridged or tuberculate between the angles; leaves either crustaceous margined or peduncles stipitate-glandular; southwestern Utah 18 — Achenes striate between the angles; leaves and peduncles otherwise (rarely glandular setose in some Crepis species); widely distributed 19 18(17). Plants depressed annuals with crustaceous-margined leaves, not stipitate- glandular; achenes abruptly beaked, transversely ridged between the ribs Glyptopleura — Plants erect, lacking crustaceous-margined leaves, conspicuously stipitate- glandular above; achenes tapering to a beak, not transversely ridged Calycoseris 19(17). Pappus bristles early deciduous, more or less united below and falling together, only a few of the stout outer ones may be persistent Malacothrix — Pappus bristles persistent or tardily deciduous, and then falling separately 20 20(19). Pappus tan to brown; involucral bracts not thickened Hieracium — Pappus white or whitish; involucral bracts somewhat thickened at base or on midrib Crepis KEY II. Corollas all tubular; no ray flowers present. 1. Heads unisexual, the pistillate heads with 1-4 flowers enclosed in involucre; in- volucre burlike or nutlike, only style tips exserted 2 — Heads perfect or unisexual; involucre not burlike or nutlike 4 2(1). Involucral bracts of the staminate heads separate; fruiting involucres burlike, covered with hooked appendages Xanthium — Involucral bracts of the staminate heads united; fruiting involucres various but, if burlike, lacking hooked appendages 3 3(2). Shrubs; fruiting involucre with several transverse, scarious wings; leaves or their lobes linear-filiform Hymenoclea — Shrubs or herbs; fruiting involucre lacking transverse wings; leaves and their lobes not linear-filiform Ambrosia 4(1). Stamens not united by their anthers; flowers always unisexual, the pistillate co- rollas none or much reduced 5 — Stamens with united anthers or rarely not united in some species with perfect flowers, at least some flowers usually perfect 7 5(4). Achenes long-villous; leaves or their lobes linear-filiform Oxytenia — Achenes not long-villous; leaves or their lobes not linear-filiform 6 6(5). Pistillate flowers subtended by large, chaffy scales simulating inner involucral bracts; achenes with pectinate or winged margins Dicoria — Pistillate flowers subtended by chaffy scales or these lacking; achenes without pectinate or toothed wings Iva 7(4). Involucral bracts with translucent, usually yellow or orange dots Porophyllum — Involucral bracts without distinct dots; pappus various, but not as above 8 8(7). Pappus of capillary bristles, at least in part, these smooth, scabrous, barbellate, or plumose 9 — Pappus lacking or, if present, not of capillary bristles 41 184 Great Basin Naturalist Vol. 43, No. 2 9(8). Leaves opposite or whorled, some or all cauline 10 — Leaves alternate, at least basally, or basal and actually alternate 13 10(9). Corollas yellow; involucral bracts in 1 series or in 2 series, but all equal in length Arnica — Corollas white, ochroleucous, flesh colored, blue, or purple; involucral bracts in 2 to several series 11 11(10). Pappus double— the outer series of short scales, the inner series of capillary bristles; shrubs with white bark Hofmeistera — Pappus single, or else plants herbaceous; shrubs or herbs 12 12(11). Achenes 5-angled or 5-ribbed; involucral bracts subequal or in 2 series ... Eupatorium — Achenes 10-angled or 10-ribbed; involucral bracts imbricated in several series of different lengths Brickellia 13(9). Leaves spinescent, usually with spiny teeth or lobes, rarely entire but then with spine-tipped apex, thistlelike 14 — Leaves entire, denticulate or lobed, lacking spines, not thistlelike 18 14(13). Corollas of some or all flowers bilabiate; basal leaf axils woolly; leaves spin- ulose-dentate; flowers pink; arid sites in Kane and Washington counties Perezia — Corollas not bilabiate; leaves not or seldom spinulose-dentate; basal leaf axils woolly; flowers pink-white or cream; various distribution 15 15(14). Pappus of 2 series of awns, the outer long and naked, the inner short and hispi- dulous; flowers yellow Cniciis — Pappus of plumose or barbellate capillary bristles; flowers not yellow 16 16(15). Pappus bristles plumose (rarely some otherwise); receptacle densely bristly .. Cirsium — Pappus bristles merely barbellate 17 17(16). Receptacle densely bristly, not fleshy or honeycombed; heads nodding Carduus — Receptacle not bristly or scarcely so, fleshy and honeycombed; heads not nod- ding Onopardum 18(13). Receptacle with dense bristles or narrow, chaffy scales between disk flowers 19 — Receptacle naked or at most short-hairy, never with dense bristles or scales 21 19(18). Involucral bracts with hooked spines; lower leaves large (resembling rhubarb), cordate at base Arctium — Involucral bracts without spines, or spines not hooked; lower leaves not large and cordate at base 20 20(19). Receptacle chaffy except in center; plants small, woolly Filago — Receptacle chaffy throughout; plants not small and woolly Centaurea 21(20). Heads unisexual; plants dioecious (staminate flowers may have styles but ovary does not develop) 22 — Heads with at least central flowers perfect 24 22(21). Plants shrubs or else woody at base, not tomentose; leaves sometimes toothed or lobed; involucral bracts not strongly scarious margined Baccharis — Plants herbaceous, more or less tomentose; leaves entire; involucral bracts strongly scarious, at least along margins 23 23(22). Pappus bristles of pistillate flowers united at base and falling together; pappus bristles of staminate flowers usually club shaped at apex; plants usually less than 30 cm tall; basal leaves commonly in a rosette; cauline leaves reduced and different in shape; leaves usually tomentose on both sides Antennaria April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 185 — Pappus of pistillate flowers separate at base and falling separately; pappus bristles of staminate flowers not club shaped at apex; plants mostly over 30 cm tall; leaves all alike, usually green and glabrate above Anaphalis 24(21). Stems longitudinally brown-striate; involucral bracts imbricate, chartaceous, the inner with scarious margins and broadly rounded apices; shrubs with yel- low flowers, of western Millard County Lepidospartum — Stems striate or not; involucral bracts scarious, hyaline, or herbaceous but not as above; herbs, or shrubs with flowers and distribution various 25 25(24). Involucral bracts scarious or hyaline (only partly so in Pluchea) 26 — Involucral bracts herbaceous, at least in the center 28 26(25). Involucral bracts subscarious; corollas purplish; plants not tomentose, slender woody shrubs Pluchea — Involucral bracts scarious; corollas rarely purplish; plants tomentose, prostrate to erect herbs 27 27(26). Plants perennial, subdioecious pistillate heads usually with a few central, per- fect flowers Anaphalis — Plants annual or perennial, not dioecious; heads all alike, the marginal flowers pistillate and central ones perfect Gnaphalium 28(25). Involucral bracts in a single series, a few very short ones may be present at the very base 29 — Involucral bracts of 2 or more series, these often of different lengths 32 29(28). Plants woody, shrubs; involucral bracts 4-6 per head Tedradymia — Plants herbaceous; bracts more than 6 per head 30 30(29). Plants annual; heads with inner flowers perfect, the outer pistillate Conyza — Plants perennial; heads with all flowers perfect 31 31(30). Style branches with a tuft of hairs near the truncate apex; involucral bracts in 1 series only (a few short bracts may be present) Senecio — Style branches without a tuft of hairs near the truncate apex; involucral bracts actually in 2 or more series Erigeron 32(28). Pappus double, the outer series of short scales, the inner ones of capillary bristles; shrubs with white bark Hofmeistera — Pappus simple or else the plants herbaceous 33 33(32). Plants annual 34 — Plants perennial 36 34(33). Plants low, depressed, scurfy pubescent herbs; leaves broadly ovate or round- ish, entire or toothed Psathyrotes — Plants not as above 35 35(34). Leaves all entire Aster — Leaves toothed or lobed, at least the lower Conyza 36(33). Involucral bracts in more or less distinct vertical rows Chrysothamnus — Involucral bracts not in vertical rows 37 37(36). Involucral bracts usually in 1 subequal series Erigeron — Involucral bracts imbricate, in 2 or more series 38 186 Great Basin Naturalist Vol. 43, No. 2 38(37). Involucral bracts not longitudinally striate; flowers commonly yellow .. Haplopappus — Involucral bracts longitudinally striate; flowers commonly cream to ofl-white, or pink to purplish 39 39(38). Flowers pink to purplish; plants of northwestern Utah Eupatorium — Flowers cream to white; plants of various distribution 40 40(39). Pappus plumose; plants perennial herbs Kuhnia — Pappus scabrous or hispidulose; plants shrubs or herbs Brickellia 41(9). Receptacle with bristles or chaffy scales among the flowers 42 — Receptacle naked or merely short-hairy 50 42(41). Receptacle densely bristly Centaurea — Receptacle with chaffy scales 43 43(42). Plants low woolly annuals; outer bracts boat shaped and enclosing the achenes 44 — Plants various, but not low and woolly; outer bracts various but not usually en- closing the achenes 45 44(43). Stem leaves opposite; style lateral Psilocarpus — Stem leaves alternate; style terminal Stylocline 45(43). Involucral bracts in 2 distinct sets — the outer herbaceous, the inner differing in shape and texture; leaves opposite, at least below, or alternate 46 — Involucral bracts not in 2 unlike sets; leaves alternate or basal 47 46(45). Leaves alternate throughout; outer involucral bracts about 5, spreading, her- baceous, the irmer (1-3 subtending pistillate flowers) larger and broader, be- coming strongly accrescent and hooded in fruit Dicoria — Leaves opposite, at least below; outer involucral bracts various, but not as above, not accrescent and hooded in fruit Thelesperma 47(45). Involucral bracts in 1 series, boat shaped, each bract enclosing a marginal flower; rays short, yellow Madia — Involucral bracts in 1 or more series, not boat shaped and enclosing marginal flowers; rays lacking 48 48(47). Plants woody shrubs; mostly along the canyons of the Colorado and Green riv- ers Encelia — Plants herbaceous; widely distributed 49 49(48). Receptacles high-conical, mostly over 3 cm long; stems leafy Rudbeckia — Receptacles merely convex, much less than 3 cm long; leaves all basal Enceliopsis 50(41). Pappus none 51 — Pappus present 54 51(50). Leaves opposite, some cauline, somewhat connate at base; plants of Grand, San Juan, and Tooele counties Flaveria — Leaves alternate or basal 52 52(51). Heads numerous, in spikes, racemes, or panicles; anthers with acute tips; re- ceptacles flat; plants woody or herbaceous Artemisia — Heads solitary on ends of stems, or sometimes corymbose or capitate; anthers with rounded tips; receptacles convex or conic; plants herbaceous, or woody only at base 53 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 187 53(52). Plants annual; heads solitary or paniculately arranged; leaves green and glabrous Chamomilla — Plants perennial; heads corymbose or capitate; leaves usually silvery-canescent Chrysanthemum 54(50). Plants dioecious shmbs Baccharis — Plants not dioecious herbs or shrubs 55 55(54). Pappus of 2-8 caducous awns; plants usually strongly glutinous Grindelia — Pappus various, but not of 2-8 caducous awns 56 56(55). Leaves and involucre conspicuously punctate with translucent oil glands .... Dyssodia — Leaves and involucre sometimes impressed-punctate, but without translucent oil glands 57 57(56). Pappus of 12 or more scale or bristlelike segments, these nearly or quite as long as achene 58 — Pappus of fewer than 12 scalelike segments or else much shorter than achene 59 58(57). Pappus of 12-16 linear, acuminate awns; involucres glutinous; leaves 3- to 5- nerved Vanclevea — Pappus of ca 35 flattened, silvery scales and bristles of different widths; in- volucres not glutinous; leaves 1-nerved Acamptopappus 59(57). Achenes strongly compressed; pappus of 1 or 2 slender awns Laphamia — Achenes not compressed or, if so, then pappus not of 1 or 2 slender awns 60 60(59). Pappus a crown with margins entire or of short scales united into a crown 61 — Pappus not as above 63 61(60). Plants annual; heads solitary or paniculately arranged; flowers all perfect; leaves green and glabrous Chamomilla — Plants perennial; heads corymbose or capitate, rarely solitary; some marginal flowers pistillate only; leaves mostly silvery-canescent 62 62(61). Plants 0.5-1 m tall; leaves doubly pinnately dissected, mainly 10-20 cm long ... Tanacetum — Plants mainly less than 0.3 m tall; leaves entire, once pinnately dissected, ter- nate, merely toothed apically, or entire, mainly less than 10 cm long ... Sphaeromeria 63(60). Involucral bracts with a thin, scarious, white, yellow, or purplish margin and tip Hymenopappus — Involucral bracts without a scarious, colored margin and tip 64 64(63). Plants scapose; leaves roundish, entire, or crenate Chamaechaenactis — Plants leafy stemmed; leaves not roundish and entire or subentire 65 65(64). Pappus scales with a strong midrib; leaves lanceolate or linear, entire; southern Utah Palafoxia — Pappus scales nerveless or essentially so; leaves, at least in part, toothed to pin- natifid; widely distributed Chaenactis KEY III. Corollas not all tubular; ray flowers present pappus of capillary bristles. I. Rays white, pink, violet, or purple, not yellow 2 — Rays yellow or orange-yellow 9 188 Great Basin Naturalist Vol. 43, No. 2 2(1). Pappus of numerous unequal bristles, alternating with shorter, lacerate scales; involucral bracts subequal; low winter annuals Monoptilon — Pappus of numerous bristles; involucral bracts imbricate or subequal; plants various, but seldom low winter annuals 3 3(2). Pappus, at least of disk flowers, of several to many rigid bristles; achenes pu- bescent with 2-forked hairs or the hairs barbed at apex Townsendia — Pappus, at least of disk flowers, of many capillary bristles, at least in part; achenes glabrous or pubescent with simple hairs 4 4(3). Rays very inconspicuous, shorter than the tube and scarcely if at all exceeding their pappus; central perfect flowers few; plants annual Conyza — Rays usually conspicuous, longer than the tube and pappus; central perfect flowers several to many; plants annual, biennial, or perennial 5 5(4). Involucres subequal, rarely somewhat graduated; rays usually narrow; style tips very short, triangular, rounded, or obtuse Erigeron — Involucres usually strongly graduated; rays comparatively broad; style tips ovate and acute to subulate, usually lanceolate 6 6(5). Plants perennial, rhizomatous, or annual, or, if from a caudex, ordinarily less than 10 cm tall (see also Aster kingii) 7 — Plants from a caudex or taproot 8 7(6). Low, white-rayed perennial herbs from spreading cordlike rootstocks, in arid sites; flowering in springtime Leucelene — Low to tall, white- to pink- or purple-rayed annual or perennial herbs from rhizomes or fibrous roots (a caudex in A. kingii); mainly flowering in summer and autumn Aster 8(6). Plants herbaceous, from a taproot, biennial or perennial; heads usually several to numerous Machaeranthera — Plants more or less woody, from a ligneous caudex; heads usually solitary and large (primary selenophytes) Xylorhiza 9(1). Leaves opposite, at least below 10 — Leaves alternate throughout 12 10(9). Plants subshrubs Laphamia — Plants herbaceous 11 11(10). Leaves with stiff marginal bristles; involucre and leaves with conspicuous oil glands; plants annual Pectis — Leaves without stiff marginal bristles; involucre and leaves without oil glands; plants perennial Arnica 12(11). Plants 1-1.5 m tall, herbaceous; heads 3-5 cm wide; rays 1-2 cm long Inula — Plants various, usually less than 1 m tall, or, if taller, woody; heads much small- er; rays seldom to 1 cm long 13 13(12). Pappus of 2-8 stiff, caducous bristles; plants usually glutinous Grindelia — Pappus of numerous, usually soft, persistent bristles 14 14(13). Pappus of about 20 twisted, flattish bristles Amphipappus — Pappus of numerous, straight, capillary bristles 15 15(16). Pappus double, the inner of numerous bristles, the outer sometimes scalelike 16 — Pappus not double, of subequal capillary bristles only 17 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 189 16(15). Leaves essentially filiform Conyza — Leaves not filiform, linear-oblong or broader Heterotheca 17(15). Involucral bracts in distinct vertical ranks 18 — Involucral bracts not in distinct vertical ranks 19 18(19). Outer involucral bracts with loose herbaceous tips; erect stems perennial; plants shrubs; leaves deciduous Chrysothamnus — Outer involucral bracts without loose herbaceous tips; erect stems annual; plants herbaceous; leaves persistent Petradoria 19(17). Involucral bracts in 1 series, frequently with some smaller bracts at base; style branches truncate apically Senecio — Involucral bracts neither in 1 series nor with smaller bracts at base; style branches without truncate tips 20 20(19). Heads small, the involucres usually less than 6 mm high, usually very numerous and densely paniculate, rarely racemose or corymbose; plants rhizomatous, fibrous rooted Solidago — Heads medium to large, the involucres usually more than 6 mm high, neither very numerous nor densely paniculate; plants with taproots, occasionally also rhizomatous Haplopappus Key IV. Corollas not all tubular; ray flowers present; pappus lacking. 1. Rays white, pink, or pink-purple, sometimes yellow at base 2 — Rays yellow, sometimes partly purplish or maroon 6 2(1). Receptacle naked 3 — Receptacle with chaffy scales 5 3(2). Leaves all basal; plants scapose Bellis — Leaves not all basal, at least some cauline; plants caulescent 4 4(3). Receptacle broad and flattish; involucral bracts with a dark brown submarginal line Chrysanthemum — Receptacle convex, conic, or hemispheric; involucral bracts without a dark brown submarginal line Chamomilla 5(3). Heads small, numerous, in dense, flattish or rounded cymose panicles; plants perennial Achillea — Heads comparatively large, solitary or few; plants annual or perennial Anthemis 6(1). Receptacles not chaffy 7 — Receptacles chaffy, at least toward the margin 12 7(6). Heads 1- or 2-flowered, in dense glomerate clusters, sessile in the forks of the stem, or terminal and leafy involucrate Flaveria — Heads several- to many-flowered, solitary on terminal peduncles 8 8(7). Plants woolly 9 — Plants not woolly 10 9(8). Rays persistent, becoming papery Baileya — Rays not persistent Eriophyllum 190 Great Basin Naturalist Vol. 43, No. 2 10(8). Involucre and leaves with translucent oil glands Pedis — Involucre and leaves without translucent oil glands 11 11(10). Rays conspicuous; involucral bracts acuminate, without scarious margins Bahia — Rays minute; involucral bracts obtuse, with scarious margins Tanacetum 12(6). Ray achenes partly or wholly enfolded by their involucral bracts; plants an- nual, glandular-viscid above Madia — Ray achenes not conspicuously enfolded by their involucral bracts or, if so, then plants perennial; plants perennial or, if annual, not glandular above 13 13(12). Involucre distinctly double, the outer bracts herbaceous, the inner ones broad- er and united to about the middle Thelesperma — Involucre not double, the bracts distinct to the base 14 14(13). Plants scapose perennials; leaves broad, silvery-pubescent, entire; heads very broad Enceliopsis — Plants leafy stemmed or subscapose; leaves various but not broad and silvery- pubescent, or if so, then sagittate; heads broad or narrow 15 15(14). Plants subscapose; leaves variously dissected or sagittate; heads broad .. Balsamorhiza — Plants with stems definitely leafy; leaves usually not dissected or sagittate 16 16(15). Plants shrubby; achenes conspicuously ciliate on the margins, notched at the apex, very flat Encelia — Plants herbaceous; achenes not conspicuously ciliate on the margins 17 17(16). Leaves doubly pinnately dissected; heads numerous in corymbose cymes Achillea — Leaves simple, entire or toothed to lobed; heads few to several 18 18(17). Achenes 2-winged; disks 15-25 mm wide; leaves white-strigose beneath, green above Verbesina — Achenes not 2-winged; disks 6-15 mm wide; leaves green on both sides .... Heliomeris Key V. Corollas not all tubular; ray flowers present; pappus of awns or scales. 1. Receptacle chaffy 2 — Receptacle not chaffy, either naked or bristly 17 2(1). Pappus scales fimbriate; ray flowers 4 or 5, white, only slightly surpassing the disk; introduced weedy plants, to be expected in Utah Galinsoga parviflora Cav. — Pappus scales or awns not fimbriate; ray flowers various in size and color; in- digenous or introduced 3 3(2). Receptacle bearing a row of chaffy scales between the ray flowers and the out- er disk flowers, otherwise naked; pappus of 10-20 slender setiform scales Layia — Receptacle chaffy throughout; pappus not of 10-20 slender scales 4 4(3). Ray achenes dorsiventrally compressed, the thickened margins attached to a contiguous pair of infertile disk flowers and the subtending bract, and falling as a unit; pulvinate herbs of eastern Utah and shrubs of southwestern Utah Parthenium — Ray achenes various, but not as above; herbs or shrubs 5 5(4). Pappus of awns only, without scales 6 — Pappus, at least in part, of scales 10 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 191 6(5). Achenes flat and obcompressed; awns retrorsely hispid Bidens — Achenes not obcompressed; awns not retrorsely hispid 7 7(6). Achenes plump; pappus of 2 to several caducous awns Helianthus — Achenes flat, very strongly compressed; pappus various 8 8(7). Plants scapose; heads large, solitary Enceliopsis — Plants leafy stemmed; heads medium sized, usually several 9 9(8). Plants shrubby; achenes narrowly white margined, the margin not continuous between weak awns Encelia — Plants herbaceous annuals; achenes strongly white margined, the margin con- tinuous between stout awns Geraea 10(5). Achenes very flat, strongly compressed 11 — Achenes not very flat, usually much thickened 13 11(10). Leaves once to twice pinnatifid Anthemis — Leaves not pinnatifid, entire or nearly so 12 12(11). Plants scapose Enceliopsis — Plants leafy stemmed Helianthella 13(10). Pappus caducous (of 2 awns and rarely some scales) Helianthus — Pappus persistent 14 14(13). Inner involucral bracts united to middle into a cup Thelesperma — Inner involucral bracts not united into a cup 15 15(14). Receptacle merely convex; rays pistillate Wyethia — Receptacle conic or cylindric; rays neuter 16 16(15). Involucral bracts subequal, in 2 or 3 series Rudbeckia — Involucral bracts unequal, in 2 series, the inner ones shorter Ratibida 17(1). Rays white or purple 18 — Rays yellow, sometimes marked with purple 24 18(17). Pappus a short crown 19 — Pappus of awns or scales 21 19(18). Leaves entire or pinnately divided Chrysanthemum — Leaves irregularly 2-3 times pinnately dissected 20 20(19). Plants annual; heads 1-2.5 cm wide; receptacle conic, hollow Chamomilh — Plants biennial or perennial; heads 3-5 cm wide; receptacle hemispheric . Matricaria 21(18). Pappus of 1 plumose awn and a denticulate crown Monoptilon — Pappus of 2 to several awns or scales 22 22(21). Plants dwarf woolly annuals Eriophyllum — Plants armual or perennial, not woolly 23 23(22). Pappus of numerous awns or scales; involucral bracts conspicuously scarious- margined Touinsendia — Pappus of 4 or 5 stiff awns; involucral bracts obscurely scarious-margined Rigiopappus 24(17). Receptacle densely bristly or hairy 25 — Receptacle naked 26 192 Great Basin Naturalist Vol. 43, No. 2 Heads very small; involucres less than 10 mm wide Gutierrezia Heads medium sized; involucres more than 10 mm wide Gaillardia Pappus of 4 hyaline scales united at the base; rays reddish purple to yellow ... Hulsea Pappus a crown, or of cauducous or persistent awns or scales; rays mostly yel- low 27 Pappus a mere crown or of caducous awns 28 Pappus persistent, of awns or scales 30 Pappus of 2-8 caducous awns; plants glutinous Grindelia Pappus a short crown; plants seldom if ever glutinous 29 Leaves entire, bristly margined basally Pedis Leaves 2- or 3-pinnate Tanacetum Pappus of 1 or 2 awns or scales (rarely more) with or without a crown Perityle Pappus of 4 to many awns or scales 31 Pappus of about 20 slender, twisted awns; rays 1 or 2 small Amphipappus Pappus of 4-16 twisted or plane awns or scales; rays usually several 32 Pappus of 4 or 5 stiff, narrowly lanceolate awns; achenes linear, transversely rugulose Rigiopappus Pappus of scales, awn-tipped scales, or setose-dissected awns 33 Pappus of several scales dissected nearly to base; dwarf woolly annuals Syntrichopappus Pappus awns or scales not dissected or else plants perennial or woody 34 Pappus of several more or less united scales; rays broad, papery, and persistent Psilostrophe Pappus not of united scales; rays not papery and persistent (occasionally so in Hymenoxys) 35 Leaves and involucre with conspicuous oil glands Dyssodia Leaves and involucre without conspicuous oil glands 36 Achenes slender, elongate-clavate 37 Achenes stouter, oblong or obovoid 38 Plants woolly Eriophyllum Plants merely strigose Platyschkuhria Involucral bracts spreading or reflexed; receptacle convex to subglobose; leaves decurrent Helenium Involucral bracts appressed; receptacle almost flat; leaves not decurrent 39 Pappus of numerous scales; stems leafy; leaves linear or linear-spatulate, entire, 2.5 mm wide or less Gutierrezia Pappus of about 5 scales; leaves lobed or, if entire, broader and mostly or en- tirely basal Hymenoxys AcAMPTOPAPPUs Gray strongly imbricate, the bracts broad, Shrubs with white bark; leaves alternate, ^-ounded, the tip greenish, the margin entire, 1-nerved; heads yellow, discoid, sub- scarious, erose; receptacle convex, fimbril- globose, cymose at tips of branches; flowers late; style branches linear; achenes sub- all fertile; involucral bracts ca 4-seriate, turbinate, densely villous; pappus persistent. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 193 of ca 35 flattened silvery scales and bristles of different widths. Acamptopappus sphaerocephalus (Harv. & Gray) Gray Goldenhead. [Haplopappus sphaerocephalus Harv. & Gray]. Low rounded shrubs to 1 m tall, much branched, glabrous throughout or scabrous along some leaf margins; leaves spatulate to almost lin- ear, 4-28 mm long, 1-5 mm wide, obtuse to acute, mucronulate, thick, sessile; heads sub- globose, 6-10 mm high; involucre 4-6 mm high. Blackbrush, indigobush, and creosote bush communities at 850 to 1375 m in Kane, San Juan, and Washington counties; Arizona, Nevada, and California; 22 (iv). Achillea L. Perennial, rhizomatous, aromatic herbs, with watery juice; stems erect or ascending; leaves alternate, 1- to 3-pinnately dissected; leaves several to many, borne in compact to open corymbose cymes; involucral bracts im- bricate in several series, chaffy, the margins scarious and hyaline; receptacle chaffy; ray flowers present, usually 3-12, pistillate, fer- tile, yellow, white, pink, or pink-purple; disk flowers mostly 10 or more, perfect, fertile; pappus none; style branches flattened; achenes compressed, callus margined, glabrous, beakless. 1. Flowers yellow; leaves coarsely twice pinnately dissected; plants cultivated A. filipendulina — Flowers white, pink, or pink-purple; leaves finely 2-3 times dissected; plants indigenous or cultivated A. millefolium Achillea filipendulina Lam. Fernleaf Yar- row. Herbs, the stems erect, 8-12 dm tall or more, longitudinally furrowed and minutely glandular; leaves 4-35 cm long, doubly pin- natifid, the lateral lobes with one large lobe on the upper side; heads numerous, borne in hemispheric or flat-topped corymbose cymes; involucre 3-4 mm high, the bracts with pale scarous margins, villous; rays about 5, to 1 mm long, yellow; disk flowers 30-40, yellow; achenes 1-2 mm long. Cultivated ornamen- tal. Salt Lake and Utah counties, and to be expected elsewhere; introduced from Asia; 2 (0). Achillea millefolium L. Milfoil Yarrow. Herbs, the rhizomes horizontal; stems ascend- ing to erect, 0.5-10 dm tall, villous-tomen- tose, simple or branched above; leaves 2-26 cm long, reduced upwards, pinnately once to thrice dissected, the segments very slender; heads numerous, borne in hemispheric or flat-topped, corymbose cymes; involucres 4-6 mm high, the bracts dark to pale mar- gined, villous to glabrate; rays usually about 5, 2-3.5 mm long, white to pink or pink- purple; disk flowers 10-20; achenes 1-2 mm long. Gravelly, sandy, and clayey soils in sagebrush, pinyon-juniper, cottonwood, juni- per, rabbitbrush, ponderosa pine, mountain brush, aspen, Douglas fir, spruce-fir, and al- pine tundra commtmities at 1070 to 3750 m in all Utah counties; widely distributed in North America; circumboreal. Two very sim- ilar taxa are present in Utah; the common, in- digenous ssp. lanulosum (Nutt.) Piper (n=18), and the introduced, cultivated, ssp. millefolium (n = 27). A trend is recognizable within ssp. lanulosum; the high elevation specimens tend to have dark involucral bracts, fewer heads, and lower stature. These alpine plants have been treated as var. alpi cola (Rydb.) Garrett, but they intergrade completely with specimens attributable to var. lanulosa. Indeed, the two extremes can be found mounted on the same herbarium sheet, taken from the same locality; 133 (xv). Agoseris Raf. Perennial scapose herbs with milky juice, from taproots; leaves all basal, entire to pin- nately lobed or merely toothed; heads soli- tary on a naked scape; involucral bracts in 2 to several series, herbaceous, or the inner ones hyaline or nearly so; receptacle usually naked; corollas all raylike, perfect, yellow to orange, often drying pinkish or purplish; pap- pus of capillary bristles; style branches semi- cylindric; achenes angular or terete, prom- inently nerved, usually beaked. 194 Great Basin Naturalist Vol. 43, No. 2 1. Plants annual; achene beak 2-3 times as long as the body; rare in Utah A. heterophylla — Plants perennial; achene beak less than half to 2 or more times as long as the body 2 2(1). Achene beak striate, mostly less than half as long as the body (longer in some var. laciniata); flowers yellow, often drying bluish to pinkish A. glauca — Achene beaks scarcely striate, more than half to 2 or more times as long as the body 3 3(2). Flowers brownish orange to yellow-orange, often drying purplish; achene beak less than twice as long as the body A. aurantiaca — Flowers yellow, often drying bluish or pinkish; achene beak more than twice as long as the body A. grandiflora Agoseris aurantiaca (Hook.) Greene Orange Agoseris. Plants 0.6-6.6 dm tall, from a simple or branched caudex; leaves 3.5-36 cm long, 0.5-3 cm broad, narrowly oblanceolate, entire to toothed or lobed, villous to glabrate; scapes villous-tomentose to nearly glabrous; involucres 10-27 mm long, 10-42 mm wide, the outer bracts villous to glabrate and cil- iate, often purple spotted; corollas brownish orange to yellow-orange, often drying pur- plish; achene body 4-8 mm long, the slender beak not striate, from more than half as long to longer than the body. Two rather weak and intergrading phases are recognized at va- rietal rank. 1. Involucres with bracts subequal or nearly so, slender, tapering, some often over 20 mm long A. aurantiaca var. aurantiaca — Involucres with bracts definitely imbricate, broad, and rounded apically or abruptly tapering A. aurantiaca var. purpurea Var. aurantiaca [Troximon aurantiacum Hook.; A. arizonica Greene; A. gracilens (Gray) Kuntze; A. longirostris Greene, type from Fish Lake]. Sagebrush, mountain brush, juniper, pinyon-juniper, and alpine meadow communities at 1375 to 3355 m in Beaver, Box Elder, Carbon, Duchesne, Juab, Salt Lake, San Juan, Sevier, Tooele, Uintah, Utah, and Wasatch counties; Alberta to British Co- lumbia, south to California and New Mexico; 33 (v). Var. purpurea (Gray) Cronq. [Troximon aurantiacum var. purpureum Gray; A. pur- purea (Gray) Greene; A. confinis Greene, type from near Marysvale]. Mountain brush, aspen, aspen-fir, and spruce-fir communities at 1700 to 3425 m in Carbon, Emery, Grand, Juab, Iron, Piute, Sanpete, Sevier, and Wasatch counties; Montana to Arizona and New Mexico; 20 (vi). Agoseris glauca (Pursh) Raf. Pale Agoseris; Mountain Dandelion. Plants perennial, 0.2-6.4 dm tall, from a simple or branched caudex; leaves 2-26 cm long, 0.2-3 cm broad, narrowly oblanceolate to linear or spatulate to elliptic, entire or toothed to lobed, villous to glabrate; involucres 12-28 mm high, 0.8-4 cm wide, the outer bracts vil- lous to glabrous, ciliate or not, sometimes purple spotted; corollas yellow, often drying bluish to pinkish; achene body 4-10 mm long, the striate beak stout, to half as long as the body (slender and to as long as the body in some var. laciniata). Three intergrading and partially sympatric varieties are present in Utah. 1. Leaves laciniately toothed or lobed; plants of broad distribution, common A. glauca var. laciniata April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 195 — Leaves entire, rarely with a few teeth or lobes; plants variously distributed, lo- cally common 2 2(1). Plants pubescent, at least below the heads; mainly of spruce-fir and alpine communities, sometimes lower A. glauca var. dasycephala — Plants glabrous throughout, mainly of lower elevation wet meadows, but some- times of high elevation meadows A. glauca var. glauca Var. dasycephala (T. & G.) Jeps. [Am- mogeton scorzoneraefolius Shrad.; A. scorzo- neraefolia (Shrad.) Greene; Troximon pumi- lum Nutt.; A. pumila (Nutt.) Rydb.; A. glauca var. pumila (Nutt.) Garrett; T. glaucum var. dasycephalum T. & G.; A. villosa Rydb.]. Sagebrush, mountain brush, aspen, spruce-fir, and alpine tundra communities at 1830 to 3385 m in Duchesne, Garfield, Iron, Kane, Piute, Salt Lake, San Juan, Sanpete, Sevier, Tooele, Uintah, and Utah counties; Alaska to Manitoba and south to Colorado. Plants of this variety pass by degree into each of the following; 29 (v). Var. glauca [Troximon glaucum Pursh; A. isomeris Greene, type from the Uinta Moun- tains]. Meadows at 2325 to 3660 m in Box El- der, Duchesne, Iron, Juab, Sanpete, Sevier, Svimmit, Uintah, and Wasatch counties; Brit- ish Columbia to Manitoba and south to Cali- fornia and Arizona; 22 (iv). Var. laciniata (D.C. Eaton) Smiley [Troxi- mon parviflorum Nutt.; A. parviflora (Nutt.) D. Dietr.; A. glauca var. parviflora (Nutt.) Rydb.; T. taracifolium Nutt.; A. taracifolia (Nutt.) D. Dietr.; Macorhynchus glaucus var. laciniatus D.C. Eaton; A. taraxacoides Greene, type from near Marysvale; A. caud- ata Greene, type from Salina Canyon; A. agr- estis Osterh.; A. glauca var. agrestis (Osterh.) Q. Jones]. Sagebrush, mountain brush, juni- per, pinyon-juniper, Douglas fir, aspen, and spruce-fir communities at 1300 to 3050 m in Beaver, Box Elder, Carbon, Daggett, Du- chesne, Emery, Garfield, Grand, Juab, Kane, Millard, Piute, San Juan, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, Wash- ington, Wayne, and Weber counties; Wash- ington to Montana and south to Arizona. The phase designated as var. agrestis blends com- pletely in our area with that treated herein as var. laciniata; 84 (xii). Agoseris grandiflora (Nutt.) Greene [Stylopappus grandiflorus Nutt.]. Plants pe- rennial, 1.5-4.5 (7) dm tall, from a simple or branching caudex; leaves 8-25 cm long, 1-3 cm broad, narrowly oblanceolate, pinnatifid to subentire, villous to glabrate; involucres 15-38 mm long, 20-43 mm wide, the outer bracts villous-tomentose to glabrate, ciliate, often suffused with purple; corolla yellow, drying bluish to pinkish; achene body 4-7 mm long, the nerveless beak more than twice as long as the body. Specimens tentatively as- signed to this species are from Cache, Iron, Tooele, and Washington counties, where they occur in sagebrush and mountain brush communities at 1830 to 2135 m; British Co- lumbia to California and Nevada; 4 (i). Agoseris heterophylla (Nutt.) Greene An- nual Agoseris. [Macrorhynchus heterophyllus Nutt.]. Plants annual, 0.3-2.5 (4) dm tall, with 1 to several scapes from the base; leaves 1-20 (15) cm long, 0.3-1.5 cm wide, narrow- ly oblanceolate, toothed or pinnatifid to en- tire, all basal, or with some not strictly basal; involucres 5-20 mm long, 4-10 mm wide, sparingly villous with multicellular hairs, the cross-walls purplish; corolla yellow, some- times turning pinkish on drying; achene body 2-5 mm long, prominently ribbed or winged, the beak 2-3 times as long as the body. Our one collection (Diehl D29, 1899 BRY) is from Salt Lake County at low elevation; British Columbia to California and Arizona; 1 (0). Ambrosia L Annual or perennial herbs or shrubs; leaves alternate or opposite, pinnately or palmately lobed, toothed, or dissected; heads unisexual, discoid; staminate heads in slender spicate, bractless racemes; involucre 5- to 12-lobed; 196 Great Basin Naturalist Vol. 43, No. 2 receptacle flat, bearing flattened filiform-se- tose bracts; staminal filaments monadelphous, the anthers scarcely united; pistillate heads borne below the fertile ones, mostly axillary, their involucres closed, nutlike, armed with prickles arranged in one or more series; pistil naked, the corolla lacking; pappus lacking. Payne, W. W. 1964. A ree valuation of the genus Ambrosia. J. Arnold Arboretum 45:401-438. 1. Plants woody shrubs of southwestern Utah 2 — Plants armual or perennial herbs, of various distribution 3 2(1). Leaves mainly less than 15 mm long, pinnately lobed, the lobes again toothed or lobed, silvery-strigose overall A. dumosa — Leaves mainly more than 20 mm long, merely toothed or lobed, the lobes not again toothed or lobed, bicolored, the upper surface green, the lower surface white-tomentose A. eriocentra 3(1). Leaves palmately lobed, the lobes serrate; plants tall coarse herbs A. trifida — Leaves pinnatifid or pinnately lobed; plants slender herbs usually less than 5 dm tall 4 4(3). Leaves bicolored, the lower surface obscured by appressed white hairs; plants low rhizomatous perennials A. tomentosa — Leaves various, but not definitely bicolored; plants from taproots or rhizomes, but, if the latter, not as above 5 5(4). Plants perennial, rhizomatous; leaves opposite A. psilostachya — Plants annual; leaves mainly alternate 6 6(5). Lower stems and leaves with pustular-based, stiff, multicellular hairs; plants of- ten with lower lateral branches decumbent-ascending; burs with spines in more than one series A. acanthicarpa — Lower stems lacking pustular-based hairs, all stems slender and curved ascend- ing-appressed; burs with spines in one series A. artemisiifolia Ambrosia acanthicarpa Hook. Bur Rag- weed. [Franseria acanthicarpa (Hook.) Co- ville]. Plants annual, 0.9-7.5 dm tall, often branching from the base, the lower branches commonly decumbent-ascending; pubescence of stiff multicellular hairs, the bases pustular; leaves mostly alternate, petiolate, the blades 0.9-4.5 cm long, 0.6-3.5 cm wide, bipinnati- fid to pinnatifid; heads numerous in terminal or axillary racemes; staminate above, pistil- late below; staminate heads short-peduncu- late, not bracteate; pistillate solitary or clus- tered in upper axils, with 2-3 series of flattened, curved spines. Blackbrush, salt desert shrub, desert shrub, pinion-juniper, and riparian communities, often in sandy sub- strates, at 850 to 2000 m, in Beaver, Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Iron, Juab, Kane, Millard, San Juan, Tooele, Uintah, Utah, Washington, and Wayne coun- ties; Washington to Saskatchewan, south to California, Arizona, and Texas; 60 (xi). Ambrosia artemisiifolia L. Common Rag- weed. [A. elatior L.]. Plants annual, mostly 3-9 dm tall, branching from above the middle; pubescence of lax multicellular hairs, the bases not pustular; leaves alternate, or the lower usually opposite, petiolate, the blades 2.5-8.5 cm long, 1.9-7.5 cm wide, 1- to 2-pinnatifid; heads numerous in terminal or axillary racemes, the staminate above, pis- tillate below, clustered or solitary, with 1 series of tuberculate spines. Moist disturbed sites at 1375 to 1500 m in Juab and Utah counties; widespread in North America; 11 (i)- Ambrosia dumosa (Gray) Payne Bur-sage. [Franseria dumosa Gray]. Shrubs, 2-6 dm tall, rounded, much branched; branchlets white, subspinescent; pubescence dense, stri- gose; leaves alternate, petiolate, the blades 9-30 mm long, 5-15 mm wide, mostly 2-pin- natifid, uniformly hairy on both sides; stami- nate heads spicate, rather few; pistillate April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 197 heads often scattered among the staminate; pistillate involucre maturing 20-35 lance- subulate spines. Creosote bush, blackbrush, and Joshua tree communities at 670 to 1000 m in Washington County; Arizona, Califor- nia, and Mexico; 28 (i). Ambrosia eriocentra (Gray) Payne [Fran- seria eriocentra Gray]. Shrubs, 3-10 (12) dm tall, aromatic, branchlets white, sub- spinescent, pubescence of white tomentum and coarse multicellular hairs; leaves alter- nate, subsessile, sinuately toothed to lobed or 1-pinnatifid, 8-40 (50) mm long, 2-20 mm wide; staminate heads more or less clustered; pistillate heads 1-flowered; pistillate in- volucre with 12-20 flattened, subulate spines. Creosote bush, blackbrush, and Joshua tree communities at 670 to 1000 m in Washington County; Arizona, Nevada, and California; 12 (ii). Ambrosia psilostachya DC. Western Rag- weed. [A. coronopifolia T. & G.]. Perennial herbs, mostly 3-6 dm tall, simple or branch- ing above the middle; pubescence of harsh, spreading, multicellular, pustular-based hairs (at least in part); leaves opposite, at least be- low, petiolate to subsessile, the blades 4-10 cm long, 2.5-4.5 cm wide, mostly once pin- natifid; staminate heads in terminal or ax- illary spicate racemes; pistillate involucres merely tuberculate or quite unarmed. Mead- ows, stream banks, and roadsides in sagebrush and other communities at 1300 to 2100 m in Davis, Juab, Millard, Salt Lake, Utah, and Weber counties; Washington to Illinois, south to Arizona and Mexico; 20 (ii). Ambrosia tomentosa Nutt. [Franseria dis- color Nutt.; F. tomentosa (Nutt.) A. Nels., not A. tomentosa Gray]. Perennial rhizomatous herbs, mostly 1-3.5 dm tall, branching from above the base; pubescence of short, stiff, ap- pressed hairs; leaves alternate, petiolate, the blades 2-15 cm long, 0.4-3.5 cm wide, 1- to 3-pinnatifid; staminate heads racemose; pis- tillate heads armed with 2 or 3 series of coarse spines. Meadows and stream banks at 1300 to 1525 m in Davis, Duchesne, and Grand counties (likely elsewhere); Wyoming and Colorado; 5 (iii). Ambrosia trifida L. Giant Ragweed. An- nual, robust herbs, 10-15 dm tall or more; pubescence spreading-hirsute to hispid, at least above; leaves opposite, petiolate, the blades palmately 3- to 5-lobed, or unlobed, mainly 5-20 cm long, 4-15 cm wide, scabrous on both surfaces, serrate; staminate involucres 3-nerved; pistillate involucres 5-10 mm long, bearing short spines at the tip. Uncommon (introduced?) weedy plants of disturbed sites in Salt Lake County (likely elsewhere); widely distributed in North America; 1 (0). Amphipappus T. & G. Low shrubs; branches white-barked, divari- cate; leaves alternate, entire, short-petiolate; heads small, radiate, few flowered, clustered at tips of branches; involucre in ca 3 series, strongly imbricate, straw colored to greenish, the bracts broad, rounded; receptacle fimbril- late; ray flowers yellow, 1 or 2, small; disk flowers 3-6, perfect; ray achenes hairy, broadly oblanceolate, compressed, their pap- pus of more or less united bristles, awns, or scales; disk achenes undeveloped, glabrous or sparingly pilose, their pappus of twisted, his- pidulous bristles or scales. Porter, C. L. 1943. The genus Amphipappus Torr. & Gray. Amer. J. Bot. 30: 481-483. Amphipappus fremontii T. & G. Chaff- bush. Shrubs 3-8 dm tall, the herbage scabrous-puberulent; leaves 5-12 mm long, 2-5 mm wide, oblanceolate to elliptic, cu- neate basally, acute to obtuse and apiculate, green; heads 4-6 mm high, the bracts green- ish medially near the apex, the margins hyal- ine and more or less erose. Joshua tree and creosote bush communities at 700 to 900 m in Washington County; Nevada, Arizona, and California. Our material belongs to var. spin- osus (A. Nels.) C. L. Porter [ssp. spinosus (A. Nels.) Keck]; 4 (i). Anaphalis DC. Perennial, dioecious or polygamo- dioecious, rhizomatous herbs, with watery juice; stems ascending to erect, simple or branched above; leaves simple, alternate, en- tire; heads several to many, in hemispheric or flat-topped corymbose cymes; involucral bracts imbricate in several rows, chaffy, scarious, white, or with a dark triangular bas- al spot; receptacle naked; corollas of disk flowers only, imperfect, whitish, the pistillate 198 Great Basin Naturalist Vol. 43, No. 2 heads sometimes bearing some central stami- chesne, Iron, Juab, Salt Lake, Summit, nate flowers, the pistillate corollas tubular- Wasatch, and Washington counties; widely filiform, the staminate corollas tubular-fun- distributed in North America; Asia; 33 (vi). nelform; pappus of capillary bristles; style branches somewhat flattened; achenes small, Antennaria Gaertn. roughened, glabrous to sparingly hairy. Anaphalis margaritacea (L.) Benth. & Perennial, dioecious herbs with stolons. Hook. Pearly Everlasting. [Gnaphalium mar- caudices, or rhizomes, the juice watery; stems garitaceum L.]. Plants 1.5-8 dm tall, the ascending to erect, usually simple; leaves stems white villous-tomentose; leaves only simple, alternate and basal, the cauline gen- gradually reduced upwards, 2.5-12 cm long, erally reduced upward; heads solitary to 0.5-2 cm wide, narrowly lanceolate to ob- many, borne in corymbose cymes; involucral long, elliptic, or oblanceolate, sessile, entire, bracts imbricate in several rows, scarious (at flat to slightly revolute, white-tomentose be- least marginally), often colored; receptacle low, commonly less pubescent and greenish naked; corollas of disk flowers only, imper- above; heads showy, the involucres 4-7 mm feet, whitish or tawny; pistillate corollas high, 5-10 mm broad, the bracts pearly- tubular-filiform, the pappus of numerous white, with a dark triangular base, glabrous; capillary bristles; staminate corollas tubular- achenes about 1 mm long. Meadows, stream- funnelform, the pappus of few clavate to bar- banks, and openings in ponderosa pine, bellate, usually flattened bristles; style lodgepole pine, box elder, and aspen commu- branches slightly flattened; achenes terete to nities at 1150 to 2700 m in Box Elder, Du- slightly compressed, glabrous or papillose. 1. Heads solitary; flowering stems usually less than 5 cm tall A. dimorpha — Heads (1) 2 to many (see A. rosulata); flowering stems often more than 5 cm tall 2 2(1). Upper leaf surface green; leaf blades broadly spatulate, rounded to obtuse A. neglecta — Upper leaf surface not notably different from the lower; blades seldom both spatulate and rounded to obtuse 3 3(2). Plants not forming mats, lacking leafy stolons, some caespitose from caudex or rhizomes 4 — Plants mat forming, with leafy stolons 6 4(3). Involucral bracts glabrous or nearly so, scarious near the base, white-opaque apically A. luzuloides — Involucral bracts densely tomentose in the lower half, opaque to dark with pale scarious apices 5 5(4). Involucral bracts blackish in aspect, the tips pale and scarious A. pulcherrima — Involucral bracts opaque white, somewhat darkened at the middle .. A. anaphaloides 6(3). Terminal scarious portion of involucral bracts dirty brownish to blackish green on at least the middle and outer ones 7 — Terminal scarious portion of involucral bracts white to pink, with a dark basal spot on some only 8 7(6). Terminal scarious portion of involucral bracts blackish green; plants usually al- pine in Uinta, Wasatch, and Tushar Mountains, and on the Markagunt Plateau A. alpina — Terminal scarious portion of bracts merely discolored and pale brown, or the inner bracts whitish at the tips; plants usually of lower elevations A. umbrinella 8(6). Flowering stems less than 5 cm tall; heads 1 or 2; plants of Garfield, Kane, and Wayne counties A. rosulata April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 199 — Flowering stems commonly more than 5 cm tall; heads usually 3 or more; plants of broad, or other, distribution 9 9(8). Involucral bracts with a black spot between the tomentose greenish base and the opaque white-scarious apex A. corymbosa — Involucral bracts lacking a conspicuous black spot 10 10(9). Involucres mostly 4-7 mm high, often bright pink; pistillate corollas mostly 2-4.5 mm long A. microphylla — Involucres mostly 7-11 mm high, seldom pink; pistillate corollas mostly 5-8 mm long A. parvifolia Antennaria alpina (L.) Gaertn. Alpine Pussytoes. Plants caespitose from a caudex, mat forming and stoloniferous, 2-13 cm tall; basal leaves 0.6-2.2 cm long, 2-6 mm wide, cuneate-oblanceolate to spatulate, acute to obtuse or rounded apically, grayish tomen- tose on both surfaces or greenish and sub- glabrous above on some leaves; heads 3-5, borne in subcapitate cymes; pistillate in- volucres 5-7 mm high, villous-tomentose be- low, the scarious tips of bracts uniformly blackish or brownish green, all rather blunt apically, often erose; staminate involucres mostly 4-5 mm high, the scarious tips of bracts often pale apically; achenes glabrous. Lodgepole pine, spruce-fir, and alpine tundra communities at 3050 to 3550 m in Daggett, Duchesne, Piute, Salt Lake, Uintah, and Utah counties; north to Alaska and east to Lab- rador; circumboreal. Our material belongs to var. media (Greene) Jeps. [A. media Greene; A. austromontana E. Nels., type from Tushar Mountains]. There is a tendency for some specimens to approach A. parvifolia in the Uinta Mountains and A. umbrinella else- where; 27 (vii). Antennaria anaphaloides Rydb. Pearly Pussytoes. Plants from a caudex, not mat forming or stoloniferous, 1.5-3.5 (5) dm tall; basal leaves 2.5-19 cm long, 4-18 mm wide, narrowly oblanceolate to elliptic, tomentose on both surfaces; heads several to many in branching or compact cymes; pistillate in- volucres 5-8 mm high, villous-tomentose be- low, the scarious tips opaque-white, all rounded or obtuse, often erose; staminate in- volucres 5-8 mm high, similar to the pistil- late; achenes glabrous. Aspen, spruce-fir, sagebrush, and mountain brush communities at 2440 to 3325 m in Daggett, Summit, and Uintah coimties; British Columbia to Mon- tana and south to Nevada and Colorado; 7 (i). Antennaria corymbosa E. Nels. Plains Pus- sytoes. [A. nardina Greene]. Plants caespi- tose, mat forming and stoloniferous, 5-26 cm tall; basal leaves 0.6-3.7 cm long, 2-6 mm wide, narrowly oblanceolate to spatulate, acute to obtuse apically, gray to greenish and tomentose on both surfaces; heads commonly 3-8, in compact to branching cymes; pistil- late and staminate involucres 4.5-6 mm high, the bracts green and tomentose basally, with a dark spot at the base of the white or sordid terminal portion; achenes puberulent. Alpine tundra, krumholz, spruce-fir, lodgepole pine, and willow-alder communities, often along stream banks and in wet meadows or bogs, at 2240 to 3355 m in Beaver, Duchesne, Gar- field, Summit, and Uintah counties; Montana and Idaho to Colorado and California(?). The main body of the species in Utah lies in the Uinta Mountains, with outliers in the Stans- bury and Tushar mountains, and in the Mark- agunt Plateau; 22 (iii). Antennaria dimorpha (Nutt.) T. & G. Low Pussytoes. [Gnaphalium dimorphum Nutt.; A. dimorpha var. macrocephala D.C. Eaton, type from Salt Lake City]. Plants caespitose, mat forming, rooting from short caudex branches, not truly stoloniferous, 1-5 (7) cm tall; basal leaves narrowly oblanceolate, 0.6-4 cm long, 1-14 mm wide, acute api- cally, grayish tomentose on both sides; heads solitary, terminal on short leafy stems; pistil- late involucres (7) 10-18 mm long, the bracts strongly imbricated, slender, attenuate, green at base, suffused with brown above the base, the apical portions yellowish to brownish scarious; staminate involucres 6-9 mm long, tomentose at the base, brown above the base, the broad apical portion hyaline to scarious; achenes puberulent. Mat-saltbush, sagebrush, juniper, oak-serviceberry, ponderosa pine, and spruce-fir-lodgepole pine communities at 200 Great Basin Naturalist Vol. 43, No. 2 1430 to 3050 m in Beaver, Daggett, Du- chesne, Iron, Juab, Millard, Salt Lake, San- pete, Sevier, Summit, Tooele, Utah, and Washington counties (and certainly else- where); British Columbia to Montana, south to California, Nevada, Colorado, and Nebras- ka; 37 (viii). Antennaria luzuloides T. & G. Rush Pus- sytoes. [A. oblanceolata Rydb.]. Caespitose from a caudex, 1.1-5 (7) dm tall; basal leaves 2-5 (8) cm long, 2-8 mm wide, greenish, to- mentose on both surfaces; heads numerous in a compact or more often branched corym- bose inflorescence; pistillate and staminate involucres similar, 4-5 mm high, glabrous to the base, the bracts brownish scarious and more or less hyaline below, opaque whitish above; achenes puberulent. Openings in as- pen-conifer and lodgepole pine-spruce com- munities at 2950 to 3050 m in Duchesne and Summit counties; British Columbia to Mon- tana, south to California, Nevada, and Colo- rado; 7 (i). Antennaria microphylla Rydb. Rosy Pus- sytoes. [A. rosea Rydb.; A. concinna E. Nels.; A. arida A. Nels.]. Plants caespitose, stoloni- ferous and mat forming, 0.4-3 (4) dm tall; basal leaves 0.5-3 cm long, 2-8 mm wide, oblanceolate to spatulate; heads 2-13 (or more), in congested to open cymes; pistillate involucres 4-7 mm high, the bracts tomen- tose below, greenish or scarious below the middle, often somewhat brownish below the scarious, whitish or pinkish, terminal portion; mainly known from pistillate individuals; achenes glabrous or sparingly hispidulous. Sagebrush, juniper, ponderosa pine, Douglas fir, lodgepole pine, spruce-fir, and alpine meadow communities at 1830 to 3450 m in Beaver, Box Elder, Carbon, Daggett, Du- chesne, Emery, Garfield, Iron, Juab, Kane, Millard, Piute, Rich, Salt Lake, Sanpete, Se- vier, Summit, Tooele, Uintah, Utah, Wasatch, Wayne, and Weber counties; Alaska to On- tario, south to California and New Mexico; 105 (xxiii). Antennaria neglecta Greene Field Pus- sytoes. [A. marginata Greene]. Plants caespi- tose, stoloniferous and mat forming, 5-15 (25) cm tall; basal leaves 1.8-3.5 cm long, 3-15 mm wide, spatulate, thinly tomentose to glabrous and green above, white-tomentose beneath; heads mainly 3-5, in compact cymes; pistillate involucres 6-11 mm high, the bracts tomentose on the greenish base, the apical scarious portion white or suffused with pink; staminate plants rare; achenes glabrous or minutely pubescent. Pinyon-juni- per and shrub communities at 1525 to 1900 m in San Juan, Utah, and Washington coun- ties; Alaska to Newfoundland, south to Cali- fornia, Arizona, and Virginia; 3 (i). Our few specimens are hardly adequate to represent this species clearly in Utah. Antennaria parvifolia Nutt. [A. aprica Greene; A. obtusita Greene]. Plants caespi- tose, stoloniferous and mat forming, 3-15 cm tall; basal leaves 0.8-3.5 cm long, 3-8 mm wide, spatulate, obtuse to acute apically, to- mentose on both sides; heads 2-6 or more; pistillate involucres 7-11 mm high, the bracts more or less tomentose on the greenish base, the scarious portion white, sordid, or pink; staminate plants rarely collected; achenes glabrous. Mountain brush, pinyon-juniper, sagebrush, ponderosa pine, aspen, lodgepole, and spruce-fir communities at 1650 to 3250 m in Beaver, Carbon, Daggett, Duchesne, Emery, Garfield, Juab, Kane, Piute, San Juan, Summit, Tooele, Uintah, Utah, and Wayne counties; British Columbia to Manitoba, south to Arizona and New Mexico; 46 (iv). Antennaria pulcherrima (Hook.) Greene Showy Pussy toes. [A. carpathica var. pulcher- rima Hook.]. Plants from a caudex, not mat forming or stoloniferous, 23-40 cm tall; basal leaves 4-19 cm long, 5-23 mm wide, narrow- ly to broadly oblanceolate to elliptic, tomen- tose on both surfaces; heads several to many in branching or compact cymes; pistillate and staminate involucres both 6.5-8 mm long, the bracts tomentose at the greenish base, the terminal scarious portion blackish to brownish or the apex whitish; achenes glabrous. Sedge-rush meadows, streamsides, and bogs at 2440 to 2800 m in Duchesne, Garfield, and Summit counties; Alaska to Newfoundland, south to Colorado; 5 (i). Antennaria rosulata Rydb. Plants caespi- tose, stoloniferous and mat forming, 1-3 cm tall; basal leaves 0.5-1.1 cm long, 2-5 mm broad, spatulate, obtuse to rounded apically, tomentose on both surfaces; heads 1 or 2, ter- minating short erect branches; pistillate in- volucres 5-9 mm high, the outer bracts April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 201 greenish and tomentose to the apex, the inner ones green at base, with scarious slender white tips; staminate involucres 4-5 mm high, the bracts densely tomentose at base, the broad scarious tips white-opaque; achenes puberulent. Ponderosa pine, aspen, Douglas fir, limber pine, sagebrush, and spruce communities, and in alpine meadows, at 2600 to 3350 m in Garfield, Kane, and Wayne counties; Colorado, New Mexico, and Arizona; 14 (i). Antennaria umbrinella Rydb. [A. dioica authors, not (L.) Gaertn.]. Plants caespitose, mat forming and stoloniferous, 2-14 mm tall; basal leaves 0.7-2 cm long, 0.2-1.5 cm wide, cuneate-oblanceolate to spatulate, acute to obtuse apically, tomentose on both sides; heads 2-6, borne in subcapitate cymes; pistil- late involucres 5-8 mm long, the bracts greenish and tomentose at the base, the scarious tips dirty brownish to pale tan, or the innermost almost white, acute to rounded, usually erose; staminate plants un- known in our region; achenes glabrous. As- pen communities and alpine meadows at 2745 to 3500 m in Duchesne, Juab, and Sum- mit counties; Alaska to Hudson Bay, south to California, Arizona, and Colorado; 4 (0). Specimens assigned here are more or less in- termediate between A. alpina and A. micro- phylla. Many more specimens are required to provide definitive information on this entity in Utah. Anthemis L. Annual or short-lived perennial, aromatic herbs from taproots, the juice watery; stems erect, commonly branched; leaves alternate, 1-3 pinnately dissected; heads solitary on the uppermost branches; involucral bracts imbri- cated in several series, chaffy, the margins scarious or hyaline; receptacle hemispheric, chaffy at least near the middle; ray flowers present, white or yellow, usually 10 or more, sterile; disk flowers numerous, perfect, fer- tile; pappus none or a short crown; style branches flattened; achenes subterete or com- pressed, not callous-margined, glabrous, beakless. Rays white; pappus lacking; disk commonly less than 10 mm broad A. cotula Rays yellow; pappus a short crown; disk commonly more than 12 mm broad A. tinctoria Anthemis cotula L. Mayweed. Plants an- nual, 1-7.5 dm tall; stems simple or branched, ill scented; leaves 1-6 cm long, twice pinnatifid, the ultimate segments lance-oblong, sparsely villous and glandular- dotted; heads borne solitary at the upper ends of the uppermost branches; ray flowers commonly 10-20, white, sterile, 5-10 mm long; disk flowers numerous; disk 4-10 (12) mm wide; receptacle chaffy only in the middle, the bracts narrowly subulate; achenes slightly flattened, glandular, the pappus lack- ing. Introduced Old World weeds of fields, roadsides, revegetated woodlands, and other disturbed sites at 1280 to 1400 m in Du- chesne, Morgan, Salt Lake, Utah, and Weber counties (likely elsewhere); widespread in North America; 7 (0). Anthemis tinctoria L. Yellow Camomile. Plants short-lived perennials, 2.5-6 dm tall; stems simple or branched; leaves 1.5-7 cm long, 1- to 2-pirmatifid, the segments oblong in outline, merely toothed or lobed, villous- tomentose below, glabrous or glabrate above, sparsely glandular-dotted; heads borne soli- tary at ends of the uppermost branches; ray flowers 20-35, yellow, fertile, 7-14 mm long; disk flowers numerous; disk 12-15 mm wide or more; receptacle chaffy throughout, the bracts narrow and with yellow awn-tips; achenes compressed; pappus a short crown. Old World cultivated ornamentals; widely planted and occasionally escaping (Salt Lake County, Garrett 8865 BRY); widespread in North America; 1 (0). Arctium L. Biennial, coarse herbs with watery juice, from a taproot; leaves rhubarblike, basal and alternate, entire or toothed; heads few to nu- merous in axillary or terminal corymbose or racemose inflorescences; flowers all tubular, perfect, the corollas pink to purplish; in- 202 Great Basin Naturalist Vol. 43, No. 2 volucres urn shaped, the bracts imbricate in achenes sHghtly compressed, more or less 3- many series, the tips slender and inwardly angled, many nerved, truncate apically; pap- hooked; receptacle flat, densely bristly; pus of numerous, scaly, deciduous bristles. 1. Heads mainly 1.5-2.5 cm thick, arranged in racemelike axillary clusters, the terminal also racemelike A. minus — Heads commonly over 2.5 cm thick, arranged in corymbose clusters, especially the terminal A. lappa Arctium lappa L. Great Burdock. Plants Sevier, Summit, Tooele, Uintah, Utah, 8-15 dm tall; basal leaves long-petiolate, the Wasatch, Wayne, and Weber counties, and blades commonly 2-5 dm long, 1-3 dm probably cosmopolitan; widespread in North broad, cordate-ovate, obtuse, thinly tomen- America; Eurasia; 23 (i). tose beneath, glabrous or nearly so above; in- florescence corymbosely disposed, the pe- Arnica L duncles glandular or glandular-hairy; heads 2.5-4 cm broad, the involucre greenish stra- Perennial herbs from rhizomes or caudices, mineus, glabrous or glandular, often sparingly the juice watery; stems erect, simple or arachnoid-tomentose. Cultivated for its branched above; leaves opposite or the up- edible roots, and persisting; introduced from permost alternate, simple, entire or toothed; Eurasia; 1 (0). heads solitary, or 3-9 (11) in corymbose clus- Arctium minus (Hill) Bernh. Burdock, ters; involucral bracts subequal or evidently Plants 5-15 dm tall; basal leaves long- biseriate, herbaceous; receptacle naked, con- petiolate, the blades commonly 1-3.5 (4) dm vex; ray flowers present, yellow or orange, long, 1-3 dm wide, cordate-ovate, obtuse, several to many, fertile, or lacking (in A. par- thinly tomentose to glabrous beneath, ryi); disk flowers numerous, perfect, fertile; glabrous above or nearly so; inflorescence pappus of barbellate or subplumose capillary racemosely disposed, the peduncles short or bristles; style branches flattened; achenes cy- lacking; heads 1-2.5 cm thick (rarely more), lindrical, 5- to 10-nerved, pubescent to gla- the bracts glabrous or glandular to definitely brate or glabrous, often glandular, arachnoid. Introduced Old World weed of Maguire, B. 1943. A monograph of the genus consequence in Cache, Juab, Millard, Piute, Arnica. Brittonia 4:386-510. 1. Heads discoid (rarely some with rays), the lateral (lower) ones spreading or re- flexed, the uppermost one erect A. parryi — Heads radiate, the lateral ones (if any) erect like the uppermost 2 2(1). Cauline leaves (4) 5-9 pairs; pappus brownish; heads often 5 or more per main stem 3 — Cauline leaves 1-4 (5) pairs; pappus white or brownish; heads mainly 1-4 per stem 4 3(2). Involucral bracts merely acute to abruptly rounded (rarely acuminate), bearing an apical or subapical tuft of hairs A. chamissonis — Involucral bracts acuminate to attenuate, not especially more hairy at the apex A. longifolia 4(2). Leaves (at least the lower) cordate, ovate, or broadly ovate-lanceolate, often cordate, truncate, or obtuse basally, seldom cuneate 5 — Leaves narrowly lanceolate to lance-oblong or lanceolate, usually cuneate ba- sally 7 5(4). Pappus brownish, subplumose; main cauline leaves obtuse to subcuneate ba- sally A. diversifolia April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 203 — Pappus white, merely barbellate; main cauline leaves usually cordate, truncate, or obtuse basally 6 6(5). Blades of main cauline leaves much longer than the petiole, or sessile; achenes glabrous throughout, or at least near the base A. latifolia — Blades of main cauline leaves subequal to or shorter than the petioles; achenes uniformly, though sometimes sparingly, hairy A. cordifolia 7(4). Pappus brownish, subplumose A. mollis — Pappus usually white or tawny, merely barbellate 8 8(7). Heads turbinate-campanulate, commonly with 7-10 rays; lower cauline leaves sessile or nearly so A. rydbergii — Heads hemispheric, commonly with 10-20 rays; lower cauline leaves often petiolate 9 9(8). Old leaf bases bearing dense brown wool in the axils; disk corollas both spread- ing hairy and stipitate-glandular A. fulgens — Old leaf bases lacking axillary tufts of hair, or with white hair only; disk corollas merely stipitate-glandular A. sororia Arnica chamissonis Less. [A. foliosa Nutt.; A. chamissonis ssp. foliosa (Nutt.) Maguire; A. foliosa var. incana Gray]. Plants 1-6 (8) dm tall, the stems erect or ascending, simple or more commonly branched in the in- florescence, sparsely to densely villous with multicellular hairs and often glandular as well; basal leaves 3-11 (15) cm long, 3-16 (20) mm wide, lanceolate to oblong or oblan- ceolate, with 3-5 main veins, pilose to villous or tomentose, tapering to a slender petiole, entire to distinctly toothed, smaller than the cauline ones and often withered by flowering time; cauline leaves (4) 5-8 (9) pairs, lanceo- late to lance-elliptic, the largest near the middle of stem or slightly below, the lower ones petiolate and with membranous con- nate-sheathing bases, the upper sessile, entire to distinctly toothed; heads (1) 3-9, the pe- duncle apex sparingly to densely villous with whitish hairs often intermixed with glands; involucres 9-15 mm high, the bracts lanceo- late, obtuse, acute, or" less commonly acumi- nate, sparsely to densely pilose, ciliate, the tips with a conspicuous tuft of whitish hairs; rays usually 10-16, yellow; achenes 4-6 mm long, hairy to glandular or glabrate; pappus brownish to straw colored, barbellate. Stream banks, gravel bars, and lake shores in aspen, willow, and spruce-fir communities at 2300 to 3350 m in Duchesne, Emery, Garfield, Iron, Kane, Salt Lake, Sanpete, Sevier, Sum- mit, Tooele, Wasatch, Washington, and Wayne counties; Alaska to Hudson Bay, south to California and New Mexico; 29 (v). Maguire (1943) treated all Utah material as ssp. foliosa (Nutt.) Maguire. Cronquist (Univ. Washington Publ. Biol. 17(5): 45-54. 1955) cited var. incana (Gray) Hulten [ssp. incana (Gray) Maguire] from Utah. Our specimens are only arbitrarily separable into two phases, differing mainly in degree of pu- bescence. The correct name at varietal level for our gray hairy plants appears to be A. chamissonis var. andina (Nutt.) Ediger & Barkley. Arnica cordifolia Hook. Plants 1.5-4 dm tall, the stems erect or ascending, simple or branched above, sparsely villous with multi- cellular hairs and often glandular as well; basal leaves smaller than the cauline, often withered at anthesis; petioles of main leaves (at least) often longer than the blades; cauline leaves 2-4 (5) pairs, the blades 2-9 cm long (from sinus to apex), 1-9 cm wide, cordate- ovate to orbicular or reniform, or the up- permost lanceolate, the largest below the middle of the stem, the lower leaves petio- late, the upper ones sessile or subsessile, ser- rate-dentate to subentire; heads 1 (3), rarely more, the peduncle apex villous with whitish hairs often intermixed with glands; involucres 14-20 mm high, the bracts lanceolate to ob- long, acuminate to acute, sparsely to densely pilose and often glandular-ciliate, the tip with a moderate tuft of hair; rays usually 10-15, yellow; achenes 4-5.5 mm long, uni- formly hairy and often glandular; pappus 204 Great Basin Naturalist Vol. 43, No. 2 white, barbellate. Sagebrush, Douglas fir, white fir, lodgepole pine, ponderosa pine, as- pen, and spruce-fir communities at 1525 to 3355 m in Beaver, Box Elder, Cache, Carbon, Daggett, Duchesne, Garfield, Iron, Juab, Piute, Salt Lake, San Juan, Sanpete, Sevier, Summit, Uintah, Utah, Wasatch, Washington, Wayne, and Weber counties (likely univer- sal); Alaska to Michigan, south to California, Arizona, New Mexico, and Nebraska; 102 (xii). The white pappus and cordate long- petiolate leaves are diagnostic for this species. Arnica diversifolia Greene Plants 1.5-4.2 dm tall, the stems erect or ascending, simple or branched above, sparsely villous with mul- ticellular hairs and often glandular, or almost glabrous; basal leaves smaller than the cau- line and often withered by flowering time, borne on slender to broadly winged petioles shorter than or subequal to the blades; cau- line leaves 2-4 (5) pairs, blades 2-8 cm long, 0.8-4 (6) cm wide, ovate or the uppermost lanceolate, the largest at the middle or be- low, becoming sessile to subsessile above, subentire or irregularly serrate; heads 1-3 or more, the peduncle apex sparsely to moder- ately villous with whitish hairs and often with glands; involucre 10-16 mm high; bracts lanceolate, acuminate to acute, sparse- ly to densely pilose and often glandular, cil- iate, the tip lacking a tuft of hairs; rays usual- ly 10-15, yellow; achenes 5-7 mm long, glabrous or sparsely and uniformly hairy; pappus brownish, subplumose. Stream sides, meadows, and scree slopes in spruce-fir and alpine tundra communities at 2560 to 3400 m in Duchesne, Grand, Piute, San Juan, San- pete, Summit, Uintah, and Utah counties; Alaska and Yukon, south to Oregon and Colo- rado; 19 (ii). This taxon is not well collected in Utah. The broad leaves and brownish sub- plumose pappus are diagnostic for these plants that might be regarded as consisting of a series of hybrid derivatives between A. mol- lis and A. cordifolia, A. latifolia, or A. ryd- bergii. More work is necessary. Arnica fulgens Pursh [A. pedunculata Rydb.]. Plants 1.5-6 (7) dm tall, the stems erect, the basal leaf axils with tufts of long brown woolly hair, otherwise stipitate- glandular and often hairy as well; basal leaves smaller than the cauline, often with- ered at anthesis, with broadly winged pet- ioles or subsessile; cauline leaves 2-4 pair, the blades oblanceolate to elliptic (often nar- rowly so), mostly 3-12 cm long, 0.6-4 cm wide, the largest ones near the base, becom- ing sessile upward, subentire to entire; heads 1-3, the peduncle apex yellowish villous; in- volucre 10-15 (18) mm high, the bracts nar- rowly elliptic to lance-elliptic, attenuate to an obtuse or acute apex, villous, the tips pu- bescent within; rays mostly 10-20, yellow to yellow-orange; achenes 4-5.5 mm long, densely hairy; pappus whitish to cream col- ored, barbellate. Dry sagebrush community at 2000 m in Daggett County; British Colum- bia to Saskatchewan, south to California, Ne- vada, and Colorado; 1 (0). Arnica latifolia Bong. [A. gracilis Rydb.; A. jonesii Rydb.]. Plants 1-4 (6) dm tall, the stems erect or ascending, simple or branched above, sparsely villous with multicellular hairs and often glandular; basal leaves smaller than the cauline, usually withered by flowering time, the petioles (if any) usually shorter than the blades; cauline leaves 2-5 pairs, the blades 2-4.5 (7) cm broad, cordate- ovate to lanceolate, the largest ones at the middle or below, the lower ones with petioles shorter than the blades, the upper ones sessile or subsessile, serrate-dentate, less commonly entire or nearly so; heads 1-5 or rarely more, the peduncle apex sparsely to moderately vil- lous with whitish or yellowish hairs and often glandular; involucres 9-17 mm high, the bracts lanceolate, acuminate to acute, sparse- ly pilose and often glandular, ciliate, lacking an apical tuft of hair; rays usually 8-12, yel- low; achenes 5-8 mm long, glabrous or sparsely hairy, or glabrous in the lower por tion; pappus white, barbellate. Lodgepole pine, spruce-fir, and alpine tundra commu- nities at 2240 to 3400 m in Duchesne, Salt Lake, Summit, and Utah counties; Alaska and Yukon to California and Colorado; 24 (v). Specimens available for study are variable. TTiey occur in the Uinta and Wasatch moun- tains and on the Tavaputs Plateau. The var. gracilis (Rydb.) Cronq. was reported from Utah by Maguire (I.e., as A. gracilis Rydb.), but has not been seen by me. It differs from the bulk of our material in its small size (1-3 dm), more numerous heads (3-9), and narrow April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 205 small involucre 9-13 mm high. The single collection cited by Maguire is from Salt Lake or Utah County. More material is necessary. Arnica longifolia D.C. Eaton in Wats. [A. caudata Rydb., type from Big Cottonwood Canyon]. Plants 3-10.5 dm tall; stems erect or sprawling, tufted from caudexlike short- ened rhizomes, simple or branched above, shortly villous to puberulent and often some- what glandular-viscid; basal leaves lacking or soon withering, the cauline ones 5.5-20 cm long, 0.6-3 cm wide, lanceolate to elliptic, with 3-5 main veins, puberulent, all sessile, 5-7 pairs, the largest near the middle of the stem, the lower ones connate-sheathing, en- tire or nearly so; heads 1-9, the peduncle apex sparingly yellowish villous; involucres 6-13 mm high, the bracts lanceolate to lance- oblong, acute to aciiminate, sparingly pilose and glandular, ciliate, the tips sparingly white-hairy; rays mainly 8-13, yellow; achenes 4.5-5.5 mm long, glabrate, or uni- formly stipitate-glandular; pappus brownish to straw colored, barbellate. Snow flushes, talus, and stream banks in lodgepole pine, as- pen, ponderosa pine, Douglas fir, white fir, and spruce-fir communities at 1890 to 3325 m in Box Elder, Cache, Garfield, Grand, Juab, Rich, Salt Lake, Sanpete, Summit, Tooele, and Washington counties; Washing- ton to Alberta, south to California, Nevada, and Colorado; 15 (i). Our material belongs to var. longifolia. Arnica mollis Hook. [A. arachnoidea Rydb., type form Big Cottonwood Canyon; A. chamissonis var. longinodosa A. Nels., type from near Marysvale; A. ovata Greene, type from Alta]. Plants 1.5-6.5 dm tall, the stems erect or ascending, loosely to com- pactly clump-forming, simple, or branched in inflorescence, puberulent to villous and glandular; basal leaves smaller than the cau- line ones, often withered at anthesis, the cau- line ones 4.5-18 cm long, 0.8-4 cm wide, ob- lanceolate to obovate, lanceolate or elliptic, the lower slenderly to broadly petiolate, be- coming sessile upwards, 3-4 pairs, the largest below the middle, the lower connate-sheath- ing, entire to irregularly denticulate; heads 1-5 (7), the peduncle apex sparingly yellow- ish villous; involucres 10-17 mm high, the bracts lanceolate to lance-elliptic, acute to attenuate, sparingly to densely villous-pilose and more or less glandular, lacking a sub- apical tuft of hair; rays mainly 12-18, yellow; pappus brownish, subplumose; achenes pu- bescent to stipitate-glandular. Meadows, bogs, stream banks, seeps, talus slopes, and rock stripes in sagebrush, ponderosa pine, lodgepole pine, Douglas fir, white fir, aspen, spruce-fir, and alpine tundra communities at 1950 to 3550 m in Box Elder, Cache, Dag- gett, Duchesne, Garfield, Juab, Kane, Piute, Rich, Salt Lake, Sanpete, Summit, Tooele, Uintah, and Utah counties; British Columbia to California, Nevada, and Colorado; 50 (xii). Arnica parryi Gray. Plants 1.5-5 (6) dm tall, erect or ascending, from elongate rhi- zomes, simple or branched in inflorescence, villous and more or less glandular; basal leaves smaller than the cauline ones, often withered at anthesis, the cauline ones long petioled below, becoming sessile upwards, the blades 2-9.5 cm long, 0.4-4 cm wide, lan- ceolate to ovate, the base obtuse to truncate or cuneate, 2-4 (5) pairs, the largest near the stem base, the lower connate-sheathing, en- tire to denticulate; heads 3-12, nodding in bud, the peduncle apex glandular-villous; in- volucres 10-16 mm high, the bracts narrowly lanceolate, acute to attenuate, glandular-vil- lous, lacking a subapical tuft of hairs; rays lacking, or rarely present, yellow; pappus brownish, barbellate to subplumose; achenes glabrous to glandular or hairy. Aspen and spruce-fir communities at 2415 to 3175 m in Carbon, Daggett (?), Garfield, Iron, Salt Lake, Sanpete, and Summit counties; British Columbia and Alberta to California and Col- orado; 9 (ii). A specimen from Piute County (Welsh et al. 14018 BRY) has ray flowers well developed. Arnica rydbergii Greene. Plants 1-2.6 dm tall, erect or ascending, from elongate rhi- zomes, sparingly villous and shortly stipitate- glandular; basal leaves smaller than the cau- line, sometimes bladeless, often withered at anthesis, the cauline ones short to long pet- ioled below, becoming sessile upwards, the blades 2-5 cm long, 0.4-1.5 cm wide, lan- ceolate to elliptic, ovate, or obovate, the base obtuse to cuneate, 2 or 3 (4) pair, the largest at or near the middle of the stem, the lower connate-sheathing, entire or denticulate; heads 1-5, the peduncle apex yellowish vil- lous, glandular; involucres 9-13 mm high, the 206 Great Basin Naturalist Vol. 43, No. 2 bracts narrowly lanceolate, acute to attenu- ate, stipitate-glandular, ciliate, lacking a sub- apical tuft of hair; rays mainly 7-10, yellow; pappus white, barbellate; achenes shortly pi- lose. Spruce-fir and lodgepole pine forests in Duchesne, Summit, and Utah counties; Brit- ish Columbia and Alberta to Oregon and Col- orado; 12 (iii). Arnica sororia Greene Plants 1.5-6 dm tall, the stems erect, the basal axils lacking tufts of hair, otherwise more or less villous and glandular; basal leaves smaller than the cauline, often withered at anthesis, with winged to narrow petioles or subsessile; cau- line leaves 2-4 pair, the blades lanceolate to elliptic, mostly 3-10 cm long, 0.5-2 cm wide, the largest ones near the base, becoming ses- sile upward, mainly entire; heads 1-3, the pe- duncle apex sparingly villous; involucres 10-15 mm high, the bracts narrowly oblong- lanceolate, attenuate, villous, the tips more or less hairy within; rays mainly 9-15, yel- low; achenes 4-6 mm long, densely short- hairy; pappus white, barbellate. Meadows and foothills in sagebrush and aspen commu- nities at 1675 to 2100 m in Cache and Rich counties; Alberta and British Columbia to Wyoming, Nevada, and California; 2 (0). Artemisia L. Annual, biennial, or perennial herbs, sub- shrubs, or shrubs from taproots, caudices, or rhizomes, the juice watery; stems decumbent to ascending or erect, simple or branched; leaves alternate or basal, entire or toothed, lobed, or divided; heads several to numerous, borne in spicate, racemose, or paniculate clusters; involucral bracts imbricate in sever- al series, dry, at least the inner with scarious margins; receptacle naked or beset with long hairs, often glandular; corollas of disk flowers only (rarely with minute bilabiate ray flowers in A. bigelovii), perfect, or sometimes the central ones sterile, the marginal merely pis- tillate; marginal corollas tubular (or bila- biate), the central ones tubular- funnelform; pappus lacking, or a short crown; style branches flattened; achenes subterete or an- gular, glabrous. Beetle, A. A. 1960. A study of sagebrush — The section Tridentate of Artemisia. Univ. Wyoming Agr. Expt. Sta. Bull. 368. 83 pp. Keck, D. D. 1946. A revision of the Artemisia vulgaris complex in North America. Proc. Calif. Acad. 25:421-468. 1. Plants shrubs or subshrubs 2 — Plants herbs 13 2(1). Heads with both ray and disk flowers, the ray flowers 2-lipped; branchlets of inflorescence spreading to reflexed; plants of rimrock areas in Colorado drain- age, rarely in southern Great Basin A. bigelovii — Heads discoid; branchlets of inflorescence variously disposed; plants seldom of rimrock, the distribution various 3 3(2). Leaves 1 to 3-pinnately or ternately dissected, the segments linear 4 — Leaves entire or toothed, or if lobed, the lobes oblong or broader, or if linear (see A. filifolia), tall shrubs of sandy areas at low elevations 6 4(3). Plants silvery-canescent; receptacle hairy; growing commonly on windswept ridges, but not always so restricted A. frigida — Plants green to gray-green; receptacle glabrous, or, if hairy, plants of low ele- vations 5 5(4). Shrubs with spreading branches, spinescent, flowering in springtime .... A. spinescens — Shrubs with erect or ascending branches, not spinescent, flowering in late sum- mer and autumn A. pygmaea 6(3). Leaves linear-filiform, less than 1 mm wide, entire, or 3-parted; tall plants of sandy low elevation sites A. filifolia — Leaves broader, entire, or the segments broader than 1 mm wide; plants of var- ious habitats and elevations 7 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 207 7(6). Leaves entire or with 1 or 2 teeth; heads borne in slender panicles; plants of high elevations A. cana — Leaves toothed or lobed at the apex; heads borne in slender spicate to broad panicles 8 8(7). Plants usually less than 3 dm tall; leaves usually less than 1 cm long; foliage dull yellow- to lead-gray or rarely silvery A. nova — Plants mainly more than 3 dm tall; leaves usually more than 1 cm long (at least some); foliage silvery-canescent 9 9(8). Leaves all, or many of them, deeply cleft into narrowly oblong lobes which may be further divided; flowers commonly 5-8 per head A. tripartita — Leaves mainly merely toothed apically; flowers various 10 10(9). Leaves coarsely and deeply 3-lobed, the lobes broad and rounded apically; in- florescence narrow, seldom over 1.5 cm wide; plants of Rich and Summit counties A. longiloba — Leaves variously 3- to 5-toothed, seldom lobed; inflorescence various; plants variously distributed 11 11(10). Inflorescence open, paniculate, commonly more than 2 cm wide; plants of broad distribution, our common sagebrush species A. tridentata — Inflorescence narrow, spicate, commonly less than 1.5 cm wide; plants less broadly distributed 12 12(11). Plants often less than 4 dm tall; leaves mainly less than 1.5 cm long; heads small; plants usually of middle elevations A. arbuscula — Plants often over 4 dm tall; leaves mostly over 2 cm long; heads large; plants of high elevations A. spiciformis 13(1). Leaves all entire, or the lower ones toothed or lobed, glabrous and green above and beneath, or white-hairy on both surfaces (see also A. carruthii and A. mich- auxiana), usually much longer than broad 14 — Leaves deeply incised, pinnatifid, or temately divided, variously pubescent, various in length-width proportions 15 14(13 ) Leaves green above and beneath; central flowers of heads with normal ovaries A. dracunculus — Leaves white-hairy above and beneath or green above; central flowers of head with abortive ovaries A. ludoviciana 15(14). Plants annual or biennial from a taproot; leaves green, essentially glabrous; ad- ventive 16 — Plants perennial from a rhizome or caudex; leaves tomentose, strigose, or pilose ... 17 16(15). Inflorescence paniculate, loose and open; heads borne on short peduncles; in- volucres 1-2 mm high A. annua — Inflorescence a spicate panicle, the branches appressed-ascending; heads ses- sile or nearly so; involucres more than 2 mm high A. biennis 17(15). Cauline leaves reduced upwards, the largest leaves in a basal rosette, silvery- villous to strigulose, scarcely tomentose and uniformly colored above and be- neath; plants from caudices, only occasionally rhizomatous 18 — Cauline leaves not especially reduced upwards, seldom with a basal rosette, variously tomentose and often bicolored; plants often rhizomatous (except in A. absinthium) 20 208 Great Basin Naturalist Vol. 43, No. 2 18(17). Pubescence of leaves loosely villous to glabrous; corollas hairy, the receptacle glabrous; plants of high elevations in the Uinta and La Sal mountains 19 — Pubescence of leaves appressed strigose or villosulose; corollas glabrous or hairy, but, if hairy, the receptacle long- villous; plants variously distributed 20 19(18). Involucres 3-4 mm high; plants of the La Sal Mountains A. parryi — Involucres 4-5.3 mm high; plants of the Uinta Mountains A. norvegica 20(19). Inflorescence a spicate raceme; receptacle and corolla long-villous; plants of high elevations A. scopulorum — Inflorescence a slender panicle; receptacle and corollas glabrous; plants of low elevations, seldom of high elevations A. campestris 21(17). Receptacle beset with numerous long hairs between the flowers; leaves about equally hairy above as below; plants introduced, weedy, of low elevations A. absinthium — Receptacle naked; leaves more or less tomentose below, usually green or greenish above, or equally tomentose on both sides; plants indigenous, not weedy, of mid-to-high elevations 22 22(21). Leaves entire or with entire lobes; plants of moderate elevations in central and southern Utah A. carruthii — Leaves bipinnatifid, the lobes again toothed; plants of high elevations in the Uinta, Wasatch, and La Sal mountains A. michauxiana Artemisia absinthium L. Absinthe. Pe- rennial fragrant herbs from a rhizomatous caudex, 5-10 (12) dm tall, appressed sericeus; leaves bi- or tripinnatifid, the main lobes again lobed or toothed, silvery-sericeus on both surfaces, with very short tangled hairs, hardly tomentose, 1.5-5.5 cm long on flower- ing stems (2-10 cm long on sterile stems) the main ultimate segments mostly 2-4 mm wide, petiolate below, shortly petiolate and less commonly divided above; involucres 2-3 mm high, the bracts scarious over the green- ish center, the margins brownish hyaline; flowers all fertile, the marginal ones pistil- late; receptacles with numerous long slender hairs; achenes glabrous. Roadsides, stream- banks, and abandoned fields in Garfield, Rich, and Utah counties; widely established in North America; adventive from Europe; 6 (ii). Artemisia annua L. Sweet Wormwood. Annual fragrant herbs, mainly 0.3-1.5 (3) m tall; stems sparingly glandular; leaves 2- or 3- pinnatifid, the main lobes again lobed, green and minutely glandular on both surfaces, 1.5-8 (10) cm long, the main ultimate seg- ments 0.5-3 mm wide, petiolate below, sub- sessile or shortly petiolate above; involucres 1.3-2 mm high; involucral bracts with green centers minutely glandular, the margins hya- line; receptacles naked; achenes glabrous. In- troduced weedy species of disturbed sites in Washington County; adventive from Eurasia, now widely naturalized in North America; 1 (0). Artemisia arbuscula Nutt. Low Sagebrush. [A. tridentata ssp. arbuscula (Nutt.) H. & C.; A. tridentata var. arbuscula (Nutt.) McMinn]. Shrubs, commonly 2-4 (5) dm tall, the vege- tative stems 1.5-10 cm long, the flowering stems erect, 8-30 cm long; leaves 0.4-1.6 cm long, shallowly 3- to 5-dentate to deeply lobed, cuneate basally, appressed canescent; inflorescence spicate, mostly less than 2 cm wide; involucres 4-6 mm long, campanulate; involucral bracts 4-8, canescent, the margins brownish-scarious; flowers 4-9, all perfect; receptacle naked; achenes glabrous. Pinyon- juniper, mountain brush, sagebrush, white fir, aspen, and spruce-fir communities at 1375 to 2550 m in Box Elder, Cache, Millard, Rich, Salt Lake, Summit, and Tooele counties; Washington to Montana, south to California and Nevada; 14 (0). A. arbuscula, or low sagebrush, has been confused with both A. tridentata and A. nova. It can be distin- guished from the former by its narrow in- florescence, and from the latter by its canes- cent involucres. Beetle (Ic.) reports April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 209 intermediates with A. longiloha, a taxon with broadly campanulate heads and bluntly lobed leaves. Artemisia biennis Willd. Biennial Worm- wood. Plants annual or biennial, with tap- roots, the stems 0.3-9 (10) dm tall or more, glabrous; basal leaves often withered by an- thesis; cauline leaves well developed, 1.5-10 (15) cm long, once pinnately divided, the seg- ments oblong to oblanceolate, again toothed, essentially glabrous, green; inflorescence spi- cate or in spicate panicles; heads numerous, crowded, sessile or subsessile, erect or nearly so; involucres 2-3 mm high, 2-4 mm broad, the bracts glabrous, greenish to yellowish, the margins hyaline; marginal flowers perfect, fertile, the corollas glabrous; receptacle and achenes glabrous. Floodplains, lake beds and shores, mud flats, and pond margins at 1375 to 2900 m in Cache, Duchesne, Garfield, Grand, Iron, Millard, Salt Lake, Sanpete, Tooele, Uintah, and Utah counties; wide- spread in North America, where presumably indigenous in the western portion; Europe; 20 (v). Artemisia bigelovii Gray Bigelow Sage- brush. Shrubs, commonly 2-7 (10) dm tall or more, the vegetative stems 1-3 dm long, the flowering stems erect, 3-4.5 dm tall; leaves 0.3-2.3 cm long, 1-7 mm wide, entire or shallowly 3-toothed, basally cuneate, ap- pressed to loosely canescent-tomentose; in- florescence narrowly paniculate, mostly less than 4 cm wide, the branches often lax and with heads tending to be pendulous; in- volucres mainly 2.5-3.5 mm high, sub- cylindric to narrowly campanulate, the bracts 5-10, silvery canescent, with narrow scarious margins; flowers 3 or 4, imperfect or some perfect, the marginal pistillate (ray) flowers bilaterally symmetrical; receptacle naked; achenes glabrous. Rimrock areas in pinyon-juniper and mixed desert shrub com- munities at 975 to 2135 m in Duchesne, Emery, Garfield, Grand, Kane, Millard, San Juan, Sevier, Uintah, and Wayne counties; California and Nevada east to Colorado, New Mexico, and Texas; 45 (xvii). Artemisia campestris L. Field Wormwood. Perennial herbs from a caudex and taproot, the stems (1.5) 2.5-7 dm tall (rarely taller), tomentose or glabrous; basal leaves well-de- veloped (often withered at anthesis), 2-12 cm long, 2- to 3-pinnatifid or ternate, the seg- ments linear to narrowly oblong or spatulate, villous or pilose to glabrous on both sides; cauline leaves reduced upwards, once pin- natifid, ternate, or entire; inflorescence of narrow to lax panicles; heads numerous, shortly pedunculate on contracted to lax branchlets, finally pendulous; involucres 2.5-3.8 mm high, 2-2.3 mm wide, the bracts glabrous, greenish to yellowish, the margin hyaline; marginal flowers pistillate, fertile, the corollas glabrous; disk flowers sterile, the ovaries abortive; receptacle and achenes glabrous. Saltbush, greasewood, sagebrush, mountain brush, and pinyon-juniper commu- nities, mainly in dunes and other sandy sites at 1250 to 2075 m in Emery, Garfield, Grand, Kane, San Juan, Sevier, Washington, and Wayne counties; Arizona, New Mexico, Colorado, Wyoming, and west to the Pacific; 24 (vii). Our material is assignable to ssp. ho- realis (Pallas) H. & C, in a broad sense, and belongs to var. scouleriana (Benth.) Cronq. [A. pacifica Nutt.; A. campestris ssp. pacifica (Nutt.) H. & C; A. forwoodii authors, not Wats.; A. caudata authors, not Michx.] in a more narrow sense. Artemisia cana Pursh Silver Sagebrush. Shrubs, commonly 2.5-12 (15) dm tall, the vegetative branches 1-3 (5) dm long; flower- ing stems erect, 1-3 dm tall; leaves 0.8-5.3 (7) cm long, linear to narrowly elliptic or ob- long, entire, or some of them toothed or deeply lobed, usually acute basally, acute to obtuse apically, appressed tomentose; in- florescence narrowly spicate or glomerate- paniculate, mostly less than 5 cm wide, often conspicuously bracteate, the branches, when present, erect, the heads erect; involucres 3.3-6.1 mm high, 3.5-6 mm wide, cam- panulate; bracts numerous, the outer silvery- canescent, with greenish median, the margins brownish scarious, rounded-erose; flowers 10-20, perfect; receptacle naked; achenes glabrous. Meadows and stream terraces, less commonly on moist slopes away from mead- ows and streams at 2270 to 3050 m in Cache, Carbon, Daggett, Duchesne, Emery, Gar- field, Iron, Juab, Kane, Piute, Rich, Sanpete, Sevier, Summit, Utah, Wasatch, and Wash- ington counties; British Columbia to Sas- katchewan, south to California, Nevada, and New Mexico. Our materials are assigned to 210 Great Basin Naturalist Vol. 43, No. 2 var. viscidula Osterh. [A. cana ssp. viscidula (Osterh.) Beetle], which differs from typical var. cana in its smaller, narrower leaves and less canescent herbage. Silver sagebrush forms intermediates with both A. tridentata var. vaseyana and A. spiciformis, within whose altitudinal range it occurs, but whose habitats are ordinarily separate; 42 (viii). Artemisia carruthii Wood ex Carruth Car- ruth Wormwood. [A. wrightii Gray; A. vul- garis ssp. wrightii (Gray) H. & C.]. Plants pe- rennial herbs, with well-developed rhizomes, the stems 2-7 dm tall, sparingly to densely tomentose; basal leaves not well developed; cauline leaves various but usually pinnatifid with linear lobes, those of innovations and sometimes the primary ones at base of flow- ering stems entire or merely lobed, 0.6-3 cm long, the lobes 0.5-1.5 (2) mm wide, linear or narrowly oblong, tomentose on both sides, or less so above; inflorescence paniculate (nar- rowly so) or spicate; heads numerous, shortly pedunculate to sessile, erect; involucres 2.3-3 mm high, 2-2.5 mm wide, the bracts spar- ingly tomentose, pale greenish with hyaline margins; marginal flowers pistillate, fertile; central flowers perfect, fertile, the corollas glabrous; receptacle and achenes glabrous. Canyon bottoms, slopes, and rock outcrops in sagebrush, mountain brush, aspen, and spruce- fir communities at 1890 to 3050 m in Emery, Iron, Piute, San Juan, Utah, and Washington counties, and likely elsewhere; east to Kansas and south to Arizona, New Mexico, and Texas. This taxon is allied to A. ludoviciana, and some specimens appear to be intermediate between them. There is justi- fication for inclusion of A. carruthii within an expanded A. ludoviciana, but no formal pro- posal is intended or implied herein. The deeply pinnatisect main foliage leaves are thought to be diagnostic. The species has not been collected in sufficient numbers as to un- derstand its distribution in any definitive manner; 16 (ii). Artemisia dracunculus L. Terragon. [A. glauca Pallas; A. dracunculus ssp. glauca (Pallas) H. & C; A. aromatica A. Nels.; A. dracunculoides Pursh]. Plants shortly rhizo- matous, perermial herbs, the stems (2) 5-12 (15) dm tall, glabrous (rarely tomentose?); leaves primarily cauline, entire or rarely a few of them cleft, 1.2-7.5 cm long, 1-6 mm wide, glabrous, green on both surfaces; in- florescence paniculate; heads numerous, short-pedunculate to subsessile, more or less pendulous; involucres 2-2.8 mm high, 2.2-3 mm wide, the bracts glabrous, greenish, with broad hyaline margins; marginal flowers pis- tillate, fertile; central flowers sterile, the ovaries abortive, the corolla glabrous (often glandular); receptacle and achenes glabrous. Rabbitbrush, sagebrush, skunkbush, wildrye, salt desert shrub, pinyon-juniper, ponderosa pine, aspen, spruce-fir, and hanging garden communities at 1220 to 3200 m in Duchesne, Emery, Garfield, Grand, Iron, Juab, Kane, Millard, Piute, San Juan, Sanpete, Sevier, Tooele, Uintah, Utah, Wasatch, Washington, and Wayne counties; Yukon southeast to Il- linois and south to Mexico. Our material fits within the concept of var. glauca (Pallas) Bess, in Hook., which is probably not sepa- rable from var. dracunculus of the Old World; 63 (xvii). Artemisia filifolia Torr. Sand Sagebrush; Old-man Sagebrush. Shrubs commonly 5-15 dm tall, the vegetative branches 1-3 dm long; flowering branches erect, 1.5-6 dm long; leaves 0.6-8 cm long, 0.3-1.5 mm wide, revolute (appearing terete) or somewhat flat- tened (and still revolute), entire or the lower ternate, appressed villous-tomentose; in- florescence paniculate, mostly more than 3 cm wide, conspicuously bracteate, the branches erect, the heads pendulous; in- volucres 1.6-2.2 mm long 1.5-2.2 mm wide, campanulate to subglobose; bracts 5-9, densely silvery canescent; flowers 3-9, the marginal ones pistillate, fertile, the central ones sterile; receptacle naked; achenes glabrous. Sandy sites in blackbrush, creosote bush, ephedra, Poliomintha, Eriogonum, rab- bitbrush, and pinyon-juniper communities at 825 to 2290 m in Emery, Garfield, Grand, Iron, Kane, San Juan, Washington, and Wayne counties; Colorado and South Da- kota, south to Arizona, Texas, and Mexico; 43 (vi). Artemisia frigida Willd. Fringed Sage- brush; Prairie Sagewort. Shrubs 0.5-4.5 dm tall, white-tomentose to strigulose; flowering stems arising from short prostrate or ascend- ing woody offsets; leaves of basal offsets April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 211 much like the stem leaves, 0.5-1.5 (2.5) cm long, 2- to 3-temately (or subpinnately) di- vided into linear segments mainly 0.3-0.8 mm wide, often with stipulelike divisions near the base, whitish pilose-tomentose throughout (fading brownish); inflorescence paniculate or less commonly borne sessile or on very short pedimcles; involucres 2-3.5 mm high, 4-6 mm broad, the bracts pilose-to- mentose, with brownish scarious margins; marginal flowers pistillate, fertile; central flowers perfect, fertile, the corolla glabrous (often glandular), yellow or tinged reddish; receptacle long-hairy; achenes glabrous. Shadscale, sagebrush, pinyon-juniper, ponde- rosa pine, mountain brush, aspen, spruce, and alpine (often on windswept ridge crests) com- munities at 900 to 3480 m in Box Elder, Car- bon, Duchesne, Emery, Garfield, Grand, Juab, Kane, Millard, Piute, San Juan, San- pete, Sevier, Summit, Uintah, Utah, Wasatch, and Wayne counties (likely elsewhere); Alaska to Quebec, south to Arizona and Kan- sas; Asia; 78 (x). Artemisia longiloba (Osterh.) Beetle Longleaf Sagebrush. [A. spiciformis (?) long- iloba Osterh.]. Shrubs, mainly 2-5 dm tall, appressed villous-tomentose; flowering stems 1-2 dm long; leaves 0.4-2 cm long, broadly cuneate, deeply 3-lobed, the lobes obtuse, ap- pressed villous-tomentose; inflorescence spi- cate, the heads several, shortly pedunculate to sessile, erect; involucres 4-6 mm high, 3-5 mm wide, the 4-12 bracts villous-tomentose, green, with brownish scarious margins; mar- ginal flowers perfect, fertile, central flowers perfect, fertile, the corolla glabrous (glandu- lar), cream colored; receptacle and achenes glabrous. Sagebrush and grass communities at 1675 to 2440 m in Rich and Summit counties; Oregon to Montana, south to Nevada and Colorado. This entity is reported to grow in tight to heavy soils (Beetle 1960) and matures seed in July and August. The plants have large heads similar to those of A. cana and the low habit of A. nova. Possibly they would best be treated within an expanded A. triden- tata, but no combination is proposed herein; 4 (0). Artemisia ludoviciana Nutt. Perennial rhizomatous herbs, the stems 2-10 dm tall (or white-tomentose or glabrate to more glabrous; leaves mainly cauline, entire, lobed, or pinnately incised, white-tomentose below, green and glabrous or tomentose above (rarely glabrous throughout), 0.8-9 cm long, 0.1-1 (2) cm wide; inflorescence spicate to paniculate; heads numerous, shortly pe- dunculate to sessile, more or less pendulous; involucres 2.5-4.5 mm high, 3-7 mm wide (or more), the bracts tomentose to glabrous, with broad scarious margins; marginal flow- ers pistillate, fertile; central flowers perfect, fertile, the corolla glabrous, yellow; recep- tacle and achenes glabrous. This is a wide spread species of many phases and habitats. In Utah there are five more or less distinctive varieties. Two of the varieties, ludoviciana and incompta are especially abundant, the re- maining three less so. Not all specimens are readily separable into the named varieties, and the following key is arbitrary. 1. Inflorescence an open panicle, often more than 8 cm wide; plants of southern and southeastern Utah 2 — Inflorescence a spicate panicle, usually less than 6 cm wide; plants of various distribution 3 2(1). Leaves mainly less than 2.5 cm long, the margin often narrowly revolute A. ludoviciana var. alhula — Leaves mainly over 2 cm long, the margins not revolute A. ludoviciana var. mexicana 3(1). Leaves entire or less commonly some of them toothed or lobed A. ludoviciana var. ludoviciana — Leaves more or less deeply parted or divided 4 4(3). Involucres 3.5-4.2 mm high, 4-7 mm wide A. ludoviciana var. latiloba — Involucres 2.5-3.5 (3.8) mm high, 2.5-5 mm wide A. ludoviciana var. incompta 212 Great Basin Naturalist Vol. 43, No. 2 Var. albula (Wooton) Shinners [A. albula Wooton, nomen novum pro A. microcephalu Wooton]. This distinctive short-leaved vari- ety has open inflorescences; it occurs in ri- parian areas with rabbitbrush, cottonwood, and copperweed at 880 to 1680 m in Emery, Garfield, Kane, San Juan, and Washington counties; Nevada and Colorado south to Mex- ico; 4 (ii). Var. incompta (Nutt.) Cronq. [A. incompta Nutt.; A. ludoviciana ssp. incompta (Nutt.) Keck]. The deeply lobed or cleft leaves and compactly spicate inflorescence are diagnos- tic. The plants occur at moderate to high ele- vations (2135 to 3500 m) in aspen, spruce-fir, willow-wet meadow, and riparian commu- nities in Cache, Carbon, Duchesne, Juab, Millard, Piute, Salt Lake, San Juan, Sanpete, Sevier, Summit, Tooele, Utah, and Wasatch counties; Oregon to Montana, south to Cali- fornia, Nevada, and Colorado; 65 (vi). This variety passes by degree into A. michauxiana at high elevations. Var. latiloba Nutt. [A. candicans Rydb.; A. ludoviciana ssp. candicans (Rydb.) Keck]. This variety is similar to var. incompta, differing in larger (higher and wider) heads. It is poorly understood in Utah, where it was reported by Keck (1946) from Cache County. Our one specimen is from Utah County, without locality data; 1 (0). Var. ludoviciana [A. gnaphaloides Nutt.; A. ludoviciana var. gnaphaloides (Nutt.) T. & G.; A. purshianus Bess, in Hook.]. The typi- cal variety is a plant with entire or cleft (rarely deeply cleft or parted) leaves and loose, but not open, inflorescences. Rabbit- brush, sagebrush-grass, mountain brush, pin- yon-juniper, ponderosa pine, and hanging garden communities at 880 to 2750 m in all Utah counties, except for Morgan, Piute, and Summit (and likely there also); British Co- lumbia to Ontario, south to California, Ari- zona, New Mexico, Texas, and Indiana; 86 (xvi). Var. mexicana (Willd.) Fern. [A. mexicana Willd.; A. ludoviciana ssp. mexicana (Willd.) Keck]. This is the long-leaved plant with open inflorescences; it forms the counterpart of var. albula. It is a component of riparian, pinyon-juniper, ponderosa pine, and aspen communities at 750 to 2600 m in Garfield, Grand, Kane, San Juan, Washington, and Wayne counties; Colorado to Missouri, south to Mexico; 17 (viii). Artemisia michauxiana Bess. Michaux Wormwood. [A. discolor Dougl. ex Bess.]. Pe- rennial herbs, the stems 0.8-4 dm tall (rarely more), white tomentose to glabrate or glabrous; leaves mainly cauline, 0.5-4 (5) cm long and about as broad, bipinnately dis- sected, the secondary segments again toothed or lobed, acute, the uppermost seldom entire, commonly green above and tomentose be- neath, but often green beneath also; in- florescence spicate; heads several to numer- ous, commonly pedicellate, erect or nodding; involucres 3.4-4.4 mm high, 3-6 mm wide, the bracts glabrous or sparingly tomentose, green, the broad margins brownish scarious and erose-ciliate; marginal flowers pistillate, fertile; central flowers perfect, fertile, the co- rolla glabrous (glandular), yellow; receptacle and achenes glabrous. Spruce-lodgepole pine and alpine tundra communities, often in boulder stripes and talus, at 2950 to 3500 m in Duchesne, San Juan, Summit, and Utah counties (Uinta, La Sal, and Wasatch moun- tains); British Columbia and Alberta south to Nevada and Wyoming; 11 (ii). Keck (1946) notes that A. michauxiana is connected through a series of intermediates with A. lu- doviciana var. incompta in Nevada speci- mens. This is true for ours also. There ap- pears to be some justification for treating A. michauxiana within an enlarged A. ludovi- ciana, but such a combination is not implied herein. Artemisia norvegica Fries Spruce Worm- wood. Perennial herbs, 2-4.1 dm tall, from a simple or branched caudex and stout taproot, the caudex branches short, clothed with per- sistent leaf bases, the flowering stems arising directly from the caudex, villous, often red- dish; leaves of basal rosettes 2-19 cm long, bi- or tripinnatifid, the segments lance-atten- uate to acute, villous on both surfaces; cau- line leaves becoming smaller upwards, often with stipulelike divisions near the base; in- florescence racemose; heads several to nu- merous, finally nodding, the peduncles to 4.5 cm long; involucres 4-5.3 mm high, 6-11 mm wide, the bracts sparingly to densely vil- lous-pilose, more or less green, the margins broadly dark brownish scarious; marginal April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 213 flowers pistillate, fertile; central flowers per- fect, fertile, the corollas long-hairy from near the base, cream colored; receptacle and ach- enes glabrous. Spruce-fir, lodgepole pine, and alpine tundra communities in Duchesne and Summit counties; Alaska east to Mackenzie, and south to California and Colorado. Our material belongs to var. piceetorum Welsh & Goodrich in Welsh; 4 (iii). Artemisia nova A. Nels. Black Sagebrush. [A. tridentata ssp. nova (A. Nels.) H. & C; A. tridentata var. nova (A, Nels.) McMinn; A. arbuscula ssp. nova (A. Nels.) Ward]. Shrubs, 1-3 (5) dm tall, the main branches spreading, the vegetative stems 1-3 dm long (rarely more); flowering stems mainly 1.5-3 (4) dm long; leaves 0.3-2.1 cm long, shallowly to deeply 3- to 5-lobed or -toothed, the lobes or teeth rounded, often lead-gray or gray green, cuneate basally, appressed canescent and of- ten minutely punctate; inflorescence narrow- ly paniculate, seldom more than 3 cm wide; involucres 3.1-5.8 mm long, 1.4-3.4 mm wide, cylindric to narrowly campanulate; bracts 8-12, canescent to glabrous, green to yellowish, the margin hyaline; flowers 3-8, all perfect; receptacle glabrous; achenes glabrous. Horsebrush, greasewood, shadscale, ephedra, juniper, sagebrush, rabbitbrush, winterfat, pinyon-juniper, and mountain brush communities at 1400 to 2600 m in Bea- ver, Box Elder, Cache, Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Iron, Juab, Kane, Millard, Piute, Rich, San Juan, Sanpete, Sevier, Summit, Tooele, Uintah, and Weber counties; Oregon to Montana, south to California, Arizona, and New Mexico; 57 (x). Black sagebrush forms intermediates with all other members of the section Tridentatae that it contacts. The intermediates form nar- row bands along lines of contact, but general- ly the habitats are mutually exclusive. There is little justification for considering black sagebrush in an expanded A. tridentata unless one is willing to accept most of the remain- der of the section as portions of that species also. Artemisia parryi Gray Parry Wormwood. Perennial herbs, 0.8-2 (4) dm tall, from a simple or branched caudex and stout taproot, the caudex branches short, clothed with per- sistent leaf bases, the flowering stems arising directly from the caudex, sparingly and loose- ly villous to glabrous, often reddish; leaves of basal rosettes 2-4 (8) cm long, twice pinnati- fid, the segments oblong to lance-oblong, sparingly and loosely villous (to glabrous?); cauline leaves becoming smaller upwards; in- florescence racemose to subspicate; heads several to numerous, commonly nodding, the peduncles 1-5 mm long; involucres 3-4 mm long, 3-5 mm wide, the bracts sparingly vil- lous to glabrate, with green to brownish middle and brownish scarious margins; mar- ginal flowers pistillate, fertile; central flowers perfect, fertile, the corollas long-hairy (to glabrous?); receptacle and achenes glabrous. Alpine sites in the La Sal Mountains (Grand and San Juan counties); Colorado; 0 (0). The species is reported for Utah by Hall and Cle- ments (I.e.), but no specimens have been seen from the state by me. Possibly it is only a phase of A. norvegica. Artemisia pygmaea Gray Pygmy Sage- brush. Shrubs 0.5-2 dm tall, from superficial woody caudexlike branches and stout tap- roots, the vegetative stems to 0.5 dm long; flowering stems erect to 2 dm tall; leaves 0.3-1 cm long, pinnately (or subbipinnately) 3- to 10-lobed, the lobes acute, yellow- to gray-green, sparingly villous to glabrous; in- florescence spicate or narrowly paniculate, less than 2 cm wide; involucres 5.2-6.3 mm high, 3-4.5 mm wide, cylindric or becoming campanulate upon drying; involucral bracts oblong, 15 or more, sparingly villous to glabrous, green, the margins stramineous- hyaline; marginal flowers lacking; central flowers 3-5, perfect, fertile, the corollas cream colored, glandular; receptacle and achenes glabrous. Black sagebrush, rabbit- brush, shadscale, greasebush, juniper, pinyon- juniper, and ponderosa pine communities at 1600 to 2300 m in Beaver, Duchesne, Emery, Garfield, Iron, Millard, Piute, Sevier, and Uintah counties (likely elsewhere); Arizona and Nevada; 27 (viii). This dwarf sagebrush occurs in peculiar edaphic situations on Green River Shale, in clay soils forming the matrix in igneous gravels, on calcareous gravels, and on dolomitic outcrops and gravels. It is often a component of commu- nities that support rare plant species. Artemisia scopulorum Gray Dwarf Sage- wort. Perennial herbs, 0.5-3.7 dm tall, from a 214 Great Basin Naturalist Vol. 43, No. 2 simple or branched caudex and stout taproot, the caudex branches short, clothed with per- sistent leaf bases, the flowering stems arising directly from the caudex, appressed pilose to loosely and sparingly villous, often reddish or purplish; leaves of basal rosettes 1.5-9 cm long, twice pinnatifid, the segments oblong to elliptic, pubescent like the stems; in- florescence spicate to racemose; heads sever- al to numerous, erect or nodding, the pe- duncles lacking, or to 2.3 cm long; involucres 3-5.2 mm high, 3-8 mm wide, the bracts vil- lous, green to brownish in the middle, the margins brown-scarious; marginal flowers pistillate, fertile; central flowers perfect, fer- tile, the corollas cream colored, long-hairy; receptacle copiously long-villous; achenes glabrous. Talus slopes, moraines, and outwash plains and terraces in alpine tundra and meadows in spruce, lodgepole pine, and Douglas fir communities at 3050 to 4000 m in Boulder, Tushar, La Sal, and Uinta moun- tains; Beaver, Duchesne, Garfield, Grand, Piute, Summit, and Uintah counties; Mon- tana, Wyoming, Colorado, and New Mexico; 30 (xi). The hairy corollas and long-villous re- ceptacles are diagnostic for this distinctive species. Artemisia spiciformis Osterh. Osterhout Sagebrush. Shrubs, mainly 5-8 dm tall, the vegetative stems 0.4-1 dm long, the flower- ing stems erect, 1.5-3.4 dm long; leaves 1.7-5.7 cm long, shallowly to deeply 3- to 5- lobed or -toothed, often widest below the teeth, the lobes acute to obtuse (or rounded) or lacking, gray-green, long-cuneate basally, appressed villous-canescent; inflorescences narrowly paniculate, usually less than 4 cm wide; involucres 5-6.3 mm long, 3.5-7 mm wide, cylindric to campanulate; involucral bracts 8-12 or more, canescent to glabrate, green, with broad yellowish brown scarious margins; flowers 6-10 or more, all perfect; receptacle and achenes glabrous. Ridge mar- gins and snow-flushes in sagebrush-grass, snowberry, aspen, spruce-fir, and Douglas fir commimities at 2680 to 3050 m in Cache (?), Duchesne, Emery, Juab, Sanpete, Summit, Tooele, and Wasatch counties (likely else- where); Colorado and Wyoming; 16 (ii). This is the material which has long passed under the name of A. rothrockii Gray in Utah. Resemblance to that species appears to be superficial, with relationships running to both A. cana and A. tridentata var. vaseyana. Its habitat is intermediate between the high ele- vation, moderately xeric conditions of var. vaseyana, and the more mesic stream terrace and valley bottoms of A. cana. Artemisia spinescens D.C. Eaton in Wats. Budsage. Shrubs, flowering in springtime, the branches spreading and often prostrate, 0.5-3 dm long or more, the vegetative stems mainly 0.3-0.8 dm long, commonly surpass- ing the flowering stems; leaves 0.4-2 cm long, petiolate, the blade palmately 3- to 5- cleft, the main divisions again cleft, sub- orbicular in outline, villous; inflorescence of short leafy-bracted racemose or spicate branches, or of solitary heads, the rachis per- sistent as a thorn; involucres 2-3.5 mm high, 3.5-5 mm wide; involucral bracts 4-8, vil- lous, green, with narrow hyaline margins; flowers 6-20 or more, the marginal ones pis- tillate, fertile, the central ones sterile; co- rollas copiously long-hairy; receptacle naked; achenes long-hairy. Silty, clayey, or gravelly, often saline, substrates in black sagebrush, shadscale, tetradymia, greasewood, black- brush, juniper, and winterfat communities at 1200 to 1925 m in Carbon, Duchesne, Emery, Garfield, Juab, Kane, Millard, Piute, San Juan, Sevier, Tooele, Uintah, and Utah counties; Oregon to Montana, south to Cali- fornia and New Mexico; 92 (vii). This low shrub is a principal browse plant for domes- tic livestock on the spring ranges of western and southern Utah. Artemisia tridentata Nutt. Big or Com- mon Sagebrush. Shrubs 4-20 (30) dm tall; branches spreading to erect, the vegetative branchlets 0.5-2 dm long; flowering stems mostly 1.5-4 dm long, usually much surpass- ing the vegetative ones; leaves 0.5-5 cm long, 3- to 5-toothed apically, or the upper ones entire, long-cuneate; inflorescence pan- iculate, 3-20 (15) cm wide; involucres 3-5 mm long, 2-4 mm wide, the bracts 10-20, green, canescent, the margins scarious; flow- ers 3-8, all perfect, the corollas cream col- ored, glandular; receptacle and achenes glabrous. Three more or less completely in- tergrading varieties are known from Utah; they tend to occupy distinctive habitats, but April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 215 intermediates form wherever they meet. Fur- ther, this taxon is known to hybridize with most if not all other members of the section Tridentatae. The help of Durrant MacArthur and Sherel Goodrich of the U.S. Forest Ser- vice is here gratefully acknowledged. They sorted our materials into their respective va- rieties following my initial attempt and gen- eral failure. While it is not possible to segre- gate all specimens, the following key will prove useful to those who must manage the sagebrush lands of Utah and the west. 2(1). Vegetative stems short, standing at about the same height, the inflorescence rather uniformly overtopping them; plants of middle and higher elevations A. tridentata var. vaseyana Vegetative stems short to long, the inflorescence not uniformly overtopping them; plants of low to moderate elevations 2 Leaves mainly to 2 cm long or more, narrowly cuneate; plants of low to mod- erate elevations A. tridentata var. tridentata Leaves mainly less than 1.2 cm long, cuneate to cuneate-flabellate; plants mainly of moderate elevations, in drier sites A. tridentata var. wyomingensis Var. tridentata Big Sagebrush. Sagebrush, juniper, pinyon-juniper, and rabbitbrush communities at 1220 to 2410 m in most, if not all, Utah counties; Washington to Mon- tana, south to California, Arizona, and New Mexico; 57 (xviii). Var. vaseyana (Rydb.) B. Boi. Vasey Sage- brush. [A. tridentata ssp. vaseyana (Rydb.) Beetle]. Sagebrush, rabbitbrush, mountain brush, pinyon-juniper, aspen, Douglas fir, ponderosa pine, and spruce-fir communities at 1830 to 3050 m in all, or nearly all, Utah coimties; Idaho to the Dakotas, south to Col- orado; 55 (viii). Var. wyomingensis (Beetle & Young) Welsh Stat. nov. [based on Artemisia triden- tata ssp. wyomingensis Beetle & Young Rho- dora 67: 405. 1965]. Wyoming Sagebrush. Shadscale, rabbitbrush, sagebrush, juniper, bitterbrush, and mountain mahogany com- mimities at 1525 to 1980 m in Box Elder, Garfield, Emery, Rich, Tooele, and Uintah counties; Wyoming and Idaho to Colorado. This is the sagebrush of drier sites at middle elevations. Its distribution is poorly under- stood; likely it is widespread. Its recognition allows management considerations by profes- sionals in the various state and federal agencies; 9 (0). Artemisia tripartita Rydb. Threetip Sage- brush. Shrubs 2-20 dm tall, the branches erect, the vegetative ones 0.3-1.5 dm long, the flowering stems 0.6-3.5 dm long; leaves 1-4 cm long, deeply 3-cleft, the linear lobes 0.5-0.8 mm wide, canescent, the lobes some- times again divided, or the upper ones entire; inflorescence paniculate, commonly 2-5 cm wide; involucres campanulate, 3-4 mm long, 1.5-4 mm wide; bracts many, imbricate, ca- nescent and more or less green, the inner with broad brownish scarious margins; flow- ers 4-8, all perfect, the corollas stramineus to cream-colored, more or less glandular; ach enes and receptacle glabrous. Sagebrush and mountain brush communities at ca 1525 to 1830 m in Box Elder and Cache counties; British Columbia to Montana, south to Cali- fornia and Colorado; 1 (0). Aster L. Annual or perennial herbs from rhizomes (suffrutescent in A. spinosus), with watery juice; stems decumbent to ascending or erect, simple or branched; leaves alternate, simple, entire or toothed; heads solitary or few to several in corymbose clusters; involucral bracts strongly imbricate to subequal (or the outer surpassing the inner), herbaceous throughout, or with scarious margins near the base; receptacle flat or merely convex, naked; rays blue, purple, pink, or white, few to numerous, pistillate; disk flowers numer- ous, perfect, fertile, yellow or tinged reddish or purplish; pappus of capillary bristles; style branches flattened, oblong to lanceolate, mostly more than 0.5 mm long; achenes mostly several nerved. 216 Great Basin Naturalist Vol. 43, No. 2 1. Plants suffrutescent, rushlike, armed with axillary or subaxillary thorns, from a deep-seated rhizome; known from Garfield County (possibly Washington also) A. spinosus — Plants herbaceous, annual or perennial, unarmed, from a taproot or rhizome; distri- bution various 2 2(1). Plants annual, from taproots 3 — Plants perennial, from rhizomes or subrhizomatous caudices, or from branch- ing caudices 4 3(2). Involucral bracts definitely acute; rays wanting or nearly so, the pistillate corollas tubular, shorter than the style A. brachyactis — Involucral bracts obtuse to obtusish; rays to 2 mm long, longer than the style ... A. frondosus 4(2). Plants with a well developed caudex; involucral bracts reflexed, at least the outer; plants of rock crevices in the Wasatch and Canyon mountains A. kingii — Plants with caudex lacking or poorly developed, rhizomatous; involucral bracts not reflexed; plants of various habitats and localities 5 5(4). Leaves all erect-ascending, thickened, to about 4 mm wide; pappus double, the outer series of very short bristles; heads solitary; plants known from Box Elder County A. scopiiloriim — Leaves various, seldom as above; pappus in one series, or rarely double; heads solitary to numerous; distribution various 6 6(5). Involucral bracts dry, chartaceous, with scarious tips (at least the innermost), with a distinctive midvein, not herbaceous (the outer sometimes so) 7 — Involucral bracts herbaceous at the tips or throughout, lacking a distinctive midvein 9 7(6). Involucral bracts (at least the outer) bluntly obtuse apically; herbage strongly glaucous; plants often of open calcareous sites A. glaucodes — Involucral bracts acute; herbage green, not glaucous; plants of various habitats 8 8(7). Rays white (drying pinkish); main leaves often over 20 mm wide; plants 6-15 dm tall, of montane areas in central northern Utah A. engelmannii — Rays purple or violet; leaves mainly less than 15 mm wide; plants 2-6 dm tall, of central northern and western Utah A. perelegans 9(5). Involucres and peduncles glandular 10 — Involucres and peduncles lacking glands or apparently so 13 10(9), Stems glabrous; leaves linear to linear-oblanceolate, 2-5 mm wide, 1.5-7 cm long; plants of saline or hot water seeps and springs A. pauciflorus — Stems puberulent to villous with multicellular hairs, or glabrous, but, if so, differing in other respects 11 11(10). Rays white; leaves glaucous; plants of central to south central Utah ..A. wasatchensis — Rays blue to purple, lavender, or violet 12 12(11). Cauline leaves clasping the stem, mainly 15-40 mm wide; involucres 8-15 mm high; plants of central northern Utah A. integrifolius — Cauline leaves not or only slightly clasping, 2-10 mm wide; involucres 5-8 mm high A. campestris April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 217 13(9). Pubescence occurring in decurrent lines below leaf bases, commonly not uni- form below the heads, or only in the inflorescence; inflorescence often conic, mostly large and leafy A. hesperius — Pubescence of stem uniform, or, if in decurrent lines, uniform below the heads and confined to the inflorescence; inflorescence few to many flowered and not usually leafy to large and leafy (see also A. eatonii) 14 14(13). Rays white; involucral bracts strigulose dorsally (rarely glabrous), with spread- ing to squarrose minutely spinulose tips; heads numerous 15 — Rays pink to purple, or less commonly white; involucral bracts mucronate at the tip; heads few to numerous 16 15(14). Rhizomes well developed, creeping; involucres 4.6-6.5 mm high, 7-9.5 mm wide (when pressed); plants of western Utah A. falcatus — Rhizomes mainly poorly developed, or reduced and caudexlike; involucres 3.8-4.9 mm high, 4.5-6 mm wide; plants of eastern Utah A. pansus 16(14). Achenes glabrous or nearly so; herbage glabrous except for lines of pu- berulence in the inflorescence, tending to be glaucous; rare plants in southeastern Utah A. laevis — Achenes pubescent, except in some A. foliaceus; herbage pubescent to almost glabrous, scarcely glaucous 17 17(16). Involucral bracts strongly imbricate, the outer ones at least obtuse or obtusish (sometimes acute), not foliaceous; pubescence below the heads harsh A. chilensis — Involucral bracts not strongly imbricate, or, if so, the bracts sharply acute, the outer ones acute, or, if obtuse, foliaceous; pubescence below heads soft or minute 18 18(17). Inflorescence a long slender leafy panicle; heads numerous; stem pubescence short, uniform; leaves mostly more than 7 times longer than wide; rays usually pink to white A. eatonii — Inflorescence an open or congested panicle; heads solitary to several; pubescence various; rays usually blue to violet 19 19(18). Involucral bracts slender, never foliaceous; leaves at midstem mostly less than 1 cm wide, mostly over 7 times longer than broad A. occidentalis — Involucral bracts various, but some of them usually enlarged and foliaceous; leaves at midstem mostly less than 7 times longer than broad A. foliaceus Aster brachyactis Blake in Tidestr. [Tripo- tubular filiform, lacking rays, much shorter Hum angustum Lindl. in Hook.; A. angustus than the styles; pappus abundant, white, (Lindl.) T. & G., not Nees; Brachyactis ang- longer than the corollas. Sandbars, terraces, ustus (Lindl.) Britt. in Britt. & Brown]. An- stream banks, marshes and pond margins, of- nual herbs, with taproots, glabrous through- ten where saline, in tamarix, rush, rabbit- out, except for leaf margins and involucral brush, and cottonwood commimities at 1220 bracts; stems 0.9-5.3 (7) dm tall; leaves 1.3-8 to 1525 m in Box Elder, Carbon, Duchesne, (12) cm long, 1-7 (9) mm wide, linear to nar- Emery, Grand, Garfield, Salt Lake, Uintah, rowly oblong, entire, the lower ones soon de- and Utah counties; British Columbia to Min- ciduous; heads few to numerous, in pan- nesota, south to Washington and Colorado; iculate to spicate inflorescences; involucres 14 (i). 5.5-9.4 (11) mm high, 7-15 (17) mm wide. Aster campestris Nutt. Meadow Aster. Pe- the bracts linear-oblong, acute to attenuate, rennial rhizomatous herbs, glandular, at least herbaceous, subequal to somewhat imbricate, in inflorescence; stems puberulent to or some outer ones often surpassing the in- glabrous, mainly 1-5 dm tall; leaves 2-8 cm ner; marginal flowers pistillate, the corollas long, 2-8 mm wide, linear to oblong, entire. 218 Great Basin Naturalist Vol. 43, No. 2 sessile, sometimes clasping, the lower ones larger and more or less petiolate, or smaller, soon deciduous; heads solitary or several to many; involucres 5-8 mm high, glandular, the bracts subequal to definitely imbricate, acute or attenuate, with long herbaceous tips; rays 15-20, violet to purple, 6-12 mm long. Meadows at 1525 to 2475 m, reported for Utah (Univ. Washington Publ. Biol. 17(5): 77. 1955), but I have seen no specimens from the state. Aster chilensis Nees Pacific Aster. Pe- rennial rhizomatous to subrhizomatous herbs, imiformly harshly strigose to strigulose, at least above; stems (0.8) 1.2-10.5 dm tall; leaves 0.6-16.5 cm long, 2-16 (20) mm wide, entire or nearly so, pubescent to glabrous, cil- iate, the lower ones more or less petiolate, of- ten deciduous at anthesis in taller plants, be- coming smaller and sessile upwards, sometimes markedly reduced-bracteate in in- florescence; inflorescence of 1 to many heads, narrow, corymbose, or open paniculate; in- volucres 5-8 mm high, 6-15 mm broad, the bracts imbricate, green tipped (machaeran- theroid), the chartaceous bases white to straw colored, the outer ones abruptly pointed but mucronate; rays commonly 15-40, purplish to violet (rarely white) or pink, 5-15 mm long; achenes pubescent. Alluvial fans, ter- races, and slopes along stream and canal banks, in hanging gardens, rabbitbmsh, sage- brush, grass-sedge, cottonwood-willow, pon- derosa pine, juniper-pinyon, mountain brush, aspen, and spruce-fir communities at 850 to 3200 m in all Utah counties; Washington to Saskatchewan, south to California and New Mexico; 189 (xxviii). The Pacific aster is a generalized taxon with no clearly diagnostic features. It is separated from its near con- geners by a group of intangible character- istics. Involucral bracts are definitely imbri- cate, with the greenish portion usually glabrous, and margins ciliate. The tips of out- er bracts are often but not always obtuse, and the tip, even when abruptly contracted is mucronate. These features, which I designate as "machaerantheroid," are shared to a great- er or lesser extent with A. eatonii, A. occiden- talis, and A. foliaceus. The harsh pubescence below the heads appears to be diagnostic, but is difficult to distinguish from the soft or merely puberulent vesture of closely related species. Not all specimens can be assigned with certainty to any of the taxa. There are two intergrading morphological phases of the Pacific aster, which are striking in their ex- tremes, but which probably represent noth- ing more than developmental gradients. There are plants with few flowers that lack distinctive reduced bracteate leaves in the in- florescence, and taller plants with more nu- merous heads and distinctively bracteate in- florescences. The inflorescences of the taller plants are mainly corymbiform, and not cy- lindroid as in A. eatonii. More work is in- dicated. Our material belongs to ssp. adscen- dens (Lindl.) Cronq. Aster eatonii (Gray) Howell Eaton Aster. [A. foliaceus var. eatonii Gray; A. oregonus authors, not (Nutt.) T. & G.]. Perennial rhizo- matous to subrhizomatous herbs, uniformly puberulent, at least above (below the heads and sometimes on upper leaves), the stems (2.7) 6-10.5 dm tall, often reddish; leaves 0.8-15 cm long, 2-25 mm wide, entire or ser- rate, puberulent to glabrous, ciliate, the lowermost shortly petiolate, often deciduous in anthesis, becoming smaller and sessile up- wards, linear to narrowly elliptic or lanceo- late to oblanceolate; inflorescence of few to numerous heads, commonly open-cylindric to conic in form; involucres 4.5-8 (10) mm high, 6-10 mm wide, the bracts more or less sub- equal to indistinctly imbricate, green tipped (but not especially machaerantheroid), the chartaceous bases white to straw colored, all or most of them mucronate; rays 20-40, com- monly pink (sometimes white), 5-12 mm long; achenes pubescent. Gravel bars, stream terraces, meadows, canal banks, hanging gar- dens, and marshes at 1370 to 2325 m in Cache, Garfield, Grand, Iron, Juab, Kane, Salt Lake, Summit, Uintah, Utah, Wasatch, and Washington counties; British Columbia to Saskatchewan, south to California, Ari- zona, and New Mexico; 48 (ix). The pink or white rays, uniform upper stem puberulence and leaves many times longer than broad are diagnostic for most specimens. Reports of A. junciformis Rydb. for Utah appear to be based on slender phases of the Eaton aster with linear leaves and slender rhizomes. Aster engelmannii (D.C. Eaton) Gray Engelmann Aster. [A. elegans var. engel- mannii D.C. Eaton]. Perennial rhizomatous April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 219 or subrhizomatous herbs, puberulent to spar- ingly villous with multicellular hairs, or somewhat glandular, the stems 2-15.2 dm tall, reddish at the base; leaves 2-13.5 cm long, 3-46 mm wide, elliptic to lanceolate, entire (or nearly so), sparingly puberulent to glabrous or sparsely villous, sessile, largest near midstem, the lowermost reduced to scales; inflorescence of 1 to numerous large heads, corymbose or conic; involucres 8-13 mm high, 11-25 mm wide, the bracts mainly strongly imbricate, with a definite midvein, commonly purplish (at least the inner), the outer sometimes green and more or less fo- liaceous, sometimes all greenish or straw col- ored to the tip, glabrous dorsally, ciliate; rays 8-23, white (drying pinkish), 12-25 mm long; achenes pubescent. Mountain brush, juniper, Douglas fir, aspen, white fir, lodgepole pine, and spruce-fir communities at 1950 to 3200 m in Cache, Carbon, Davis, Duchesne, Juab, Salt Lake, Sanpete, Summit, Utah, and Wasatch counties; British Columbia and Al- berta, south to Nevada and Colorado; 57 (vi). Aster falcatus Lindl. [A. miiltiflorus var. commutatus T. & C; A. commutatus (T. & G.) Gray]. Perennial rhizomatous herbs, vil- lous or villous-hirsute with multicellular hairs, the stems 2.8-7.5 dm tall; leaves 1.2-6 (8) cm long, 2-8 mm wide, entire, antrorsely scaberulous on both surfaces (or glabrous), sessile, linear to narrowly oblong, often spin- ulose-mucronate, the lowermost often lacking at anthesis; inflorescences several- to many- headed, cylindroid; involucres 4.6-6.5 mm high, 7-9.5 mm wide, the bracts strongly to only somewhat imbricate, with a green tip, scaberulous to glabrous dorsally and ciliate; rays mainly 17-25, white (drying pale laven- der in some), 6-8 mm long; achenes pu- bescent. Oak, sagebrush, and ponderosa pine communities at 1525 to 2135 m in Box Elder, Kane, Utah, and Washington counties; Alaska to Minnesota, south to California, New Mexi- co, and Kansas; 7 (i). The species is closely al- lied to A. pansus (q.v.), which has smaller heads. Aster foliaceus Lindl. in DC. Leafybract Aster. Perennial rhizomatous or sub- rhizomatous herbs, uniformly and shortly soft-villous below the heads, uniformly vil- lous to glabrous below, or in lines below leaf bases, the stems 1.3-7 dm tall; leaves 1,8-16 cm long, 3-34 mm wide, entire or nearly so, strigose to glabrous, ciliate, the lower ones petiolate (often lacking at anthesis), becom- ing smaller and sessile (and more or less clasping) upwards; inflorescence of 1-19 (50) corymbosely arranged large and showy heads; involucres 6-12 mm high, 10-20 mm wide, the bracts imbricate to slightly so, fo- liaceous or slender, green with pale white to yellowish or brownish chartaceous bases (at least the inner), acute to obtuse or rounded, mucronate; rays mainly 15-50, pink to purple, blue, or violet, 9-16 (20) mm long; achenes hairy. The leafybract aster is a por- tion of an assemblage that includes the con- cept of A. subspicatus Nees. Both A. fo- liaceus and A. subspicatus were described from coastal Alaska (Unalaska and Yakutat Bay, respectively). Brownish bases of in- volucral bracts, commonly serrate leaves, and reddish pappus are supposedly diagnostic for A. subspicatus, which is not known from Utah, but some specimens of A. foliaceus have one or more of these features. In the Alaska Flora (Welsh 1974), I treated both species under the older name of A. sub- spicatus. Now, I follow tradition so as to avoid creation of synonyms should further study indicate a better course of action. Three more or less distinctive infraspecific taxa are present in Utah. 1. Involucral bracts foliaceous, 2-6 mm broad; plants uncommon A. foliaceus var. canbyi — Involucral bracts not especially foliaceous, mainly less than 2 (2.5 ) mm wide; plants common to uncommon 2 2(1). Plants mainly 0.5-2.5 dm tall, decumbent or ascending; bracts often purple margined or tipped; known from high elevations, rare A. foliaceus var. apricus — Plants often more than 2 dm tall, erect; bracts seldom as above; known from low to high elevations, common A. foliaceus var. parryi 220 Great Basin Naturalist Vol. 43, No. 2 Var. apricus Gray Meadows in spruce-fir forest at 3050 to 3660 m in Summit County; British Columbia to Montana, south to Cali- fornia and Colorado; 1 (0). Var. canbyi Gray Mountain brush, aspen, and spruce-fir communities at 1950 to 2900 m in Iron, Juab and Salt Lake counties; Washington to Wyoming, south to California and New Mexico; 5 (0). Var. parryi (D.C. Eaton) Gray [A. adscen- dens var. parryi D.C. Eaton; A. foliaceus var. frondeus Gray]. Meadows and openings in as- pen, spruce, lodgepole pine, and Douglas fir communities at 1890 to 3265 m in Cache, Duchesne, Garfield, Iron, Juab, Piute, Salt Lake, Sanpete, Summit, Tooele, Uintah, and Utah counties; Washington to Wyoming, south to California and New Mexico; 36 (viii). This is the phase of the leafybract aster that simulates A. occidentalis (q.v.), but which seldom has long peduncles, dark blue- purple ray corollas, and much reduced upper stem leaves of that species. Aster frondosus (Nutt.) T. & G. Leafy As- ter. [Tripolium frondosum Nutt.]. Annual herbs from taproots; stems 0.2-3.6 cm tall; leaves 1-6 cm long, 2-12 mm wide, linear to oblong or oblanceolate, entire, the lower ones sometimes deciduous; heads few to nu- merous, in a narrow paniculate to spicate in- florescence; involucres 5-9 mm high, 6-13 mm wide, the bracts oblong to narrowly ob- lanceolate, obtuse or obtusish, herbaceous, subequal to moderately imbricate; marginal flowers pistillate, the rays developed, pink, to 2 mm long; pappus abundant, white, longer than the disk corollas. Lake shores, seep mar- gins, wet meadows, and stream banks in salt- grass, tamarix, Russian olive, rabbitbrush, and greasewood communities at 1250 to 2270 m in Beaver, Duchesne, Garfield, Grand, Juab, Kane, Salt Lake, San Juan, Utah, and Wayne counties; Washington to Wyoming, south to California and New Mexico; 18 (ii). Aster glaucodes Blake Blueleaf Aster. Pe- rennial rhizomatous herbs, glabrous and glaucous, or puberulent to glandular in the inflorescence; stems 1.1-7 dm tall; leaves 1.4-12.5 cm long, 4-25 mm wide, entire, lance-oblong to oblong or elliptic, glaucous, glabrous, sessile and clasping, the lower often lacking at anthesis, reduced upwards; heads few to numerous in corymbose in- florescences; involucres 6-9 mm tall, 7-9 mm wide, the bracts imbricate, dry, chartaceous throughout or sometimes some of them greenish, the midvein prominent, commonly suffused with pink or purple, mainly obtuse to less commonly acute apically; rays 10-20, white or pink, 11-17 mm long. There are two varieties within our specimens. Peduncles and/ or involucres glandular-pubescent; plants of Washington and adjacent western Kane counties A. glaucodes var. pulcher Pedimcles and involucres lacking glandular pubescence; plants widespread A. glaucodes var. glaucodes Var. glaucodes This is the common phase of the species, often on calcareous substrates at higher elevations and in saline seeps at moderate to lower elevations in sagebrush, pinyon-juniper, mountain brush, ponderosa pine, ryegrass, spruce-fir, Douglas fir, lodge- pole pine, and hanging garden communities at 1220 to 3050 m in Cache, Carbon, Du- chesne, Emery, Garfield, Grand, Kane, San Juan, Sanpete, Sevier, Summit, Tooele, Uin- tah, Utah, Washington, and Wayne counties; Idaho and Wyoming, south to Arizona and Colorado; 56 (ix). Var. pulcher (Blake) Kearney & Peebles Note: This taxon was published at subspecific rank by Blake, and was inadvertantly accept- ed at varietal rank by Kearney and Peebles (Arizona Flora 872. 1951), without citation of the basionym, A. glaucodes ssp. pulcher Blake, Proc. Biol. Soc. Washington 35: 174. 1922. Salt desert shrub, sagebrush, pinyon- juniper, and ponderosa pine communities at 825 to 2136 m in Washington and adjacent western Kane counties; Arizona; 6 (0). Aster hesperius Gray Siskiyou Aster. [A. laetivirens Greene]. Perennial rhizomatous herbs, villous with multicellular hairs in de- current lines from leaf bases, or less com- monly almost glabrous and with decurrent lines below the heads; stems 3.6-9.5 (15) dm April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 221 tall; leaves 3-17 (21) cm long, 5-27 mm wide, entire or serrate, glabrous or scabrous, ciliate, the lower ones commonly petiolate, often deciduous at an thesis, becoming small- er, sessile and more or less clasping upwards, sometimes much reduced in inflorescence; heads few to numerous in open to narrow subcorymbose inflorescences; involucres 4.5-7 (8) mm high, 7-12 mm wide, the bracts imbricate to subequal, green tipped, the chartaceous base white to straw colored, all acute and mucronate; rays commonly 20-50, pink to blue or white, 6-14 mm long; ach- enes hairy. Wet meadows, canal banks, and stream sides with sedges, rabbitbrush, willow, and other riparian communities at 850 to 2135 m in Box Elder, Cache, Duchesne, Gar- field, Grand, Kane, Millard, Summit, Utah, Wasatch, and Washington counties; Alberta to Saskatchewan, south to California, Ari- zona, New Mexico, and Missouri; 32 (vii). This plant occurs at lower elevations in Utah and has been confused with A. foliaceus, with which some plants share the subequal bracts. It has also been mistaken for A. chilensis, with which it is partially sympatric. The lack of uniformly disposed hair in the in- florescence appears to be diagnostic. Aster integrifolius Nutt. Thickstem Aster. Perennial subrhizomatous herbs, glandular villous with multicellular hairs, at least above; stems 2.3-6.4 (7) dm tall; leaves 2.5-19 cm long, 8-50 mm wide, entire, ob- lanceolate to elliptic or lanceolate, glandular- villous, ciliate, the lower ones petiolate, be- coming smaller, sessile and clasping upward; heads few to several (numerous), large and showy, in elongate to subcorymbose clusters; involucres 8-13 (14) mm high, 12-23 mm wide, the bracts mainly subequal, green or suffused with purple, glandular dorsally, fo- liaceous or not; rays commonly 10-25, dark purple, 10-15 mm long. Meadows and moist woods in sedge-willow, sagebrush, Douglas fir, and spruce communities at 2275 to 3125 m in Rich, Salt Lake, Summit, and Wasatch counties; Washington and Montana, south to California and Colorado; 8 (0). Aster kingii D.C. Eaton [Machaeranthera kingii (D.C. Eaton) Cronq. & Keck]. Pe- rennial herbs from a caudex and taproot, the caudex branches clothed with blackish or dark brown marcescent leaf bases, these scarious and ashy when young; stems 3-12 (15) cm long, more or less villous below, stipitate-glandular above; basal leaves 0.8-12 cm long, 3-22 mm wide, petiolate, the pet- iole bases expanded and scarious, the blades oblanceolate or spatulate, glabrous or glandu- lar, or less commonly hispidulous or merely puberulent on one or both sides; heads 1-5, racemosely or corymbosely arranged; in- volucres 8-11 mm high, 10-16 mm wide; bracts glandular to shortly stipitate-glandu- lar, herbaceous above the middle, scarious below, often suffused purplish, especially the inner, the tips of at least the outer reflexed; rays 15-27, white (often fading pale pink), 8-17 mm long, 1.5-2.8 mm wide; achenes ca 3.5 mm long. Douglas fir-white fir, mountain brush, and Cottonwood communities at 1839 to 3050 m in Juab, Millard, Salt Lake, and Utah counties; endemic; 21 (i). The southern populations have at least some toothed leaves and stems with longer stipitate-glandular hairs; they belong to var. barnebyana (Welsh & Goodrich) Welsh comb. nov. [based on: Machaeranthera kingii var. barnebyana Welsh & Goodrich Brittonia 33: 299. 1981]; 6 (0). Attempts to segregate genera within the Astereae are often fraught with diffi- culties. This is especially true of that core of genera involving Haplopappus, Machaeran- thera, Xylorhiza, and Aster. Cronquist and Keck (1957. Brittonia 9: 231-329) reconsti- tuted the genus Machaeranthera, and includ- ed within that expanded generic definition those species treated elsewhere herein as Machaeranthera and Xylorhiza. Included within the series Integrifoliae of section Xy- lorhiza was Aster kingii. Watson (1978. Madrono 25: 205-210) has shown the chromosome number to be 2n=18 for Aster kingii, and he notes that its placement within Machaeranthera section Xylorhiza "is pheno- logically, ecologically, morphologically, and chromosomally anomalous. ..." The chromo- some numbers reported for Xylorhiza are 2n = 12 or 24; that of Machaeranthera, in a restricted sense, is 2n = 8, 10, or 16; that of Aster is mainly 2n = 18. The taproots and squarrose involucral bracts suggest an al- liance with Machaeranthera, shorn of Xylo- rhiza, but the similarity seems superficial, es- pecially in light of different chromosome numbers. Some asters in a strict sense, i.e., 222 Great Basin Naturalist Vol. 43, No. 2 A. alpigenns Rydb., have a caudex, with the rhizome attenuated. The logical conclusion of such an attenuation is the caudex of A. kingii, and the squarrose bracts seem to have been secondarily derived, being present to a greater or lesser degree in other Aster species as well as in Machaeranthera. Hence, it seems best to treat this taxon within Aster. Aster laevis L. Smooth Aster. Sub- rhizomatous perennial herbs, glabrous or nearly so; stems mainly 5-12 dm tall; leaves 0.8-14 cm long, 2-30 mm wide, entire or ser- rate, linear-subulate to lanceolate or elliptic, the lower ones petiolate, often lacking at an- thesis, becoming smaller, sessile, and more or less clasping upwards; heads numerous, in co- rymbose inflorescences; involucres 5-8 mm high, 7-12 mm broad (when pressed), the bracts slender, green tipped, the chartaceous bases straw colored to brownish or white, acute and mucronate; rays 15-30, blue or purple, 6-9 mm long; achenes glabrous. Ri- parian communities at ca 1400 m in Grand (and San Juan?) County; Yukon to Maine, south to Oregon, New Mexico, and Georgia; 1 (i). This plant is rare in collections from Utah, due presumably to the paucity of late season collections from southeastern Utah. Aster occidentalis (Nutt.) T. & G. Western Aster. [Tripolium occidentale Nutt.]. Rhizo- matous or subrhizomatous perennial herbs, uniformly, softly, and often loosely villous (at least above); stems 0.9-8.5 dm tall; leaves 1-15 cm long, 1-20 mm wide, entire or toothed, glabrous or nearly so, ciliate, the lower ones petiolate, sometimes lacking at anthesis, rather abruptly smaller and finally sessile upwards; inflorescence mainly of 1-7 (rarely to 15), corymbosely arranged large and showy heads; involucres 5-12 mm high, 7-20 mm wide, the bracts imbricate to sub- equal, slender, green, with pale yellowish to white or brownish chartaceous bases (at least the inner), mainly acute, mucronate; rays 20-50, blue to purple, 6-15 mm long; ach- enes hairy. Meadows and stream sides in lodgepole pine, cottonwood, willow, aspen, and spruce-fir communities at 2175 to 3175 m in Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Piute, Sanpete, Sevier, Sum- mit, and Wasatch counties; Mackenzie to Colorado and California; 43 (ix). This species shares the features of soft loose pubescence and general aspect with the partially sympa- tric A. foliaceus. The very slender and abruptly reduced cauline leaves are diagnos- tic in most instances. Aster pansus (Blake) Cronq. Elongate Aster. Subrhizomatous herbs, villous or villous-hir- sute with multicellular hairs, the stems 3-12 (or more) dm tall; leaves 1-6 cm long, 2-8 mm wide, entire, antrorsely scaberulous on both surfaces, sessile, linear to narrowly ob- long, often spinulose-mucronate, the lower- most commonly lacking at anthesis; in- florescence paniculate to secund-paniculate, narrow; involucres 3.8-4.9 mm high, 4.5-6 mm wide, the bracts strongly imbricate, green tipped, scaberulous dorsally and cil- iate; rays mainly 15-25, white, 3-8 mm long; achenes hairy. Drainages, meadows, seeps, and hanging gardens at 1220 to 1890 m in Daggett, Grand, San Juan, and Uintah coun- ties; British Columbia to Montana, south to Colorado and Nebraska; 6 (iv). This species forms the basis for inclusion in previous bo- tanical works of the name A. ericoides L. in the Utah flora. It is closely allied to A. fal- catus, but differs in the smaller heads, taller stature, and eastern distribution. Aster pauciflorus Nutt. Alkali Aster. [A. thermalis Jones, type from Monroe Hot Springs]. Subrhizomatous perennial herbs, glabrous below, stipitate-glandular above and in inflorescence; stems 2-7.5 dm tall; leaves 1.1-12.5 cm long, 1-4 mm wide, entire, acicular to lance-linear or linear, glaucous, glabrous, all sessile or the lowermost petio- late, reduced upwards; heads few to several in corymbose inflorescences; involucres 4.3-7 mm long, 7-10 mm wide, the bracts imbri- cate to subequal, glandular dorsally, green throughout, narrow and acute; rays mainly 20-35, blue to purple, 5-12 mm long; ach- enes hairy. Hot springs, stream terraces, and salt grass meadows, often in saline or alkaline substrates at 1300 to 2135 m in Box Elder, Duchesne, Emery, Juab, Kane, Millard, San- pete, Sevier, Salt Lake, and Utah counties; Saskatchewan to Nevada, Arizona, and Mexi- co. This distinctive glandular aster has been collected in full anthesis on 27 April growing in hot water at Monroe Hot Springs in Sevier County. It continues to flower into October; 21 (vi). April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 223 Aster perelegans Nels. & Macbr. Nuttall Aster. [EucepJialus elegans Nutt.; A. elegans (Nutt.) T. & G., not Willd.]. Subrhizomatous perennial herb, puberulent to glabrate (some.- times glandular); stems 3-7 dm tall; leaves 1.3-6.5 cm long, 3-14 mm wide, entire, ob- long to oblong-lanceolate or elliptic, scabrous, firm, sessile, the lowermost reduced in size; heads 3-16, in corymbose in- florescences; involucres 7-10 mm high, 7-12 mm wide, the bracts chartaceous, imbricate, with prominent midvein, and acute to obtuse apex, the margins hyaline and ciliate, more or less puberulent dorsally; rays 5-16, dark purple, 7-13 mm long; achenes hairy. Sage- brush, mountain brush, Douglas fir, aspen, and limber pine communities at 1725 to 3050 m in Carbon, Duchesne, Juab, Millard, Salt Lake, Wasatch, and Weber counties; Oregon to Montana, south to Nevada; 21 (iv). Aster scopulorum Gray Crag Aster. [Chry- sopsis alpina Nutt., not A. alpinus L.]. Pe- rennial subrhizomatous herbs with a woody caudex, villous on stems and peduncles; stems 4-12 cm tall; leaves 5-12 (15) mm long, 1-3 mm wide, overlapping, elliptic to oblong or linear, firm, scabrous or puberulent, often with some villous hairs above, spinulose- mucronate; heads solitary, pedunculate; in- volucre 7-11 mm high, 8-12 mm wide, the bracts imbricate, sparingly villous-hirsute and glandular, with a prominent midvein in the lower half, greenish, with chartaceous border and hyaline margins, acute; rays mainly 8-15, blue or purplish, 6-15 mm long; achenes hairy. Sagebrush community at 2440 to 2745 m in Box Elder County; Oregon to Montana, south to California and Nevada; 8 (0). Aster spinosus Benth. Mexican Devilweed. Suffrutescent, rushlike plants from a deeply placed rhizome, glabrous; stems 6-12 (or more) dm tall, with axillary or supraxillary thorns to 1.5 cm long; leaves 2-4 cm long, 2-5 mm wide, firm, entire to toothed, re- duced above to scales; heads solitary at ends of branches, or some axillary; involucres 4-6 mm high, 6-8 mm wide, the bracts imbricate, slender, acute to acuminate, green, with prominent scarious margin; rays 15-30, white, very short; achenes glabrous. Riparian communities at below 1130 m in Garfield and probably Washington counties; Califor- nia to Texas, south to Central America; 1 (0). The plant was collected at the mouth of Ti- caboo Canyon, along the Colorado River in Glen Canyon (Lindsay 20, 1958 UT), at a site now inundated by Lake Powell. This is one of a series of extirpations related to construc- tion of Glen Canyon Dam. The plant should be sought in the St. George vicinity. Aster wasatchensis (Jones) Blake Mark- agunt Aster. Subrhizomatous perennial, glandular-puberulent; stems 3.5-6.5 dm tall; leaves 1.8-8.5 cm long, 6-24 mm wide, en- tire, lanceolate to oblong, or oblanceolate, glandular-puberulent to glabrous, firm, more or less glaucous, the lowermost often smaller and commonly lacking at an thesis; heads sev- eral to numerous, more or less corymbosely arranged; involucres 8-11.5 mm long, 10-20 mm wide, the bracts herbaceous throughout or the inner with scarious bases, glandular dorsally, abruptly acute to attenuate, api- cally; rays 15-25, white or pink, 10-20 mm long; achenes hairy. Pinyon-juniper, aspen, limber pine, and spruce-fir communities at 1890 to 3050 m in Garfield, Iron, Millard, and Piute counties; endemic. This remarkable aster is unique in Utah in having foliaceous or subfoliaceous glandular involucral bracts and glaucous leaves; 14 (vi). Atrichoseris Gray Annual scapose herbs, with milky juice, from taproots; leaves all basal, sinuate-den- tate, often spotted; heads on slender pe- duncles, few to numerous, corymbosely ar- ranged; involucre of about 12-15 subequal but biseriate, lance-linear scarious-margined bracts and some shorter outer bracts; recep- tacle naked; corollas all raylike, perfect, white; pappus lacking; achenes oblong, with corky-thickened ribs. Atrichoseris platyphylla Gray Tobacco- weed; Gravel Ghost. Plants 3-10 dm tall (or more), from slender taproots; leaves 1.2-10.5 cm long, 0.5-6 cm wide, obovate to broadly spatulate, tapering abruptly to a broad pet- iole, sinuate-dentate, the teeth mucronate- cuspidate, glabrous, often mottled, more or less glaucous; involucres 6-8 mm high, 12-16 mm wide, the outer bracts ovate-lanceolate, hyaline, more or less scurfy, the inner ones lance-acuminate, with broad hyaline margins; 224 Great Basin Naturalist Vol. 43, No. 2 corollas white, 8-20 mm long; achenes white, with corky ridges. Joshua tree, ambrosia, yucca, cholla communities at 670 to 750 m in Washington County; California and Arizona; 4(i). Baccharis L. Dioecious shrubs; leaves alternate, entire or toothed; heads discoid, many flowered, the corollas white, turbinate, borne in corymbose or paniculate clusters; involucres imbricate, the bracts chartaceous, whitish; pistillate heads with tubular-filiform obscurely toothed or truncate corollas, the pappus of copious capillary bristles; staminate heads of tubular 5-toothed corollas, the pappus (often scanty) of usually twisted clavellate scales; recep- tacle naked; style branches flattened; achenes subcylindric, 5- to 10-ribbed. 1. Branches fastigiate, deeply sulcate and more or less ridged, the leaves com- monly deciduous at anthesis; achenes 10-ridged 2 — Branches not especially fastigiate, commonly spreading to ascending; leaves commonly persistent at flowering time; achenes 5- or 10-ribbed 3 2(1). Main leaves linear; pistillate pappus to 10 mm long or more in fruit ... B. sarothroides — Main leaves obovate-spatulate; pistillate pappus to 3 mm long in fruit. ..B. sergilloides 3(1). Leaves long-cuneate basally, thickened, entire or few toothed toward apex; branches often subfastigiate; achenes 10-nerved; plants of Virgin and Colorado drainages 4 — Leaves not especially long-cuneate basally, commonly thin, entire, or toothed from below the middle; achenes 5-nerved 5 4(3). Staminate involucres 3.5-5.3 mm long, 3.7-4.8 mm wide; pistillate involucres 7.3-8.5 mm long; pappus 11-13 mm long; plants of Washington and Kane counties B. emoryi — Staminate involucres 5.3-6 mm long, 5-10 mm wide; pistillate involucres 6-6.5 mm long; pappus 8-9.5 mm long; plants of Emery, Grand, Garfield, and San Juan counties B. salicina 5(3). Leaves mainly entire; panicles terminating short lateral branches; plants of Washington County B. viminea — Leaves usually serrate; panicles terminating main stems; plants of Washington and Kane counties B. glutinosa Baccharis emoryi Gray in Torr. Shrubs, mainly 1-2 (3) m tall, the branches green to olive or brownish, ascending, subfastigiate, more or less glutinous; leaves 1.2-8.5 cm long, 3-20 mm wide, spatulate-oblanceolate to elliptic or linear, cuneate to a slender pet- iole, thick, entire or sparingly and irregularly toothed, obtuse to acute apically; heads nu- merous in a conic to pyramidal panicle; pis- tillate involucres 7.3-8.3 mm high, 4.5-7 mm wide, the bracts in several series, scarious, of- ten glutinous, with thickened green or brown to reddish tips and hyaline margins; stami- nate involucres 3.7-5.3 mm high, 3.7-4.8 mm wide; pistillate corollas 4.5-5.5 mm long, the pappus 11-13 mm long; achenes 10-ribbed. Stream and canal banks and hanging gar- dens at 825 to 1220 m in Kane and Washing- ton counties; Arizona, Texas, and California; Mexico; 19 (iv). Baccharis glutinosa Pers. Shrubs, mainly 1-3 m tall, the branches straw colored to brownish or greenish, ascending-spreading, not fastigiate, glutinous; leaves 1.2-12.5 (15) cm long, 4-18 mm wide, elliptic to narrowly lanceolate, acuminate to attenuate, cuneate to a short petiole, evenly serrate to entire; heads numerous in terminal cymose panicles (less commonly in lateral ones) with pistillate and staminate heads about the same size; in- volucres 3.5-4.5 mm high, 4-5.5 (7.5) mm wide; corollas 2.2-3 mm long, the pistillate pappus 3.5-4.5 mm long; involucral bracts in several series, chartaceous, greenish in the April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 225 center, the margins scarious, not glutinous; achenes 5-ribbed. Stream bars and banks, and in seeps, at 670 to 1130 m in Kane and Wash- ington counties; Colorado and Nevada to Texas and California; South America; 6 (ii). Baccharis salicina T. & G. Shrubs, mainly 1.5-3 m tall, the branches green to brownish, subfastigiate, glutinous; leaves 1.4-8 cm long, 4-18 mm wide, elliptic to oblanceolate or linear, cuneate to a short petiole, thick or thin, entire or sparingly toothed or lobed mainly near the apex, acute to rounded api- cally; heads few to numerous in axillary and/ or terminal panicles; pistillate involucres 6-6.5 mm high, 4-6 mm wide, the bracts in several series, scarious, often glutinous, with thickened greenish to reddish tips and hya- line margins; staminate involucres 5.3-6 mm high, 5-10 mm wide; pistillate corollas 2.5-3.5 mm long, the pappus 8-10 mm long; achenes 10-ribbed. Stream banks and hanging gardens at 1220 to 1525 m in Emery, Gar- field, Grand, and San Juan counties; Colo- rado to Kansas, south to New Mexico and Texas; 7 (iv). Our material of B. salicina has long been mistaken for B. emoryi, to which it is allied. The shorter pistillate involucres and broader staminate involucres are diagnostic. Baccharis sarothroides Gray Broom Bac- charis. Shrubs, mainly 1-3 m tall, the branches green to brown, fastigiate, glu- tinous, finally almost leafless; leaves 1-3.5 cm long, 2-5 mm wide, linear to oblong, entire, ridged; heads solitary at tips of fastigiate branches forming a hemispheric panicle; pis- tillate involucres 6-8 mm high, 5-10 mm wide, the bracts in several series, cream col- ored; staminate involucres 3-4 mm high, 4-8 mm wide, the bracts green apically; pappus 6-11 mm long; achenes 10-ribbed. Reported for Washington County (UT), where presum- ably it occurs along streams; California to New Mexico; 0 (0). Baccharis sergilloides Gray Squaw Water- weed. Shrubs, mainly 0.3-2 m tall, the branches green to brown, fastigiate, glu- tinous, finally almost leafless; leaves 0.5-2.5 cm long, 1-10 mm wide, spatulate to obo- vate, entire or few toothed, thick; heads nu- merous, borne in conic to pyramidal panicles; involucres 2.5-3.5 mm high, 2.5-3.5 mm wide, the bracts in several series, straw col- ored, or with thickened brownish centers; pappus 2.5-3 mm long; achenes 10-ribbed. Stream bars and banks at 670 to 825 m in Washington County; California and Arizona; 3(i). Baccharis viminea DC. Mule-fat. Shrubs, mainly 2-3 m tall, the branches green to straw colored or brownish, spreading-ascend- ing, not fastigiate, glutinous; leaves 0.8-9.5 cm long, 2-9 mm wide, elliptic to lance-el- liptic or narrowly oblong, attenuate to acute, cuneate to a short petiole, entire to evenly serrate; heads few to many in terminal cy- mose panicles on short lateral branches, with pistillate and staminate heads about the same size; involucres 3-5.7 mm high, 6-9 mm wide; corollas 2.5-3.8 mm long; pistillate pappus 5-6 mm long; involucral bracts in several series, chartaceous, commonly with reddish centers, the margins scarious, not glu- tinous; achenes 5-ribbed. Stream bars and banks at 650 to 900 m in Washington Coun- ty; California and Arizona; 10 (ii). Bah I A Lag. Biennial or short-lived perennial herbs with watery juice, arising from taproots; stems erect or ascending, puberulent; leaves alternate, once to twice ternately divided; heads few to numerous, in corymbose pan- icles; involucral bracts subequal, in 1 or 2 series, greenish; ray flowers present, yellow, pistillate, fertile; disk flowers perfect, fertile; pappus none; style branches flattened; ach enes 4-angled, 12-nerved. Bahia dissecta (Gray) Britt. [Amauria dis- secta Gray]. Biennial or short-lived perennial herbs, the stems 2-8 dm tall, minutely pu- berulent; leaves 1-10 cm long, the blade 1- to 3-ternately divided, oval to cordate in out- line, strigulose; peduncles glandular hairy; in- volucres hemispheric, 3.4-6 mm high, 8-12 mm wide, the bracts more or less glandular hairy (or merely villous), greenish, abruptly contracted to a broadened apex; rays mainly 10-15, yellow, 4.5-9 mm long; achenes glabrous. Sagebrush, pinyon-juniper, moun- tain brush, aspen, lodgepole pine, ponderosa pine, and spruce communities at 1700 to 2930 m in Beaver, Garfield, Grand, Kane, Se- vier, Uintah, Washington, and Wayne coun- ties; Nevada to Wyoming, south to Califor- nia, Arizona, and Mexico; 24 (iii). Those species treated elsewhere in this work as Pla- tyschkuhria belong to Bahia in a broad sense 226 Great Basin Naturalist Vol. 43, No. 2 and are probably best treated in the latter genus, but their combination is not implied here. Baileya Harv. & Gray Annual, biennial, or perennial herbs from taproots, with watery juice; stem erect, white-tomentose; leaves alternate, 1- or 2- pinnatifid to entire; heads solitary or few in cymose clusters; involucral bracts subequal, white-tomentose; receptacle naked; ray flow- ers persistent, yellow, pistillate, fertile; disk flowers perfect, fertile; pappus none; style branches short, truncate; achenes oblong or clavate, striate. 1. Ray flowers 7 or fewer; plants slender annuals with involucres less than 8 mm wide B. pauciradiata — Ray flowers 20 or more; plants annual, biennial, or perennial, with involucres 10-26 mm wide 2 2(1). Rays 11-22 mm long; peduncles (4.5) 12-32 cm long in anthesis; involucres 5.7-7.5 mm high, 13-26 mm wide B. multiradiata — Rays 8-10 mm long; peduncles 1-8 (11) cm long in anthesis; involucres 3-5.5 mm high, 10-16 mm wide B. pleniradiata 0.8-5 dm tall, white-tomentose; leaves 0.8-12 Baileya multiradiata Harv. & Gray Bien- cm long, the blades 1- to 2-pinnately lobed to nial or short-lived perennial herbs; stems entire, obovate to linear, white-tomentose; 1.9-5 (5.2) dm tall, white-tomentose; leaves peduncles 1-8 (11) cm long in anthesis, 0.8-10 cm long, the blade 1- to 2-pinnately white-tomentose; involucres 3.5-5.5 mm lobed to entire, ovate-oval to linear, white- high, 6-13 mm wide, the bracts slender, tomentose; peduncles (4.5) 13-32 cm long in greenish, white-tomentose; rays 18-58, yel- anthesis, white-tomentose; involucres 5-7.5 low, 8-10 mm long; achenes glabrous. Creo- mm high, 13-26 mm wide, the bracts slender, sote bush, blackbrush, shadscale, mesquite, greenish, white-tomentose; rays 25-40 or sagebrush, and pinyon-juniper communities more, yellow, 11-22 mm long; achenes at 820 to 1100 m in Washington County; Ne- glabrous. Creosote bush, Joshua tree, bur- vada and California to Texas; Mexico; 29 (ii). robush, blackbrush, and sagebrush commu- nities at 670 to 1320 m in western Kane and Washington counties; Nevada and California Balsamorhiza Nutt. south to Mexico; 36 (iii). Baileya pauciradiata Harv. & Gray An- Perennial scapose or subscapose herbs from nual herbs; stems mainly 1.5-4 (6) dm tall, taproots, the juice watery; leaves mainly bas- densely floccose-lanate; leaves 3-10 cm long, al, simple and entire or variously pinnatifid, the blades entire or the lower irregularly pin- reduced and bractlike upward; heads solitary, natifid (or bipinnatifid), linear or linear-Ian- or few to several; involucral bracts in several ceolate, white-tomentose; peduncles 2-5 cm series, imbricate or subequal, herbaceous; re- long in anthesis, tomentose; involucres 5-6 ceptacle chaffy, convex, the bracts enclosing mm high, 5-8 mm broad, the bracts slender, the achenes; ray flowers present, pistillate, greenish, loosely tomentose; rays 5-7, yellow, fertile, usually yellow; disk flowers numerous, 5-8 mm long; achenes glabrous. Reported perfect, fertile, yellow; pappus none; style from Washington County by Meyer (1976), branches slender; achenes compressed. Note: where it was collected at Warner Valley The genus is notorious for the lack of genetic Spring; California, Arizona, and Mexico; 0 (0). barriers to hybridization. Any two taxa can Baileya pleniradiata Harv. & Gray An- intergrade where they occur together, nual to short-lived perermial herbs; stems Leaves sagittate, with entire margins B. sagittate Leaves pinnatifid or variously cleft 2 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 227 2(1). Leaves mainly 3-6 dm long, with segments mainly 5-12 cm long, these entire or few lobed or toothed B. macrophylla — Leaves mainly 1-3 dm long, with segments mostly 1-5 cm long, these entire or variously lobed or toothed 3 3(2). Involucral bracts abruptly tapering to a long-attenuate apex; stem leaves rela- tively well developed, pinnatifid or bipinnatifid; reported for northern Utah, but no specimens have been seen B. hirsuta Nutt. — Involucral bracts gradually tapering to an attenuate apex; stem leaves lacking or small and inconspicuous B. hookeri Balsamorhiza hookeri Nutt. Hooker Bal- ^^^ ^^^^t^ lance-linear, evenly tapering to the samroot. Perennial scapose herbs from a ^P^^ °' somewhat enlarged at the base, long- thick taproot, mainly 0.9-4.5 (5.2) dm tall; ^^1^^*^' glandular to tomentose dorsally; rays leaves 6-30 cm long, (0.3) 1.5-11 cm wide, "^^'^^ly ^^-l^' Y^^^^w, 16-40 mm long; ach- pinnatifid or bipinnatifid, the segments to 5.5 ^"^^ glabrous. Phases of this taxon are known cm long; peduncles naked or with a few in- *« ^^^ mtermediates with B. sagittata, and conspicuous, linear, entire or pinnatifid Presumably with B. macrophyllum. Ours are bracts near the base; heads solitary; in- separable into two modestly distinctive volucres 13-24 mm high, 21-47 mm wide, varieties. Involucres densely villous-tomentose dorsally; plants of Daggett, Duchesne, and Uintah counties B. hookeri var. neglecta Involucres glandular to glabrous dorsally; plants of broad distribution, occa- sionally of Daggett and Duchesne counties B. hookeri var. hispidula Var. hispidula (Sharp) Cronq. [B. hispidula Sharp]. This is the common phase of the spe- cies in Utah, and it has been confused with B. hirsuta Nutt., with which it is compared in the key. Bunchgrass, sagebrush, mountain brush, juniper, pinyon-juniper, and salt desert shrub communities at 1240 to 2745 m in Bea- ver, Box Elder, Daggett, Duchesne, Juab, Salt Lake, Tooele, Utah, Wasatch, and Washing- ton counties; Nevada, Idaho, and Wyoming (?); 31 (ii). Var. neglecta (Sharp) Cronq. [B. hirsuta var. neglecta Sharp] .Salt desert shrub, sage- brush, pinyon-juniper, and ponderosa pine communities at 1640 to 2625 m in Daggett, Duchesne, and Uintah counties; Nevada, Idaho, and Wyoming (?); 18 (ii). Plants of this variety form hybrids with B. sagittata. Balsamorhiza macrophylla Nutt. Cutleaf Balsamroot. Perermial scapose herbs from a thick taproot, mainly 3-7 dm tall; leaves 15-60 cm long, 3.7-25 cm wide, pinnatifid, the segments entire, few toothed or lobed, up to 12.5 cm long; peduncles sparingly long shaggy-villous, naked, or with one to few re- duced leaves near the base; heads solitary; in- volucres 23-35 mm high, 30-60 mm wide, the bracts lance-linear, attenuate, long- ciliate, glandular and more or less long-vil- lous dorsally; rays 9-14, yellow, 30-55 mm long; achenes glabrous. Mountain brush and sagebrush or bunchgrass communities at 1525 to 2290 m in Box Elder, Cache, Salt Lake, Summit, Utah, and Weber counties; Idaho to Montana and Wyoming; 7 (0). Balsamorhiza sagittata (Pursh) Nutt. Ar- rowleaf Balsamroot. [Bupthalmium sagitta- tum Pursh]. Perennial scapose herbs, from thick taproot, mainly 1.5-8 dm tall; leaves (including long slender petioles) 5-45 cm long, 1.5-15 cm wide, sagittate, entire, or the cauline ones from near the summit to near the middle of the subscapose stem and linear to elliptic; peduncles villous-tomentose; heads solitary (or with additional reduced ones); involucre 15-30 mm long, 20-50 mm wide, the bracts lance-linear, attenuate, vil- lous-tomentose; rays 8-25, yellow, 25-60 mm long; achenes glabrous. Sagebrush, mountain brush, pinyon-juniper, ponderosa pine. 228 Great Basin Naturalist Vol. 43, No. 2 Douglas fir, aspen, and fir communities at 1340 to 3020 m in Beaver, Box Elder, Cache, Davis, Garfield, Iron, Juab, Kane, Millard, San Juan, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, and Washington coun- ties; British Columbia to Montana and South Dakota, south to California, Nevada, and Colorado; 43 (vii). Bellis L. Scapose perennial herbs, with fibrous roots and short stolons, the juice watery; stems leafless, simple; leaves all basal, simple, pet- iolate, toothed to entire; heads solitary; in- volucral bracts in 2 subequal series, her- baceous; receptacle conic to hemispheric, naked; rays white, pink, or purple, numerous, pistillate; disk flowers numerous, perfect, yel- low; pappus lacking; style branches flattened; achenes flattened, usually 2-nerved, pubescent. Bellis perennis L. European Daisy. Plants 0.2-2 dm tall; leaves all basal, with short to long petioles, the blades 0.7-3 (4) cm long, 5-25 mm wide, obovate to oval or orbicular, dentate to entire, obtuse to rounded or emarginate apically, pubescent on both sides with coarse spreading hairs; scapes pubescent with ascending hairs; heads solitary; in- volucres 4-7 mm high, 9-15 mm wide, the bracts ovate to broadly lanceolate, rounded to obtuse apically, sparsely hairy dorsally, of- ten suffused with purple, mostly 8-10 mm long, 1.5-2.5 mm wide; pappus lacking; ach- enes flattened. Cultivated ornamental, escap- ing and persisting in lawns of lower valleys in Salt Lake and Utah counties; adventive from Europe; 4 (0). BiDENS L. Annual herbs with fibrous roots, or rooting along the lower stem, the juice watery; stems decumbent to erect, commonly branched; leaves opposite, simple or pinnately com- pound; heads few to several in cymose in- florescences; involucral bracts in 2 series, the outer herbaceous, the inner somewhat pet- aloid and striate; receptacle flat or slightly convex, chaffy throughout, the chaff similar to the inner involucral bracts; ray flowers present, yellow, neutral or pistillate, or lack ing; disk flowers numerous, perfect, fertile, yellow; pappus of (1) 2-4 awns or teeth, these retrorsely barbed, persistent; style branches flattened; achenes flattened, pu- bescent, usually 2- to 4-awned. Sherff, E. E. 1937. The genus Bidens. Field Mus. Pub. Bot. 16:1-709. Leaves simple, the middle and upper ones (at least) sessile or subsessile B. cernua Leaves pinnately compound, with 3-5 leaflets, all petiolate B. frondosa Bidens cernua L. Bur-marigold. Plants 1-13 dm tall, the stems sparingly sp reading- hairy to glabrous; leaves simple, 1.5-15 cm long, 0.5-4 cm wide, narrowly lanceolate to lance-ovate, coarsely serrate to subentire, glabrous; heads nodding in age; outer in- volucral bracts 5-8, green, foliaceous, un- equal, spreading or reflexed, the inner bracts erect, mostly 6-15 mm long; rays 6-8, yel- low, or lacking; achenes mainly 5-7 mm long, tan, the 2-4 awns retrorsely barbed. Wet meadows, bogs, stream banks, bars, and shores, at 1300 to 2380 m in Cache, Garfield, Juab, Kane, Salt Lake, Sevier, Summit, Uin- tah, and Tooele coimties; widely distributed in the Northern Hemisphere; 27 (iii). Bidens frondosa L. Devil's Beggarticks. Plants 2-12 dm tall, the stems short-hairy to glabrous; leaves petiolate, pinnately com- pound with 3-5 leaflets, these 2-10 cm long, 0.5-3 cm wide, lanceolate, serrate; heads erect in age; outer involucral bracts 5-8, green, subfoliaceous, subequal, erect or spreading, the inner bracts erect, mostly 5-8 mm long; rays usually lacking; achenes 5-9 mm long, dark brown to black, the 2 awns barbed. Marshes, pond and lake shores, bars, wet meadows, and irrigation canals at 1190 to 1650 m in Davis, Grand, Salt Lake, and Utah counties; widespread in North America; 12 (ii). Note: The panboreal weed, Bidens tri- partita L., might occur in our area. It is dis- tinguished from B. frondosa in its simple but trifid leaves, and from B. cernua in its petio- late trifid leaves. Brickellia Ell. Nom. Cons. Perennial herbs, subshrubs or shrubs; leaves alternate or opposite, simple; heads cam- April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 229 panulate or cylindric, cymose or paniculate, discoid; flowers all perfect, fertile; involucral bracts imbricate in several series, striate; re- ceptacle almost flat, naked; style branches flattened, with long-papillate appendage; achenes 10-ribbed; pappus of barbellate, smooth, or subplumose bristles. Robinson, B. L. 1917. A monograph of the genus Brickellia. Mem. Gray. Herb. 1: 1-151. 1. Leaves spinulose-serrate, or spinulose tipped; low rounded shrubs of Washing- ton and San Juan counties B. atractyloides — Leaves entire or toothed, not spinulose; herbs, subshrubs, or tall shrubs of vari- ous distribution 2 2(1). Plants herbaceous; heads reflexed, broadly campanulate; leaves sagittate- to cordate-ovate, longer than broad B. grandiflora — Plants, shrubs, or subshrubs; heads narrowly cylindric, or, if campanulate, erect; leaves ovate to linear, if cordate, about as broad as long or broader 3 3(2). Leaves petiolate, the blades cordate-ovate to ovate or suborbicular, 1-5 cm broad B. calif ornica — Leaves sessile or subsessile, linear to narrowly lanceolate, or, if broader, mainly less than 1 cm broad 4 4(3). Leaves linear to lanceolate or narrowly elliptic; shrubs 6-15 dm tall or more; flowers 3-5 per head B. longifolia — Leaves ovate to oval or oblong to linear; shrubs or subshrubs less than 5 dm tall; flowers many per head 5 5(4). Leaves 5-10 times longer than broad or more, entire or nearly so, sessile; in- volucres 10-20 mm high B. oblongifolia — Leaves only somewhat longer than broad, often toothed or lobed, at least some evidently petiolate; involucres 8-12 mm high B. microphylla Brickellia atractyloides Gray Shrubs, much branched, mostly 3-5 dm tall, the branchlets greenish to straw colored, soon gray; leaves alternate, short-petiolate, the blades 0.6-3.2 cm long, 0.3-2.2 cm wide, lance-ovate to ovate, obtuse to rounded ba- sally, spinulose-serrate to entire, acuminate and spinulose tipped apically, thick and prominently veined, glabrous or minutely glandular puberulent; heads solitary, termi- nating the branches; peduncles 1-5.2 cm long, glandular-puberulent; involucres 10-13.5 mm high, 8-16 mm wide, the outer bracts ovate-lanceolate, acuminate apically, many veined; the inner narrower, glandular- puberulent dorsally; flowers 50-75 or more; achenes black, 3.8-4.2 mm long, hirtellous on the ribs. Rock crevices and talus slopes, creosote bush, blackbrush, and indigo bush communities at 820 to 1130 m in San Juan (confluence of San Juan and Glen Canyon arms of Lake Powell) and Washington coun- ties; Nevada and Arizona; 9 (i). The type is from the Colorado River (Utah?), Palmer sn, 1870 (US!). Brickellia californica Gray [Bulbostylis californica T.& G.; Coleosanthus californicus (T. & G.) Kuntze]. Subshrubs, mainly 5-10 dm tall, the branchlets whitish to brownish; leaves alternate, petiolate, the blades 1.7-5.2 cm long, 1.3-4.5 cm wide, cordate-ovate to ovate or orbicular, truncate to cordate ba- sally, crenate-serrate, rounded to obtuse api- cally, the veins not prominent, glandular- scabrous; heads clustered in a leafy-bracteate panicle; sessile or shortly pedunculate; in vol ucres 5.5-8 mm high, 4-7 mm wide, the out- er bracts very short, rounded apically, few veined, the inner long and slender, often suf- fused with red or purple, glabrous; flowers 8-18; achenes straw colored, 2.5-3.5 mm long. Canyons and rock outcrops at 825 to 2135 m in Garfield, Kane, San Juan, Utah, and Washington counties; Colorado to California and south to Texas and Mexico; 15 230 Great Basin Naturalist Vol. 43, No. 2 Brickellia grandiflora (Hook.) Nutt. [Eu- patorium grandiflorum Hook.]. Perennial herb, from a caudex and taproot, the stems green to straw colored, 2.5-9.5 dm tall; leaves alternate, petiolate, the blades sagit- tate to cordate-ovate, 1.5-9 (11) cm long, 0.6-6.5 cm wide, cordate to truncate basally, serrate to doubly so, attenuate to acuminate apically, the veins not prominent, minutely puberulent or hirtellous; heads several to nu- merous in short corymbose panicle, com- monly reflexed; involucres 7-12 mm high, 6-10 mm wide, the outer bracts lance-acumi- nate, the inner abruptly acuminate, pu- berulent dorsally; flowers mostly 20-40 (70); achenes brown to black, 3.5-4.5 mm long, hirtellous. Pinyon-juniper, mountain brush, ponderosa pine, aspen, Douglas fir-white fir, spruce, and bristlecone pine communities at 1640 to 3200 m in Beaver, Duchesne, Gar- field, Iron, Juab, Kane, Salt Lake, San Juan, Tooele, Utah, and Washington counties; Washington east to Missouri, south to Mexi- co; 34 (vii). Brickellia hngifolia Wats. [Coleosanthus longifolia (Wats.) Kuntze]. Shrubs, with stems and white to tan bark, mainly 10-15 dm tall; leaves alternate, sessile or subsessile, 1.2-13.5 cm long, 3-8 mm broad, lance-linear to lance-elliptic, obtuse to acute basally, attenu- ate apically, the veins not prominent. glabrous, glandular-resinous; heads numerous in panicles; involucres 3.4-6.2 mm high, 2.3-4 mm wide, the outer bracts ovate, acute, the inner, longer and slender, glabrous; flowers 3-5; achenes 1.8-2.4 mm long, brown, glabrous. Canyon bottoms, stream margins, seeps, and hanging gardens at 750 to 1590 m in Emery, Garfield, Grand, Kane, San Juan, Washington, and Wayne counties; Cali- fornia, Nevada, Arizona; 23 (viii). Brickellia microphylla (Nutt.) Gray [Bul- hostylis microphyllus Nutt.]. Shrubs or sub- shrubs, with tan to whitish bark, mainly 2-7 dm tall; leaves alternate, shortly petiolate to subsessile or sessile, 3-14 (20) mm long, 1-9 (12) mm wide, ovate to suborbicular, toothed to entire, commonly glandular-villous or -his- pidulose, the veins not especially prominent, rounded to acute apically; heads solitary or few at tips of branches, racemosely arranged in leafy-bracteate panicles; involucres 7-10.3 mm high, 4-8.5 mm wide, the outer bracts oval to ovate, with thickened glandular tips, the inner often lacking glands and more or less 3-lobed or 3-veined; flowers 8-18; ach- enes 3.5-4.3 mm long, blackish, hirtellous or glabrous. Two distinctive phases, which have been treated at specific level, are present in Utah. There is justification for treating them at specific rank, but they are similar in vege- tative features and general aspect. Flowers 8-11 per head; involucres 7-10 mm long; 4-7.5 mm wide; plants of the Green, Colorado, and Virgin river systems B. microphylla var. scabra Flowers (12) 17-18 per head; involucres 8.5-10.3 mm long, 6.5-10 mm wide; plants of the Great Basin B. microphylla var. watsonii Var. scabra Gray Blackbrush, rabbitbrush, sagebrush, shadscale, Grayia, greasewood, juniper, and pinyon-juniper communities mainly on sandstone outcrops at 885 to 2170 m in Daggett, Duchesne, Emery, Garfield, Grand, Kane, San Juan, Uintah, and Wash- ington counties; Colorado, Nevada, Arizona. Our material is uniformly hispidulose-glandu- lar along upper stems at least, and has 8-11 flowers per head; 35 (xi). Note: A peculiar specimen from San Juan County (Anderson A-6 BRY) has heads nearly all clustered at branch tips. Var. watsonii (Robins.) Welsh comb. nov. [based on Brickellia watsonii Robins. Mem. Gray Herb. 1:42. 1917]. Sagebrush, shadscale. mountain brush, and juniper communities at 1525 to 2440 m in Juab, Millard, Sevier, Tooele, and Utah counties; Nevada and Cali- fornia. All modern floras distinguish B. micro- phylla by its heads "about 22-flowered." Our material fits well within the concept of B. watsonii Robins., which has heads "18-flow- ered." Specimens from the Great Basin of Utah are uniformly 18-flowered, except in depauperate heads that vary downward to 12 flowers per head. Stems are villous to glandu- lar-villous, with the type of B. watsonii Rob- ins. (Watson 494 US!) at the villous end of a cline; 7 (ii). Brickellia oblongifolia Nutt. Subshrubs or subherbaceous, with green to tan branches. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 231 mainly 1-5.5 dm tall; leaves alternate, sessile or nearly so, 0.9-4 cm long, 1-11 (15) mm wide, elliptic to oblong, or lance-oblong, en- tire or essentially so, glandular-hispidulous, the veins not especially prominent, acute to attenuate or obtuse apically; heads solitary and terminating branches, or corymbosely ar- ranged; involucres 10.8-15 mm long, 12-22 mm wide, the bracts all acute to acuminate, glabrous or glandular to glandular- puberulent; flowers (11) 26-40 (50); achenes 4.8-5.8 mm long, blackish, hispidulous. Gray- ia, shadscale, rabbitbrush, blackbnish, desert almond, juniper, pinyon-juniper, and pon- derosa pine communities at 1280 to 2500 m in Beaver, Duchesne, Emery, Garfield, Juab, Kane, Millard, San Juan, Sevier, Uintah, Utah, Wasatch, Washington, and Wasatch coimties; British Columbia to Montana, south to California, Arizona, and New Mexico. Our material is assignable to var. linifolia (D.C. Eaton) Robins. [B. linifolia D.C. Eaton, type from Jordan Valley, American Fork] which is distinguished by its achenes being his- pidulous, not glandular-hispidulous or glandular. The segregation is tenuous at best; 41 (x). Calycoseris Gray Annual subscapose or caulescent herbs, with milky juice, from taproots, beset with tacklike stipitate glands above; leaves mostly basal, pinnately parted; heads solitary or few on leafy-bracteate peduncles; involucral bracts in 2 series, herbaceous, the inner with hyaline margins; receptacle with capillary bristles; corollas all raylike, yellow or white tipped; achenes fusiform, 5- or 6-ribbed, ta- pering to a short beak, this produced apically into a low denticulate cup; pappus abundant, white, of barbellate capillary bristles falling attached. 1. Rays white, with pink or purple dots or streaks dorsally; stipitate glands pale ... C. wrightii — Rays yellow; stipitate glands purple C. parryi Calycoseris parryi Gray Annual herbs, mainly 0.7-3 dm tall, the stems simple or with spreading branches; leaves basal and al- ternate along stem, pinnately parted, the lobes linear, reduced and entire above, glabrous except for a few tangled long hairs on lower surface; peduncles mainly 0.5-4 cm long, clad with tacklike, long-stipitate, purple or purplish-black glands; involucres 11-15 mm high, 8-14 mm wide (when pressed), the bracts linear-subulate to lance- subulate, more or less stipitate-glandular, at- tenuate apically; rays yellow, 10-20 (25) mm long; pappus surpassing the achene. Creosote bush and Joshua tree communities, reported for Utah by Munz (1959. Calif. Flora p. 1300); to be expected in Washington County; California and Arizona. Measurements are from Arizona and California materials; 0 (0). Calycoseris wrightii Gray Annual herbs, mainly 1.4-4 dm tall, the stems commonly with spreading branches; leaves basal and al- ternate along the stem, pinnately parted, the lobes linear, reduced and subentire upward, glabrous except for a few long tangled hairs on lower surface; peduncles mainly 0.3-5 cm long, clad with tacklike long-stipitate pale glands; involucres 12-17 mm long, 12-20 mm wide, the bracts linear-subulate to lance- subulate, more or less stipitate-glandular, at- tenuate apically, rays 10-25 mm long, white, with pink or purple markings dorsally; pap- pus shorter than achene. Creosote bush and Joshua tree communities in Washington County; California, Nevada, Arizona; 1 (0). Carduus L. Biennial or annual herbs with taproots, the juice watery; stems erect, simple or branched; leaves alternate simple, pinnatifid to bipinnatifid or merely pinnately lobed, of- ten decurrent, spiny; heads solitary or few, borne in corymbose cymes; involucral bracts imbricated in several series, spine tipped; re- ceptacle hemispheric, densely bristly; disk flowers only present, perfect, red-purple, with long slender lobes; pappus of barbellate bristles; style branches connate, shortly hairy at base of branches; achenes compressed. Carduus nutans L. Nodding Thistle; Musk Thistle. Rank biennial or annual herbs, most- ly 0.6-20 (25) dm tall; stems arachnoid-to- mentose to glabrate; leaves alternate, decur- rent, 3-40 cm long, 0.5-20 cm wide (or 232 Great Basin Naturalist Vol. 43, No. 2 more), lance-linear to elliptic, glabrous, or to- mentose along veins beneath; heads com- monly solitary, nodding; involucres 20-30 mm long, 30-80 mm wide, the bracts 2-8 mm wide, ovate-lanceolate to lanceolate, glabrous or nearly so, spinose tipped, at least the outermost reflexed near the middle, the midrib prominent; flowers red-purple; ach- enes 3.5-4.5 mm long, smooth, marked with vertical lines, umbonate. Disturbed sites along roads and in fields and pastureland at 1340 to 2440 m in Daggett, Juab, Salt Lake, Sanpete, and Utah counties, and probably universal; introduced Old World plants, now widely established in the United States; 15 Centaurea L. Annual, biennial, or perennial herbs with taproots or rhizomes, the juice watery; stems erect or ascending; leaves alternate, entire to pinnatifid; heads solitary, or few to numer- ous, discoid; involucral bracts imbricate in several series, spine tipped or some of them enlarged and with scarious or hyaline erose to lacerate or pectinate appendages; recep- tacle bristly; flowers all tubular, perfect, or the marginal ones sterile and falsely sub- radiate; purple, blue, yellow, pink, or white; pappus of bristles, scales, or none; style branches more or less connate, with a thick- ened often hairy ring at the base; achenes obliquely or laterally attached to receptacle. Note: This is a large genus, mainly of the Mediterranean region of the Old World, but with some indigenous to North America, Aus- tralia, and South America. All of ours are in- troduced, and the potential for other in- troductions in this remarkable genus is great. In Flora Europaea, our species are treated within three genera: Amberboa (Pers.) Less. (C. moschata L.), Acroptilon Cass (C. repens L.), and Centaurea for the others. 1. Involucral bracts definitely spine tipped, at least some with spines 1-20 mm long 2 — Involucral bracts definitely not spine tipped, or, if shortly spinose as in C. maculosa and C. scabiosa, the heads 6-25 mm wide .' 5 2(1). Stem definitely winged, the leaf bases decurrent; pappus present (central flow- ers, at least) 3 — Stems angled, not winged; pappus none 4 3(2). Apical spine of involucral bract 5-9 mm long; plants arachnoid when young; flowers all with evident pappus C melitensis — Apical spine of involucral bract 11-20 mm long; plants persistently tomentose; flowers in center only with a pappus C. solstitialis 4(2). Apical spine of bracts 5-15 mm long or more C. calcitrapa — Apical spine of bracts 1-4 mm long C. virgata 5(1). Leaves entire or merely toothed, not pinnatifid 6 — Leaves pirmatifid or deeply pinnately lobed 8 6(5). Leaves linear to lance-linear, entire or nearly so, less than 1 cm wide C. cyanus — Leaves various, but, if as above, plants rhizomatous 7 7(6). Plants rhizomatous; leaves mainly 2-10 mm wide; pappus evident, 6-11 mm long C. repens — Plants not rhizomatous; leaves 6-15 mm wide; pappus 2-5 mm long C. jacea 8(5). Leaves merely pinnately lobed; involucral bracts entire or nearly so C. moschata — Leaves pinnately divided, the lobes linear to narrowly oblong; involucral bracts pectinately lobed 9 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 233 9(8). Involucres 15-25 mm wide; lobes of leaves often again toothed or lobed C. scabiosa — Involucres mainly 6-10 mm wide; lobes of leaves usually entire C. maculosa Centaurea calcitrapa L. Star-thistle. Bien- nial herbs, from taproots, the stems usually branched, 1-8 dm tall, arachnoid-villous to glabrate; leaves 0.5-4.5 cm long, pinnatifid, the lobes linear to oblong, attenuate, or the upper ones entire; heads few to numerous; involucres urn shaped, 10-18 mm high, mainly 8-12 mm wide, the bracts weakly spinose-ciliate, with a stout apical spine mainly 5-30 mm long; flowers few, purple; pappus none. Roadside weeds, Utah County (Wadley & Holmgren 381 UT); introduced from Eurasia; 1 (0). Centaurea cyanus L. Bachelor's Button; cornflower. Annual or biennial herbs from taproots, the stem usually branched, mostly 1-8 (12) dm tall, arachnoid-tomentose; leaves 2-10 (13) cm long, 1-8 mm wide, entire or some with slender lobes, attenuate; heads few to numerous; involucres hemispheric, 10-16 mm high, 10-23 mm wide, the bracts with a tapering pectinate or fringed tip, often purplish suffused, the central apical tooth not especially spinose; flowers several, blue, purple, pink, or white, the marginal ones en- larged, irregular; pappus 2-3 mm long. Culti- vated ornamental, now established in dis- turbed sites in Cache, Salt Lake, Tooele, Utah, Wasatch, and Washington counties; ad- ventive from Europe; 6 (0). Centaurea jacea L. Perennial herbs from taproots, the stems simple or branched from the middle, mostly 5-12 dm tall, glabrous or somewhat arachnoid; leaves entire or toothed to shallowly lobed, the basal ovate to lanceo- late, petiolate, becoming smaller upward; heads few to numerous; involucre 12-18 mm high, 12-15 mm wide, ovoid, the bracts with orbicular appendages, scarious, brown, dark- er in middle, the outer denticulate to pecti- nate-lacerate, the inner less so and often bi- fid; flowers purple or white, the outer more or less radiate; pappus none or very short. Cultivated ornamental, now established in Salt Lake County; adventive from Europe; 0 (0). Note: The large headed C. montana L., is cultivated in Utah. It has wedge-shaped in- volucral bracts and decurrent large leaves. Centaurea maculosa Lam. Biennial or short-lived perennial, the stems simple or commonly branched above the middle, mainly 3-10 (15) dm tall, tomentose and sparingly scabrous-puberulent; leaves 1-9 cm long, pinnatifid, the lobes linear to lanceolate or oblong, entire or variously toothed or lobed, reduced and bracteate in the in- florescence; heads few to many, hemispheric to vase shaped; involucres 10-13 mm high, 10-13 mm wide, the bracts with short dark pectinate tip, the central tooth produced as a spine to 0.5 mm long; flowers pink or pur- plish, rarely white, the marginal ones radiate; pappus to 2 mm long, rarely lacking. Road- sides in Beaver, Juab, and Tooele counties; adventive from Europe; 3 (i). Centaurea melitensis L. Annual or bien- nial, the stems sparingly branched from middle or below, 1.5-8 dm tall, winged by decurrent leaf bases; basal and lower cauline leaves oblanceolate, toothed to lyrate-pin- natifid or sinuately lobed, reduced upward, finally entire; heads solitary, terminating branches, or 2 or 3 in clusters; involucres 8-15 mm high, 8-12 mm wide, tapering api- cally, the middle and outer bracts spine tipped, the spines 5-8 mm long; flowers yel- low, all alike; pappus 1.5-3 mm long. Adven- tive Old World species of disturbed sites in Salt Lake County (Without collector UT); 1 (0). Centaurea moschata L. [Amberboa mos- chata (L.) DC.]. Annual herbs; simple or sparingly branched, mainly 3-7 dm long, sparingly tomentose; leaves 1-9.5 cm long, 1-3 cm wide, pinnatifid, the lowermost pet- iolate, becoming sessile upward; heads soli- tary, on peduncles 8-15 cm long or more; in- volucres vase shaped, 12-14 mm high, 18-22 mm wide, the bracts oval, with purplish mar- gins, only the inner with broad, reflexed, en- tire appendage; flowers pink; pappus shorter to about equaling the achenes. Cultivated or- namental, escaping and persisting in Wash- ington County; adventive from Asia; 1 (0). Centaurea repens L. Russian Knapweed. [C. picris Pallas ex Willd.; Acroptilon repens 234 Great Basin Naturalist Vol. 43, No. 2 (L.) DC.]. Perennial rhizomatous herbs, most- ly 3-8 dm tall, arachnoid-tomentose to gla- brate; leaves in a basal rosette and cauline, the basal leaves often withered by flowering time, the cauline mainly 1-6 cm long, 2-12 mm wide, entire or serrate; heads few to nu- merous, terminating branches; involucre 9-15 mm high, 5-12 mm wide, more or less um shaped, middle and outer bracts broad, glabrous, with broader rounded, subentire hyaline tips, the inner bracts narrow, taper- ing, and with plumose hairy tips; flowers pink to purplish, all alike; pappus bristle sub- plumose, 6-11 mm long. Introduced Old World primary noxious weed, now widely es- tablished at 1220 to 2380 m in Cache, Dag- gett, Duchesne, Emery, Garfield, Grand, Kane, San Juan, Salt Lake, Tooele, Uintah, and Utah counties; widespread in North America; adventive from Eurasia; 28 (ii). Centaurea scabiosa L. Perennial herbs, mostly 5-15 dm tall, scabrous-puberulent; leaves 4-20 cm long or more, the lowermost long-petiolate, once to twice pinnatisect, the segments linear to oblong, entire or dentate- serrate to lobed, the upper pinnately divided, sessile; heads few to several, terminating branches; involucres 13-20 mm high, 18-25 mm wide, ovoid-globose; bracts ovate, glabrous or arachnoid, the appendages triangular-ovate, brown or black, with pale brown teeth; flowers purple, alike or nearly so; pappus 4-5 mm long. Cultivated orna- mental, persisting and escaping, Salt Lake County and probably elsewhere; adventive from Europe; 2 (0). Centaurea solstitialis L. Annual or bien- nial, grayish tomentose, the stems 1-6 (10) dm tall, evidently winged; leaves mainly 1-12 (20) cm long, 0.1-3 (5) cm wide, the basal ones lyrate to pinnatifid, the cauline ones progressively smaller and entire up- wards, linear to linear-subulate; heads few to numerous, terminating branches; involucres 8-15 mm high, 7-15 mm wide, urn shaped, the middle and outer bracts with central apical spines 10-20 (30) mm long, the inner with a small hyaline appendage; flowers yel- low, all alike; pappus of marginal flowers none, that of the central ones 3-5 mm long. Roadsides and abandoned fields at 915 to 1900 m in Wasatch, Washington, and Weber counties; adventive from Europe; 4 (i). Centaurea virgata Lam. Perennial, from a caudex, more or less grayish tomentose, the stems 4-9 dm tall, branched above; leaves mainly 0.5-15 cm long, 0.1-6 cm wide, the basal ones petiolate, once to twice pinnately divided, the lobes linear, these often again toothed or lobed; cauline leaves smaller, ses- sile, and lobed to entire; heads several to nu- merous, terminating short branches; in- volucre 7-10 mm high, 3-5 mm wide, the bracts pale or suffused with red or purple, with a slender apical spine 1-2 mm long; flowers pink; pappus about 1.5 mm long. Roadsides and other disturbed sites in Grand, Juab, and Utah counties at 1525 to 1830 m; adventive from Eurasia; 5 (i). Chaenactis DC. Annual, biennial, or perennial herbs, from taproots; leaves alternate or mainly basal, pinnately dissected to entire; heads solitary or few to several, borne in corymbose cymes, discoid, the flowers white, or cream to pink, all perfect, the marginal ones sometimes en- larged and raylike; involucral bracts in 1-3 series, herbaceous; receptacle flat, naked; pappus of 4-20 hyaline scales; style branches slightly compressed; achenes clavate, terete or more or less compressed. Stockwell, p. 1940. A revision of the genus Chaenactis. Contr. Dudley Herb. 3:89-168. 2(1). Plants perennial from a simple or branching caudex; 2-9 cm tall; stemless or with few short internodes; of high elevations C. alpina Plants annual or biennial, rarely perennial, the caudex seldom developed; stems mainly 10-30 cm tall, or, if less, plants definitely not perennial; dis- tribution usually of middle and lower elevations 2 Basal rosette well developed; plants biennial or short-lived perennials; pappus scales 10-16 C. douglasii April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 235 — Basal rosettes poorly, if at all, developed; plants annual; pappus scales 4 or 5 (rarely 8) 3 3(2). Lower and upper cauline leaves simple, the middle ones few lobed; Washington County C. fretnontii — Lower, middle, and upper leaves pinnately divided, or only the uppermost simple 4 4(3). Heads mostly 15-22 mm high; flowers pink, much surpassing the involucre; anthers included C. macrantha — Heads mostly 8-10 mm high; flowers white or cream, only slightly surpassing the involucre; anthers exserted 5 5(4). Involucral bracts blunt or nearly acute apically; plants widely distributed C. stevioides — Involucral bracts long-attenuate and bristle tipped apically; plants of Washington and Millard counties C. carphoclina Chaenactis alpina (Gray) Jones Alpine Dusty-maiden. [C. douglasii var. alpina Gray]. Perermial, from a simple or branched, sometimes soboliferous caudex, 3.5-9 cm tall; stems with few contracted internodes, very short, or not developed; leaves 1.3-5 cm long, pinnately divided, the lobes again toothed or lobed, 1-7 mm long, gray tomen- tose to glabrate; heads solitary or sometimes 2, the peduncles tomentose or glandular, 0.5-6 cm long; involucres (7.5) 10-13 mm long, (8) 10-17 mm wide, the bracts often suffused with purple, glandular or tomentose; corolla purplish to white, glandular or spar- ingly tomentose; pappus of 10 oblong-spatu- late rounded hyaline scales, in 2 series; ach- enes 6-8 mm long, hairy. Boulder stripes and talus in alpine tundra or upper montane com- munities at 2980 to 3965 m in Duchesne, Salt Lake, Summit, and Utah counties; Oregon to Montana, California and Colorado. Our ma- terials are separable into two more or less distinctive phases; a glandular phase, with distribution mainly in the Wasatch Moun- tains, which is var. alpina [including C. ru- bella Greene; C. alpina var. rubella (Greene) Stockwell], and a tomentose phase, mainly from the Uinta Mountains, which might be assignable to var. leucopsis (Greene) Cock- erell [C. leucopsis Greene]. More work is nec- essary, including evaluation of the type speci- men of var. leucopsis; 10 (i). Chaenactis carphoclina Gray Annual, from a taproot, 6-28 (40) cm tall; stems well developed, more or less flexuous; leaves 0.8-5.6 cm long, mealy-puberulent, 1- to 2- pinnatifid, the segments linear-filiform, 1-20 mm long; heads few to numerous, on slender farinose to glandular peduncles 0.4-3 cm long; involucres 6-9 mm high, 6-15 mm wide, the bracts lance-attenuate into slender, bristlelike tips, glandular; flowers white to cream; pappus of central flowers usually of 4 lance-acuminate scales, those of marginal flowers sometimes shorter; achenes 3.5-4.5 mm long, hairy. Larrea community at 850 to 1000 m in Washington County; California, Nevada, Arizona; 10 (0). Chaenactis douglasii (Hook.) H. & A. Douglas Dusty-maiden. [Hymenopappus douglasii Hook.; C. achilleaefolia H. & A.; C. douglasii var. achilleaefolia (H. & A.) A. Nels.; C. douglasii var. montana Jones; C. brachiata Greene, type from Springdale; C. brachiata var. stansburyi Stockwell, type from Stansbury Island]. Biennial or short- lived perennial, from a taproot, seldom with a caudex, mainly 5-50 (60) cm tall, sparsely to densely tomentose; stems with few to many well developed internodes; leaves 0.6-12 (15) cm long, 1-3 pinnatifid, the lobes 1-3 cm long, tomentose to glabrate; heads solitary or several in a corymbose cyme; in- volucre 7-16 mm high, 8-25 mm wide, the bracts glandular to glandular-tomentose, ob- long to narrowly oblanceolate or linear, blunt apically; flowers white to pink; pappus of 10-16 scales in 2 series; achenes 6-8 mm long, hairy. Shadscale, sagebrush, pinyon- juniper, mountain brush, ponderosa pine, white fir, Douglas fir, aspen, and limber pine communities at 1340 to 3050 m in all Utah 236 Great Basin Naturalist Vol. 43, No. 2 counties; British Columbia to Montana, south to California, Arizona, and Colorado. It does not seem reasonable to attempt to segregate our materials into varieties. The variability apparently does not demonstrate geographic correlation; 132 (xx). Chaenactis fremontii Gray Annual or win- ter annual, from a taproot, 10-25 (40) cm tall, glabrate or sparingly tomentose when young; leaves 0.6-6.5 cm long, the lower and upper simple, linear, the middle few lobed, glabrous; heads solitary to several on tomen- tose to glabrate (glandular?) peduncles 1-5 cm long; involucres 8-10 mm high, 10-12 mm wide, glabrous or tomentose, attenuate but not caudate; flowers white to pinkish, the outer ones enlarged; pappus of central flow- ers of 4 scales; achenes hairy. Creosote bush and Joshua tree communities at 670 to 885 m in Washington County; Arizona, Nevada, California; 2 (0). Chaenactis macrantha D.C. Eaton Annual or winter annual, from a taproot, mainly 6-25 cm tall, branching from the base or simple, floccose-tomentose to glabrate; leaves 0.5-5 cm long, 1- to 2-pinnatifid, the lobes to 1 cm long, broad, floccose to glabrate; heads solitary to several, on tomentose peduncles 0.5-5 cm long; involucres 12-17 mm high, 8-22 mm wide, the bracts oblong-lanceolate, rather abruptly short-acuminate, tomentose; corollas pink to white, all about alike; an- thers included; pappus of 4 linear-oblong scales and 2-4 short outer ones or these lack- ing; achenes hairy. Shadscale, pinyon-juniper, creosote bush, and blackbrush communities at 885 to 2135 m in Beaver, Juab, Kane, Mil- lard, Tooele, and Washington counties; Cali- fornia, Nevada, Arizona; 17 (iii). Chaenactis stevioides H. & A. Annual or winter annual, from a taproot, mainly 4-42 cm tall, branching from the base or simple, more or less tomentose; leaves 0.3-10 cm long, 1-2 pinnatifid, the lobes to 2.5 cm long, linear to oblong, sometimes all or nearly all simple in depauperate specimens; heads soli- tary to several on glandular peduncles 0.3-3 cm long; involucres 6-11 mm high, 8-22 mm wide, the bracts oblong-lanceolate to linear, acute to shortly acuminate apically, glandu- lar; corollas white to cream, the outer ones enlarged; pappus of 4 oblong-lanceolate scales; achenes hairy. Creosote bush, black- brush, mat-atriplex, shadscale, indigo bush, and juniper communities at 915 to 1891 m in Beaver, Carbon, Duchesne, Emery, Garfield, Grand, Juab, Kane, San Juan, Washington, and Weber counties; Wyoming south to Ne- vada, west to California; 63 (vi). Chamaechaenactis Rydb. Perennial scapose herbs from a long-pilose caudex, clothed with marcescent leaf bases, and taproot, with watery juice; leaves all bas- al, petiolate, simple; heads solitary; in- volucres turbinate, the bracts subequal or the outer shorter; receptacle naked; rays none; disk flowers perfect, fertile, cream colored to pink; pappus of hyaline scales; style branches flattened, papillate; achenes 4-angled, hairy. Chamaechaenactis scaposa (Eastw.) Rydb. [Chaenactis scaposa Eastw.] Plants 2-9 cm tall, the scapes long-villous; leaves petiolate, the blades 0.4-1.8 cm long, 3-13 (15) mm wide, lance-oblong to ovate, to oval or or- bicular, obtuse to rounded apically, obtuse to truncate basally, villous beneath, strigose to strigulous or villous above; heads solitary; in- volucre 7-17 mm high, 10-23 mm wide, the bracts oblong or linear-oblong, the outer densely villous, green or suffused with red- purple, the margin hyaline; corollas cream to pink; pappus scales oblanceolate-spatulate, rounded; achenes black, hirsute-pilose. Shad- scale, galleta, pygmy sagebrush, mountain brush, pinyon-juniper, and ponderosa pine communities at 1580 to 2565 m in Carbon, Duchesne, Emery, Garfield, Grand (?), and San Juan counties; Arizona and Colorado; 40 (V). Chamomilla S.F. Gray Annual herbs, aromatic in some; leaves al- ternate, 2- or 3-pinnatifid, with linear fili- form ultimate segments; heads radiate or dis- coid, solitary or corymbose; involucral bracts greenish-chartaceous, the margins hyaline, in 2 or 3 series, subequal to imbricate; recep- tacle conic, hollow, naked; marginal flowers pistillate; rays (when present) white, the cen- tral disk flowers perfect and fertile, the style branches truncate, tufted-hairy apically; pap- pus a short crown of minute scales, or vesti- gial or lacking; achenes subcylindric, the ven- tral face with 3-5 narrow ribs, the dorsal face smooth and convex. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 237 1. Heads radiate; disk corollas 5-lobed; involucre 11-25 mm in diameter C. recutita — Heads discoid; disk corollas 4-lobed; involucre 4-10 mm in diameter ... C. suaveolens Chamomilla recutita (L.) Rauschert Chamomile. [Matricaria chamomilla L.]. An- nual herbs; stems 0.2-4 (6) dm tall, erect or ascending, branched above; herbage glabrous or puberulent; leaves 2-6 cm long; heads soli- tary or more commonly few to many and corymbosely arranged; involucres saucer shaped, 3-4 mm high, 11-25 mm wide, the bracts subequal, the margins broadly hyaline, the midstripe greenish to brownish; rays 10-20, white, 4-10 mm long. Moist disturbed soils at low to moderate elevations in Salt Lake and Wasatch counties; adventive from Europe; 2 (0). Chamomilla suaveolens (Pursh) Rydb. [Matricaria inatricarioides (Less.) Porter]. An- nual herbs; stems 0.4-4 dm tall, erect or as- cending, branched from the base or simple; herbage glabrous or pubescent; leaves 1-5 (9) cm long; heads few to many, paniculately ar- ranged; involucres saucer shaped, 2-6 mm high, 4-10 mm wide, the bracts subequal to somewhat imbricate, the margins hyaline, the midstripe greenish; rays lacking; disk flowers 4-lobed. Disturbed sites at 1310 to 2810 m in Box Elder, Cache, Carbon, Rich, Salt Lake, Sevier, Utah, and Weber counties; adventive from Europe; 16 (0). Chrysanthemum L. Perennial herbs from a rhizome or a cau- dex, with watery juice; stems erect or nearly SO; leaves alternate, serrate to pinnatifid; heads solitary or few to numerous in open corymbose clusters; involucral bracts imbri- cate, in 2-4 series, greenish or straw colored, the margins brownish-scarious; receptacle naked; ray flowers white, numerous, pistil- late, fertile, or lacking; disk flowers numer- ous, perfect, fertile, yellow; pappus lacking or a short crown; style branches flattened; achenes several nerved, beakless, glabrous. 1. Leaves finely serrate; heads usually numerous, small, commonly rayless..C. balsamita — Leaves coarsely serrate or pinnatifid; heads larger, fewer, commonly with rays 2 2(1). Heads solitary or few; involucres 7-10 mm high; rays 1-2 cm long; leaves ser- rate to more or less once pinnatifid C. leucanthemum — Heads several to numerous; involucres 3-4.5 mm high; rays 2-6 mm long C. parthenium Chrysanthemum balsamita L. Costmary. [Balsamita major Desf.]. Perennial herbs, from a caudex, commonly 5-10 (12) dm tall; stems strigose, at least above; leaves petiolate below, sessile or subsessile above, the blades 0.9-10 (15) cm long, 0.6-5 (8) cm wide, ellip- tic to oblanceolate, finely serrate, strigose; heads numerous, corymbose; involucres 3.7-4.6 mm high, 6-8 mm wide, the bracts oblong, sparingly strigose, the tip hyaline; ray flowers (when present) 4-6 mm long. Fields, roadsides, and cemetaries at 1370 to 2135 m Salt Lake, Summit, Tooele, and Utah counties; escaped from cultivation, now widely established in the United States; 5 (i). Chrysanthemum leucanthemum L. Oxeye- daisy. [Leucanthemum vulgare Lam.] Pe- rennial rhizomatous or subrhizomatous herbs, commonly 2-8 (10) dm tall; stems glabrous or nearly so, mainly simple; leaves petiolate be- low, becoming smaller and sessile above, the blades 0.8-5 cm long, oblanceolate to obo- vate or linear, serrate, crenate, or pinnately lobed, glabrous or villosulose; heads solitary; involucres 7-10 mm high, 15-23 mm wide, the bracts lance-ovate to oblong-linear, with brown margins, hyaline apically; rays mainly 15-30, white, 10-22 mm long; pappus none. Roadsides, fields, and other disturbed sites at 1525 to 2135 m in Salt Lake, Utah, Wasatch, and Weber counties; widespread in North America; adventive from Eurasia; 6 (0). Chrysanthemum parthenium (L.) Bemh. [Matricaria parthenium L.; Leucanthemum parthenium (L.) Gren. & Godron; Pyrethrum parthenium (L.) Sm.; Tanacetum parthenium (L.) Schultz-Bip.]. Perennial herbs with cau- dex and taproot; commonly 3-9 dm tall; 238 Great Basin Naturalist Vol. 43, No. 2 stems glabrous, or puberulent above; leaves Chrysothamnus Nutt. petiolate, becoming smaller, but still petio- ^ i -.l i.-. u i .u r i. r , , 1 1 ? r> ,- o 1 nV. A c Shrubs with white bark, or the suriace ob- late above, the blades 0.5-8 cm long, 0.6-4.5 J, ^ ^ ^V.- U 1 J 1 .-. , 1 , , 1 1 scured by a tomentum, this orten giandular- (6) cm wide, pinnatirid or doubly so; heads . \ ,. . ,. ^ ui ^ ^ ' ' ^ ■' resinous; leaves alternate, linear to oblong, or several to numerous, the inflorescence cor- ^^^^^^^^^^ ^^^^^^e, entire; heads white or yel- ymbose; involucres 3-4.5 mm high, 7-10 mm j^^ ^^^^^^^ ^^^^.^.^^ ^^ contracted to open wide, the bracts oblong, with a dark center, paniculate inflorescences; flowers perfect, otherwise scarious except the tip hyaline; £gj.tile; involucral bracts imbricate, more or rays 10-20, white, 4-8 mm long; pappus a \q^^ keeled, in 4 or 5 vertical or obscure crown or none. Cultivated ornamental, es- ranks, chartaceous or coriaceous, or the tip caping and persisting at 1525 to 1950 m in herbaceous; receptacle naked; style branches Carbon, Salt Lake, Utah, and Weber coun- flattened; achenes slender, flattened, angled, ties; widely established in the United States; or terete, hairy or glabrous; pappus of nu- adventive from Europe; 5 (0). merous capillary bristles. 1. Flowers white; leaves terete; plants of western tier of counties (except Iron and Washington) C. alhidus — Flowers yellow; leaves various, but, if terete, of Washington County or rarely elsewhere 2 2(1). Leaves terete, resinous punctate; stems more or less fastigiate; plants of Wash- ington County C. paniculatus — Leaves commonly more or less flattened, resinous-punctate or not; stems not especially fastigiate; plants of broad or other distribution 3 3(2). Stems obscured by a tomentum, this often impregnated with resinous-glandular material 4 — Stems glabrous or puberulent, the surface readily apparent 5 4(3). Involucral bracts long-attenuate, membranous; inflorescence more or less race- mose C. parryi — Involucral bracts obtuse to acute, rarely attenuate, but, if so, chartaceous; in- florescence cymose C. nauseosus 5(3). Leaves lanceolate to lance-oblong, not contorted; shrubs mainly 6-20 dm tall; plants of the Uinta and Navajo basins C. linifolius — Leaves linear, oblong, or lanceolate, but, if lanceolate, twisted and shrubs mainly less than 6 dm tall; distribution various 6 6(5). Achenes hairy 7 — Achenes lacking hairs, sometimes glandular, or, if sparingly hairy, the in- volucre over 10 mm long 8 7(6). Involucral bracts acuminate-cuspidate; leaves 1-2 mm wide C greenei — Involucral bracts acute to obtuse; leaves various C viscidiflorus 8(6). Flowers 10-12 mm long, surpassed by the pappus; plants of Emery, Wayne, and San Juan counties C. pulchellus — Flowers 7-9 mm long, surpassing or subequal to the pappus; distribution various 9 9(8). Involucral bracts strongly ranked; involucres 9.2-13 mm long C. depressus — Involucral bracts not strongly ranked; involucres 6.2-7.5 mm long C. vaseyi April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 239 Chrysothamnus albidus (Jones) Greene Alkali Rabbitbrush; White Rabbitbush. [Bige- lovia alhida Jones]. Shrubs, mainly 5-10 dm tall, more or less fastigiately branched, white barked, glabrous, resinous-viscid, aromatic; leaves 0.5-3.5 cm long, terete, 0.5-1 mm thick, glandular-punctate, mucronate, crowded, often with axillary fascicles; heads clustered at branchlet apices; involucres 6.8-9 mm high, 3-7 mm wide, the bracts ob- scurely 4- to 5-ranked, the outer ones lance- ovate, thickened in lower half, abruptly sub- ulate-attenuate, the inner oblong, acuminate to acute, the margin hyaline, glandular to to- mentose; corollas white, 6-7.5 mm long; ach- enes 4-4.5 mm long, pilose and glandular; pappus abundant. Local in salt grass, pickle- weed, and alkali-saccaton communities at 1450 to 1650 m in Beaver, Box Elder, Juab, Millard, and Tooele counties; California, Ne- vada; 8 (iii). Chrysothamnus depressus Nutt. Dwarf Rabbitbrush. Low, spreading shrubs, the as- cending to erect, subherbaceous stems 0.6-3 dm tall, white barked, scabrous-puberulent or glandular-puberulent; leaves 0.4-2 cm long, 1-4 (5) mm wide, flat, narrowly lanceolate to oblanceolate or spatulate, flat, scabrous- puberulent, obtuse, rounded or sharply apicu- late; heads clustered at branch apices; in- volucres 9.2-13 mm high, 4.5-7 mm wide, the bracts in 4 or 5 definite vertical ranks, keeled, lance-attenuate, the subulate tip soft, the outer more or less herbaceous (sometimes suffused with purple) and the inner with broad hyaline margins; corollas yellow, 7.5-9 mm long; achenes (5) 6-7 mm long, glabrous or sparingly stipitate-glandular; pappus off- white to brownish, abundant. Sagebrush, salt desert shrub, juniper, pinyon-juniper, moun- tain brush, ponderosa pine and alpine fir communities at 1550 to 2900 m in Carbon, Duchesne, Emery, Garfield, Iron, Juab, Kane, Millard, Piute, San Juan, Sanpete, Sevier, Summit, Uintah, Utah, Wasatch, Washington, and Wayne counties; Colorado, New Mexico, Arizona, and Nevada; 34 (iv). Chrysothamnus greenei (Gray) Greene Greene Rabbitbrush. Low, ascending to erect shrubs, with subherbaceous stems from a woody crown, mainly 1-3.5 dm tall, white- barked, glabrous; leaves 0.3-3.5 cm long, 0.8-1.2 mm wide, flat, linear, glabrous or scabrous-ciliate; heads numerous, corym- bosely clustered at branch tips; involucres 5-7.1 mm high, 2.5-4 mm wide, the bracts obscurely ranked, the outer ones herbaceous- thickened near the tip, gradually acuminate- cuspidate, the inner ones abruptly narrowed, glabrous or more or less tomentose, narrowly if at all hyaline-margined; corollas yellow, 3.5-4.8 mm long; achenes 3.3-4 mm long, pi- lose. Rabbitbrush, black sagebrush, shadscale, winterfat, sagebrush, and pinyon-juniper communities at 1280 to 2745 m in Carbon, Duchesne, Emery, Garfield, Grand, Juab, Millard, Piute, Tooele, Uintah, Utah, and Wayne counties; Colorado, New Mexico, Ari- zona, and Nevada; 53 (vi). This entity forms intermediates with phases of C. viscidiflorus. Chrysothamnus Unifolius Greene Spread- ing Rabbitbrush. Tall shrubs, the branches erect-ascending, mainly 8-20 (35) dm tall, white barked, glabrous; leaves 0.9-7.7 cm long, 1-9 mm wide, flat, plane (not contorted or rarely somewhat so), thick, oblong to ellip- tic or narrowly lanceolate, glabrous, scabrous-ciliate, attenuate to acute; heads nu- merous, corymbosely arranged at branch tips; involucres 4.3-7.2 mm long, 1.8-3 mm wide, the bracts indistinctly ranked, the outer dis- tinctly herbaceous at tip, the inner often merely glandular thickened at midrib, all ob- tuse to rounded, glabrous; corollas yellow, 4.5-5.8 mm long; achenes 2.1-2.8 mm long, pilose. Stream banks and terraces, irrigation canals, seeps and springs in riparian commu- nities at 1130 to 2535 m in Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Kane, San Juan, Sanpete, Sevier, Uintah, and Wayne counties; Montana to Arizona and New Mexico; 54 (xvii). Chrysothamnus nauseosus (Pallas) Britt. Rubber Rabbitbrush. Low to tall shrubs, the branches erect-ascending, mainly 2-20 (30) dm tall, the bark obscured by a tomentum, this often resinous-glandular impregnated; leaves 0.6-7 (10) cm long, 0.5-5 (10) mm wide, 1- to 3-nerved, tomentose to glabrate or glabrous, subcylindric to flat, if the latter then commonly plane, linear to narrowly ob- long, acute to apiculate apically; heads nu- merous, in terminal paniculate cymes; in- volucres (6) 6.5-11.5 (13) mm high, 1.5-7.2 mm wide, the bracts obscurely to definitely ranked, the outer ones sparingly tomentose to 240 Great Basin Naturalist Vol. 43, No. 2 glabrous, the inner commonly glabrous, ob- complex in Utah is represented by a diverse long, chartaceous to more or less herbaceous- assemblage of more or less geographically thickened, obtuse to acute or shortly acumi- and ecologically segregated races, which are nate apically; corollas yellow or yellow-or- placed in some 14 varieties. The following ange, 6-10.3 (12) mm long; achenes 2.5-5.5 arbitrary key will serve to identify most mm long, glabrous or hairy. The nauseosus specimens. 1. Shrubs usually 3 dm tall or lower; plants local endemics in Piute, Sanpete, Sevier, Carbon, Emery, Daggett, and Duchesne counties 2 — Shrubs usually more than 3 dm tall, seldom lower, but then of different distri- bution 4 2(1). Involucres glabrous, 8.5-9.5 mm high; plants of Emery, Carbon, Wasatch and Duchesne counties C. nauseosus var. psilocarpus — Involucres tomentose or glabrous, 10-12 (13.5) mm high; plants of Sanpete, Sevier, and Piute counties 3 3(2). Involucres glabrous; corollas 7.8-9 mm long; plants local on Arapien shale in Sanpete and Sevier counties C. nauseosus var. iridis — Involucres tomentose; corollas 10-12 mm high; plants local in Piute County .... C. nauseosus var. glareosus 4(1). Achenes and ovaries glabrous 5 — Achenes and ovaries pilose 8 5(4). Flowers 5-8 mm long; involucres 7-8.5 (9) mm long, 1.5-3 mm wide (when pressed) C. nauseosus var. abbreviata — Flowers 8.3-10 mm long; involucres 9-11 mm long, 3.7-7 mm wide (when pressed) 6 6(5). Involucres subcylindric; plants of dunes and deep sands of western Utah and in the Uinta Basin C. nauseosus var. turbinatus — Involucres tapering to the base; plants of south central and southeastern Utah 7 7(6). Achenes 5-5.5 mm long; plants low, commonly less than 5 dm tall; known from San Juan and Emery counties C. nauseosus var. bigelovii — Achenes 2.5-4 mm long; plants taller, commonly over 5 dm tall; known from Kane County C. nauseosus var. nitidus 8(4). Involucres over 10 mm long; corollas 9.5-10.5 mm long 9 — Involucres 6.5-8.6 (9.5) mm long (to 11 mm long in var. junceus); corollas 5-8.6 (10) mm long 10 9(8). Involucres cylindric, the bracts neither strongly keeled nor ranked; plants of dime areas in western and northeastern Utah C. nauseosus var. turbinatus — Involucres tapering, clavate, the bracts strongly keeled and aligned; plants of Kane County C. nauseosus var. arenarius 10(8). Leaves 3-5 (10) mm wide; plants of central to north central Utah C. nauseosus var. salicifolius — Leaves 0.5-3 mm wide; plants of various distribution 11 11(10). Corolla lobes commonly long-pilose (glabrate in age); leaves often deciduous by anthesis; plants of southeastern Utah C. nauseosus var. junceus — Corolla lobes glabrous; leaves present or absent at anthesis; distribution vari- ous 12 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 241 12(11). Corolla lobes 0.4-0.9 mm long C. nauseosus var. gnaphaloides — Corolla lobes 1-2 mm long 13 13(12). Leaves (1) 3- to 5-nerved, commonly 1-3 mm wide C. nauseosus var. glabratus — Leaves 1-nerved, commonly 0.5-1.5 mm wide 14 14(13). Leaves and/ or stems usually grayish or whitish tomentose or green, not especially yellow-green; involucres more or less tomentose C. nauseosus var. albicaulis — Leaves and/or stems usually yellowish-green, the tomentum commonly resinous-matted; involucres glabrous C. nauseosus var. consimilis Var. abbreviatus (Jones) Welsh comb. nov. [based on: Bigelovia leiosperma var. abbre- viata Jones Proc. Calif. Acad. II, 5: 693. 1895; type from Clear Creek Canyon, Sevier County; C. nauseosus var. leiosperma (Gray) Hall; C. nauseosus ssp. leiospermus (Gray) H. & C; Bigelovia leiosperma Gray, type from St. George.] Blackbrush, Grayia, shadscale, black sagebrush, Vanclevea, pinyon-juniper, and ponderosa pine communities at 1070 to 2745 m in Emery, Garfield, Grand, Kane, Millard, Piute, Sevier and Washington coun- ties; Nevada, California; 15 (v). The materials from Emery and Grand counties have leaves that are very slender and subterete. The con- dition is presumably derived from in- trogression with var. bigelovii. Var. albicaulis (Nutt.) Rydb. [C. nauseosus var. albicaulis Nutt.]. Saltgrass, sagebrush, pinyon-juniper, and ponderosa pine commu- nities at 1310 to 2290 m in Box Elder, Cache, Carbon, Juab, Kane, Millard, Morgan, Salt Lake, San Juan, Uintah, Utah, Wasatch, and Weber counties; Oregon to Wyoming, south to California, Nevada, and New Mexico; 24 (i). This taxon forms intermediates with var. glabratus. In low elevation phases of saline substrates the stems are white-pannose. Var. arenarius (L.C. Anderson) Welsh comb. nov. [based on: C. nauseosus ssp. are- narius L.C. Anderson Phytologia 38: 311. 1978.]. Sagebrush, juniper, and pinyon-juni- per communities at 1675 to 1830 m in Kane County; Arizona; 3 (i). This is a plant of deep sandy alluvium. Var. bigelovii (Gray) Hall [C. nauseosus ssp. bigelovii (Gray) H. & C.; Linosyris (Chrysothamnus) bigelovii Gray]. Grayia and pinyon-juniper communities 1460 to 1950 m in Emery and San Juan (Lavender Mesa) counties; Arizona, Colorado, New Mexico; 2 (i). More collections of this entity are required. Var. consimilis (Greene) Hall [C. nau- seosus ssp. consimilis (Greene) H. & C; C. consimilis Greene]. Saline meadows, riparian zones, and terraces in saltgrass-alkali sacca- ton, shadscale, sagebrush, rabbitbrush, moun- tain brush, pinyon-juniper, and ponderosa pine communities at 1280 to 3000 m in all Utah counties except Grand and San Juan; Oregon to Wyoming, south to California, Arizona and New Mexico; 100 (xxv). This is the common narrow-leaved phase with cone- shaped panicles. They occur frequently in sa- line moist sites, such as the travertine mounds at Monroe Hot Springs. Var. glabratus (Gray) Cronq. [Bigelovia graveolens var. glabrata Gray; C. nauseosus ssp. graveolens (Gray) Piper; C. nauseosus var. graveolens (Gray) Hall]. Desert willow- baccharis, willow-cottonwood, greasewood- tamarix, sagebrush, shadscale, mountain brush, and ponderosa pine communities at 750 to 2475 m in Summit, Wasatch, Utah, Sanpete, Sevier, Piute, Iron, and Washington counties, and in all counties east of those; Idaho to North Dakota, south to Arizona, and New Mexico; 88 (xxiii). Var. glareosus (Jones) Welsh stat. nov. [based on: Bigelovia glareosa Jones Zoe 2: 247. 1891, type from Marysvale; C. nau- seosus ssp. glareosa (Jones) H. & C.]. The type specimen is lost, and the ultimate dis- position of this taxon is uncertain; it should be sought in the canyon north of Marysvale, on Tertiary igneous substrates; endemic; 0 (0). Var. gnaphaloides (Greene) Hall [C. speci- osus var. gnaphaloides Greene; C. nauseosus ssp. hololeucus (Gray) H. & C, in part]. Shadscale, pigmy sagebrush, rabbitbrush, sagebrush, and pinyon-juniper communities at 1070 to 2380 m; known in all Utah coun- ties except Box Elder, Daggett, Duchesne, Kane, Morgan, Rich, Summit, and Wayne, 242 Great Basin Naturalist Vol. 43, No. 2 and likely in them also; California, Nevada, and Arizona (?); 73 (vii). This taxon is a near ally of ssp. hololeucus (Gray) H. & C., and should that taxon be placed within a quad- rinomial, then the var. gnaphaloides would be placed within it. However, no such com- bination is implied or proposed herein. Var. iridis (L.C. Anderson) Welsh stat. nov. [based on: C. nauseosus ssp. iridis L.C. Anderson Great Basin Nat. 41:311. 1981, type from Rainbow Hills, Sevier County]. Rabbitbrush-sagebrush community on an in- cipient seep in Arapien shale at ca 1980 m in Sevier Co.; endemic; 2 (i). Var. junceus (Greene) Hall [C. nauseosus ssp. junceus (Greene) H. & C; Bigelovia jun- cea Greene]. Blackbrush, shadscale, rabbit- brush, matchweed, and pinyon-juniper com- munities at 1220 to 1800 m in Emery, Garfield, Grand, Kane, San Juan and Wayne counties; Arizona; 18 (iv). The nonglandular, clear straw-colored, long involucres with bracts usually aligned are distinctive of this variety. Var. nitidus (L.C. Anderson) Welsh stat. nov. [based on: C. nauseosus ssp. nitidus L.C. Anderson Phytologia 38: 313. 1978]. Van- clevea-ephedra community at about 1250 m in Kane County; Arizona; 1 (0). This variety has the general aspect of vars. bigelovii and abbreviata. It is a taller plant than either, and differs otherwise as set forth in the key. Var. psilocarpus Blake [C. nauseosus ssp. psilocarpus (Blake) L.C. Anderson]. Sage- brush and salina wildrye communities at 1925 to 2290 m in Carbon, Duchesne, Emery, and Wasatch counties; endemic; 5 (0). These peculiar low shrubs occasionally produce taller intermediates with var. gla- bratus (qv.) Var. salicifolius (Rydb.) Hall [C. salici- folius Rydb., type from Strawberry Valley; C. nauseosus ssp. salicifolius (Rydb.) H. & C.]. Sagebrush, pinyon-juniper, mountain brush, and aspen communities at 1310 to 2870 m in Box Elder, Carbon, Duchesne, Emery, Juab, Salt Lake, Sanpete, Sevier, Summit, Tooele, Utah, and Wasatch counties; endemic; 19 (iii). This entity forms intermediates with var. glabratus, and might represent nothing more than a broad-leaved extension of that taxon. Var. turbinatus (Jones) Blake [Bigelovia turbinata Jones, type from Kane County; C. nauseosus ssp. turbinatus (Jones) H. & C.]. Rabbitbrush, saltbush, ephedra, juniper, and greasewood communities at 1370 to 1710 m in Beaver, Iron, Juab, Kane, Millard, and Uin- tah counties; Nevada(?); 10 (iii). Both glabrous and pilose achenes occur in this dis- tinctive taxon. It shares the feature of villous corolla lobes with the sand-loving var. jun- ceus of the Navajo Basin. The Uintah Basin materials differ in the more keeled and atten- uate involucres and flowers that are more ex- serted from the involucre. Chrysothamnus paniculatus (Gray) Greene [Bigelovia paniculata Gray]. Tall shrubs, the branches subfastigiate, mainly 6-20 dm tall, the bark green, becoming tan to gray in age, resinous-punctate; leaves 0.4-3 cm long, about 0.5 mm wide, linear- filiform, terete, mucronate apically; heads numerous, in usually conic panicles; in- volucres 4.8-6.5 mm high, 2-3 mm wide, the bracts indistinctly ranked, chartaceous-in- durate, scarcely if at all glandular, thickened at midrib, obtuse, glandular; corollas yellow, 5.5-6 mm long; achenes 1.8-3.4 mm long, pi- lose. Roadsides, stream banks, terraces, and slopes in creosote bush, Joshua tree, and bac- charis communities at 670 to 1220 m in Washington County; Nevada, Arizona, Cali- fornia; 9 (iii). The plants begin to flower in October and continue into November. Chrysothamnus parryi (Gray) Greene Low to moderate shrubs, the branches not es- pecially fastigiate, mainly 2-6 dm tall, the bark pannose-tomentose or the tomentum glandular-resinous; leaves 0.6-6 (8) cm long, 1-2 mm wide, 1- to 3-nerved, green, viscid or sometimes tomentulose, flat, usually plane, linear to narrowly oblong; heads several to many, the inflorescences tending to be elon- gate and subracemose; involucres 9-14.5 mm high, 4-8 mm wide, the bracts obscurely to definitely ranked, puberulent to glabrous, the outer usually with elongate herbaceous tips, the inner chartaceous, with glandular-thick- ened midrib, abruptly to gradually acumi- nate-attenuate or attenuate; corollas yellow or creamy yellow, 8-10 mm long; achenes 3.3-7.5 mm long, pilose. Plants of the parryi complex form hybrid derivatives with phases of C. nauseosus, and with other named segre- gates within the complex. Except for varie- ties parryi and nevadensis, only arbitrary April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 243 segregation appears possible. Thus, the con- low seems to best reflect the nature of C. par- servative treatment as outlined be- ryi in Utah. 1. Flowers usually more than 10 per head C. parryi var. parryi — Flowers commonly 5-9 per head 2 2(1). Involucral bracts mainly 24-28; plants of southwestern Utah C. parryi var. nevadensis — Involucral bracts mainly 12-22; plants of south central, central, and north- eastern Utah C. parryi var. attenuatus Var. attenuatus (Jones) Kittell in Tidestr. & Kittell [Bigelovia howardii var. attenitata Jones, type from near Marysvale; C. parryi ssp. attenuatus (Jones) H. & C; C. affinis. A. Nels.; C. parryi ssp. affinis (A. Nels.) L.C. Anderson; Linosyris howardii Parry in Gray; C. parryi ssp. howardii (Parry) H. & C; C. parryi var. howardii (Parry) Kittell in Tidestr. & Kittell]. Meadows, sagebrush, juniper, pin- yon-juniper, mountain brush, ponderosa pine, and aspen communities at 1740 to 2930 m in Beaver, Carbon, Daggett, Duchesne, Gar- field, Grand, Iron, Kane, Piute, Sanpete, Se- vier, Uintah, Utah, Wasatch, and Wayne counties; Wyoming and Nebraska, south to Arizona and New Mexico; 55 (xv). The how- ardii phase differs supposedly in the brac- teate leaves overtopping the inflorescence and in the pale colored flowers; both charac- ters fail as diagnostic features. Var. nevadensis (Gray) Kittell in Tidestr. & Kittell [Linosyris howardii var. nevadensis Gray; C. parryi ssp. nevadensis (Gray) H. & C.]. Sagebrush, juniper, pinyon-juniper, mountain brush, and ponderosa pine commu- nities at 1830 to 2565 m in Beaver, Iron, Mil- lard, and Washington counties; Arizona; 10 (ii). The var. nevadensis differs only in degree from var. attenuatus, with which it is con- tiguous, if not partially sympatric, to the east. Should the two be combined, then the cor- rect name will be var. nevadensis, since that name has priority in rank. Plants with leaves overtopping the inflorescence occur; techni- cally they would key to the howardii phase of var. attenuatus. Var. parryi [Linosyris parryi Gray]. Pon- derosa pine and spruce-fir communities at 2075 to 2625 m in Beaver, Emery (?), Gar- field, Kane, Millard, and Washington coun- ties; Wyoming, Colorado, New Mexico, and Nevada; 9 (ii). Chrysothamnus pulchellus (Gray) Greene Low to moderately tall shrubs, the branches not fastigiate, mainly 5-10 dm tall, the bark white, becoming tan or brown in age, glabrous or puberulent above; leaves 0.4-3 cm long, 1-2 mm wide, linear to narrowly oblanceolate, glabrous or puberulent, flat or revolute, mucronate; heads few to many, in corymbose panicles; involucres 11.5-15 mm high, 4.5-6 mm wide, the bracts distinctly aligned, more or less herbaceous toward the apex, glandular, attenuate to sharply acute; corollas yellow, 9-10 (14) mm long; achenes 3.8-4.5 mm long, sparingly hirsute and glandular. Shadscale, blackbrush, ephedra, pinyon-juniper, and ponderosa pine commu- nities at 1370 to 2350 m in Emery, Wayne, and San Juan counties; Arizona to Kansas, south to Mexico; 4 (i). Our material belongs to var. baileyi (Woot. & Standi.) Blake [ssp. baileyi (Woot. & Standi.) H. & C.]. Chrysothamnus vaseyi (Gray) Greene [Bigelovia vaseyi Gray]. Low shrubs, mainly 1-3 dm tall, the branches not especially fasti- giate, the bark green, becoming whitish tan or finally gray in age, puberulent; leaves 0.3-3.7 cm long, 0.8-3 mm wide, linear to oblong or narrowly oblanceolate, glabrous or glandular, flat, plane, mucronate; heads nu- merous in compact terminal cymes; in- volucres 6.2-7.5 mm high, 3-6 mm wide, the bracts more or less aligned, commonly her- baceous or thickened near the apex, glandu- lar, obtuse, the margins fimbriate-hyaline; co- rolla yellow, 4.8-7 mm long; achenes 2.6-4 mm long, glabrous. Meadows, sagebrush, rab- bitbrush, juniper, mountain brush, and pon- derosa pine communities at 1675 to 2900 m in Beaver, Carbon, Emery, Garfield, Juab, Kane, Iron, Piute, San Juan, Sanpete, Sevier, and Utah counties; Nevada, Wyoming, Colo- rado, New Mexico; 21 (ii). 244 Great Basin Naturalist Vol. 43, No. 2 Chrysothamnus viscidiflorus (Hook.) Nutt. Low to moderate shrubs, mainly 2-10 dm tall, the branches fastigiate or not, the bark green to tan or white, finally gray in age, glabrous or puberulent; leaves 0.3-4.5 (6) cm long, 0.5-4 (10) mm wide, 1- to 5- nerved, linear to oblong, elliptic or oblan- ceolate, often twisted, mucronate; heads nu- merous, in compact to open terminal cymes; involucres 5-7.5 mm high, 2-4 mm wide, the bracts not well aligned, commonly her- baceous or thickened near the apex (at least the outer), glandular or puberulent, obtuse, or abruptly acute, the margin narrow, hya- line; corollas yellow, 3.8-6 mm long; achenes 3-4 mm long, pilose. The viscidiflorus com- plex is separable into two groups on the basis of pubescence of upper stems or the lack of pubescence. The segregation is not complete, because pubescence or its absence is not an absolute criterion. There is a cline in the amount of pubescence from abundant to few (or none), and the adoption of a position that one hair equals pubescence and, therefore one part of the complex and not the other, will lead to absurdity. Within the hairy phase of the complex are two more or less dis- tinctive but largely sympatric varieties. The "glabrous" portion of the species is more dif- ficult to separate into its constituent entities. Anderson (Great Basin Nat. 40: 117-20, 1980) reviewed this portion of the complex; concluding that there are three taxa involved, i.e. ssp. axillaris, ssp. viscidiflorus var. vis- cidiflorus, and ssp. viscidiflorus var. steno- phyllus. Only arbitrary separation of the three is possible, and segregation of the ax- illaris phase is problematical. In my view it is not practical to attempt recognition of more than two taxa, i.e. var. stenophyllus (includ- ing axillaris) and var. viscidiflorus. They are aH recognized herein at varietal level, but probably would best fit within an expanded ssp. viscidiflorus as varieties (a course not in- tended or implied herein). The following key will allow for identification of most specimens. 1. Stems (at least above) and/or leaves puberulent to hispidulous 2 — Stems and leaves glabrous, or the leaves ciliate, or rarely with a few short hairs on stems or with glandular excrescences in the inflorescence 3 2(1). Leaves 0.5-2 mm wide; stems finely puberulent above C. viscidiflorus var. puberulus — Leaves 2-5 mm wide; stems hispidulous-puberulent above C. viscidiflorus var. lanceolatus 3(1). Leaves 0.5-1.5 mm wide; plants mainly 2-3 dm tall C. viscidiflorus var. stenophyllus — Leaves mainly 1-4 mm wide (or more); plants mainly 3-10 dm tall C. viscidiflorus var. viscidiflorus Var. lanceolatus (Nutt.) Greene [C. lan- ceolatus Nutt.; C. viscidiflorus ssp. lanceo- latus (Nutt.) H. & C.]. Sagebrush, pinyon- juniper, mountain brush, aspen, Douglas fir, lodgepole pine, spruce-fir, and alpine mead- ow communities at 1375 to 3200 m in all Utah counties except Kane and Washington, and likely there also; British Columbia to South Dakota, and south to California, Ne- vada, Arizona, and New Mexico; 112 (xii). Var. puberulus (D.C. Eaton) Jepson [Lino- syris viscidiflora var. puberula D.C. Eaton; C. viscidiflorus ssp. puberulus (D.C. Eaton) H. & C.]. Rabbitbrush, black sagebrush, shadscale, sagebrush, pinyon-juniper and pon- derosa pine communities at 1460 to 2200 m in the western tier of counties, east to Piute, Sevier, Emery, Carbon, Utah, and Salt Lake counties; Oregon and Idaho south to Califor- nia, Nevada, and Arizona; 44 (vii). Var. stenophyllus (Gray) Hall [Bigelovia douglasii var. stenophylla Gray; C. vis- cidiflorus ssp. stenophylla (Gray) H. & C; C. axillaris Keck; C. viscidiflorus ssp. axillaris (Keck) L.C. Anderson]. Ephedra, blackbrush, rabbitbrush, sagebrush, galleta, shadscale, and pinyon-juniper communities at 1280 to 2075 m in all Utah counties except Piute, Sevier, April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 245 Sanpete, Carbon, Duchesne, Wasatch, Utah, Cirsium Mill, Salt Lake, Davis, Weber, Morgan, Summit, and Cache; Oregon to Wyoming and south to Annual, biennial, or perennial, caulescent California,, Nevada, Arizona, and Colorado; ""' acaulescent, spiny herbs from taproots, 04 / -jx with caudices or rhizomes in some, the juice Var.' viscidifhrus [Crinitaria viscidiflora ^^^^^y' ^^^^^^ ^^^^^ ^"^ ^^"^i"^' alternate; Hook.; C. viscidiflorus var. pumilus authors, ^^^^' fo^it^^X *« ^^^^^f 1' involucral bracts in not (Nutt.) Jeps. (?). Rabbitbrush, shadscale, ^^^^^^^ ^^"^^' subequal to imbricate, some or sagebmsh, pinyon-juniper, mountain brush, "^^f ^^ ^^^"^ ^P^^e tipped; receptacle dense- white fir, ponderosa pine, and aspen commu- ^X bristly; corollas all discoid, pink, purple, nities at 1460 to 2900 m in all or nearly all '^^' ""' ^^^^"^y ^^^^e, perfect or imperfect; Utah counties; Washington to Nebraska, P^PP"^ ^^ plumose bristles (or those of the south to California, Nevada, Arizona, and outermost flowers merely barbellate); style Colorado; 100 (xx). The var. viscidiflorus ^^^^ ^ thickened minutely hairy ring below forms intermediates with all other taxa in the ^^^ ^^^^^X ^o""^*^ 1°^^^' ^^^enes glabrous, species, and with C.greend also. ?^"^^!^ °' ^-angled, 4- to many-nerved. Note: This is a particularly complex genus taxonomically, with both introduced and in- CiCHORiuM L. digenous species. The indigenous members Perennial herbs, with milky juice, from ^'^ especially difficult, due in part to hy- taproots; leaves alternate, toothed to pinnati- bridization, mainly within species groups, fid; heads sessile or subsessile, numerous, ^^ following treatment is tentative, but rep- borne in clusters at nodes of a spicate, simple, resents an attempt to categorize the variation or branched inflorescence; involucral bracts P^'^^^"* ^^ Utah plants and to provide a le- biseriate, the outer shorter; corollas all ray- gitimate name for each. Several taxa pre- Uke, perfect; pappus of 2 or 3 series of scales, ^^o^^^X reported from the state are excluded, sometimes minute; achenes angular or some- ""' ^^y ^'^ ^'^^^^"^ ^^t^*'^ *e constituent what compressed, glabrous. t^^a. All involucral measurements are in Cichorium intybus L. Chickory. Plants pressed condition! 3-10 dm tall or more, hirsute or glabrous; ^o^^^, R. J. and C. Frankton. 1963a. Cyto- lower leaves petiolate, the blades 6-20 cm taxonomic notes on some Cirsium spe- long, 1-5 (7) cm wide, sinuate-dentate to run- ^^^j ?f J^^ ;^^^tf " V^l*. ' ■I ■ \c^ u ■ u A nad. J. Bot. 41: 1553-1567. cmate-pmnatirid, becommg smaller and ses- ^r:r>r^,\ * i r- ^- c ^- ■ ., ^ J £• 11 1 ..• 11 1963b. A clarification of Ctrstum sue upward, some finally subentire; heads r ;• j /-• • j j- o , 11 ir, ^ r ■ foliosum and Cirstum drummondii. Ca- large and showy, 1-3 per node of in- ^^^^ . g^^ 42: 451-461. florescence; flowers pure blue, rarely white; ^ggg Cytotaxonomy of Cirsium hook- involucre 9-15 mm high, the outer bracts erianum and related species. Canad. J. chartaceous at base, herbaceous apically; gQj 43. 597_613 achenes 2-3 mm long. Roadsides and dis- I973. xhe Cirsium arizonicum com- turbed sites at 1340 to 2135 m in Duchesne, plex of the southwestern United States. Iron, Kane, Salt Lake, Tooele, and Utah Canad. J. Bot. 52: 543-551. counties; widespread in North America; na- Petrak, F. 1917. Die nordamerikanischen tive of Eurasia; 8 (i). The herb C. endiva L. is Arten der Gattung Cirsium. Beih. Bot. grown in Utah; the extent is not known. Centralbl. (Abt. 2), 35: 223-567. 1. Flowers mainly imperfect; heads unisexual; plants perennial, from rhizomes; introduced weed of consequence C. arvense — Flowers perfect; plants biennial or perennial, seldom if ever with rhizomes 2 2(1). Leaves roughly hispid above, green; stems conspicuously winged decurrent; plants biennial, introduced C. vulgare — Leaves villous, floccose, arachnoid, tomentose, or glabrous, white to gray or green; stems not winged-decurrent, except in some species; plants indigenous biennials or perennials 3 246 Great Basin Naturalist Vol. 43, No. 2 3(2). Basal rosettes to 10 dm across, the mature leaves commonly 10-30 cm wide, green, glabrate or glabrous on both sides; heads small, with long, tapering, re- curved spines; plants of hanging gardens in southeastern Utah, rarely below them C. rydbergii — Basal rosettes rarely to 5 dm across, the mature leaves usually less than 8 cm wide, floccose, tomentose, arachnoid, or glabrous on one or both sides; plants seldom of hanging gardens in southeastern Utah 4 4(3). Bracts, at least the innermost, conspicuously dilated (but not lacerate), or definitely tan to silvery in appearance, contrasting with the overall aspect of the bracts; plants commonly of meadows C. scariosum — Bracts all spinose, or the innermost occasionally twisted to contorted at the tips, but not especially dilated or conspicuously different in color or texture from the overall aspect of bracts (see C. centaureae); plants of various habitats 5 5(4). Involucral bracts (at least the outer) pinnately spinose; plants green, with yel- lowish spines, of high elevations in the Wasatch, Tushar, and Uinta mountains C. eatonii — Involucral bracts not, or rarely, pinnately spinose (except in C. clavatum, C. scopuhrum, and C. ownbeyi); plants of low to high elevations, but, if pinnately spinose, of other distribution or of low elevations 6 6(5). Heads 1.8-2.7 cm high, and about as wide; inner bracts with coarsely lacerate margins; plants of lower middle elevation meadows C. centaureae — Heads 1.5-3 cm high, 1.5-4.5 (6) cm wide; inner bracts not lacerate; leaves thinly textured, finely to coarsely spined, definitely tomentose or glabrous; plants of various distribution 7 7(6). Herbage definitely white- to gray-tomentose (or rarely green); involucres 1.5-2 cm high, 1.5-2.5 cm wide; known from white shale outcrops in the Uinta Basin C. bamebyi — Herbage green, or white- to gray-tomentose; involucres mainly longer and broader, but if not, then of different distribution 8 8(7). Stems definitely winged-decurrent; heads mainly 1.3-2 cm high, 1.2-3.2 cm wide; herbage white- to gray-tomentose; plants of Sanpete and Washington counties 9 — Stems not winged, or if so, the herbage green and glabrous or nearly so, or the heads commonly larger; plants of various distribution 10 9(8). Leaves of upper stem merely spinose-toothed, tapering from base to apex; plants of Washington County only C. virginensis — Leaves of upper stem definitely lobed, the lobes spinose-toothed, with parallel sides from base to near apex; plants not of Washington County C. subniveum 10(8). Herbage glabrous or glabrate, green 11 ^ Herbage tomentose, floccose-tomentose, gray or white, or only the upper leaf surfaces green 16 11(10). Flowers bright red or carmine; corolla lobes 15-18 mm long; spines of middle involucral bracts 7-11 mm long or more; plants of San Juan County C. rothrockii — Flowers pink, pink-purple, or white; corolla lobes less than 15 mm long; spines of middle involucral bracts 1-6 mm long; plants of various distribution 12 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 247 12(11). Outer bracts not pinnately spinose; mainly low elevation plants, usually in gypsiferous soils, in the Navajo Basin C. calcareum — Outer bracts more or less pinnately spinose; plants of the Navajo and Uinta ba- sins 13 13(12). Stems strongly winged almost or quite the length of upper intemodes; main upper leaves tripinnatifid; plants of lower elevations in northern Uintah and Daggett counties C. ownbeyi — Stems not winged, or rarely some intemodes with incipient wings; main upper leaves pinnatifid to bipinnatifid; plants of moderate to high elevations in the southern end of the Uinta Basin and southward 14 14(13). Involucral bracts ciliate with long yellowish or brownish multicellular hairs; spines of bracts 6-15 mm long or more; plants of the east Tavaputs Plateau and La Sal Mountains C. scopulorum — Involucral bracts more or less ciliate with whitish hairs or a tomentum; spines of bracts mainly 3-7 mm long; plants from the Tavaputs Plateau and south westward 15 15(14). Involucral bracts scabrous dorsally, at least the innermost; herbage not at all tomentose; plants of the Henry Mountains C. calcareum — Involucral bracts not scabrous dorsally; herbage more or less tomentose; plants not of the Henry Mountains C. clavatum 16(11). Heads campanulate, mainly 3.5-6.5 cm wide at anthesis, or, if narrower, bracts commonly glandular-thickened dorsally 17 — Heads turbinate to subcylindric, mainly 2-3.5 cm wide at anthesis; involucral bracts seldom glandular-thickened dorsally 18 17(16). Involucral bracts appearing brown to gray-brown, the spines arising from the body of the bract, not from spreading long-attenuate herbaceous terminal por- tions; bracts of inflorescence usually prominent; plants of broad distribution C. undulatum — Involucral bracts appearing green or fresh green or at least herbaceous, the spines arising from the apex of spreading long-attenuate terminal portions; bracts of inflorescence much reduced; plants of various distribution C. neomexicanum 18(16). Corollas bright red or carmine; plants from Garfield and Iron counties south- ward C. arizonicum — Corollas pale pink, pink, rose-purple, or white; plants from Garfield and Iron counties northward 19 19(18). Involucral bracts (at least the inner) tapering, wedge-shaped, definitely scabrous roughened on dorsal surface, often suffused with red or purple C. calcareum — Involucral bracts smooth dorsally, seldom only somewhat scabrous, not con- spicuously tapering, and seldom conspicuously suffused with red or purple C. wheeleri Cirsium arizonicum (Gray) Petrak Ari- lobed or toothed, the main spines 1-6 mm zona Thistle. [Cnicus arizonicus Gray]. Bien- long, white to grayish tomentose below, more nial or short-lived perennial herbs from a or less tomentose and greenish to green taproot, the caudex sometimes developed; above; stems 4-7.5 dm tall, more or less floc- leaves of basal rosettes 7-36 cm long, bi- cose-tomentose; cauline leaves 3-35 cm long, pinnately lobed or parted, the lobes again 1-8 cm wide, with lobing and vesture similar 248 Great Basin Naturalist Vol. 43, No. 2 to the basal, reduced and less deeply lobed upward; involucres 22-30 mm high, 20-50 mm wide, subcylindric to turbinate, the bracts tomentose at margins, and over back, smooth and often shiny medially, rarely glandular-thickened, the apical portions, es- pecially of the inner definitely scabrous; spines yellowish, 3-10 (15) mm long; corollas crimson to carmine, 25-34 mm long, the tube 8-13 mm long, throat 1.5-11 mm long, the lobes 10-19 mm long. Two more or less dis- tinctive but intergrading phases are present. Heads subcylindric to turbinate; spines 3-10 mm long; plants mainly of the Colorado drainage system (also in western Garfield, and in Iron counties) C. arizonicum var. arizonicum Heads turbinate to broadly so; spines 3-15 mm long or more; plants mainly of the Great Basin and Virgin drainages (also in eastern Iron and western Garfield counties) C. arizonicum var. nidulum Var. arizonicum Salt desert shrub, pinyon- juniper, ponderosa pine, spruce-fir, and hang- ing garden communities at 1220 to 3050 m in Garfield, Iron, Kane, Piute, San Juan, and Washington counties; Arizona; 26 (iv). Var. nidulum (Jones) Welsh comb, no v. [based on: Cnicus nidulus Jones Proc. Calif. Acad. II. 5: 705. 1895]. Pinyon-juniper, mountain brush, aspen, ponderosa pine, Douglas fir, white fir, and spruce-fir commu- nities at 1890 to 3200 m in Beaver, Garfield, Iron, Kane, San Juan and Washington coun- ties; Arizona, Nevada; 37 (iii). Relationships apparently lie with C. rothrockii, C. cal- careum, and, to a lesser extent, with C. wheeleri. Cirsium arvense (L.) Scop. Creeping or Canada Thistle. [Serratula arvensis L.]. Pe- rennial rhizomatous herbs, the stems mostly 5-10 dm tall, glabrous or sparingly tomen- tose; leaves 3-15 cm long, 1-6 cm broad, deeply pinnatifid or lobed to merely toothed, glabrous to tomentose above and beneath; heads several to many, mainly unisexual; in- volucres 10-20 (25) mm high, 10-25 mm wide, the bracts lance-ovate, at least the out- er ones and often all of them spine tipped, to- mentose to glabrous; corollas pink-purple to white; pappus of pistillate heads longer than the corollas, that of staminate heads shorter than the corollas; achenes 3-5 mm long. Roadsides, fields, and other disturbed sites, but also invading native plant communities, at 1280 to 2535 m, probably in all Utah counties; widespread in North America; ad- ventive from Eurasia; 42 (iii). We have two phases of creeping thistle in Utah; the one with merely toothed (unlobed) leaves is var. mite Wimm. & Grab., and the common one with deeply lobed leaves is var. horridum Wimm. & Grab. This common weed and the bull thistle are our only two introduced thistles in the genus Cirsium, which makes up a huge assemblage in the Old World. We can expect more introductions. Cirsium barnebyi Welsh & Neese in Welsh Barneby Thistle. Perennial herbs from a caudex and taproot, the caudex clothed with black marcescent leaf bases; leaves of basal rosettes 11-25 cm long, bipinnately lobed or parted, the lobes again lobed or toothed, the main spines 3-5 mm long, whit- ish- to grayish tomentose on both sides; stems 3-5 dm tall, whitish tomentose (rarely green); cauline leaves 2-30 cm long, 1-8 cm wide, with lobing and vesture similar to the basal, reduced and less deeply lobed upwards; in- volucres 15-22 mm high, 20-30 mm wide, turbinate, the bracts glabrate or sparingly arachnoid on margins, glutinous dorsal ridge inconspicuous, smooth medially, the apical portions of the inner often contorted, not scabrous dorsally; spines 2-7 mm long, flat- tened apically, more or less spreading; co- rollas bluish pink. Sagebrush, juniper, cryp- tantha, ephedra, wildrye, and rabbitbrush communities at 1525 to 2257 m in Uintah County; endemic; 7 (iii). The Barneby thistle is apparently related to the undulatum complex. Cirsium calcareum (Jones) Woot. & Standi. Cainville Thistle. [Cnicus calcareus Jones, type from Cainville]. Perennial herbs from a caudex and taproot, the caudex with brownish black to castaneous marcescent leaf bases; leaves of basal rosettes 6-35 cm long, pinnatifid to bipinnatifid, glabrous and green or tomentose on one or both surfaces, the April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 249 main spines 3-8 mm long; stems mainly 2-5 dm tall, glabrous or more or less floccose-to- mentose, winged-decurrent or not; cauline leaves 3-28 cm long, 0.8-7 cm wide, bipin- natifid, with lobing and vesture like the basal, reduced upward, the main spines 3-8 mm long; involucres 19-34 mm long, 15-45 mm wide, the bracts ovate-lanceolate to linear, more or less tomentose at the margins, smooth and often shiny medially, the dorsal ridge glandular-thickened or not, the apical portions of at least the inner scabrous; spines straw colored, 1.5-6 mm long; corollas pink to blue-pink. The calcareum complex is a portion of the arizonicum group of thistles, and has long been misinterpreted. There are three more or less confluent varieties present in Utah. Specimens collected are few, espe- cially in the critical southeastern portion of the state. More work is indicated. 2(1). Herbage permanently tomentose, the leaves grayish tomentose beneath C. calcareum var. pulchellum Herbage green, the leaves rarely sparingly tomentose along the midveins be- neath 2 Leaves definitely decurrent, the stems winged 2-6 cm below leaf base; plants of San Juan and Wayne counties C. calcareum var. calcareum Leaves not or scarcely decurrent; plants of other distribution C. calcareum var. bipinnatum Var. bipinnatum (Eastw.) Welsh stat. nov. [based on: Cnicus drummondii var. bipinna- tum Eastw. Zoe 4: 8. 1893]. Aspen, Douglas fir, and riparian communities at 1130 to 3150 m in Garfield, Kane, and San Juan counties; Colorado, New Mexico, and Arizona; 8 (i). Var. calcareum [Cirsium pulchellum var. glabrescens Petrak type from Elk Mountains, San Juan County]. Riparian communities at 1460 to 2200 m in Carbon, San Juan, and Wayne counties; endemic (?); 4 (i). Jones (Proc. Calif. Acad. II., 5: 704. 1895) cited two collections with the protologue; i.e., Jones 5695bh from Bromide Pass in the Henry Mountains and Jones 5696 from Cainville. His description best fits the Cainville mate- rials, and that collection is here selected as lectotype. The material from Bromide Pass seems best to fit var. bipinnatum (q.v.). A pe- culiar plant with thin leaves that are glabrous on both sides and subentire is known from Cedar Canyon (Atwood and Higgins 5918 BRY). How it fits into the scheme of Utah thistles is not known, but the plant appears to be intermediate between this and some other thistles. The status of the Cainville thistle, as strictly interpreted, beyond Utah is unknown; it seems likely that it does not occur outside the state. Var. pulchellum (Greene) Welsh comb, nov. (based on: Carduus pulchellus Greene ex Rydb. Fl. Colorado 400, 401. 1906). Rabbit- brush, sagebrush, tamarix, rabbitbrush, pin- yon-juniper, and aspen communities at 1340 to 2745 m in Carbon, Emery, Garfield, Grand, Kane, San Juan, Uintah, Utah, and Wayne counties; Colorado, New Mexico, Ari- zona; 41 (vii). Both winged and wingless stems are present within our material. There are plants from the San Rafael Swell with winged stems and they are similar to C. och- rocentrum Gray of New Mexico, but they ap- pear to be transitional in every way with the wingless plants. And it seems probable that they are not conspecific with that plant as it occurs beyond Utah. Possibly they do war- rant taxonomic recognition. Further collec- tions are necessary. Cirsium centaureae (Rydb.) K. Schum. [Carduus centaureae Rydb.]. Fringed Thistle. Perennial herbs from a simple caudex and taproot, the caudex with chestnut leaf bases; leaves of basal rosette 2-28 cm long, 1-8 cm wide, pinnatifid, the lobes often again toothed, tomentose below, thinly tomentose to glabrous above, the main spines 1-5 mm long; stems 3-12 dm tall, not succulent, arachnoid or glabrous; cauline leaves with lobing and vesture like the basal, the spines 3-8 mm long; involucres 18-27 mm high, and about as wide, the outer bracts lance- ovate, the inner with coarsely lacerate mar- gins, usually dilated in the upper half, tomen- tose to glabrous on the margins, the dorsal 250 Great Basin Naturalist Vol. 43, No. 2 ridge not well developed, the longest spines 2-5 mm long, straw colored; flowers white to pink or purple. Montane communities at 3355 m in San Juan Co.; Wyoming and Colo- rado; 2 (0). Cirsium clavatum (Jones) Petrak Fish Lake Thistle. [Cnicus clavatus Jones, type from Fish Lake]. Perennial or biennial herbs from a taproot, and often with a caudex, the caudex clothed with marcescent chestnut- brown leaf bases; leaves of basal rosettes 2.5-22 cm long, bipinnately parted to merely toothed, green on both sides or more or less tomentose below, the main spines 1-6 mm long; stems 3-10 dm tall, glabrous or thinly tomentose; cauline leaves 3-26 cm long, 0.5-7 cm wide, with lobing and vesture like the basal, reduced and less lobed above; in- volucres 18-23 (32) mm high, 22-30 (55) mm wide, the bracts more or less villous-tomen- tose on margins, the outer ones usually pin- nately spiny, smooth medially, the dorsal ridge not especially glandular, apical por- tions of the inner ones often scabrous, some- times slightly dilated-erose; spines yellowish, 3-8 (18) mm long; corollas white or less com- monly pink. Sagebrush, meadow, aspen, Douglas fir, and spruce-fir communities at 2135 to 3200 m in Beaver, Carbon, Emery, Garfield, Grand, Kane, Piute, Sanpete, Se- vier, Uintah, and Wayne counties; endemic; 27 (viii). The Fish Lake thistle is apparently related to the allopatric C. eatonii. It is more or less transitional to C wheeleri, and prob- ably other taxa, especially those with scabrous inner bracts. Rarely some have de- current leaf bases, and when the pinnately spinose bracts are poorly developed, this thistle approaches C. calcareum. Moore and Frankton (1965) proposed that C clavatum was a hybrid between C. eatonii and C. cen- taureae. However, despite its possible origin from hybridization, the taxon seems to be or- ganized on about the same basis as other thistles. Further, its distribution is distinct from that of the putative parents. There does not seem to be justification for recognition of this entity as a hybrid. Cirsium eatonii (Gray) Robins. Eaton Thistle. [Carduus eatonii Gray]. Perennial herbs from a simple or rarely branched cau- dex and taproot, the caudex clothed with brownish black to brown marcescent leaf bases; leaves of basal rosette 4-20 cm long, more or less bipinnatifid, green and glabrous or nearly so on both sides, the main spines 1.5-4 mm long; stems 1.5-5 dm tall, glabrous or nearly so; cauline leaves 3-25 cm long, 0.6-5.5 cm wide, with lobing like the basal, reduced upward; involucres 20-37 mm high, 25-50 mm wide, the bracts ovate-lanceolate to lance-linear, tomentose to long-villous marginally (rarely overall), the outer ones usually pinnately spiny, smooth to roughened medially, the dorsal ridge not developed, the apical portions of the inner ones sometimes contorted; spines 5-18 mm long, straw col- ored; corollas pink to white. Three more or less distinctive varieties are present. 2(1). Involucral bracts copiously gray- to brown-villous with multicellular hairs; co- rollas ocroleucous; plants of the Uinta Mountains from Lake Fork eastward C. eatonii var. murdockii Involucral bracts merely white-tomentose or rarely with short multicellular hairs; corollas mainly pink or rose; plants of western Uinta Mountains, and elsewhere 2 Involucral bracts commonly suffused with dark purple; involucres not ob- scured by outer spinose bracts; plants of the Tushar Mountains C. eatonii var. harrisonii Involucral bracts green or variously purplish; involucres with copious pinnate spines, mainly obscuring the surface of inner bractlets; plants of western Uinta and Wasatch mountains, and Great Basin ranges C. eatonii var. eatonii Var. eatonii [C. eriocephalum var. leio- cephalum D.C. Eaton; this is the basionym for C. eatonii in a strict sense, which was re- named by Gray in honor of D.C. Eaton who collected with Sereno Watson in 1869]. The lectotype came from the head of the Bear River, in Summit County (Watson 691, 1869 US!), with syntypical material being taken April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 251 under the same number in Cottonwood Can- yon (now Salt Lake County). Lodgepole pine and spruce communities upwards into alpine tundra at 2375 to 3420 m in Duchesne, Juab, Salt Lake, Summit, Tooele, and Weber coun- ties; Nevada and Colorado; 31 (iv). Speci- mens from the Deep Creek Mountains have few lateral spines on the outer bracts, and ap- proach C. clavatum in technical features. More material is needed to determine their status and relationships. Var. harrisonii Welsh Talus slopes and al- pine meadows at 2975 to 3450 m in Beaver and Piute counties; endemic; 6 (v). This low phase of the Eaton thistle stands geographi- cally apart from the remainder of the species, isolated on the islandlike Tushar Mountains. Var. murdockii Welsh The plants grow in talus slopes and on rock stripes at 3230 to 3660 m in Daggett, Duchesne, and Uintah counties; endemic; 7 (iii). This variety has been regarded as constituting a portion of C. tweedyi (Rydb.) Petrak. That entity was re- viewed by Moore and Frankton (1965) and was mapped to include northeastern Utah in its range. However, no specimens were cited from Utah. I have seen the type of that taxon, and other material within its range in north- western Wyoming, and they differ in pu- bescence of involucral bracts being merely white tomentose along the margins. Cirsium neomexicanum Gray Biennial herbs from taproots; leaves of basal rosette 5-25 cm long (or more), pinnatifid, the lobes again toothed or lobed, white tomentose be- low and less so above, the main spines 1-6 mm long; stems 6-15 dm tall, whitish tomen- tose; cauline leaves 1.5-35 cm long, 0.5-7 cm wide, tomentose, appearing filmy greenish white, lobed like the basal ones, rather abruptly reduced upward, finally minute spiny bracts; involucres 20-30 mm high, 40-65 mm wide, the bracts green or greenish, narrowly lanceolate, tomentose marginally (or overall), the outer ones often reflexed, the inner minutely serrulate-ciliate, long-attenu- ate apically, the spine a continuation of the attenuation, smooth medially, the glandular dorsal ridge more or less well developed, the apical portions of the inner often contorted; spines 1-9 mm long, yellowish; corollas creamy white. 1. Involucral bracts green throughout, the attenuate apex not differing in texture from the body of the bract C. neomexicanum var. neomexicanum — Involucral bracts not green throughout, the attenuate apex differing in texture from the body of the bract C. neomexicanum var. utahense Var. neomexicanum Creosote bush, Joshua tree, blackbrush, shadscale, sagebrush, and pinyon-juniper communities at 915 to 2050 m in Beaver, Garfield, Grand, Juab, Kane, Millard, San Juan, Tooele, and Washington counties; Nevada, Arizona, New Mexico; 26 (vii). This is one of the most distinctive spe- cies of thistle in Utah. The tall slender stems, with one or few large heads with creamy white flowers, stand in candelabra form in the arid portions of western and southern Utah. Ghostlike stalks of previous years per- sist for a time, reminding one of the regime which allowed their growth. Var. utahense (Petrak) Welsh comb. nov. [based on: C. utahense Petrak Beih. Bot. Centr. 35(2): 470. 1917.] Salt desert shrub, sagebrush, pinyon-juniper, and mountain brush communities at 1220 to 2300 m in Cache, Carbon, Emery, Millard, Rich, Salt Lake, Tooele, and Utah counties; Colorado (?); 24 (ii). This taxon has long been confused with C. undulatum with which it shares the grayish tomentum, large heads, and tall stat- ure. They have been separated previously on the basis of glandular development of the dorsal ridge; a feature which is, unfortu- nately, not diagnostic. The long-attenuate bract apices from which the spines arise are apparently distinctive for this taxon. It is es- sentially intermediate between undulatum and neomexicanum in a strict sense. The type is from Silver Reef, Washington County, but the main area of distribution for this variety is apparently along the Wasatch Mountains in northern Utah. Cirsium ownbeyi Welsh Ownbey Thistle. Perennial herbs from caudex and taproot, the caudex with marcescent dark brown leaf bases; leaves of basal rosettes 5-13 cm long, 1.5-3 cm wide, tripinnatifid, green on both sides, sparingly tomentose along lower side of 252 Great Basin Naturalist Vol. 43, No. 2 midrib; cauline leaves with vesture and lob- ing like the basal; stems 5-7 dm tall, winged- decmrent, sparingly tomentose; involucres 1.8-2.5 cm high, 1.5-2.5 cm wide, the out- ermost bracts more or less pinnately spinose, lance-attenuate, smooth medially, the dorsal ridge not well developed, not scabrous, spar- ingly tomentose along margins, the inner more or less contorted apically; spines 3-8 mm long; corollas rose-pink. Juniper, sage- brush, and riparian communities at 1678 to 1891 m in Daggett and Uintah counties; en- demic; 2 (i). Relationships of the Ownbey thistle apparently lie with C. eatonii. Cirsium rothrockii (Gray) Petrak Rothrock Thistle. [Cnicus rothrockii Gray; Cnicus roth- rockii var. diffusus Eastw., type from Willow Creek, San Juan County]. Perennial or bien- nial herbs from a caudex and taproot, the stems 5-8 dm tall, sparingly tomentose or glabrate to glabrous; cauline leaves 3.5-30 cm long, 2-9 cm wide, bipinnatifid, green and glabrous or nearly so on both sides, car- ried well to the inflorescence; involucres (19) 23-28 (34) mm long, 20-35 mm wide, the bracts lanceolate to lance-linear, more or less tomentose along the margins, smooth me- dially, the dorsal ridge not or only somewhat glandular, sometimes purplish apically, the apical portions of the inner definitely scabrous, the spines 7-17 mm long; corollas red to carmine. Mixed shrubs and ponderosa pine woods at 1830 to 2560 m in San Juan County; Arizona; 3 (0). This entity is poorly known in Utah; its relationship is with both C. calcareum and C. arizonicum. It is a green subglabrous plant with red flowers and long involucral spines. Cirsium rydbergii Petrak Rydberg Thistle. [Cirsium lactucinum Rydb., type from Bluff]. Perennial herbs from a definite caudex and taproot, the caudex clothed with blackish brown leaf bases; leaves of basal rosette mainly 30-90 cm long, 15-40 cm wide, bi- pinnatifid, the lobes narrow to very broad, glabrous to glabrate on both surfaces, the main spines 2-11 mm long; stems 6-12 dm tall or more, glabrous; cauline leaves glabrous, less lobed and much reduced up- wards; involucres 10-17 mm high (not mea- suring the reflexed outer bracts), 13-26 mm wide, the outer bracts lance-ovate, rather abruptly contracted into recurved spines 3-25 mm long, sparingly tomentose mar- ginally; dorsal glandular ridge lacking, the in- ner attenuate, not scabrous; flowers pink. Hanging gardens, or rarely in canyons below them, at 1125 to 1525 m in Grand, Kane, San Juan, Wayne (and probably in Garfield) counties; Arizona (?).; 15 (v). Both C ryd- bergii Petrak and C. lactucinum Rydberg are based on the same type collection from the hanging gardens near Bluff. The Rydberg thistle is a plant with huge basal rosettes, tall slender flowering stems, and small heads. Cirsium scariosum Nutt. Meadow Thistle. [Carduus lacerus Rydb., type from near Mid- way; Carduus olivescens Rydb., type from the Aquarius Plateau; Cirsium acaule var. americanum Gray; Cnicus drummondii var. acaulescens Gray; C foliosum authors, not T. & G.; C. drummondii authors, not T. & G.]. Perennial herbs from a simple caudex and taproot, the caudex with chestnut leaf bases; leaves of basal rosette 2-28 cm long, 1-8 cm wide, merely spiny toothed to bipinnatifid, the lobes often again toothed, tomentose to glabrate below, thinly tomentose to glabrous above, the main spines 1-5 mm long; stems lacking, or 1-12 dm tall or more, often suc- culent and edible, arachnoid to glabrous; cauline leaves (when stems present) bipin- natifid or merely pinnatifid, the spines 3-35 mm long, with vesture like the basal; in- volucre 22-35 mm high, 20-65 mm wide, the outer bracts lance-ovate, the inner progres- sively more lance-attenuate, smooth me- dially, the margins smooth to minutely scabrous, tomentose to glabrous on margins, the dorsal ridge not well developed, the long- est spines mainly 2-5 mm long, straw col- ored, the inner with tips more or less con- torted, dilated, or fimbriate, usually whitish or silvery; flowers white to pink or pink- purple. Our specimens fall into two rather distinctive varieties. 1. Heads 25-35 mm high, 35-80 mm wide; inner bracts slender, sometimes con- torted, not especially dilated; plants mainly 6-12 dm tall C. scariosum var. thorneae — Heads 22-30 mm high, 20-40 mm wide; inner bracts often dilated or con- torted, sometimes fimbriate; plants 0-6 dm tall C. scariosum var. scariosum April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 253 Var. scariosum [Cirsium acaule var. ameri- canum Gray]. This taxon, as here interpreted, consists of an amazingly diverse assemblage that has passed under a series of names in- cluding those cited above; and, if it is demon- strated that C. foliosum (Hook.) DC. is ac- tually conspecific, that name has priority. Saline seeps and salt marshes, stream sides, terraces, and other meadowlands at 1310 to 3175 m in Carbon, Duchesne, Emery, Gar- field, Juab, Millard, Salt Lake, Sanpete, Se- vier, Summit, Tooele, and Utah counties; British Columbia to Montana, south to Cali- fornia, Arizona, and Colorado; 43 (x). This phase of C. scariosum has passed under the names C. acaulescens (Gray) Schum., C. colo- radoense (Rydb.) Cockerell; C tioganum (Congdon) Petrak, C drummondii T. & G., and C. foliosum. Nomenclature is still un- clear, and more work is indicated. Our highly variable material is transitional from acaules- cent to caulescent within populations, with stems, when present, fleshy and edible. This is our common thistle of meadowlands, and it is unfortimate that nomenclatural entangle- ments have not allowed selection of an unequivocal name. Reported for the state is C. parryi (Harrington, Flora of Colorado, 1952), but I have seen no specimens of that entity from Utah. It would key to C. scario- sum in the present work. It has densely arachnoid involucral bracts, with at least the innermost dilated-fringed at the tips; flowers are greenish yellow and the leaves are gla- brate on both surfaces. Var. thorneae Welsh Stream terraces and seeps or springs at 1650 to 2475 m in Beaver, Garfield, Iron, Kane, Millard, and Piute counties; endemic (?); 10 (vi). In addition to the features noted above, the cauline leaves are thick, with coarse veins, and spines 8-35 mm long. Cirsium scopulorum (Greene) Cockerell in Daniels [Carduus scopulorum Greene]. Pe- rennial herbs from taproots; leaves of basal rosettes 3-28 cm long, 0.8-8 cm wide, with spines 2-6 mm long, unlobed to bipinnatifid, tomentose below, glabrate to glabrous and green above; stems mainly 3-7 dm tall, spar- ingly arachnoid, not winged-decurrent; cau- line leaves mainly bipinnatifid, or the upper ones merely pinnatifid, green above, glabrous to sparingly tomentose below, rather gradu- ally reduced upward; heads in a compact subglobose terminal cluster; involucres 30-35 mm high, 30-55 mm wide, the bracts lance- attenuate, abundantly villous marginally, with long yellowish to brownish multicellular hairs, the outer ones usually pinnately spiny, the dorsal crest not glandular, smooth me- dially, the apical portions of the inner ones often contorted; spines 10-18 mm long, yel- lowish; corollas pale yellow to cream. Sage- brush, aspen, and spruce-fir communities at 2135 to 3000 m in Grand, San Juan (?), and Uintah counties; Colorado; 3 (0). Cirsium subniveum Rydb. Perennial herbs from taproots; basal rosettes not seen; stems mainly 6-10 (13) dm tall, tomentose, winged- decurrent; cauline leaves 3-25 cm long or more, 1-6 cm wide, pinnatifid, tomentose on both sides, or less so above, the bases decur- rent; involucres 17-25 mm high, 20-30 mm wide, the bracts ovate-lanceolate, smooth medially, the glandular dorsal ridge more or less developed, none scabrous, tomentose marginally; spines 3-5 mm long; corollas ap- parently white to cream. Pinyon-juniper community at 1890 m in Rich and Sanpete counties; Oregon to Montana; 2 (0). Cirsium undulatum (Nutt.) Spreng. Gray Thistle. [Carduus undulatus Nutt.]. Perennial herbs from a simple caudex and taproot, the caudex more or less clothed with persistent leaf bases; leaves of basal rosette mainly 7-25 cm long, 1.5-6 cm wide, merely toothed to bipinnatifid, tomentose on both sides, white- tomentose below, white to greenish above, the main spines 1-6 mm long; stems 2-10 (12) dm tall, tomentose; cauline leaves bipin- natifid or the upper ones merely pinnatifid, with vesture as in the basal ones, rather grad- ually reduced upward; involucres (15) 20-30 mm high, 20-60 mm wide, the bracts brown or brownish, lance-ovate to lanceolate, to- mentose on margins or overall, the dorsal ridge strongly glutinous to undeveloped, the spinose tips spreading, with yellowish spines mainly 2-5 (10) mm long, smooth medially, the apical portion of the innermost more or less contorted; corollas pink, pink-purple, or creamy white. 254 1. Great Basin Naturalist Vol. 43, No. 2 Heads mainly less than 2.5 cm wide, even the largest, commonly (1) 3-10 or more per stem C undulatiim var. tracyi Heads mainly more than 2.5 cm wide, at least the largest, commonly 1-3 per stem C. undulatum var. undulatum Var. tracyi (Rydb.) Welsh comb. nov. [based on: Carduus tracyi Rydb. Bull Torrey Bot. Club. 32: 133. 1905]. Sagebrush, moun- tain brush, juniper, aspen, and Douglas fir communities at 1525 to 2900 m in Duchesne, Emery, Grand, Juab, Summit, and Uintah counties; Colorado; 26 (iii). This variety grades with the type variety, and separation is at least partially arbitrary. Var. undulatum Desert shrub, sagebrtish, pinyon-juniper, mountain brush, ponderosa pine, and aspen communities at 1400 to 2600 m in Cache, Carbon, Daggett, Duchesne, Emery, Garfield, Juab, Grand, Rich, San Juan, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, and Weber counties; British Columbia to Minnesota, south to Arizona, New Mexico, and Missouri; 59 (vi). Cirsium virginensis Welsh Virgin Thistle. Perennial(?) herbs from taproots; leaves of basal rosettes 6-35 cm long, 1-5 cm wide, unlobed, pubescent like the cauline ones, with spines 1-4 mm long; stems 6-15 dm tall, tomentose, winged by definitely decurrent leaf bases; cauline leaves 1.5-15 cm long or more, sinuate-dentate to pinnatifid, whitish tomentose on both sides, or greenish above, often reduced to spiny bracts upwards; in- volucres 13-20 mm tall, 12-32 mm wide, the bracts ovate-lanceolate to narrowly lanceo- late, brownish to straw colored, or often suf- fused with purple, tomentose marginally (or overall), the outer not especially reflexed, the inner serrulate or entire, smooth medially, the glandular dorsal ridge more or less devel- oped, the apical portions of the inner often contorted; spines 2-6 (8) mm long, yellowish; corollas pink to lavender (or white?). Saline seeps and stream terraces at 850 to 950 m in Washington Co.; Arizona; 9 (i). The small heads and long decurrent leaf bases are diag- nostic. The relationships of the Virgin thistle are unknown. It does not appear to be closely related to other species groups represented in our area. Cirsium vulgare (Savi) Ten. Bull Thistle. [Carduus vulgaris Savi]. Biennial herbs from taproots; leaves of basal rosette mainly 5-25 cm long, 2-8 cm wide, merely doubly ser- rate-dentate to doubly pinnatifid, tomentose beneath, coarsely hispid above; stems mainly 3-12 (15) dm tall, spiny-winged by decurrent leaf bases; cauline leaves mainly bipinnatifid, with vesture as in the basal ones; involucres 28-40 mm high, 35-70 mm wide, the bracts narrowly lanceolate, with spreading spine- tips, tomentose marginally, the dorsal ridge not developed, the inner sometimes con- torted apically; spines 1-4 mm long, yellow- ish; corollas rose-purple. Meadows, fields, roadsides, and other disturbed sites from 1340 to 2745 m in most, if not all, Utah counties; widespread in North America; 52 (i). Cirsium wheeleri (Gray) Petrak Wheeler Thistle. [Cnicus wheeleri Gray]. Perennial or biennial herbs from a simple or branched caudex and taproot, the caudex clothed with persistent brown to dark brown leaf bases; leaves of basal rosettes mainly 7-20 cm long, 1-5 cm wide, once to twice pinnatifid, or merely toothed or spinose-serrate, grayish or whitish tomentose below, thinly so to glabrous and green above, the main spines 0.5-4 mm long; stems 2.5-7 dm tall; cauline leaves 2-25 (32) cm long, 0.5-5 (7) cm wide, with lobing and vesture similar to the basal, carried well to the inflorescence, though re- duced above; involucres 20-27 mm high, 20-35 mm wide, the bracts lance-ovate to lance-linear, more or less tomentose along the margins, smooth medially, the dorsal ridge not or only somewhat glandular, some- times purplish tipped, the apical portions of at least the inner more or less scabrous; co- rollas pink to pink-purple, or less commonly white. Mountain brush, pinyon-juniper, white fir, aspen, and spruce-fir communities at (1980) 2165 to 3150 m in Beaver, Emery, Garfield, Iron, Juab, Kane, Millard, San Juan, Sanpete, and Sevier counties; Colorado, New Mexico, and Arizona. Our materials appar- ently intergrade with C. undulatum, C. nidu- lum, and possibly C. scariosum. The moder- ate sized heads, usually pink or pink-purple flowers, low stature, essentially nonglandular April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 255 bracts, and usually green upper leaf surface appears to be diagnostic. The phases from Cedar Canyon (Iron County), with merely spinose unlobed leaves, are striking, but probably not more than minor variants; 39 (iv). Cnicus L. Annual caulescent spiny herbs from tap- roots, the juice watery; leaves alternate; heads solitary, terminating branches; in- volucral bracts in several series, spine tipped, the inner ones pinnately spiny; receptacle densely bristly; corollas all discoid, yellow, perfect; pappus in 2 series, the outer smooth, long, alternating with short sparingly pecti- nate ones; style with a ring of hairs at base of divergent branches; achenes terete, strongly ribbed, glabrous. Cnicus benedictus L. Blessed Thistle. Plants 1-5 dm tall or more, branching from near the base; stems villous; leaves mainly 8-15 cm long, pinnatifid, more or less glandular and sparingly villous, the spines 0.5-1.5 mm long, the lower ones petiolate, becoming sessile above; involucres 3-4 cm high, closely subtended and obscured by the foliose bracteate upper leaves; corollas yel- low. Waste places and gardens at 885 m in Washington County; widespread in the U.S.; adventive from Europe; 1 (0). CoNYZA Less. Nom. Cons. Annual herbs from taproots, with watery juice; stems erect, commonly branched; leaves alternate, simple; heads numerous, in cylindric to conic panicles; involucral bracts more or less imbricate, herbaceous medially; receptacle flat or nearly so, naked; rays min- ute, white or purplish, scarcely surpassing the pappus; disk flowers seldom more than 20, perfect, fertile; pappus of capillary bristles; achenes 1- or 2-nerved or nerveless. Conyza canadensis (L.) Cronq. Horse- weed. [Erigeron canadensis L.]. Annuals, mainly 0.5-10 dm tall, glabrous or spreading- hairy; leaves 2-8 (10) cm long, 2-8 cm wide, linear to oblanceolate, ciliate-serrate, often deciduous by late anthesis; heads numerous, inconspicuous; involucres 2-3.5 (4) mm high, (2.5) 3-7 mm wide, the bracts lance-subulate, the midvein glandular-thickened, herbaceous medially, glabrous or strigose; rays white or purplish. Weedy species, often in riparian or other moist disturbed sites at 850 to 2135 m in all (?) Utah counties; widespread in North America; Europe; 30 (vi). Our material be- longs to var. glabrata (Gray) Cronq. Crepis L. Annual, biennial, or perennial caulescent or subacaulescent herbs, from taproots, with milky juice; leaves basal and cauline, alter- nate, pinnatifid to toothed or entire; heads few to numerous, in corymbose or paniculate clusters; involucral bracts in 1 or 2 series, herbaceous; receptacle naked; corollas all raylike, perfect, yellow or yellowish; pappus of numerous white capillary bristles; achenes terete or nearly so, 10- to 20-ribbed, often beaked. 1. Plants annual, adventive, of disturbed sites C. capillaris — Plants perennial, indigenous, neither weedy nor of disturbed sites 2 2(1). Leaves and stems glabrous (or glandular-hispid only above); plants sub- acaulescent or subscapose 3 — Leaves and stems more or less tomentose or puberulent to setose or glandular hispid; plants caulescent 4 3(2). Plants less than 10 cm tall, soboliferous, of high elevations C. nana — Plants mainly 15-40 cm tall, never soboliferous, of lower-elevation meadows ... C. runcinata 4(2). Heads narrowly cylindric; involucral bracts 5-7 (8), the inner commonly glabrous; flowers mostly 5-10 C. acuminata — Heads narrowly to broadly campanulate; involucral bracts 8-15, tomentose and often setose-hispid; flowers mostly 8-60 5 256 Great Basin Naturalist Vol. 43, No. 2 5(4). Leaf segments linear to narrowly lanceolate, entire or nearly so, the terminal lobe more than 5 cm long; achenes commonly green C. atrabarba — Leaf segments narrowly lanceolate to triangular, some usually toothed, the ter- minal lobe less than 5 cm long; achenes mainly yellowish to brownish 6 6(5). Involucres more than twice longer than broad; leaves usually green, runcinate- pinnatifid C intermedia — Involucres less than twice longer than broad, or leaves not green or not run- cinate-pinnatifid 7 7(6). Involucre and stems not or sparingly setose, but, if setose, the setae gland- tipped C. occidentalis — Involucre and/or stems conspicuously setose, the setae not glandular C. modocensis Crepis acuminata Nutt. Perennial herbs, communities at 1890 to 2870 m in Daggett, 2.5-8.5 dm tall, with 1-several stems from a Garfield, Salt Lake, Summit, Tooele, Uintah, caudex, the caudex clothed with dark brown and Utah counties; British Columbia and Al- marcescent leaf bases; herbage more or less berta, south to Nevada and Colorado; 8 (i). tomentose to glabrate; basal and lowermost The species is evidently uncommon in Utah, cauline leaves 8-33 (40) cm long, 2-12 cm It is known to form apparent hybrids with C. wide, petiolate, the blade elliptic to oblan- acuminata, and probably with other taxa as ceolate in outline, pinnatifid to runcinate- well. pinnatifid, the lobes triangular to narrowly Crepis capillaris (L.) Wallr. Slender subulate, sometimes toothed or lobed; heads Hawksbeard. Annual or biennial herbs, the mainly 20-75 or more, cylindric, 5- to 10- stems erect, simple or branched, mostly 1-6 flowered; involucres (8) 9-13.5 (16) mm high, dm tall, sparingly spreading-hairy; basal 3-7 mm wide, the inner ones 5-8, glabrous or leaves 3-20 cm long, 0.5-3 cm broad, lan- sometimes shortly villous-tomentose, the out- ceolate to oblanceolate, denticulate to pin- er bracts much shorter, commonly tomen- natifid or bipinnatifid, glabrous or pubescent tose; corollas 10-18 mm long, yellow; ach- with stiff spreading hairs, especially along the enes yellow to brown, narrowed above, lower midvein, petiolate; cauline leaves re- Sagebrush, mountain brush, white fir, aspen, duced upward, sessile and auriculate-clasp- and spruce-fir communities at 1430 to 2900 ing; heads (1) several to numerous, mostly 20- m in most if not all Utah counties; Washing- to 60-flowered, borne in an open in- ton to Montana, south to California, Arizona, florescence; involucres 5-8 mm high, 5-14 and New Mexico; 69 (viii). mm wide, the inner bracts lance-attenuate, Crepis atrabarba Heller [C. occidentalis 8-16, tomentose, often glandular-hairy, var. gracilis D.C. Eaton]. Perennial herbs, glabrous within, the outer bracts lance-linear; 2-4.5 dm tall, with 1-several stems from a achenes 2-5 mm long, pale brown to straw caudex, the caudex with dark brown to colored; achenes 2-5 mm long, pale brown to purplish marcescent leaf bases; herbage gray straw colored, not beaked. Ruderal weed of villous-tomentose to glabrate, basal and Salt Lake County; widely scattered in North lowermost cauline leaves 6-22 cm long, America; adventive from Europe; 1 (0). 1.5-4 cm wide, petiolate, the blade lance- Crepis intermedia Gray Gray Hawksbeard. elliptic in outline, pirmatifid, the lobes linear [C. barbigera Leiberg, in part]. Perennial or linear-subulate, the terminal lobe 5-9 cm herbs, 2.5-7 dm tall, with 1-several stems long, entire; heads mainly 2-15, cam- from a caudex, the caudex clothed with pale panulate, commonly 10- to 40-flowered; in- to dark brown marcescent leaf bases; herbage volucres 9-15 mm long, 7-13 mm wide, the more or less tomentose or villous; basal and inner ones 8-10, usually grayish tomentulose lowermost cauline leaves 15-30 cm long, and often with few glandless black setae; co- 2-10 cm wide, petiolate, the blade elliptic to rollas 10-18 mm long, yellow; achenes usual- oblanceolate in outline, pinnatifid to run- ly greenish, attenuate at the apex. Sagebrush, cinate-pinnatifid, the lobes triangular to lin- ponderosa pine, Douglas fir, and white fir ear-subulate, sometimes toothed or lobed, the April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 257 terminal lobe less than 5 cm long; heads mainly 10-60, campanulate, 7- to 16-flow- ered; involucres 11-20 mm high, 6-12 mm wide, the inner ones 7-12, tomentulose (rarely glabrate), sometimes setose with non- glandular setae, the outer bracts much short- er; corollas 13-20 mm long, yellow; achenes mainly yellowish or brownish, narrowed above. Sagebrush, pinyon-juniper, and moun- tain brush communities at 1525 to 2575 m in Beaver, Cache, Duchesne, Garfield, Salt Lake, Sevier, Utah, Wasatch, and Washing- ton counties; Washington to Alberta, south to California, Nevada, and Colorado; 16 (iii). The intermedia assemblage consists of a series of apomictic intermediates involving C. acuminata as one of the parental types, and one or more of the other taxa (i.e., occiden- talis or modocensis) to complete the complex. Included here is the concept of C. barbigera as it has been applied in Utah; it consists of a similar hybrid sequence of polyploid apo- micts from outside our area. Crepis modocensis Greene Modoc Hawks- beard. Perennial herbs, 1.5-3.7 dm tall, with 1 -several stems from a caudex, the caudex clothed with pale to brown marcescent leaf bases (the stem base often yellow); herbage more or less tomentose; basal and lowermost cauline leaves 9-25 cm long, 2-5 cm wide, petiolate, the blade elliptic to oblanceolate in outline, bipinnatifid, the lobes linear to lance-subulate, again toothed or lobed, the terminal lobe less than 5 cm long; heads 1-9, 10- to 60-flowered; involucres 11-16 mm high, 11-23 mm wide, the inner bracts 10-15, tomentulose, commonly setose, the setae not glandular, the outer bracts much shorter; corollas 13-22 mm long, yellow; ach- enes greenish black to reddish brown, attenu- ate. Sagebrush, pinyon-juniper, and mountain brush communities at 1640 to 3175 m in Bea- ver, Box Elder, Cache, Daggett, Juab, Mil- lard, Rich, Salt Lake, Sanpete, Sevier, Tooele, Uintah, and Utah counties; British Columbia to California, Nevada, and Colo- rado; 24 (0). The peculiar numerous slender lateral lobes of the deeply dissected or parted leaf blades are diagnostic. Crepis nana Richards. Dwarf Hawksbeard. Perennial caespitose herbs, the stems much branched, often soboliferous, mostly 0.2-1.1 dm tall, contracted, usually obscured by the leaves, glabrous; basal leaves mainly 1-7.5 cm long, 0.2-1.8 cm wide, the blades spatu- late to orbicular, elliptic, or ovate, glabrous, petiolate; cauline leaves similar to the basal, not clasping; heads few to numerous, mostly 4- to 12-flowered, borne in a compact cush- ionlike inflorescence; involucre 7-12 mm high, 3-6 mm wide, the inner bracts narrow- ly oblong, 8-12, greenish or blackish, glabrous, the outer much shorter; achenes brownish, ribbed, shortly beaked. Alpine communities, mainly in talus, at 3050 to 3425 m in Juab, Piute, and Utah or Salt Lake counties; Alaska to Labrador, south to Cali- fornia and Utah; 10 (iii). Crepis occidentalis Nutt. Western Hawks- beard. Perennial herbs, 1-4 dm tall, with 1- several stems from a caudex, the caudex clothed with brown marcescent leaf bases (the stem base often yellow); herbage tomen- tose; basal leaves mainly 6-30 cm long, 1-5 cm wide, petiolate, the blade lanceolate to elliptic in outline, pinnatifid to bipinnatifid, the lobes triangular to oblong or linear-sub- ulate, usually again toothed or lobed, the ter- minal lobe less than 5 cm long; heads 2-25, 12- to 30-flowered; involucres 10-20 mm high, 6-15 mm wide, the inner bracts (7) 8-13 (18), tomentose, the outer ones much shorter; corollas 10-22 mm long, yellow; ach- enes pale to dark brown, not much attenuate apically. There are three rather weak and ar- bitrarily recognizable varieties of this species, with some geographical correlation, in Utah. Intermediates occur between the varieties and with other taxa as well. 2(1). Largest heads 12- to 14-flowered, with 8 or 9 involucral bracts; plants mainly of the Great Basin C. occidentalis var. costata Largest heads with more than 15 flowers, with 10-13 involucral bracts; plants of various distribution 2 Involucres with few glandular setae, or none; plants mainly 2-3 dm tall, of the Great Basin C. occidentalis var. pumila 258 Great Basin Naturalist Vol. 43, No. 2 Involucres with few to numerous glandular setae; plants mainly 1-2 dm tall, of the Colorado drainage system, less commonly in the southern Great Basin C. occidentalis var. occidentalis Var. costata Gray Sagebrush, pinyon- juniper, mountain brush, and aspen commu- nities at 1525 to 2200 m in Box Elder, Juab, Millard, Salt Lake, Tooele (type from Stans- bury Island), Utah, and Washington counties; British Columbia to California and Colorado; 23 (0). Var. occidentalis Shadscale, rabbitbrush, sagebrush, pinyon-juniper, and ponderosa pine communities at 1280 to 2565 m in Bea- ver, Daggett, Duchesne, Garfield, Kane, Piute, Sanpete, Sevier, San Juan, and Wash- ington counties; Oregon to Wyoming, south to California and New Mexico; 25 (v). Var. pumila (Rydb.) Babcock & Stebbins [Crepis pumila Rydb.]. Sagebrush, pinyon- juniper, and mountain brush communities at 1700 to 2100 m in Millard and Tooele coun- ties; 7 (0). Crepis runcinata (James) T. & G. [Hier- acium runcinatum James]. Perennial herbs, 1.5-5 (7) dm tall, with 1-several stems from a caudex, the short caudex clothed with brown marcescent leaf bases; herbage glabrous or hispid above (puberulent in some), not to- mentose; basal leaves mainly 2-25 cm long, 1-6 (8) cm wide, petiolate or not, spatulate to oblanceolate, or the blades ovate to oval, oblong, or oblanceolate, more or less pinnati- fid to lobed or entire, commonly glaucous; heads 1-30, with 20-50 flowers; involucres campanulate, 8-16 mm high, 6-15 mm wide or more, the inner bracts mainly 10-16, pu- berulent or hispid, the outer ones much shorter; corollas 9-18 mm long, yellow; ach- enes light to dark brown, attenuate, or short- ly beaked. Three distinctive varieties are present. 2(1). Involucres merely puberulent; plants mainly of saline meadows C. runcinata var. glauca Involucres hispid with black hairs (resembling species of Hieracium); plants of saline or nonsaline sites 2 Basal leaves definitely petiolate, the blade 2-4 times longer than broad C runcinata var. hispidulosa Basal leaves broadly winged-petiolate, the blade 4-8 times longer than broad ... C. runcinata var. runcinata Var. glauca (Nutt.) Welsh stat. nov. [based on: Crepidium glaucum Nutt., Trans. Amer. Phil. Soc. II. 7: 436. 1841]. Meadows, lake shores, seeps, and hot springs in salt grass, rush, alkali sacaton, and common reed com- munities at 1220 to 2200 m in Carbon, Dag- gett, Duchesne, Emery, Grand, Juab, Kane, Millard, Piute, San Juan, Sevier, Tooele, Uin- tah, Utah, and Wayne counties; Idaho to Sas- katchewan, south to Arizona and New Mexi- co; 34 (xii). This variety has been collected in full flower on 27 April at Monroe Hot Springs. Var. hispidulosa Howell ex Rydb. Sedge- willow and meadow communities at 1370 to 2535 m in Box Elder, Duchesne, Kane, Piute, Rich, Sanpete, Sevier, Summit, and Utah counties; Washington to Montana, south to California and Colorado; 14 (ii). The mead- ows are seldom saline where this plant occurs. Var. runcinata [C. runcinata var. alpicola Rydb.]. Bogs in Salt Lake and Utah counties; Manitoba to Minnesota, south to Idaho and New Mexico; 1 (0). This variety is evidently uncommon in Utah. DicoRiA T. & G. Annual herbs; leaves alternate or the lower ones opposite, simple, entire or toothed; heads unisexual or perfect, discoid; involucral bracts strongly dimorphic, the ca 5 outer ones small, herbaceous, the inner subtending the 1 or 2 pistillate flowers, subscarious, ac- crescent, much larger than the outer at matu- rity; chaff narrow, tardily deciduous; pis- tillate flowers without corolla; staminate flowers with funnelform corolla, the anthers distinct; achenes plano-convex, black, toothed to pectinately wing margined; pap- pus lacking. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 259 Foliose bracts of inflorescence orbicular to broadly ovate; plants of Washing- ton County D. canescens Foliose bracts of inflorescence lance-ovate to lanceolate; plants not of Wash- ington County D. brandegei Dicoria brandegei Gray [D. paniculata Eastw.; D. wetherillii Eastw., a monstrous form]. Plants branched from the base up- wards, 1.5-5.5 dm tall, the herbage white-pi- losulose to strigose, the hairs multicellular; lower cauline leaves linear to lanceolate, more or less hastately lobed, toothed, or sub- entire, 1-7 cm long (including petiole), 0.2-1.5 cm wide; foliose bracts linear to ob- long, lanceolate or ovate, rarely if ever or- bicular, the blades 0.6-4 cm long; outer in- volucral bracts oblong, 1.5-3 mm long, the inner ones suborbicular, glandular-pu- berulent, accrescent in fruit; achenes 5-8 mm long, the winged margin toothed to pecti- nate, black like the body or pale. In dunes and other sandy sites, in wavy-leaf oak, eriogonum, amsonia, old-man sagebrush, rab- bitbrush, ephedra, and vanclevea commu- nities at 1130 to 1830 m in Emery, Garfield, Grand, Kane, San Juan, and Wayne counties; Arizona, New Mexico, and Colorado (?); 21 (V). Dicoria canescens Gray in Torr. Plants branched from base upward, 2.5-9 dm high, the herbage white-pilosulose to strigose and glandular, the hairs multicellular; lower cau- line leaves deltoid-lanceolate, dentate, 1-5 cm long; foliose bracts ovate to orbicular, the blades 0.6-1.5 cm long; outer involucral bracts oblong, 2-3 mm long, the inner ones suborbicular, glandular-puberulent, accres- cent in fruit, to 10 mm long or more; achenes 5-6 mm long, the winged margin toothed to pectinate, black like the body, or pale. Dunes and other sandy sites in blackbrush and creo- sote bush communities at 825 to 1000 m in Washington County; Arizona, Nevada, and California; 4 (1). Our material belongs to ssp. clarkae (Kennedy) Keck. Dyssodia Cav. Annual or perennial herbs or subshrubs from taproots, the juice watery; herbage with conspicuous translucent oil glands; stems striate, numerous; leaves opposite or alter- nate, entire to pinnatisect; heads solitary at branch ends, or few to several in cymose clusters; involucral bracts in 2 series, distinct or united, and usually with a much shorter outer set; receptacle flat or convex, pu- berulent; ray flowers yellow, pistillate, fer- tile; disk flowers fertile; pappus of 10-15 bristle-tipped scales, or these dissected into 3 or more bristles; style branches with a short, conic appendage. Strother J. L. 1969. Systematics of Dyssodia Cavanilles (Compositae: Tageteae). Univ. Calif. Publ. Bot. 48: 1-88. 1. Plants annual; leaves bipinnatisect; stems villosulous D. papposa — Plants perennial, herbs or subshrubs; leaves simple or merely pinnatisect; stems hispidulous 2 2(1). Heads borne on elongate merely bracteate peduncles; leaves pinnately 5-lobed, shortly hispid; pappus scales tipped with usually a solitary bristle D. pentachaeta — Heads sessile or essentially so; leaves simple, entire or rarely irregularly few lobed; glabrous or merely ciliate; pappus scales with 3-5 bristles D. acerosa Dyssodia acerosa DC. Dogweed. Plants glands; ray flowers 7-8, lemon-yellow; disk suffruticose, 10-25 cm tall, forming compact flowers 18-25, pale yellow; pappus of ca 20 clumps, from taproots; herbage glabrous or scales, each dissected into 3-5 bristles; ach- villosulous; leaves opposite (or alternate enes dark brown, 3-3.5 mm long, strigose. above), simple or irregularly lobed, 3-18 mm Blackbrush communities at 1130 to 1350 m long, 0.5-1 (2) mm wide, glandular, ciliate or in Garfield, Washington, and San Juan coun- glabrous; heads sessile or subsessile; in- ties; Arizona and New Mexico, south to Mex- volucres turbinate-cylindric, 5-7 mm high, ico; 5 (ii). 3-4 mm wide; involucral bracts ca 13, con- Dyssodia papposa (Vent.) A.S. Hitchc. nate, each bract with conspicuous orange [Tagetes papposa Vent.]. Plants annual, 1.5-4 260 Great Basin Naturalist Vol. 43, No. 2 dm tall; herbage glabrous to sparingly pu- berulent; leaves opposite below, alternate above, 1.5-3 (5) cm long, pinnatisect into 11-15 lobes, these sometimes again lobed; heads shortly pedunculate to subsessile; in- volucres turbinate to campanulate, 6-10 mm high, and about as wide; involucral bracts 6-12, oblanceolate, with yellowish oil glands, connate only at the base; ray flowers 8 or fewer, yellow-orange; disk flowers mainly 20-40, dull yellow; pappus of ca 20 scales, each dissected into 5-10 bristles; achenes black, 8-35 mm long. Sandy roadsides at 1450 to 1500 m in Carbon, Duchesne, San- pete, and Tooele counties; through much of the United States and Mexico; 4 (0). Dyssodia pentachaeta (DC.) Robins. [Hymenatherum pentachaetum DC] Plants suffruticose, 8-28 cm tall, forming rounded clumps, from taproots; leaves opposite, pin- nately parted into 3-5 rigid linear lobes, 0.5-2 cm long, sparingly hirtellous; pe- duncles 1-8 cm long; involucres turbinate, 4.8-6 mm high, 5-10 mm wide; involucral bracts in 2 series, connate for much of their length, with distinctive yellow oil glands; ray flowers usually 13, bright yellow; disk flow- ers 50-70, dull yellow; pappus usually of 10 scales, these awnless or with 1-3 awns; achenes brown, 2.2-3 mm long, hispid to glabrous. Blackbrush, ephedra, shadscale, creosote bush, and Joshua tree communities at 700 to 1220 m in Garfield, Kane, San Juan, and Washington counties; Nevada and Cali- fornia to Texas and Mexico; South America; 30 (vi). Our material has been assigned to var. belinidium (DC.) Strother [D. thurberi (Gray) Woot. & Standi.]. Encelia Adams Shrubs; stems ascending to erect, grayish or whitish, the branchlets commonly pu- bescent; leaves alternate, simple, petiolate, entire or toothed; heads solitary or in cymose clusters, radiate or discoid; involucral bracts in 2 or 3 series; receptacle convex to flat, chaffy, the scales clasping the achenes and falling with them; ray flowers (when present) sterile, yellow; disk flowers perfect, yellow; pappus lacking (or of 2 awns); achenes flat, obovate, villous-ciliate and pubescent on the surfaces. Blake, S. F. 1913. A recision of Encelia and some related genera. Proc. Amer. Acad. 49: 358-376. 1. Leaves white-tomentulose; peduncles glabrous; heads in branching cymes; plants rare in Washington County E. farinosa — Leaves strigose to hispid, green; peduncles scabrous to strigose; heads soli- tary at branch ends; plants of Washington County, and elsewhere E. frutescens Encelia farinosa Gray Incienso. Plants mainly 3-10 dm tall, aromatic; leaves clus- tered at apex of current stems, 2-8 cm long, ovate, entire or toothed, silvery tomentose, petiolate; peduncles elongate, cymosely branched or simple; heads showy, the disk 1-1.5 cm wide; involucres 4-7 mm high, vil- lous and glandular dotted; rays 8-12 mm long, orange-yellow; achenes narrowly obo- vate. Blackbrush community at 1280 m in Washington County; Nevada, Arizona, and California; Mexico; 1 (0). Encelia frutescens Gray Bush Encelia. [Simsia frutescens Gray]. Plants mainly 3-12 (15) dm tall; leaves scattered along current stems, the blades commonly 0.5-2.5 cm long, 0.3-2 dm wide, ovate to orbicular or lanceo- late, entire or toothed, strigose to hispid with pustular-based hairs; heads showy or not, the disk 1-3 cm wide; involucres 6-10 mm high, strigose or glandular; rays lacking or 1-16 (or more), 2-12 mm long, yellow; achenes obo- vate. Two distinctive varieties are present in Utah. Herbage strigose, also with some pustular-based hairs; involucral bracts abrupt- ly caudate-acuminate, strigose; plants of Washington County £. frutescens var. virginensis Herbage hispid with pustular-based hairs; involucral bracts gradually attenu- ate, more or less glandular (sometimes strigose) E. frutescens vslt. frutescens April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 261 Var. frutescens [E. frutescens var. resinosa Jones in Blake] Talus and slickrock in black- brush and shadscale communities at 1130 to 1830 m in Emery, Grand, Kane, and San Juan counties; Arizona, California; 15 (vii). There is a cline of glandularity in leaves from defi- nitely glandular in the southern portion of the range in Utah to no glands at all in the northern material. Also, our plants vary from discoid to radiate. Var. virginensis (A. Nels.) Blake [E. virgi- nensis A. Nels.]. Creosote bush, Joshua tree, and blackbrush communities at 760 to 1325 m in Washington County; Nevada, Arizona, and California; 23 (i). Enceliopsis (Gray) A. Nels. Perennial scapose or subscapose herbs, from tuberous roots or taproots and sub- terranean to superficial caudex; herbage pi- losulose to velutinous; leaves all basal (rarely some reduced bracteate ones along the scape), the blades spatulate, lanceolate, ob- lanceolate, ovate, or orbicular; heads solitary; involucral bracts in 2 or 3 series, herbaceous throughout; receptacle flat to convex, chaffy, the scales clasping the achenes; rays yellow, sterile (but apparently pistillate), or lacking; disk flowers numerous, perfect, fertile, yel- low; pappus of 2 awns and with or without small scales between, or none; achenes flat- tened, blackish. 1. Heads discoid; herbage pilose-hirsutulose; plants arising from a subterranean tuberous root E. nutans — Heads radiate; herbage tomentulose; plants arising from a superficial caudex 2 2(1). Petioles broadly winged, mainly shorter than the blades; plants reported from the Virgin Narrows section of Washington County, but none have been seen by me E. argophyllus (D. C. Eaton) A. Nels. — Petioles slender, not or only narrowly winged, mainly longer than the blades; plants commonly in eastern and west central portions of the state E. nudicaulis Enceliopsis nudicaulis (Gray) A. Nels. [Encelia nudicaulis Gray]. Scapose, caespi- tose perennials from a superficial, branching caudex, 10-43 cm tall, the herbage tomentu- lose, silvery white; petioles 0.7-17 cm long, narrowly if at all winged; leaf blades 2-9 cm long, 1.3-10 cm wide, ovate to elliptic, or- bicular or spatulate, cuneate to subcordate basally, obtuse to rounded apically; scapes of- ten with a reduced foliose bract; involucres 1.3-2.2 cm high, 3-5.6 cm wide, the bracts ovate-lanceolate to lanceolate or linear-lan- ceolate, attenuate to acuminate; rays 13-21, yellow, 22-38 mm long; achenes 10-12 mm long, long silky-pilose, cuneate, black or dark brown; pappus commonly of 2 awns con- nected by a crown of short connate scales (or none). Commonly on gypsiferous semibarren knolls in blackbrush, rabbitbrush, ephedra, shadscale, grayia, and pinyon-juniper com- munities in Beaver, Emery, Garfield, Grand, Millard, Piute, San Juan, Sevier, Uintah, and Wayne counties; Idaho, Nevada, Arizona, and California; 66 (xiii). Enceliopsis nutans (Eastw.) A. Nels. [En- celia nutans Eastw.]. Scapose, discoid pe- rermials, 10-25 cm tall, from a subterranean caudex (2-15 cm long) and tuberous root to 4 cm thick, the herbage strigose to pilosulose (antrorsely on the upper surface, retrorsely so below), green; petioles 2-6.5 cm long, often narrowly winged; leaf blades 2-7.5 cm long, 1.4-6 cm wide, ovate to orbicular or spatu- late, cuneate basally, obtuse to rounded api- cally; scapes not bracteate; involucres 0.9-1.5 cm high, 2.5-4 cm wide, the bracts lance-at- tenuate; rays lacking; achenes 9-11 mm long, oblanceolate, long silky-pilose, brown; pap- pus lacking. Mainly in finely textured soils in shadscale, budsage, galleta, and ephedra com- munities at 1310 to 1830 m in Carbon, Du- chesne, Emery, Grand, Uintah, and Wayne counties; Colorado (a Colorado Basin endem- ic); 35 (iii). Erigeron L. Annual, biennial, or perennial herbs from caudices, rhizomes, stolons, or taproots, with watery juice; stems decumbent to ascending or erect, rarely prostrate; leaves alternate, simple, entire, toothed, or pinnatifid to pal- matifid; heads solitary or few to numerous in corymbose or paniculate inflorescences; in- 262 Great Basin Naturalist Vol. 43, No. 2 volucral bracts equal, or slightly to definitely flattened, 2 (rarely 4-14) -nerved. Note: This imbricate, slender, herbaceous (or scarcely is a large and complex genus. The species, al- herbaceous) throughout; receptacle flat, though mainly distinctive, are distinguished naked; rays white, pink, purple, bluish, or by minute features that can be interpreted yellow, numerous, pistillate, or lacking; disk variously. The genus is a near congener of flowers numerous, yellow or tinged reddish; both Aster and Conyza, and is not always sep- pappus of capillary bristles, sometimes with arable from either. an outer series of short bristles or scales; style Cronquist, A. C. 1947. Revision of the branches with lanceolate and acute or North American species of Erigeron triangular and obtuse appendages; achenes north of Mexico. Brittonia 6: 121-302. 1. Plants with yellow ray flowers, known from Box Elder County E. linearis — Plants with ray flowers pink, pink-purple, blue, blue-purple, or white, but not yellow, of various distribution 2 2(1). Plants annual, biennial, or short-lived perennials from usually slender taproots, lacking rhizomes (except in some specimens of E. proselyticus) or woody caudices KEY I — Plants definitely perennial, often from rhizomes or caudices 3 3(2). Plants silvery pubescent; achenes with 4 or more nerves; involucral bracts defi- nitely imbricate KEY II — Plants green, or less commonly silvery pubescent; achenes with 2 nerves, or, if with more nerves, involucral bracts subequal 4 4(3). Involucres wooUy-villous to spreading villous, or villous-hirsute with at least some long spreading multicellular hairs KEY III — Involucres merely glandular, glabrous, puberulent, or with appressed simple or multicellular hairs, rarely with some spreading long hairs near the base KEY IV Key I. Plants armual, biennial, or short-lived perennials from slender taproots, lacking rhizomes or woody caudices. 1. Pistillate corollas very numerous, filiform, the rays short, erect, not exceeding the disk, or the inner ones tubular and lacking rays 2 — Pistillate corollas few to numerous (rarely lacking), the tube generally cylin- dric, the rays well developed and spreading, rarely reduced or absent 3 2(1). Cauline leaves narrowly lanceolate to oblong, or less commonly linear; rayless pistillate flowers present between the ray and disk flowers; inflorescence corymbose, the peduncles curved-ascending, or the heads solitary E. acris — Cauline leaves linear to oblong; rayless pistillate flowers lacking; inflorescence racemose, the peduncles erect or nearly so, or the heads solitary E. lonchophyllus 3(1). Pappus of ray and disk flowers unlike, that of the disk flowers composed of bristles and short outer setae, that of the ray flowers lacking bristles; plants tall adventive weedy species 4 — Pappus of ray and disk flowers alike, consisting of bristles, sometimes also with outer setae or scales; plants indigenous, low to tall 5 4(3). Foliage ample; plants mainly 6-12 (15) dm tall; pubescence of stem long and spreading (at least below); plants introduced, weedy E. annuus — Foliage sparse; plants mainly 3-7 dm tall; pubescence various; plants to be sought in Utah E. strigosus Muhl. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 263 5(3). Plants diffusely branched, annual; leaves linear to linear-oblong; hairs of stem short and incurved; pappus simple E. hellidiastrum — Plants various, but seldom as above, or, if so, the pappus double 6 6(5). Disks mainly over 1 cm wide; stems commonly simple, with solitary or few heads, and broad cauline leaves E. glabellus — Disks mostly less than 1 cm wide; stems commonly branched, often with sever- al to many heads 7 7(6). Stems with hairs all spreading E. divergens — Stems with hairs appressed or ascending, or glabrous 8 8(7). Leaves entire; plants with sterile flagellate branches E. flagellaris — Leaves pinnately lobate or toothed, or if entire then the plants lacking sterile flagellate branches 9 9(8). Ray flowers commonly 40-80; plants psammophytes of eastern Washington and western Kane counties E. religiosus — Ray flowers commonly 25-40; plants of various substrates in eastern Washing- ton and Kane counties 10 10(9). Involucres 2.5 mm high or less; stems 1-11 cm tall; peduncles sparingly villous; plants known from seeps and moist sandstone in Zion National Park E. sionis — Involucres 2.5-3 mm high; stems 14-25 cm long; peduncles hirsute; plants from limestone and sandstone outcrops in eastern Iron and adjacent Kane counties E. proselyticus KEY II. Plants perennial, silvery pubescent; achenes 4 (or more) -nerved; involucral bracts imbricate. 1. Achenes glabrous, with 8-14 nerves; caudex clothed with marcescent leaf bases, the midribs evident in age; plants of higher elevations in southern Utah £. canus — Achenes more or less hairy, with 3-8 nerves; caudex lacking marcescent leaf bases, or if these present then the midribs not evident; plants of low to moderate elevations, more widely or otherwise distributed 2 2(1). Involucres villous-hirsute with multicellular spreading hairs, the bracts more or less glandular apically; achenes with 3-5 nerves E. pulcherrimus — Involucres more or less strigose with simple hairs; achenes 4- to 8-nerved 3 3(2). Basal leaves evident, tufted, persistent; heads one per stem; plants through much of Utah E. argentatus — Basal leaves mostly withered at anthesis, not forming a conspicuous tuft; plants mainly of southeastern Utah E. utahensis KEY III. Plants perennial; achenes mostly 2-nerved; bracts mostly subequal, villous with woolly or spreading multicellular hairs. 1. Plants with pinnatifid or palmatifid or merely lobed leaves, low-spreading, more or less mat or clump forming, of high elevations 2 264 Great Basin Naturalist Vol. 43, No. 2 — Plants with entire leaves, or, if some of them lobed, otherwise differing; low to tall, of various elevations 4 2(1). Plants soboliferous, the caudex divided into elongate spreading branches; leaves merely toothed, or if lobed not as below E. vagus — Plants not soboliferous, the caudex branches short; leaves pinnately to palmately lobed or divided 3 3(2). Leaves pinnately lobed; plants of the La Sal Mountains E. mancus — Leaves palmately lobed or divided; plants widespread E. compositus 4(1). Involucres long- and shaggy-villous, the hairs sometimes obscuring the bract surface from middle to base 5 — Involucres hirsute to shortly villous, or, if long and shaggy-villous, the hairs not obscuring the bract surface even in the lower portion 7 5(4). Plants 4-7 dm tall or more; known from southeastern Utah E. elatior — Plants mainly 0.3-1 dm tall; distribution various 6 6(5). Hairs of involucre with black or dark purple cross-walls; basal leaves rounded to refuse apically; plants of the La Sal mountains E. melanocephalus — Hairs of involucre with pale cross-walls or some with bright reddish purple to dark purple cross-walls; basal leaves acute to abruptly obtuse apically; plants of the Uinta, Deep Creek, Tushar, and La Sal Mountains E. simplex 7(4). Cauline leaves ample, usually lanceolate or broader; plants tall, erect (more or less asterlike) 8 — Cauline leaves usually much reduced, subulate, linear, oblong oblanceolate, or, if broader, the plants not tall or not erect 11 8(7). Hairs of involucre with black cross-walls near their bases; rays white; plants rare, known from the Wasatch Mountains E. coulteri — Hairs of involucre with pale cross-walls; rays white, pink, or purple; abun- dance and distribution various 9 9(8). Plants with cauline leaves well developed and equably distributed, only gradu- ally reduced upward, the middle ones as large as or larger than the lower ones E. speciosus — Plants with cauline leaves rather abruptly reduced upward, those of the middle smaller than the lower ones 10 10(9). Involucres glandular or viscid toward the apex; stems curved at base E. formosissimus — Involucres seldom if at all glandular or viscid; stems erect E. glabellus 11(7). Ray flowers lacking E. aphanactis — Ray flowers present 12 12(11). Plants subscapose, the bracteate leaves very small; caudex branches with per- sistent leaf bases 13 — Plants not subscapose, the leaves merely reduced upward; caudex branches with or without persistent leaf bases 16 13(12). Stems and involucres with long, contorted villous hairs; plants of Box Elder and Daggett counties E. nanus — Stems and involucres strigose, pilosulose, or hispidulous, the hairs appressed or ascending to spreading; distribution various 14 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 265 14(13). Leaves linear; herbage strigose; rays 7-11 mm long; plants mainly of lower elevations in the Navajo and Great basins E. compactus — Leaves narrowly oblanceolate to spatulate; herbage strigose to pilosulose or hispidulous; rays 4-8.2 mm long; plants of the Uinta Basin and Wasatch Plateau 15 15(14). Involucres long-villous with spreading multicellular hairs; rays 6.8-8.2 mm long; plants of the Wasatch Plateau E. carringtonae — Involucres short-hispidulous; rays 4-6.5 mm long; plants of the Uinta Basin E. untermannii 16(12). Caudex branches robust, 1-2.5 cm thick; plants of western Beaver and Wash- ington counties E. wahwahensis — Caudex branches mainly less than 1.5 cm thick, or, if broader, of different dis- tribution 17 17(16). Stems spreading-hairy 18 — Stems strigose, or with ascending hairs 21 18(17). Leaves linear to linear-oblanceolate; plants without a prominent caudex, mainly of lower elevations E. pumilus Leaves oblanceolate to spatulate; plants with prominent caudex, of low to middle or higher elevations 19 19(18). Stems glandular, with sand grains adhering; plants of lower elevations in Emery and Wayne counties E. maguirei — Stems lacking glands; plants of moderate and higher elevations 20 20(19). Stems commonly purplish at the base; leaves thin; plants of broad or other dis- tribution E. eatonii — Stems green throughout; leaves thickish; plants of the Uinta and Wasatch mountains E. goodrichii 21(17). Caudex with spreading subrhizomatous branches, with numerous fibrous roots; stems and lower leaf bases purplish; plants sod forming, of higher elevations E. ursinus — Caudex not subrhizomatous, seldom if ever with roots; stem and leaf bases not purple; plants of various elevations 22 22(21). Stems decumbent, sharply bent from apex of caudex 23 — Stems ascending to erect, not sharply bent from caudex apex 24 23(22). Herbage glaucous, sparingly hairy; rays 15-22; basal leaf bases greatly ex- panded, long-ciliate; plants of eastern Washington County E. canaani — Herbage green, not especially glaucous, sparingly to moderately hairy; basal leaf bases not greatly expanded, short-ciliate; plants of broad distribution ....£. eatonii 24(22). Cauline leaves moderately well developed, the basal ones linear-oblanceolate; involucres 9-12 mm wide; plants of lower elevations E. engelmannii — Cauline leaves much reduced, the basal ones spatulate; involucres less than 8 mm wide 25 25(24). Basal leaves acute or acutish; rays blue to red-purple; pappus subequal to disk corollas; plants widely distributed E. tener — Basal leaves obtuse to rounded; rays white to pink; pappus shorter than disk corollas; plants of the Bear River Range, Cache County E. cronquistii 266 Great Basin Naturalist Vol. 43, No. 2 KEY IV. Plants perennial, green; achenes mostly 2-nerved; involucres mostly lacking; long, spreading multicellular hairs. 1. Plants substoloniferous; leaves spatulate; involucres mainly less than 8 mm wide; plants of hanging gardens in San Juan County E. kachinensis — Plants not substoloniferous; leaves various; involucres mainly over 8 mm wide; plants not or seldom of hanging gardens, variously distributed 2 2(1). Cauline leaves ample, usually lanceolate or broader; plants tall and erect (more or less asterlike) 3 — Cauline leaves definitely reduced upward, mostly linear to oblanceolate, or broader in some low species; stems often spreading or decumbent 5 3(2). Rays mainly 2-3 mm wide E. peregrinus — Rays 1-2 mm wide 4 4(3). Cauline leaves glabrous or minutely glandular, not ciliate, subequal to or short- er than the internodes E. superbus — Cauline leaves ciliate or otherwise pubescent, sometimes also glandular, usual- ly longer than the internodes E. speciosus 5(2). Pubescence of the stem widely spreading or glandular-scabrous 6 — Pubescence of the stem appressed, ascending, or lacking 8 6(5). Involucre canescent with fine white hairs, sometimes also glandular .... E. caespitosus — Involucre glandular and more or less spreading hairy or strigose 7 7(6). Stems hirsute with short spreading hairs, conspicuously decumbent; involucres glandular and spreading hairy E. jonesii — Stems glandular-scabrous, ascending or erect; involucres glandular (rarely sparingly strigose) E. nauseosus 8(5). Basal leaves broadly oblanceolate or usually broader, the blade well-developed, usually abruptly contracted to the petiole 9 — Basal leaves linear to oblanceolate or spatulate, tapering gradually to the pet- iole 11 9(8). Rays purple; achenes 4- to 7-nerved; pappus simple E. peregrinus — Rays various; achenes 2-nerved (occasionally more, but rays then pale and pap- pus double) 10 10(9). Stems essentially scapose, the upper bracts linear; plants known from the Wasatch Mountains E. garrettii — Stems subscapose, the upper bracts oblong; plants rather broadly distributed .... E. leiomeris 11(8). Peduncles and involucres densely glandular, not hairy; stems glabrous or essen- tially SO; plants of the Wasatch Mountains E. arenarioides — Peduncles not glandular, or, if so, the stem more or less hairy; involucres and distribution various 12 12(11). Bases of basal leaves neither enlarged nor of different texture than the blades; blades linear or linear-filiform; plants known from Cache and Daggett coun- ties E. filifolius April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 267 — Bases of basal leaves somewhat enlarged, membranous or thickened, or other- wise different from above; blades not linear 13 13(12). Leaves glabrous or nearly so, the hairs, if present, short and appressed .... E. leiomeris — Leaves hairy, the hairs spreading or curved-ascending 14 14(13). Plants subscapose; cauline leaves reduced to acicular bracts; plants of the Uinta Basin and west Tavaputs Plateau E. nematophyllus — Plants caulescent; cauline leaves well developed 15 15(14). Stems decumbent-ascending, commonly curved at the base; basal leaves sheathing basally; heads mainly solitary E. abajoensis — Stems erect or nearly so; basal leaves not especially sheathing; heads mainly 2-4 E. awapensis Erigeron abajoensis Cronq. Abajo Daisy. Perennial herb, with a taproot and stout cau- dex, the caudex branches clothed with brown marcescent leaf bases; stems decumbent to spreading at the base, 5-20 cm long, strigose to strigulose, the hairs ascending; basal leaves oblanceolate, 2-7 cm long, 2-6 mm wide, more or less sheathing basally; cauline leaves several to many, oblong to lance-oblong, mostly 0.6-2.5 cm long, 1.5-4 mm wide; heads solitary, less commonly 2-4; involucres 4-5.2 mm high, 7-12 mm wide, the bracts subequal or slightly imbricate, somewhat thickened dorsally, greenish brown, strigose to strigulose, the hairs multicellular; rays about 40-60, pink-purple to blue (or white), 3-8 mm long, 1-1.8 mm wide; pappus double, the inner of 12-20 bristles, the outer of setae or scales; achenes 2-nerved, hairy. Pinyon-juniper, ponderosa pine, and spruce- fir communities at 2135 to 3450 m in Gar- field, Piute, San Juan, and Wayne counties; endemic; 4 (i). Erigeron acris L. Bitter Fleabane. Short- lived perennial, with a slender taproot and poorly developed caudex; stems erect or de- cumbent at the base, 8-32 cm tall, spreading- hairy and more or less glandular; basal leaves spatulate-oblanceolate, 0.5-6.5 cm long, 2-10 mm wide, entire or sparingly toothed; cau- line leaves several to many, oblong to nar- rowly oblanceolate, lanceolate, or linear, mostly 0.8-7 cm long, 1-8 mm wide; heads solitary, or more commonly few to numerous, on short to elongate peduncles; involucres 4.5-8 mm high, 9-17 mm wide, the bracts imbricate, not especially thickened, green or tinged pink apically in some, sparingly hairy with spreading to ascending stiff multi- cellular hairs and beset with short glandular processes; rays numerous, pink or white, erect, about 2-4.5 mm long, the inner pistil- late flowers eligulate, with corolla tubular; pappus of ca 25-35 slender barbellate white to reddish bristles, surpassing the disk co- rollas; achenes 2-nerved, sparingly hairy. Lodgepole pine, spruce, and fir communities at 2800 to 3500 m in Duchesne, Summit, and Uintah counties; Alaska to Labrador, south to California, Colorado, Michigan, and Maine; circumboreal; 11 (ix). Varietal status of our few specimens is unclear. One of the speci- mens has few heads and has essentially eg- landular bracts, one is monocephalus and has glandular involucres, and the others are poly- cephalus and have glandular involucres. Names available are var. asteroides (Andrz.) DC. and var. debilis Gray, but there appear to be three rather poorly differentiated taxa involved. Decisions as to proper names must await further study. Erigeron annuus (L.) Pers. Annual Flea- bane. Plants annual, with slender taproots; stems erect, 6-12 (15) dm tall, sparingly to densely hirsute with long spreading hairs, be- coming appressed upward; basal leaves com- monly withered at anthesis, ovate to sub- orbicular, petiolate; cauline leaves numerous, lanceolate to oblong, mainly 1.5-8 cm long, 3-20 mm wide, serrate to entire; heads sever- al to numerous, in a leafy inflorescence; in- volucres 7.5-12 mm wide, 3-5 mm high, the bracts subequal or the outer somewhat short- er, greenish to brownish, acuminate-attenu- ate, glandular and sparingly villous-hirsute with multicellular hairs; rays ca 80-125, 268 Great Basin Naturalist Vol. 43, No. 2 white (rarely bluish), 4-10 mm long, 0.5-1 mm wide; pappus double; achenes 2-nerved, hairy. Roadsides, fields, and other disturbed sites at 1370 to 1830 m in Utah and Wasatch counties; widespread in the United States; Europe; 8 (ii). Erigeron aphanactis (Gray) Greene Hairy Daisy. [E. concinnus var. aphanactis Gray]. Perennials with definite branching caudex; stems decumbent to ascending or erect, 5-20 (30) cm tall, sparingly to copiously spreading- hirsute with multicellular hairs; basal leaves narrowly oblanceolate to spatulate, 0.5-8 cm long, 1-6 mm wide, petiolate; cauline leaves well developed or essentially lacking; heads solitary or several; involucres 7-15 mm wide, 3.5-6 mm high, finely to coarsely spreading- hirsute and sometimes also finely glandular; bracts subequal or somewhat imbricate, slen- der, acuminate, green or greenish brown, the midrib thickened; pistillate flowers present, tubular, eligulate, or sometimes with rays shorter than the disk; pappus double; achenes 2-nerved, sparsely hairy. This species is rep- resented in Utah by two rather weak varieties. 1. Plants essentially scapose; corolla lobes sometimes becoming reddish or pur- plish E. aphanactis var. congestus — Plants with leafy stems; corolla lobes commonly yellowish E. aphanactis var. aphanactis Var. aphanactis Salt desert shrub, sage- brush, pinyon-juniper, sagebrush, and moun- tain brush communities at 1300 to 2700 m in Beaver, Garfield, Juab, Piute, Sanpete, Se- vier, Washington, and Wayne counties; Ore- gon and Idaho south to California, Arizona, and Colorado; 21 (iii). Var. congestus (Greene) Cronq. [E. con- gestus Greene]. Juniper-black sagebrush, sagebrush, and aspen communities at 1830 to 2600 m in Garfield and Sevier counties; Cali- fornia; 3 (0). Erigeron arenarioides (D.C. Eaton) Gray [E. stenophyUus D.C. Eaton, not H. & A.; As- ter arenarioides D.C. Eaton ex Gray]. Pe- rennial herbs, with definite branching cau- dex, the caudex branches clothed with brownish marcescent leaf bases; stems as- cending to erect, 6-25 (30) cm tall, slender, glabrous or glandular below the heads; leaves glabrous or sparingly strigose, the basal ones linear-filiform to linear-oblanceolate, 1.5-6 (8) cm long, 0.5-2 (4) mm wide, entire; heads solitary or 2 or 3 (rarely more); involucres 7-9 mm wide, 3.7-5 mm high, the bracts im- bricate in several series, greenish brown, fine- ly glandular, the tips often purplish; rays 10-25, blue, 4-8 mm long, 0.8-1.8 mm wide; pappus of about 10-16 bristles, and with a few short setae; achenes 2-nerved, sparsely strigose. Crevices in limestone and quartzite outcrops, rarely in beach sand, at 1300 to 2440 m in Salt Lake, Tooele, Utah, and We- ber coumties; endemic; 8 (0). Erigeron rgentatus Gray Silver Daisy. [Wyomingia argentata (Gray) A. Nels.]. Pe- rennial herbs, with definite branching cau- dex, the caudex branches more or less clothed with brown marcescent leaf bases, the mid- ribs not especially persistent; stems erect, 9-28 (40) cm tall, finely strigose and silvery to gray-green; basal leaves tufted, spatulate to oblanceolate, 1.5-7 cm long, 1-4 (6) mm wide, petiolate, entire; cauline leaves re- duced upward; heads solitary; involucres 10-18 mm wide, 5.5-9 mm high, the bracts strongly imbricate, silvery strigose with ap- pressed antrorse hairs; rays ca 20-50, blue, lavender, or pink to white, 9-15 mm long, 1.6-2.8 mm wide; pappus double; achenes pi- lose. Salt desert shrub, sagebrush, pinyon- juniper, and mountain brush communities at 1600 to 2440 m in Beaver, Box Elder, Emery, Garfield, Iron, Juab, Millard, Piute, Sanpete, Sevier, Tooele, and Utah counties; Nevada, California; 29 (iii). Erigeron awapensis Welsh Awapa Daisy. Perennial herbs from a branching caudex, the caudex branches clothed with ragged brown marcescent leaf bases; stems erect or nearly so, 10-24 cm long, strigose, the hairs ascend- ing; basal leaves 1.5-7 cm long, 2-8 mm wide, not especially sheathing; cauline leaves well developed, oblong to linear, mostly 1-4 cm long, 2-4 mm wide; heads 2-4, rarely solitary; involucres 3-9 mm wide, 3.7-4.5 mm high, the bracts more or less imbricate, thickened near the base dorsally, greenish, April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 269 strigulose, the hairs multicellular; rays 35-45, pink-purple to pink (or white?), 5-6 mm long, 0.9-1.8 mm wide; pappus apparently simple, of 15-20 slender bristles, and with a few inconspicuous shorter setae in some; ach- enes 2-nerved, hairy. Pinyon-juniper and sagebrush communities at 2135 to 2260 m in Garfield and Wayne counties; endemic; 2 (1). Erigeron bellidiastrum Nutt. Plants annual (or biermial), the stems 3.5-32 (50) cm tall, erect or ascending, often intricately branched; herbage strigulose with incurved multicellular hairs; leaves mainly cauline, 0.5-4 cm long, 1-3 (6) mm wide, linear to ob- lanceolate, entire (or sparingly toothed to pinnatifid), petiolate, becoming sessile up- ward; heads solitary to numerous; involucres 5-11 mm wide, 3-5 mm high, the bracts hir- tellous with spreading curved multicellular hairs, thick, greenish, subequal, or the out- ermost shorter; rays ca 30-70, pink or white, 4-6 mm long, ca 1 mm wide; pappus of ca 15 deciduous bristles; achenes 2-nerved, hairy. Vanclevea-ephedra, blackbrush, and pinyon- juniper communities at 1125 to 1830 m in Garfield, Grand, Kane, San Juan, and Wash- ington counties; Wyoming and South Dakota to New Mexico and Texas; 23 (ii). Erigeron caespitosus Nutt. Tufted Daisy. [E. caespitosus var. laccoliticus Jones]. Pe- rermial herbs with a branching caudex, the caudex branches clothed with brown or blackish marcescent leaf bases; stems decum- bent at the base, 4-25 (30) cm tall, hirtellous with short spreading hairs (especially above); basal leaves oblanceolate to spatulate, gener- ally rounded to obtuse apically, 1-9 cm long, 2-13 mm wide, 1- to 3-nerved; cauline leaves reduced upward; heads solitary or few to sev- eral; involucres 9-18 mm wide, 4-7 mm high, the bracts subequal to imbricate, thick- ened on the back, green, strigose to pilose with multicellular hairs spreading laterally from the midrib; rays ca 30-100, blue, pink, or white, 5-15 mm long, 1-2 mm wide; pap- pus double; achenes 2-nerved, hairy. Sage- brush, pinyon-juniper, aspen, lodgepole pine, spruce, and tundra communities at 2135 to 3570 m in Daggett, Duchesne, Emery, Gar- field, Salt Lake, Sevier, Summit, Uintah, Utah, Wasatch, and Wayne counties; Alaska and Yukon south to Arizona, New Mexico, and Nebraska; 40 (viii). This is a variable spe- cies with many phases in Utah, each differing in stature, nature of vesture, size of heads, and other features that fail singly and in com- bination as diagnostic criteria. The species in- terfaces with E. nauseosus, E. abapensis, and probably with other taxa. Erigeron canaani Welsh Canaan Daisy. Perennial herbs from a simple (or branched?) caudex, this clothed with brown marcescent leaf bases, the taproot prominent; stems 7-20 cm tall, decumbent to ascending, sometimes purplish at the base, sparingly pubescent with ascending hairs; leaves pubescent like the stem, the basal ones tufted, 1-nerved, 1.4-9 cm long, 0.7-1 mm wide, linear, in- volute, sharply acute, conspicuously ex- panded and long-ciliate basally; cauline leaves numerous, reduced upward; heads 1-3; involucres 9-13 mm wide, 5.3-6.5 mm high; bracts imbricate, conspicuously glandu- lar and sparingly to moderately villous-pilose with multicellular hairs, green or variously suffused with purple; rays 15-22, white or pinkish, 3.5-5 mm long, 1.8-2.1 mm wide; pappus single, of ca 20 slender bristles; ach- enes 2-nerved, hairy. Ponderosa pine commu- nity at 1585 to 2075 m in Washington Coun- ty; endemic; 2 (i). The Canaan daisy is similar in general aspect to E. eatonii (q.v.). The in- volute linear glaucous leaves and few ray flowers appear to be diagnostic. Erigeron canus Gray Hoary Daisy. Pe- rennial herbs, with branching caudex, the caudex branches clothed with persistent leaf bases, the marcescent midribs prominent; stems erect or nearly so, 5-30 (35) cm tall, appressed strigose; basal leaves oblanceolate, mostly 1-6 cm long and 1-5 (7) mm wide, hairy like the stems; cauline leaves reduced upward; heads solitary (rarely up to 4 ); in- volucres 9-16 mm wide, 5-7 mm high, the bracts strigulose with ascending to spreading multicellular hairs and more or less glandu- lar, imbricate; rays ca 30-40, blue or white, 7-12 mm long, 0.8-1.4 mm wide; pappus double; achenes ca 8- to 14-nerved. Gravelly substrates of the Cedar Breaks (Wasatch) Formation in ponderosa pine and sagebrush communities at 2300 to 2500 m in Garfield County; Wyoming and South Dakota to Ari- zona and New Mexico; 3 (i). 270 Great Basin Naturalist Vol. 43, No. 2 Erigeron carringtonae Welsh Carrington Daisy. Pulvinate perennial herbs with a plu- ricipital caudex, the branches clothed with conspicuous brown to straw colored or ashy marcescent leaf bases; leaves mainly basal, thickish, 0.6-3.5 cm long, 1-5 mm wide, spatulate to oblanceolate, strigose to pilosu- lose, obtuse to rounded apically; scapes 2.5-8 cm tall; heads solitary; involucres 9.8-15 mm wide, 5.8-7 mm high, the bracts imbricate, suffused with purple or green, the inner greenish with scarious margins, spreading-vil- lous with long multicellular hairs; rays 18-30, pink to pink-purple, 6.8-8.2 mm long, 1.4-2.3 mm wide; pappus double, the inner of 25-35 barbellate bristles, the outer of short setae; achenes 2-nerved, pilose. Meadows and escarpment margins, commonly on Flagstaff Limestone at 3050 to 3355 m in Emery and Sanpete counties; endemic; 6 (i). Erigeron compactus Blake [E. pulvinatus Rydb.]. Perennial pulvinate herbs with a branching caudex, the caudex branches clothed with marcescent leaf bases; leaves mainly basal, 4-20 mm long, 0.6-1.4 mm wide, linear, finely strigose; scapes 2-10 cm tall; heads solitary; involucres 7-17 mm wide, 5-8.5 mm high, the bracts more or less imbricate, straw colored or greenish brown to green, hispidulous with short spreading hairs; rays mainly 15-50, white or pink, 7-11 mm long, 1.4-2.5 mm wide; pappus double; achenes 2-nerved. Two geographically segre- gated races are recognizable as varieties. Leaves yellowish green; involucral bracts appressed strigose; plants of the Great Basin E. compactus var. compactus Leaves grayish green; involucral bracts spreading-hispidulose; plants of the Colorado Drainage system E. compactus var. consimilis Var. compactus Pinyon-juniper community at 1830 to 2135 m in Beaver, Box Elder, Mil- lard, and Tooele counties; Nevada and Cali- fornia; 4 (0). Var. consimilis (Cronq.) Blake Salt desert shrub and pinyon-juniper communities in Daggett, Duchesne, Emery, and Wayne counties; Arizona; 11 (i). The general aspect of this variety is similar to that of E. pulcher- rimus (q.v.), with which it is sympatric in much of its range; the 2-nerved achenes and low subscapose stems are diagnostic. Erigeron compositus Pursh Fern-leaf Daisy. Perennial caespitose cushion plants, with a shortened pluricipital caudex, the cau- dex branches densely clothed with brown marcescent leaf bases; herbage glandular and more or less spreading-hairy; leaves mainly basal, mostly 2- or 3-temately lobed or dis- sected, 0.5-7 cm long; cauline leaves few and reduced upward, simple or ternate; stems subscapose, 2-20 (25) cm tall; heads solitary; involucres 8-20 mm wide, 5-10 mm high, the bracts glandular and spreading-hairy, com- monly purplish at the tips; rays lacking, or developed and 20-60, blue, pink, or white, to 12 mm long and 2 mm wide; pappus simple; achenes 2-nerved, villous-hirsute. Sagebrush, rabbitbrush, aspen, aspen-fir, lodgepole pine, spruce-fir, and alpine tundra communities at 2375 to 3965 m in Beaver, Box Elder, Dag- gett, Duchesne, Emery, Garfield, Iron, Juab, Millard, Piute, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, Wayne, and Weber counties; Alaska to Greenland, south to California, Arizona, Col- orado, South Dakota, and Quebec; 77 (xvi). This is an extremely variable apomictic spe- cies, with rare sexual individuals. Our mate- rial has been assigned to var. glabratus Ma- coun, which is separable from the type variety only problematically. Erigeron coulteri T.C. Porter in Port. & Coult. Coulter Daisy. Perennial herbs from a rhizome or caudex; stems more or less spreading-hairy, mainly 1-6 dm tall; basal and cauline leaves ample or the cauline ones somewhat reduced, entire or toothed, the largest 6-15 cm long, 1-2.5 cm wide, oblan- ceolate to elliptic, lanceolate, oblong, or ovate; heads solitary or 2 or 3; involucres 10-15 mm wide, 6-10 mm high, the bracts densely white hirsute below with hairs hav- ing purplish black cross-walls, at least near the base, glandular to the tips; rays 40-80, ca 10-15 mm long, white to pink-purple; pap- pus simple; achenes sparsely hairy. Moist slopes in Salt Lake and Utah counties; Ore- gon to Wyoming, south to California, Ne- vada, and New Mexico; 8 (0). April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 271 Erigeron cronquistii Maguire Cronquist Daisy. Perennial herb, with short caudex branches clothed with brown leaf bases; stems 1.5-7 cm long, sparingly strigose; basal leaves 0.5-4 cm long, spatulate to oblanceo- late or elliptic, petiolate, sparingly strigose; cauline leaves few or wanting; heads solitary, sometimes 2; involucres 5-8 mm wide, 3-5 mm high, glandular and spreading-hirsute, the bracts imbricate, green, often suffused with purple; rays 10-25, white or pale pink, 5-6 mm long, 1.3-2.1 mm wide; pappus single, or with a few shorter outer ones; ach- enes 2-nerved, sparingly hairy. Limestone cliffs at 1750 to 2600 m in the Bear River Range, Cache County; endemic; 2 (0). This beautiful, tiny plant is a near congener of E. tener (q.v.). Erigeron divergens T. & G. Spreading Daisy. [E. divaricatus Nutt., not Michx.]. An- nual, biennial, or short-lived perennial herbs from taproots; stems branched from the base and above, pubescent with spreading hairs, 0.5-5 (7) dm tall; basal leaves oblanceolate to spatulate, mainly 1-7 cm long, 2-10 mm wide, spreading-hairy, petiolate, usually lack- ing at anthesis; cauline leaves reduced up- ward; heads several to numerous; involucres 7-11 mm wide, 4-5 mm high, finely glandu- lar and hirsute with long, spreading hairs, the bracts green, attenuate; rays ca 75-150, blue, pink, or white, ca 5-10 mm long, 0.5-1.2 mm wide, sometimes scarcely developed; pappus double; achenes 2 (4) -nerved, sparsely hairy. Riparian, rabbitbrush, sagebrush, pinyon-juni- per, mountain brush, ponderosa pine, and as- pen-spruce communities at 975 to 2900 m in Beaver, Cache, Daggett, Davis, Duchesne, Emery, Garfield, Grand, Iron, Kane, Millard, Piute, Salt Lake, San Juan, Sevier, Tooele, Uintah, Utah, Wasatch, Washington, and Weber counties; British Columbia to South Dakota, south to California, Arizona, and Mexico; 109 (xiii). Our materials have been segregated into two weak varieties differen- tiated as follows: var. cinereus Gray, with earliest flowers borne on long naked pe- duncles and plants later with long leafy sto- lons; and var. divergens, with earliest heads on leafy peduncles and plants not developing leafy stolons. The var. cinereus is evidently rare in Utah; 5 (ii). Erigeron eatonii Gray Eaton Daisy. [£. ea- tonii f. molestus Cronq., type from the Stans- bury Mountains]. Perennial herbs, from a short simple or branched caudex, this clothed with brown marcescent leaf bases, the tap- root prominent; stems 5-38 cm tall, decum- bent to ascending, usually purplish at the base, strigose or rarely more or less hirsute; leaves pubescent like the stem, the basal ones tufted, 1- (or more commonly) 3-nerved, acute, mainly 1.2-12 (15) cm long, 1-10 mm wide; cauline leaves numerous, reduced up- ward; heads 1-3 (7); involucres 8-15 mm wide, 5-8 mm high, the bracts imbricate, conspicuously glandular and more or less hir- sute with spreading-ascending multicellular hairs, green or the tips purplish; rays about 20-50, white to blue or pink, mainly 4-10 mm long, 1-2.5 mm wide; pappus single or with a few short outer setae; achenes 2 (3) -nerved. Sagebrush, mountain brush, pinyon- juniper, ponderosa pine, aspen, spruce-fir, and alpine tundra communities at 1890 to 3630 m in all Utah counties except for Box Elder and Morgan; Oregon to Wyoming, south to California, Arizona, and Colorado; 171 (xxv). This is a widespread and variable species, with variants differing in size, in head dimensions, and in nature of the pu- bescence. The hirsute phase from the Stans- bury Mountains has been designated as f. mo- lestus Cronq. Erigeron elatior (Gray) Greene Tall Daisy. [£. grandiflorus var. elatior Gray]. Perennial herbs, from a short caudex (seldom collected); stems mainly 4-7 dm tall, often purplish be- low, leafy throughout, spreading-hairy and more or less glandular above; leaves mainly 2.2-10 cm long, 6-28 mm wide, the lower- most smaller than the middle ones and com- monly withered at anthesis, ovate-lanceolate to lanceolate, entire, the lower petiolate, be- coming sessile and somewhat clasping up- ward; heads 1-3 (6); involucres 12-20 mm wide, 7-11 mm high, the bracts densely woolly-villous with long, flattened, shiny, multicellular hairs, some of which may have reddish purple cross-walls, subequal, long-at- tenuate apically, the tips glandular, purple, and reflexed; rays ca 75-150, pink or pink- purple (white), 12-20 mm long, 0.8-1.6 mm wide; pappus double; achenes 2-nerved, hairy. Meadows and openings in mountain 272 Great Basin Naturalist Vol. 43, No. 2 brush and spruce-fir communities at 2440 to 3050 m in the La Sal Mountains of Grand and San Juan counties; Colorado and Wyom- ing. This is a beautiful asterlike plant with equably leafy stems and densely villous in- volucres; 4 (0). Erigeron engelmannii A. Nels. Engelmann Daisy. Perennial herbs, with short branching caudex, this clothed with straw-colored to brown marcescent leaf bases; taproot defi- nite; stems 3-24 (30) cm tall, decumbent to erect, strigose or the hairs ascending, multi- cellular; basal leaves 1-6 (10) cm long, 1.5-5 mm wide, linear-oblanceolate, the blades hairy like the stems, the basal margins long and coarsely ciliate; cauline leaves reduced but well distributed upward; heads 1-4; in- volucres 7-12 mm wide, 4-7 mm high, the bracts hirsute and more or less glandular, subequal, green, with brownish midrib and scarious apices; rays ca 35-100, white (rarely pink or blue), 5-12 mm long, 0.6-2 mm wide; pappus double; achenes 2-nerved, hairy. Salt desert shrub, sagebrush, rabbit- brush, and pinyon-juniper communities at 1370 to 2200 m in Box Elder, Cache, Dag- gett, Duchesne, Grand, Juab, Millard, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uin- tah, and Utah counties; Oregon to Wyoming and Colorado; 28 (iii). Erigeron filifolius Nutt. Thread-leaf Daisy. Perennial herbs, with branching woody caudex, the caudex branches clothed with brownish marcescent leaf bases; stems 10-30 (50) cm tall, more or less strigose; leaves 1-8 cm long, 0.3-3 mm wide, linear or filiform, strigose, the cauline ones distributed along the stem but smaller than the basal ones; heads 1-several; involucres 5-15 mm wide, 4-6 mm high, the bracts villous to stri- gose and commonly glandular as well, sub- equal or somewhat imbricate, greenish; rays ca 15-75, blue to pink or white, 3-12 mm long, 1-2 mm wide; pappus single or with a few outer setae; achenes 2-nerved, more or less hairy. The species is reported from Utah (Cache County, Logan, C. P. Smith 1737 RM) by Cronquist (1947), also Daggett Coun- ty; British Columbia and Montana to Califor- nia and Nevada; 1 (0). Erigeron flagellaris Gray Trailing Daisy. Biennial or short-lived perennials, with a poorly developed caudex (if at all) and slen- der taproot; herbage strigose or with spread- ing hairs at stem base; stems 3-25 (40) cm tall, the fertile ones terminated by a solitary head, the sterile ones developed as leafy sto- lons; basal leaves 1-5 cm long, 1.5-8 mm wide, oblanceolate to spatulate; cauline leaves smaller upward, linear to oblanceo- late; heads solitary; involucres 7-13 mm wide, 3.5-5 mm high, the bracts with ap- pressed or spreading hairs, glandular, green to purplish; rays mostly ca 50-100, white, pink, or blue, 5-10 mm long, 0.8-1 mm wide; pappus double; achenes 2-nerved, hairy to al- most glabrous. Sagebrush, juniper, ponderosa pine, aspen, spruce-fir, and alpine meadow commimities at 1980 to 3180 m in Beaver, Carbon, Daggett, Duchesne, Emery, Gar- field, Grand, Iron, Kane, Millard, San Juan, Sanpete, Sevier, Summit, Washington, and Wayne counties; British Columbia to Ne- vada, Arizona, and Texas; 63 (xi). Erigeron formosissimus Greene Pretty Daisy. [E. frucetorum Rydb.]. Perennial herbs, with a simple or sparingly branched subrhizomatous caudex; herbage variously hirsute, glandular, or glabrous, the stems more or less glandular above, mainly 1.5-3 (4.5) dm tall; basal leaves the largest, mainly 2-10 (15) cm long, 4-10 (15) mm wide, ob- lanceolate to spatulate; cauline leaves com- monly much reduced upward, lanceolate to oblong or ovate; heads 1-6; involucres 10-20 mm wide, 5-8 mm high, the bracts subequal, linear, acuminate, glandular, and more or less hirsute; rays ca 75-150, 8-15 mm long, ca 1 mm wide, blue, pink, or white; pappus double; achenes 2-nerved, hairy. Meadows in aspen and mountain brush communities at 2440 to 1840 m in Grand, Iron, Salt Lake, San Juan, and Sevier counties; Alberta south to Arizona and New Mexico; 5 (i). The spe- cies is poorly known in Utah (reports of the species in Iron county are from Cronquist 1947). Erigeron garrettii A. Nels. Garrett Daisy. [£. controversus Greene]. Perennial sub- scapose herbs, with branching caudex, the caudex branches clothed with brown leaf bases; stems 3-23 cm tall, sparingly strigose; basal leaves 1.2-12 cm long, 3-13 mm wide, oblanceolate to spatulate, glabrous, sparingly April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 273 ciliate; cauline leaves lacking or greatly re- duced; heads solitary; involucres 8-17 mm wide, 5-8 mm high, the bracts finely strigose and obscurely glandular, moderately imbri- cate; rays ca 20-35, white to pink, 7-13 mm long, 1.4-2.7 mm wide; pappus double; ach- enes 2-nerved, hairy. Moist cliff faces and crevices at 2750 to 3570 m in Salt Lake, Utah, and Wasatch coimties; endemic; 17 (0). Erigeron glabellus Nutt. Smooth Daisy. Perennial or biennial herbs with simple or branched caudices, the caudex, when present, clothed with brown to blackish leaf bases; herbage strigose to hirsute; stems 1-6.5 dm tall, erect or nearly so; basal and lower leaves mainly 3-15 cm long, 3-18 mm wide, oblan- ceolate, entire or toothed, petiolate; middle cauline leaves lanceolate to linear, reduced upward; heads 1-12 (15), borne on bracteate peduncles; involucres 10-20 mm wide, 5-9 mm high, the bracts subequal to slightly im- bricate, acuminate, strigose to strigulose; rays ca 125-175, blue to pink, or white; pappus double; achenes 2-nerved, hairy. Meadows and stream sides at 1370 to 1770 m in Bea- ver, Cache, Daggett, Davis, Duchesne, Salt Lake, Uintah, Utah, and Wasatch counties; Alaska and Yukon, south to Utah, Colorado, South Dakota, and Wisconsin; 12 (0). This is a tall handsome daisy of lower elevations in Utah. Erigeron goodrichii Welsh Goodrich Daisy. Perennial herbs from a stout taproot and caudex, the caudex branches with dark brown marcescent leaf bases; stems 3-10 cm tall, decumbent-ascending to erect, spread- ing-hairy; basal leaves 0.4-6 cm long, 1.2-5 mm wide, narrowly oblanceolate, the veins not apparent, pilosulose, obtuse apically; cauline leaves more or less developed, but much reduced upward; heads solitary; in- volucres 10.5-18 mm wide, 6.4-7.8 mm high; bracts imbricate, spreading villous-pilose with multicellular hairs, thickened basally, green or the apices suffused purplish, the in- ner with scarious margins, the attenuate apices more or less glandular and sometimes spreading; rays 40-65, pink-purple to pink or white, 6.8-10.4 mm long, 1.5-2 mm wide; pappus apparently single, of 20-30 minutely barbellate bristles; achenes 2-nerved, pilose. Engelmann spruce krummholz and meadow communities, often on rock outcrops or talus at 3050 to 3400 m in Duchesne, Summit, Uintah, and Utah counties; endemic; 8 (0). Erigeron jonesii Cronq. Jones Daisy. Pe- rennial herbs, from a branching or simple caudex, the caudex branches clothed with blackish or dark brown marcescent leaf bases; herbage hirsute with short spreading hairs; stems mainly 10-25 cm tall, conspicuously decumbent and often purplish at the base; basal leaves 3-nerved, 1.5-8 cm long, 3-12 mm wide, oblanceolate to elliptic or spatu- late, petiolate, entire or toothed; cauline leaves smaller than the basal; heads 1-4; in- volucres 9-15 mm wide, 5-7 mm high, the bracts glandular and spreading-hairy, slightly thickened dorsally, more or less imbricate, green, with tips often purplish; rays ca 25-50, blue, pink, or white, 4-8 mm long, 1.4-1.8 mm wide; pappus single or with a few short outer setae; achene's 2-nerved, hairy. Sagebrush, pinyon-juniper, mountain brush, and alpine meadow communities at 1890 to 3350 m in Juab, Tooele, and Wash- ington counties; Nevada; 5 (iii). The Jones daisy simulates E. eatonii in habit and stature, but the definite spreading hairs of the herb- age are apparently definitive in most in- stances. Possibly it would best be treated at some infraspecific rank within E. eatonii. Erigeron kachinensis Welsh & Moore Kachina Daisy. Perennial herbs, from a short thick branching or simple caudex, the caudex branches clothed with brown marcescent leaf bases; herbage glabrous throughout; stems 6-18 cm tall, decumbent to erect; basal leaves 1.3-5 cm long, 2-13 mm wide, oblan- ceolate to obovate or spatulate, the blade ta- pering to the petiole, rounded or retuse api- cally, entire; cauline leaves 5-11, reduced upward; heads solitary or 2-4, the involucres 5-6 mm broad, 3.2-4 mm high, the bracts distinctly imbricate, some purplish at the tip, glabrous or minutely glandular; rays 10-15, white or pinkish, 3.5-5.5 mm long, 0.9-1.1 mm wide; pappus double; achenes 2-nerved, hairy. Seeps and hanging gardens at 1680 to 1890 m in White (type from Natural Bridges National Monument) and Dark Canyons, San Juan County, Utah, and Montrose County, Colorado; 3 (ii). This distinctive dwarf daisy is a Colorado Plateau endemic. Erigeron leiomerus Gray Glaber Daisy. Perennial herbs, from a branching caudex. 274 Great Basin Naturalist Vol. 43, No. 2 the caudex branches clothed with brown marcescent leaf bases; herbage glabrous or merely strigose; stems 4-12 (15) cm tall, de- cumbent to erect; basal leaves 1.3-7 cm long, 2-11 (15) mm wide, oblanceolate to spatulate or obovate, rounded to retuse apically, en- larged and often purplish basally, glabrous or strigose to glabrate; cauline leaves reduced upward, usually several, becoming acutish; head solitary, the involucres 7-13 mm wide, 4-6 mm high, the bracts somewhat imbricate, purplish overall or at tips, finely glandular; rays ca 15-60, purplish to blue or white, 6-11 mm long, 1.5-2.5 mm wide; pappus double; achenes 2-nerved, short-hairy. Talus slopes, boulder fields, and meadows in spruce and lodgepole pine and alpine tundra com- munities at 2950 to 3750 m in Beaver, Box Elder, Cache, Daggett, Duchesne, Juab, Piute, Salt Lake, Summit, Tooele, and Uintah counties; Nevada and Idaho to Wyoming, Colorado, and New Mexico; 27 (x). Erigeron linearis (Hook.) Piper [Dauco- pappus linearis Hook.]. Perennial herbs from a pluricipital caudex, the branches of the caudex clothed with broad clasping brownish marcescent leaf bases; herbage strigose; stems 5-20 cm tall; basal leaves 1-9 cm long, 0.5-3 mm wide, linear to linear-oblanceolate, acute, the bases enlarged, more or less sheathing, straw colored and strongly ciliate; cauline leaves reduced upward; heads soli- tary or 2 or 3; involucres 8-13 mm wide, 4-7 mm high, strigose-villous with multicellular hairs and more or less glandular; bracts sub- equal to somewhat imbricate, green or green- ish to straw colored, attenuate, thickened dorsally; rays ca 20-45, yellow, 4-11 mm long, 1.3-2.5 mm wide; pappus double, the inner of 10-20 barbellate bristles, the outer of scales; achenes 2-nerved, short hairy. Sage- brush and juniper communities at 1675 to 2000 m in Box Elder County; British Colum- bia, Washington, and Oregon, east to Idaho and Wyoming, and south to Nevada; 1 (0). Erigeron hnchophyllus Hook. Short-lived perennial or biennial (?) herbs, with slender taproots and subfibrous roots from a poorly developed caudex; stems decumbent to erect, 5-55 (60) cm long, sparsely to densely spreading-hairy; basal leaves oblanceolate to spatulate, 1.2-11 (15) cm long, 2-12 mm wide; cauline leaves several to many, mostly 0.6-8 cm long, 2-6 mm wide; heads few to numerous, rarely solitary, borne on nearly erect peduncles; involucres 4-9 mm high, 7-17 mm wide, the bracts evidently imbri- cate, not especially thickened basally, green- ish to brownish or yellowish, the tips com- monly purplish, sparsely to moderately strigulose with multicellular hairs; rays nu- merous, white or pinkish, about 2-4 mm long, lacking inner eligulate pistillate co- rollas; pappus of ca 20-30 slender barbellate white bristles, surpassing the disk corollas; achenes 2-nerved, sparsely hairy. Marshes, stream banks, seeps, and wet meadows at 1370 to 2900 m in Beaver, Daggett, Du- chesne, Garfield, Grand, Juab, Piute, Rich, Salt Lake, Sanpete, Sevier, Summit, Tooele, Utah, and Washington counties; Alaska and southern Yukon, south to California and New Mexico, and east to Quebec and South Da- kota; 39 (vii). Erigeron maguirei Cronq. Maguire Daisy. Perennial herbs, with a branching caudex, the caudex branches clothed with brown to straw-colored marcescent leaf bases; herbage spreading hirsute; stems 7-18 cm high, de- cumbent to sprawling or erect; basal leaves 2-5 cm long, 3-8 mm wide, oblanceolate to spatulate, rounded apically; cauline leaves well developed, but somewhat reduced up- ward, becoming acutish; heads solitary or 2-5; involucres 5-6.5 mm high, 7-11 mm wide, the bracts imbricate, not much thick- ened, green or yellowish, the inner less pu- bescent and with scarious purplish tips, all finely glandular also; rays 12-20, white or pinkish, ca 6-8 mm long, 1.1-2 mm wide; pappus of 13-25 slender barbellate sordid bristles, with a few shorter outer ones; ach- enes 2-nerved, hairy. Canyon bottoms in Wingate (?) and Navajo formations at 1640 to 1740 m in Emery and Wayne counties; en- demic; 5 (ii). For the past four decades the Maguire daisy was known officially from the type locality in the San Rafael Swell in Emery County. Now, other material has been discovered at BRY and relocated in the field, which is distinguishable only technically from specimens at the type locality. These latter specimens tend to have more heads per stem, have narrower ray corollas, and shorter disk corollas. All of these may be the result of ecological responses, but they are recognized as var. harrisonii Welsh. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 275 Erigeron mancus Rydb. [E. pinnatisectus (Gray) A. Nels. var. insolens Macbr. & Pays.]. Pulvinate caespitose subscapose perennials from a usually branched caudex, the caudex clothed with dark brown to straw-colored marcescent leaf bases; herbage more or less hirtellous and puberulent or minutely glandu- lar; stems mainly 2-6 cm long, erect or as- cending; basal leaves 1.2-4 cm long, 2-4 mm wide, pinnatifid, the lobes lanceolate, some- times again lobed; cauline leaves much re- duced; heads solitary; involucres 5-6.5 mm high, 7-12 mm wide, glandular, villous with multicellular hairs, the bracts subequal, some- what thickened basally, the acuminate tips often purplish; ray flowers lacking; pappus simple or nearly so, of 20-30 bristles; achenes 2-nerved, hairy. Alpine forb and grass-sedge communities at 3050 to 3660 m in the La Sal Mountains, astride the Grand-San Juan County line; endemic; 3 (0). Erigeron melanocephalus (A. Nels.) A. Nels. [£. uniflorus var. melanocephalus A. Nels.]. Perennial herbs, from a simple or branched caudex, the caudex branches clothed with dark brown marcescent leaf bases; herbage more or less villous with mul- ticellular hairs; stems commonly 5-12 cm tall, erect; basal leaves 0.8-6 cm long, oblan- ceolate to spatulate, rounded or retuse api- cally; cauline leaves much reduced upward; heads solitary; involucres 10-14 cm wide, 5-9 mm high, the bracts more or less densely villous with multicellular hairs, the cross- walls black or dark purple, equal, attenuate, green, with purplish tips or purplish through- out; rays 50-70, white or pink, 7-11 mm long, 1.2-2 mm wide; pappus single, of ca 20-25 bristles; achenes 2-nerved, sparsely hairy. Alpine meadows at 3355 to 3720 m in Grand and San Juan counties (La Sal Moun- tains); Wyoming, Colorado, and New Mexi- co; 4 (0). Specimens from the Uinta Moun- tains, which have involucral hairs with purple cross-walls, have been assigned here previously, but they seem to represent noth- ing more than phases of E. simplex (q.v.). Erigeron nanus Nutt. Dwarf Daisy. [E. in- amoenus A. Nels.]. Perennial herbs, from a branching caudex, the caudex branches clothed with imbricate ashy to straw-colored marcescent leaf bases; stems 3-8 cm high, vil- lous with contorted multicellular hairs, sub- scapose; basal leaves linear-oblanceolate. 1.2-4 cm long, 1-2 mm wide, hirtellous to sparingly villous or glabrous, ciliate toward base with spreading long hairs, the bases con- spicuously enlarged; heads solitary; in- volucres 7-13 mm wide, 5-8 mm high, long- villous with multicellular hairs and more or less finely glandular; bracts subequal, the midstripe brown to purplish, the margins green to scarious or purplish; rays 15-35, purplish, 5-10 mm long, 1.3-2.4 mm wide; pappus of 15-25 bristles and some outer setae; achenes 2-nerved, hirsute. Sagebrush and sagebrush-grass communities, often on windswept ridges, at 2135 to 3270 m in Box Elder and Daggett counties; Idaho and Wyoming; 5 (0). Erigeron nauseosus (Jones) A. Nels. Marys- vale Daisy. [E. caespitosus Nutt. var. nau- seosus Jones, type from near Marysvale]. Pe- rennial herbs, from a stout-" branching brittle caudex, the branches clothed with dark brown marcescent leaf bases, the taproot sim- ilarly colored; stems 6-25 cm tall, ascending to erect, glandular-scabrous; basal leaves 2.3-10 cm long, 2-15 mm wide, oblanceolate to spatulate, rounded apically, tapering to the petiole, commonly 3-nerved; cauline leaves well developed, only gradually re- duced upward; heads solitary, rarely 2; in- volucres 8-17 mm wide, 5-8 mm high, finely glandular (rarely sparingly strigose as well); bracts imbricate, somewhat thickened, often purplish, attenuate; rays 30-60, white or purplish, 6-12 mm long, 1.3-2 mm wide; pappus double, the inner of 12-23 bristles, the outer of inconspicuous setae; achenes 2- nerved, hairy. Crevices in limestone, quartz- ite, and igneous outcrops, and in talus in pin- yon-juniper, sagebrush, mountain brush, and Douglas fir- white fir communities at 1830 to 2900 m in Beaver, Garfield, Millard, Piute, and Sevier counties; White Pine County, Ne- vada; a Great Basin endemic; 24 (iii). Erigeron nematophyllus Rydb. Needleleaf Daisy. Perennial herbs, from a branching caudex, the caudex branches clothed with fibrous ashy to brown marcescent leaf bases; herbage strigose to subglabrous; stems 4-15 cm tall; basal leaves 1-8 cm long, 1-3 mm wide, linear to linear-oblanceolate, ciliate near the enlarged sheathing base; cauline leaves few and reduced, not especially ex- ceeding the basal cluster; heads solitary; in- volucres 6-13 mm wide, 4-6.5 mm high; 276 Great Basin Naturalist Vol. 43, No. 2 bracts more or less imbricate, moderately strigulose, green or brown, the inner often with scarious margins and purplish tips; rays 15-55, white (less commonly pink), 4-8 mm long, 1.2-2.3 mm wide; pappus of ca 15-25 bristles; achenes 2-nerved, shortly hairy. Sagebrush, mountain brush, and pinyon-juni- per communities, often on Green River Shale, at 2280 to 2870 m in Carbon, Daggett, Duchesne, and Uintah counties; Wyoming and Colorado; 7 (i). Erigeron peregrinus (Pursh) Greene Strange Daisy. [E. callianthemus Greene; E. peregrinus ssp. callianthemus (Greene) Cronq.; £. regalis Greene; E. peregrinus var. eucallianthemus Cronq.; E. peregrinus var. scaposus (T. & G.) Cronq.; E. salsuginosus var. scaposus T. & G.]. Perennial herbs, from a rhizome, the rhizome sometimes short, dark brown; stems 0.9-5.5 (7) dm tall, glabrous or sparingly to moderately villous below, often densely villous below the heads; basal leaves 2-16 (20) cm long, 0.8-3.2 (4.5) cm wide, ob- lanceolate to spatulate or obovate, tapering or abruptly contracted to the petiole, obtuse or rounded to acute apically, glabrous or rarely sparingly villous, ciliate; cauline leaves reduced upward, becoming sessile and more or less clasping; heads solitary, or 2-6; in- volucres 12-22 (25) mm wide, 6-9 (11) mm high; bracts subequal, reflexed at the attenu- ate apices, glandular and purplish through- out; rays ca 30-75, 8-17 (25) mm long, 1.8-4 mm wide, rose-purple to white; pappus of ca 20-30 bristles, sometimes with a few outer setae; achenes 4- to 7-nerved, sparingly hairy. Aspen, spruce-fir, lodgepole pine, and sedge communities at 2280 to 3570 m in Box Elder, Cache, Duchesne, Garfield, Grand (?), Salt Lake, San Juan, Summit, Uintah, Wasatch, Weber, and Washington counties; Alaska south to California and New Mexico; 57 (x). Our materials were segregated by Cronquist (1947) into a dwarf alpine var. scaposus (T. & G.) Cronq. and a taller montane var. eu- callianthemus. On the basis of the rather abundant materials at hand, there does not seem to be any means of recognition of those taxa, except arbitrarily. Thus, all our speci- mens are herein considered as belonging to ssp". callianthemus (Greene) Cronq. var. callianthemus. Erigeron proselyticus Nesom Professor Daisy. [E. flagellaris Gray var. trilobatus Ma- guire ex Cronq.]. Perennial herbs, from a sub- rhizomatous or substoloniferous caudex, the caudex branches with weakly persistent brown marcescent leaf bases; stems 14-25 cm tall, decumbent to ascending or erect, spar- ingly strigose; basal leaves 0.5-6.5 (7.5) cm long, 2-11 mm wide, oblanceolate to spatu- late or linear, entire to pinnately few toothed or lobed, glabrous to sparingly strigose, acute to obtuse or rounded apically; cauline leaves gradually to abruptly reduced upward, entire or the lower few toothed; heads 3 to several; involucres 3.5-7 mm wide, 2.5-4.5 mm high, sparingly to moderately hirtellous; bracts subequal, brown, suffused with purple, or the inner greenish, with chartaceous margins; rays 22-46, white to purplish, 5.4-8.5 mm long, 1-1.4 mm wide; pappus double, the in- ner of 10-19 bristles, the outer of short setae; achenes 2- or 4-nerved, sparsely hairy. Bristlecone pine, spruce-fir, and aspen com- munities on sandstone and marly limestone formation:; at 2440 to 3050 m in Iron and Kane counties; endemic; 8 (i). Erigeron pulcherrimus Heller Basin Daisy. Perennial herbs, from a branching caudex, the caudex branches with exfoliating brown- ish bark, not especially clothed with per- sistent leaf bases; herbage silvery or grayish strigose; stems (5) 9-32 (35) cm tall, erect; basal leaves 0.8-7 cm long, 1-3 (5) mm wide, linear to linear-oblanceolate; cauline leaves reduced upward, but generally developed to stem middle or above; heads solitary; in- volucres 10-20 mm wide, 6-9 mm high, coarsely villous with spreading-ascending, multicellular hairs, obscurely glandular api- cally; bracts imbricate, greenish, the midrib often brown, the margins chartaceous, acumi- nate-attenuate, especially the inner; rays ca 25-60, white, pink, or violet, 8-15 mm long, 2-3.7 mm wide; pappus of ca 30-50 bristles, the outer series more or less developed; ach- enes (2-) 3- to 5-nerved, densely hairy. Salt desert shrub and pinyon-juniper communities on saline and seleniferous clays, clay-silts, and gravelly pediments at 1310 to 2105 m in Car- bon, Duchesne, Emery, Grand, Uintah, and Wayne counties; Wyoming, Colorado, and New Mexico. Our materials have been treated as belonging to a wide-leaved (1.5-5 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 277 mm) var. wyomingia (Rydb.) Cronq. and a narrow-leaved (1-1.5 mm) var. pulcherrima. However, only arbitrary segregation appears to be possible, and it seems best not to at- tempt recognition of infraspecific taxa; 61 (xiv). Erigeron pumilus Nutt. Vernal Daisy. Pe- rennial herbs, arising from a caudex, the branches clothed with ashy to brown marces- cent leaf bases; herbage more or less hirsute with spreading hairs; stems 4-50 cm tall, leafy or subscapose; basal leaves 0.4-8 cm long, mostly 2-5 mm wide, linear-oblanceo- late to oblanceolate; cauline leaves well de- veloped, somewhat reduced, or much re- duced upward, or almost lacking; heads solitary or few to numerous; involucres 7-15 mm wide, 4-7 mm high, sparingly to densely spreading-villous with multicellular hairs; bracts subequal, acuminate to attenuate, green, with brownish midrib; rays mostly 50-100, white or pink to lavender, 6-15 mm long, 0.7-1.5 mm wide (or more); pappus double, the inner of 7-20 coarse bristles, the outer of evident bristles or scales; achenes 2- nerved, sparsely to moderately hairy. Black- brush, shadscale, sagebrush, pinyon-juniper, and mountain brush communities at 885 to 2960 m in all Utah counties; Washington to Saskatchewan, south to California, Arizona, New Mexico, and Kansas. Our highly variable material was segregated on technical charac- teristics by Cronquist (1947) into two sub- species, each with two varieties. The bulk of the Utah specimens belong to ssp. con- cinnoides Cronq., segregated in large mea- sure from the much less common and more northern ssp. intermedius Cronq. by the few- er (7-15, not 13-20) inner pappus bristles and evidently puberulent (not glabrous or slightly puberulent) corolla tubes. The varieties inter- medius (var. euintermedius Cronq.) and grac- ilior Cronq. of ssp. intermedius are only arbi- trarily separable by stem thickness and head number. The weakly segregated varieties within ssp. concinnoides, var. concinnoides (var. euconcinnoides Cronq.) and var. con- densatus (D. C. Eaton) Cronq., differ in de- gree of development of cauline leaves, with the former having more equably leafy stems and the latter tending to be subscapose. It seems best to treat our material as belonging to two variable taxa; ssp. intermedius and ssp. concinnoides; 212 (xxvi). Erigeron religiosus Cronq. Religious Daisy. Short-lived perennial (or biennial?) herbs from a slender taproot and poorly developed caudex; herbage more or less strigose and glandular below the heads; stems 6-35 cm tall, decumbent-ascending to erect; basal leaves 2-5.5 (7) cm long, 2-8 mm wide (or more), oblanceolate to spatulate, entire or some pinnately toothed or lobed; cauline leaves gradually reduced upward; heads 2 to numerous; involucres 5.5-7.5 mm wide, 2-3.5 mm high, sparingly to moderately villous and more or less glandular; bracts with brown midrib, somewhat thickened, scarious api- cally; rays 35-85, white or pinkish, 3.4-6.8 mm long, 0.5-1.4 mm wide; pappus double, the inner of 6-12 bristles, the outer of short setae; achenes 2-nerved, sparsely hairy. Pon- derosa pine-oak and pinyon-juniper commu- nities at 1525 to 1830 m in Kane and Wash- ington (type from Clear Creek Canyon) counties; endemic; 9 (iv). Erigeron simplex Greene Greene Daisy. Perennial herbs, from a simple or branched caudex, the caudex clothed with dark brown marcescent leaf bases; herbage more or less viscid-villous with multicellular hairs; stems commonly 2-15 (20) cm tall; basal leaves 0.8-6 (8) cm long, 2-10 (13) mm wide, oblan- ceolate to spatulate, obtuse to abruptly acute or mucronate apically; cauline leaves re- duced; heads solitary; involucres 8-22 mm wide, 5-10 mm high, moderately to densely villous and somewhat viscid, the hairs with clear to reddish purple or purplish black cross-walls; bracts equal, suffused with purple or green, appressed or some reflexed; rays 50-125, blue-purple to pink (or white), 7-11 mm long, 1.2-2.5 mm wide; pappus double, the inner of ca 10-15 barbellate bristles, the outer of conspicuous setae; achenes 2-nerved, sparsely hairy. Lodgepole pine, Engelmann spruce, alpine fir, and alpine meadow and tundra communities at 3355 to 3660 m (in Deep Creek, Tushar, La Sal, and Uinta mountains) in Beaver, Daggett, Duchesne, Juab, Piute, San Juan, Summit, and Uintah counties; Oregon to Montana, south to Ne- vada, Arizona, and New Mexico; 30 (vi). Our variable materials include specimens with purplish-black cross-walls of the multicellular hairs, especially on the involucres and below the heads. These have been placed with the 278 Great Basin Naturalist Vol. 43, No. 2 similar and related E. melanocephalus (q.v.), but differ in shape of lower leaves and gener- al aspect of the plants. Erigeron sionis Cronq. Zion Daisy. Low perennial herbs, with short stoloniferous branches arising from a slender taproot; stems 1.5-13.5 cm long, decumbent to erect, glabrous or appressed pubescent; basal leaves 0.5-3.5 cm long, 2-10 mm wide, oblanceo- late to obovate, entire or more commonly 3- to 5-lobed, glabrous or sparsely strigose; heads solitary or 2 to several; involucres 5-7 mm wide, 2-3 mm high, glandular and sparsely to moderately spreading-hairy; bracts suffused purplish or the inner green with chartaceous margins; rays 23-38, white, the midstripe below purplish, 3.9-6.1 mm long, 1-1.6 mm wide; pappus double, the in- ner of 7-13 bristles, the outer of slender setae; achenes 2-nerved, sparsely pubescent. Seeps and hanging gardens in ponderosa pine and riparian communities in Navajo and Wingate sandstones at 1350 to 1600 m in Zion National Park, Washington and Kane (?) counties; endemic; 3 (i). Erigeron speciosus (Lindl.) DC. Oregon Daisy. Rhizomatous perennial herbs with the caudex more or less developed; stems 1.5-9 cm tall, ascending to erect, spreading-hairy to subglabrous or glandular above; basal leaves often lacking at anthesis, the lower- most cauline ones oblanceolate to spatulate, petiolate, commonly 5-15 cm long, 4-20 mm wide; middle cauline leaves lanceolate to oval, oblanceolate, or elliptic, 2-11 cm long, 5-28 mm wide; upper leaves gradually to markedly reduced, lanceolate to obliquely ovate, ciliate, the surfaces glabrous, spread- ing-hairy, or glandular (or a combination); heads 1-15 (or more); involucres 11-22 mm wide, 5.5-9 mm high, glandular, with a few long hairs, or more or less spreading-hairy; bracts subequal, acuminate or attenuate, the tips more or less spreading, often suffused purplish; ray flowers ca 75-150, pink, pink- or blue-purple, or white, 7-18 mm long, 0.7-1 mm wide; pappus double, the inner of 20-30 bristles, the outer of more of less evi- dent setae; achenes 2- to 4-nerved, hairy. The speciosus complex in Utah, as herein inter- preted, consists of four variable, and more or less intergrading, largely sympatric in- fraspecific taxa. All have been treated pre- viously at specific rank, or they have been treated within E. speciosus, in part. Cronquist (1947) discussed the problem of in- termediacy in the complex but hesitated to combine the taxa because "such a treatment would distort the facts as well as being un- wieldy." It is here contended that they are unwieldy apart; it seems therefore best to combine them as follows: 1. Leaves spreading-hairy on one or both surfaces; involucres spreading-hairy and more or less glandular E. speciosus var. mollis — Leaves glabrous on both surfaces or minutely glandular, or with minute stri- gose hairs, rarely with a few spreading multicellular hairs 2 2(1). Leaves glandular on the surfaces (especially the upper ones), and also ciliate .... E. speciosus var. uintahensis — Leaves glabrous on both surfaces, ciliate 3 3(2). Involucral bracts merely glandular, rarely also somewhat spreading-hairy; up- per leaves often ovate E. speciosus var. macranthus — Involucral bracts glandular and commonly also spreading-hairy; upper leaves lance-attenuate E. speciosus var. speciosus Var. macranthus (Nutt.) Cronq. [£. grandi- florus Nutt., not Hook.; E. macranthus Nutt.]. Sagebrush, snowberry, aspen, spruce- fir, and alpine meadow communities at 1760 to 3420 m in Beaver, Box Elder, Cache, Car- bon, Duchesne, Emery, Garfield, Grand, Juab, Kane, Millard, Piute, Rich, Salt Lake, Sanpete, Sevier, Tooele, Utah, Wasatch, Washington, and Weber counties; Washing- ton and Alberta south to Nevada, Arizona, and New Mexico; 104 (xv). This is our most common phase, but it is only arbitrarily sepa- rable from var. speciosus, to which it is com- pletely transitional. Var. mollis (Gray) Welsh comb. nov. [based on: Erigeron glabellus var. mollis Gray April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 279 Proc. Acad. Nat. Sci. Philadelphia 1863: 64. 1864.]. Aspen, spruce-fir, and meadow com- munities at 2070 to 3050 m in Carbon, Du- chesne, Garfield, Grand, Juab, Salt Lake, San Juan, Sanpete, Sevier, Uintah, Utah, and Wasatch counties; Montana to South Dakota, and south to New Mexico and Nebraska; 20 (i). This variety includes what has tradition- ally been called E. subtrinervis Rydb. Var. speciosus [Stenactis speciosa Lindl.]. Mountain brush, sagebrush, ponderosa pine, aspen, spruce-fir, and alpine meadows at 2040 to 3300 m in Duchesne, Garfield, Grand, Iron, Juab, Piute, San Juan, Sanpete, Sevier, and Utah counties; British Columbia and Montana, south to Nevada and New Mexico; 23 (iv). Var. uintahensis (Cronq.) Welsh comb, nov. [based on: Erigeron uintahensis Cronq. Bull. Torrey Bot. Club 70: 270. 1943]. Sage- brush, mountain brush, ponderosa pine, as- pen, lodgepole pine, spruce-fir, and alpine meadow communities at 2070 to 3420 m in Beaver, Carbon, Daggett, Duchesne, Juab, Piute, Sanpete, Sevier, Summit, Uintah, Utah, and Wasatch counties; Wyoming; 39 (vi). The glandular condition of the leaves varies in amount and position, and the Uinta phase passes by degree into other taxa of the speci- osus complex. Because of the intergradation it seems best that this most distinctive por- tion of the variation should be treated within an expanded E. speciosus. Erigeron superbus Greene ex Rydb. Splen- did Daisy. Rhizomatous perennial herbs and with the caudex more or less developed, the perennating branches bearing brown marces- cent leaf bases; herbage glabrous or glandular above and villous in some below the heads; stems 1-6 dm tall, erect; basal leaves smaller than the cauline and commonly present at anthesis, 3-15 cm long, 6-25 (33) mm wide, oblanceolate to obovate or spatulate, petio- late; middle cauline leaves somewhat smaller than the lower ones, oblong to elliptic or lan- ceolate, glandular (glabrous), the uppermost sessile and glandular, rarely some denticulate, not ciliate; heads 1-7; involucres 11-19 mm wide, 7-10 mm high; bracts subequal, glandular, sometimes with long spreading hairs near the base, acuminate, sometimes suffused purplish; rays 40-95, 1-2 mm wide. 12-20 mm long, rose-purple or white; pap- pus double, the inner of 20-25 pinkish or tawny bristles, the outer of setae; achenes 2- nerved, hairy. Aspen, Douglas fir, lodgepole pine, and spruce-fir communities at 2250 to 3050 m in Carbon, Garfield, Kane, Piute, San Juan, Summit, and Uintah counties; Wyom- ing south to Arizona and New Mexico; 18 (v). Erigeron tener Gray Thin Daisy. Perennial herbs, from a branching caudex, the slender branches with ashy to brownish marcescent leaf bases; herbage strigose; stems slender, decumbent, ascending, or erect, 3-15 cm tall; basal leaves 1-7.5 cm long, oblanceolate to elliptic, rhombic, or obovate, petiolate, acute to obtuse apically; cauline leaves much re- duced; heads solitary or 2 or 3; involucres 6-10 (12) mm wide, 3.5-5 mm high, glandu- lar and with spreading multicellular hairs; bracts imbricate, somewhat thickened, brownish, the inner membranous or some- what scarious, sometimes suffused with purple; rays ca 15-40, purplish or white, 4-8 mm long, 1-1.7 mm wide; pappus double, the inner of 15-30 bristles, usually with slen- der outer setae; achenes 2-nerved, hairy to subglabrous. Sagebrush, mountain brush, pin- yon-juniper, and white fir-Douglas fir com- munities, often on limestone outcrops at 1980 to 2900 m in Beaver, Juab, Millard, Rich, Sanpete, Tooele, and Utah counties; Oregon to Wyoming south to California and Nevada; 10 (i) Erigeron untermannii Welsh & Goodrich Untermann Daisy. Perennial pulvinate herbs with an intricately branched caudex, the cau- dex branches mainly basal, 0.8-3.3 cm long, 1-4 mm wide, narrowly oblanceolate to spatulate, pilosulose with ascending, often curved, hairs; scapes 2-6 cm tall; heads soli- tary; involucres 7-11 mm wide, 5-5.7 mm high, the bracts more or less imbricate, green, or the inner somewhat chartaceous, the mar- gins hyaline, the tips suffused with purple (sometimes throughout), densely hispidulous with short spreading hairs; rays 14-26, white, 4-6.5 mm long, 1.5-2.1 mm wide; pappus apparently single, of ca 20 slender fragile bristles; achenes 2-nerved, pilose. Pinyon- juniper community on calcareous shales and sandstones of the Uinta and Green River for- mations at 2135 to 2380 m in Duchesne and Uintah counties; endemic; 4 (0). 280 Great Basin Naturalist Vol. 43, No. 2 Erigeron ursinus D.C. Eaton Bear Daisy. Perennial rhizomatous sod-forming herbs, the perennating organs arising from short super- ficial branches clothed with brown marces- cent leaf bases; herbage subglabrous to stri- gose or variously ascending- or spreading- hairy; stems ascending, 5-25 (30) cm tall; basal leaves 1.2-12 cm long, 2-11 mm wide, oblanceolate to oblong, commonly acute or acutish apically, ciliate, the surfaces glabrous or variously hairy; cauline leaves reduced up- ward; heads solitary or 2 or 3; involucre 9-19 mm wide, 5-7 mm high, glandular and spreading-hairy with multicellular heads; bracts subequal, green or suffused purplish at the usually reflexed tips; rays ca 30-100, pink or blue-purple, 6-15 mm long, 1-2 mm wide; pappus double, the inner of ca 10-20 bristles, the outer of setae or scales; achenes 2-nerved, hairy. Sagebrush, aspen, lodgepole pine, and spruce-fir communities, often in forb-grass or forb-sedge meadows at 2440 to 3660 m in Beaver, Cache, Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Iron, Juab, Kane, Piute, San Juan, Sanpete, Sevier, Summit, Uintah, Utah, Wasatch, and Wayne counties; Idaho and Montana, south to Nevada and Arizona; 95 (x). Erigeron utahensis Gray Utah Daisy. Pe- rennial herbs from a branching caudex, the branches with grayish marcescent leaf bases and usually densely clothed with white vil- lous-pilose hairs; stems 10-60 cm tall, erect, appearing grayish or silvery due to strigose hairs; basal and lowermost cauline leaves 1.5-10 cm long, 1-6 mm wide, linear-oblan- ceolate, commonly withered or lacking at an- thesis; cauline leaves gradually reduced up- ward; heads solitary or few to many; involucres 5-15 mm wide; 3-7 mm high, stri- gose and often glandular apically; bracts im- bricate, brownish, the inner with scarious margins; rays ca 10-40, blue, pink, or white, 4-18 mm long, 1-2.7 mm wide; pappus double, the inner of ca 5lO-30 bristles, the outer of setae; achenes 4-nerved, more of less pilose. Two rather weakly separable varieties are present in Utah, as follows: 1. Stem bases not densely white-pilose; involucres mainly less than 8 mm wide; plants uncommon E. utahensis var. sparsifolius — Stem bases densely white-pilose; involucres commonly more than 10 mm wide; plants common E. utahensis var. utahensis Var. sparsifolius (Eastw.) Cronq. [E. spar- sifolius Eastw. and Wyomingia vivax A. Nels, both types from San Juan County] . Sandstone outcrops in salt desert shrub and pinyon-juni- per communities, often in shaded mesic areas, at 1220 to 1900 m in Emery, Garfield, Kane, and San Juan counties; Colorado and Arizona; 9 (iv). Var. utahensis [E. stenophyllus var. tetra- pleuris Gray]. Creosote bush, blackbrush, warm desert shrub, pinyon-juniper, and Mountain brush communities at 900 to 2000 m in Emery, Garfield, Grand, Iron, Kane, San Juan, Washington, and Wayne counties; Col- orado and Arizona; 75 (vii). Erigeron vagus Payson Payson Daisy. Caespitose perennial herbs, from a diffuse caudex, the branches commonly soboliferous; herbage moderately villous and glandular; leaves mainly basal, tufted at the apex of the caudex branches, 0.5-2.5 cm long, palmately 3-toothed or -lobed; heads solitary, sub- scapose; involucres 8-16 mm wide, 5-7.5 mm high, spreading-hairy and more or less glandular; bracts subequal, commonly suf- fused purplish at the attenuate apices; rays ca 25-35, white or pink, 4-7 mm long, 1-2 mm wide; pappus simple, of about 20 bristles; achenes 2-nerved, sparingly hairy. Ponderosa pine western bristlecone pine, and sedge-forb communities at 2375 to 3660 m in Garfield, Grand, Iron, and San Juan counties; Califor- nia east to Colorado; 9 (0). Erigeron wahwahensis Welsh Wah Wah Daisy. Perennial herbs, from a branched cau- dex, the caudex branches with conspicuous fibrous brown to ash-colored marcescent leaf bases; stems 15-40 cm long, decumbent to as- cending; basal leaves 3-18 cm long, 4-13 mm wide, linear-oblanceolate to oblanceolate or elliptic, 3-nerved, petiolate, appressed to spreading-hairy with curved hairs; cauline leaves reduced, sessile, and bracteate above; heads solitary or 2 or 3; involucres 13-17 mm wide, 6-7 mm high, spreading-villous with multicellular hairs, glandular apically; bracts April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 281 imbricate, green, the tips reddish, thickened basally; rays 30-40, pink or white, 5.5-7 mm long, 1.7-2.2 mm wide; disk corollas 3.5-4.2 mm long, the tube ca 2 mm long, the lobes 0.4 mm long; pappus of 15-20 bristles, with inconspicuous outer setae; achenes 2-nerved, short-hairy. Sagebrush, oak-maple, and pin- yon-juniper communities at 1670 to 2440 m in Beaver and Washington counties; endem- ic; 7 (iii). The Wah Wah daisy stands be- tween the distributions of E. pnesii and E. eatonii, and it shares features of both. The specimens examined from Washington Coun- ty have appressed strigose stems, and are highly variable. Those from the Wah Wah Mountains have spreading hairy stems. Addi- tional work is indicated. Eriophyllum Lag. Annual or perennial woolly herbs; leaves alternate, entire or toothed to lobed; heads solitary or corymbosely clustered; radiate; rays few, pistillate and fertile, yellow or white; involucres campanulate or hemispher- ic; bracts 1 (apparently 2) -seriate, firm, erect; receptacle flat to low-conic, naked; disk flowers perfect, fertile, the tube glandu- lar or hairy; pappus of firm nerveless chaffy scales; style branches flattened; achenes 4- angled. Constance, L. 1937. A systematic study of the genus Eriophyllum. Univ. California Publ. Bot. 18: 69-136. 1. Plants perennial E. lanatum — Plants annual 2 2(1). Rays white; pappus of unequal scales E. lanosum — Rays yellow; pappus of equal scales or reduced to a short crown E. wallacei Eriophyllum lanatum (Pursh) Forbes Pe- 2.5-4.5 mm long, slender, sparsely strigulose. rennial herbs, the herbage tomentose; stems Creosote bush, blackbrush, and Joshua tree erect or decumbent from a ligneus base, communities at 700 to 900 m in Washington mainly 10-20 cm tall; leaves mainly 1-4 cm County; California, Nevada, and Arizona; 13 long, entire or 3- to 5-toothed or -lobed; (i). heads solitary or corymbose on naked pe- Eriophyllum wallacei (Gray) Gray [Bahia duncles 3-10 cm long; involucres cam- wallacei Gray]. Annual tomentose herb; panulate, 6-10 mm wide, 6-8 mm high; stems mainly 1-8 cm tall, simple or branched bracts 5-8 (10), carinate, distinct, the tips from the base; leaves 0.5-1.5 cm long, spatu- erect; rays 5-8 (10), yellow, 6-10 mm long, late to obovate, entire or 3-lobed; heads soli- 2-5 mm wide; pappus of 8-10 variable tary, turbinate-cylindric, on short peduncles; scales; achenes 2.5-4 mm long, 4-angled, var- involucres 4-6 mm wide, 5-7 mm high; iously glabrous, hairy, or glandular. Sage- bracts 6-10, ovate, distinct; rays 5-10, yel- brush community (reported from Utah in the low, 3-4 mm long, 2.5-3.5 mm wide; pappus Pacific Northwest Flora); British Columbia to of 6-10 scales or none; achenes ca 2 mm Montana, south to California, Nevada, and long, linear, hairy or glabrous. Larrea, black- Wyoming; 0 (0). Our material likely belongs brush and Joshua tree communities at 700 to to var. integrifolium (Hook.) Smiley. 900 m in Washington Co.; California, Ne- Eriophyllum lanosum (Gray) Gray [Ac- vada, Arizona, and Mexico; 32 (iii). Hnolepis lanosa Gray]. Annual floccose-to- „ ^ mentose herbs; stems mainly 2-10 cm tall, simple and erect or branching from the base; Perennial herbs; leaves alternate, opposite, leaves 0.5-1.8 cm long, 1-2 mm wide, linear or whorled, simple; heads discoid, the flowers to linear-oblanceolate, entire or essentially all perfect and tubular; involucres cylindric so; heads turbinate, solitary on named pe- to campanulate, the bracts striate, imbricate; dimcles 0.5-5 cm long; involucres 5-6.5 mm receptacle naked, mainly flat; anthers obtuse wide, 5-7 mm high; bracts 8-10, oblong, and entire basally, or minutely sagittate; style acute, distinct or nearly so; rays 5-10, white, branches with short stigmatic lines and an 3-5 mm long, 2.5-3.5 mm wide; pappus of ca elongate papillate appendage; pappus of nu- 5 slender hyaline awn-tipped scales; achenes merous capillary bristles; achenes 10-nerved. 282 Great Basin Naturalist Vol. 43, No. 2 1. Leaves alternate; plants of Box Elder County E. occidentale — Leaves opposite or whorled; plants of various distribution 2 2(1). Leaves opposite; flowers white to cream E. herbaceum — Leaves whorled; flowers purple or purplish E. maculatum Eupatorium herbaceum (Gray) Greene [E. in Box Elder and Tooele counties; Washing- ageratifolium var.? herbaceum Gray]. Pe- ton to Idaho, south to California and Nevada; rennial herbs from a woody caudex; stems 2 (0). 4-7 dm tall, branched above; herbage „ ^ scabrous-puberulent; leaves mainly opposite, the blades L5-6 cm long, 0.5-4 cm wide, White-tomentose annual herbs; leaves en- ovate, the bases cordate or truncate, coarsely tire, alternate; heads discoid, small, in capi- crenate-serrate, acute; heads numerous, in tate clusters; involucre reduced, the bracts dense corymbose clusters; involucres 3.5-5 resembling those of the receptacle; outer mm wide, 3-4 mm high; bracts green, pu- flowers pistillate, fertile, with tubular-flli- berulent, subequal; corollas white; achenes form corolla, in several series, the outer black, 1.5-2 mm long. Ponderosa pine and epappose and subtended by concave, partly spruce-fir communities at 1585 to 2745 m in enclosing bracts, the inner bractless and with Piute and Washington counties; California pappus of capillary bristles; central flowers and Arizona; 2 (ii). 2-5, apparently perfect, but often sterile, Eupatorium maculatum L. Joe-Pye Weed, bractless, with capillary bristles; achenes sub- [£. bnineri Gray]. Robust perennial herbs terete, nerveless. from short subrhizomatous caudices; stems Filago californica Nutt. Annual herbs, the mainly 6-15 dm tall, branching in the in- stems erect, simple or branched above, 0.5-3 florescence; herbage puberulent and glandu- dm tall; leaves 0.8-2 cm long, narrowly ob- lar-dotted; leaves in whorls of 3 or 4, mainly long to oblanceolate; heads ovoid, 3-4 mm 6-25 cm long and 1.5-7 cm wide, lanceolate high, subequal to involucrate leaves; bracts of to lance-elliptic or lance-ovate, sharply ser- outer pistillate flowers 8-10, tomentose, boat rate; heads numerous in corymbose clusters; shaped, the tips hyaline, the inner ones thin- involucres 3.5-5 mm wide, 6.5-9 mm high, ner and less hairy, the inner florets ca 12-20; the outer puberulent, the inner glabrous dor- inner achenes papillose. Warm desert shrub sally, often ciliate, purplish to straw colored; at 915 to 1070 m in Washington County; Ari- flowers purple; achenes ca 3 mm long, green zona and California; 3 (2). to brown, glandular-dotted. River and canal p, , banks, wet meadows, bogs, and seeps at 1370 •' to 1865 m in Box Elder, Cache, Kane, Uin- Annual herbs; leaves opposite, sessile, more tah, and Utah counties; British Columbia to or less connate; heads several to numerous, in Newfoundland, south to New Mexico, II- compact corymbose clusters; involucres cy- linois, and Michigan. Our material belongs to lindric; bracts carinate, striate, 2-5, subequal; var. bruneri (Gray) Breitimg; 15 (i). receptacles naked; ray flowers pistillate, fer- Eupatorium occidentale Hook. Perennial tile, commonly 1 per head, yellowish, incon- herbs from a rhizome and with a branching spicuous; disk flowers 2-5, perfect, fertile, caudex; stems 1.5-7 dm tall, often branched yellowish; anthers not caudate at the base; above; herbage scabrous-puberulent; leaves pappus none; achenes 8- to 10-ribbed, alternate, the blades mainly 1.5-6 cm long, glabrous. 0.6-3 cm wide, deltoid or deltoid-ovate, ser- Flaveria campestris J.R. Johnst. Plants rate or subentire; heads numerous, in com- 12-85 cm tall, simple or branched, glabrous pact corymbose clusters; involucres 3-5 mm or hairy at the nodes; leaves 1-8 cm long, wide and as high; bracts subequal, pu- 0.4-1.5 cm wide, lance-oblong to linear, ser- berulent, green or suffused with purple; flow- rate to subentire, commonly 3-veined, ers pink or purplish; achenes ca 3 mm long, glabrous; inflorescence leafy bracted; in- brown, glandular-dotted. Rock crevices and volucres 5-8 mm high, the longer inner talus (usually in quartzite) at 2135 to 2745 m bracts mostly 3, strongly keeled, glabrous; April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 283 rays ca 1-2 mm long; achenes black, ca 3 mm long. Sand bars, stream banks, and seeps at 1220 to 1680 m in Grand and Tooele coun- ties; Colorado to Missouri, south to New Mexico and Texas; 7 (iii). Gaillardia Foug. Perennial (or biennial or annual) herbs; leaves alternate or mainly basal, entire or pinnatifid; heads radiate, the rays yellow, 3- lobed, neuter or sometimes pistillate and fer- tile; involucres 2- or 3-seriate, herbaceous, more or less spreading, reflexed in fruit; re- ceptacle convex, with numerous setae; disk flowers perfect, fertile; anthers auricled at the base; pappus of 5-10 scarious, awned scales; achenes broadly obpyramidal, long- hairy. 1. Disk flowers purple or purplish 2 — Disk flowers yellow 3 2(1). Base of involucral bracts densely long-villous or the corolla lobes 5-11 mm long, or both; plants mainly montane in northeastern Utah G. aristata — Base of involucral bracts not especially hairy, the corolla lobes mainly less than 5 mm long; plants of lower elevations in southeastern to south- western Utah G. pinnatifida 3(1). Stems with well developed, pinnately dissected cauline leaves; plants of canyon bottoms of the Tavaputs Plateau G. flava — Stems subscapose, or, if the cauline leaves well developed, merely toothed or lobed, and plants mainly of other distribution 4 4(3). Pappus scales broadly oblong or oval, awnless or abruptly short awned; plants annual, reported for southern Utah G. arizonica Greene — Pappus scales oblong-lanceolate, awned; plants perennial, rarely flower- ing the first year 5 5(4). Leaves mainly basal, entire or rarely some of them toothed or lobed G. parryi — Leaves cauline, toothed, lobed or entire G. spathulata Gaillardia aristata Pursh Blanketflower. Perennial herbs from a slender taproot; stems 20-80 cm tall, commonly foliose to middle or above, less commonly with basal leaves only; leaves 1.5-16 cm long, 3-25 mm wide, ob- long to oblanceolate or elliptic, entire or toothed to pinnatifid, puberulent and spar- ingly long-villous with multicellular hairs; heads solitary or few, long peduncled; disk mainly 2-2.5 cm wide, purple; involucral bracts (and/or peduncle apex) commonly long-villous basally, green or suffused with purple, attenuate; rays 6-16, yellow, often purplish at the base, the lobes 5-12 mm long; setae of receptacle well developed; disk co- rollas densely woolly-villous, the hairs with reddish purple cross-walls, often obscuring the attenuate lobes; pappus of slender lance- attenuate scales, the caudate apex entire; achenes ca 1.5 mm long, ruffous-pilose. Pin- yon-juniper, ponderosa pine, aspen, lodge pole pine, and spruce-fir communities at 2135 to 2870 m in Daggett and Uintah coun- ties; British Columbia to Saskatchewan, south to Oregon, Colorado, and South Dakota; 9 (ii). A specimen by Neese (5711 BRY) is only sparingly villous on the basal portion of the bracts. The species is known from cultivation in Utah and Emery counties; 3 (0). Gaillardia flava Rydb. Perennial herbs from a subrhizomatous woody caudex; stems 20-50 cm tall, foliose to the middle or above; leaves 2-5 cm long, 4-25 mm wide, pin- nately incised, minutely puberulent and glandular-punctate; heads solitary, on pe- duncles to 25 cm long; disk 17-32 mm wide, yellow; involucral bracts sparingly to moder- ately villous, green, caudate-attenuate; rays 8-12, yellow, the lobes 3-5 mm long; setae of receptacle well developed, coarse and spi- nescent; disk corollas sparingly villous, the hairs with colorless cross-walls, the lobes 284 Great Basin Naturalist Vol. 43, No. 2 acute; pappus scales oblong to oblanceolate, abruptly contracted to a barbellate appen- dage; achenes ca 1-1.5 mm long, yellowish pilose. Stream terraces and valley bottoms, commonly in cottonwood, willow, and tama- rix communities at 1280 to 1650 m in Emery (type from Lower Crossing) and Grand coun- ties; endemic; 6 (v). The plants are extremely resinous glandular, with a very bitter-fla- vored exudate. Gaillardia parryi Greene [G. acaulis Gray]. Perennial herbs from a woody caudex; stems 10-35 cm tall; foliose basally, less com- monly with some leaves cauline; leaves 2.5-9 cm long, 8-25 mm wide, petiolate, the blades ovate to elliptic, sparingly puberulent, mi- nutely glandular-punctate, entire or irregu- larly lobed, obtuse; heads solitary on scapose peduncles; disks 17-32 mm wide, yellow; in- volucral bracts sparingly villous, green, atten- uate; rays ca 8-12, yellow, the lobes 3-5 mm long; setae of receptacle copious, surpassing achenes; disk corollas sparingly villous, hairs with translucent cross-walls, the lobes acut- ish; pappus scales lanceolate, rather abruptly contracted to a smooth bristle; achenes ca 1.5 mm long, yellowish pilose. Pinyon-juniper and ponderosa pine communities, often in distiu-bed sites, at 1525 to 1830 m in Garfield, Kane, and Washington counties; northern Arizona; 7 (i). Gaillardia pinnatifida Torr. [G. meamsii Rydb.; G. crassifolia Nels. & Macbr., type from LaVerkin; G. gracilis A. Nels., type from Diamond Valley; and G. straminea A. Nels., type from LaVerkin]. Perennial (less commonly biennial or annual) herbs, the cau- dex seldom well developed; stems 8-55 cm tall, foliose to the middle, less commonly all leaves basal; leaves 1-7.5 cm long, 2-15 mm wide, petiolate; blades ellliptic to oblanceo- late or linear-oblong, puberulent and minute- ly glandular-punctate, pinnatifid to entire, acute to obtuse; heads solitary, on long pe- duncles; disks 15-35 mm wide, purple; in- volucral bracts moderately to sparingly vil- lous, green or suffused purplish, caudate- attenuate; rays 7-12, yellow, the lobes 2-5 mm long; setae of receptacle spinescent; disk corollas sparingly villous, hairs with trans- lucent or reddish cross-walls, the lobes acute; pappus scales oblanceolate, abruptly con- tracted to a scabrous awn; achenes ca 2 mm long, white-pilose. Blackbrush, shadscale, ephedra-vanclevea, and pinyon-juniper com- munities at 915 to 1830 m in Carbon, Emery, Garfield, Grand, Kane, San Juan, Washing- ton, and Wayne counties; Colorado and Ari- zona to Texas and Mexico; 75 (ix). Gaillardia spathulata Gray Perennial herbs from a taproot and caudex; stems 6-35 cm tall, commonly foliose to middle or above; leaves 1-7.5 cm long, 0.4-2.3 cm wide, petiolate to sessile; blades oblanceolate to elliptic or ovate to oval, sparingly villous and glandular-punctate, entire or variously toothed or lobed, obtuse; heads solitary or few, on long peduncles; disks 18-33 mm wide, yellow; involucral bracts moderately to densely villous-pilose, green, lance-attenuate; rays 7-10, yellow, the lobes 2-4 mm long; setae of receptacle short, spinescent; disk co- rollas shortly villous on the obtuse lobes, the hairs with colorless cross-walls; pappus scales oblong-lanceolate, abruptly contracted to a scabrous awn; achenes ca 3.5 mm long, yel- lowish pilose. Salt desert shrub and shrub- grass communities at 1220 to 2320 m in Car- bon, Emery, Garfield, Grand, and Wayne counties; endemic; 58 (xi). Geraea T. & G. Annual herbs; leaves alternate; heads radi- ate, showy, solitary or few in a corymbose panicle; involucres hemispheric, 2- or 3-se- riate; bracts white-ciliate; receptacle convex, the bracts clasping the achenes; rays neuter, yellow; pappus of two awns, connected by a low whitish crown; disk achenes flat, cu- neate, villous-ciliate, black. Geraea canescens T. & G. Desert Sun- flower. Annual herbs; stems 2-6 dm tall, simple or branched, white-hirsute, glandular; leaves 1-7 cm long, 0.8-4 cm wide, lanceo- late to oblanceolate or ovate, acute to obtuse, entire or few toothed, reduced upward; heads showy, borne on slender, often bracteate pe- duncles; involucres 10-25 mm wide, 7-12 mm high; bracts green, strongly ciliate, lance-acuminate; rays 10-21, yellow, 7-20 mm long; achenes 6-7 mm long. Warm desert shrub communities at 700 to 900 m in Washington County; Nevada, Arizona, and California; 3 (0). April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 285 Glyptopleura D.C. Eaton Low annual herbs; leaves rosettiform, with a few-toothed, white, crustaceous margin; heads many, short peduncled, the flowers all raylike, white or pale yellowish (or drying pinkish); involucres of 7-12 scarious- margined bracts subtended by a basal group of pinnatifid or toothed bractlets; pappus of capillary white bristles in several series, the outer falling separately; achenes oblong, 5- angled, each face with 2 rows of tubercles, abruptly beaked. 1. Ray flowers showy, long exserted, 1.5-2.5 cm long; plants of Washington County G. setulosa — Ray flowers inconspicuous, only shortly exserted, mainly less than 10 mm long; plants broadly distributed G. marginata Glyptopleura marginata D.C. Eaton De- pressed annual herbs; stems 0.5-4 cm long; leaves crowded on the short stems, mainly 0.5-4 cm long, pinnatifid, the margins white- crustose, extended into irregular white pro- cesses; involucres 10-13 mm high, urceolate; bracts green, the margins hyaline; bractlets with white, irregular, branched processes, crustose at the apex; rays mainly 4-7 mm long, withered and pinkish on drying; ach- enes 4-5 mm long, tan, sculptured. Desert shrub communities at 1240 to 1590 m in Bea- ver, Box Elder, Iron, Piute, San Juan, Sevier, and Uintah counties; Oregon and Nevada; 9 (i)- Glyptopleura setulosa Gray Low annual herbs; stems 1.5-6 cm long; leaves crowded on the short stems, mainly 0.3-5 cm long, pirmately lobed, the margins white-crustose into teeth; involucres 10-13 mm high, urceo- late; bracts green or purplish tipped, the bracts with expanded apices bearing simple or coalescent processes; rays mainly 1.5-2.5 cm long, pale yellowish, showy; achenes 4-5 mm long, tan, sculptured. Larrea, blackbrush, and Joshua tree communities at 700 to 915 m in Washington County; Arizona, Nevada, and California; 6 (0). Gnaphalium L. Annual or perennial tomentose herbs; leaves alternate, entire; heads discoid, the flowers white, yellowish, or suffused with pink, borne in spikes, corymbs, or panicles; involucres campanulate to ovoid; bracts im- bricate, scarious apically (at least); receptacle naked; outer flowers numerous, slender and pistillate, the few inner ones broader and perfect; style branches of inner flowers flat- tened, truncate, the stigmatic portion not sharply differentiated; anthers caudate; pap- pus of capillary bristles; achenes small, nerveless. 1. Heads large, mostly 4-7 mm high; clusters of heads not or rarely surpassed by leafy bracts; plants often over 20 cm tall 2 — Heads small, the involucres 2-4 mm long; clusters of heads commonly sur- passed or equaled by leafy bracts; plants mainly 4-20 cm tall 3 2(1). Leaves strongly decurrent; bracts of involucre yellowish or fading yellowish G. chilense — Leaves not strongly decurrent; bracts of involucre pearly white G. wrightii 3(1). Leaves spatulate to oblong, mainly 3-8 mm wide; plants loosely tomentose G. palustre — Leaves linear to narrowly oblanceolate, mainly 1-3 mm wide; plants rather closely tomentose 4 4(3). Leafy bracts commonly less than 1.5 cm long, more loosely tomentose than the following G. exilifolium — Leafy bracts commonly more than 1.5 cm long, the tomentum appressed G. uliginosum 286 Great Basin Naturalist Vol. 43, No. 2 Gnaphalium chilense Spreng. Annual or biennial herbs, the tomentose stems 15-40 cm tall or more; leaves 1.5-7 cm long, 2-8 mm wide, oblong to linear or the lowermost oblanceolate, decurrent, tomentose, reduced upward; heads numerous, in capitate clusters at stem apices; involucres 4-7 mm high, the bracts yellowish, tomentose only at the base. Disturbed, often moist sites at 1370 to 1770 m in Daggett, Duchesne, Kane, Salt Lake, and Utah counties; British Columbia to Mon- tana, south to California, Arizona, and Texas; 5(0). Gnaphalium exilifoUum A. Nels. [G. grayi Nels. & Macbr.]. Annual herbs; stems 8-25 cm tall, simple or branching from the base, tomentum appressed or somewhat loose; leaves 0.4-4 cm long, 1-3 mm wide, linear to linear-oblanceolate; heads clustered, in capi- tate cymes or spicate, subtended by leafy bracts that surpass them; involucres ca 3 mm high; bracts with hyaline brownish tips, to- mentose at the base. Sedge-grass commimity, known in Utah from Wasatch (Lewis sn 1975 BRY) and Washington (Albee 2936b BRY); Colorado, New Mexico, and Arizona; 2 (0). This plant simulates G. uliginosum, with which it has been synonymized by some workers. More material is necessary to pro- vide a definitive solution as to its proper tax- onomic position. Gnaphalium palustre Nutt. Annual herbs; stems 3-20 (30) cm tall, simple or more com- monly much branched, loosely tomentose; leaves 1-3.5 cm long, 2-6 (10) mm wide, ob- long to oblanceolate; heads clustered in capi- tate terminal or axillary cymes, subtended by leafy bracts that equal or surpass them; in- volucres 3-4 mm high; bracts brown, usually with whitish tips, tomentose below. Tamarix- willow, mountain brush, ponderosa pine, Douglas-fir, and sedge-grass communities, of- ten on sand bars, lake shores, and pond mar- gins, at 1370 to 2600 m in Cache, Garfield, Iron, Juab, Millard, Piute, Salt Lake, Sanpete, Sevier, Tooele, Uintah, Utah, Washington, and Wayne counties; British Columbia and Alberta, south to California and New Mexico; 31 (iii). Gnaphalium uliginosum L. Annual herbs; stems 3-15 (25) cm tall, simple or more com- monly much branched, closely tomentose; leaves 1-5 cm long, 1-3 mm wide, linear to linear-oblanceolate; heads clustered in capi- tate terminal or axillary cymes, subtended by leafy bracts that much surpass them; in- volucres 3-4 mm high; bracts brown with pale tips, tomentose below. Lake and pond margins and other disturbed sites at 2410 to 2830 m in Garfield and Sevier counties; 4 (0). Gnaphalium wrightii Gray Perennial herbs; stems 3-8 dm tall, branched in the in- florescence; leaves 1.5-7 cm long, lance-lin- ear, the lower ones spatulate; panicle open, with capitate clusters of heads not subtended or surpassed by bracteate leaves; involucres 5-6 mm high; bracts pearly white, tomentose below. Ponderosa pine and live oak commu- nities at 1585 to 1830 m in Washington County; California to Texas, south to Mexico; 3(0). Grindelia Willd. Annual, biennial, or perennial herbs, some- times woody at the base; leaves alternate, simple, more or less resinous-punctate, usual- ly sessile, often clasping; heads radiate or dis- coid, the rays 10-45, pistillate, fertile, yel- low; involucres imbricate, more or less resinous; bracts thickish, with pale appressed base and often squarrose or revolute her- baceous tips; receptacle naked, flattish; disk flowers fertile, yellow; style branches with slender hispidulous appendages; pappus of 2-8 stiff, often curved, deciduous awns; ach- enes compressed to angular, glabrous. 1. Heads discoid 2 — Heads radiate 3 2(1). Plants perennial; involucral bracts much thickened apically G. fastigiata — Plants annual or biennial; involucral bracts only somewhat thickened G. aphanactis April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 287 3(1). Involucral bracts, at least middle and upper ones, with appressed or erect tips, these not revolute or thickened G. laciniata — Involucral bracts spreading or recurved apically, often thickened apically 4 4(3). Rays mostly 12-25, rarely more; leaves entire or sharply toothed, not callous- serrulate; achenes usually with one or more knobs on the apical margin; plants perennial G. nana — Rays mostly 25-40, rarely fewer; leaves regularly callous-serrulate to sharply toothed or entire; achenes mainly lacking apical knobs; plants biennial or perennial G. squarrosa Grindelia aphanactis Rydb. Biennial herbs; stems 1.5-9 dm tall, uniformly leafy, glabrous; leaves mainly 2.5-7 cm long, 2-12 mm wide, oblong or oblanceolate, entire, crenulate-serrate or denticulate to pinnatifid, glabrous, the margin scabridulous; heads dis- coid, campanulate; involucres 7-20 mm high, 10-28 mm wide, resinous, mostly in 5 or 6 series, the upper portion loosely to moder- ately reflexed, glabrous; pappus awns 2 or 3; achenes 2.3-3 mm long, brown, mainly trun- cate apically. Weedy species of disturbed sites in Kane and San Juan counties; Colo- rado, Arizona, and Texas; 1 (0). Grindelia fastigiata Greene Perennial herbs; stems 5-10 dm tall or taller, glabrous; leaves mainly 1.5-13 cm long, 10-18 mm wide, oblanceolate to lance-oblong, entire or denticulate to dentate or serrate, glabrous; heads discoid; involucres campanulate, 10-14 mm high, 9-17 mm broad; bracts in ca 6 series, only the upper third or fourth spread- ing, with revolute, thickened tips; pappus awns 2 or 3; achenes oblong, 3.5-5 mm long. Sandy terraces and washes at 1125 to 1375 m in Emery, Grand, and San Juan counties; Col- orado; 9 (iv); a Plateau endemic. Grindelia laciniata Rydb. Perennial herbs; stems 2.5-4.5 dm tall, glabrous; leaves mainly 2-6 cm long, 3-1.5 mm broad, pinnatifid or the upper subentire or entire, narrowly ob- lanceolate to oblanceolate, glabrous; heads radiate; involucres 7-10 mm high and wide; bracts with upper one-third to one-half spreading, glabrous; pappus awns 3-5; ach- enes 2.5-3.5 mm long. Sandy washes in San Juan County (type from San Juan County); Arizona; 1 (0). Grindelia nana Nutt. Low Gumweed. [G. brownii Heller; G. nana f. brownii (Heller) Steyermark]. Perennial herbs; stems 0.8-6.5 (8) dm tall, glabrous; leaves mainly 1.5-10 cm long, 5-30 mm wide, oblanceolate, scarcely clasping; heads radiate; involucres campanulate; bracts in 5-7 series, reflexed or revolute in the upper third to fifth; rays 11-28, yellow, 5-11 mm long; pappus awns 2; achenes 3.5-4 mm long. Ruderal weed at ca 1585 to 1650 m in Cache County; Wash- ington to Montana, south to California and Idaho; 4 (0). Grindelia squarrosa (Pursh) Dunal Curly Gumweed. [Donia squarrosa Pursh; G. squar- rosa f. depressa Steyermark, type from Salt Lake County]. Perennial or biennial herbs; stems 1-8 (10) dm tall, glabrous; leaves most- ly 2-5 cm long, oblong, regularly callous toothed, sometimes sharply toothed or entire, the upper clasping; heads radiate, strongly resinous; bracts with the green tips strongly rolled back; rays 25-40, yellow, 7-15 mm long; pappus awns 2 or 3 (to 6); achenes 2.3-3 mm long. Two more or less distinctive varieties are present in Utah. 1. Main upper cauline leaves 2-4 times longer than broad, oblong-ovate to oblong G. squarrosa var. squarrosa Main upper cauline leaves 5-8 times longer than broad, narrowly oblong to oblanceolate G. squarrosa var. serrulata Var. serrulata (Rydb.) Steyermark [G. ser- rulata Rydb.]. Salt desert shrub, sagebrush, saline meadow, and mountain brush commu- nities at 1310 to 1420 m in all Utah counties; Wyoming south to New Mexico and Arizona, and introduced widely elsewhere; 72 (vii). Var. squarrosa [G. serrulata f. depressa Steyermark, type from west of Salt Lake 288 Great Basin Naturalist Vol. 43, No. 2 City]. Waste places at 1300 to 2135 m in naked or bristly, convex; disk flowers few to Duchesne, Juab, Salt Lake, Utah, and many, yellow, perfect or sterile; pappus of Wasatch counties; widespread mainly to the 10-12 unequal scales; achenes obovoid or ob- east of our area; 6 (0). long, pubescent. Lane, M. 1982. Generic limits of Xantho- C T FRRF7IA T ap cephaluTTi, Gutierrezia, Amphiachris, Gymnosperma, Greenella, and Thurovia Perennial shrubs or subshrubs, glutinous, (Compositae: Asteraceae). Systematic glabrous or hirtellous; leaves alternate, linear, Botany 7: 405-417. often punctate; heads radiate, small, numer- Solbrig, O. T. 1960. Cytotaxonomic and ous; rays pistillate or neutral, yellow, or lack- evolutionary studies in the North Amer- ing; involucres cylindric to turbinate, the ican species of Gutierrezia (Compos- bracts imbricate, chartaceous; receptacles itae). Contr. Gray Herb. 188: 1-63. 1. Heads cylindric, the ray and disk flowers 1 or 2 each G. microcephala — Heads turbinate, with more than 4 flowers 2 2(1). Ray and disk flowers 3-8 each; involucres 2-3 mm thick; heads often clus- tered at ends of branches; plants widespread G. sarothrae — Ray flowers 4-10, disk flowers 5-23; involucres 2-7 (9) mm thick; heads soli- tary or in pairs at branch ends; plants of restricted distribution 3 3(2). Disk flowers 5-12, 3.5-4.5 mm long; ray flowers 2-5 mm long; plants of Uintah County G. pomariensis — Disk flowers 15-23, ca 3 mm long; ray flowers 5-7 (10) mm long; plants of eastern Millard County G. petradoria Gutierrezia microcephala (DC.) Gray cephalum petradoria Welsh & Goodrich Brit- Thread Snakeweed. [Brachyris microcephala tonia 33: 301. 1981]. Goldenrod Snakeweed. DC; G. sarothrae var. microcephala (DC.) Perennial, suffrutescent; stems herbaceous ex- Benson; Xanthocephalum microcephalum cept at the base, hirtellous, simple below the (DC.) Shinners]. Rounded shrub, 30-100 cm inflorescence, loosely caespitose, from a stout tall; stems slender, grayish to straw colored taproot and branching, mostly underground, or green above, from a woody crown; leaves woody caudex; leaves arranged singly along dimorphic, the cauline 2-5 cm long, 2-4 mm the stems, linear, 0.5-4.5 cm long, 1-3 (4) wide, linear or linear-lanceolate, and with mm wide, reduced upward, secondary fas- shorter, narrower fasciculate axillary ones, of- cicled leaves in some lower axils; heads soli- ten one or both lacking at anthesis; heads tary or in pairs on bracteate peduncles, or clustered at branch ends, sessile; involucre some almost sessile; involucres 5-9 mm high, 3-4 mm long, 1-1.5 mm wide, cylindric; 3-7 mm wide (to 9 when pressed), cam- bracts fewer than 10, lanceolate, the tip panulate, the bracts ca 20, in 3 (4) series, greenish, slightly thickened; ray flowers 1 or greenish, the tips thickened; ray flowers 2, 3-4 mm long; disk flowers 1-3, 2-3 mm 4-10, 5-10 mm long, 1-4 mm wide, when long; pappus of ca 8 scales; achenes of disk fresh; disk flowers 15-23, ca 3 mm long; pap- flowers abortive, those of ray flowers fertile, pus scales ca 10-12; achenes 3-4 mm long, 2-3 mm long, hairy. Blackbrush, vanclevea- pubescent, abortive in disk flowers. Sage- ephedra, saltbush, purple sage, rabbitbrush, brush, oakbrush, mountain mahogany, and and pinyon-juniper communities at 850 to white fir communities at 1920 to 2590 m in 1830 m in Emery, Garfield, Grand, Juab, eastern Millard County (Canyon and Pavant Kane, Millard, San Juan, Utah, Washington, ranges); type from the Canyon Mountains; and Wayne counties; Nevada and California endemic; 9 (0). to Colorado, south to Texas and Mexico; 27 Gutierrezia pomariensis (Welsh) Welsh (viii). comb. nov. [based on: Gutierrezia sarothrae Gutierrezia petradoria (Welsh & Good- var. pomariensis Welsh Great Basin Nat. 30: rich) Welsh comb. nov. [based on: Xantho- 19. 1970; Xanthocephalum sarothrae var. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 289 pomariense (Welsh) Welsh]. Orchard Snake- weed. Rounded subshrubs; stems 1.2-4.5 dm tall, several to many from a persistent woody base; leaves 1.5-5.2 cm long, 0.5-2.5 mm wide, linear, entire, glabrous or scabrous, glandular- punctate; heads in corymbose in- florescences, solitary or 2 or 3 clustered at stem ends; involucres 5-7.5 mm high, 2-5 mm broad, turbinate to cylindric; bracts broadly obtuse, with a greenish subapical spot, resin coated; ray flowers 5-9, the co- rollas 2-5 mm long; disk flowers 5-12, the corollas 3.5-4.5 mm long; pappus scales ca 5-8; achenes 1-2 mm long, hairy. Mixed desert shrub community at 1460 to 2135 m in Duchesne and Uintah (type from Dinosaur National Monument) counties; endemic; 17 (iv). Gutierrezia sarothrae (Pursh) Britt. & Rus- hy Broom Snakeweed. [Solidago sarothrae Pursh; Xanthocephalum sarothrae (Pursh) Shinners]. Rounded shrubs; stems 9-90 cm tall, profusely branched from the base, other- wise in the inflorescence, from a woody cau- dex and stout taproot; leaves dimorphic, the main cauline ones 2-7 cm long, 1-3 mm wide, linear to linear-lanceolate, the fascicled secondary ones in lower axils, entire, glabrous to tomentulose; heads in corymbose in- florescences, usually in clusters of 3-10 at branchlet ends, seldom solitary; involucres 3-4.5 mm high, 2-3.5 mm wide, turbinate; bracts narrow, acute, with green thickened tip; ray flowers 3-7, yellow, 2-5 mm long; disk flowers mostly 3-8, 2-3 mm long; pap- pus of 8-10 scales; achenes 1-2 mm long, hairy. Warm desert shrub, sand sagebrush, live oak, sagebrush, rabbitbrush, mountain brush, and pinyon-juniper communities, often in disturbed sites, at 760 to 2440 m in prob- ably all Utah counties; British Columbia east to Saskatchewan and south to Mexico. Our variable material adjusts to disturbances and increases on grazed native rangelands; it is not considered to be palatable; 208 (xlv). Haplopappus Cassini Annual or perennial herbs, subshrubs, or shrubs, usually resinous or glandular; leaves alternate, entire or toothed to lobed; heads discoid or radiate, usually small to large, vari- ously clustered or solitary; involucres cylin- dric to turbinate or campanulate, the bracts imbricate, not aligned; receptacle flat to con- vex, naked; rays yellow when present; disk flowers perfect, yellow; pappus of barbellate capillary bristles; achenes angled or striate to smooth. Hall, H. M. 1928. The genus Haplopappus. Carnegie Institution of Washington. 391 pp. 1. Plants low, rounded, branched shrubs, or tall slender shrubs or subshrubs 2 — Plants aimual or perennial herbs, branched or unbranched 11 2(1). Heads borne on stems 2.5-5 dm long; plants of saline sandy drainages or sand- stone outcrops in southern Utah 3 — Heads borne on stems less than 2.5 dm long; plants of various substrates and distribution 4 3(2). Plants definitely shrubby, the mature branchlets ashy gray or white, of sand- stone outcrops and canyons H. scopulorum — Plants shrubby only at the base, the branchlets straw colored to greenish; plants of saline drainage bottoms and terraces H. drummondii 4(2). Stems of the season white-tomentose; involucres 10-13 mm long, the bracts only somewhat imbricate; plants commonly of high elevations H. macronema — Stems glabrous, glandular, or hairy, not tomentose; involucres mainly less than 10 mm long, but, if longer, otherwise differing 5 5(4). Involucres campanulate, 8-12 mm long; heads showy, the rays 8-10 mm long; plants of lower elevations in Washington County H. linearifolius — Involucres turbinate to cylindric or campanulate, commonly less than 8 mm long; heads not especially showy, the rays mainly 2-5 mm long, or lacking 6 290 Great Basin Naturalist Vol. 43, No. 2 6(5). Leaves densely glandular punctate, linear; ray flowers present; plants known from Washington County H. laricifolius — Leaves not glandular-punctate, narrowly to broadly oblanceolate or oblong; ray flowers lacking; plants more broadly or otherwise distributed 7 7(6). Heads 12-22 mm high, the bracts subequal, the outer herbaceous and the inner chartaceous and with broad hyaline margins; plants of limestone outcrops in the Paunsagunt and Markagunt plateaus, or mainly of igneous outcrops in the Pine Valley Mountains 8 — Heads 5.5-9.5 mm high, the bracts imbricate in several series, variously her- baceous or chartaceous, but seldom any with broad hyaline margins; plants of various substrates and distribution 9 8(7). Involucral bracts 1-nerved; achenes evenly though sparingly hairy; plants of the Pine Valley Mountains, Washington County H. crispus — Involucral bracts 3-nerved; achenes glabrous except for a few hairs apically; plants of the Paunsagunt and Markagunt plateaus H. zionis 9(7). Leaves densely stipitate-glandular, oblanceolate, acute, the margins not espe- cially repand H. watsonii — Leaves lacking stipitate glands, narrowly oblanceolate to oblong, or, if oblan- ceolate, the margins repand-undulate 10 10(9). Leaves oblanceolate, 2-5 mm wide H. cervinus — Leaves narrowly oblanceolate to oblong, 0.5-2 mm wide H. nanus 11(1). Leaves strongly 3-nerved and veiny, thick and leathery; caudices thick, woody, branched 12 — Leaves not 3-nerved and veiny; caudices simple, or, if branched, not woody 13 12(11). Bracts obtuse to rounded or less commonly acutish, strongly imbricate; plants of the Colorado drainage system H. armerioides — Bracts acute to attenuate, subequal to strongly imbricate; plants widespread .... H. acaulis 13(11). Leaves with lobes or teeth spinulose tipped; involucral bracts spinulose tipped 14 — Leaves entire or toothed, but then not spinulose tipped; involucral bracts not spinulose tipped 15 14(13). Involucral bracts glabrous or glandular dorsally; leaves pinnatifid; plants perennial H. spinulosus — Involucral bracts strigose dorsally, also minutely ciliate; leaves lobed or merely toothed to entire; plants annual H. gracilis 15(13). Stems mainly 20-40 cm tall, loosely tomentose above; involucres 15-30 mm wide; plants evidently rare H. croceus — Stems 5-20 cm tall, or, if taller, not or seldom loosely tomentose, or the heads smaller 16 16(15). Heads racemosely or spicately arranged; stems erect or nearly so, not strongly bent at the base H. racemosus — Heads solitary or corymbosely (rarely racemosely) arranged; stems strongly bent at the base 17 17(16). Involucres 12-15 mm high, 20-30 mm wide; plants not hairy in the leaf axils ... H. dementis — Involucres 5-10 mm high, 10-20 mm wide, or, if larger, the plants with hair tufts in basal leaf axils 18 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 291 18(16). Involucral bracts herbaceous throughout; achenes glabrous; plants rare, known only from the Tushar Mountains H. apargoides — Involucral bracts herbaceous only apically; achenes hairy; plants locally com- mon in saline meadows H. lanceolatus Haplopappus acaulis (Nutt.) Gray Stem- less Goldenweed. [Chrysopsis acaulis Nutt.; C. caespitosa Nutt.]. Perennial caespitose herbs from a thick ligneous pluricipital cau- dex and stout taproot, the caudex branches clothed with brown to ashy marcescent leaf bases and leaves; herbage resinous, scabrous to glabrous; stems mainly 5-20 cm tall; basal leaves 0.3-6 cm long, 1.5-10 mm wide, rigid, narrowly to broadly oblanceolate, sharply mucronate, 1- to 3-nerved; cauline leaves few, developed or reduced upward; heads solitary (rarely 2); involucres hemispheric, 6-10 mm high, 8-20 mm wide; bracts in 3 series, more or less mucronate; rays 6-15, 8-12 mm long, 2-4 mm wide; pappus white to brownish; achenes silky-villous or glabrous. This is a variable taxon, with sever- al morphological phases. Despite the tenden- cy for some of the variations to be correlated geographically, it seems best to regard our materials as consisting of two mainly sym- patric varieties. 1. Cauline leaves well developed, often the main foliage leaves; herbage merely resinous-glandular; plants of the Great Basin H. acaulis var. glabratus — Cauline leaves usually much reduced, surpassed in size by the basal ones; her- bage scaberulous or merely resinous-glandular; plants more widely distributed H. acaulis var. acaulis Var. acaulis Sagebrush-grass, pinyon-juni- per, mountain brush, ponderosa pine, western bristlecone, and spruce-fir communities at 1430 to 2685 m in Beaver, Box Elder, Cache, Daggett, Emery, Garfield, Juab, Kane, Mil- lard, Sanpete, Sevier, Summit, Tooele, Uin- tah, and Utah counties; Oregon to Wyoming, south to California, Nevada, and Colorado; 60 (ix). There is a narrow-leaved glabrous phase of this taxon in the southern portion of Duchesne County, mainly on Green River Shale. Possibly it deserves recognition at some taxonomic rank. Var. glabratus D.C. Eaton [Stenotus fal- catus Rydb., type from Iron County; S. lati- folius A. Nels., type from Utah County]. Black sagebrush, wildrye, pinyon-juniper, mountain brush, and grass-shrub communities at 1525 to 2900 m in Beaver, Iron, Juab, Mil- lard, Tooele, and Utah counties; Saskatche- wan south and west to California and Ne- vada; 22 (iv). Haplopappus apargoides Gray Perennial shortly caulescent herbs, 3-8 (15) cm tall, from a taproot and simple or branched cau- dex, this clothed with brown marcescent leaf bases; basal leaves mainly 2-6 cm long, 2-6 mm wide, lanceolate to narrowly oblanceo- late; cauline leaves reduced upward, sessile. the margins scabrous or ciliate; herbage spar- ingly long-villous with multicellular hairs; heads solitary; involucres hemispheric, 8-12 mm high, 10-14 mm broad; bracts imbricate, lanceolate to oblong, acute, cuspidate, her- baceous almost or quite to the base, glabrous dorsally, the margins long-ciliate; ray flowers 15-40, yellow, 8-15 mm long; pappus tawny; achenes glabrous. Alpine tundra community at 3355 m in Piute County (Tushar Moun- tains); California and Nevada; 1 (i). The spec- imen examined (Welsh and Thorne 12982 BRY) is tentatively assigned to this species, which is known otherwise only from the east- ern Sierra Nevada and adjacent Nevada. Haplopappus armerioides (Nutt.) Gray [Stenotus armerioides Nutt.]. Perennial caespitose herbs from a thick ligneous plu- ricipital caudex and stout taproot, the caudex branches clothed with brown to ashy marces- cent leaf bases and leaves; herbage resinous- glandular, otherwise glabrous or with scabrous leaf margins; stems 0.5-20 cm tall; basal leaves 1.5-8 cm long, 1.5-10 mm wide, rigid, linear to oblanceolate, sharply mu- cronate; 1- to 3-nerved; cauline leaves few, reduced upward; heads solitary (rarely 2); in- volucres campanulate, 8-13 mm high, 10-18 mm wide; bracts in 3 or 4 series, imbricate. 292 Great Basin Naturalist Vol. 43, No. 2 oblong to oval or obovate, obtuse, sometimes lobed below the apex, greenish near the apex, glabrous; rays 8-12, 10-12 mm long, yellow, 3-5 mm wide; pappus white; achenes silky- villous. This distinctive species is represented in Utah by two phases, which are more or less morphologically distinctive and geo- graphically correlated. Stems mainly 3-8 cm tall; leaves linear to linear-oblanceolate, mainly 1-3 mm wide; plants of the Green River Formation, Uintah County H. armerioides var. gramineus Stems usually over 8 cm tall; leaves oblanceolate, mainly 3-10 mm wide; plants widespread H. armerioides var. armerioides Var. armerioides Blackbrush, black sage- brush, pigmy sagebrush, salt desert shrub, pinyon-juniper, mountain brush, and pon- derosa pine communities at 1340 to 2120 m in Carbon, Daggett, Duchesne, Emery, Gar- field, Grand, Kane, San Juan, Sevier, and Uintah counties; Montana to Arizona, east to New Mexico and Nebraska; 85 (xii). Var. gramineus Welsh & F. J. Smith Desert shrub and pinyon-juniper commu- nities at ca 1585 to 1895 m in Duchesne and Uintah counties; endemic; 17 (0). Haplopappus cervinus Wats. Shrubs, 1-4 dm tall, much branched; branchlets grayish to straw colored; leaves 6-18 mm long, 2.2-6 mm wide, oblanceolate, straight or curved, entire or repand-undulate, attenuate basally, cuspidate apically, glabrous or resinous; heads few, cymose; peduncles 3-10 mm long; involucres 6.5-7.5 mm high, 5-8 mm wide; bracts imbricate in several series, the outer greenish ones narrowly acuminate with straight or spreading tips, the irmer char- taceous ones narrowly oblong, acute or cuspi- date, all glabrous but resinous; ray flowers 5-7, yellow, 2.5-4 mm long, ca 1 mm wide; disk flowers 5-11, glabrous or the tube spar- ingly puberulent; pappus tawny; achenes stri- gose. Black sagebrush, shadscale, pinyon-juni- per, and mountain brush communities at 1670 to 2440 m in Millard and Sevier coun- ties; Arizona; 5 (0). The type came from a place called Antelope Canyon (possibly in present-day western Millard County). More collections are needed. Haplopappus dementis (Rydb.) Blake [Pyrrocoma dementis Rydb.; P. subcaesia Greene, type from Panguitch Lake; P. la- pathifolia Greene, type from "Utah"]. Pe- rennial herbs from a simple caudex and stout taproot, the subrhizomatous caudex clothed with brown, often shredded marcescent leaf bases; stems 10-30 (40) cm tall, decumbent- ascending from an abruptly curved base, vil- lous; basal leaves mostly 2-15 cm long, 4-17 mm wide, oblanceolate, glabrous or sparingly puberulent, entire or dentate, tapering to a petiole, acute; cauline leaves reduced up- ward, sessile and somewhat clasping; heads solitary (rarely 2 or 3); involucres broadly hemispheric, 8-16 mm high, 18-30 mm wide; bracts in several series, oblong to lanceolate, green throughout or the base chartaceous, villous; ray flowers 30-60, yellow or golden, 8-14 mm long; pappus tawny; achenes hairy. Grass-sagebrush, spruce-fir, sedge-forb, and meadow communities at 2590 to 3390 m in Beaver, Daggett, Duchesne, Emery, Garfield, Iron, Kane, Piute, Sanpete, Summit, and Uin- tah counties; Wyoming to Colorado; 39 (vi). Specimens from Utah that have been deter- mined as H. integrifolius Gray apparently fall here, including the type of Pyrrocoma la- pathifolia, which was discussed by Hall (1920). Involucral bracts vary from her- baceous throughout to chartaceous at the base. More work is indicated. Haplopappus crispus L.C. Anderson Shrubs, much branched from the base, 3-5 dm tall (or more); branchlets covered with short-stalked glands; leaves 1.5-3 cm long, 3-8 mm wide, entire, green, spatulate to ob- long-oblanceolate, acuminate, the margins undulate-crisped, glutinous with low glands, not crowded below the inflorescence; heads 1 or 2, more commonly more, per branch, loosely paniculate to congested and cymose; involucres campanulate, the heads 12.5-16 mm long, 5-9 mm wide; bracts in several series, finely glandular, the outermost green, leaflike; rays lacking; disk flowers 14-24, pale yellow; pappus tawny; achenes 6.5-8.5 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 293 mm long, sparsely but evenly hairy. Ponder- osa pine, fir, manzanita, and aspen commu- nities at (915?) 2471 to 3050 m in Washing- ton and Millard (?) counties; endemic; 8 (0). Haplopappus croceus Rydb. Perennial herbs, mainly 2-6 cm tall, from a simple cau- dex and stout taproot, the caudex clothed with fibrous marcescent leaf bases; basal leaves 8-20 cm long, 6-25 (40) mm wide, el- liptic to oblanceolate, petiolate, entire or un- dulate, obtuse to acutish, glabrous or pu- berulent; cauline leaves reduced upward, sessile, more or less clasping; heads solitary (rarely more); involucres hemispheric, 12-18 mm high, 20-30 mm wide; bracts in several series, ovate to oblong or oblanceolate, her- baceous apically, chartaceous to leathery ba- sally; ray flowers 25-70, burnt orange, 10-25 mm long; pappus brownish; achenes glabrous or pilose. Mountain brush community at ca 2470 m in San Juan (La Sal Moimtains) and Washington (Kolob Reservoir) counties; Wyoming south to Arizona and New Mexico; 2(0). Haplopappus drwnTnondii (T. & G.) Blake [Linosyris drummondii T. & G.]. Perennial subshrub, the stems subherbaceous, arising from a woody base, 25-75 cm tall, straw col- ored to tan, longitudinally striate, glabrous; leaves 1.5-7.5 cm long, 1-16 mm wide, en- tire or irregularly lobed, linear to spatulate, glabrous, resinous; heads few to numerous, borne in corymbose cymes, peduncled; in- volucres turbinate, 6-8 mm high, 4-7.2 mm wide; bracts in 4 or 5 series, lance-oblong, co- riaceous, with a thick green or brownish sub- apical spot, acute, resinous; ray flowers lack- ing; pappus tawny; achenes silky. Saline riparian areas in greasewood, saltgrass, rab- bitbrush, saltbush, and tamarix communities at 1050 to 1800 m in Emery, Garfield, Grand, Kane, and San Juan counties; Colo- rado, Arizona, New Mexico, and Texas; 25 vn Haplopappus gracilis (Nutt.) Gray [Die- teria gracilis Nutt.]. Annual herbs, 3-25 (30) cm tall, commonly branched from near the base; leaves 4-25 mm long, 1-3 mm wide, linear to narrowly spatulate, spinulose-den- tate to piimatifid, white-strigose, progressive- ly/reduced and entire upward; heads solitary or few to several and corymbosely arranged; involucres 6-8.5 mm high, 8-12 mm wide; bracts in 5 or 6 series, linear-lanceolate, awn tipped, herbaceous medially, strigulose, not glandular; rays 15-30, yellow, 6-9 mm long; strigulose, not glandular; rays 15-30, yellow, 6-9 mm long; pappus tawny to white; ach- enes pilose. Larrea-gutierrezia, ponderosa pine, and spruce-fir communities at 850 to 960 m in Iron, Kane, and Washington coun- ties; California to Colorado, south to Mexico; 10 (0). Haplopappus lanceolatus (Hook.) T. & G. [Donia lanceolata Hook.; H. tenuicaulis D.C. Eaton; H. lanceolatus var. tenuicaulis (D.C. Eaton) Gray; Pyrrocoma subviscosa Greene; H. lanceolatus ssp. subviscosus (Greene) Hall; Donia uniflora Hook.; H. uniflorus (Hook.) T. & G.]. Perennial herbs from a simple cau- dex and stout taproot, the caudex clothed with brown to ashy marcescent, often fibrous, leaf bases; stems decumbent-ascending, abruptly bent at the base, 5-68 cm long; bas- al leaves 3-16 cm long, 3-35 mm wide, ellip- tic-oblong or lanceolate, glabrous or tomen- tose, petiolate, entire or dentate to lobed, often densely tomentose in the axils; cauline leaves reduced upward, finally sessile and clasping; heads solitary or few to several, and subcorymbose or less commonly racemose; involucres hemispheric, 5-12 mm high, 10-18 mm wide; bracts imbricate in 3 or 4 series, with green tips, glabrous or tomentu- lose; ray flowers 10-45, yellow, 5-10 mm long; pappus tawny; achenes densely hairy. Saline meadows at 1300 to 2500 m in Beaver, Cache, Carbon, Duchesne, Emery, Garfield, Iron, Juab, Millard, Piute, Rich, Salt Lake, Sevier, Tooele, and Utah counties; Oregon to Saskatchewan, south to California, Nevada, Colorado, and Nebraska; 42 (xi). This is a highly variable taxon of saline meadows through much of our area. Heads vary from solitary to numerous, from solitary to cor- ymbosely or racemosely arranged. Vesture is lacking or tomentose, or rarely glandular. Recognition of taxonomic categories within the variation appears to be only arbitrarily possible, and it seems best to treat our speci- mens conservatively. Haplopappus laricifolius Gray Rounded shrubs 3-8 dm tall; branchlets resinous, yel- lowish, becoming gray in age; leaves 5-18 mm long, 1-1.5 mm wide, thick, linear, res- inous-punctate; heads few to several in com- pact cymes, shortly pedunculate; involucres 294 Great Basin Naturalist Vol. 43, No. 2 campanulate, 3-5 mm high, 3-6 mm wide; bracts imbricate in ca 3 series, narrowly ob- long, acute, yellowish or hyaline, glabrous or puberulent-ciliate; ray flowers 3-6, yellow, 4-5 mm long; disk flowers 9-16, glabrous or minutely pubescent; pappus tawny; achenes white hairy. Saltgrass seep margin in warm desert shrub at 1220 m in Washington Coun- ty; Arizona to Texas and Mexico; 2 (ii). Haplopappus linearifolius Gray [H. interi- or Gov.; H. linearifolius ssp. interior (Gov.) Hall]. Shrubs, mainly 4-10 (12) dm tall; branchlets yellowish, resinous, becoming gray in age; leaves 6-28 mm long, 1-2.5 mm wide, thickish, linear to narrowly oblanceolate, res- inous-punctate; heads few to many, solitary on naked peduncles mainly 2-7 cm long; in- volucres hemispheric, 8-10 mm high, 10-18 mm wide; bracts biseriate, lance-linear, acute or acuminate, herbaceous medially, some- times minutely glandular; rays 12-18, yellow, 9-15 mm long, 4-5 mm wide; disk flowers numerous; pappus white; achenes densely hairy. Joshua tree, creosote bush, blackbrush, juniper, live oak, and sagebrush communities at 700 to 1375 m in Washington Gounty; Galifomia, Nevada, Arizona, and Baja Gali- fomia; 30 (ii). Our material is assignable to var. interior (Gov.) Jones. Haplopappus macronema Gray [Mac- ronema discoideum Nutt.]. Shrubs, mainly 1-5 dm tall; branchlets white-tomentose; leaves 8-32 mm long, 2-7 mm wide, oblan- ceolate to oblong, entire or more commonly undulate-crisped, acute to obtuse, mucronate, glandular-scabrous; heads solitary or 2 to sev- eral; involucres campanulate, 9-13 mm high, 6-12 mm wide; bracts subequal, the outer few herbaceous, oblong, the inner lance- acuminate, chartaceous, glandular-scabrous; ray flowers lacking; disk flowers 10-25; pap- pus tawny; achenes villous. Douglas fir, lodgepole pine, spruce-fir, and alpine tundra communities at 2135 to 3420 m in Beaver, Box Elder, Duchesne, Garfield, Iron, Juab, Piute, Salt Lake, Sanpete, Sevier, Tooele, Salt Lake, and Utah counties; Oregon to Wyoming, south to Galifornia, Nevada, and Colorado; 38 (vi). Haplopappus nanus (Nutt.) D.G. Eaton [Ericameria nana Nutt.]. Compact shrubs, mainly 1-3 (5) dm tall; branchlets yellowish, resinous, becoming gray in age; leaves 3-18 mm long, 0.5-2 mm broad, narrowly oblan- ceolate to linear, entire, acute, resinous but not punctate; heads solitary or few to several in compact cymes, sessile or shortly peduncu- late; involucres narrowly turbinate, 5.5-8.5 mm high, 3-7 mm wide; bracts imbricate in 4 or 5 series, the outer often greenish me- dially, the inner chartaceous, with hyaline margins, glabrous; rays 1-7, yellow, 2-3 mm long; disk flowers 4-10; pappus tawny; ach- enes villous or glabrous. Desert shrub, shrub- grass, and juniper or pinyon-juniper commu- nities at 1310 to 2820 m in Beaver, Juab, Mil- lard, Piute, Sevier, Tooele, and Washington counties; Oregon, Galifomia, Nevada, and Idaho; 18 (iii). Haplopappus racemosus (Nutt.) Torr. [Homopappus racemosus Nutt.]. Perennial herbs, from a simple caudex and stout tap- root, the caudex clothed with fibrous marces- cent leaf bases; stems 20-60 (100) cm tall, erect, not abruptly bent at the base (in ours); basal leaves mainly 6-25 cm long, 5-30 mm wide, the blades elliptic to oblong or oblan- ceolate, petiolate, rigidly erect, entire or toothed, glabrous or puberulent; cauline leaves reduced, sessile, clasping; heads ra- cemose, in panicles or spikes, shortly pe- dunculate; involucres 8-12 mm high, 4-18 mm wide; bracts in 3 or 4 series, with green tips and coriaceous bases, abruptly pointed apically; rays 10-35, yellow, 5-12 mm long; pappus tawny; achenes hairy or glabrous. Sa- line meadows at 1370 to 1470 m in Millard and Utah counties; Oregon to Idaho, south to Galifornia and Nevada; 2 (i). Utah lies at the eastern margin of the range of this species complex, in which Hall (1928) recognized nine subspecies. Our material is hardly repre- sentative of the variation within the assem- blage of forms that lie to the west of this re- gion. One of our specimens (Welsh et al. 14514 BRY) belongs to the spiciform narrow- headed var. sessiliflorus (Greene) Welsh stat. nov. (based on: Pyrrocoma sessiliflora Greene Leafl. Bot. Obs. & Grit. 2: 12. 1909), and the other is a paniculiform large-headed phase apparently nearest to var. prionophyllus (Greene) Welsh stat. nov. (based on Pyrro- coma prionophylla Greene Leafl. Bot. Obs. & Grit. 2: 12. 1909). Much more material is re- quired to evaluate the nature of the speci- mens in Utah. Racemose phases of the closely April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 295 related H. lanceolatus (q.v.) have been mis- taken for H. racemosus. The erect or suberect stems and stiffly erect leaves appear to be di- agnostic for our specimens of H. racemosus. Haplopappus scopulorum (Jones) Blake [Bigelovia nienziesii var. scopulorum Jones, type from Zion Canyon; H. scopulorum var. hirtellus Blake, type from Cedar Canyon]. Shrubs, mainly 3-10 dm tall; branchlets green to straw colored or white, glabrous; leaves 0.7-7.8 cm long, 1-8 mm wide, nar- rowly lanceolate to oblong, entire, 3-nerved, glabrous, the margins scabrous, attenuate to a spinulose apex; heads few to many, borne in loose to subcompact cymes, peduncled; in- volucre narrowly campanulate, 6.5-9.5 mm high, 3-5.5 mm wide; bracts in 5 or 6 series, oblong, chartaceous and pale, or the tips greenish or often brownish, rounded-obtuse, glabrous, not resin coated; ray flowers lack- ing; disk flowers 10-20; pappus white; ach- enes white-pilose. Pinyon-juniper, mountain brush, and ponderosa pine commimities at 1370 to 1830 m in Iron, Kane, San Juan, and Washington counties; Arizona; 10 (vii). Haplopappus spinulosus (Pursh) DC. [Amellus spinulosus Pursh]. Perennial herbs from a ligneus caudex; stems mainly 12-50 (60) cm tall, branching above the base; leaves 0.5-6 cm long, 1-10 mm wide, pinnatifid to bipinnatifid or the upper ones entire, or merely toothed, spinulose; heads solitary, or few in corymbose clusters; involucres 5-8 mm high, 8-12 mm wide; bracts in 4-6 series, linear, awn-tipped, herbaceous me- dially, glandular, not strigulose; rays 15-50, yellow, 8-10 mm long; pappus brownish; achenes pilose. Desert shrub commimity at ca 1300 m in San Juan County (Atwood 7175 BRY); Alberta to Minnesota, south to Califor- nia, Arizona, New Mexico, Texas, and Mexi- co; 1 (0). Haplopappus watsonii Gray Shrubs, 1-4 dm tall; herbage stipitate-glandular; branch- lets yellowish, becoming whitish to straw col- ored or grayish in age; leaves 4-28 mm long, 3-10 mm wide, oblanceolate to obovate or spatulate, entire or undulate, abruptly cuspi- date-acuminate apically; heads several to nu- merous (rarely some solitary) in loose cymes, the peduncles 1-7 mm long; involucres sub- cylindric to narrowly campanulate, 5.5-8 mm high, 3-76 mm wide; bracts in ca 5 series, the outer ones greenish, the inner chartaceous or greenish at the tips; rays 5-10, yellow, 4-6 mm long; disk flowers 5-15; pap- pus brownish; achenes hairy. Rock outcrops (limestone, sandstone, or quartzite) in desert shrub, pinyon-juniper, mountain brush, and ponderosa pine commimities at 1310 to 3440 m in Beaver, Box Elder, Cache, Davis, Juab, Millard, Salt Lake, Summit, Tooele, and We- ber counties; Nevada and Utah; 33 (ii). Our material belongs to one of a vicarious pair of infraspecific taxa within the Great Basin known as var. rydbergii (Blake) Welsh comb, nov. (based on: H. rydbergii Blake Contr. U.S. Natl. Herb. 25: 545. 1925, nom. nov. pro Macronema obovatum Rydb. Bull. Torrey Bot. Club. 27: 68. 1900, type from City Creek Canyon). The var. rydbergii differs in having fewer disk flowers (5-15 not 15-25). Other supposedly diagnostic features (i.e., the green outer involucral bracts) fail, being present to a greater or lesser degree in both phases. The type variety has not been discov- ered in Utah, but should be expected in the western border region. Haplopappus zionis L.C. Anderson Shrubs, mainly 1-3 dm tall; herbage minutely and shortly stipitate-glandular; leaves 0.8-3.5 (4) cm long, 2-4.5 (7) mm wide, oblong to nar- rowly oblanceolate, 1 -nerved, entire, abrupt- ly mucronate; heads solitary or 2 or 3, in cy- mose clusters, peduncled; involucres cylindric-campanulate, 12-15 mm high, 6-12 mm wide; bracts subequal, herbaceous (outer) and greenish, the inner chartaceous or with a subapical green spot and broadly hyaline margins; rays lacking; disk flowers 8-21; pap- pus tawny; achenes glabrous below, strigose apically. Ponderosa pine and spruce-fir com- munities, commonly on limestone members of the Cedar Breaks (Wasatch) Formation, at 2440 to 3050 m in Garfield, Iron, and Kane counties; endemic; 5 (i). Helenium L. Annual or perennial herbs; leaves alter- nate, glandular-pimctate, decurrent or clasp- ing; heads solitary or few to numerous in co- rymbose clusters, radiate, yellow; involucral bracts in 2 or 3 series, the bracts subequal or the inner shorter and narrower, herbaceous 296 Great Basin Naturalist Vol. 43, No. 2 or essentially so, soon deflexed; receptacle naked, convex or conic; rays pistillate or neu- ter; disk flowers numerous, perfect; pappus of 5-10 scarious or hyaline scales; achenes truncately obpyramidal, 4- or 5-angled, with as many intermediate ribs. 1. Leaves sessile, clasping; stems not winged; plants of aspen communities and upward H. hoopesii — Leaves decurrent; stems winged below the leaf bases; plants of riparian com- munities at lower elevations H. autumnale Helenium autumnale L. Common Sneeze- weed. Perennial herbs; stems mainly L5-10 (12) dm tall, puberulent and glandular, cor- ymbosely branched above; leaves 1.5-15 cm long, 3-35 (40) mm wide, serrate to entire, glandular-punctate; heads 3 to many, the disk hemispheric to subglobose, yellow, 1-2 cm wide; rays 10-20, yellow, mainly 8-12 mm long, soon reflexed; pappus scales lance- ovate, with slender awn-tip as long as the body; achenes ca 1.5 mm long, hirsute and glandular. Cattail-willow, tamarix-grease- wood, and sedge-rush communities at 1220 to 1830 m in Box Elder, Daggett, Emery, Rich, Uintah, and Utah counties; British Columbia to Quebec, south to Arizona, and Florida; 18 (iii). This species is poisonous to livestock. Helenium hoopesii Gray Orange Sneeze- weed. [Heleniastrum hoopesii (Gray) Kuntze; Dugaldia hoopesii (Gray) Rydb.]. Perennial herbs, mainly 2-8 (10) dm tall, with a sub- rhizomatous caudex and fibrous roots; herb- age more or less villous-tomentose to gla- brate; basal leaves 2-30 cm long, 0.5-5 cm wide, oblanceolate, tapering to a clasping base; cauline leaves reduced upward, oblan- ceolate to elliptic or lanceolate, entire; heads 2-11, in loose corymbs; disks hemispheric, 2-3.5 cm wide; involucres 5-8 mm high, the bracts lanceolate to elliptic; rays 13-21, yel- low or yellow-orange, 15-35 mm long, finally reflexed; pappus scales hyaline, lanceolate, attenuate; achenes 3-4 mm long, hairy. Sage- brush, mountain brush, aspen, and spruce-fir communities, often in openings or riparian zones, at 1830 to 3200 m in Beaver, Box El- der, Carbon, Duchesne, Garfield, Iron, Juab, Piute, San Juan, Sevier, Summit, Utah, Wasatch, and Washington counties; Oregon to Wyoming, south to California, Arizona, and New Mexico; 69 (xiv). This is a poisonous plant, causing spewing sickness in sheep. Helianthella T. & G. Perennial herbs; leaves simple, opposite or alternate, entire; heads radiate, solitary or few to several in loose subcorymbose clusters; bracts imbricate to subequal, more or less herbaceous; receptacle plano-convex, chaffy throughout, the persistent bracts clasping the achenes; disk flowers numerous, fertile, yel- low, or purple; rays yellow; pappus of 2 slen- der awns and short scales; achenes strongly compressed at right-angles to involucral bracts. Weber, W. A. 1952. The genus Helianthella (Compositae). Amer. Midi. Naturalist 48: 1-35. 2(1). Heads 3-12 or more, mainly less than 20 mm broad; rays 7-13 mm long, in- conspicuous; disk flowers normally purple H. microcephala Heads solitary or 2 or 3, mainly over 20 mm broad; rays 15-30 mm long, showy; disk flowers yellow 2 Heads erect; involucral bracts lance-oblong, short-ciliate H. uniflora Heads nodding; involucral bracts oblong-ovate, long-ciliate with multicellular hairs H. quinquenervis Helianthella microcephala (Gray) Gray [Encelia microcephala Gray]. Perennial herbs; stems 20-65 cm tall; herbage ap- pressed hispidulous; basal leaves mainly 4-30 cm long, 0.5-3 cm wide, petiolate, the blades elliptic to lanceolate, scabrous and harshly ciliate, acute to obtuse; cauline leaves re- duced upward; heads 3-12 or more; bracts imbricate in ca 3 series, oblong to lanceolate or oblanceolate, strigose and roughly ciliate April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 297 and glandular; rays 8-10, yellow, 7-13 mm long; disk flowers commonly purple; achenes 7-8 mm long, long-pilose. Desert shrub, pin- yon-juniper, ponderosa pine, mountain brush, and Douglas fir-limber pine communities at 1220 to 2745 m in Carbon, Duchesne, Emery, Garfield, Grand, Kane, San Juan, Se- vier, and Uintah counties; Colorado and Ari- zona; 34 (x). In one plant from Navajo Moun- tain the disk flowers are apparently yellow. A single collection from west of Richfield (Welsh et al. 17487 BRY) is the only record examined for the Great Basin. Helianthella quinquenervis (Hook.) Gray [Helianthus quinquenervis Hook.]. Perennial herbs; stems 5-15 dm tall, glabrous or villous above; basal leaves 0.3-40 cm long, 0.8-4 cm wide, petiolate, the blades elliptic to oblong or oblanceolate, entire, obtuse to acute; cau- line leaves often enlarged to near stem middle then reduced, becoming subsessile or sessile, the largest (at least) prominently 5- nerved; heads nodding, solitary or 2 or 3; disk 2.5-4 cm wide; bracts ovate-lanceolate, acuminate, long-ciliate; rays 12-21, yellow, 15-35 mm long; achenes 8-10 mm long, pi- lose. Sagebrush, aspen, ponderosa pine, and spruce-fir communities at 2115 to 3175 m in Carbon, Daggett, Duchesne, Emery, Gar- field, Grand, San Juan, Sevier, Summit, Uin- tah, and Wasatch counties; Oregon to South Dakota, south to Nevada, Arizona, and New Mexico; 13 (0). Helianthella uniflora (Nutt.) T. & G. [He- lianthus uniflorus Nutt.]. Perennial herbs from a branching caudex; stems mainly 3-10 dm tall, glabrous below or more or less spreading-hairy throughout; basal leaves 3-15 cm long, 0.6-5.5 cm wide, petiolate, the blades oblanceolate to elliptic or lanceolate, entire, obtuse to acute; cauline leaves often enlarged to near stem middle, then reduced, becoming sessile or subsessile, the largest prominently 3-nerved; heads erect, solitary or 2 or 3; disk 1.5-3 cm wide; bracts lance- linear, acuminate or obtuse, scabrous- puberulent, shortly ciliate; rays 13-17, yel- low, 2-4.5 cm long; achenes 6-7 mm long, pilose. Sagebrush, pinyon-juniper, mountain brush, ponderosa pine, aspen, and spruce-fir communities at 1525 to 3175 m in Beaver, Box Elder, Cache, Carbon, Davis, Duchesne, Garfield, Grand, Iron, Millard, Morgan, Piute, Rich, Salt Lake, Sanpete, Sevier, Sum- mit, Tooele, Uintah, Utah, Wasatch, and We- ber counties; Alberta to Montana, south to Nevada and Colorado; 64 (v). Helianthus L. Annual or perennial herbs; leaves simple, opposite below, usually alternate above; heads radiate, showy, solitary or few in cor- ymbs; involucral bracts imbricate or sub- equal, herbaceous; receptacle flat to convex, chaffy throughout, its bracts clasping the achenes; ray flowers conspicuous, yellow, neuter; disk flowers yellow or reddish, fertile; pappus usually of 2 main awns, scalelike at base, sometimes with additional scales pres- ent; achenes narrowly obovate in outline, 4- angled or obcompressed. Blauer, a. C. 1965. Helianthus (Compositae) in Utah. Proc. Utah Academy 42: 240-251. Heiser, C. B. Jr. 1947. Hybridization be- tween sunflower species Helianthus an- nuus and H. petiolaris. Evolution 1: 249-262. 1. Plants perennial; disk flowers yellow; leaves mainly opposite, lanceolate to linear-lanceolate H. nuttallii — Plants annual; disk flowers reddish brown to purplish; leaves mainly alternate; leaves lanceolate to ovate 2 2(1). Involucral bracts linear to narrowly lanceolate; pappus of numerous unequal scales 3 — Involucral bracts lanceolate to ovate; pappus commonly of 2 distinct awns 4 3(2). Involucral bracts surpassing the disk; pappus scales linear; stems moderately hirsute to glabrous; plants widespread H. anomalus 298 Great Basin Naturalist Vol. 43, No. 2 — Involucral bracts subequal to the disk; pappus scales ovate; stems markedly hirsute; plants of Washington County H. deserticola 4(3). Leaves canescent on both sides; involucral bracts narrowly lanceolate; plants of San Juan County H. niveus — Leaves hispid to strigose; involucral bracts lanceolate to ovate; plants wide- spread 5 5(4). Involucral bracts ovate, rather abruptly narrowed to an acuminate tip, the cen- tral ones inconspicuously hairy; leaves often cordate and with serrate margins .. H. annuus — Involucral bracts lanceolate, tapering to the tip, the central ones often white bearded apically; leaves seldom cordate, usually entire H. petiolaris Helianthus annuus L. Common Sun- flower. Annual herbs; stems commonly hispid and rough, 3-40 dm tall, simple or branched; leaves alternate above (3) 5-40 cm long, 2-40 cm wide, lance-ovate to broadly ovate, acute to obtuse, serrate, truncate or cordate (alter- nate) basally, hispid to hispidulous on both sides, petiolate; heads solitary or few; disks mainly 2-5 cm wide; involucral bracts lance- ovate to ovate, attenuate to caudate, hispid to hispidulous, ciliate; disk corolla lobes purplish red (rarely yellow); pappus of 2 awnlike ovate-lanceolate scales; achenes glabrous to strigose. Saltgrass-muhly grass, desert shrub, pinyon-jimiper, and mountain brush communities, commonly where dis- turbed, at 1200 to 2440 m, probably in all Utah counties; widespread in the United States, Canada, Mexico, and elsewhere; 70 (xv). Our common weedy sunflower is assign- able to ssp. lenticularis (Dougl.) Cockerell; the cultivated large-headed phase to var. macrocarpus (DC.) Cockerell. Helianthus anomalus Blake Sand Sun- flower. Annual herbs; stems sparingly hispid to glabrate, 5-70 cm tall; leaves mainly alter- nate, petiolate, the blades L2-10 cm long, 0.4-4 cm wide, narrowly lanceolate to lance- ovate, yellowish green, acute, cuneate to ob- tuse basally, hispidulous to hispid on both sides; heads solitary or few, showy; disks mainly 12-24 mm wide; involucral bracts lin- ear, commonly 10-25 mm long and 2-3 mm wide, hispid above, definitely hispid-ciliate, at least below, often some much surpassing the disk; disk corolla lobes purple; pappus of 2 large linear scales and numerous similar subequal scales; achenes 3.5-5.5 mm long, appressed pilose. Blackbrush, ephedra, purple-sage, vanclevea, psorothamnus, and pinyon-juniper communities, commonly in dunes or other sandy sites, at 1150 to 1830 m in Emery, Garfield, Grand, Juab, Kane, Mil- lard, San Juan, Tooele, and Wayne (type from near Hanksville) counties; Arizona; 33 (xi). This is a Colorado Plateau endemic, with an extension onto dunes of the eastern Great Basin. Helianthus deserticola Heiser Desert Sun- flower. Annual herbs; stems strongly hispid (at least below), 2-12 dm tall; leaves mainly alternate, petiolate, the blades 2-6 cm long, 0.5-2 cm wide, lanceolate to lance-ovate, green, acute, cuneate to obtuse basally, his- pid to hispidulous on both sides; heads few to several (solitary), showy; disks 1.3-2.5 cm wide; involucral bracts linear, 8-14 mm long, 1.5-2.5 mm wide, hispid dorsally and ciliate, subequal to the disk; disk corolla lobes purple; pappus of 2 large lanceolate to lance- ovate scales and ca 10 smaller ones; achenes 4-5 mm long, pilose. Blackbrush, creosote bush, matchweed, and live oak communities at 850 to 1070 m in Washington County; Ari- zona and Nevada; 2 (0). This is an obscure taxon, despite the passage of two decades since its description; more specimens are re- quired. Perhaps it is too nearly allied to H. anomalus, with which it shares some mor- phological features. Helianthus niveus (Benth.) Brandegee Snowy Sunflower. [Encelia nivea Benth.]. Annual herbs; stems mainly 5-15 dm tall, ca- nescent and hispid; leaves mainly alternate, petiolate, the blades 3-12 cm long, 0.9-6 cm wide or more, lanceolate to ovate, entire or serrate, mostly acute apically, canescent on both sides; heads solitary or few to many; disk 1-2.5 cm wide; bracts narrowly lanceo- late, 1.5-2.5 mm wide, 8-12 mm long, sub- equal to the disk; disk corolla lobes purplish; April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 299 pappus of 2 lanceolate scales and several shorter scales. Sandy sites at ca 1375 m in San Juan Counties; Texas to California and Mexico; 1 (0). Our limited material is assign- able to ssp. canescens (Gray) Heiser [H. pet- iolaris var. canus Britt.]. Helianthus nuttallii T. & G. Nuttall Sun- flower. [H. bracteatus E.E. Watson, type from Logan; H. giganteus var. utahensis D.C. Eaton, type from Wasatch Mountains; H. utahensis (D.C. Eaton) A. Nels.]. Perennial rhizomatous herbs with tuberous roots; stems 3-20 dm tall or more, glabrous or sparingly scabrous or hispid; leaves mainly opposite, shortly petiolate, the blades 4-16 cm long, 0.8-3 cm wide, narrowly lanceolate, acute to attenuate, entire or denticulate, cuneate ba- sally, scabrous on both sides; heads solitary or few to many; disks mainly 12-28 mm wide; bracts lance-linear, 1.5-3 mm wide, subequal to, or surpassing, the disk, attenuate, ap- pressed pubescent and more or less ciliate; disk corolla lobes yellow; pappus of 2 narrow awnlike scales; achenes 3-4 mm long, glabrous. Seeps, springs, wet meadows, and canal banks at 1280 to 2200 m in Cache, Car- bon, Duchesne, Garfield, Juab, Rich, Salt Lake, Summit, Tooele, Uintah, Utah, Wasatch, Washington, and Weber counties; British Columbia to Saskatchewan, south to Nevada, Arizona, and New Mexico; 18 (0). Note: The perennial sunflower, H. tuherosus L., is grown for its edible roots in our area. It persists following cultivation and is difficult to eradicate. The leaves are broadly lanceo- late to ovate. Helianthus petiolaris Nutt. Prairie Sun- flower. Annual herbs; stems 0.5-12 dm tall, strigose to hispid or glabrous; leaves mainly alternate, petiolate, the blades 1-8 cm long, 4-25 (30) mm wide, lanceolate to ovate, acute to obtuse, entire or rarely serrate, cu- neate to truncate basally, hispidulous to stri- gose; heads solitary or few; disk 10-25 mm wide; involucral bracts 2-5 mm wide, 7-15 mm long, lanceolate, acuminate or attenuate, hispidulous, usually short-ciliate; disk corolla lobes purplish; pappus of 2 lanceolate awn- like scales; achenes 3-4.5 mm long, hairy. Salt desert shrub, desert shrub, pinyon-juni- per, and riparian communities, often where disturbed, at 1220 to 1920 m in Beaver, Duchesne, Emery, Garfield, Grand, Kane, Millard, San Juan, Sevier, Uintah, Wasatch, Washington, and Wayne counties; Alberta to Maine, south to California, Arizona, New Mexico, Texas, Louisiana, and South Caro- lina; 62 (xxii). Our material has been assigned to ssp. fallax Heiser. The material appears to be indigenous in the Colorado drainage sys- tem, but the rare specimens in the Great Ba- sin seem to be adventive. Heliomeris Nutt. Annual or perennial herbs; leaves opposite (at least below), simple; heads radiate, soli- tary or cymose; involucres 2- or 3-seriate; rays yellow, neuter, pubescent dorsally; re- ceptacles chaffy, the chaffy bracts clasping the achenes; disk flowers fertile; pappus none; achenes laterally compressed, 4-angled. 1. Plants perennial, widespread in montane habitats, less commonly in saline low elevation sites H. multiflora — Plants annual, restricted in low elevation saline habitats 2 2(1). Plants subscapose, with long naked peduncles; leaves ovate to lanceolate . H. soliceps — Plants caulescent, the peduncles bracteate or leafy; leaves linear 3 3(2). Leaves canescent with appressed hairs; plants of southern Utah H. longifolia — Leaves hispidulous; plants of central and western Utah H. hispida Heliomeris hispida (Gray) Blake Hairy Goldeneye. [H. multiflora var. hispida Gray; Gymnolomia hispida var. ciliata Robins. & Greenm., type from Utah; Viguiera ciliata (Robins. & Greenm.) Blake]. Annual herbs; stems simple or variously branched, 10-70 cm tall, hispidulous; leaves 0.6-9 cm long, 1-3 mm wide, linear, hispid and hispid- ciliate, acute; heads solitary or 2-5 or more; disks 7-15 mm wide, the corollas yellow; rays 300 Great Basin Naturalist Vol. 43, No. 2 ca 9-15, yellow, 6-13 mm long; involucral bracts 5.5-10 mm long, lance-attenuate, his- pid and coarsely ciliate; pappus lacking; ach- enes ca 2.5 mm long, glabrous. Saline mar- shes and meadows at ca 1300 to 1470 m in Millard, Salt Lake, and Utah counties; Ari- zona, New Mexico, and Mexico; 9 (ii). Heliomeris longifolia (Robins. & Greenm.) Cockerel! [Gymnolomia longifolia Robins. & Greenm.; Viguiera longifolia (Robins. & Greenm.) Blake]. Annual herbs; stems simple or variously branched, 14-60 cm tall, stri- gose; leaves 1-6 cm long, 1.2-7.5 mm wide, linear to oblong, strigose, rarely hispid-ciliate near the bases, acute; heads solitary, or 2 to numerous; disks 7-10 mm wide, the corollas yellow; rays ca 8-10, yellow, 6-12 mm long; involucral bracts lance-acuminate to -attenu- ate, strigose, not especially ciliate; pappus lacking; achenes 2-2.5 mm long, brown, glabrous. Salt desert shrub and pinyon juni- per communities at 1150 to 1525 m in Kane and Washington counties; Arizona to Texas and Mexico; 7 (iii). Our material is assignable to var. annua (Jones) Yates [Gymnolomia multiflora var. annua Jones, type from Utah?]. Heliomeris multiflora Nutt. Showy Gold- eneye. [Viguiera multiflora (Nutt.) Blake]. Perennial herbs, from a woody taproot and pluricipital caudex; stems 2-10 (13) dm tall, strigose to scabrous-puberulent; leaves lan- ceolate to linear, mainly opposite, entire or serrate, 1-8 (10) cm long, 2-20 (25) mm wide, short-petiolate, plane or revolute, acute to obtuse; heads commonly 2 to several; disk 6-14 mm wide; involucral bracts linear or narrowly lanceolate, strigose; rays 10-14, yel- low, 7-18 mm long; pappus lacking; achenes 1.2-1.8 mm long, brown, glabrous. Two weakly discernible varieties are included in our material. Leaves commonly over 5 mm wide, plane; plants of mesic montane sites H. multiflora var. multiflora Leaves commonly less than 5 mm wide, the margins revolute; plants of arid plains and mountains H. multiflora var. nevadensis Var. multiflora Sagebrush, juniper, cotton- wood, pinyon-juniper, aspen, and spruce-fir communities, often in riparian sites, at 1340 to 2870 m in all Utah counties; Montana south to California, Arizona, and New Mexi- co; 137 (xvi). Var. nevadensis (A. Nels.) Yates [Gymo- nolmia nevadensis A. Nels.]. Shadscale, mat- atriplex, pinyon-juniper, and mountain brush communities at 1370 to 2135 m in Grand, Juab, Uintah, and Washington counties; Ne- vada; 13 (i). Heliomeris soliceps (Bameby) Yates Tropic Goldeneye. [Viguiera soliceps Bameby]. An- nual herbs, 10-41 cm tall; stems branched be- low, terminating in subscapose, merely brac- teate peduncles that overtop the foliage; leaves opposite below, the blades 15-38 mm long, 6-20 mm wide, ovate to lanceolate, strigose, 3-nerved, petiolate, obtuse to cu- neate, becoming smaller upwards; peduncles 7-28 cm long; involucres biseriate, the bracts lance-acuminate, acute, 5-6 mm long, stri- gose; rays 10-12, yellow, 10-15 mm long; pappus lacking; achenes 2.8-3.3 mm long, blackish. Mat-saltbush community on Tropic Shale Formation at 1400 to 1470 m in Kane County; endemic. This is a striking species, forming masses of yellow blossoms in years of adequate rainfall; 5 (ii). Heterotheca Cass. Annual, biennial, or perennial herbs; leaves alternate, simple, entire; heads radiate; in- volucres campanulate to hemispheric; bracts numerous, narrow, imbricated in several series; receptacle convex, naked; rays yellow, pistillate and fertile; pappus of capillary bristles; disk flowers numerous, the pappus present and usually double, the inner of ca- pillary bristles, the outer (when present) of short scales or bristles; achenes hairy. Wagenknecht, B. L. 1960. Revision of Het- erotheca, Section Heterotheca (Com- positae). Rhodora 62:61-76, 97-109. 1. Plants low, creeping, arising from subrhizomatous caudex branches; heads nod- ding, solitary or 2 or 3, known from sandy sites in Garfield, Kane, and Wash- ington (?) counties H. pnesii April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 301 — Plants various in habit, the caudex branches, if present, not rhizomatous; heads few to numerous, seldom nodding, distribution various 2 2(1). Plants perennial, from a woody root crown, the stems numerous, forming rounded clumps, common H. villosa — Plants annual or biennial, the root crown herbaceous; stems solitary or few, not forming rounded clumps, rare 3 3(2). Upper leaves cordate-clasping basally; involucres glandular-puberulent and canescent H. psammophila — Upper leaves tapering to a sessile base; involucres glandular, pubescent, not canescent H. grandiflora Heterotheca grandiflora Nutt. Telegraph herbs; stems stout, 5-12 (20) dm tall, branch- Weed. Annual or biennial herbs; stems stout, ing above, hispid hirsute and glandular up- branched above, 5-12 (20) dm tall, hirsute, ward; leaves mainly 1-7 cm long, ovate to glandular-pubescent; leaves 2-6 cm long, lance-oblong, serrate to subentire, the lower 0.8-2.5 cm wide (or more), ovate to elliptic, petiolate, the upper cordate-clasping; heads oblong, or oblanceolate, serrate, the lower numerous; involucres 8-12 mm high, glandu- petiolate and lobed at base; heads numerous; lar and canescent; rays 20-30, yellow, mainly involucres 7-9 mm high; rays 25-35, 6-8 mm 3-7 mm long; pappus tawny; achenes 2.4-3.8 long, ca 1 mm wide, the tube hairy; disk mm long. Sandy roadside at ca 970 to 1350 m flowers numerous, slender; pappus tawny, in Grand and Washington counties; Arizona Sandy roadside at ca 915 m in Washington to Texas, south to Mexico; 1 (i). The speci- County; California and Arizona; 1 (i). Our men from Grand County (Welsh & Moore material has the pappus merely tawny, not 2745) is missing from BRY, the Washington brick red as reported elsewhere for the County locality is reported by Meyer, species. Heterotheca villosa (Pursh) Shinners Hairy Heterotheca jonesii (Blake) Welsh & At- Goldenaster. [Amellus villosus Pursh; Chry- wood Jones Goldenaster. [Chrysopsis caespi- sepsis villosa (Pursh) Nutt. ex DC.]. Perennial tosa Jones, not Nutt.; Chrysopsis jonesii herbs, from a ligneous root-crown and tap- Blake]. Perennial caespitose herbs from a root; stems several to numerous, forming creeping subrhizomatous caudex; stems 4-8 rounded clumps, mainly 1.5-5 dm tall; herb- cm tall, loosely villous; leaves 5-11 mm long, age hirsute to strigose and more or less 1.5-4 mm wide, petiolate, the blades obovate glandular; leaves 0.5-5 cm ong, 2-10 mm to spatulate, pilose; heads solitary or 2 or 3; ^i^^' oblanceolate to spatulate or elliptic, ., c % c uucin A green or silvery to gray-green, petiolate or involucres 5-7.5 mm high, 6-10 mm wide, => , ... / . & -^ & ' r ^, , ^ Till ^ • subsessiie; heads few to numerous, mainly co- the bracts narrowly lance-oblong, stngose-pi- , . , -7 m u- u -7 io , , , 1. -^ . 11.1 n rymbose; involucres 7-10 mm high, 7-12 mm lose, the hyaline margins reddish; ray flowers ^.^^_ ^^^^^^ lance-linear, green or char- 5-13, yellow, 4-6 mm long, 1.5-2.5 mm ^^^^^^^ ^^^ .^^ ^^^^.^^^ sometimes red- wide; pappus tawny; achenes 2-3 mm long, ^^^. ^^^^ ^q_25 yellow, 6-10 mm long; pap- hairy. Ponderosa pine, manzanita, and ^^^ ^^wny; achenes 2-3 mm long, hairy. Our Douglas fir communities, on sandstone or in materials represent only a small portion of sand, at 1580 to 2745 m in Garfield, Kane, the vast array of variation within the villosa and Washington (?) counties (the type pre- complex. Three infraspecific taxa are appar- sumably came from Springdale); endemic; 7 ent among our specimens, but application of (ii). names is difficult. The following treatment is Heterotheca psammophila Wagenkn. [H. therefore tentative, with a definitive treat- subaxillaris, authors]. Annual or biennial ment awaiting monographic study. 1. Leaves green or gray-green, the surface apparent through the spreading to sub- appressed hairs; plants widespread H. villosa var. hispida — Leaves silvery or grayish, the surface seldom apparent through the usually ap- pressed hairs; plants restricted 2 302 Great Basin Naturalist Vol. 43, No. 2 2(1). Stems mainly appressed hairy, or with some hairs ascending to spreading H. villosa var. villosa — Stems with appressed and spreading contorted long hairs H. villosa var. foliosa Var. foliosa (Nutt.) V.L. Harms [Chry- Var. villosa Blackbrush, pinyon-jimiper, sopsis foliosa Nutt.]. Mountain brush and and ponderosa pine communities at 1090 to bunchgrass (Agropyron et al.) communities at 2785 m in Garfield, Kane, Washington, and 1280 to 2135 m in Davis, Duchesne, Salt Wayne counties; widespread in western U.S.; Lake, San Juan (?), Utah, and Weber coun- 8 (iii). ties; widespread in western U.S.; 9 (0). Var. hispida (Hook.) V.L. Harms [Dip- Hieracium L. lopappus hispidus Hook.; Chry sopsis hispida (Hook.) Nutt.; C. villosa var. scabra Eastw. Perennial rhizomatous herbs with milky (?), type from San Juan County; C. viscida juice; leaves alternate or basal and still alter- var. cinerascens Blake, type from Beaver nate, entire or toothed, simple; heads few to Canyon]. Shadscale-rabbitbrush, ephedra-ly- numerous, in corymbose clusters; flowers all cium-dropseed, sagebrush-grass, pinyon-juni- raylike, yellow to orange or white; involucres per, mountain brush, ponderosa pine-manza- cylindric to hemispheric; bracts more or less nita, and aspen commimities at 1150 to 2745 imbricate; receptacle naked; pappus usually m in all Utah counties; widely distributed in of brownish capillary bristles; achenes terete the western U.S.; 105 (xii). or prismatic, more or less strongly ribbed. 1. Flowers white; stems long-setose basally if at all, the petioles and leaves long- setose on the lower midrib H. albiflorum — Flowers yellow; stems long-setose throughout or only above; leaves variously setose or glabrous 2 2(1). Leaves glabrous or short-hairy, 2-10 cm long, mainly basal, the stems merely bracteate H. gracile — Leaves long-hairy, mainly 10-20 cm long, the cauline ones well developed, re- duced above H. cynoglossoides Hieracium albiflorum Hook. White stems erect, 20-75 cm tall, pubescent with Hawkweed. Perennial herbs; stems erect, long, coarse, loose or spreading white setae 15-75 cm tall, long-hairy at the base, becom- that dry yellowish; basal leaves commonly ing glabrous upward; basal leaves 2-12 (17) withered at anthesis; lower cauline leaves cm long, 0.8-4.5 cm wide, oblanceolate, pet- 10-25 cm long, 1-3 cm wide, petiolate, the iolate, entire or remotely toothed, long-hairy blades oblanceolate to elliptic, long-hairy; on petioles and midvein, commonly long- middle and upper leaves reduced, sessile; ciliate; cauline leaves sessile, reduced up- heads yellow, few to many, 15- to 40-flow- ward; stellate hairs lacking; heads few to ered, corymbose, the inflorescence more or many, on slender peduncles, the inflorescence less open; involucres 7-12 mm high; bracts open; involucres 6-11 mm high, blackish greenish, the margins chartaceous to hyaline, green, glandular or sparingly long-hairy, or minutely stellate and stipitate-glandular; glabrous; flowers 12-35, white; pappus taw- pappus tawny. Grass-forb, aspen, and spruce- ny. Lodgepole pine, spruce, and spruce-fir fir communities at 200 to 2990 m in Box El- communities at 1980 to 3420 m in Daggett, der. Salt Lake, Summit, Utah, Wasatch, and Duchesne, Sanpete, Uintah, and Utah coun- Weber counties; British Columbia and Al- lies; Alaska and Yukon to Saskatchewan, berta, south to Oregon and Wyoming; 18 (0). south to California, Nevada, and Colorado; Hieracium gracile Hook. Slender Hawk- 12 (ii). weed. Perennial herbs; stems erect, 8-40 cm Hieracium cynoglossoides Arv.-Touv. tall, tomentulous to glabrous; basal leaves Houndstongue Hawkweed. [H. griseum 2-10 cm long, 0.4-2 cm wide, petiolate, Rydb.; H. scouleri, authors]. Perennial herbs; blades oblanceolate, entire or denticulate. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 303 stipitate-glandular to glabrous or less com- monly with a few long blackish setae; cauline leaves much reduced; heads solitary or more commonly greenish black, stellate hairy, and with long black setae; pappus tawny. Lodge- pole pine, spruce-fir, and grass-forb commu- nities at 3050 to 3390 m in Duchesne, Sum- mit, and Uintah counties; Alaska and Yukon to Mackenzie, south to California and New Mexico; South America; 13 (v). wide, sessile, and more or less clasping; heads conspicuous, in racemose or corymbose clus- ters; bracts with long-attenuate, often reddish tips, subequal to the disk; rays reddish to purple or yellowish, hairy and glandular; pappus scales unequal, lacerate; achenes 6-7 mm long. Pinyon-juniper community at 2135 m in the Beaverdam Mountains, Washington County (Higgins 1410 BRY); Nevada and California; 1 (0). HOFMEISTERIA Walp. Shrubs; leaves opposite below, alternate above, simple, petiolate; heads discoid, few to several in terminal corymbose clusters; in- volucre campanulate; bracts striate, narrow, imbricated; receptacle naked; disk flowers whitish; pappus of 10-12 scabrous bristles and other short scales; achenes 5-angled, cal- lous-thickened. Hofmeisteria pluriseta Gray Arrowleaf. Shrubs, low, rounded, and intricately branched, mostly 3-8 dm tall; branchlets green, glandular-puberulent, becoming white barked in age; leaves long petioled, the pet- ioles 0.8-4 cm long, the blades hastately lobed to entire, 4-10 mm long, 2-4 mm wide; heads small; involucres 4-9 mm high; bracts 3-lined, acuminate; disk flowers whit- ish. Reported for Utah in Munz (Flora of Cal- ifornia, p. 267); to be sought on rock out- crops at lower elevations in Washington County; Nevada, California, and Arizona; 0 (0). HULSEA T. & G. Perennial viscid-pubescent aromatic herbs; leaves alternate, simple; heads radiate; in- volucres hemispheric, the bracts subequal in 2 or 3 series, herbaceous, finally reflexed; re- ceptacle convex, naked; ray flowers yellowish to purplish, pistillate, fertile; disk flowers perfect, fertile; pappus of 4 hyaline scales united at the base; achenes compressed, an- gled, villous. Hulsea heterochroma Gray Perennial herbs from a stout taproot; herbage viscid- villous, scented, 3-10 (12) dm tall; basal leaves oblanceolate or spatulate, tapering to a broadly petioled base, dentate; cauline leaves mainly 3-10 cm long, 1.5-3.5 cm Hymenoclea T. & G. Xerophytic shrubs; leaves alternate, linear, usually entire; heads discoid, small, numer- ous, mostly glomerate-paniculate, with both sexes in each leaf axil, the staminate above the pistillate; staminate heads several-flow- ered; pistillate heads 1-flowered; involucre becoming indurated and beaked in fruit, the bracts persistent as scarious wings; pappus none. Hymenoclea salsola T. & G. Burrobrush. Shrubs, 6-12 (15) dm tall; branchlets green, becoming straw colored to gray in age; herb- age yellow green, resinous, glabrous or scabrous; leaves 2-5 cm long, linear, entire; staminate heads 2-3 mm high, 2.5-5 mm wide, the bracts obtuse to rounded, ciliate on the hyaline margin; pistillate heads mainly 6-9 mm high at maturity, the middle and up- per bracts with white, chartaceous, broadly rounded, erose margins, longitudinally veined. Blackbrush, creosote bush, and Joshua tree communities at 670 to 900 m in Wash- ington County; Nevada, Arizona, and Cali- fornia; 19 (ii). Hymenopappus L'Her Perennial herbs; leaves alternate or mainly basal (and still alternate), mainly pinnatifid; heads discoid, the flowers perfect; involucral bracts in 2 or 3 series, subequal, at least the inner with broad rounded scarious or hyaline margins; receptacle flat, naked or rarely chaffy; corollas yellow or white; anthers sag- ittate; pappus of several membranous scales; achenes 15- to 20-nerved, 4- or 5-angled. Turner, B. L. 1956. A cytotaxonomic study of the genus Hymenopappus (Compos- itae). Rhodora 58:163-308. Hymenopappus filifolius Hook. Perennial subscapose herbs; stems 5-60 (100) cm tall. 304 Great Basin Naturalist Vol. 43, No. 2 tomentose to glabrate; basal leaves 3-20 cm long, twice pinnately dissected, the ultimate divisions mainly 2-25 mm long, minutely punctate; cauline leaves lacking or several, much reduced upward; heads solitary or more commonly few to numerous, turbinate to campanulate, with 10-59 flowers or more, on peduncles 0.5-10 cm long or more; in- volucral bracts mainly 3-14 mm long; co- rollas yellow or white, 2-7 mm long; pappus of narrowly oblong scales; achenes 3-7 mm long, densely hairy. This is a polymorphic species, which consists of a series of geo- graphic and/or edaphically correlated in- fraspecific taxa. Those taxa peripheral to the main body of the species in the Colorado Plateau province are the most distinctive. The following treatment differs from that of Turner (1956) and represents a more conserv- ative approach. 1. Basal leaf axils sparingly tomentose or glabrous; stems scapose, or with 1 or 2 leaves; plants of high elevations H. filifolius var. alpestris — Basal leaf axils prominently white-tomentose; stem leaves often more than 2; plants of middle and lower elevations 2 2(1). Corollas 2-3 mm long; flowers fewer than 30; plants of Daggett and Uintah counties H. filifolius var. luteus — Corollas 3-7 mm long, or, if shorter, not of Daggett or Uintah counties; flow- ers in main heads often more than 30; leaves more coarsely dissected; plants of various distribution 3 3(2). Flowers white; achene hairs 0.5-1 mm long; plants of Washington County H. filifolius var. eriopodus — Flowers yellow; achene hairs 1-2 mm long; plants more widely distributed 4 4(3). Leaves mainly basal; plants of the Great Basin H. filifolius var. nanus — Leaves cauline and basal; plants of the Colorado drainage system H. filifolius var. cinereus Var. alpestris (Maguire) Shinners [H. nu- dipes var. alpestris Maguire; H. nudipes Ma- guire; H. filifolius var. nudipes (Maguire) Turner]. Ponderosa pine, western bristlecone pine, sagebrush-grass, limber pine, aspen, and alpine tundra communities, commonly on limestone or thermally modified igneous out- crops, at 2445 to 3450 m, in Beaver, Carbon, Duchesne, Emery, Iron, Garfield, Kane, Mil- lard, Piute, Sanpete, Sevier, Summit, Utah, and Washington counties; Wyoming; 46 (ix). This is the most distinctive of the varieties within H. filifolius in Utah. Var. cinereus (Rydb.) Johnst. [H. cinereus Rydb.; H. lugens Greene; H. filifolius var. lu- gens (Greene) Jepson; H. filifolius var. me- gacephalus Turner, as to Utah materials; H. pauciflorus Johnst., type from near Bluff; H. filifolius var. pauciflorus (Johnst.) Turner; H. tomentosus Rydb., type from St. George; H. filifolius var. tomentosus (Rydb.) Turner; H. niveus Rydb., type from Springdale]. Black- brush, warm desert shrub, salt desert shrub, sand sagebrush-ephedra, pinyon-juniper, pon- derosa pine, and sagebrush communities at 1065 to 2685 m in Carbon, Daggett, Du- chesne, Emery, Garfield, Grand, Kane, San Juan, Uintah, Washington, and Wayne coun- ties; Colorado, California, Arizona, New Mexico, and Texas; 144 (xxix). The variety cinereus, as interpreted herein, includes three largely sympatric phases that were treated by Turner (1956) at varietal rank. Although there is a tendency for these phases to be geographically correlated, they are con- nected completely by series of intermediates, and they can be segregated only arbitrarily. Turner noted that herbarium specimens of var. megacephalus from eastern Utah, inter alia, carried a "hodge-podge of annotations: H. lugens, cinereus, pauciflorus, tomentosus, eriopodus, etc.," and further that "the possi- bility exists that the variety [megacephalus] here typified includes only the individuals from Clark County, Nevada, and vicinity, and that most of the remaining material to the east represents either a weakly defined separate variety or a common area of exten- sive hybridization and introgression among the several peripheral taxa mentioned April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 305 above, ..." A phase of the cinereus complex from Washington County has cauline leaves well developed and plant bases appearing bulbous due to a copious tomentum. These plants are apparently intermediate between var. tomentosus (Rydb.) Turner, in a narrow sense, and var. eriopodus (A. Nels.) Turner, which share the feature of the "bulbous" bases. Var. eriopodus (A. Nels.) Turner [H. eriop- odus A. Nels., type from Diamond Valley]. Pinyon-juniper community at 1675 to 2135 m in Washington County; California to Nevada; 3 (0). This variety is evidently rare in Utah, and might best be treated within an ex- panded var. cinereus (q.v.). Var. luteus (Nutt.) Turner [H. luteus Nutt.]. Salt desert shrub, mixed cool desert shrub, and pinyon-juniper communities at 1525 to 1830 m in Daggett, Summit, and Uintah counties; Wyoming and Colorado; 7 (ii). The small flowers and finely divided leaves appear to be diagnostic for this variety. Var. nanus (Rydb.) Turner [H. nanus Rydb.]. Black sagebrush-rabbitbrush, pinyon- juniper, and ponderosa pine communities at 1490 to 2300 m in Beaver, Garfield, Iron, Juab, Millard, Piute, Sevier, Tooele, and Washington counties; Nevada, California, and Arizona; 26 (v). Hymenoxys Cass. Perennial or biennial herbs from a taproot and commonly with a pluricipital caudex; stems simple or branched; leaves basal or bas- al and cauline, simple and entire or pinnately to ternately divided; heads radiate, peduncu- late; involucres hemispheric; bracts in 2 or 3 series, the outer distinct or connate basally, subequal or imbricate, herbaceous or carti- laginous; receptacle naked, hemispheric; ray flowers yellow, pistillate, fertile, prominently veined, 3-toothed; disk flowers perfect, fer- tile; pappus scales usually 5, hyaline, nerved or nerveless, the nerve often produced into an awn; achenes obpyramidal, more or less 5- angled, appressed hairy. 1. Leaves entire, essentially all basal 2 — Leaves pinnatifid or palmatifid, or some entire, the cauline ones well developed 5 2(1). Involucral bracts sparsely pubescent or glabrous apically, the margins thin and scarious or hyaline 3 — Involucral bracts moderately to densely villous-pilose or some rarely glabrous, the margins not at all or only narrowly scarious 4 3(2). Plants depressed pulvinate-caespitose, acaulescent; outer involucral bracts re- curved, thickened and reddish apically; disks less than 10 mm wide H. lapidicola — Plants merely caespitose, scapose; outer involucral bracts erect, not thickened and seldom reddish apically; disks over 10 mm wide H. torreyana 4(2). Plants pulvinate-caespitose; caudex branches clothed with a marcescent thatch of erect-ascending leaf bases; leaves mainly linear, cuspidate apically H. depressa — Plants seldom pulvinate-caespitose; caudex branches without a definite thatch of ascending or erect leaf bases; leaves various, sometimes cuspidate apically ... H. acaulis 5(1). Disks 18-30 mm wide or more; involucral bracts similar, distinct, in 2 or 3 in- definite subequal series; herbage villous-tomentose; plants of high elevations .... H. grandiflora — Disks 7-22 mm wide; involucral bracts in 2 dissimilar series, the outer thick- ened and united at the base 6 6(5). Plants silvery-canescent; leaves entire or 3-cleft, the blades or segments 1.5-4 mm wide H. subintegra — Plants green, or, if silvery-canescent, the leaves commonly 3- to 5-cleft 7 7(6). Plants apparently biennial, with an evident basal rosette and taproot; cauline leaves numerous, gradually reduced upward, temate or palmatifid H. cooperi 306 Great Basin Naturalist Vol. 43, No. 2 — Plants perennial, from a taproot and caudex; cauline leaves rather well devel- oped, palmatifid to entire 8 8(7). Stems merely glandular or glandular-scabrous; plants of low elevation saline meadows in western Utah H. lemmonii — Stems more or less villous; plants of various habitats, but seldom of saline meadows and not of western Utah 9 9(8). Stems few to several from a pluricipital caudex; leaf bases conspicuously long- villous below; leaf segments mainly 1-2.5 mm wide H. richardsonii — Stems solitary or few from a simple or branched caudex; leaf bases glabrous or only somewhat hairy; leaf segments 2-6 mm wide H. helenoides Hymenoxys acaulis (Pursh) Parker [Gail- lardia acaulis Pursh]. Perennial caespitose herbs from a short multicipital caudex, the caudex branches clothed with short brownish or blackish marcescent leaf bases, 2-50 cm tall, villous to glabrous; leaves 1-6 cm long, 2-8 mm wide, all basal or some cauline, glandular-punctate or epunctate, linear to disk 7-20 mm broad; bracts distinct, in 2 or 3 subequal series, 4-9 mm high; rays 5-9, yel- low, 6-15 mm long; pappus scales 2.5-4.5 mm long, acute or shortly awned; achenes 2.5-4.5 mm long. This is a complex entity, consisting of a series of morphological phases, which are more or less geographically or edaphically correlated. oblanceolate, entire; heads solitary (rarely 2); 1. Plants with 1-4 (or more) cauline leaves; stems simple or branched //. acaulis var. ivesiana — Plants scapose, with cauline leaves lacking or rarely with 1; scapes un- branched 2 2(1). Leaves linear to linear-oblanceolate, conspicuously glandular-punctate, spar- ingly long-hairy to glabrous; plants of the Colorado drainage system H. acaulis var. arizonica — Leaves narrowly to broadly oblanceolate, inconspicuously glandular-punctate, merely punctate, or epunctate, or plants of the Great Basin, densely pilose to villous or glabrous 3 3(2). Leaves epunctate or nearly so, glabrous or less commonly silky-hairy; plants of the Colorado drainage system H. acaulis var. caespitosa — Leaves punctate, silky-hairy, or less commonly glabrous; plants of the Great Basin H. acaulis var. acaulis Var. acaulis Sagebrush, mixed desert shrub, pinyon-juniper, and bunchgrass com- munities, often on windswept ridges, at 1525 to 2990 m in Beaver, Box Elder, Juab, Mil- lard, Sanpete, Sevier, Tooele, and Washing- ton counties; Idaho east to Saskatchewan, south to Nevada, Colorado, and Texas; 42 (viii). Specimens from the Great Basin might not belong to var. acaulis in a strict sense, and perhaps should be regarded as a separate variety. The problem cannot be solved on the basis of Utah specimens alone. Var. arizonica (Greene) Parker [Tetra- neuris arizonica Greene]. Salt and sandy desert shrub, pinyon-juniper, sagebrush, blue grama, aspen, Douglas fir, white fir, and pon- derosa pine communities at 1220 to 3175 m in Carbon, Daggett, Duchesne, Emery, Grand, Kane, San Juan, and Uintah counties; Colorado and Arizona; 108 (xvi). Var. caespitosa (A. Nels.) Parker [Tetra- neuris acaulis var. caespitosa A. Nels.; Tetra- neuris epunctata A. Nels., type from Dyer Mine]. Shadscale-eriogonum, black sage- brush, sagebrush, pinyon-juniper, mountain brush, and alpine tundra, often on plateau margins and windswept ridges, at 1585 to 3510 m in Carbon, Daggett, Duchesne, Emery, Grand, Sanpete, Sevier, Summit, and Uintah counties; Wyoming south to New Mexico; 87 (viii). Var. ivesiana (Greene) Parker [Tetraneuris ivesiana Greene; H. ivesiana (Greene) Park- er]. Sand sagebrush, ephedra, pinyon-juniper, April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 307 and ponderosa pine communities at 1150 to 2505 m in Garfield, Grand, Kane, San Juan, and Wayne counties; Colorado, New Mexico, and Arizona; 66 (xii). This variety approaches phases of the partially sympatric var. arizo- nica in stature, and it is possible to confuse some specimens when cauline leaves are lacking and the stems are unbranched. The varieties acaulis, arizonica, and caespitosa are tetraploids, i.e., 2n = 60, whereas var. ive- siana is diploid, i.e., 2n = 30. Because of this difference, Parker (1960. Leafl. W. Bot. 9: 93) elevated this taxon to specific rank. Hymenoxys cooperi (Gray) Cockerell [Ac- tinella cooperi Gray; A. biennis Gray, type from Washington County?]. Biennial or short-lived perennial herbs; stems 16-60 (80) cm tall, leafy, simple below, branched in a corymbose inflorescence above, often red- dish, scurfy villous, canescent; basal rosette leaves mainly 2-10 cm long, pinnately di- vided, the linear lobes often again divided, mainly 1-1.5 mm wide; stem leaves longer than the internodes; heads (1) 3-50; in- volucres 5-6 mm high, 10-24 mm wide, hemispheric; bracts thickened and united ba- sally, more or less pubescent and glandular; rays 7-13, yellow, 6-15 mm long; pappus scales acuminate; achenes 2-3 mm long, densely pilose. Sagebrush and pinyon-juniper communities at 975 to 2380 m in Garfield, Juab, Kane, and Washington counties; Ne- vada, California, and Arizona; 18 (i). Hymenoxys depressa (T. & G.) Welsh & Reveal [Actinella depressa Gray]. Pulvinate- caespitose scapose perennial herbs from a multicipital caudex, the caudex branches clothed with conspicuous, commonly erect marcescent leaf bases (often forming a thatch), 1-4 cm tall; scapes villous; leaves 0.4-3 (4) cm long, 1-2 (4) mm wide, linear to oblanceolate, the outer sparingly if at all glandular-punctate, the inner definitely so, sparingly villous to glabrous, cuspidate; heads solitary; disk 6-10 mm wide; involucres 4-6 mm high; bracts in 2 or 3 subequal series, long villous, the margins nonscarious, the apices erect; rays 5-7, yellow, 3-6 mm long; pappus scales 2-3 mm long, long-acuminate; achenes 2-3 mm long. Ephedra, sagebrush, shadscale, and pinyon-juniper woodland at 1340 to 2170 m in Duchesne, Emery, and eastern Sevier counties; endemic? There is justification for inclusion of H. depressa with- in the H. acaulis complex, at some in- fraspecific rank. And the plants have been suggested as merely depauperate phases of that group. However, if they are ecologically controlled variations, they should be ex- pected through much of the range of H. acaulis; but they are not. Dwarf forms of H. acaulis, especially of the var. caespitosa, have been mistaken for this species, but they are more hairy, have usually broader leaves, and lack glandular punctate. There is also a ques- tion of typification; the type of H. depressus was taken by Fremont, on his second expedi- tion in the Rocky Mountains. Fremont evi- dently traversed the area occupied by H. depressa in 1845, and the material could have come from western Emery County; 24 (iv). Hymenoxys grandiflora (T. & G.) Parker [Actinella grandiflora T. & G.]. Perennial herbs from a taproot and usually simple cau- dex, this clothed with brown marcescent leaf bases; stems mainly 5-25 cm tall, 1 to sever- al, simple or branched basally, densely vil- lous; leaves basal and cauline, 2-10 cm long, 2- or 3-times ternately or palmately divided, the lobes linear, villous to glabrate; heads solitary; disk 1.5-3 cm wide or more; in- volucral bracts subequal, in 2 or 3 series, 8-14 (16) mm high, densely villous-tomen- tose; rays 15-50, yellow, 25-35 mm long; pappus scales 3.5-7 mm long, attenuate; achenes 3-5 mm long. Sedge-forb commu- nities at or above timberline, often in talus or rockstripes, at 3050 to 3660 m in Duchesne, Grand, Salt Lake, San Juan, Summit, and Utah counties (Uinta, Wasatch, and La Sal mountains); Idaho to Montana, south to Colo- rado; 14 (ii). This is a strikingly beautiful yel- low sunflower of alpine tundra in our mountains. Hymenoxys helenioides (Rydb.) Cockerell [Picradenia helenioides Rydb.]. Perennial herbs from a simple or branched caudex, this clothed with broad brown marcescent leaf bases; stems mainly 25-45 cm tall, simple be- low, branched above, scurfy and more or less villous; leaves basal and cauline, mainly 5-15 cm long, entire or 2- to 5-lobed, the lobes mainly 3-8 mm wide, finely glandular-punc- tate, glabrous or puberulent; heads 3-13, in corymbose clusters; disks 10-21 mm wide; in- volucres 6.5-8 mm high, the outer bracts 308 Great Basin Naturalist Vol. 43, No. 2 green, connate in the lower portion, more or less villous and glandular; rays 5-11, yellow, 8-19 mm long; pappus scales 2.5-3.5 mm long, acuminate; achenes 2.5-3 mm long. Mountain brush, sagebrush, and aspen com- munities, often in meadows, at 2440 to 2990 m in Emery, Garfield, Sanpete, and Sevier counties; Colorado and Arizona; 10 (i). This handsome plant has long remained obscure in Utah, partially due, no doubt, to its resem- blance to Helenium hoopesii (q.v), with which it occurs in the aspen communities of central and southern Utah. Hymenoxys lapidicola Welsh & Neese Pul- vinate caespitose herbs from a multicipital caudex, this densely clothed with brown mar- cescent leaf bases, acaulescent; leaves all bas- al, 0.3-1.2 cm long, 0.8-2 mm wide, narrow- ly oblanceolate, the inner conspicuously glandular-punctate, the blades glabrous, the axils long-villous; heads solitary, immersed in the leaves; disks 5.5-9 mm wide; involucres 5-8 mm high; bracts distinct, in 2 or 3 sub- equal series, sparingly villous and suffused reddish, the margins scarious, the tips more or less squarrose-spreading and somewhat thickened; rays 5 or 6, yellow, 5-6 mm long; pappus scales lance-acuminate, 2.3-3 mm long; achenes 2-2.5 mm long, pilose. Pinyon- juniper and ponderosa pine-manzanita com- munities, often in rock crevices, at 1830 to 2476 m in Uintah County; endemic; 4 (0). Hymenoxys lemmonii (Greene) Cockerel! [Picradenia lemmonii Greene; H. lemmonii ssp. greenei Cockerell, type from Washington County (?)]. Perennial herbs from a taproot and short ligneus caudex, the caudex clothed with brown to straw-colored or purplish clasping leaf bases; stems 20-60 cm tall, 1 to few, glabrous; leaves cauline and basal, 2-15 cm long, pinnately parted, the lobes linear, 2-3 mm wide, glabrous, glandular-punctate; cauline leaves longer than the internodes, the uppermost often unlobed; heads 5-12; in- volucres 4.5-7 mm high, hemispheric, 8-14 mm wide; outer bracts green, sparsely scurfy and glandular, thickened dorsally and con- nate below; rays 6-10, yellow, 6-13 mm long; pappus lance-attenuate, 1.6-2 mm long; achenes 2.5-3 mm long, pilose. Saline rabbitbrush-alkali sacaton meadows at 1660 m in Millard, Tooele, and Washington (?) counties; Nevada and California; 3 (i). Hymenoxys richardsonii (Hook.) Cock- erell Colorado Rubberweed. [Picradenia rich- ardsonii Hook.]. Perennial caespitose herbs from a pluricipital ligneus caudex, the caudex branches clothed with a thatch of marcescent brown leaf bases, usually with villous leaf axils; stems few to numerous, 6-40 (50) cm tall, simple below, branched; leaves basal and cauline 2-12 (15) cm long, ternate or with 5-7 linear segments, or some entire, pu- bescent to glabrous; involucres hemispheric, 5-8 mm high, the outer bracts connate ba- sally, thickened dorsally, green or char- taceous, more or less villous; rays 9-14, yel- low, 8-20 mm long; pappus 2-4.5 mm long, acuminate; achenes 2.5-4 mm long, pilose. Salt desert shrub, cool desert shrub, pinyon- juniper, sagebrush, mountain brush, pon- derosa pine, aspen, fir, and western bristle- cone pine communities at 1460 to 2870 m. Our material falls into two varieties, a low plant with 1-5 large heads of Daggett and Uintah counties, belonging to var. richard- sonii, and a taller plant with 3-20 smaller heads of Beaver, Carbon, Duchesne, Emery, Garfield, Kane, Millard, Piute, Sanpete, Se- vier, Uintah, and Wayne counties, belonging to var. florabunda (Gray) Parker; Alberta and Saskatchewan to Arizona and Texas; 129 (xvii). The plants are considered poisonous to sheep, cattle, and goats. Hymenoxys subintegra Cockerell Per- ennial (or biennial) herbs from a taproot; stems solitary or few, 10-60 cm tall, branched above; herbage silvery canescent; basal leaves often withered at flowering; cauline leaves numerous, 1.5-8 cm long, 2-4 mm wide, entire or 2- or 3-lobed; heads few to several; disks 9-12 mm wide; involucres 5-7 mm high; outer bracts connate basally, villous; rays ca 12-20, yellow, 5-10 mm long; pappus scales lance-acuminate, 2.8-3.2 mm long; achenes ca 3 mm long. Ponderosa pine, aspen, and spruce-fir communities in Sanpete (Maguire 20049 BRY) and Washington (re- ported by Meyer) counties; Arizona; 1 (0). Hymenoxys torreyana (Nutt.) Parker [Ac- tinella torreyana Nutt.]. Perennial caespitose scapose herbs from a stout pluricipital cau- dex, the caudex branches densely clothed with brown to straw-colored or ashy leaf bases, 3-10 cm tall, villous; leaves 1-7.5 (9) April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 309 cm long, 2-6 mm wide, all basal, glandular- punctate, narrowly oblanceolate, entire; heads solitary; disk 12-20 mm wide; in- volucres hemispheric, 5-10 mm high; bracts distinct, in 2 or 3 subequal series, less pu- bescent to glabrous apically, the margins scarious, not thickened apically, the tips erect, sometimes reddish; rays 10-16, yellow, 8-20 mm long; pappus scales ovate-acumi- nate, 2.8-3.5 mm long; achenes 2-3 mm long. Pinyon-juniper, sagebrush, and mountain brush communities at 1830 to 2200 m in Daggett and Uintah counties; Wyoming; 6 (0). Hypochaeris L. Perennial subscapose herbs from taproots, the juice milky; leaves primarily basal, simple, pirmately lobed to pinnatifid, the cauline leaves small and bractlike; heads soli- tary or few in a branching inflorescence; in- volucral bracts in several series, greenish black, the irmer ones with hyaline margins; receptacle chaffy; corollas of ray flowers only, perfect, yellow or purplish on the dor- sal surface; pappus of plumose capillary bristles; style branches semicylindrical; achenes several-nerved, subterete, minutely roughened, long beaked. Hypochaeris radicata L. Cat's-ears. Plants 1.5-5 dm tall, the stems simple or branched above, glabrous or spreading-hairy below; basal leaves 3-16 (25) cm long, 0.5-3.5 (5) cm broad, oblanceolate, pinnately toothed or piimatifid, sparsely to moderately spreading- hairy above and below, rounded to obtuse apically, tapering to a broad petiole basally; cauline leaves alternate, minute or lacking; heads solitary, or more commonly 2-5; pe- duncles glabrous; involucres 5-15 mm high, 7-20 mm wide; bracts glabrous or stiffly hairy along the midribs; corollas numerous, longer than the bracts; achenes 4-7 mm long, the beak mostly 2-3 mm long. Weedy species of disturbed soils in Davis and Salt Lake counties; widespread in North America; ad- vent! ve from Europe; 2 (0). Inula L. Perennial tomentose herbs; leaves basal and cauline, alternate; heads radiate, large, hemispheric, few to numerous in cymose clusters; involucral bracts imbricate in sever- al series; receptacle naked; ray flowers pistil- late, yellow, 3-toothed; disk flowers perfect, fertile; anthers sagittate at the base; style branches of disk flowers linear; pappus of capillary bristles; achenes 4- or 5-ribbed. Inula helenium L. Elecampane. Perennial herbs, mainly 6-20 dm tall, from thick roots; stems simple below; basal leaves 2-5 dm long, petiolate, the blades ovate to oblong, denticulate, rough-hairy above, velvety be- neath; cauline leaves reduced upward, cor- date-clasping, acute; heads large and showy; involucres 15-23 mm high, 30-50 mm wide; outer bracts foliaceous, ovate; ray flowers nu- merous, 18-30 mm long, narrow; achenes glabrous. Canal banks and moist meadows at 1370 to 1830 m in Sanpete and Utah coun- ties; widespread in North America; adventive from Eurasia; 2 (i). IvaL. Annual or perennial herbs; leaves opposite, at least below; heads discoid, the pistillate flowers few, with corolla tubular or lacking; involucres campanulate; bracts subequal or imbricate in 1-3 series, sometimes with a short inner series subtending the achenes; re- ceptacle chaffy, the receptacular bracts lin- ear to spatulate; staminate flowers with abor- tive pistils, the styles undivided, the filaments monadelphous; anthers obtuse basally, almost distinct; pappus none; achenes compressed. 1. Leaves sessile or shortly petiolate, entire; plants rhizomatous, mainly less than 40 cm tall, of saline low-elevation sites 7. axillaris Leaves petiolate, serrate; plants taprooted annuals, mainly much over 40 cm tall, ruderal weeds /. xanthifolia Iva axillaris L. Poverty Weed. Perennial herbs from elongate rhizomes; stems 15-50 (60) cm tall, branched from the base; herbage strigose to strigulose and more or less glandu- lar; leaves opposite below, alternate above, 0.8-4.5 cm long, 4-15 mm wide, elliptic to 310 Great Basin Naturalist Vol. 43, No. 2 obovate or lanceolate; heads numerous in ter- minal bracteate spicate clusters, nodding, 3-7 mm wide; bracts connate, shallowly 4- or 5- lobed; pistillate flowers 4-8, perfect; achenes 2-3 mm long, glandular. Commonly in saline riparian sites in the warm desert shrub, salt desert shrub, pinyon-juniper, and aspen com- munities at 760 to 2440 m in all Utah coun- ties; British Columbia to Manitoba, south to California, New Mexico, and Oklahoma; 60 (viii). Iva xanthifoUa Nutt. Marsh-elder. Coarse perennial herbs, mainly 4-25 dm tall, simple or branched, essentially glabrous below, glandular above; leaves opposite below, pet- iolate, the blades 4-20 cm long and about as wide, broadly ovate to lance-ovate, serrate and sometimes lobed, green above, canescent beneath; heads 2-4 mm thick, numerous, borne ebracteate in paniculate clusters; in- volucral bracts distinct, ovate; pistillate flow- ers 5; achenes sparsely pilose apically, ca 2 mm long. Ruderal weeds of disturbed soils at 1370 to 2290 m in Beaver, Duchesne, Emery, Iron, Kane, Millard, Salt Lake, Sevier, Sum- mit, Uintah, Utah, and Wayne counties; Al- berta to Saskatchewan, south to Washington, Arizona, and New Mexico; widely distributed elsewhere; 22 (iii). KUHNIA L. Perennial herbs from a woody caudex and taproot; stems branched, erect or ascending; leaves alternate or some lower ones opposite; entire or lobed; heads discoid, several to nu- merous in paniculate clusters; involucres campanulate; involucral bracts in 4-7 series, the outer ones only graduated; receptacle naked; disk flowers perfect, fertile, whitish; style tips flattened, clavate; pappus of plu- mose bristles; achenes 10-ribbed. Kuhnia chlorolepis Woot. & Standi. Per- ennial clump-forming herbs; stems 30-75 cm tall, much branched, minutely hairy; leaves 8-50 mm long (or more), 1-3 mm wide, en- tire or with a pair of basal lobes, linear; in- volucres 8-12 mm high; bracts linear to nar- rowly oblong, striate; corollas 6-7.5 mm long; achenes 4.8-5.2 mm long, dark brown, short-hairy. Rabbitbrush community in inter- mittent stream courses at 1890 to 2045 m in Uintah County; Colorado to Arizona, New Mexico, Texas, and Mexico; 2 (i). Lactuca L. Annual, biennial, or perennial herbs; leaves alternate, entire or pinnatifid; flowers all raylike, yellow, blue, or white; heads panic- ulately arranged; involucres cylindrical; bracts imbricate in several series; receptacle flat, naked; pappus copious, of white or brownish capillary bristles; achenes oval, ob- long, or linear in outline, compressed, ribbed on each face, short- to long-beaked. 1. Plants perennial, rhizomatous; rays long-exserted, blue L. tatarica — Plants annual or biennial; rays not long-exserted, yellow (often fading blue) or blue to white (in L. biennis) 2 2(1). Achenes prominently 1-nerved on each side 3 — Achenes prominently several nerved on each side 4 3(2). Involucres 10-15 mm high in fruit; pappus 5-7 mm long; achenes 4.5-6.5 mm long, including the beak L. canadensis — Involucres 15-22 mm high in fruit; pappus 7-12 mm long; achenes 7-10 mm long L. ludoviciana 4(2). Involucres cylindrical at anthesis; flowers not fading blue; plants cultivated and occasionally escaping L. sativa — Involucres tapering to the apex at anthesis; flowers fading blue 5 5(4). Achenes with a long filiform beak as long as or longer than the body of the achene; pappus white L. serriola — Achenes with a short beak much shorter than the body of the achene; pappus brownish L. biennis April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 311 Lactuca biennis (Moench) Fern. [Sonchus biennis Moench]. Annual or biennial, glabrous or hairy (on midvein of leaves) herbs; stems erect, mainly 6-20 (35) dm tall; leaves mainly 10-40 cm long, 4-20 mm wide, pinnatifid or merely toothed; heads 13- to 50- flowered, nimierous, arranged in a narrow paniculate inflorescence; rays bluish to white; pappus brownish; achenes 4-5.5 mm long, prominently several nerved on each face, beakless or short beaked. Moist sites at ca 1800 m in Salt Lake County (Arnow 2561 BRY, UT); Alaska to Newfoundland, south to California, Colorado, and North Carolina; 1 (0). Lactuca canadensis L. Annual or biennial, glabrous or hirsute herbs; stems erect, 3-25 dm tall; leaves mainly 10-35 cm long, 1-12 cm wide, entire to pinnatifid; heads mostly 13- to 22-flowered, arranged in open pan- icles; rays yellow (fading blue); pappus white; achenes black, obovate, transversely rugose and with 1 prominent longitudinal vein on each face, 4.5-6.5 mm long, includ- ing the beak from half as long to as long as the body. Weedy species of moist sites at ca 1155 m in Kane County (Atwood 4118 BRY); widespread in U.S.; 1 (0). Lactuca ludoviciana (Nutt.) Riddell [Son- chus ludovicianus Nutt.]. Biennial or short- lived perennial herbs; stems 6-15 dm tall or more; leaves 10-35 cm long or more, mainly 1-10 (20) cm wide, commonly pinnatifid and weakly spinose-toothed, setose-hispid on the lower midrib, the uppermost auriculate- clasping; heads numerous in an open pan- iculate cluster, the peduncles bracteate; in- volucres 15-22 mm high in fruit; heads most- ly 20- to 50-flowered, the flowers yellow or sometimes blue, fading blue; pappus white, 7-10 mm long at maturity; achenes flattened, blackish, with a longitudinal median nerve on each face, transversely rugulose, 4-5 mm long. Collected once in Salt Lake County (without collector, UT); widespread in the northwestern U.S.; 1 (0). Lactuca sativa L. Lettuce. Annual herbs; stems erect, mostly 5-12 dm tall, glabrous; leaves mainly 10-30 cm long and as broad, undulate-crisped and serrate, glabrous; in- volucres 7-10 mm high; heads ca 15-flow- ered, the flowers yellow, not fading blue, numerous in a paniculate cluster; pappus white; achenes brownish, oblanceolate in out- line, flattened, hispid apically, 3.5-4.5 mm long, with 5-7 longitudinal nerves on each face, the beak 2.5-3.5 mm long. Cultivated food plant in much of Utah; introduced from Europe; 2 (0). Lactuca serriola L. Prickly Lettuce. [L. scariola, scarriola, orthographic variants]. Biennial or winter annual herbs; stems erect, 3-18 dm tall, hispid below or glabrous over- all; leaves mainly 3-30 cm long, 1-10 cm wide, pinnatifid or pinnately lobed, or merely spinose-toothed, the blades vertically oriented (twisted at the base), setose-hispid on main veins beneath; involucres 7-15 mm high at maturity; heads mostly 6- to 12-flow- ered, the flowers yellow, fading blue, several to numerous in a paniculate cluster; pappus white; achenes brown, the body obovate to oblong in outline, flattened, hispid along margin apically, 3-4.5 mm long, with 5-8 longitudinal nerves on each face, the beak 3-7 mm long. Ruderal weeds at 850 to 2440 m, probably in all Utah counties, widely dis- tributed in the U.S.; adventive from Europe; 37 (v). This species invades lower elevation range lands, where it is eaten by wildlife and livestock. It is reported to produce fertile hybrids with L. sativa (q.v.). Lactuca tatarica (L.) CA. Mey. Blue Let- tuce. Perennial rhizomatous herbs; stems 2-12 dm tall, glabrous; leaves 4-20 cm long, 5-35 mm wide, linear to lanceolate or ob- long, entire, toothed, lobed, or pinnatifid, short-petiolate below, sessile above; in- volucres 10-20 mm high; heads cylindric, 15- to 50-flowered, the flowers blue, numerous in an elongate paniculate cluster; pappus white; achenes black to pale, oblong-lanceolate in outline, flattened, 4-7 mm long, with several longitudinal nerves on each face, the beak ca 2 mm long. Marshes, canal and stream banks, and roadsides at 1370 to 2440 m in Cache, Daggett, Duchesne, Garfield, Grand, Iron, Juab, Kane, Millard, Piute, Salt Lake, Sevier, Tooele, Uintah, Utah, and Weber counties; Alaska to Minnesota, south to California and Missouri; 39 (vii). Our material belongs to ssp. puhhella (Pursh) Stebbins [Sonchus pul- chellus Pursh], the North American phase of a circumboreal species. 312 Great Basin Naturalist Vol. 43, No. 2 Lapsana L. Annual herbs from taproots, the juice milky; leaves alternate, simple, subentire to toothed or lyrate-pinnatifid; heads numerous; involucral bracts in 2 series, the inner ones large and keeled, the outer minute, greenish; receptacle naked; corollas of ray flowers only, perfect, yellow; pappus none; style branches semicylindrical; achenes subterete, several-nerved, tapering at both ends, beakless. Lapsana communis L. Nipplewort. Plants mostly 2.5-10 dm tall, the stems erect, simple or branched, pubescent with stipitate glands or glabrous; leaves mostly 3-10 cm long and 1.4-5 (7) cm wide, the blades subentire to toothed, or the lower ones lyrate-pinnatifid, sparsely hairy to glabrous above and below; heads numerous, the peduncles glabrous or nearly so; involucres 5-8 mm high, 3-9 mm broad, the bracts glabrous; flowers mostly 10-14; achenes 3-5 mm long. Weedy species of disturbed sites in Salt Lake County (Arnow 4747, BRY; UT); widely established in North America; adventive from Eurasia; 1 (0). Layia H. & A. Annual herbs from taproots; leaves mainly alternate, subentire to toothed or pinnatifid; heads radiate, solitary or few to several, sub- corymbose; ray and disk flowers both fertile; involucres campanulate to broadly hemi- spheric; bracts with thin margins abruptly di- lated below, enclosing the ray achenes; re- ceptacle plano-convex, chaffy marginally; ray flowers 8-24, yellow or with the tips white; pappus of numerous bristles, awns, or scales, the bristles often plumose below; ray achenes obcompressed, commonly glabrous and epap- pose; disk achenes pubescent and pappose. Ray flowers yellow with a white tip; pappus setae merely scabrous; anthers black; plants rare in San Juan County L. platyglossa Ray flowers white; pappus setae plumose; anthers yellow; plants locally com- mon, widespread L. glandulosa Layia glandulosa (Hook.) H. & A. Tidy- tips. [Blepharipappus glanduhsus Hook.]. Plants slender, the stems simple or branched, 0.8-3 dm tall or more, often reddish, with long spreading-ascending multicellular setae; leaves 0.8-6 cm long, 1.5-16 mm wide, often mainly basal, hispid, toothed to lobed, the cauline ones reduced upward and finally en- tire; heads solitary or 2 to numerous; in- volucres 6-9 mm high, 10-18 mm wide; bracts hispid and with some tacklike purplish black stipitate glands; rays white, 6-15 mm long; disk flowers numerous; ray achenes 3-4 mm long; disk achenes 3-6 mm long; pappus of 10-12 white flattened setose scales plu- mose to above the middle with straight capil- lary and tangled woolly hairs. Sagebrush- grass, grassland, and pinyon-juniper commu- nities at 1370 to 1865 m in Daggett, Garfield, Juab, Kane, Millard, Salt Lake, Sanpete, Tooele, Utah, and Washington counties; Brit- ish Columbia, south to Baja California and Arizona; 24 (i). Layia platyglossa (Fisch. & Mey.) Gray [Callichroa platyglossa Fisch. & Mey.]. Plants slender, the stems erect, simple or branched. setose with long, multicellular hairs, often reddish; leaves mainly 1-6 cm long, 2-7 mm wide, with long, slender, spreading multi- cellular hairs, the cauline leaves reduced up- ward and finally entire; heads solitary or few; involucres 6-12 mm high, 12-20 mm wide; bracts hairy like the leaves and with some tacklike purplish black stipitate glands; rays yellow with white tips, 6-18 mm long; disk flowers numerous; ray achenes 3-4 mm long; disk achenes 3-5 mm long; pappus of scabrous setae. Dunes at ca 1375 m in San Juan County (Harrison 2545 BRY); Califor- nia. Our material apparently belongs to var. breviseta Gray [ssp. campestris Keck], and this is apparently the only known station for the species east of California. The collection was taken in 1927. The plants resemble those of Gaillardia, in a general way, and our ma- terial has been filed for more than four de- cades in a folder of that genus. Lepidospartum Gray Shrubs; leaves alternate, linear, entire; heads several to numerous, in corymbose or April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 313 racemose clusters; heads discoid, the flowers perfect, yellow; involucres subcylindric; bracts chartaceous, inbricate in several series, rounded apically (at least the inner); recep- tacle flat, naked; anthers sagittate; style branches flattened; pappus of copious capil- lary bristles; achenes oblanceolate in outline, long-pilose. Lepidospartum latisquamum Wats. Ne- vada Broomshrub. Shrubs mainly 6-15 dm tall or more; branchlets with prominent lon- gitudinal striae, the striae glandular, the in- tervening areas tomentose; leaves 0.5-3 cm long, linear, 0.5-1 mm wide, apiculate; heads 4- to 7-flowered; involucres 8-10 mm high, 3.5-6 mm wide; bracts chartaceous, tomen- tose, the outer apiculate, very short, the inner broadly rounded and more or less hyaline margined; achenes 4-5 mm long, long-pilose with copious white hairs 3-4 mm long. Rab- bitbrush community along a wash at 1705 to 1740 m in Millard County (Pine Valley); Ne- vada and California; 7 (iii). Leucelene Greene Perennial rhizomatous herbs; leaves alter- nate, simple, entire, linear or subulate; heads radiate, solitary or few to many; involucres turbinate; bracts imbricate in several series, green, the margins scarious; ray flowers white or tinged pink, pistillate; disk flowers perfect, fertile, yellow; pappus of capillary bristles; achenes subcylindric or somewhat compressed. Shinners, L. H. 1946. Revision of the genus Leucelene Greene. Wrightia 1:82-89. Leucelene ericoides (Torr.) Greene Rose- heath. [Inula ? ericoides Torr.; L. arenosa Heller; Aster bellus Blake; A. leucelene Blake; A. hirtif alius Blake]. Perennial herbs from a branching caudex and rhizome, simple or more commonly branched, 3-17 cm tall, strigose and more or less glandular; leaves 2-10 mm long, 1-2 (3) mm wide, linear to spatulate, becoming subulate upward; heads solitary or few to many; involucres 5-7 mm high, 5-12 mm wide; bracts in 3-5 series; rays 12-25, white to pink, 3-6 mm long; achenes appressed-hairy. Blackbrush, desert shrub, salt desert shrub, pinyon-juniper, and ponderosa pine communities at 1370 to 2595 m in Beaver, Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Iron, Juab, Kane, Millard, Piute, Salt Lake, San Juan, Sanpete, Sevier, Tooele, Uintah, Utah, Washington, and Wayne counties; Nevada and California, east to Kansas, south to Arizona and Mexico; 145 (xvii). Lygodesmia D. Don Perennial or annual herbs with milky juice; leaves alternate or mainly basal and still al- ternate, entire or pinnatifid; heads solitary or few to many in corymbose or paniculate clus- ters; flowers all raylike, pink to pink-purple or white; involucres cylindric; bracts mostly 5-9, with some more or less reduced outer ones; receptacle naked; pappus of numerous capillary bristles; achenes linear, subterete, prominently several nerved. 1. Rays 10-12 mm long, ca 4 mm wide; pappus 6-9 mm long L. juncea — Rays 15-25 mm long, 6-10 mm wide; pappus 12-17 mm long 2 2(1). Flowers white (or pink?, and drying pinkish); stems ligneous, branching from the base, forming rounded clumps; leaves stiff, spreading; plants of Emery and Grand counties L. entrada — Flowers pink or pink-purple; stems various, but, if branched from the base, the leaves eitiier lax or the plants of different distribution L. grandiflora Lygodesmia entrada Welsh & Goodrich Entrada Rushpink. Perennial herbs from a subterranean caudex, branching from the base, the branches ligneus and wiry, mainly 25-45 cm tall; leaves entire, linear or acicu- lar, 5-30 (70) mm long; peduncles with nu- merous bracts, 12-20 cm long; involucral bracts hyaline-margined, the outer 5-10 mm long, fimbrillate, the inner ca 6, 16-18 mm long, puberulent at the apex; rays white, ca 3 cm long; pappus barbellate, sordid, 10-15 mm long; achene ribs glabrous. Juniper and mixed desert shrub communities at 1340 to 1465 m in Emery and Grand (type from near 314 Great Basin Naturalist Vol. 43, No. 2 Courthouse Wash) counties; endemic; 3 (i). The status of this entity is unclear; certainly it is a portion of the grandiflora complex. Further work is indicated. Lygodesmia grandiflora (Nutt.) T. & G. Showy Rushpink. [Erythremia grandiflora Nutt.]. Perennial herbs from deeply placed elongate rhizomes; stems 0.6-5 dm tall, simple or branched from the base or above; leaves alternate, 0.5-10 cm long or more, 1-5 mm wide, attenuate, gradually to abruptly reduced upward; involucres cylindric, 18-25 mm high, densely hairy to glabrous (?), the outer mostly short and ovate to lance-ovate, the inner 5-9 equal, narrowly oblong; rays 5-10, pink, pink-purple, or rarely white, mostly 2-4 cm long; pappus of numerous barbellate tawny bristles; achenes 12-18 mm long, ribbed, glabrous. Our material consists of three taxa, which have been regarded at specific rank. Intermediates between the taxa suggest a more conservative approach. 2(1). Main involucral bracts 8 or 9; flowers 8-12; plants of east central and north- eastern Utah L. grandiflora var. grandiflora Main involucral bracts 5 or 6; flowers 5-7 (10); plants of southeastern and western Utah 2 Uppermost leaves reduced to linear scales mainly 3-10 mm long; achenes 13-19 mm long, smooth on the lower surface L. grandiflora var. dianthopsis Uppermost leaves not reduced to scales, mainly 20-40 mm long; achenes 10-13 mm long, rugose on the lower surface L. grandiflora var. arizonica Var. arizonica (Tomb) Welsh comb. nov. [based on: Lygodesmia arizonica Tomb Sida 7:530. 1970]. Blackbrush-ephedra and Indian ricegrass-dropseed communities at 1125 to 1590 m in Kane and Wayne counties; Ari- zona; 7 (ii). Var. dianthopsis (D.C. Eaton) Welsh comb. nov. [based on: Lygodesmia juncea var. dianthopsis D.C. Eaton in Watson Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5:200. 1871; L. diantliopsis (D.C. Eaton) Tomb]. Sagebrush-grass, pinyon-juniper, and moun- tain brush communities at 1370 to 2440 m in Beaver, Cache, Kane, Millard, Salt Lake, Se- vier, and Utah counties; Nevada; 22 (iii). In- termediate specimens transitional to var. arizonica occur in south central Utah. Var. grandiflora [L. grandiflora var. stricta Maguire, type from south of Price]. Shad- scale, sagebrush, pinyon-juniper, mountain brush, ponderosa pine, and aspen-sagebrush communities at 1460 to 2750 m in Carbon, Daggett, Duchesne, Emery, Garfield, Grand, and Uintah counties; Wyoming south to New Mexico; 37 (vii). The var. stricta is a phase with stiffly erect leaves, but seems to repre- sent only an ecological variant. Specimens of intermediate nature occur southward with both varieties arizonica and dianthopsis. Lygodesmia juncea (Pursh) D. Don [Pre- nanthes juncea Pursh]. Perennial glabrous herbs from a deeply placed elongate root (rhizome?); stems mainly 1.5-6 dm tall, much branched; leaves stiff, entire, mainly 1-4 cm long, 1-4 mm wide, the upper ones reduced to subulate scales; heads few to several, mainly 5 (4-10) -flowered; flowers pink or less commonly white; involucres 9-16 mm high, with 4-8 main bracts and several short- er outer ones; pappus tawny; achenes ca 5-7 mm long, several nerved. Our few specimens from sandy sites in mixed desert shrub and juniper communities at ca 1400 to 1590 m in Emery and Juab counties; British Columbia to Minnesota, south to Arizona and Arkansas; 3 (0). This is mainly a Great Plains species, with disjunct populations westward, often in sandy habitats. Machaeranthera Nees Annual, biennial, or perennial herbs from taproots; leaves alternate, entire or pinnatifid to toothed or lobed, spinulose apically and the teeth, when present, spinulose; heads soli- tary or 2 to numerous; involucral bracts in several series, herbaceous apically, char- taceous or coriaceous basally, mainly squar- rose; rays pistillate and fertile, pink, laven- der, pink-purple, or white, or lacking; receptacle naked; anthers not caudate; pap- pus of capillary bristles; achenes narrowly oblong in outline. Cronquist, a. and D. D. Keck. 1957. A re- constitution of the genus Machaeran- thera. Brittonia 9:231-239. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 315 1. Heads discoid; leaves spinose-toothed M. grindelioides — Heads radiate; leaves various, but not conspicuously spinose-toothed 2 2(1). Plants perennial, from a definite caudex, of montane sites, commonly on gran- ite, limestone, or quartzite Aster kingii D.C. Eaton (q.v.) — Plants biennial, winter annual, or annual (rarely short-lived perennial), the caudex not well developed; plants of various habitats and substrates 3 3(2). Leaves pinnately dissected; plants annual M. tanacetifolia — Leaves merely toothed to entire; plants mainly biennial or short-lived per- ennial 4 4(3). Involucral bracts with green tip commonly equaling or longer than the char- taceous base, the long-tapering apices often reflexed M. bigelovii — Involucral bracts with green tip much shorter than the chartaceous base, the reflexed to erect tips shortly attenuate to acute M. canescens Machaeranthera bigelovii (Gray) Greene [Aster bigelovii Gray; M. mucronata Greene, sensu Utah materials]. Short-lived perennial (biennial in some?) herbs from a taproot, a caudex not or only poorly developed; stems 11-35 cm long, puberulent below, becoming glandular to stipitate-glandular above; basal leaves often withered at anthesis; cauline leaves oblanceolate to linear or oblong, mainly 1-7.5 cm long, 1.5-8 mm wide, the surfaces glabrous and more or less glandular or stipitate-glandular, ciliate, entire to spi- nose-toothed; heads few to many in corym- bose inflorescences; involucres 9-12 mm high, 12-23 mm wide; bracts lance-linear, at- tenuate apically, the green apex subequal to the coriaceous base, especially in the outer bracts, commonly spreading-reflexed, glandu- lar and glandular-ciliate; rays 21-31, violet or pink-purple, 10-15 mm long, 2-4.2 mm wide; pappus off-white; achenes glabrous or sparingly strigose, 2.5-4.2 mm long. Moun- tain brush, aspen, spruce-fir, and alpine meadow communities at 2440 to 3355 m in Garfield, Iron, Kane, and Washington coun- ties (Henry Mountains, Markagunt Plateau, and Kolob Terrace); Colorado, New Mexico, and Arizona; 18 (iii). Machaeranthera canescens (Pursh) Gray [Aster canescens Pursh]. Biennial (winter an- nual) or short-lived perennial herbs from a taproot, a caudex seldom developed; stems 8-60 cm tall or more, variously glabrous, glandular, or puberulent; basal leaves with- ered or persistent at anthesis; cauline leaves linear to oblong or oblanceolate, 1-10 cm long, 1-22 mm wide, the surfaces glabrous, puberulent, or glandular, commonly ciliate, entire or toothed; heads few to many in pa- niculate to corymbose clusters; involucres 6-10 (12) mm high, 6-18 mm wide; bracts linear to oblong, attenuate to abruptly atten- uate, the green apex commonly much shorter than the coriaceous base, sometimes spread- ing-reflexed, glandular and or puberulent; rays 15-25, pink to pink-purple or white, 5-12 mm long, 1.5-2.5 mm wide; pappus off- white; achenes pilose, ca 2.5 mm long. The canescens complex consists of a series of in- tergrading taxa, which, in the extreme, are distinctive and geographically or edaphically correlated. Many names have been applied to members of the complex, and specimens of- ten bear annotations of several of the names involved. This is partially in recognition of the intermediate nature of the specimens and partially due to the quality of diagnostic cri- teria. It seems best to treat the materials from Utah as belonging to a single poly- morphic species, consisting of four inter- grading varieties. Leaf surfaces glabrous, the upper leaves commonly glandular to stipitate- glandular; upper branches usually with numerous bracteate leaves; plants of southeastern Utah, rarely elsewhere M. canescens var. vacans Leaf surfaces puberulent, the upper leaves sometimes also glandular; upper branches lacking bracteate leaves or variously so; plants of broad distribution, but not of southeastern Utah 2 316 2(1). Great Basin Naturalist 3(2). Vol. 43, No. 2 Upper branches with numerous bracteate leaves; rosette leaves abruptly and angularly lobed or toothed; plants biennial, of central and southwestern Utah ... M. canescens var. leucanthemifolia Upper branches seldom especially bracteate; rosette leaves various; plants biennial or short-lived perennial, of various distribution 3 Involucral bracts 1-1.5 (2) mm broad, abruptly attenuate apically; plants often perennial, mainly of higher middle elevations M. canescens var. commixta Involucral bracts 0.5-1 mm wide, rather gradually attenuate apically; plants often biennial, mainly of lower to middle elevations M. canescens var. canescens Var. canescens [M. pulverulenta (Nutt.) Greene]. Salt desert shrub, mixed desert shrub, pinyon-juniper, mountain brush, as- pen-sagebrush, Douglas fir, and lodgepole pine communities at 1250 to 2900 m in Bea- ver, Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Iron, Juab, Kane, Millard, Piute, Salt Lake, Sanpete, Sevier, Summit, Uintah, and Washington counties; British Co- lumbia to Saskatchewan, south to California, Arizona, and Colorado; 102 (xiii). This is a variable complex of forms that differ in sev- eral morphological features, but further seg- regation seems unwarranted. I have been unable to distinguish M. tephrodes (Gray) Greene from among our rather large collection. Var. commixta (Greene) Welsh comb. nov. [based on: Machaeranthera commixta Greene Pittonia 4:71. 1899, type from the Henry Mountains; M. latifolia A. Nels., type from Big Cottonwood Canyon, M. leptophylla Rydb., type from Logan; M. paniculata A. Nels., type from Parleys Canyon; M. rubri- caulis Rydb.; Aster rubrotinctus Blake]. Mountain brush, aspen, Douglas fir, sage- brush, spruce-fir, and alpine meadow com- munities at 1705 to 3420 m in Beaver, Cache, Carbon, Duchesne, Emery, Garfield, Iron, Juab, Millard, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, and Wayne counties; Wyoming and Colorado; 82 (v). Var. leucanthemifolia (Greene) Welsh comb. nov. [based on: Aster leucanthemi- folius Greene Erythaea 3:119. 1895; M. leu- canthemifolia (Greene) Greene]. Blackbrush, mixed desert shrub, pinyon-juniper, mountain brush, and ponderosa pine communities at 915 to 2135 m in Beaver, Iron, Juab, Sanpete, Sevier, Utah, and Washington counties; Ne- vada and Arizona; 43 (xiii). This plant is mainly a xerophyte of sandy and silty habi- tats at lower elevations in the Great Basin and lower Virgin River drainage systems; it is transitional at higher elevations with the pre- ceding varieties. Phases of var. canescens from northeastern Utah have been regarded as portions of this variety, but they seem not to fit the concept of var. leucanthemifolia, whose type is from Mineral County, Nevada. Var. vacans (A. Nels.) Welsh comb. nov. [based on: Machaeranthera pulverulenta var. vacans A. Nels. Bot. Gazette 56:70. 1913, type from San Juan County, Utah]. Salt desert shrub, mixed desert shrub, pinyon-juni- per, and ponderosa pine communities at 1155 to 2380 m in Carbon, Emery, Garfield, Grand, Kane, Juab, San Juan, and Washing- ton counties; Colorado, Arizona, and New Mexico; 61 (xix). This material has been treated as M. linearis Rydb., a glabrous- leaved phase of M. canescens whose type came from Yellowstone Park, Wyoming. Work of a monographic nature is necessary for the entire canescens complex. Additional research might indicate an older name at va- rietal rank for this taxon. Machaeranthera grindelioides (Nutt.) Shinners [Eriocarpum grindelioides Nutt.; Haplopappus nuttallii T. & G.]. Perennial herbs from a woody caudex and stout tap- root, the caudex branches more or less clothed with marcescent leaf bases; stems 2-30 cm tall, pilosulose or spreading-hairy below, stipitate-glandular and/or hairy above; basal leaves withered or persistent at anthesis; cauline leaves oblanceolate to spatulate or oblong, mainly 0.5-4.5 cm long, 2-12 mm wide, serrate, the teeth with spi- nulose tips 1-3 mm long, the surfaces pilosu- lose and/ or stipitate-glandular; heads solitary or few to many in corymbose clusters; in- volucres 6.5-9.5 mm high, 8-15 mm wide; April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 317 bracts narrowly oblong, attenuate to an acute apex, the apical portion green or brown, the base chartaceous, erect, glandular; rays lack ing; pappus off-white to brownish; achenes densely pilose, ca 3 mm long. Two distinctive varieties are present in Utah. 1. Plants dwarf, often monocephalous; leaves commonly clustered at stem bases; plants of semibarren habitats in the Great Basin M. grindelioides var. depressa — Plants seldom dwarf, often with more than 1 head; leaves mainly cauline; plants seldom of the Great Basin M. grindelioides var. grindelioides Var. depressa (Maguire) Cronq. & Keck [Haplopappus nuttallii var. depressa Ma- guire, type from Millard County]. Mixed desert shrub, pinyon-juniper, and mountain brush communities at 1465 to 2320 m in Bea- ver, Juab, and Millard counties; Nevada, a Great Basin endemic; 24 (xiv). Var. grindelioides Blackbrush, mixed desert shrub, sagebrush, pinyon-juniper, and mountain brush communities at 1340 to 3175 m in Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Juab, Kane, Millard, Rich, San Juan, Sanpete, Sevier, Summit, Uintah, and Utah counties; Montana to Saskatche- wan, south to Nevada, Arizona, and New Mexico; 98 (xiii). There is a tendency for leaves of plants from the Great Basin to be more glandular than for those in the main body of distribution in the Colorado drainage system. Machaeranthera tanacetifolia (H.B.K.) Nees [Aster tanacetifolius H.B.K.]. Annual (winter annual) herbs; stems 8-50 cm tall, glandular-puberulent and more or less vil- lous; leaves 1-6 cm long, 1- or 2-pinnatifid, the segments ending in spinulose bristles; heads 1 to many, in corymbose clusters; in- volucres 8-12 mm high, 10-18 mm wide, hemispheric; bracts lance-linear, attenuate. chartaceous basally, green apically, spread- ing, the reflexed tips glandular; rays 11-23 (36), pink- or blue-purple, 11-14 mm long; pappus off-white; achenes ca 2.5 mm long, pilose. Mixed desert shrub, salt desert shrub, and pinyon-juniper communities at 1125 to 1830 m in Emery, Garfield, Grand, Juab, Kane, San Juan, Sevier, Utah, Wasatch, Washington, and Wayne counties; 31 (vi). The plants are somewhat weedy, colonizing disturbed sandy and silty soils. The similar M. parviflora Gray [Aster parvulus Blake] is re- ported for Utah by various authors. It differs in having once-pinnatifid leaves, involucres 4-6 mm long, and rays 5-7 mm long. No ma- terial has been seen from Utah by me. Madia Molina Annual or biennial tar-scented herbs from taproots; leaves opposite below, alternate above, simple, entire; heads radiate, the rays pistillate, fertile, yellow, or inconspicuous; involucral bracts uniseriate, equal, enfolding the ray achenes; receptacle flat or convex, with a single series of bracts between the ray and disk flowers; disk flowers perfect; pappus none, a short crown, or a few scales; achenes finely striate, commonly incurved, compressed. 1. Heads turbinate-ovoid, 6-12 mm wide (when pressed); rays 4-7 mm long, showy M. gracilis — Heads ellipsoid, 2-5 mm wide (when pressed); rays to 2.5 mm long, or lacking . M. glomerata Madia glomerata Hook. Tarweed. Annual herbs; stems mainly 8-40 (60) cm tall; herb- age strigose and with long setiform multi- cellular hairs on leaf bases and on stems above, and stipitate-glandular upward, mal- odorous; leaves 1.2-9 cm long, 1.5-7 mm wide, linear; heads in dense terminal clusters or sometimes open; involucres 5.5-9 mm high, 2-5 mm wide; rays inconspicuous. mostly 1.5-2.5 mm long, yellow or purplish; disk flowers 1-10; achenes 5-nerved, glabrous. Sagebrush, mountain brush, aspen, spruce-fir, grass-forb, and alpine meadow communities at 1830 to 3175 m in Cache, Carbon, Davis, Duchesne, Emery, Iron, Juab, Piute, Salt Lake, Sanpete, Sevier, Summit, Tooele, Utah, Wasatch, Washington, and Weber counties; Alaska to Saskatchewan, 318 Great Basin Naturalist Vol. 43, No. 2 south to California, Arizona, and Colorado; 38 (iv). Madia gracilis (Smith) Keck [Sclerocarpus gracilis Smith in Rees]. Annual herbs; stems mainly 10-60 (100) cm tall; herbage pilosu- lose, becoming hirsute with long multi- cellular hairs upward, stipitate-glandular with dark capitate glands on the peduncles and sepals; leaves 1-10 cm long, 2-7 (10) mm wide, linear to elliptic or oblong; heads sev- eral to many in an open corymbose cluster involucres 6-11 mm high, 6-12 mm wide rays conspicuous 5-13, yellow, 4-7 mm long disk flowers 15-35; achenes often mottled. Opening in mountain brush community at ca 1925 m in Salt Lake County; British Colum- bia to Montana, south to California; 1 (0). Malacothrix DC. Annual (winter annual) or perennial herbs from taproots with milky juice; leaves alter- nate or mainly basal, mostly pinnatifid; heads of ray flowers only, long-peduncled, solitary or few to several and more or less corymbose; flowers yellow; involucres campanulate; bracts subequal in 2-4 series, with a few short outer ones; receptacle flat, setose or naked; rays 5-lobed; pappus of capillary bristles, these more or less united at the base and falling together or with some persistent; achenes columnar, glabrous, 10- to 15-ribbed, crowned or denticulate at the summit. Williams, E. W. 1957. The genus Malacoth- rix (Compositae). Amer. Midi. Natural- ist 58:494-512. 1. Leaves merely dentate, elliptic to oblong or lanceolate, the cauline ones clasp- ing basally; involucral bracts orbicular to ovate and with broad scarious mar- gins M. coulteri — Leaves pinnatifid or incised to pinnately lobed, the cauline ones not especially clasping; involucral bracts linear to narrowly lanceolate 2 2(1). Leaves linear-filiform or pinnately dissected into linear segments M. glahrata — Leaves with triangular to oblong lobes or teeth, these sometimes attenuate but not linear 3 3(2). Involucres longer than broad (when pressed); persistent pappus setae 1 or 2; stems decidedly tapering upward; plants rare, in Washington County . M. clevelandii — Involucres broader than long (when pressed); persistent pappus setae 1-5 or lacking; stems various 4 4(3). Leaves with lateral lobes regularly toothed; involucres mainly less than 10 mm long; achenes 2-2.8 mm long; pappus bristles all deciduous M. sonchoides — Leaves with lateral lobes irregularly toothed or lobed; involucres more than 10 mm long; achenes 3-4 mm long; pappus often with 1 or few persistent bristles M. torreyi Malacothrix clevelandii Gray Cleveland Malacothrix. Annual herbs; stems mainly 10-40 cm tall, often branched, glabrous, commonly reddish; leaves basal and cauline, 1-10 cm long, 5-15 mm wide, oblanceolate to elliptic, pinnately lobed or merely toothed; heads few to many in a sub- corymbose cluster; involucres campanulate, 6-7 mm high; main involucral bracts linear, glabrous, green, the tips often purple, the margins narrowly scarious; rays yellow, ca 2-3 mm long; pappus deciduous or with 1 or 2 persistent bristles; achenes ca 2 mm long, slender, striate. Pinyon-juniper and live oak communities at ca 1375 m in Washington County; California, Nevada, and Arizona; 2(0). Malacothrix coulteri Harv. & Gray in Gray Snakeshead Malacothrix. Annual (win- ter annual) herbs; stems mainly 10-50 cm tall, often branched, glabrous and straw col- ored to whitish tan; leaves basal and cauline 1.2-10 cm long, oblong to oblanceolate or lanceolate, the cauline ones clasping basally; heads few to several, corymbose; involucres hemispherical, 10-15 mm high; bracts imbri- cate, suborbicular to ovate, with broad scarious margins, the midline broad, purplish; rays yellow to off-white, 5-18 mm long; pap- pus with 1-4 persistent bristles; achenes April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 319 2-2.8 mm long, striate. Warm desert shrub community at ca 950 m in Washington County (Galway sn BRY); Arizona and Cali- fornia; 2 (0). Malacothrix glabrata (D.C. Eaton) Gray [M. califomica var. glabrata D.C. Eaton]. An- nual (winter-armual) or biennial herbs; stems mainly 10-60 cm tall, often branched from the base and above, glabrous; leaves basal and cauline, 0.5-15 cm long, pinnately lobed, glabrous or more or less villous, with rachis and lobes linear to linear-filiform, the cauline ones similar to the basal except reduced up- ward; head solitary or more commonly few to many and subcorymbosely arranged; in- volucres broadly campanulate 10-14 mm high, the main bracts linear to narrowly ob- long, with narrow hyaline margins, glabrous, the outer bracts commonly more or less vil- lous; rays yellow, 10-20 mm long; pappus with usually 2 persistent bristles; achenes 2-3 mm long, striate. Joshua tree, blackbrush, Vanclevea-ephedra, and pinyon-juniper com- munities at 700 to 1525 m in Kane, Millard, San Juan, and Washington counties; Oregon to Idaho, south to California and Arizona; 22 (0). Malacothrix sonchoides (Nutt.) T. & G. [Leptoseris sonchoides Nutt.]. Annual or win- ter aimual herbs; stems mainly 6-37 cm tall, often branched from the base and above, glabrous or with short yellowish glandular hairs in the inflorescence; leaves basal and cauline, 0.7-12 cm long, 1-28 mm wide, the basal ones at least pinnatifid and the lobes regularly toothed; heads solitary or more commonly few to many and subcorymbosely arranged; involucres campanulate 7.5-10.2 mm high, 6.5-12 (14) mm wide, the main bracts lance-oblong to linear, with narrowly hyaline margins, glabrous, the outer some- times with yellowish stipitate glands; rays yellow, 7-12 mm long; pappus bristles all de- ciduous; achenes 2-2.8 mm long, striate. Blackbrush, krameria-psorothamnus, mixed desert shrub, sagebrush, and pinyon-juniper communities at 915 to 1856 m in Beaver, Duchesne, Emery, Garfield, Grand, Juab, Kane, Millard, San Juan, Tooele, Uintah, Washington, and Wayne counties; California -and Nevada, east to Nebraska and New Mexi- co; 72 (vi). Malacothrix torreyi Gray [M. sonchoides var. torreyi (Gray) Williams]. Annual or win- ter annual herbs; stems mainly 8-29 cm tall, often branched from the base and above, glabrous, or with yellowish stipitate glands in the inflorescence; leaves basal and cauline, 1.7-9.5 cm long, 5-27 mm wide, the basal ones at least pinnatifid, and the lobes irregu- larly toothed or lobed, often more or less white villous; heads solitary, or more com- monly few to several or many and sub- corymbosely arranged; involucres broadly campanulate, 10.5-15 mm high, 12-21 mm wide, the main bracts lance-linear, with hya- line margins, glabrous or some with stipitate yellowish glands, the outer bracts often stipi- tate-glandular; rays yellow, 10-14 mm long; pappus all deciduous or with 1-5 persistent setae; achenes 3-4 mm long, striate. Shad- scale, greasewood, other salt desert shrub, and mixed desert shrub communities at 1460 to 1925 m in Beaver, Box Elder, Carbon, Du- chesne, Emery, Garfield, Grand, Juab, Mil- lard, Piute, Salt Lake, Sevier, Tooele (type from Great Salt Lake), and Uintah counties; Oregon to Wyoming, south to California and Arizona; 28 (i). Matricaria L. Biennial or perennial herbs; leaves alter- nate, 2- to 3- pinnatisect, with ultimate seg- ments linear- filiform; heads radiate, few to many in corymbose clusters; involucres broadly campanulate, the bracts in several series, the margins scarious; receptacle hemi- spheric, solid, naked; rays pistillate, white; disk flowers 5-lobed, perfect, yellow; pappus a small crown; achenes laterally compressed, with 3 smooth ribs on the ventral surface and 1 or 2 (rarely more) resin glands at the apex of the dorsal face. Note: Tentatively I have chosen to follow authors of Flora Europaea (Vol. 4) in segregating Chamomilla (q.v.) from Matricaria. The genera are much alike and are separated mainly on technical char- acteristics that are discernible only when fruit is mature. Matricaria maritima L. Biennial or, less commonly, perennial, essentially unscented herbs; stems 1-6 dm tall, glabrous or nearly so; leaves 1-8 cm long, the ultimate segments long and slender; heads several to many, the 320 Great Basin Naturalist Vol. 43, No. 2 disk 8-15 mm wide; rays 10-25, white, 6-13 ternate or all basal, entire or pinnatifid; mm long. Ruderal weed of moist sites at 1830 heads many flowered, erect or nodding in to 2135 m in Salt Lake, Sanpete, and Sevier bud; involucres cylindric to campanulate, the counties; widespread in North America; ad- innermost bracts lance-attenuate, subequal, ventive from Europe; 2 (i). the outer ones shorter and imbricate; recep- tacle naked; corollas all raylike, showy, yel- MiCROSERis D. Don 1«7 t° yellow-orange (fading bluish); pappus or awn-tipped scales or or plumose capillary Annual or perennial, scapose or caulescent bristles; achenes columnar to fusiform, not or taprooted herbs with milky juice; leaves al- only short beaked, ca 10-ribbed. 1. Plants annual; pappus of 5 scales, entended into scabrous bristles apically M. lindleyi — Plants perennial; pappus of numerous plumose capillary bristles arising from short scales M. nutans Microseris lindleyi (DC.) Gray [Calais lindleyi DC.; Microseris linearifolia (Nutt.) Schultz-Bip; Uropappus linearifolius Nutt.]. Armual herbs from slender taproots; herbage puberulent or glabrate; stems lacking or more or less developed, the scapose pe- duncles 10-25 cm high; leaves 6-15 (30) cm long, pinnately lobed to entire, linear to nar- rowly elliptic; heads many flowered, erect, the main bracts lance-attenuate, 15-30 mm long, subequal, the outer ones shorter and un- equal; rays yellow (drying blue); pappus 10-20 mm long, silvery, deciduous, of 5 lance-linear scales, each terminating in a scabrous awn from a bifid apex; achenes dark brown, 9-13 mm long, tapering apically, scabrous on the ribs. Blackbrush, creosote bush, and pinyon-juniper communities at 915 to 1375 m in Washington County; Washing- ton to Idaho, south to Baja California and Arizona; 5 (i). Microseris nutans (Geyer) Schultz-Bip [Scorzonella nutans Geyer]. Perennial herbs from tuberous roots; herbage glabrous or sparsely scurfy; stems more or less developed, the scapose peduncles mainly 12-40 (60) cm high, pinnately lobed to entire, linear to el- liptic, lanceolate, or oblanceolate; heads soli- tary or 2-5, many flowered, nodding in bud, the main bracts 10-20 mm long, lance-atten- uate, subequal, the outer bracts shorter and unequal; rays yellow (drying lavender or blue); pappus of numerous narrow scales, each with a plumose terminal bristle. Sage- brush, pinyon-juniper, mountain brush, Douglas fir, and aspen communities at 1675 to 2745 m in Box Elder, Cache, Daggett, Davis, Juab, Millard, Rich, Salt Lake, San- pete, Sevier, Summit, Uintah, Utah, and We- ber counties; British Columbia to Montana, south to California, Nevada, and Colorado; 33 (ii). MoNOPTiLON T. & G. ex Gray Annual herbs, branched from base, the herbage hispid; leaves alternate, spatulate, entire; heads radiate, solitary on branch tips, closely subtended by upper leaves; involucre campanulate, the bracts subequal, linear, her- baceous; receptacle flat, naked; ray flowers pistillate, white to pink; disk flowers perfect, fertile, yellow (purplish); pappus of a short scarious cup and 1 apically plumose bristle, or of numerous bristles alternating with short scales; achenes compressed, marginally nerved, pubescent. 1. Pappus of usually several nonplumose bristles alternating with scales; disk corollas sparsely if at all pilose; reported for Utah by Abrams and Ferris (Illustrated Flora of the Pacific States), but not seen by me . M. bellioides (Gray) Hall — Pappus consisting of minute scales and a single apically plumose bristle; disk corollas densely pilose below M. bellidiforme Monoptilon bellidiforme T. & G. in Gray Depressed annual branching herbs; stems 1-5 cm high; leaves 4-10 mm long, 0.5-2.5 mm wide, narrowly oblanceolate; heads showy; April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 321 involucres 4-5 mm high; bracts linear, atten- uate or acuminate, hirsute, and minutely glandular; rays 12-20, ca 4-5 mm long, the tube pilose; pappus of 1 apically plumose bristle and several shorter scales, or the pap- pus rarely lacking; achenes ca 2 mm long. Warm desert shrub at 700 to 900 m in Wash- ington County; California, Nevada, and Ari- zona; 2 (0). Onopabdum L. Biennial caulescent spiny herbs from tapr- oots, the juice watery; leaves basal and caul- ine, alternate, winged-decurrent; heads soli- tary or few to several; involucral bracts in several series, imbricate, spine tipped; recep- tacle flat, fleshy, honeycombed, often with short bristles on the partitions, not densely bristly; corollas all discoid, reddish purple or pink, perfect; pappus bristles barbellate; ach- enes glabrous, subquadrangular, 4- or 5- ribbed. Onopardum acanthium L. Biennial herbs; stems mainly 5-15 (30) dm tall; leaves of bas- al rosettes 5-50 cm long or more and 2-15 cm wide, pinnately lobed and serrate-den- tate, tomentose on both surfaces, but less so above, spinose; cauline leaves pinnatifid, to- mentose to glabrate, strongly winged-decur- rent along the stem length; involucres 25-35 mm high, 30-65 mm wide, the bracts lance- attenuate, with spreading spine tips, tomen- tose to glabrate marginally, the inner erect; spines 3-5 mm long, yellowish; corollas red- dish purple to pink. Ruderal weeds at low elevations in Millard, Tooele, Utah, Wasatch, and Washington counties; adventive from Europe; 6 (ii). This handsome but trouble- some thistle is spreading through the state, but less vigorously than the musk thistle, Car- duus nutans (q.v.). tips; pappus lacking; achenes obovoid, dense- ly villous, 1 -ridged on each face. Oxytenia acerosa Nutt. Copperweed. Pe- rennial herbs; stems erect, mainly 5-12 dm tall, broomlike in the inflorescence, striate; leaves 3-15 cm long, pinnately 3- to 7-lobed, or the upper ones simple; herbage strigulose; heads 3-4 mm wide, erect or ascending; in- volucral bracts herbaceous, strigulose; ach- enes 1.5-2 mm long, black, long villous-pi- lose. Saline riparian areas and near seeps and springs at 1220 to 2135 m in Carbon, Emery, Garfield, Grand, Kane, San Juan, and Wash- ington counties; Colorado, New Mexico, Ari- zona, Nevada, and California; 25 (vii). Cop- perweed is poisonous to livestock. Palafoxia Lag. Annual herbs; leaves alternate, entire; heads discoid, few to several, corymbose or paniculate; involucres cylindric; bracts in 1 series, herbaceous; receptacle flat, naked; flowers white, all alike or the outer with im- equal lobes; pappus scales 4-8, slender, un- equal, with a strong nerve; achenes linear, quadrangular. Palafoxia linearis (Cav.) Lag. Spanish Needle. [Ageratum lineare Cav.]. Annual herbs; herbage hispid with slender multi- cellular hairs, glandular upward; stems com- monly branched above the base, 1-7 dm tall; leaves petiolate, the blades 1-7.5 cm long, 2-8 mm wide, linear-lanceolate, long-attenu- ate; involucres 12-18 mm high, glandular, and more or less hispid, 10- to 20-flowered, the corollas white with pink exserted styles; pappus scales usually 4; achenes strigose. Warm desert shrub community at 700 to 1000 m in Washington County; California to Arizona and Mexico; 5 (0). Oxytenia Nutt. Perennial riparian herbs from a ligneus caudex; leaves alternate, pinnately divided or some entire, the segments linear-filiform, in- volute; heads discoid, numerous, in elongate paniculate inflorescences; marginal flowers 5, pistillate, inner flowers 10-30, staminate; flowers yellowish white; involucral bracts 5, orbicular, mucronate; receptacle chaffy, the chaffy bracts slender, with dilated villous Parthenium L. Herbs or shrubs; leaves alternate, entire or lobed; heads solitary or few and more or less clustered, inconspicuously radiate; ray flow- ers 5, white, pistillate, fertile, persistent; disk flowers staminate; receptacle plano-convex, chaffy throughout; pappus of 2 or 3 awns or scales; ray achenes dorsiventrally com- pressed, rotund in outline, the margins thick- ened into riblike structures attached to the 322 Great Basin Naturalist Vol. 43, No. 2 contiguous pair of infertile disk flowers and the subtending bract, the achene, the 2 at- tached flowers, and the bract falling as a unit. Rollins, R. C. 1950. The guayule rubber plant and its relatives. Contr. Gray. Herb. 179: 1-73. Plants shrubs, the internodes apparent; heads seldom solitary; known from Washington County P. incanum Plants pulvinate-caespitose herbs, the internodes not apparent; heads solitary; plants of eastern Utah P. ligulatum Parthenium incanum H.B.K. Aromatic shrub, 4-10 dm tall, much branched, the branchlets loosely tomentose, becoming gla- brate; leaves short-petioled, the blades 0.5-5 cm long, 0.4-1.5 cm wide, lobed, white-to- mentose below, less so above; heads several to many, corymbose, 3-5 mm wide, outer in- volucral bracts oblong, acute, villous, the in- ner ones suborbicular, membranous; rays white, emarginate to incised, 1-2 mm long; pappus of 2 or more pubescent awns; achenes black, oblanceolate, 1.5-2 mm long, pu- bescent on the ventral surface. Limestone cliffs in creosote bush-blackbrush community at ca 1220 m in Washington County (Higgins 4102 BRY); Arizona to Texas, south to Mexi- co; 1 (0). Parthenium ligulatum (Jones) Bameby [P. alpinum var. ligulatum Jones, type from The- odore (Duchesne); Bolophyta ligulata (Jones) W.A. Weber]. Pulvinate caespitose to merely caespitose acaulescent mound- forming herbs to ca 3 cm high, from a taproot and branched caudex, the caudex branches densely clothed with brownish marcescent leaf bases and of- ten with ashy leaves of the previous year; leaves 3-20 mm long, 1.5-4 mm wide, spatu- late to oblanceolate, strigose; heads solitary at branch ends, sessile, 5-7 mm high, 4.5-6 mm wide; outer bracts oblong, densely pu- bescent apically; pappus scales distinct or ad- nate to the corolla tube; rays white, 1-2 mm long, emarginate; achenes oblanceolate, densely hairy, 4-5 mm long, 2-3 mm wide. Barren or semibarren calciferous or gypsi- ferous outcrops of the Green River, Uinta, Ferron, and Carmel formations in salt desert shrub and pinyon-juniper communities at 1705 to 2135 m in Daggett, Duchesne, Emery, and Uintah counties; Colorado (a Colorado Plateau endemic); 42 (iv). This amazing plant is one of a series of edaphi- cally restricted mound-formers in semibarren habits on shales and clays of arid sites in Utah. It belongs to a closely related assem- blage of two or three taxa within section Bo- lophytum, and has been regarded at specific status within the segregate genus Bolophyta. Its phylogenetic position was reviewed by Rollins (1950), and its status within Parthe- nium, seems to represent best its generic affinities. Pectis L. Annual herbs; leaves opposite, entire, glandular-dotted; heads radiate, few to sever- al in cymose clusters; involucres turbinate to subcylindric; bracts 3-12 in one series, ex- panded basally, enclosing the ray flowers, of- ten with translucent glands; receptacle naked; ray flowers perfect, yellow; disk flow- ers few; anthers entire, obtuse at base; style branches short, hispidulous; pappus of short- plumose bristles on disk flowers, that of ray flowers a short crown of united scales; ach- enes terete. Pectis papposa Harv. & Gray in Gray Chinch-weed. Annual herbs; stems dichoto- mously branched, often decumbent-spread- ing, 5-20 (25) cm long, the herbage yellowish green; leaves 6-40 (60) mm long, 0.5-2 mm wide, with a few setae at the base, glabrous, bearing oval to elliptic large yellowish glands; heads on peduncles mainly 0.3-1 (2) cm long; involucres gibbous at the base, rounded dorsally, sparingly glandular like the leaves; ray flowers yellow, 7-9, ca 4-6 mm long; achenes 4-5 mm long, stipitate-glandu- lar. Sandy soil in warm and sandy desert shrub communities at 850 to 1650 m in Kane, San Juan, and Washington counties; Califor- nia to New Mexico, and south to Mexico; 9 (i)- Perezia Lag. Perennial herbs from a caudex, this clothed with rusty woolly hairs; leaves alternate, simple, clasping; heads numerous in corym- bose cymes, apparently discoid; involucres April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 323 campanulate, strongly imbricate; flowers per- fect, pink to pink-purple, the corollas bila- biate, the outer lip 3-toothed, the inner lip recurved, 2-toothed; anthers appendaged; style branches flattened, truncate apically; pappus of white capillary bristles; achenes subterete, minutely glandular. Perezia wrightii Gray [Acourtia wrightii (Gray) Reveal & King]. Perennial herbs; stems 4-6 (10) dm tall, often purplish at the base, the rusty hairs at stem base copious; leaves lance-oblong to ovate or lanceolate, spinulose-dentate, glandular-puberulent on both sides, the lower ones petiolate, becom- ing sessile and clasping upward; involucres 5-8 mm high and about as broad, the bracts graduated, the outer ones ovate, the inner lance-oblong, obtuse, green, the margins of- ten purplish, ciliate; corollas pink to pink- purple; achenes 4.8-5.2 mm long. Warm desert shrub and juniper communities at 915 to 1525 m in Kane, San Juan, and Washing- ton counties; Arizona to Texas, south to Mex- ico; 6 (i). Perityle Benth. Annual herbs or perennial subshrubs; leaves mostly opposite below, alternate above, simple, petiolate; heads few to numer- ous, corymbose, radiate or discoid; involucres hemispheric or turbinate, the bracts some- what keeled, in 1 or 2 subequal series; recep- tacle flat, naked; ray flowers (when present) pistillate, white or yellow; disk flowers per- fect; anthers subentire to auriculate at the base; style branches linear or subulate; ach- enes flattened; pappus of scales, or of 1 or 2 awnlike bristles, or lacking. Powell, A. M. 1973. Taxonomy of Perityle section Laphamia (Compositae-Helen- ieae-Peritylinae). Sida 5: 61-128. 1974. Taxonomy of Perityle section Perityle (Compositae-Peritylinae). Rho- dora 76: 229-305. 1. Plants annual P. emoryi — Plants subshrubs 2 2(1). Heads radiate; plants glandular-hispidulous, of the Great Basin P. stansburyi — Heads discoid; plants villous or glandular-hispidulous, of the Colorado or Vir- gin drainages 3 3(2). Herbage short-villous and more or less glandular; pappus bristles 1; plants of Washington County P. tenella — Herbage hispidulous; pappus of 3 (4) unequal bristles; plants of Grand County P. specuicola Perityle emoryi Torr. in Emory Emory Rock-daisy. Annual herbs, mainly 2-5 dm tall; stems erect or spreading, commonly branched above, puberulent; leaves mostly alternate, petiolate, the bases 0.5-4 cm long, 0.6-3 (5) cm wide, ovate, cordate, or sub- orbicular, toothed, lobed, cleft, or divided, the lobes again toothed or lobed, hirsute to glandular-pubescent; heads radiate; in- volucres 5-6 mm high and usually broader; rays 8-12, white, 1.5-5 mm long; disk flowers numerous; pappus vestigial or a crown of scales and 1 slender bristle; achenes 2-3 mm long, the flattened faces nearly glabrous, the margin thickened and bearing short stiff hairs. Sand sagebrush community at lower elevations in Washington County (Tanner sn 1941 BRY); Nevada, California, Arizona, and Mexico; 1 (0). Perityle specuicola Welsh & Neese Alcove Rock-daisy. Perennial suffruticose herbs, mainly 50-75 cm tall; stems sprawling or pendulous, much branched; herbage glandu- lar-hispidulose; leaves mostly alternate, short- petiolate, the blades 3-6 mm long, 1.5-3 mm wide, ovate-elliptic, entire, hispidulous; heads few to many in a branching corymbose in- florescence; involucres 3.5-5 mm high, 5-6 mm wide; bracts 11-16, oblong to elliptic, keeled; ray flowers lacking; disk flowers nu- merous, ca 2.5 mm long, whitish (?); pappus of 3 unequal scabrous bristles and often with 1 apically flattened and sigmoid scale; ach- enes 3-3.8 mm long, the faces flattened, glabrous, the margin thickened and with short ascending hairs. Hanging garden com- munities at ca 1220 m in Grand County; en- demic; 2 (ii). 324 Great Basin Naturalist Vol. 43, No. 2 Perityle stansburyi (Gray) Macbride Stansbury Rock-daisy. [Laphamia stansburyi Gray, type from Stansbury Island]. Suffruti- cose perennials, clump-forming, 7-30 cm tall and as broad or more; herbage glandular-hir- tellous; leaves mainly alternate, the blades 3-14 mm long, 1.5-12 mm wide, broadly ovate to deltoid or orbicular, typically few to several lobed; petioles 1-14 mm long; heads few to many in a branching corymbose in- florescence; involucres 5-6.5 mm high, 5-10 mm wide; bracts 16-21, lanceolate to oblan- ceolate, strongly keeled; ray flowers 10-14, yellow, 3-5.5 mm long; disk flowers numer- ous, yellow, 4-5 mm long; pappus of 1 stout bristle and a very short crown of hyaline scales; achenes 2.3-3.5 mm long, with thin callous margins, short-pubescent on margins and on faces. Limestone, dolomite, and ig- neous ignimbrite (ashflow tuff) outcrops, in mixed desert shrub, pinyon-juniper, and mountain brush communities, at 1280 to 1895 m in Beaver, Juab, Millard, Salt Lake, Sanpete, Sevier, and Juab counties; Nevada (a Great Basin endemic); 39 (v). Perityle tenella (Jones) Macbride Jones Rock-daisy. [Laphamia palmeri Gray, type from Beaverdam, Arizona?, not P. palmeri Wats.; L. palmeri var. tenella Jones, type from Springdale]. Suffruticose perennials, clump-forming, 9-25 cm tall and as broad or more; herbage villous and glandular; leaves mainly alternate, the blades 4-13 mm long, 3-15 mm wide, deltoid-ovate, the base obtuse to truncate or cordate; petioles 1-8 mm long; heads solitary or few to many, corymbose; in- volucres 4-6.5 mm long, 5-10 mm wide; bracts 11-18, lance-elliptic, keeled; ray flow- ers absent; disk flowers numerous, yellow, 3-4 mm long; pappus of a single bristle; ach- enes 2.5-3 mm long, with thin callous mar- gins, short-pubescent on margins and on faces. Joshua tree, creosote bush, blackbrush, warm desert shrub, pinyon-juniper, and pon- derosa pine communities at 915 to 2135 m in Washington County; Arizona; a Mohave en- demic; 7 (0). Plants from the Beaverdam Moimtains have heads that average larger, but they seem not to differ otherwise from the typical materials taken near Zion Nation- al Park. Petradoria Greene Suffrutescent perennials from a taproot and woody caudex; stems herbaceous, leafy; leaves basal and cauline, alternate, entire, 3- to 5-veined, coriaceous; heads radiate (in ours), congested at branch ends in an open corymbose inflorescence; involucres cylin- dric; bracts in several series, in more or less vertical ranks; flowers 4-7, yellow, the co- rollas glabrous; pappus of brownish capillary bristles; achenes somewhat compressed, glabrous. Anderson, L. C. 1964. Studies on Petradoria (Compositae); anatomy, cytology, tax- onomy. Trans. Kansas Acad. Sci. 66: 632-684. Petradoria pumila (Nutt.) Greene Rock Goldenrod. [Chrysoma pumila Nutt.]. Plants from a well-developed caudex, the caudex branches clothed with dark to ashy or tan marcescent leaf bases; leaves 1.5-12 cm long, 1-11 mm wide, oblanceolate to lanceolate, elliptic, or linear; cauline leaves reduced up- ward; heads numerous; involucres 5-9.5 mm high, 1.3-2.8 mm wide; involucral bracts 10-21, in 3-6 series, more or less keeled; flowers 2-8, the rays 1-3, yellow, 4-9 mm long; achenes 4-5 mm long, glabrous, 6- to 9- nerved. Shadscale, mixed desert shrub, pin- yon-juniper, sagebrush, and ponderosa pine communities at 1525 to 3050 m in all (?) Utah counties; Idaho and Wyoming, south to Nevada, California, Arizona, and New Mexi- co; 100 (xv). Most of our specimens belong to the broad-leaved var. pumila, but one speci- men from Emery County (Harris 546 BRY) seems to be clearly allied to var. graminea (Woot. & Standi.) Welsh comb nov. [based on: Petradoria graminea Woot. & Standi. Contr. U.S. Natl. Herb. 16: 183. 1913; ssp. graminea (Woot. & Standi.) L.C. Anderson]. That taxon has been known previously only from Arizona. Platyschkuhria (Gray) Rydb. Perennial herbs from a woody caudex and rootstock; leaves alternate, simple, co- riaceous, often impressed-punctate; heads few to many in a cymose paniculate cluster, radiate, campanulate to hemispheric; in- volucral bracts subequal in 2 series; recep- tacle essentially flat, naked; rays pistillate. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 325 fertile, yellow; disk flowers numerous, per- fect; anthers more or less sagittate basally; pappus of 8-16 scales with midribs some- times produced apically; achenes narrowly obpyramidal and 4-sided, hairy or glabrous on the sides. Ellison, W. L. 1971. Taxonomy of Platysch- kuhria (Compositae). Brittonia 23: 269-279. Platyschkuhria integrifolia (Gray) Rydb. [Schkuhria integrifolia Gray; Bahia nudi- caulis Gray; B. integrifolia (Gray) Macbride]. Perennial herbs; stems solitary or few to sev- eral, mainly 12-55 cm tall; herbage white- strigulose or stipitate-glandular, especially above; main leaves near the stem base, pe- tiolate, the blades 1.5-9.5 cm long, 0.5-4 cm wide, ovate to lanceolate, elliptic, or oblan- ceolate; cauline leaves reduced upward, fi- nally merely bracteate; heads (1) 2-10; rays 7-11, yellow, 6-14 mm long; achenes 5-8 mm long. Three rather distinctive varieties are present in eastern Utah, as indicated below. 1. Stems leafy almost or quite to the apex; plants of San Juan County P. integrifolia var. oblongifolia — Stems leafy mainly below the middle; plants not known from San Juan County 2 2(1). Pubescence of upper stems merely white-strigulose; involucral bracts caudate- attenuate P. integrifolia var. ourolepis — Pubescence of upper stems stipitate-glandular; involucral bracts mainly obtuse to acute P. integrifolia var. desertorum Var. desertorum (Jones) Ellison [Bahia desertorum Jones, type from Cisco]. Salt desert shrub, pinyon-juniper, and mountain brush communities, mainly in saline sub- strates, at 1280 to 2565 m in Carbon, Du- chesne, Emery, Garfield, Grand, Sevier, Uin- tah, and Wayne counties; Colorado; a Colorado Plateau endemic. A report by Elli- son (1971) of var. integrifolia (a Wyoming- Montana endemic) belongs here; 39 (xi). The var. desertorum is closely allied with var. in- tegrifolia, as indicated by pubescence and bract shape similarities. This variety is transi- tional with var. ourolepis. Var. oblongifolia (Gray) Ellison [Schkuhria integrifolia var. oblongifolia Gray; Bahia oblongifolia (Gray) Gray; Platysch- kuhria oblongifolia (Gray) Rydb.]. Desert shrub communities in San Juan County; Ari- zona, Colorado, New Mexico; 0 (0). The vari- ety is reported from San Juan County, but no specimens have been seen by me. Var. ourolepis (Blake) Ellison [Bahia ouro- lepis Blake, type from Green River]. Salt desert shrub and pinyon-juniper communities at 1280 to 1830 m in Duchesne, Emery, Grand, and Uintah counties; endemic; 24 (iii). The main body of this variety lies in Uintah County. Pluchea Cass. Shrubs; leaves alternate, simple, entire, se- riceus; heads discoid, few to several, aggre- gated in terminal cymose clusters; involucres campanulate; bracts imbricate in several series, scarious, the outer ones sericeus; re- ceptacle flat or concave, naked; outer flowers pistillate, numerous, their filiform corollas 3- or 4-toothed; central flowers perfect but the innermost sterile, their tubular corollas 5- toothed; anthers sagittate basally; pappus of outer flowers merely capillary bristles, those of inner flowers clavate apically. Pluchea sericea (Nutt.) Gov. Arrowweed. [Polypappus sericeus Nutt.; Tessaria sericea (Nutt.) Shinners]. Shrubs with slender, erect, willowlike branches, mainly 0.8-3 m tall, se- riceus throughout, longitudinally striate; leaves 0.8-4.5 cm long, 2-9 mm wide, ellip- tic to narrowly lanceolate or lanceolate, en- tire, sessile; heads more or less conspicuous; involucres 3.5-5 mm high, 4-7 mm wide; outer bracts ovate to ovate-lanceolate, abruptly acute, deciduous, often purplish; pistillate flowers purplish, numerous; perfect flowers purplish, fewer; achenes glabrous; pappus bristles of perfect flowers dilated api- cally. Riparian areas at 460 to 1220 m in Garfield, Kane, San Juan (?), and Washington 326 Great Basin Naturalist Vol. 43, No. 2 counties; California to Texas, south to Mexi- co; 22 (iii). The genus Pluchea, in a broad sense, includes annual and perennial herbs and shrubs. Tessaria, when segregated from Pluchea, consists of the shrubby species that have dimorphic corollas and the inner per- fect flowers with apically flared pappus bristles. The residue within Pluchea contains only herbs with uniformly 4-lobed corollas and pappus of uniform barbellate capillary bristles. I follow tradition in maintaining our species in Pluchea. PoROPHYLLUM (VaiU.) Adans. Suffruticose perennial; leaves alternate or opposite, simple, with at least some elliptic to oval oil glands; heads discoid, solitary, or few to several in corymbose clusters; in- volucres cylindric, the bracts usually 5, in subequal series, glandular like the leaves; re- ceptacle naked; flowers perfect, fertile, purplish; anthers rounded basally; style branches slender, hirtellous, subulate; pappus of scabrous bristles; achenes slender, striate. Porophyllum gracile Benth. Odora. Rounded bushy perennials from a woody base; stems much branched, 1.5-4 dm tall; herbage dark green or often purplish, glaucous, odoriferous; leaves 1-4 cm long, linear-filiform, entire; involucre subcylindric, 10-15 mm long; bracts 5, dark green, tinged purplish, oblong, the hyaline margin pink, gibbous basally, bearing conspicuous glands, especially apically; corollas purplish, white; pappus bristles pinkish; achenes 8-9 mm long, hispidulous. Desert shrub communities in Washington County (Cottam 5522 UT); California to Arizona and Mexico; 1 (0). Prenanthella Rydb. Annual herbs; leaves basal and alternate, simple, pinnately lobed, toothed, or pinnati- fid; heads small, few to numerous; involucres campanulate; bracts in 2 series, the inner sub- equal, 3 or 5, the outer much reduced, 1 or 2, herbaceous; flowers all raylike, 4 or 5; ach- enes 5-ribbed; pappus of white capillary bristles. Prenanthella exigua (Gray) Rydb. [Pre- nanthes exigua Gray; Lygodesmia exigua (Gray) Gray]. Annual; stems branched from the base, forming rounded clumps, 7-24 (30) cm tall; inflorescence paniculate, comprising more than half the plant height; lower leaves 1-4 (6.5) cm long, 3-12 (20) mm wide, spatu- late to oblanceolate, the rosette often with- ered at anthesis; cauline leaves reduced up- ward, finally bracteate scales; herbage sparingly stipitate-glandular; involucres 3-5.5 mm long, 1.2-3.5 mm wide; inner bracts ob- long, herbaceous, apically constricted in bud; rays pink or white, 1.5-2 mm long; achenes 3-3.5 mm long, 5-ridged, scabrous; pappus of white capillary bristles. Blackbrush, creosote bush, other warm desert shrub, salt desert shrub, and pinyon-juniper communities at 850 to 1925 m in Beaver, Carbon, Emery, Garfield, Grand, Juab, Kane, Millard, San Juan, Tooele, Uintah, and Washington coun- ties; California, Nevada, Colorado, Arizona, and New Mexico; 20 (ii). Psathyrotes Gray Annual or perennial (?) herbs; leaves alter- nate, petiolate, simple, entire or lobed to toothed; heads discoid, the flowers yellow or purplish in age; involucres campanulate; bracts biseriate, the outer often shorter or otherwise different; receptacle flat, naked; anthers minutely sagittate; style branches flattened; achenes hairy; pappus of capillary bristles. 2(1). Plants lanate-tomentose as well as scurfy; outer involucral bracts expanded api- cally, oblong-obovate; reported for Utah by Munz (A California Flora), but not seen by me P. ramosissima (Torr.) Gray Plants scurfy and less commonly somewhat tomentose; outer involucral bracts tapering apically, lanceolate 2 Leaves entire; herbage scurfy and with long-piliferous multicellular hairs P. pilifera Leaves toothed; herbage scurfy but not long-piliferous P. annua Psathyrotes annua (Nutt.) Gray Mealy Ro- settes [Bulbostylis annua Nutt.]. Annual or winter annual herbs forming low rounded cushions, mainly 2-18 cm tall; leaves petio- April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 327 late, the blades 5-17 mm long, 5-20 mm wide, orbicular to fan shaped, dentate; her- bage scurfy; heads few to numerous, corym- bose; involucres 5.5-7.5 mm high, 5-8 mm wide; outer bracts lanceolate to oblong, more or less constricted above the middle, scurfy and ciliate; disk corollas 3.5-4.2 mm long, yellow, becoming purplish; achenes 2-2.5 mm long, pilose. Warm desert shrub, salt desert shrub, and pinyon-juniper commu- nities, commonly on limestone and dolomitic gravels, at 790 to 1740 m in Juab, Millard, Tooele, and Washington counties; Idaho south to California, Nevada, and Arizona; 17 IV Psathyrotes pilifera Gray Annual or win- ter annual herbs forming hemispheric cush- ions, mainly 5-15 cm tall; leaves petiolate, the blades 5-15 mm long, 4-16 mm wide, obovate, ovate, or oval-elliptic, entire; herb- age scurfy and piliferous with long multi- cellular hairs; heads few to many, corymbose; involucres 8.5-10 mm high, 4-5.5 mm wide; outer bracts lanceolate, seldom constricted above the middle, scurfy and with long pili- ferous setae marginally; disk corollas 6-6.5 mm long, yellow, becoming purplish; achenes 3.8-4.8 mm long. Warm desert shrub and salt desert shrub, commonly on gypsiferous sub- strates of the Moenkopi and Chinle forma- tions, at 760 to 2260 m in Grand, Kane, and Washington counties; Arizona; endemic; 9 PSILOCARPUS Nutt. Low floccose-wooUy armual herbs; leaves opposite, simple, entire; heads discoid, sub- globose; involucre per se essentially lacking; receptacle chaffy, subglobose; pistillate flow- ers numerous, imbricate, each enclosed by and deciduous with its subtending bract, woolly, with sides meeting in the center, bearing below the rounded tip on inner side a scarious appendage; corollas filiform; pappus lacking; perfect flowers few, central, ebrac- teate, the corollas 4- or 5-toothed, epappose; anthers sagittate. Psilocarphus brevissimus Nutt. Low white-woolly annuals; stems simple or with decumbent-prostrate branches mainly 1.5-20 cm long; leaves 5-15 mm long, 1-3 mm wide, spatulate to lanceolate, apiculate; heads solitary or clustered, long-woolly, ca 5-7 mm thick, subtending leaves about as long as the head or longer; pistillate flowers 20-34 or more, the enclosing bracts 2.5-3.2 mm long, woolly, the appendage horizontally produced to erect, ca 0.5 mm long; perfect flowers ca 6-10; achenes subcylindric, terete, 1.3-2 mm long. Lake and reservoir beds at ca 1710 m in Cache and Salt Lake counties; Washington to Montana, south to California, Mexico, and South America; 6 (0). PSILOSTROPHE DC. Perennial herbs or shrubs; leaves alternate, simple, entire or merely lobed; heads few to many, corymbose; involucres campanulate; bracts in 1 series, subequal; receptacle naked; ray flowers yellow, pistillate, becoming pa- pery and persistent; disk flowers perfect, 5- lobed; anthers obtuse basally; style branches truncate; pappus of 4-6 hyaline scales; achenes obtusely angled, glabrous or hairy. 1. Plants shrubby; stems closely white-tomentose; of Washington County P. cooperi — Plants herbaceous, from a definite caudex; stems glabrous or loosely tomentose; not of Washington County 2 2(1). Stems loosely tomentose; involucres densely white villous-tomentose; plants of Grand County P. bakeri — Stems glabrous, or tomentose only at the base; involucres sparingly tomentose, greenish; plants of Wayne, Garfield, and Kane counties P. sparsiflora Psilostrophe bakeri Greene Perennial herbs from a caudex; stems 10-35 cm tall, densely white-woolly below, loosely tomen- tose upward; leaves 0.8-10 cm long, 2-15 mm wide, spatulate to oblanceolate, entire or lobed, pubescent like the stems; cauline leaves reduced upward; involucres loosely villous-tomentose, 5-9 mm high, ca 3-4 mm 328 Great Basin Naturalist Vol. 43, No. 2 wide; rays 3-6, yellow, 6-12 (15) mm long; pappus scales ca 2 mm long, rounded; achenes whitish, ca 2-2.5 mm long. Sandy warm desert shrub community at ca 1285 m in Grand County (Trotter 101 BRY); Colo- rado (a Plateau endemic?); 1 (0). Psilostrophe cooperi (Gray) Greene Paper- flower. [Riddellia cooperi Gray]. Shrubs; stems closely white-tomentose, mainly 30-60 cm tall; leaves 1.2-7 cm long, linear, entire, sparingly tomentose, finally glabrate; in- volucres tomentose, 6-8 mm high, 5-8 mm wide; rays 4-8, yellow, 8-20 mm long; pap- pus scales ca 2 mm long, acute; achenes whit- ish, 4.5-7 mm long. Joshua tree, creosote bush, blackbrush, and pinyon-juniper com- munities at 915 to 2135 m in Washington County; Nevada, California, Arizona, and New Mexico; 11 (ii). Psilostrophe sparsiflora (Gray) A. Nels. [Riddellia tagetina var. sparsiflora Gray]. Perennial herbs from a caudex; stems 14-60 cm tall, densely to moderately pilose basally. sparingly villous-tomentose upward; leaves 0.9-11.5 (14.5) cm long, 1-11 mm wide, spatulate to oblanceolate or linear, pubescent like the stems or glabrate; involucres 4.5-6 mm high, 4-6 mm wide; rays usually 3, yel- low, 6-12 mm long; pappus scales 1.5-2.5 mm long, acutish; achenes yellowish, 2.5-3 mm long. Salt desert shrub, pinyon-juniper, and sagebrush communities at 1430 to 2045 m in Garfield, Kane, and Wayne counties; Arizona, New Mexico, and Mexico; 42 (vi). Rafinesquia Nutt. Annual herbs; stems fistulous; leaves alter- nate, pinnatifid; heads 2 to several, large, showy, with white or rose-tinged flowers; in- volucres essentially cylindric; bracts 7-15, in 2 series, the inner subequal, the outer ones much shorter, obtuse or subcordate basally; flowers all raylike; pappus white or tawny, of 8-15 slender long-plumose bristles; achenes terete, obscurely few ribbed, attenuate into a beak. Rays mainly 5-8 mm long; achene beak as long as the body; plumose hairs of pappus straight; plants reported for Utah by Munz (A California Flora), but not seen by me R. californica Nutt. Rays mainly 12-18 mm long; achene beak shorter than the body; plumose hairs of pappus crinkled; plants of Washington County R. neomexicana Rafinesquia neomexicana Gray Desert Chicory. Annual (winter annual) herbs; stems mainly 15-40 (50) cm tall, simple or branched, often growing up through shrubs; basal leaves 1.2-9 cm long, pinnatifid, often withered at anthesis; cauline leaves sessile and auriculate-clasping, reduced upward; in- volucre 15-25 mm high, 5-9 mm wide; main bracts lance-attenuate, the margins scarious, the outer ones more or less cordate basally; rays 12-18 mm long, white or suffused with pink, 5-toothed or -lobed apically; pappus bristles white, the bases flattened, plumose to near the apex; achenes 12-15 mm long, pa- pillate-puberulent. Joshua tree, creosote bush, blackbrush, and desert almond commu- nities at 700 to 1070 m in Washington Coun- ty; California to Texas and Mexico; 6 (i). Ratibida Raf. Perennial herbs from a caudex and stout taproot; leaves alternate, pinnatifid; heads radiate, solitary or few and corymbose; rays neuter, commonly yellow (sometimes purple in part or throughout); involucre in 1 series, green; receptacle columnar, chaffy through- out, the bracts more or less clasping the achenes; disk flowers perfect, fertile; anthers sagittate; style branches flattened; achenes compressed at right angles to the involucral bracts, glabrous, the margins sometimes cil- iate; pappus of an evident tooth and some- times with a second one. Ratibida columnifera (Nutt.) Woot. & Standi. Prairie Coneflower. [Rudbeckia col- umnifera Nutt.; R. columnaris Pursh]. Pe- rennial herbs; stems mainly 3-6 (12) dm tall, several, often branched above, strigose; leaves 2-9 cm long, pinnatifid, with the ter- minal division often the largest; heads borne on slender peduncles 6-18 cm long, the disk grayish in bud, purplish brown in flower, 1.5-3 cm long; rays 3-7, yellow (or purple), 1-3 (4.5) cm long, spreading or reflexed; pap- pus an evident awn tooth on the inner angle April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 329 of the achene, often also a shorter one on the outer edge; achenes ciliate and more or less winged on the inner edge. Salt desert shrub and sagebrush communities at 1585 to 2565 m in Garfield, Millard, and Washington counties; British Columbia to Minnesota, south to Arizona, Texas, and Mexico; 4 (i). Our material appears to be adventive from the main body of the species in the prairies and plains provinces to the east of Utah. RiGiOPAPPUs Gray Annual herbs; leaves alternate, linear, en- tire; heads radiate, solitary or few to several, cymose; rays pistillate, fertile, yellow, incon- spicuous; involucres campanulate; bracts ob- long-alternate, subequal, partly clasping out- er achenes; receptacle flat, with a row of bristles between ray and disk flowers; disk flowers perfect, fertile, yellow; anthers not toothed; style branches flattened; pappus usually of 3-5 awnlike scales (or lacking); achenes linear, transversely rugulose. Rigiopappus leptocladus Gray Wireweed. Slender annual herbs; herbage puberulent to glabrate; stems 2-20 (30) cm tall; lateral branches, when present, very slender, over- topping the early flowers; leaves 0.3-2 (3) cm long, linear, the lower often withered at an- thesis; heads small; involucres 4-7 mm high and about as broad; bracts herbaceous, glabrous, thickened dorsally; rays few, yel- lowish, 1.5-2 mm long; pappus scales linear- subulate, ca 3 mm long; achenes 5-6 mm long. Reported for Utah by Cronquist (Flora of the Pacific Northwest), but not seen by me; 0 (0). RUDBECKIA L. Perennial caulescent herbs; leaves alter- nate, serrate or pinnately to palmately lobed; heads radiate or discoid, the rays (when pres- ent) neuter, commonly yellow; involucral bracts in 2 or 3 series, mainly unequal, her- baceous, spreading or reflexed; receptacle conic or columnar, chaffy throughout, the bracts clasping the achenes; disk flowers fer- tile; anthers obtuse or sagittate basally; style branches flattened; pappus a crown or none; achenes quadrangular or flattened at right angles to the involucral bracts. 1. Heads radiate; disks 1-2 cm wide and about as long, little elongate in fruit; plants of San Juan County H. laciniata — Heads discoid; disks 1.5-2.5 cm wide, mostly 2-5 cm long, elongating in fruit; plants not of San Juan County 2 2(1). Leaves laciniately lobed; plants glabrous or merely scabrous-ciliate on leaf margins; known from Iron and Washington counties R. montana — Leaves crenate-serrate, dentate, undulate, or entire, not lobed; plants evidently short-hairy to almost glabrous; known from mountains of central northern to south central Utah R. occidentalis Rudbeckia laciniata L. Cutleaf Cone- flower. Perennial herbs; stems erect from a coarse ligneus base, mainly 5-10 (20) dm tall, glabrous or scabrous-ciliate; leaves petiolate, the blades laciniate-pirmatifid to palmatifid, mainly 4-15 cm long and as broad; heads showy, the disk 1-2 cm wide and about as high; rays yellow, 6-16, deflexed-spreading, 2-5 cm long; pappus a short crown. Moist meadows at 1890 to 2200 m in San Juan County; Montana to Quebec, south to Ari- zona and Florida; 2 (0). Rudbeckia montana Gray? Perennial herbs; stems erect, from a short sub- rhizomatous caudex, 6-12 dm tall, glabrous; leaves petiolate, the blades laciniate-pinnati- fid, mainly 4-20 cm long and about as broad; heads discoid, the disk 1.5-2.5 cm wide, 3-5 cm high; rays lacking; pappus an irregularly margined, almost toothed crown. Iron and Washington counties; Colorado; 2 (0). Rudbeckia occidentalis Nutt. Western Coneflower. Perennial herbs; stems erect from a coarse ligneus rhizome, mainly 5-20 dm tall, glabrous or strigulose; leaves petiol- ate, the blades 5-20 cm long, 2.5-10 cm wide, ovate to ovate-lanceolate or lanceolate, attenuate to acuminate, entire, crenate-ser- rate, or dentate; heads discoid, the disks 1.5-2.5 cm wide, 3-6 cm long; rays lacking; 330 Great Basin Naturalist Vol. 43, No. 2 pappus a short crown. Mountain brush, as- margins scarious or hyaUne, or variously col- pen, grass-tall forb, and spruce-fir commu- ored; receptacle flat or convex, naked; ray nities at 2135 to 3175 m in Cache, Carbon, flowers yellow or orange, or sometimes lack- Duchesne, Emery, Piute, Salt Lake, Sanpete, ing; pappus or capillary bristles; style Sevier, Summit, Tooele, Utah, Wasatch, and branches flattened; achenes subterete, 5- to Weber counties; Washington to Montana, 10-nerved, glabrous or pubescent. Note: This south to California and Wyoming; 42 (iii). genus consists of a series of species that inter- grade freely when they are in contact with bENECio L. others of the group. Because of hybridization Annual, biennial, or perennial herbs with the species lines tend to be blurred, and it is rhizomes, caudices, or taproots, the juice wa- not possible to place all specimens with con- tery; stems erect, ascending, or decumbent at fidence. Keys are, and have been, based on the base; leaves alternate, simple, entire, features that are subject to interpretation; the toothed, or lobed to pinnatifid; heads soli- present one is not an exception, being tenta- tary, or few to many in corymbose cymes; in- tive at best. volucral bracts in 1 series, often with smaller Barkley, T. M. 1978. Senecio. N. Amer. bractlets at the base, green throughout or the Flora II. 10:50-139. 1. Leaves pinnatilobate with linear-filiform divisions or entire and linear-filiform; stems with leaves only gradually reduced upward, often more or less woody below 2 — Leaves variously lobed, toothed, or entire, but the segments and leaves not linear- filiform; stems with leaves various, seldom, if at all, woody at the base 3 2(1). Heads cylindric, subcylindric, or narrowly campanulate; main involucral bracts 8-13, the outer ones much reduced and inconspicuous; plants glabrous ..S. spartioides — Heads campanulate to broadly campanulate; main involucral bracts 13-21, the outer ones conspicuous, or, if inconspicuous, the plants tomentose S. douglasii 3(2). Heads nodding, especially in bud, or, if erect, with both distinctly black triangular tips on involucral bracts and cauline leaves prominently clasping 4 — Heads erect, even in bud; plants various but not as above 7 4(3). Heads discoid 5 — Heads radiate 6 5(4). Heads 8-12 mm high, 6-9 mm wide, narrowly campanulate, conspicuously pedunculate S. pudicus — Heads 12-20 mm high, 14-20 mm wide, broadly campanulate, short- pedunculate S. bigelovii 6(4). Heads erect, the involucral bracts black-tipped S. crassulus — Heads nodding, the involucral bracts often suffused with purple throughout, but not especially black tipped S. amplectens 7(3). Plants annual or winter annual, introduced weedy species S. vulgaris -^ Plants perennial, indigenous species 8 8(7). Stems uniformly leafy to the inflorescence, or the leaves concentrated upward 9 — Stems few leaved, or the upper leaves definitely reduced in size and distribu- tion 12 9(8). Stems 1-3 dm tall, more or less sprawling, arising from a subrhizomatous cau- dex S. fremontii — Stems mostly 2-15 dm tall, erect or ascending, arising from a rhizome or a caudex 10 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 331 10(9). Plants mainly 2-4 dm tall; leaves pinnatifid to lobed or laciniate; involucral bracts with dark tips S. eremophilus — Plants mainly 5-10 dm tall; leaves dentate to serrate; involucral bracts uni- formly greenish or brownish 11 11(10). Leaf blades acute to obtuse basally, the teeth all about alike S. serra — Leaf blades truncate to obtuse basally, or more or less hastately lobed, the lowermost teeth often the largest S. triangularis 12(8). Plants glaucous tall herbs, semiaquatic; leaves entire or denticulate, thick and leathery S. hydrophilus — Plants not or seldom glaucous, terestrial; leaves entire, toothed, or pinnatifid, not thick and leathery 13 13(12). Rays orange or orange-red (see also S. pauperculus) S. crocatus — Rays yellow or lacking 14 14(13). Heads discoid; plants tomentose, soboliferous S. fendleri — Heads radiate (or rarely discoid in some individuals); plants tomentose, gla- brate, or glabrous, not soboliferous (except in S. werneriifolius) 15 15(14). Leaves pinnatifid, at least the cauline ones, or the basal ones commonly roimded apically or oblanceolate to ovate or oval in outline 16 — Leaves serrate to entire, the basal ones variously shaped, but mainly acute to attenuate apically 20 16(15). Basal leaves distinctly pinnately divided, the lobes often again toothed S. multilobatus — Basal leaves merely toothed to subentire 17 17(16). Basal and lower cauline leaves entire to dentate, but not pinnatifid S. hartianus — Basal and lower cauline leaves toothed to pinnatifid 18 18(17). Middle and upper cauline leaves clasping with large auriculate bases S. dimorphophyllus — Middle and upper cauline leaves without a prominent clasping or auriculate base 19 19(18). Basal leaves obovate to oblanceolate or ovate, rounded apically, thickish; plants of dryish habitats S. streptanthifolius — Basal leaves oblanceolate to elliptic, obtuse, but usually pointed apically, thin; plants of meadows S. pauperculus 20(15). Cauline leaves rounded and more or less clasping basally, long-attenuate api- cally, entire or denticulate S. integerrimus — Cauline leaves tapering to the base or with a few basal clasping lobes in some, usually not attenuate apically 21 21(20). Stems subscapose, the cauline leaves none or few and bractlike; plants often so- boliferous or with a branching rhizomatous caudex S. werneriifolius — Stems more or less leafy, the cauline leaves gradually reduced upward, but hardly bractlike; plants seldom as above 22 22(21). Involucral bracts ca 8 or fewer; heads 5-6 mm wide, mainly 20-60 per in- florescence S. atratus — Involucral bracts mostly 13-21; heads 8-12 mm wide or more, fewer or larger than above 23 332 Great Basin Naturalist Vol. 43, No. 2 23(22). Achenes glabrous; plants often less than 20 cm tall 24 — Achenes hirtellous or hispidulous; plants often over 20 cm tall 25 24(23). Main leaves regularly and evenly pinnatifid or pinnatisect; plants often with slender rhizomes S. fendleri — Main leaves entire to dentate, not as above; plants shortly rhizomatous S. canus 25(23). Main leaves 10-15 cm long or more, entire or denticulate; plants 50-70 cm tall, of northern Utah S. sphaerocephahis — Main leaves 2-8 cm long, dentate, serrate, or subentire; plants mainly 15-35 cm tall, of southern Utah S. neomexicanus Senecio amplectens Gray Alpine Ground- sel. Perennial short-rhizomatous herbs; stems ascending to erect, mainly 8-30 cm tall; herbage glabrous or sparingly tomentose; main leaves middle and lower cauline, the lower ones broadly petiolate, more or less clasping the stem, the blades 3-10 cm long, 0.8-3 cm wide, dentate to shallowly lobed; cauline leaves becoming short-petiolate or sessile upward, finally bractlike; heads 1-5 (rarely more), conspicuously nodding, corym- bose; involucres broadly hemispheric, 10-15 mm long and about as wide or wider; bracts mainly ca 21, usually brown, with scarious margins, glabrous; outer bracts to about half as long as the inner; rays 7-17, yellow, 10-25 mm long; pappus white; achenes glabrous. Spruce-fir and alpine tundra communities, of- ten in talus or on ridge margins, at 3050 to 3570 m in Beaver, Grand, Piute, San Juan, Sanpete, and Utah counties (Wasatch, Tush- ar, and La Sal mountains, and Wasatch Plateau); Montana, Wyoming, Colorado, and Nevada; 24 (v). Our material belongs to var. holmii (Greene) Harrington [S. holmii Greene; Ligularia holmii (Greene) W.A. Weber]. Senecio atratus Greene Perennial sub- rhizomatous herbs from a branching caudex; stems erect or ascending, 2-8 dm tall; herb- age floccose-tomentose; basal and lower cau- line leaves petiolate, mainly 8-30 cm long, 1-4 cm wide, the blade oblanceolate or ob- long, conspicuously dentate to subentire; cauline leaves gradually reduced upward, be- coming sessile or subsessile and finally brac- teate; heads ca 15-60, in more or less com- pact corymbose clusters; involucres 6-8 mm high, 3-6 mm wide; main bracts 8 or fewer, greenish to brownish, the margins scarious, the tips black, tufted-hairy apically; rays 3-5, yellow, 4-8 mm long; pappus white; achenes glabrous. Aspen, spruce-fir, mixed conifer, and tall forb communities at 2440 to 3335 m in Duchesne, Garfield, Iron, San Juan, Salt Lake, Sanpete, and Uintah counties; Colo- rado and New Mexico; 22 (iv). Senecio bigelovii Gray in Torr. Bigelow Groundsel. Perennial subrhizomatous herbs; stems erect, mainly 3-8 (10) dm tall; herbage floccose-tomentose to glabrate or glabrous; main leaves cauline, largest below, reduced gradually upward, petiolate below, sessile and clasping to auriculate above, mostly 7-15 cm long, 0.6-3 (5) cm wide, the blades oblan- ceolate to oblong or elliptic, subentire to ser- rate; heads 3-8, nodding, racemosely ar- ranged; involucres 8-12 mm long, 12-25 mm wide; bracts mainly ca 21, usually brown, with scarious margins, the outer to half as long as the inner, all sparingly tomentose; ray flowers lacking; achenes glabrous. Mountain brush, ponderosa pine, aspen, and spruce-fir communities at 2745 to 3175 m in San Juan County; Wyoming south to New Mexico and Arizona; 2 (0). Our material has been as- signed to var. hallii Gray; the type variety is more southern. Senecio canus Hook. Gray Groundsel. [S. piirshianus Nutt.; S. convalliiim Greenm., type from Rabbit Valley]. Perennial short- rhizomatous herbs, often with a caudex; stems 8-30 cm tall (rarely more), erect or as- cending; herbage wooUy-tomentose; basal leaves petiolate, the blades 1-5 cm long, 3-30 mm wide, lanceolate to oblanceolate, elliptic or ovate, entire or denticulate, obtuse to rounded apically; cauline leaves reduced upward, the upper ones often clasping, final- ly bracteate, occasionally lobed in some in- trogressant forms; heads mainly 2-10, sub- umljellate or corymbose; involucres 3-8 mm long, 4-10 mm wide; main bracts 13-21, lance-attenuate, greenish or with brownish April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 333 midstripe, glabrous or tomentose; outer bracts very short; rays 8-13, yellow, 5-10 mm long; achenes glabrous. Pinyon-juniper, sagebrush, Douglas fir, aspen, spruce-fir, and alpine timdra commvmities, often in talus or on windswept ridges, at 2105 to 3815 m in Beaver, Box Elder, Cache, Carbon, Daggett, Duchesne, Garfield, Iron, Juab, Millard, Piute, San Juan, Sanpete, Sevier, Summit, Uintah, and Utah counties; British Columbia to Manitoba, south to California, Nevada, Colorado, and Kansas; 56 (xiii). This attrac- tive grayish white species forms inter- mediates with S. multilobatus, S. streptanthi- folius, and S. werneriifolius. Senecio crassulus Gray Perennial short- rhizomatous herbs, often with a caudex; stems 15-50 cm tall or more, erect; herbage glabrous; lower leaves broadly petiolate, the main ones 3-15 cm long, 0.6-3 (5) cm wide, lanceolate to elliptic or oblanceolate, dentate to entire; cauline leaves reduced upward, be- coming sessile and clasping; heads solitary or 2-12, corymbose; involucres 8-13 mm high, 12-21 mm wide; main bracts 8-21, oblong to lance-oblong, greenish to brown, with scarious margins, the tips black and tufted- hairy; outer bracts to half as long as the inner or more; rays 8-13, yellow, 5-12 mm long; achenes glabrous. Aspen, lodgepole pine, and spruce-fir communities, often in forb-grass meadows, at 1830 to 3355 m in Box Elder, Cache, Carbon, Duchesne, San Juan, Salt Lake, Sanpete, Sevier, Summit, and Utah counties; Oregon to Montana, south to New Mexico; 34 (iv). Senecio crocatus Rydb. Perennial sub- rhizomatous herbs, the caudex more or less developed; stems erect, mainly 2-8 dm tall; herbage glabrous or with minute hairs in the inflorescence; basal leaves with long slender petioles, the blades 1-8 cm long, 1-4 cm wide, ovate to oblong, lanceolate, or elliptic, subcordate to acute basally, often rounded apically, entire to crenate-dentate; cauline leaves reduced upward, becoming lobed or sublyrate, sessile and sometimes auriculate and/or clasping; heads mainly 3-30; in- volucres 4-8 mm long, 5-8 mm wide; main bracts 13-21, lance-oblong, green or suffused with red or purple; outer bracts very short; rays 6-13, orange or orange-red; pappus white; achenes glabrous. Rush-grass, willow, aspen-forb, and lodgepole pine communities at 2195 to 2990 m in Cache, Duchesne, Rich, Summit, and Utah counties; Colorado; 23 (i). One specimen from Rich County (Thome 1465 BRY) is apparently intermediate with S. eremophilus. The species is remarkably like the next. Senecio dimorphophyllus Greene Per- ennial subrhizomatous herbs; stems erect, mainly 30-70 cm tall; herbage glabrous or es- sentially so; basal leaves with long slender petioles, the blades 1-7 cm long, 1-5 cm wide, oval to oblong or elliptic, subcordate to acute basally, commonly rounded apically; cauline leaves becoming sessile, lyrate-pin- natifid, and auriculate-clasping, the auricles often lobed; heads mainly 2-25, sub- umbellately to corymbosely arranged; in- volucres 5-8 mm high, 6-10 mm wide; main bracts 13-21, lance-attenuate, green, some- times suffused reddish, the tips not black, tufted-hairy; outer bracts very short; rays 8-13, yellow, 5-8 mm long; pappus white; achenes glabrous. Two weakly discernible va- rieties are present in Utah. Cauline leaves merely lobed to subentire; plants of the La Sal Mountains S. dimorphophyllus var. intermedius Cauline leaves sharply lobed; plants of Uinta Mountains and Wasatch Plateau S. dimorphophyllus var. dimorphophyllus Var. dimorphophyllus Aspen-tall forb and spruce-fir communities at 1860 to 3265 m in Duchesne, Emery, Sanpete, and Utah coun- ties; Wyoming and Colorado; 9 (0). Utah ma- terials approach S. crocatus in most morpho- logical features, including the tall stature. If the flower color is discounted and the larger heads are not definitive, then the specimens could be considered as a portion of S. cro- catus. Some specimens from Duchesne Coun- ty appear to be transitional to S. sphaerocephalus. Var. intermedius T.M. Barkley Wet mead- ows at 3050 to 3115 m in the La Sal Moun- tains, San Juan County (type from Geyser Pass); endemic; 2 (0). 334 Great Basin Naturalist Vol. 43, No. 2 Senecio douglasii DC. Suffrutescent pe- rennials; stems erect or ascending, mainly 3-8 (10) dm tall; herbage glabrous or tomen- tose; leaves simple and linear-filiform or pin- natifid into linear-filiform segments, 2-11 cm long, 0.8-3 mm wide; heads few to numer- ous, in paniculately branched subcorymbose cymes; involucres campanulate, mainly 5-10 mm long, 6-14 mm wide; main bracts 13-21, lance-oblong, green, with scarious margins, minutely tufted-hairy apically; the outer bracts short and inconspicuous or to half as long as the inner ones; rays 8-17, yellow, 10-18 mm long; pappus white; achenes hairy. Two infraspecific taxa, previously treated at specific rank with some justifica- tion, are present in Utah. 1. Herbage grayish or whitish tomentose; outer involucral bracts short and incon- spicuous; plants rather broadly distributed S. douglasii var. longilobus — Herbage green, glabrous or essentially so; outer involucral bracts to about half as long as the inner ones; plants of Washington County ... S. douglasii var. monoensis Var. longilobus (Benth.) L. Benson [S. longilobus Benth.; S. filifolius var. iamesii T. & C, nom. illeg.]. Warm desert shrub, salt desert shrub, sagebrush-rabbitbrush, saltgrass, and pinyon-jimiper communities at 1095 to 2200 m in Beaver, Duchesne, Garfield, Iron, Kane, Millard, Piute, San Juan, Sevier, Wash- ington, and Wayne counties; Arizona to Texas; 52 (x). Var. monoensis (Greene) Jepson [S. mon- oensis Greene]. Creosote bush, blackbrush, other warm desert shrub, and pinyon-juniper communities at 760 to 1465 m in Washington Coimty; California to Texas; 23 (vi). Senecio eremophilus Richards. Perennial subrhizomatous herbs; stems rather equably leafy, erect or ascending, mainly 2.5-9 dm tall; herbage glabrous or essentially so; lower leaves often deciduous or withered at an- thesis; cauline leaves 2-15 cm long (or more), 0.4-5 (7) cm wide, oblanceolate to elliptic, or lanceolate in outline, pinnatifid or pinnately lobed or toothed, the lower ones petiolate, becoming sessile upward; heads several to numerous, corymbose; involucres 5-8 mm high, 6-10 mm wide; main bracts 8-17, lance-oblong, brownish or greenish, with scarious margins, blackish tips, and hair- tufted apices; outer bracts very short; rays 7-10, yellow, 5-10 mm long; pappus white; achenes glabrous or puberulent along the ribs. Grass-forb,- ponderosa pine, aspen, lodgepole pine, spruce-fir, and alpine timdra communities, at 1615 to 3450 m in Beaver, Carbon, Duchesne, Emery, Garfield, Grand, Iron, Juab, Piute, San Juan, Salt Lake, San- pete, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, Washington, and Wayne counties; British Columbia and Mackenzie south to Arizona and New Mexico; 104 (xiv). Our ma- terial belongs to var. kingii (Rydb.) Greenm. [S. kingii Rydb., type from Cottonwood Can- yon]. This plant forms intermediates with S. spartioides. Senecio fendleri Gray Perennial rhizo- matous herbs, with a caudex more or less de- veloped; stems mainly 5-30 cm tall, erect or ascending; herbage floccose-tomentose; basal leaves petiolate, the blades 1-6 cm long, 4-20 mm wide, pinnatifid or pinnately lobed; cauline leaves reduced upward, becoming sessile, finally bracteate; heads 3 to many, co- rymbose; involucres 4-6 mm high, 5-8 mm wide; main bracts ca 13, lance-attenuate, greenish, the margins scarious or hyaline, minutely hairy apically, more or less tomen- tose below; outer bracts very short; ray flow- ers lacking; pappus white; achenes glabrous. Ridge tops on limestone barrens near Musi- nea Peak, at ca 2960 to 3295 m in Sanpete County (Lewis 4274, 5516 BRY); Wyoming south to New Mexico; 2 (0). Our specimens approach S. canus, more or less. Senecio fremontii T. & G. Perennial herbs, subrhizomatous or from a caudex and tap- root; stems 0.6-4 dm tall; herbage glabrous; leaves cauline, 1-6 cm long, 0.5-2 cm wide, oblanceolate to obovate, shortly petiolate or sessile and somewhat clasping, dentate to subentire; heads 1-5; involucres 6-12 mm high, 7-12 mm wide; main bracts 8-17, lance-oblong or lance-attenuate, green or brown, the margins scarious, tufted hairy api- cally; outer bracts short and inconspicuous or April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 335 to half as long as the inner ones; rays 7-10, yellow, 5-12 mm long; pappus white; achenes glabrous or hairy. Two rather weak varieties are present. 1. Involucres mostly 8-10 mm high; stems mostly less than 20 cm high S. fremontii yar. fremontii — Involucres 10-12 mm high; stems often over 30 cm tall S. fremontii var. blitoides Var. blitoides (Greene) Cronq. [S. blitoides Greene]. Alpine communities, often in talus or on rock outcrops, at 2745 to 3355 m in Salt Lake (?), Tooele, and Utah counties; Wyoming to Colorado; 9 (0). Var. fremontii Spruce-lodgepole pine and alpine timdra communities at 3050 to 3965 m in Duchesne, San Juan, Salt Lake, Summit, Uintah, and Utah counties; British Columbia and Alberta, south to Oregon and Wyoming; 16 (v). Senecio hartianus Heller Perennial herbs from a subrhizomatous or stoloniferous cau- dex; stems erect, 2-5 dm tall; herbage floc- cose-tomentose, sometimes glabrate; basal leaves petiolate, the blades 1-5 cm long, 0.5-3 cm wide, oval to obovate or elliptic, serrate or crenate, rounded apically; cauline leaves reduced upward, subpinnatisect to en- tire; heads 3-12, corymbose; involucres 4-7 mm high and as broad; main bracts 13-21, lance-attenuate, greenish, the tips glabrous; rays ca 10-13, yellow, 5-8 mm long; achenes glabrous. Ponderosa pine community at ca 2290 m in Kane County (Atwood 7425 BRY); Arizona; 1 (0). Senecio hydrophilus Nutt. Water Ground- sel. Perermial subaquatic herbs from a caudex and fibrous roots; stems erect, mainly 4-10 dm tall; herbage glacuous, blue-green; basal and lower cauline leaves petiolate, the broad petioles with clasping bases, the blades 5-35 cm long or more, 1-10 cm wide, elliptic to oblanceolate, entire or denticulate, thick and leathery; cauline leaves reduced upward, be- coming sessile, finally bracteate; heads nu- merous in a branching corymbose cluster; in- volucres 5-8 mm long, 4-76 mm wide; main bracts 8-13, oblong or lance-attenuate, yel- lowish, the tips often black, tufted-hairy; rays 3-5 or lacking, yellow, 3-8 mm long; pappus white; achenes glabrous. Stream banks, pond margins, and wet meadows at 1375 to 2745 m in Carbon, Garfield, Juab, Kane, Piute, Salt Lake, Sanpete, Sevier, Sujnmit, and Utah counties; British Columbia, south to Califor- nia and Colorado; 25 (iv). Senecio integerrimus Nutt. Wet-the-bed. Perennial herbs with a short subrhizomatous caudex; stems mainly 1-6 (7) dm tall, erect; herbage arachnoid- villous or glabrate; basal and lower cauline leaves broadly petiolate, 3-20 cm long, 0.8-4 cm wide, lanceolate to elliptic or oblanceolate to oblong, entire or serrate to dentate, rounded to obtuse api- cally; cauline leaves reduced upward; heads few to many, in a corymbose to subumbellate cyme; involucres 6-12 mm high, 8-18 mm wide; main bracts 13-21, lance-attenuate, green, with scarious margins and black tips, the tips tufted-hairy; outer bracts very short; rays 8-13 (or lacking), yellow, 4-15 mm long; pappus white; achenes glabrous. Sagebrush, pinyon-juniper, forb-grass, mountain brush, ponderosa pine, aspen, and spruce-fir com- munities at 1460 to 3660 m in probably all Utah counties; British Columbia to Montana, south to California; 132 (xiv). Presumed hy- brids with S. dimorphophyllus are known (Hansen sn 1976 BRY). Senecio multilobatus T. & G. Uinta Groundsel. [S. lapidum Greenm., type from Silver Reef]. Perennial (or biennial?) herbs from a taproot; stems mainly 1-6 dm tall; herbage glabrous, glabrate, or tomentose throughout or only in axils of basal leaves; basal leaves 2-12 cm long, 0.3-3.5 cm wide, spatulate to obovate in outline, pinnatifid to lyrate-pinnatifid, the segments variously again toothed, petiolate; cauline leaves re- duced upward, finally bracteate; heads few to many, corymbose or subumbellate; in- volucres 4-9 cm high, 4-10 mm wide; main bracts 13-21, lance-attenuate or oblong-at- tenuate, the margins scarious, the apices hair tufted; rays 7-13, yellow, 4-10 mm long, or lacking; pappus white; achenes glabrous. Blackbrush, sandy desert shrub, pinyon-juni- per, sagebrush, mountain brush, ponderosa pine, aspen, lodgepole pine, and spruce-fir 336 Great Basin Naturalist Vol. 43, No. 2 communities at 915 to 3420 m in all Utah counties (type from the Uinta River); Idaho and Wyoming to California, Arizona, and New Mexico; 312 (xliv). This widespread and common species forms presumed hybrids with S. streptanthifolius and S. neo- mexicanus. Senecio neomexicanus Gray Perennial (or biennial?) herbs from a taproot; stems 14-40 cm tall, erect; herbage tomentose; basal and lower cauline leaves petiolate, the blades 1-5 cm long, 0.6-2 cm wide, oblanceolate to obo- vate or oval, dentate, serrate or subentire, toothed to obtuse apically; cauline leaves re- duced upward, toothed to lobed or entire, bracteate in inflorescence; heads few to many, corymbose or subumbellate; involucres 4-7 mm high, 5-12 mm wide; main bracts 13-21, lance-attenuate, green or brown, with scarious margins, not especially hairy api- cally; rays 8-13, yellow, 4-10 mm long; pap- pus white; achenes pubescent. Sagebrush, mountain brush, ponderosa pine, and aspen communities at 2105 to 3050 m in Garfield, Kane, San Juan, and Wayne counties; Colo- rado, New Mexico, and Arizona; 10 (0). Our materials are assigned to var. mutahilis (Greene) Barkley [S. mutabilis Greene]. Through this variety there is virtually a com- plete intergrading series into S. wernerii- folius, S. streptanthifolius, and S. multi- lobatus (Barkley 1978). Senecio pauperculus Michx. Perennial herbs from a subrhizomatous caudex; stems erect, mainly 2-4 dm tall; herbage glabrous or somewhat tomentose in axils of basal leaves; basal leaves petiolate, the blades mainly 2-6 cm long, 0.5-3 cm wide, oblan- ceolate to elliptic, obovate or ovate, crenate, dentate, or subentire; cuneate basally, toothed to obtuse apically; cauline leaves re- duced upward, becoming sessile, pinnatifid, not especially auriculate, finally bracteate; heads few to many, corymbose or sub- umbellate; involucres 4-8 mm long, 5-9 mm wide; main bracts 13-21, lance-attenuate, of- ten with scarious margins, the tips not espe- cially tufted-hairy; outer bracts very short; rays 8-13, yellow or yellow-orange, 4-10 mm long; pappus white; achenes glabrous or pu- berulent along the angles. Lodgepole pine and spruce-fir communities, usually in moist meadows, at 2345 to 2745 m in Daggett, Garfield, and Rich counties; Alaska to Lab- rador, south to Oregon and Georgia; 4 (0). Our material is intermediate to both S. strep- tanthifolius and S. crocatus. Senecio pudicus Greene [S. cernuus Gray, not L.f.; Ligularia pudica (Greene) W.A. Weber]. Perennial herbs from a sub- rhizomatous caudex; stems 20-50 cm tall, erect; herbage glabrous; basal and lower cauline leaves petiolate, the blades 3-15 cm long, 0.5-3 cm wide, lanceolate to oblanceol- ate or narrowly elliptic, tapering basally, acute apically, entire or shallowly dentate; cauline leaves reduced upward, finally brac- teate; heads few to many, nodding; in- volucres 5-9 mm long and as broad; main bracts 8-13, lance-oblong, green to brown, the margins scarious, tufted-hairy apically; outer bracts very short; ray flowers lacking; pappus white; achenes glabrous. Aspen, spruce-fir, and alpine tundra communities at 2650 to 3480 m in Carbon and Garfield coun- ties; Colorado; 11 (i). Senecio serra Hook. Perennial herbs from a caudex, with coarse, felt-covered roots; stems equably leafy, erect, 4-15 dm tall (or more), glabrous or sparingly tomentose; leaves 3-15 cm long, 0.4-4 cm wide, short-petiolate, the blades lanceolate to narrowly lanceolate or linear, dentate to subentire; heads several to numerous, corymbose; involucres 4-11 mm high, 2-10 mm wide; main bracts 8-13, lance-oblong, greenish to brownish, the mar- gins scarious, black-tipped, hair tufted; outer bracts very short; rays 5-8, yellow, 3-10 mm long; pappus white; achenes glabrous or es- sentially so. Two rather distinctive varieties are present. Involucral bracts 4-6 mm long, 2-6 mm wide; disk flowers ca 12; plants of central and northern Utah S. serra var. serra Involucral bracts 6-8 mm long, 6-10 mm thick; disk flowers ca 20; plants of San Juan County S. serra var. admirabilis Var. admirabilis (Greene) A. Nels. [S. ad- mirabilis Greene]. Ponderosa pine commu- nity at ca 1830 m in San Juan County; Wyoming and Colorado; 1 (0). April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 337 Var. serra Sagebrush, mountain brush, as- pen, forb-grass, lodgepole pine, and spruce- fir communities at 1830 to 3035 m in Box El- der, Cache, Davis, Duchesne, Juab, Rich, Salt Lake, Summit, Utah, and Weber counties; Washington to Montana, south to California and Nevada; 42 (vi). Senecio spartioides T. & G. Broom Groundsel. Perennial herbs from a taproot; stems equably leafy, erect or ascending, 2-10 dm tall or more, often in clumps; herbage glabrous; leaves 2-10 cm long or more, lin- ear, simple and entire or with linear lobes, mainly 1-3 mm wide (wider in some hybrid derivatives); heads several to many in branch- ing corymbose cymes; involucres subcylindric to narrowly campanulate, 5-10 mm high, 4-8 mm wide; main bracts 8-13, lance-linear, green, the margins scarious, not tufted-hairy; outer bracts very short; rays 4-8, yellow, 7-12 mm long; pappus white; achenes white- hairy. Two intergrading varieties are present. 1. Leaves simple and unlobed, or, if lobed, lower cauline leaves often over 4 mm wide; plants widespread S. spartioides var. spartioides — Leaves commonly with 4-6 lateral lobes, seldom if ever more than 2.5 mm wide; plants of southeastern Utah S. spartioides var. multicapitatus Var. multicapitatus (Greenm. in Rydb.) Welsh comb. nov. [based on: Senecio multi- capitatus Greenm. in Rydb. Bull Torrey Bot. Club 33: 160. 1906]. Warm desert shrub and pinyon-juniper communities, often in saline riparian sites, at 1220 to 1895 m in Garfield, Grand, San Juan, and Wayne counties; Colo- rado, Arizona, New Mexico, and Texas; 11 (vii). Barkley (1978) hesitated to combine S. multicapitatus with S. spartioides, because of field distinctions. They are, however, much alike and evidently lack diagnostic criteria that will allow segregation of all specimens. Further, specimens intermediate between S. spartioides and S. eremophilus bear "multi- capitatus" leaves. I follow a moderate course in maintaining this taxon at varietal level. Var. spartioides [S. incurvus A. Nels., type from Zion National Park]. Warm desert shrub, pinyon-juniper, sagebrush, mountain brush, and aspen communities, often in sand, at 1155 to 2870 m, in Beaver, Duchesne, Emery, Garfield, Grand, Iron, Kane, Piute, San Juan, Sanpete, Sevier, Uintah, Washing- ton, and Wayne counties; Wyoming to South Dakota, south to California and New Mexico; 75 (xxv). Intermediates are formed with S. eremophilus. Senecio sphaerocephalus Greene [S. lugens var. hookeri D.C. Eaton, type from Summit (?) County]. Perennial herbs from a short stout rhizome; stems erect or ascending, 3-8 dm tall; herbage tomentose; basal leaves pet- iolate, the blades 4-15 cm long, 1-3.5 cm wide, oblanceolate to elliptic, entire or den- ticulate, obtuse apically; cauline leaves re- duced upward, becoming sessile, finally brac- teate; heads few to many, corymbose; in- volucres 3-7 mm long, 6-12 mm wide; main bracts 13-21, oblong- to ovate-lanceolate, greenish or brownish, with scarious margins, the tips black, hair-tufted apically; outer bracts very short; rays 8-13, yellow, 4-10 mm long; pappus white; achenes hairy. Lodgepole pine and spruce-fir communities, in meadows, at 2315 to 3205 m in Daggett, Duchesne, Summit, and Wasatch counties; Oregon and Montana, south to Nevada and Wyoming; 10 (i). Senecio streptanthifolius Greene [S. aqua- riensis Greenm., type from Aquarius Plateau; S. jonesii Rydb., type from Alta; S. leonardii Rydb., type from American Fork Canyon; S. malmstenii Blake in Tidestr., type from Wasatch Mountains; S. ruhricaulis var. aph- anactis Greenm., type from Logan; S. wardii Greene, type from Fish Lake Mountain]. Pe- rennial herbs from a taproot and simple or branched and infrequently subrhizomatous caudex; stems erect, mainly 8-47 cm tall; herbage glabrous or rarely sparingly tomen- tose; leaves thickish; basal leaves petiolate, the blades 1-5 cm long, 0.3-3 cm wide, ob- lanceolate to obovate, suborbicular, elliptic, or ovate, crenate, dentate, or subentire, less commonly lobed; cauline leaves reduced up- ward, commonly some of them pinnatifid, fi- nally bracteate; heads few to many, corym- bose to subumbellate; involucres 4-8 mm high, 5-12 mm wide; main bracts 8-21, lance-oblong, green or brownish, the margins scarious, sparingly hair-tufted apically; outer 338 Great Basin Naturalist Vol. 43, No. 2 bracts very short; rays 8-13, yellow, 5-8 mm long; pappus white; achenes glabrous. Sage- brush, mountain brush, ponderosa pine, as- pen, lodgepole pine, spruce-fir, and alpine tundra communities, often in meadows, at 1370 to 3415 m in Box Elder, Cache, Carbon, Daggett, Duchesne, Emery, Garfield, Grand, Juab, Kane, Millard, Salt Lake, Sanpete, Se- vier, Summit, Tooele, Uintah, Utah, Wasatch, Washington, Wayne, and Weber counties; Yukon to Northwest Territories, south to Cal- ifornia and New Mexico; 107 (vii). This spe- cies forms a plexus around which revolves such species as S. pauperculus, S. multi- lobatus, S. neoniexicanus, and S. canus, as judged from morphological intermediates, which are presumed to be hybrids. Senecio triangularis Hook. Arrowleaf Groundsel. Perennial herbs from a caudex and more or less well-developed rhizome; stems equably leafy, erect, 2.5-12 dm tall or more; herbage glabrous or sparingly tomen- tose; leaves petiolate, the blades mainly 3-15 cm long, 0.5-6 cm wide, lance-oblong to triangular, abruptly contracted or subhastate at the base, dentate to sinuate dentate or sub- entire, finally bracteate in the inflorescence; heads few to many, subcorymbose; involucres 6-12 mm high, 8-17 mm wide; main bracts 8-12, lance-attenuate, the margins often scarious, tufted-hairy apically; outer bracts very short; rays 5-9, yellow, 6-15 mm long; pappus white; achenes glabrous. Aspen- mountain brush, Douglas fir-white fir, lodge- pole pine, and spruce-fir communities at 1765 to 3265 m in Cache, Duchesne, Grand, Salt Lake, Summit, Utah, and Weber coun- ties; Alaska and Yukon, south to California and New Mexico; 25 (viii). Senecio vulgaris L. Common Groundsel. Plants annual or biennial, with fibrous roots, 1-5.5 dm tall, the stems glabrous or sparingly villous; basal leaves smaller than the main cauline ones, often withered by anthesis; cauline leaves not much reduced upward, 2-10 cm long, 0.5-4.5 cm wide, irregularly piimatifid, the lobes again toothed, glabrous or more or less villous, especially along the veins beneath, the lower ones petiolate, the upper ones becoming sessile and auriculate- clasping; heads few to many; involucres 5-8 mm high, 4-10 mm wide; the outer bracts short and black tipped, the inner lance-linear, green, with scarious margins, black tipped; ray flowers lacking; pappus white; achenes hairy. Weedy species of disturbed sites in Salt Lake and Utah counties; adventive from Eu- rope; 6 (0). Senecio werneriifolius (Gray) Gray [S. aureus var. werneriifolius Gray]. Plants com- monly rhizomatous or soboliferous herbs; stems erect or ascending, 3-18 cm tall; herb- age tomentose, often glabrate or glabrous in age; basal leaves petiolate, the blades 0.6-3 cm long, 0.4-2 cm wide, oval to elliptic, obo- vate, or oblanceolate, thickish, sometimes revolute; cauline leaves few, commonly in- conspicuous and bracteate; heads 1-4; in- volucres 4-10 mm long, 7-15 mm wide; main bracts 13-21, lance-oblong, green or suffused with purple, the margins scarious, hair tufted apically; outer bracts to half as long as the in- ner; rays 8-13, yellow, 4-10 mm long; pap- pus white; achenes glabrous. Ponderosa pine, western bristlecone pine, aspen-conifer, and spruce-fir communities, often in semibarrens, at 2375 to 3600 m in Beaver, Duchesne, Gar- field, Iron, Juab, Piute, Salt Lake, Summit, and Utah counties; Idaho and Montana, south to California, Nevada, and Arizona; 28 (ii). SOLIDAGO L. Perennial herbs from a caudex or rhizome; leaves alternate, simple; heads numerous, radiate, yellow, borne in paniculate, race- mose, or cymose clusters; involucres imbri- cate in several series or subequal, commonly chartaceous or with the tips green; recep- tacle flat, naked; ray flowers fertile; disk flowers perfect, fertile; anthers subentire ba- sally; style branches with lanceolate ap- pendages; pappus of capillary bristles; achenes few nerved, pubescent. 2(1). Heads in corymbs or flat-topped cymes; leaves punctate; plants of lower eleva- tions riparian habitats [Euthamia] S. occidentalis Heads racemose or panicled; leaves not punctate; plants of various habitats 2 Stems glabrous 3 Stems puberulent with short incurved hairs or villous with multicellular hairs 4 April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 339 3(2). Plants definitely rhizomatous; involucres 2.5-4 mm long; plants of lower ele- vations S. missouriensis — Plants subrhizomatous; involucres 4-6 mm long; plants of higher elevations S. spathulata 4(2). Stems villous with multicellular hairs; petioles long-ciliate S. multiradiata — Stems puberulent with short incurved hairs; petioles scabrous or strigose mar- ginally 5 5(4). Involucres 6-11 mm high, the outer bracts subfoliaceus S. parryi — Involucres 2-5 mm high, the bracts not subfoliaceus 6 6(5). Leaves very numerous and much longer than the internodes, gradually attenu- ate or acuminate, not dimorphic, strongly 3-nerved S. canadensis — Leaves not very numerous, often less than twice as long as the internodes, acute or rounded apically, often dimorphic, with lateral nerves obscure or moderately apparent 7 7(6). Leaves sparingly hairy to glabrous, the margins rough-hairy; plants widespread and common S. sparsiflora — Leaves cinereus-puberulent with disoriented hairs, the margins hairy like the surfaces; plants more restricted and less common S. nana Solidago canadensis L. Goldenrod. [S. al- tissima L.; S. lepida DC.]. Perennial herbs from creeping rhizomes; stems 3-12 dm tall or more; herbage puberulent with short in- curved hairs, or the stems glabrous below; basal leaves often deciduous or withered at anthesis; cauline leaves numerous and crowded, 2-10 cm long or more, 3-20 mm wide, lanceolate to lance-linear, or narrowly elliptic, tapering to a sessile base, 3-nerved, serrate to entire, attenuate to acuminate api- cally; inflorescence commonly (but not al- ways) of recurved branches with secund heads; involucres 2-5 mm high and about as broad, the bracts lance-attenuate, scarious or greenish; rays 10-17, yellow, 1-3 mm long. Riparian and other mesic sites at 350 to 2290 m in all Utah counties; widespread in North America; 87 (xvi). This plant serves as host for a peculiar red and black leaf beetle. A phase of the species is cultivated as an orna- mental in Utah. Designation of varietal level in Utah seems academic. The species is tran- sitional to S. sparsiflora. Solidago missouriensis Nutt. Missouri Goldenrod. Perennial herbs from creeping rhizomes; stems* 2-5 (9) dm tall; herbage glabrous or sparingly puberulent in in- florescence only; basal leaves oblanceolate, often withered at anthesis; main cauline leaves 2-13 cm long, 0.4-1.5 cm wide, oblan- ceolate to elliptic or linear, tapering to a ses- sile base, mainly 3-nerved, entire or essen- tially so, acute to obtuse apically; in- florescence compact, with ascending branches, somewhat or not at all secund; in- volucres mostly 3-5 mm high and as broad, the bracts lance-attenuate, greenish to scarious; rays 7-13, yellow, 2-3 mm long. Ri- parian communities at 1525 to 2475 m in Box Elder, Carbon, Daggett, Duchesne, Emery, Salt Lake, Sanpete, Summit, Uintah, and Wasatch counties; British Columbia to On- tario, south to Arizona, Texas, and Tennessee; 13 (ii). Solidago multiradiata Ait. Low Gold- enrod. [S. ciliosa Greene]. Perennial herbs from a rhizome or rhizomatous caudex; stems 5-45 cm tall; herbage villous with multi- cellular hairs, at least on upper stem and pet- iole bases; basal and lower cauline leaves 1.5-14 cm long, 5-24 mm wide, oblanceolate to spatulate or elliptic, tapering to a con- spicuously ciliate petiole, obscurely 3-nerved, entire or serrate, rounded to obtuse apically; inflorescence loosely to densely corymbose; involucres 4-6 mm high, 5-7 mm wide, the bracts lance-oblong, green apically, with prominent midvein; rays ca 13, yellow, 4-5 mm long. Aspen, lodgepole pine, spruce-fir, and alpine tundra communities at 2745 to 3660 m in Beaver, Cache, Carbon, Duchesne, Garfield, Grand, Iron, Juab, Kane, Piute, San Juan, Salt Lake, Sanpete, Sevier, Summit, 340 Great Basin Naturalist Vol. 43, No. 2 Uintah, and Utah counties; Alaska to Quebec, south to California and New Mexico; 84 (xvii). Our specimens belong to var. scopulo- rum Gray. Solidagu nana Nutt. Dwarf Goldenrod. [S. radulina Rydb., type from Cottonwood Can- yon]. Perennial herbs from a rhizome or sub- rhizomatous caudex; stems 13-48 cm tall; herbage densely canescent with fine hairs of mixed orientation; basal and lower cauline leaves petiolate, 1.5-9 cm long, 0.7-2.3 cm wide, oblanceolate to spatulate, tapering to a petiole, weakly 3-nerved, entire or slightly toothed, rounded to obtuse apically; cauline leaves definitely reduced upward; in- florescence corymbose, seldom if at all se- cund; involucres 4-6 mm high and about as broad; rays 5-8, yellow, 3-4 mm long. Desert shrub upward to spruce-fir communities, mainly in riparian or wet meadow sites, at 1460 to 2745 m in Duchesne, Kane, Salt Lake, Sevier, Summit, Uintah, Utah, and Wasatch counties; Idaho to Montana, south to Arizona and Colorado; 13 (i). Solidago occidentalis (Nutt.) T. & G. Western Goldenrod. [Euthamia occidentalis Nutt.]. Perermial herbs from elongate rhi- zomes; stems erect, branched above, mainly 4-12 (20) dm tall; herbage essentially glabrous; leaves numerous, sessile, linear to lance-linear, 2-10 cm long, 1-10 mm wide; inflorescence usually large, leafy-bracted, broadly roimded; involucres 3.5-4.5 mm high and about as broad, the bracts narrowly ob- long, greenish apically, the midnerve con- spicuous; rays 15-30, yellow, 1.5-2.5 mm long. Riparian habitats at 850 to 1650 m in Box Elder, Cache (?), Carbon, Duchesne, Emery, Garfield, Grand, Juab, Kane, San Juan, Salt Lake, Uintah, Utah, Washington, and Weber counties; British Columbia and Alberta, south to California, New Mexico, and Nebraska; 42 (x). I follow tradition by in- cluding this taxon in Solidago; it might best be treated in Euthamia. Solidago parryi (Gray) Greene Parry Goldenrod. [Haplopappus parryi Gray; H. parryi var. minor Gray, type from Alta]. Per- ennial rhizomatous herbs; stems erect or as- cending, 8-50 cm tall; herbage scabrous to hispidulose; basal and cauline leaves petio- late, mainly 3-20 cm long, 0.9-3.8 cm wide, oblanceolate to elliptic, entire, obtuse to rounded apically; cauline leaves becoming sessile and smaller upward, more or less clasping; heads few to many in compact branched cymes; involucres 8-11 mm high, 7-14 mm wide; outer bracts ovate to ovate- lanceolate, green, ciliate, the bases often scarious; inner bracts narrower and with scarious or hyaline margins; rays 12-20, yel- low, 5-8 mm long. Aspen, tall forb, lodge- pole pine, spruce-fir, and alpine tundra com- munities at 2285 to 3570 m in Beaver, Carbon, Duchesne, Emery, Garfield, Juab, Kane, Millard, Piute, San Juan, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, and Wasatch counties; Wyoming, New Mexico, Arizona; 50 (ix). Solidago sparsiflora Gray [S. garrettii Rydb., type from Big Cottonwood Canyon]. Perennial rhizomatous herbs; stems erect or ascending, mainly 15-50 dm tall; herbage puberulent (often sparingly so on leaf sur- faces); leaves cauline or basal, oblanceolate to elliptic or spatulate, mainly 1-10 cm long, 2-25 mm wide, entire or less commonly some of them serrate, acute to attenuate or obtuse to rounded apically, often dimorphic, with the upper ones reduced in size; inflorescence a pyramidial to conic or cylindric cluster, compact or with branches curved and heads secund; involucres 4-6 mm high and about as broad; bracts oblong to subulate, chartaceous basally, green apically, the midvein con- spicuous; rays 5-10 or more, yellow, 3-4 mm long. Pinyon-juniper, mountain brush, sage- brush, aspen, ponderosa pine, and spruce-fir communities at 1125 to 3050 m in all Utah counties (except Box Elder and Morgan); Wyoming and South Dakota, south to Ari- zona and Nevada; 141 (xix). Our materials are far from uniform; in the hanging gardens of southeastern Utah they are transitional to S. canadensis (having more ray flowers), and at high elevations they are more or less inter- mediate with S. spathulata. Possible addition- al influence of S. mollis Bartl. and/or S. nemoralis Ait. is indicated, although neither of them is known from the state currently. Solidago spathulata DC. Perennial herbs from a subrhizomatous caudex; stems 5-30 cm tall (rarely more), erect or ascending; herbage glabrous or somewhat scabrous and often glutinous above; basal leaves oblanceo- late to spatulate, 2-15 cm long, 8-30 mm April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 341 wide, serrate to entire, obtuse to rounded apically; cauline leaves reduced upward, fi- nally sessile and more or less clasping; in- florescence compact to elongate, narrow, the heads not secund; involucres 4-6 mm high and as broad or more; bracts oblong, scarious or greenish along the prominent midvein; rays 5-10, yellow, 2.5-4 mm long. Aspen, spruce-fir, and alpine tundra communities at 2440 to 3510 m in Beaver (?), Daggett, Du- chesne, Emery, Garfield, Grand, San Juan, Sevier, Summit, and Uintah counties; Alaska to Quebec, south to California, Arizona, and New Mexico; 21 (v). Two completely inter- grading phases, regarded as varieties, are present in Utah; a tall montane phase known as var. neomexicana (Gray) Cronq. [S. multi- radiata var. neomexicana Gray], and a dwarf alpine phase known as var. nana (Gray) Cronq. [S. humilis var. nana Gray; S. decum- bens Greene]. SONCHUS L. Annual or perennial herbs from taproots or deep-seated, rhizome-like roots, the juice milky; leaves chiefly cauline, alternate, simple, entire to lobed or pinnatifid; heads few to several; involucral bracts imbricate in several series, green or greenish (drying brownish), the inner ones with hyaline mar- gins; receptacle naked; corollas of ray-flow- ers only, yellow, perfect; pappus of capillary bristles; style branches semicylindrical; achenes compressed, several to many nerved, beakless, glabrous. 1. Plants perennial, spreading from rhizomelike roots; involucres more than 14 mm long in fruit 2 — Plants armual from taproots; involucres less than 14 mm long in fruit 3 2(1). Involucres and peduncles bearing coarse stipitate glands S. arvensis — Involucres and peduncles glabrous or tomentose, not stipitate-glandular S. uliginosus 3(1). Leaves sharply and narrowly toothed, and sometimes lobed; achenes not trans- versely wrinkled, merely longitudinally nerved S. asper — Leaves sharply and broadly toothed, or merely toothed and lyrate pinnatifid; achenes transversely wrinkled and longitudinally nerved S. oleraceus Sonchus arvensis L. Field Sow-thistle. Plants perennial with deep-seated rhizome- like roots; stems 4-10 dm tall or more, pu- bescent with coarse stipitate glands, at least above, and often glabrous below; leaves 5-40 cm long, 0.8-10 cm broad, more or less pin- natifid, auriculate-clasping basally, acute to obtuse apically, prickly margined; heads few to several, the peduncles stipitate-glandular; involucres 14-20 mm high and 10-30 mm broad in fruit, the bracts lance-oblong to lance-linear, glandular like the peduncles; rays yellow, mostly 10-20 mm long; achenes transversely wrinkled. Weedy species of dis- turbed soils at 1370 to 2135 m in Cache, Duchesne, Salt Lake, and Utah counties; widely distributed and considered as a "noxious" weed in North America; adventive from Europe; 10 (0). Sonchus asper (L.) Hill Spiny Sow-thistle. Plants annual from taproots; stems 3-10 dm tall, pubescent with coarse stipitate glands, at least above, often glabrous below (less com- monly throughout); leaves 3-15 cm long, 1-5 cm broad, merely lobed or lobeless, auricu- late-clasping basally, acute to acuminate or less commonly obtuse apically, the margins armed with slender sharp prickles; heads few to several, the peduncles stipitate-glandular or glabrous; involucres 9-14 mm long and 10-16 mm wide in fruit, the bracts lance-ob- long to lance-linear, glabrous or with few stipitate glands; rays yellow, mostly 5-10 mm long; achenes 2-3 mm long, several nerved, not transversely wrinkled. Weed of disturbed sites at 760 to 2135 m in Box Elder, Du- chesne, Garfield, Grand, Kane, Millard, Piute, San Juan, Salt Lake, Sevier, Tooele, Uintah, Utah, and Washington counties; widespread in North America; adventive from Europe; 27 (ii). Sonchus oleraceus L. Common Sow-thistle. Plants annual from taproots, the stems 2-10 dm tall or more, glabrous throughout or 342 Great Basin Naturalist Vol. 43, No. 2 sometimes with stipitate glands above; leaves 4-20 cm long, 0.6-10 cm broad, more or less lyrate-pinnatifid, auriculate-clasping basally, acute to obtuse apically, irregularly and broadly toothed, the teeth weakly prickly; heads few to several, the peduncles glabrous or stipitate-glandular; involucres 10-13 mm high and 8-20 mm broad in fruit, the bracts lance-linear to lance-oblong, glabrous or with a few stipitate glands; rays yellow, mostly 8-12 mm long; achenes 2-3 mm long, several nerved and transversely wrinkled. Weeds of disturbed sites at 850 to 2135 m in Duchesne, Garfield, Salt Lake, Utah, and Washington counties; widely distributed in North Ameri- ca; adventive from Europe; 9 (i). Sonchus uUginousus Bieb. Meadow Sow- thistle. Plants perermial from deeply seated rhizomelike roots; stems 4-10 dm tall or more; herbage glabrous or obscurely tomen- tose; leaves 5-40 cm long, 0.8-10 cm wide, pinnatifid, auriculate-clasping basally, acute to obtuse apically, prickly margined; heads few to several, the peduncles glabrous; in- volucres mainly 14-16 mm high and 10-20 mm broad in fruit; bracts lance-linear to ob- long, glabrous or tomentose; rays yellow, mostly 10-20 mm long; achenes 2-3.5 mm long, several nerved, transversely wrinkled. Weeds of disturbed sites at 1220 to 2260 m in Daggett, Duchesne, Garfield, Grand, Juab, Salt Lake, Uintah, and Utah counties; wide- spread in North America; adventive from Eu- rope; 22 (ii). Authors of Flora Europaea (Tut- in et al. 1976) treat this entity as S. arvensis ssp. uliginosus (Bieb.) Nyman. Amow et al. (Flora of the Central Wasatch Front, Utah) discount the usefulness of stipitate glands as diagnostic features, noting that glandular and eglandular plants occur together in the same populations, and that glands are not corre- lated with other features. On a statewide basis the plants act like legitimate taxa, and the eglandular plants do seem to have some- what smaller heads. Sphaeromeria Nutt. Perennial herbs or subshrubs; leaves alter- nate or mainly basal, simple and entire or pinnatifid to palmatifid; heads discoid, few to several, corymbose to subcapitate; involucres hemispheric to campanulate; bracts in 2 or 3 series, imbricate to subequal; receptacle con- ic or concave, naked; outer flowers pistillate, fertile; disk flowers perfect, fertile; pappus lacking or a short crown; achenes usually 5- to 10-ribbed, glabrous or glandular. Holmgren, A. H., L. M. Shultz, and T. K. Lowrey. 1976. Sphaeromeria, a genus closer to Artemisia than to Tanacetum (Asteraceae: Anthemidae). Brittonia 28: 255-262. 2(1). Plants pulvinate-caespitose; heads capitately arranged on subscapose branches; known from Garfield County S. capitata Plants caulescent subshrubs; heads in paniculate or corymbose clusters on leafy branches, not of Garfield County 2 Leaves pinnatifid, at least some, tomentose; heads paniculate; plants of Wash- ington County S. ruthiae Leaves entire or pinnatifid, glabrous; heads corymbose; plants not of Washing- ton County S. diversifolia Sphaeromeria capitata Nutt. [Tanacetum capitatum (Nutt.) T. & G.]. Pulvinate-caespi- tose herbs; herbage canescent with malpi- ghian hairs; stems subscapose, 2-12 (20) cm tall; leaves mainly basal, 4-10 mm long, 1- or 2-palmately lobed, the cauline entire and re- duced upward; heads few to numerous in a compact headlike cluster; involucres 3-5 mm high, the broad bracts with hyaline margins; corollas 2.5-3 mm long. With western bristlecone pine on Cedar Breaks limestone, at ca 2380 m in Garfield County; Montana and Wyoming; 1 (0). Sphaeromeria diversifolia (D.C. Eaton) Rydb. [Tanacetum diversifolium D.C. Eaton]. Subshrubs, mainly 1-4 dm tall; herbage glabrous; leaves simple, entire, or some of them pinnately lobed, 8-55 mm long, 0.5-5 mm wide, linear; heads several to many in compact to open corymbose clusters; in- volucres 3-4 mm high, the broad bracts with hyaline margins; corollas 2-2.5 mm long. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 343 Juniper, mountain brush, mixed conifer, and aspen communities upward to alpine tundra, often in rock crevices, at 1370 to 3205 m in Davis, Juab, Millard, Salt Lake, Tooele, and Utah counties; Nevada; 33 (i). This is a Great Basin endemic. Sphaeromeria ruthiae Holmgren, Shultz, and Lowrey Subshrubs, mainly 3-7 dm tall; herbage tomentose-canescent with malpi- ghian hairs; leaves pinnately lobed or the up- per ones entire, 1-9 cm long, 2-4 mm wide or more; heads several to many, paniculate; involucres 3-5 mm high, the broad bracts with hyaline margins; corollas 1.8-2 mm long, yellow. Crevices in Navajo Sandstone, ponderosa pine community, in Washington County; endemic; 3 (0). Stephanomeria Nutt. Nom. Cons. Annual, biennial, or perennial herbs with milky juice; leaves alternate, often pinnatifid; flowers all raylike, perfect, pink or white; in- volucres cylindric; main bracts few, subequal; outer bracts much shorter; receptacle naked; pappus of plumose bristles (barbellate in S. spinosa); achenes 5-angled or -ribbed. 1. Plants annual, from slender taproots S. exigua — Plants perennial, a caudex often more or less developed 2 2(1). Plants spinescent; pappus barbellate S. spinosa — Plants unarmed; pappus plumose 3 3(2). Involucres 12-15 mm high; heads with 10 or more flowers S. parryi — Involucres 5-12 mm high (rarely higher); heads with 3-9 flowers 4 4(3). Main leaves runcinate-pinnatifid; plants commonly 1-2 dm tall S. runcinata — Main leaves entire or pinnatifid, often deciduous at anthesis; plants commonly 2-8 dm tall 5 5(4). Stems very slender; leaves filiform to linear, entire or dentate; pappus bristles white (rarely brownish), plumose to the base S. tenuifolia — Stems not very slender; leaves linear-subulate, often pinnatifid or lobed; pap- pus brownish, scabrous toward the base S. pauciflora Stephanomeria exigua Nutt. Annual or biennial (winter annual) herbs from slender taproots; herbage glabrous or puberulent; stems 5-60 cm tall, erect and commonly branched from the base upward, often fis- tulous; main leaves 1-6 cm long, pinnatifid to bipinnatifid, deciduous or withered by an- thesis; cauline leaves soon reduced and brac- teate upward; heads more or less corymbose, terminating bracteate branchlets; involucres 5-10 mm high, 3-4.5 mm wide; main bracts usually 3-5; rays pink or white, 3-5 mm long; pappus of white to off-white bristles plumose in the upper half; achenes 3-4 mm long, tuberculate. Warm, mixed cool, and salt desert shrub, and pinyon juniper commu- nities, often in sand, at 850 to 2230 m in Bea- ver, Emery, Garfield, Grand, Kane, Millard, San Juan, Sevier, Tooele, Uintah, Utah, Washington, and Wayne counties; Oregon to Wyoming, south to California and New Mex- ico; 84 (x). Stephanomeria parryi Gray Perennial herbs; stems 1 to few, weak, branching, 8-25 cm tall; herbage glabrous; leaves 2-8 cm tall, runcinate-pinnatifid, thickish, the lobes weakly spinulose-tipped; heads terminating very short bracteate branches, 10- to 14-flow- ered; involucres 12-15 mm high; rays whit- ish, 15-20 mm long; pappus bristles tawny, scabrous at the base only; achenes 3-4 mm long, not rugose. Blackbrush community at ca 1460 m in Kane County (Atwood & Allen 2822a BRY); California to Arizona; 1 (0). Stephanomeria pauciflora (Torr.) A. Nels. in Coult. & Nels. [Prenanthes? pauciflora Torr.]. Perennial herbs (or somewhat woody below) from a caudex, branched from the base, mostly 30-60 cm tall; herbage glabrous; main leaves 2-7 cm long, runcinate-pinnati- fid, the lobes weakly spinulose-toothed; heads terminating short to elongate branchlets, 3-to 5-flowered; involucres 8-10 high, 3-5 mm wide; main bracts 5; rays pink or white, 344 Great Basin Naturalist Vol. 43, No. 2 mainly 4-7 mm long; pappus bristles brown- ish, plumose except at the base; achenes 3.5-7 mm long, striate, more or less wrinkled. Warm, salt, and mixed desert shrub, and juni- per communities, often in sandy soil, at 760 to 1525 m in Beaver, Garfield, Grand, Juab, Kane, Millard, San Juan (?), Tooele, and Washington counties; California to Kansas, south to Texas and Mexico; 26 (v). Stephanomeria runcinata Nutt. Perennial herbs from a caudex; stems branched from the base, mostly 8-25 (30) cm tall; herbage glabrous, scabrous, or sparingly villous; main leaves 2-7 cm long, runcinate-pinnatifid, the lobes merely cuspidate; heads terminating naked or sparingly bracteate branchlets, com- monly 5-flowered; involucres 9-12 mm high, 3.5-7 mm wide; rays pink, mainly 8-12 mm long; pappus bristles white, plumose almost to the base; achenes 4-5 mm long, tubercu- late. Salt Desert shrub and pinyon-juniper communities at 1250 to 2535 m in Daggett, Duchesne, Emery, Grand, Uintah, and Wayne counties; Montana to Nebraska and Colorado; 17 (i). Stephanomeria spinosa (Nutt.) Tomb [Lygodesmia spinosa Nutt.]. Perennial herbs from a woody caudex, the caudex branches clothed with brownish marcescent leaf bases, the axils copiously villous-hairy; stems 11-52 cm tall, thorny; herbage glabrous upward or the branches puberulent; leaves linear 0.5-7 cm long, 1-3 mm wide, reduced to bracteate scales upward, often lacking at anthesis; heads terminal on short lateral branches or sessile, 3- to 5-flowered; involucres 5.7-10 mm high, 3-5 mm wide; main bracts oblong to lance-oblong, green or often suffused with purple; outer bracts proportionately broader; rays pink, 3-5 mm long; pappus bristles off- white, scabrous throughout; achenes 4-6.5 mm long, smooth. Desert shrub, sagebrush- grass, pinyon-juniper, mixed conifer, and as- pen communities, often in moist sites, at 1675 to 3050 m in Beaver, Emery, Garfield, Juab, Kane, Millard, Piute, Sevier, Tooele, Wash- ington, and Wayne counties; British Colum- bia to Montana, south to California and Ari- zona; 41 (ii). Stephanomeria tenuifolia (Torr.) Hall Slender Wirelettuce. [Prenanthes? tenuifolia Torr.]. Perennial herbs from a woody caudex; caudex branches lacking or with few marces- cent leaf bases, not hairy; stems 25-100 cm tall or more; herbage glabrous or puberulent; leaves filiform to linear, 1-8 (11) cm long, 1-3 (8) mm wide, entire or dentate, much re- duced upward; heads terminating elongate or short lateral bracteate branchlets, 5-flowered; involucres 8-11.2 (16) mm high, 3-5 mm wide; main bracts lance-oblong, green, pu- berulent or glabrous; outer bracts very short; rays 4-8 (10) mm long, pink; pappus bristles white, dull white, or less commonly brown- ish, plumose to the base; achenes 4-6 mm long, longitudinally ribbed, smooth. Two more or less distinctive phases are present, recognizable as varieties. Involucre 10-16 mm high, the bracts attenuate; basal leaves bipinnatifid, at least some; plants of Uintah County S. tenuifolia var. uintaensis Involucres mainly 8-11.2 mm high, the bracts not especially attenuate; basal leaves seldom if ever bipinnatifid; plants of rather broad distribution S. tenuifolia var. tenuifolia Var. tenuifolia Desert shrub, hanging gar- den, pinyon-juniper, mountain brush, pon- derosa pine, and white-fir communities, at 1155 to 2746 m in Beaver, Duchesne, Emery, Garfield, Grand, Iron, Kane, Piute, San Juan, Sevier, Uintah, Washington, and Wayne counties; British Columbia to Montana, south to California, Arizona, and Texas; 46 (xvi). The great sprawling plants of the canyon- lands portion of Utah might be worthy of tax- onomic consideration; sometimes they ap- proach S. pauciflora in having tawny pappus bristles. Var. uintahensis Goodrich & Welsh Pon- derosa pine community at ca 2490 m in Uin- tah County; endemic; 2 (0). Stylocline Nutt. Woolly annual herbs; stems commonly branched; leaves alternate, simple, entire; heads discoid, leafy bracted; involucre per se lacking; outer receptacular bracts subtending and enclosing pistillate flowers; receptacle cylindric; pistillate flowers many, deciduous with the enclosing bract, the bract apex hya- line; corollas filiform; pappus none; perfect April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 345 flowers (functionally staminate) few, sur- rounded by linear hyaline bracts; corollas tubular, the ovaries vestigial; pappus of 3-5 deciduous bristles; anthers sagittate basally; achenes ellipsoid, few nerved. Stylocline micropoides Gray Desert Nest- straw. Annual woolly herbs; stems usually branched, 4-12 cm tall; leaves 4-12 mm long, 0.5-1.5 mm wide, acute; bracteate leaves 6-10 mm long, 1.5-2.5 mm wide, lan- ceolate; heads clustered at branch tips, dense- ly woolly; pistillate flowers with bracts boat shaped, densely long-woolly, hyaline mar- gined; staminate flowers with pappus of 3-5 deciduous bristles; achenes ellipsoid, ca 1.5 mm long. Blackbnish, bursage, and indigo bush communities at 915 to 1160 m in San Juan and Washington counties; California to New Mexico, south to Mexico; 3 (i). Syntrichopappus Gray in Torr. Aimual herbs; stems simple or branched; leaves alternate (or some opposite below), simple, entire or lobed; heads radiate, many, terminating branchlets; involucres sub- cylindric; bracts few, in 1 series, partly en- closing ray achenes; receptacle flat, naked; ray flowers pistillate, fertile, yellow; disk flowers perfect, fertile, yellow; anthers ob- tuse at base; style branches flattened; pappus of barbellate bristles; achenes 5-angled. Syntrichopappus fremontii Gray Annual herbs, 2-14 cm tall; herbage floccose-tomen- tose; leaves 5-22 mm long, narrowly spatu- late to spatulate, rounded to 3-lobed apically, cuneate basally; heads few to many; in- volucres 5-6 mm high, 3-4 mm wide; bracts 5, oblong, greenish, with scarious margins, abruptly acute apically; rays 5, yellow, 2-5 mm long; disk corollas numerous, yellow; pappus of white barbellate bristles falling to- gether. Joshua tree, creosote bush, black- brush, sagebrush, and juniper communities at 760 to 1375 m in Washington County; Cali- fornia, Nevada, Arizona; 6 (i). Tanacetum L. Perennial herbs from a rhizome; leaves al- ternate, 2- to 3-pinnatifid; heads discoid, nu- merous, corymbose; flowers perfect; in- volucres hemispheric; bracts in 2 or 3 series, more or less imbricate, the margins scarious; receptacle low-convex, naked; anthers entire at the base; pappus a minute crown; achenes 5-angled, truncate. Tanacetum vulgare L. Tansy. Aromatic, glabrous or sparingly tomentose perennials, 3-10 (15) dm tall; leaves 6-15 cm long, ses- sile or subsessile, the blades 2- to 3-pinnatifid; heads many, discoid, yellow; involucres ca 4-5 mm high and 6-10 mm broad; bracts lan- ceolate; marginal flowers 3-lobed; inner flow- ers 5-lobed; achenes glandular, 5-angled, ca 1 mm long. Weedy species of disturbed soils at 1370 to 1985 m in Emery, Uintah, and Utah counties; widespread in the U.S.; adventive from Europe; 3 (0). Taraxacum Hall. Perennial scapose herbs with milk juice, from taproots; leaves all basal, pinnatifid to subentire; heads solitary on a scape; in- volucral bracts in 2 series, herbaceous, the outer shorter, the inner often dilated or ap- pendaged apically, usually with broad hya- line or scarious margins, at least basally; re- ceptacle naked; corollas of ray flowers only, perfect, yellow; pappus of capillary bristles; style branches semicylindric; achenes angular or terete, prominently nerved or ribbed, usu- ally spinulose or with ridges near the body apex, glabrous, beaked. 2(1). Irmer involucral bracts commonly dilated or bearing appendages apically, over 10 cm long; plants indigenous, of high elevations T. ceratophorum Inner invofucral bracts usually not dilated or with appendages apically; plants various 2 Outer bracts reflexed or spreading, the inner ones 12-18 mm long; achenes straw colored to olive drab or brownish; plants adventive T. officinale Outer bracts erect, the inner ones 6-10 mm long; achenes black to grayish; plants indigenous at high elevations T. lyratum 346 Great Basin Naturalist Vol. 43, No. 2 Taraxacum ceratophorum (Ledeb.) DC. Rough Dandelion. [Leontodon ceratophorus Ledeb.]. Plants mostly 4-10 cm tall, from a simple or branched caudex; leaves 4-8 cm long, 0.7-2 cm broad, subentire to toothed; scapes sparingly villous, moderately so below the head; involucres 12-17 mm high in flow- er, the outer bracts ovate to lanceolate, ap- pressed or ascending, the inner ones lance- oblong, attenuate, the apex dilated or appendaged; rays yellow; achene bodies 3-7 mm long, straw colored to olive-drab or brownish, the beak usually 2-4 times longer than the body; pappus white. Spruce krumm- holz and sedge-forb meadows at 3230 to 3660 m in Daggett, Duchesne, and Uintah counties (Leidy Peak); Alaska to Yukon, east to the Atlantic, south to California, New Mexico, and Massachusetts; circumboreal; 2 (0). Taraxacum lyratum (Ledeb.) DC. Alpine Dandelion. [Leontodon lyratus Ledeb.]. Plants mostly 2-8 cm tall, from a simple or branched caudex; leaves 1-6 cm long, 0.3-1 cm wide, pinnately lobed to pinnatifid or subentire; scapes glabrous or nearly so; in- volucres 6-10 mm high, the outer bracts lan- ceolate-ovate, appressed or ascending-spread- ing, the inner ones lance-oblong to oblong, scarcely or slightly dilated; rays yellow (fad- ing bluish); achene bodies 3-6 mm long, black or grayish, the beak subequal to the body; pappus white. Alpine tundra and meadows in spruce-fir communities at 3325 to 3965 m in Duchesne and Summit counties; Alaska and Yukon, south to Nevada, Arizona, and Colorado; Asia; 5 (i). Taraxacum officinale Weber ex Wiggars Common Dandehon. Plants mostly 3-60 cm tall, from a simple or branched caudex; leaves 5-40 cm long, 1-10 cm wide, pin- nately lobed to pinnatifid, the terminal lobe broader than the lateral ones; scapes villous to subglabrous, often moderately to densely villous below the head; involucres 15-25 mm high in flower, the outer bracts lance-acumi- nate, reflexed, the inner ones lance-attenuate, not or scarcely dilated apically, rarely appen- daged; rays yellow, or bluish externally; ach- ene bodies 3-4 mm long, straw colored to olive drab, the beak usually 2-4 times longer than the body; pappus white. Ubiquitous brightly flowered weedy species at 885 to 3205 m throughout Utah; widespread in North America; adventive from Eurasia; 65 (xiii). This handsome plant is among the earliest of our spring flowers, and among the last to bloom in autumn. Tetradymia DC. Armed or unarmed shrubs; stems pannose- tomentose; leaves alternate, entire, foliaceous or modified as spines, with secondary leaves fasciculate in the axils; heads discoid, corym- bose or racemose; involucres cylindric to tur- binate or hemispheric; receptacle naked; bracts 4-6, equal or nearly so; flowers 4-8, yellow or cream; style branches truncate to rounded or conic apically; anthers sagittate basally; pappus of capillary bristles or barbel- late scales; achenes striate. Strother, J. L. 1974. Taxonomy of Tetra- dymia (Compositae: Secenioneae). Brit- tonia 26:177-202. 1. Heads solitary or 2 or 3, axillary; primary leaves modified as spines 2 — Heads several to many in terminal corymbose clusters; primary leaves fo- liaceous or modified as spines 3 2(1). Spines commonly recurved, mainly 5-20 mm long, pannose-tomentose; achenes 6-8 mm long; plants widespread, not of Washington County T. spinosa — Spines straiglit, mainly 20-40 mm long, glabrescent; achenes 4-5 mm long; plants of Washington County T. axillaris 3(2). Primary leaves modified as persistent spreading, straight or recurved spines 5-25 mm long T. nuttallii — Primary leaves not modified as persistent spines, if at all spinescent then ap- pressed-ascending 4 April 1983 4(3). Welsh: Utah Flora: Compositae (Asteraceae) 347 Primary leaves linear-subulate, spinescent apically, appressed-ascending, to- mentose; secondary leaves obtuse apically, glabrous or essentially so T. glabrata Primary leaves various but not spinescent, not contrasting in shape and pubescence with the secondary ones T. canescens Tetradymia axillaris A. Nels. Longspine Horsebrush. Spiny shrubs, mainly 4-12 dm tall; branchlets evenly white-pannose; pri- mary leaves modified as persistent spines 1-5 cm long, straight or becoming curved, to- mentose at first, becoming glabrate; second- ary leaves linear to spatulate, 2-12 mm long, essentially glabrous; heads solitary or 2 or 3, from nodes of the previous year; involucres 8-11 mm high; bracts 5, subequal, tomentose; flowers 5-7, pale yellow, the corollas 7.5-9 mm long; pappus of slender bristles; achenes 4.5-5.5 mm long; achenes pilose, the hairs 9-11 mm long. Salt and warm desert shrub communities at 850 to 1375 m in Washington County; Nevada and California; 18 (ii). Our material belongs to var. longispina (Jones) Strother [T. spinosa var. longispina Jones, type from St. George]. Tetradymia canescens DC. Gray Horse- brush. [T. linearis Rydb., type from Iron County]. Unarmed shrubs, mainly 1-9 dm tall; branchlets white-pannose except for gla- brate streaks below the primary leaves; pri- mary leaves 0.5-4 cm long, 1-6 mm wide, lanceolate to oblanceolate or spatulate, to- mentose; secondary leaves similar to the pri- mary ones but shorter and narrower; heads few to several at branch tips; involucres 6-8 mm high or more; bracts 4, subequal, tomen- tose; flowers 4, yellow to cream, the corollas 7-11 mm long; pappus of white or tawny bristles; achenes 2.5-5 mm long, glabrous or hairy. Sagebrush-grass, mountain brush, pon- derosa pine, mixed conifer, and aspen com- munities at 1525 to 3150 m throughout Utah; British Columbia to Montana, south to Cali- fornia, Arizona, and New Mexico; 75 (viii). Tetradymia glabrata T. & G. Shrubs, mainly 3-12 dm tall; branchlets pannose ex- cept for glabrate or glabrous streaks below the primary leaves; primary leaves mainly 5-15 mm long, 0.8-1.4 mm wide, linear-sub- ulate, spinose tipped, soon deciduous; second- ary leaves linear to narrowly spatulate, glabrous or thinly tomentose; heads few to many on branch tips; involucres 7-10 mm high; bracts 4, subequal, tomentose to glabrous; flowers 4, yellow to cream, the co- rollas 9-10 mm long; pappus of white bristles; achenes 3-5 mm long, hirsute. Shad- scale, greasewood, sagebrush, rabbitbrush, and juniper communities at 1370 to 2370 m in Emery, Juab, Millard, Sanpete, Sevier, Tooele, and Wayne counties; Oregon and Idaho, south to California and Nevada; 44 (V). Tetradymia nuttallii T. & G. Nuttall Horsebrush. Spinescent shrubs, 3-12 dm tall; branchlets white-pannose except for glabres- cent streaks below the primary leaf bases; primary leaves modified as persistent straight or recurved spines 5-25 mm long, tomentose to glabrous; heads in terminal clusters of (2) 3-6; involucres 6-9 mm high; bracts 4, equal; flowers 4, yellow, the corollas 8-10 mm long; pappus of white or tawny bristles; achenes 4-6 mm long, hirsute. Shadscale, greasewood, sagebrush-rabbitbrush and pinyon-juniper communities at 1370 to 1830 m in Box Elder, Daggett, Duchesne, Juab, Millard, Tooele, and Uintah counties; Wyoming and Nevada; 25 (i). Tetradymia spinosa H. & A. Spinescent shrubs, 3-12 dm tall; branchlets evenly pan- nose; primary leaves modified as spines, 5-20 mm long, tomentose, finally glabrate; second- ary leaves linear to spatulate, glabrous or gla- brescent; heads borne singly or in pairs, later- ally, on stems of the previous season; involucres 8-12 mm high; bracts 4-6, sub- equal, tomentose; flowers 5-8, yellow, the corollas 6-10 mm long; pappus of slender bristles, white; achenes 6-8 mm long, hairy, the trichomes 9-12 mm long. Mixed desert shrub, shrub-grass, and pinyon-juniper com- munities at 1250 to 1925 m in Carbon, Dag- gett, Duchesne, Emery, Garfield, Grand, Juab, Millard, Salt Lake, Uintah, and Utah counties; Oregon to Montana and Wyoming, south to California, Nevada, and New Mexi- co; 15 (vi). 348 Great Basin Naturalist Vol. 43, No. 2 Thelesperma Less. Perennial glabrous or sparingly puberulent herbs; leaves opposite, pinnately to pal- mately parted, or the upper ones entire; heads pedunculate, solitary or few per stem; involucres hemispheric to campanulate; bracts in 2 unlike series, the outer ones spreading and distinct, the inner ones con- nate to the middle and calyxlike; receptacle flat, chaffy with broad scarious scales; rays present (or lacking), neuter, yellow; disk flowers perfect, fertile; anthers not caudate basally; pappus of 2 retrorsely hispid awns, a crown, or lacking; achenes oblong to linear. Plants 30-80 cm tall; rays normally lacking; pappus of 2 awns; known from San Juan and Washington counties T. megapotamicum Plants 3-35 cm tall; rays normally present; pappus a crown or none T. suhnudum Thelesperma megapotamicum (Spreng.) Kuntze Greenthread. [Bidens megapotamica Spreng]. Perennial herbs from a caudex and stout root; stems 30-80 cm tall; leaves mainly 2-7 cm long, once or twice pinnatifid, the lobes linear, or the uppermost simple; outer bracts 4-6, oblong to ovate, obtuse, much shorter than the inner; inner bracts 6-12 mm high, connate to above the middle, the lobes with narrow scarious margins; rays lacking; disk flowers yellow (or brownish); pappus of 2 or 3 retrorsely hispid awns; outer achenes somewhat papillose dorsally. Desert shrub commimity at ca 915 to 1375 m in San Juan and Washington counties; Wyoming to Ne- braska, south to Arizona, Texas, and Mexico; 1(0). Thelesperma subnudum Gray Perennial herbs from a taproot and less commonly with a caudex and creeping rootstock; stems 3-35 cm tall, subscapose; leaves mainly at base of stem, 1.5-9 cm long, pinnately to sub- palmately lobed or some or all of them en- tire; petioles often ciliate and blades more or less puberulent; involucres 6.3-14 mm high, 9-22 mm wide; outer bracts oblong to lan- ceolate, with narrow scarious margins, to half as long as the inner ones; inner bracts united to below the middle, conspicuously scarious- margined; rays present and bright yellow, 10-28 mm long and 6-18 mm wide, or lack- ing; disk flowers yellow; pappus a toothed crown or lacking; achenes glabrous or hairy apically, 3.5-4.5 mm long. This taxon is vari- able being radiate or discoid, in division of leaves, and in position of leaves along the stem. They occur mainly at elevations below 2135 m elevation. A dwarf alpine phase oc- curs above that elevation, and because of its small size, lack of rays, and apparent eco- typical differences these plants are herein designated at varietal level. 1. Plants mainly 3-7 cm tall; involucres 6.3-9 mm high, 9-14 mm wide; heads discoid T. subnudum var. alpinum — Plants mainly 9-35 cm tall; involucres 8-14 mm high, 12-22 mm wide; heads commonly radiate T. subnudum var. subnudum, Var. alpinum Welsh Pinyon-juniper, mountain brush, and western bristlecone pine communities at ca 2745 m in Wayne County; endemic; 2 (0). Var. subnudum Mixed desert shrub, salt desert shrub, and pinyon-juniper commu- nities at 1065 to 2135 m in Carbon, Du- chesne, Garfield, Grand, Iron (type from Red Creek), Kane, San Juan, Uintah, Washington, and Wayne counties; Colorado, Arizona, and New Mexico; 109 (xiv). TOWNSENDIA Hook. Annual, biennial, or perennial herbs, cau- lescent or acaulescent; leaves alternate, en- tire or rarely lobed or toothed; heads radiate, solitary or few, terminating branches, or ses- sile; receptacle convex, naked; involucres campanulate to hemispheric; bracts in 2-7 series; rays pistillate, fertile, the corollas white, pink, or yellow; disk flowers perfect, yellow; disk pappus of barbellate capillary bristles; ray pappus similar to that of the disk April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 349 or shortened; achenes 2- or 3-ribbed, com- Contr. Gray Herb. 183:1-151. pressed, usually hairy. Reveal, J. L. 1970. A revision of the Utah Beaman, J. H. 1957. The systematics and species of Townsendia (Compositae). evolution of Townsendia (Compositae). Great Basin Nat. 30:23-52. 1. Plants caulescent, the internodes apparent, annual or biennial (short-lived perennial) 2 — Plants acaulescent, the internodes not elongating, perennial 5 2(1). Plants annual or winter annual; disk pappus shorter than disk-corollas; plants of southeastern Utah (Navajo Basin) T. annua — Plants biennial or short-lived perennials; disk pappus subequal to or longer than the disk corollas 3 3(2). Achenial hairs unevenly branched; ray flowers usually dark pink-purple dor- sally; plants biennials of western Utah T. florifer — Achenial hairs glochidiate; ray flowers variously colored, but if dark pink- purple dorsally then the plants perennial and of different distribution 4 4(3). Stems gray-white, the pubescence dense; plants of broad distribution, perennial T. incana — Stems thinly strigose, evident beneath the hairs; plants of the Uinta Basin, bien- nial T. strigosa 5(1). Involucral bracts linear to narrowly lanceolate, in 5-7 series 6 — Involucral bracts lanceolate to ovate or elliptic, in 2-5 series 9 6(5). Involucral bracts hair tufted apically, linear, acuminate; plants of Carbon, Duchesne, and Daggett counties T. hookeri — Involucral bracts not hair tufted apically, narrowly lanceolate, acute; plants variously distributed 7 7(6). Rays glandular dorsally; leaves canescent; plants of Duchesne and Uintah counties T. mensana — Plants glabrous or sparingly pubescent dorsally; leaves greenish or grayish- canescent; plants not or seldom of Duchesne and Uintah counties 8 8(7). Disk pappus 3-6 mm long; leaves green, the midveins not conspicuous; plants of the Wasatch Plateau and Uinta Mountains T. leptotes — Disk pappus 6-11 mm long; leaves grayish canescent, the midveins con- spicuous; plants of Sevier, Iron, Wayne, and Garfield counties T. exscapa 9(5). Rays yellow ventrally, densely glandular and often purplish dorsally; ray pap- pus 1-2 mm long; plants of Emery and eastern Sevier counties T. aprica — Rays white or pink or bluish, or rarely yellow ventrally, but, if yellow, the ray pappus 2-4.5 mm long and plants of other distribution 10 10(9). Plants green or greenish; flowers often bluish or purplish to pink, mainly of higher elevations in mountains and plateaus T. montana — Plants grayish canescent or whitish; flowers seldom bluish or purplish, usually white to pink or yellowish ventrally; mainly of low elevations 11 11(10). Involucral bracts sparingly strigose; ray pappus 2-4.5 mm long; plants mainly of western Utah T. jonesii — Involucral bracts moderately strigose; ray pappus 0.3-0.6 mm long; plants mainly of eastern Utah T. incana 350 Great Basin Naturalist Vol. 43, No. 2 Townsendia annua Beaman Caulescent annual or winter annual herbs, 2-18 cm tall; herbage strigose; leaves of basal rosettes soon withered or poorly developed; cauline leaves 5-28 mm long, 1-5 mm wide, oblanceolate to spatulate or linear, sparingly to moder- ately strigose, green or greenish; heads soli- tary or few; involucres 4.5-7 mm long, 6-14 mm wide; bracts in 2-4 series, green or suf- fused with purple, scarious, ciliate; rays 13-34, the corollas white or pink to lavender, 4-8 mm long, 1-2.3 mm wide, glabrous; disk corollas yellow, 2.2-3.5 mm long; achenes 1.9-2.6 mm long, pubescent with glochidiate hairs; ray pappus 0.2-0.8 mm long, that of disk flowers 1.8-3 mm long. Sandy desert shrub and blackbrush communities at 1125 to 1590 m in Carbon, Emery, Garfield, Grand, Kane, and San Juan counties; Colorado, Ari- zona, New Mexico, and Texas; 23 (v). Townsendia aprica Welsh & Reveal Pulvi- nate-caespitose acaulescent perennial herbs from a caudex, 1.5-2.5 cm tall; leaves 7-13 (16) mm long, 1-3.5 mm wide, spatulate to oblanceolate, strigose; heads sessile, sub- mersed in the leaves; involucres 4-8 mm high, 7-13 mm wide; bracts in 3-4 series, lanceolate, fimbriate, red-scarious, hyaline- ciliate, the outermost sparsely strigose; rays 13-21, the corollas yellow to golden ven- trally, purplish dorsally and glandular, 4-7 mm long; disk corollas yellow, 3.7-4.5 mm long; achenes 2-2.5 mm long, 2-ribbed, the hairs glochidiate; ray pappus 0.7-1 mm long; pappus of disk flowers 4-5 mm long. Salt desert shrub and pinyon-juniper commu- nities, commonly on clay or clay-silt expo- sures of the Mancos Shale (Blue Gate Mem- ber), at 1860 to 2440 m in Emery and adjacent Sevier (type from south of Fremont Junction) counties; endemic; 10 (ii). The yel- low flowers and short pappus of ray flowers are diagnostic. Toumsendia exscapa (Richards.) T.C. Por- ter [Aster? exscapa Richards.]. Caespitose acaulescent perennial herbs from a simple or branched caudex, 2-3.5 cm high; leaves 0.6-5 cm long, 1-3.5 mm wide, oblanceolate to lin- ear, acute and mucronate apically, strigose, with midvein apparent; involucres 10-18 mm high, 15-30 mm wide; bracts in 4-7 series, linear to narrowly lanceolate, ciliate on scarious margins, sparingly strigose to glabrous; ray flowers 21-40, the corollas white or pinkish, 8-15 mm long, 1.2-3 mm wide; disk corollas yellow; achenes 2- or 3- ribbed, pubescent with glochidiate hairs; ray pappus 4-8 mm long; disk pappus 6-12 mm long. Ponderosa pine, mountain sagebrush, and spRice-fir communities, often in mead- ows, at 2135 to 3295 m in Garfield, Iron, Se- vier, and Wayne counties; British Columbia to Manitoba, south to Nevada, Arizona, Mex- ico, and Texas; 8 (ii). Townsendia florifer (Hook.) Gray [Erig- eron? florifer Hook.; T. watsonii Gray, type from Stansbury Island; T. scapigera var. am- higua Gray, type from Rabbit Valley; T. florifer var. communis Jones, type from Marysvale]. Caulescent winter annual or biennial herbs 3-20 cm tall; basal leaves 6-50 mm long, 3-12 mm wide, spatulate; cauline leaves narrowly oblanceolate to linear, 10-40 mm long, 1-5 mm wide, strigose, petiolate, grayish; heads solitary or few; involucres 6.5-13 mm high, 15-30 mm wide; bracts in 3 or 4 series, green or suffused with purple, scarious, ciliate; rays 13-34, the corollas white or pink ventrally, dark pink or laven- der dorsally, 7-12 mm long, 1.2-3 mm wide, often glandular; disk corollas yellow, 3.3-6 mm long; achenes 3.3-4.5 mm long, pu- bescent with unequally forked hairs; ray pap- pus 1-6 mm long; disk pappus 3.5-7.5 mm long. Mixed desert shrub communities at 1280 to 1985 m in Beaver, Box Elder, Gar- field, Juab, Millard, Sanpete, Sevier, Tooele, Utah, and Wayne counties; Washington to Idaho, Oregon, and Nevada; 56 (vii). Townsendia hookeri Beaman Caespitose acaulescent perennial herbs from a simple or branched caudex, 2.5-3.5 cm high; leaves 10-40 mm long, 1-2.5 mm wide, linear to linear-oblanceolate, strigose; involucres 9-13 mm high, 9-14 mm wide; bracts in 5-7 series, linear to lance-linear, tufted-hairy api- cally, green or suffused with purple, strigose; rays 13-34, the corollas 6-9 mm long, 1-1.9 mm wide, white or pink ventrally, pinkish dorsally, glabrous; disk corollas yellow, 4.5-6 mm long; achenes 3.5-4.5 mm long, pu- bescent with glochidiate hairs; ray pappus 1-1.5 mm long; disk pappus 5.5-8.5 mm long. Sagebrush, sagebrush-grass, and mixed conifer communities at 2165 to 2716 m in Carbon, Daggett, Duchesne, and Uintah April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 351 counties; Yukon to Saskatchewan, south to South Dakota and Colorado; 5 (0). Townsendia incana Nutt. [T. incana var. amhigua Jones, type from Thompson]. Sub- caulescent to acaulescent caespitose herbs, the caudex often branched; stems con- spicuously white strigose, mainly 2-6 cm high, forming clumps to 2 dm wide; leaves 5-40 mm long, 1-5 mm wide, spatulate to oblanceolate, strigose; heads solitary or few; involucres 7-11 mm high, 8-20 mm wide; bracts in 3 or 4 series, lanceolate, green, the margins scarious and ciliate, strigose; rays 13-34, the corollas white ventrally, pink to lavender dorsally, 6-10 mm long, 1.5-3 mm wide; achenes 2.5-4.5 mm long, pubescent with glochidiate hairs; ray pappus 0.3-0.6 mm long; disk pappus 4-7.5 mm long. Black- brush, salt desert shrub, mixed desert shrub, pinyon-juniper, and sagebrush commimities at 1310 to 2290 m in Beaver, Carbon, Dag- gett, Duchesne, Emery, Garfield, Grand, Iron, Kane, Piute, San Juan, Sevier, Uintah, and Wayne counties; Wyoming to Nevada, Arizona, and New Mexico; 183 (xxiii). This is the common townsendia of the Colorado drainage system in Utah; its Great Basin counterpart is T. pnesii, from which it can be distinguished by the white strigose stems and shorter ray pappus. Townsendia jonesii (Beaman) Reveal [T. mensana var. jonesii Beaman, type from Mammoth]. Subcaulescent to acaulescent caespitose herbs, the caudex commonly branched; stems not conspicuously white stri- gose, mainly 2-4 cm tall, forming clumps to 1 dm wide; leaves 10-40 mm long, 1-4 mm wide, oblanceolate to spatulate or almost lin- ear, strigose; heads mostly solitary; involucres 9-12.5 mm high, 8-14 mm wide; bracts in 4 or 5 series, lanceolate, green or suffused purple, sparsely strigose; rays 13-21, the co- rollas white to pink, cream, or yellow ven- trally, pink to red-purple dorsally, glandular, 4-7 mm long; disk corollas yellow, ca 3.5 mm long; achenes 3-5.5 mm long; pubescent with glochidiate hairs; ray pappus 2-4.5 mm long; disk pappus 5-8 mm long. Two weak, but geographically and edaphically correlated, varieties are present. 1. Ray flowers yellow to lemon-yellow ventrally; plants of gypsiferous substrates in Sevier and Piute counties T. pnesii var. lutea — Ray flowers pink to white or cream ventrally; plants of various substrates, rather broadly distributed T. pnesii var. pnesii Var. jonesii Sagebrush, shadscale, rabbit- brush, pinyon-juniper, mountain brush com- munities at 1525 to 2745 m in Beaver, Juab, Millard, Sanpete, and Sevier counties; Ne- vada; 13 (ii). The type of T. mensana var. pnesii consists of strictly acaulescent plants with very slender leaves and smallish heads; it is unmatched in the specimens examined, and it is understandable why the taxon was placed initially with T. mensana. Var. lutea Welsh Salt desert shrub and juniper communities at ca 1675 to 1830 m in Sevier and Piute counties (on Arapien shale and clays in volcanic rubble); endemic; 6 (i). Toumsendia leptotes (Gray) Osterh. [T. se- ricea var. leptotes Gray]. Perennial acaules- cent herbs from a simple or more commonly branched caudex, 1-3 cm tall; herbage spar- ingly strigose, greene; leaves 0.6-4 cm long, 1.3-2.6 mm wide, linear to narrowly oblan- ceolate; involucres 5-10 mm high, 9-14 mm wide; bracts in 4-7 series, lanceolate to lin- ear, the margins scarious, ciliate, often suf- fused purple; rays 13-34, the corollas white, cream, or pink ventrally, sometimes lavender dorsally, 6-10 mm long, 1.2-2 mm wide; disk corollas yellow, 3-5 mm long; achenes pu- bescent with glochidiate hairs; ray pappus 0.8-6.5 mm long; disk pappus like the ray pappus. Montane sagebrush and grass-forb communities, often on ridge crests and plateau margins at 2680 to 3145 m in Du- chesne, Sanpete, and Summit counties (Uinta Mountains and Wasatch Plateau); Idaho and Montana, south to California, Nevada, and New Mexico; 7 (0). Townsendia mensana Jones Perennial acaulescent herbs from a simple or more commonly branched caudex 1-2.5 cm high; herbage strigose; leaves 3-17 mm long, 0.6-1.3 mm wide, narrowly oblanceolate to linear; involucres 5-9 mm high, 76-10 mm wide; bracts in 4 or 5 series, lanceolate, the margin scarious and ciliate; rays 13-21, the corollas whitish, cream, or pinkish, glandular 352 Great Basin Naturalist Vol. 43, No. 2 dorsally, 5-7.5 mm long, 0.9-1.4 mm wide; disk corollas yellow, 3.5-4.8 mm long; achenes pubescent with glochidiate hairs; ray pappus 2.5-4 mm long; disk pappus 5-6.5 mm long. Salt desert shrub, pinyon-juniper, and sagebrush commimities, especially on barren and semibarren sites, at 1705 to 2715 m in Duchesne (type from near Duchesne, then Theodore) and Uintah counties; Colo- rado (?); a Uinta Basin endemic; 38 (v). Townsendia montana Jones Perennial acaulescent or rarely subcaulescent herbs from a simple or branched caudex, sometimes with soboliferous rhizomatous branches, from a taproot, 2-6 cm high; herbage glabrate to strigose; leaves 5-40 mm long, 2-8 mm wide, spatulate, thickish; involucres 6-12 mm high, 8-20 mm wide; bracts in 3-6 series, oblong, obovate, oblanceolate or lanceolate, glabrous or sparingly strigose, the margins scarious; ciliate, often suffused with purple; rays 12-30, the corollas blue, pink, lavender, or white, 6-12 mm long, 1-3.5 mm wide; ach- enes 3.7-5.2 mm long, glabrous or sparingly pubescent with bifurcate or glochidiate hairs; ray and disk pappus alike, 3-5.5 mm long. Three more or less distinctive varieties are present. 1. Heads usually sessile; leaves mainly 1-3.5 mm wide, rather abruptly obtuse apically; plants of Garfield and Kane counties T. montana var. minima — Heads usually at least shortly pedunclulate; leaves mainly broader (at least some), rounded to obtuse; plants not of Garfield or Kane counties 2 2(1). Leaves rounded apically, broadly spatulate; plants of calciferous outcrops in southern Duchesne, Wasatch, and Sanpete counties T. montana var. caelilinensis — Leaves obtuse to subacute apically; plants of various substrates in the Uinta and Wasatch mountains , T. montana var. montana Var. caelilinensis Welsh Pinyon-juniper, spruce-fir, and limber pine communities on Flagstaff Limestone and Green River forma- tions at 2135 to 3735 m in southern Du- chesne, Wasatch, and Sanpete counties; en- demic; 13 (i). Var. minima (Eastw.) Beaman [T. minima Eastw., type from Bryce Canyon]. Ponderosa pine, western bristlecone, limber pine, and Douglas fir-white fir communities, on white and pink members of the Cedar Breaks For- mation, at 2375 to 3115 m in Garfield and Kane counties; endemic; 14 (i). Var. montana [T. dejecta A. Nels., type from Dyer Mine]. Spruce-fir and lodgepole pine communities at 3050 to 3510 m in Cache, Juab, Salt Lake (type from Alta), Summit, and Uintah counties; Idaho, Mon- tana, and Wyoming; 2 (0). Townsendia strigosa Nutt. Caulescent biennial herbs; stem-s branched from the base and above, 3-15 cm long; herbage strigose to strigulose; basal leaves 1.2-4.5 cm long, 1.2-7 mm wide, oblanceolate to spatulate, more or less persistent; cauline leaves mostly smaller and narrower, often clustered below and overtopping the heads; involucres 5-10 mm high, 7-20 mm wide; bracts in 3 or 4 series, lance-ovate to lanceolate, the margins scarious, ciliate, strigose; rays 12-30, the co- rollas white to pink, sometimes darker dor- sally, 5-14 mm long, 1.5-3 mm wide; disk co- rollas 3.3-5 mm long; achenes pubescent with glochidiate hairs; ray pappus 0.5-1.6 mm long; disk pappus 3.3-5 mm long. Salt desert shrub, mixed desert shrub, and pinyon- juniper communities at 1460 to 1895 m in Daggett, Duchesne, and Uintah counties; Wyoming; 14 (ii). Tragopogon L. Biennial (annual or perennial) herbs from thickened taproots, the juice milky; leaves al- ternate, entire, clasping basally; heads soli- tary or few and corymbose; flowers all ray- like, perfect, yellow or purple; involucres cylindric or campanulate; bracts uniseriate, equal; receptacle naked; pappus of plumose bristles united at the base; achenes 5- to 10- nerved, slender-beaked or the outer beakless. OwNBEY, M. 1950. Natural hybridization and amphiploidy in the genus Tragopogon. Amer. J. Bot. 37:487-499. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 353 1. Peduncles scarcely if at all inflated, even in fruit; achenes 15-25 mm long (in- cluding the beak); bracts subequal to the rays; plants rare in Utah T. pratensis — Peduncles strongly inflated apically; achenes 25-36 mm long (including the beak); bracts usually longer than the rays; plants locally common 2 2(1). Rays purple; involucral bracts mainly 8 or 9 T. porrifolius — Rays yellow; involucral bracts usually 13 T. dubius Tragopogon dubius Scop. Biennial herbs; stems erect, 3-10 dm tall, simple or branched; leaves mainly 5-25 cm long, linear-subulate from an expanded base, floccose, becoming glabrate; peduncles enlarged and fistulous be- low the heads; involucres cylindric to cam- panulate; bracts commonly 13 (8 on later heads), 2.5-4 cm long in flower, 4-7 cm long in fruit; rays pale lemon yellow, shorter than the bracts; achenes 25-36 mm long; pappus whitish to tawny. Disturbed soils and in low quality range sites at 1370 to 3205 m in all Utah counties; widely distributed in the U.S.; adventive from Europe; 58 (vi). Tragopogon porrifolius L. Oyster-plant; Salsify. Biennial herbs; stems erect, 3-10 dm tall, simple or branched above; leaves mainly 5-30 cm long, linear-subulate, the apex not recurved; peduncles enlarged and fistulose below the heads; involucres cylindric to campanulate; bracts commonly 8 (5-11), 2.5-4 cm long in flower, 4-7 cm long in fruit; rays purple, subequal to or shorter than the bracts; achenes 25-35 mm long; pappus brownish. Cultivated plants, escaping and persisting on canal banks, in moist meadows, and along roadsides at 1370 to 2595 m in Carbon, Millard, Salt Lake, Sanpete, Summit, and Weber counties; widespread in much of the U.S.; introduced from Europe; 10 (0). Tragopogon pratensis L. Biennial herbs; stems erect, 1.5-8 dm tall, simple or branched; leaves mainly 5-30 cm broad, ta- pering from a broadly expanded base to 2 cm wide, recurved apically; peduncles not espe- cially enlarged in flower or in fruit; in- volucres campanulate; bracts commonly 8, 12-24 mm long in flower, 18-38 mm long in fruit; rays chrome-yellow, equaling or sur- passing the bracts; achenes 15-25 mm long; pappus off-white. Disturbed sites in Rich, Salt Lake, and Summit counties; widespread in the U.S.; adventive from Europe; 2 (0). Vanclevea Greene Shrubs; branchlets glutinous-resinous, green to tan, finally white to gray barked; leaves alternate, sessile, entire or serrate, fal- cately curved; heads discoid, yellow, solitary or cymose; involucres campanulate; bracts in 4 or 5 series, imbricate, glutinous; receptacle naked, resinous; styles long-exserted, the branches flattened, papillose; achenes cla- vate, 5-angled; pappus of 12-16 linear per- sistent slender scales. Vanclevea stylosa (Eastw.) Greene [Grin- delia stylosa Eastw., type from San Juan County]. Shrubs, mainly 5-12 dm tall; branchlets glutinous-resinous; bark tan to white or grayish black in age; leaves 0.6-3.5 cm long, 1-9 mm wide, narrowly lanceolate to oblong or elliptic, commonly entire, atten- uate to a spinulose tip; heads solitary or more commonly few to many in corymbose or cy- mose clusters; involucres 8-10 mm high, 9-15 mm wide; bracts lanceolate to lance-at- tenuate, sometimes abruptly acuminate and recurved apically, resin coated; corollas yel- low to cream, 6-7 mm long; achenes 4-5 mm long, compressed, glutinous and spreading hairy. Four-wing saltbush, ephedra, sand dropseed, Indian ricegrass, blackbrush, and juniper communities, in sand, at 1125 to 1620 m in Emery, Garfield, Grand, Kane, San Juan, and Wayne counties; Arizona (a Colo- rado Plateau endemic); 32 (viii). The genus is monotypic. Verbesina L. Annual (biennial or perennial?) herbs; leaves opposite, at least below, simple, toothed; heads radiate, showy; involucres biseriate, about equal, herbaceous; receptacle convex, chaffy, the bracts enfolding the ach- enes; rays yellow or yellow-orange, pistillate; disk flowers perfect, fertile; anthers subentire basally; style branches with hispidulous ap- pendages; pappus of 2 slender awns; achenes flattened, 2-winged. Verbesina encelioides (Cav.) Benth. [Xime- nesia encelioides Cav.]. Annual herbs; stems 4-10 dm tall, cinereous-strigose, often 354 Great Basin Naturalist Vol. 43, No. 2 branched above; lowest leaves opposite, al- ternate upward, petiolate, often with stipule- like appendages at base; blades 1.2-10 cm long, 0.7-6 cm wide, ovate to lanceolate, acute to attenuate, irregularly toothed, stri- gose beneath, green and sparingly strigose above; involucres 7-12 mm high, 15-25 mm wide; bracts lance-ovate to lance-linear, her- baceous, strigose; rays 10-15, yellow or yel- low-orange, 8-20 mm long; pappus of 2 short slender awns; achenes thickly 2-winged, pu- bescent. Sagebrush, rabbitbrush, saltgrass, pinyon-juniper, and ponderosa pine commu- nities, often in disturbed sites, at 1280 to 2260 m in Beaver, Garfield, Juab, Kane, San Juan, and Washington counties; Montana to California and Texas; 20 (v). Most of our ma- terial belongs to var. exariculata Robins. & Greenm. The bright flowers contrast sharply with the grayish-strigose pubescence, result- ing in a strikingly beautiful plant. Wyethia Nutt. Perennial herbs from thick taproots; stems erect or ascending; leaves alternate, simple; heads large, solitary or several, radiate; in- volucral bracts in 2-4 series, herbaceous or coriaceous; receptacle convex, chaffy, the bracts folded, persistent; rays yellow, pistil- late, fertile; disk flowers perfect, yellow; pappus a crown of scales or lacking; achenes trigonal or 4-angled, glabrous or pubescent. Weber, W. A. 1946. A taxonomic and cy- tological study of the genus Wyethia, family Compositae, with notes on the related genus BalsaTuorhiza. Amer. Midi. Nat. 35:400-452. 1. Leaves mainly cauline, the basal reduced or lacking, scabrous-roughened; plants of sandy desert shrublands W. scahra — Leaves basal and cauline, the basal often larger than the cauline ones, smooth or, if rough-hairy, not of lower elevations 2 2(1). Herbage glabrous, resinous; upper leaves rounded and clasping basally W. amplexicaulis — Herbage hirsute to glabrate; upper leaves petiolate W. arizonica Wyethia amplexicaulis (Nutt.) Nutt. Mulesears. [Espeletia amplexicaulis Nutt.]. Perennial herbs; stems mostly 2.5-9 dm tall, glabrous; basal leaves 12-40 cm long, 2-15 cm wide, entire or dentate, petiolate, res- inous; cauline leaves smaller, sessile, rounded and clasping basally; heads large, solitary or several; involucres hemispheric, 25-35 mm high, 25-50 mm wide; outer bracts fo- liaceous, subequal; rays 6-16, yellow, 2.5-4.5 cm long; pappus a crown, sometimes pro- longed into filiform awns; achenes 8-10 mm long, glabrous. Sagebrush, oak,, pinyon-juni- per, aspen-fir, and forb-grass communities at 1525 to 2745 m in Box Elder, Cache, Juab, Millard, Morgan, Salt Lake, Sanpete, Sevier, Summit, Tooele, Utah, Weber, and Washing- ton counties; Washington to Montana, south to Nevada and Colorado; 38 (ii). Wyethia arizonica Gray Perennial herbs; stems mainly 30-80 cm tall, spreading hairy, especially upward; basal leaves 15-40 cm long or more, 3-15 cm wide, petiolate, the blades oblanceolate to elliptic or lanceolate; cauline leaves smaller, attenuate basally to a short petiole; heads large, solitary or several; involucres hemispheric or campanulate, 20-30 mm high, 15-40 mm wide; outer bracts foliaceous, subequal; rays 6-16, yel- low, 2.5-4 cm long; pappus a crown, some- times prolonged into filiform awns; achenes 8-10 mm long, glabrous. Pinyon-juniper, oak, and ponderosa pine communities at 1430 to 2440 m in Grand, Kane, San Juan, and Wash- ington counties; Colorado, New Mexico, and Arizona; 9 (0). Wyethia scabra Hook. Robust, clump- forming perennial herbs; stems several to many, 1.5-6 dm high or more, scabrous and hispidulose; leaves mainly cauline, the lower ones rudimentary, 3-15 cm long, 3-17 mm wide, elliptic to oblong or linear, scabrous; heads solitary or few, terminating stems and branches; involucres hemispheric, 20-40 mm high, 20-55 mm wide; bracts lanceolate to linear, attenuate to caudate-attenuate; rays 10-23, yellow, 18-40 mm long; pappus a crown; achenes 6-8 mm long, glabrous. April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 355 Three more or less distinctive varieties are present. Diagnostic features are based on the nature of surface and habit of the involucral bracts, which in the typical, common phase is almost sufficiently variable as to include the others. 1. Involucral bracts long-attenuate from short dilated bases, ciliate with multi- cellular hairs, glabrous but with shiny resin droplets dorsally; plants of Kane County W. scabra var. attenuata — Involucral bracts variable in shape, ciliate or not, scabrous to pubescent and more or less glandular dorsally, but seldom if ever with resin droplets 2 2(1). Involucral bracts closely imbricate, the outer recurved-spreading, pubescent with short fine hairs; plants of San Juan, Grand, and eastern Kane counties W. scabra var. canescens — Involucral bracts various, scabrous to long-hairy dorsally; plants rather widely distributed W. scabra var. scabra Var. attenuata W. A. Weber Ponderosa pine, oak, and pinyon-juniper (less commonly in desert shrub) communities, in sand, at 1370 to 1985 m in Kane County (type from north of Kanab); Arizona; 13 (iii). This handsome plant is a botanical motif of the Coral Pink dunes area, and is also present on East Clark Bench. Var. canescens W. A. Weber. Warm desert shrub and mixed desert shrub communities at 1125 to 1680 m in Grand, Kane, and San Juan counties; Colorado, Arizona, and New Mexico; 4 (i). This is a variable entity transi- tional to the typical variety, especially in Grand and eastern Kane counties. Var. scabra Bl^ckbrush, vanclevea- ephedra, other mixed desert shrub, pinyon- juniper, and ponderosa pine communities at 1220 to 2625 m in Carbon, Daggett, Du- chesne, Emery, Garfield, Grand, Kane, and Uintah counties; Wyoming; 48 (vii). Xanthium L. Aimual herbs with fleshy large cotyledons and a taproot; leaves alternate, petiolate, the blades broad, rough-hairy; heads unisexual, discoid, or the corolla lacking; staminate heads uppermost, many flowered; involucral bracts in 1-3 series, separate; receptacle cy- lindric, chaffy; fikments monadelphous, the anthers separate; pistil vestigial, the styles unbranched; involucre of pistillate heads en- closing the 2 flowers, forming a 2-chambered bur armed with hooked prickles, the corolla lacking; achenes large, solitary in each cham- ber; pappus none. Xanthium strumarium L. Cocklebur. [X. italicum Moretti; X. pensylvanicum Wallr.]. Annual monoecious herbs; stem 1.5-10 dm tall or more, simple or branched, scabrous, often purple mottled; leaves petiolate, the blades mainly 2-12 cm long and about as broad, ovate to oval or orbicular, obtuse to cuneate or cordate basally, scabrous, dentate and often lobed; heads in few to many short axillary clusters; burs broadly cylindric to ovoid, 1-3.5 cm long, with 2 more or less in- curved beaks apically, covered with stout hooked prickles. Weedy species of cultivated and other disturbed lands, at 850 to 1925 m in much of Utah; adventive (?) from the east- ern U.S. or possibly from Europe; 33 (iii). The seedlings are poisonous to livestock, and they produce dermatitis in some people. Xylorhiza Nutt. Subshrubs or suffrutescent perennial herbs; branchlets green to straw colored or whitish; leaves alternate, simple; heads solitary at branch ends; involucres campanulate to hemispheric; bracts imbricate in several series, herbaceous to largely scarious, erect; ray flowers pistillate, fertile, yellow; achenes somewhat compressed, hairy; pappus of taw- ny to whitish capillary bristles. Note: Mem- bers of this genus are all primary or second- ary selenium indicators. Cronquist, a. and D. D. Keck. 1957. A re- constitution of the genus Machaeran- thera. Brittonia 9:231-239. Watson, T. J. 1977. The taxonomy of Xylor- hiza (Asteraceae-Astereae). Brittonia 29:199-216. 356 1. 2(1). 3(2). Great Basin Naturalist Vol. 43, No. 2 4(2). Leaves linear to linear-filiform, the margins entire and more or less involute; plants of Kane and Garfield counties X. confertifolia Leaves serrate to serrate-dentate, or, if entire, of other distribution (except X. cronquistii) 2 Leaves serrate to serrate-dentate (at least some); plants of south central and southwestern Utah, and of canyons of the Colorado 3 Leaves entire; plants of eastern Utah 4 Leaves only sparingly serrate, linear-oblanceolate to elliptic; involucral bracts shortly attenuate, short-villous dorsally; plants of north central Kane County .... X. cronquistii Leaves sharply serrate-dentate, narrowly oblanceolate, elliptic, oblong, or lan- ceolate; involucral bracts long-attenuate, glandular or villous-pilose dorsally; plants of canyons of the Colorado and southwestern Utah X. tortifolia Peduncles mainly less than 5 cm long; stems usually leafy to much above the middle X. glabriuscula Peduncles mainly more than 5 cm long; stems usually to the middle or below ... X. venusta Xyhrhiza confertifolia (Cronq.) T.J. Wat- son [Macliaeranthera glabriuscula var. con- fertifolia Cronq., type from NE of Henrie- ville]. Perennial herbs from a woody caudex and taproot, with rootstocks sometimes de- veloped; stems 9-23 cm high, sparingly pi- lose to glabrate and sparingly to densely stipitate-glandular; leaves 1-4.5 cm long, 1-2.5 mm wide, linear, pilose to glabrate, commonly involute; peduncles 1.8-14 cm long; involucres 9-12 mm high, 12-18 mm wide; bracts lanceolate to lance-acuminate, pilose to glabrate and glandular; rays 4-12, white, 9-18 mm long, 2-4 mm wide; disk flowers yellow, the corollas 6-9 mm long; pappus of capillary bristles to 6.5 mm long; achenes 3.5-6 mm long, pubescent. Salt desert shrub and pinyon-juniper communities at 1675 to 1985 m in Garfield and Kane counties; endemic; 6 (i). Xylorhiza cronquistii Welsh & Atwood in Welsh Cronquist Woody-aster. Subshrubs, forming rounded clumps, from a stout tap- root; stems numerous, whitish, ca 30 cm tall, villous at the nodes, almost glabrous other- wise; leaves 2.5-5 cm long, 2.5-5 mm wide, linear-lanceolate, sparingly serrate-dentate to entire, sparsely villous, ciliate, the midrib prominent; heads solitary on branches; in- volucre 10-12.5 mm high, 13-19 mm wide; bracts oblanceolate to lance-attenuate, acute to acuminate, herbaceous above the middle, chartaceous below, short-villous and glandu- lar dorsally; rays white, 14-16, 20-25 mm long; achenes compressed, villous; pappus of capillary bristles to 7.2 mm long. Pinyon- juniper community, on the Kaiparowits For- mation, at 1890 to 2075 m in Kane County; endemic; 1 (0). Xylorhiza glabriuscula Nutt. Subshrubs or suffrutescent perennial herbs from a woody caudex and taproot; stems 7-37 cm tall, vil- lous to glabrous; leaves 1-7.5 cm long, 1-9 mm wide, villous to glabrate, lanceolate to narrowly lanceolate or oblanceolate; heads solitary at branch ends; involucres 9-13 mm high, 15-27 mm wide; bracts lanceolate, at- tenuate to acute or acuminate, herbaceous above the middle, scarious below, villous to glabrous; rays 10-22, white to bluish or purplish, 11-20 mm long; achenes com- pressed, villous; pappus of capillary bristles to 5 mm long. Two allopatric varieties are present. Leaves with attenuate bases; rays white; plants of Daggett County X. glabriuscula var. glabriuscula Leaves with truncate or rounded bases; rays bluish, purplish, or white; plants of San Juan Coimty X. glabriuscula var. linearifolia April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 357 Var. glabriuscula [Aster glabriuscula (Nutt.) T. & G.; Machaeranthera glabriuscula (Nutt.) Cronq. & Keck]. Salt and mixed desert shrub communities at ca 1525 to 2135 m in Daggett County; Colorado, Montana, South Dakota, and Wyoming; 0 (0). Var. linearifolia T.J. Watson Salt desert shrub community, mainly on Chinle and Moenkopi formations, in Grand and San Juan counties; endemic; 3 (iii). Xyhrhiza tortifolia (T. & G.) Greene Sub- shrubs; stems 15-50 cm tall or more, villous or tomentose and more or less stipitate- glandular; leaves 1-10 cm long, 4-20 mm wide, lanceolate to elliptic or oblanceolate, villous to tomentose and glandular, spinulose- dentate; heads terminating branches; in- volucres mainly 12-20 mm high and 15-30 mm wide; bracts narrowly lance-attenuate to -acuminate, herbaceous above, scarious be- low; rays 17-60 or more, bluish or purplish to white, 10-33 mm long, 1.8-5.5 mm wide; pappus of capillary bristles to 9 mm long; achenes compressed, pilose. Two varieties are present. 1. Involucres merely glandular dorsally; plants of canyons of the Colorado X. tortifolia var. imberbis — Involucres villous-pilose as well as glandular; plants of Washington County X. tortifolia var. tortifolia Var. imberbis (Cronq.) T.J. Watson [Mach- aeranthera tortifolia var. imberbis Cronq.]. Blackbrush, pinyon-juniper and sagebrush communities at 1220 to 2290 m in Garfield, Grand, Kane, San Juan, and Wayne counties; Arizona (Colorado canyons endemic); 32 (viii). Var. tortifolia [Haplopappus tortifolius T. & G.; Aster abatus Blake; Machaeranthera tortifolia (T. & G.) Cronq. & Keck]. Black- brush and other warm desert shrub commu- nities at 760 to 1010 m in Washington Coun- ty; Arizona, Nevada, and California; 10 (i). Xylorhiza venusta (Jones) Heller [Aster venustus Jones, type from Cisco]. Suffrutes- cent to herbaceous perennial herbs from a woody caudex and taproot; stems mainly 10-40 cm tall, glabrous to densely pilose; leaves 2.4-9 cm long, 2-17 mm wide, oblan- ceolate to spatulate, villous to glabrate, at- tenuate basally; heads terminating branches; peduncles 5-20 cm long; involucres 10-18 mm high, 18-50 mm wide; bracts lance-at- tenuate to caudate-acuminate, herbaceous above, scarious below; rays 12-36, white or bluish to purplish, 12-27 mm long; pappus bristles to 10 mm long; achenes sericeus. Salt desert shrub communities at 1250 to 1985 m in Carbon, Daggett, Emery, Garfield, Grand, San Juan, Uintah, and Wayne counties; Colo- rado (a Colorado Plateau endemic); 99 (xv). HAPLOPAPPUS CRISPUS AND H. ZIONIS (ASTERACEAE): NEW SPECIES FROM UTAH Loran C. Anderson' Abstract. — The new species, Haplopappus crispus and H. zionis of section Macronema, are formally described and illustrated. They are endemic to southern Utah. Also, H. bloomeri ssp. compactus is raised to species. Chromo- .some numbers of all three are n = 9. Aspects of anatomy are detailed. Comparisons are made to H. bloomeri and H. suffntticosus. Relationships are discussed, and a key to the species is given. The only comprehensive monograph of Haplopappus is that of H. M. Hall (1928). In recent years the generic integrity of Hap- lopappus has been questioned (see Anderson 1980 for review). Data from anatomy, cy- tology, and chemistry suggest it is a poly- phyletic assemblage. Nevertheless, a suitable taxonomic reorganization of the group has not been achieved. Therefore, I choose to de- scribe new taxa under the name Haplopappus even though the species will very probably be placed in some other genus at a later date. Some years ago I found plants in southern Utah that appeared to be H. bloomeri Gray ssp. compactus Hall; a chromosome count for the collection, Anderson 3358, was published under that name (Anderson et al. 1974). Other collectors had identified similar plants as H. suffruticosus (Nutt.) Gray. The com- bined collections actually represent two new species of section Macronema, and, addition- ally, H. bloomeri ssp. compactus should be elevated to species level. Methods and Materials Fresh and dried materials were processed for anatomical study as in Anderson (1970a). Five heads from personal collections, along with one to five heads from other collections, were measured (as in Anderson 1964) for in- volucral and floral data. Cytological methods are those of Anderson (1966). Voucher specimens for anatomical (a) and morphological (m) studies are: H. bloomeri: Anderson 1620, m (FSU), Anderson 2018, a and m (FSU), Anderson 2943, a (FSU), Ander- son 4539, m (FSU), Reveal 1070, m (FSU); H. compactus: Ackerman 30797, m (FSU), An- derson 6186, a and m (FSU), Clokey 8570, m (UTC); H. crispus: Anderson 5504, a and m (FSU), Cottam 1526, m (BRY), Maguire 13386, m (UTC), Stanton in 1927, m (BRY); H. suffruticosus: Anderson 1023, m (FSU), Anderson 2920, a (FSU), Anderson 2970, a and m (FSU), Goodrich 10133, m (BRY), Shultz 3738, m (UTC), Yoder-Williams 1311, a (FSU), Wiggins 9298, m (UTC); and H. zi- onis: Anderson 3358, m (FSU), Anderson et al. 5094, a and m (FSU), Arnow 107, m (UT). Taxonomy Haplopappus compactus (Hall) L. C. Ander- son, comb. nov. Basionym: Haplopappus bloomeri Gray ssp. compactus Hall. Carnegie Inst. Publ. 389:199, fig. 68, 1928. Type. - Nevada: Clark Co., Charleston [Spring] Mountains, E. C. Jaeger on 12 Sep 1925 (holotype: POM!; isotype: UC!). Synonymy: Haplopappus bloomeri Gray var. compactus (Hall) Blake in Clokey. Univ. Calif. Publ. Bot. 24:231. 1951. The only description of this taxon is that given by Hall (1928). It is amplified and emended with the following: woody shrubs to 5 dm tall; leaves oblanceolate-spatulate, 2-3.5(4) cm long, (2)2.5-3(5) mm wide; heads (involucres) 12-14.8 mm long, 4.5-5.5 mm wide, phyllaries 18-24; disk flowers 11-16, golden yellow (fading lighter), corollas 9.1-11.4 mm long, tubes glandular, strongly dilated at point of staminal departure (at 50 percent of total corolla length), lobes 0.8-1.6 mm long, styles 12.5-14.5 mm long. 'Department of Biological Science, Florida State University, Tallahassee, Florida 32306. 358 April 1983 Anderson: New Haplopappus 359 In addition to specimens cited by Hall (1928) and Clokey (1951), the following rep- resent this species: NEVADA: Clark Co., above McWilliams Campground at 8500 ft, Lee Canyon, Spring Mountains, L. C. Ander- son 6186 (BRY, FSU, UNLV, UTC), A. D. Blauer & £. D. McArthur N-144, N-145 (FSU, SSLP), W. E. Niles 3160 (FSU, UNLV); top of ridge, Charleston Mountain, 11,500 ft, T. Dawson 7 (UC); top of Sheep Mountains, 9300 ft, T. L. Ackemian 30797 (FSU, NY). The last collection is a range extension for H. compactus; the taxon was previously thought to be endemic to the Spring (or Charleston) Mountains. Haplopappus crispus L. C. Anderson, sp. nov. Frutices lignosi 3-4(5) dm alti; caules fo- liosi, graciles, erecti, tenuiter glandulosi; folia spatulata vel oblanceolata et acuminata, mar- ginibus undulatis et crispis, (1.5)2-2.5(30) cm longa, (3)5-6(8) mm lata, in ramis altioribus solum parvo minora; inflorescentia cyma so- lute paniculata vel aliquanto congesta, tem- poribus paucis solum uno capitulo vel duobus per ramum; capitula campanulata, (12.5)13-14(15) mm longa, (5)6.5-7(9) mm lata, foliis superioribus saepissime separata; phyllaria (24)26-30(35); nuUi radii flosculi; disci flosculi (14)15-20(24), luridi, coroUis (9.5)10-10.8 mm longis, tubulo aliquanto di- lato, lobis 1-1.5 mm longis, stylis 14-18 mm longis, lineis stigmaticis multo brevioribus quam appendicibus; achenia 6.5-8.5 mm longa et raro pubescentia. Type.— Utah: V^ashington Co., weathered andesite with manzanita in mountain mahog- any-fir woods with few pine and aspen along Whipple Valley Trail at 8100 ft, above Pine Valley, 19 air mi NE of St. George in Pine Valley Mountains, T39S, R14W, El/2 Sec 29, 18 Sep 1981, L. C. Anderson 5504 (holo- type: BRY; isotypes: FSU, MO, NY, RSA, UC, UTC). Woody shrubs, much branched at base, 3-4(5) dm tall (or taller?); leafy stems slender, erect, covered with short-stalked glands; leaves entire, alternate, green, spatulate to oblong-oblanceolate, acuminate, margins wavy-crispate, glutinous with low glands (not prominently stalked as in H. suffruticosus), (1.5)2-2.5(3) cm long, (3)5-6(8) mm wide. Fig. 1. Haplopappus crispus (Anderson 5504): a., flowering branch with crispate leaves and rather naked peduncles; b., involucral bracts, outermost with large green tips, inner ones with narrowed tips, innermost without green tips; c, disk flower with gradually flaring corolla and sparsely strigose achenes. A., x 1; b. and c, x 4. only slightly reduced toward the in- florescence, but generally not crowding the heads as in H. zionis; inflorescence a loosely paniculate to somewhat congested cyme, oc- casionally reduced to one or two heads per branch (Fig. la); heads campanulate, (12.5)13-14(15) mm long, (5)6.5-7(9) mm wide; phyllaries (24)26-30(35), 1-nerved, finely glandular, outermost bracts mostly green, leaflike, inner bracts stramineous, abruptly or gradually narrowed into a slender green tip, innermost bracts stramineous, erose-ciliate apically, but not as pronounced as in H. zionis; ray flowers absent; disk flow- ers (14)15-20(24), pale yellow, corollas (9.5)10-10.8 mm long, tubes glandular pu- bescent, not abruptly dilated as in H. com- pactus and H. zionis, lobes 1-1.5 mm long, 360 Great Basin Naturalist Vol. 43, No. 2 styles 14-18 mm long, branches slender, stig- matic lines much shorter than appendages (22-36 percent of total branch length); achenes cylindric, 6.5-8.5 mm long, sparsely but evenly pubescent, pappus 8-9.5 mm long; n = 9. Infrequent, generally in moder- ately open settings in spruce-pine associ- ations, (3000?) 8100-10,000 ft, apparently en- demic to mountains of southwestern Utah. August- September . Additional specimens examined.— Utah: Millard Co., Pine Valley, W. D. Stanton in 1927 (BRY); Washington Co., south end Pine Valley Mountains, 9200 ft, B. Albee 2911a (UT), Anderson Valley area, north end Pine Valley Mountains, 8400 ft, B. Albee 2911b (UT), Pine Valley Mountains, T39S, R14W, /. L. Gentry & E. Jensen, Jr., 2245 (BRY, NY, UTC), R. K. Gierisch 542 (UC, UTC), Pine Valley Mountains, 8500 ft, W. P. Cottam 5699 (UT), 10,000 ft, B. Maguire & B. L. Richards, Jr., 13386 (UC, UTC), Santa Clara, 3000 ft, W. P. Cottam 1526 (BRY, NY, UT). The last collection is so far out of range alti- tudinally that it may represent a chance in- troduction; the population has not been found again to determine its persistence. The collection is of further interest for the nota- tion "along ditch banks, 8-10 ft tall"; also, it has heads with greater numbers of phyllaries and flowers than the other collections. Haplopappus zionis L. C. Anderson, sp. nov. Frutices humiles et diffundentes 1-3 dm alti; corpulenti caules decumbentes vel as- cendentes, foliosi in inflorescentiam et glandulosi-hispiduli; folia spatulata et acumi- nata, (2.5)3-3.5(4) cm longa, 2.4-4(7) mm lata; inflorescentia cyma paniculata et con- gesta et foliosa; capitula turbinata vel ang- uste campanulata, (14)16-19(22) mm longa, 6-8 mm lata, bracteis exterioribus in forma folii apicibus longis et acuminatis, interior- ibus apice villosis, phyllariis (17)20-22(23); nulli radii flosculi; disci flosculi (10)12-18(21), flavi, corollis (9.5)9.8- 10.8(11.5) mm longis, tubulis subito dilatis, lobis 1.2-2 mm longis, stylis 18-19 mm long- is, lineis stigmaticis multo brevioribus quam appendicibus; achenia 7-8 mm longa et ferme glabra. Type.— Utah: Iron Co., moderately bare, weathered pink limestone member of Wasatch Formation in aspen, spruce, limber pine, bristlecone pine association near top of Cedar Canyon, 9800 ft, 13.5 air mi SE of Cedar City, 30 Aug 1980, L. C. Anderson, S. L. Welsh, and M. Chatterley 5094 (holotype: BRY; isotypes: ASU, DS, FSU, MO, NY, RSA, UC, UTC). Low, branching, spreading shrubs, 1-3 dm tall; stems stout, decumbent or ascending, glandular-hispidulous, leafy into the in- florescence (Fig. 2a); leaves entire, alternate, green, spatulate, acuminate, margins not un- dulate or crispate, glandular but less glu- tinous than H. crispus, (2.5)3-3.5(4) cm long, 2.5-4(7) mm wide; inflorescence a congested, leafy paniculate cyme; heads turbinate to narrowly campanulate, (14)16-19(22) mm long, 6-8 mm wide, lateral heads fewer flow- ered than central ones; phyllaries (17)20-22(23), 3-nerved, finely glandular, outermost bracts leaflike and much longer than other bracts, inner bracts stramineous with long green tips, innermost bracts stra- mineous with praemorse, prominently villous tips (Fig. 2b); ray flowers absent; disk flowers (10)12-18(21), golden yellow, corollas (9.5)9.8-10.8(12) mm long, tubes abruptly di- lated at point of staminal departure (at 40 percent of total corolla length), sparsely glandular, lobes 1.2-2 mm long, styles 18-19 mm long, branches slender, stigmatic lines much shorter than appendages (23-31 per- cent of total branch length); achenes cylin- dric, 7-8 mm long, essentially glabrous with few hairs apically (Fig. 2c), pappus 9-9.5 mm long; n = 9. Infrequent, usually on gravelly sandy clay from limestones, often with man- zanita on rather barren slopes in spruce, fir, pine associations, 8,000-10,000 ft, apparently endemic to mountains of southern Utah. July- September. Additional specimens examined: Utah, Garfield County, vie. summit between Esca- lante and Widtsoe, Escalante Mountains, 9000 ft, W. P. Cottam 6562 (UT), 9100 ft, L. C. Anderson 3358 (BRY, FSU, KSC, NY, UC, UTC), 10,000 ft, E. Neese & S. White 3964 (BRY), 2 miles south of Pine Lake, Escalante Moimtains, E. Neese & S. White 3844; Iron County, Cedar Canyon, southeast of Cedar City, 8000 ft, L. Arnow 107 (UT), R. Foster 5297 (BRY), vie. Midway Summit, west of Cedar Breaks, 10,000 ft, B. Maguire & B. L. April 1983 Anderson: New Haplopappus 361 Fig. 2. Haplopappus zionis (Anderson et al. 5094): a., flowering branch with foHage overtopping flowering heads; b., involucral bracts, outermost leaflike and 3- nerved, inner ones with elongate green tips, innermost with prominently villous tip; c, disk flower with abrupt- ly dilated corolla tube, long style, and nearly glabrous achene with very few hairs just below pappus, a., x 1; b. and c, X 4. Richards, Jr. 13389 (UTC); Co. unknown, Dixie National Forest, R. K. Gierisch 217 (UTC). Cytology and Anatomy Chromosome numbers of section Mac- ronema of Haplopappus are monotonously uniform at n = 9 with an apparent absence of polyploidy and aneuploidy (Anderson et al. 1974). The new species are also n = 9. Original counts for H. compactus and H. crispus were made from root tip squashes from Niles 3160 and Anderson 5504, respec- tively. An earlier count for H. zionis (Ander- son 3358) was listed as H. bloomeri ssp. com- pactus in Anderson et al. (1974). Meiotic behavior has not been observed for any of the new species, but all three have pollen fertilities above 98 percent (as deter- mined by staining in cotton blue in lactophe- nol). Embryo sac development appears nor- mal (Polygonum type) in H. compactus, H. crispus, and H. suffruticosus; it was not stud- ied in H. bloomeri and H. zionis. Mature em- bryo sacs are long and narrow (280-380 jam long) and 8-nucleate without multiplication of antipodals as found in many Chryso- thamnus (Anderson 1970b). Aspects of floral anatomy for the new spe- cies and possibly related taxa are summarized in Table 1 (following format in Anderson 1970a); frequency categories are: -I- + , abun- dant; -I- , frequent; -, rare; and 0, absent. General information on the other taxa of Haplopappus is in Nelson (1982). Achenes of H. zionis have the greatest amount of vas- culature, whereas those of H. crispus have the least. Secretory canals vary from abun- dant to absent at selected levels in the ach- enes and corollas. They are always absent in the styles of these species but present in H. macronema (Anderson 1970a). Achenes of H. bloomeri, H. compactus, and H. zionis are essentially glabrous but with a few twin hairs just below the pappus attach- ment (the last being the "most glabrous"). A few short glandular trichomes (60-66 jum long) are hidden among the abundant twin hairs on H. suffruticosus achenes, and larger glandular hairs (190 jum) occur rarely on achenes of H. bloomeri. Glandular trichomes on achenes were so rare that they were omitted from Table 1, and they do not occur in the new species. Pappus bristles are posi- tioned in three closely spaced rows on ach- enes of H. zionis and in two in the other four 362 Great Basin Naturalist Vol. 43, No. 2 Table 1. Comparisons for range and aspects of morphology of selected taxa oi F Haplopappus , section Macronema. H. bloomeri H. compactui ( H. crispus H. suffruticosus H. zionis Range Calif., W Nev., SNev. SW Utah Calif., Nev., ] [daho. SUtah Oregon, S Wash. W Wyo., SW Mont Elevation, ft 3500-9600 7900-11,500 (3000)8100-10,000 7500-12,300 8000-10,000 Habit of growth tall, 4-5 dm tall, 4-5 dm medium, 3-4 dm low, 1-3 dm low, 1-3 dm Leaf size, cm x mm 2-6 X 0.5-3 2-4 X 2-5 1.5-3 X 3-8 1-3 X 1.5-5 2.5-4 X 2.5-7 Leaf shape filiform-narrowly oblanceolate oblanceolate- spatulate ■ spatulate- oblanceolate oblong-spati ilate spatulate Leaf margins smooth smooth crispate crispate smooth Head length, mm 7.8-12.2(16) 12-15 12.5-15 8-16 14-22 Head width /length ratio .29- .35 .36-.40 .45-.48 .45-.60 .38-.41 Phyllary number 17-36 18-24 24-35 17-31 17-23 Ray flower number (0)1-3 0 0 (0)1-8 0 Disk flower number 4-8(12) 11-16 14-24 15-40 10-21 Disk flower length, mm 7.3-9.4(10.9) 9.1-11.4 9.5-10.8 8.5-11 9.5-12 Corolla tube straight dilated ± dilated straight dilated Corolla lobe length, mm 0.9-1.9 0.8-1.6 1.0-1.5 1.0-1.7 1.2-2.0 Style branch, mm 2.5-4.3 3.8-4.4 3.8-4.8 2.6-4.6 4.2-5.2 Achene length, mm 6-7 7.5-8.5 6.5-8.5 7.5-8 7-8 Achene pubescence glabrous glabrous sparsely strigose villous-sericeus glabrous species (contrary to the meaning of the ge- neric name). The five species have glandular corolla tubes with the trichomes, "g" in Table 1, being similar to those in Chrysothamnus (fig. 20-22, Anderson 1970a); however, those in H. zionis are less abimdant, are longer, and have narrower glandular heads as in Figure 19. Additionally, corollas of H. crispus have a few very long, nonglandular villi like Fig. 17 of Anderson (1970a). Nectaries at the base of the style are most prominent in H. zionis and least so in H. crispus. The new species have somewhat thicker corollas than do H. bloomeri and H. suffruticosus as determined by cell number radially through the tubes between adjacent vascular bundles. Nodal anatomy is trilacunar, three trace for the species. In leaf anatomy, they are sim- ilar in that leaves are isolateral with three rows of palisade cells facing each epidermis. Bundle sheath extensions are present only on the midvein— a feature characteristic of all taxa of Haplopappus with a chromosome base of X = 9, whereas those with x = 4, 5, or 6 have leaves with bimdle sheath exten- sions on lateral veins as well. A possible ex- ception would be H. parryi, (x = 9 group), which has many bundle sheath extensions on bifacial leaves, but it is better placed in Sol- idago (Anderson and Creech 1975). Some variation exists in leaf thickness. Of leaves studied, those of H. compactus and H. zionis have blades averaging 336 jum thick with midveins 480 and 350 jum, respectively. Leaves of H. crispus are thicker, 355 /xm in the blade and 400 at the midvein. Much thin- ner leaves occur in H. suffruticosus with blades 202 and 250 {Anderson 2970 and 2920, respectively), and both with midveins about 260 jLim thick. Glandular trichomes on leaves of H. com- pactus and H. crispus average 77 jum long with nearly spherical heads 50 jum and 38 /xm wide, respectively. Glands on H. zionis Table 2. Features of floral anatomy for selected taxa of Haplopappus, section Macronema. Ovarian bundle number Secretory canal distribution Achene Corolla Taxon Average Range I II III IV V H. bloomeri H. compactus H. crispm H. suffruticosus H. zionis 5.5 5.5 5.0 5.2 7.0 5-6 5-6 5 5-6 5-9 + -1- + + + + + + -1- + + + + + + + + + + 0 -1- + + + + + + + + + + + + + + April 1983 Anderson: New Haplopappus 363 leaves, thoughsomewhat less abundant, have larger buttressed bases and average 100 /xm long with heads 38 jum wide. For H. suffruti- cosus, glands on Anderson 2970 leaves aver- age 80 jum long with spherical heads, whereas those on Anderson 2920 have long slender stalks and heads, averaging 220 jum long with the elongate heads 48 jum long and 29 jum wide. Glands on H. hhomeri are nearly ses- sile, only 38-42 ju,ni long with spherical heads 29 jLim in diameter. Relationships Table 2 gives ranges of the species and summarizes some aspects of their morphol- ogy based on my observations; some mea- surements differ from those of Hall (1928). Reveal 1070 is an unusual form of H. bhom- eri, with exceptionally large heads with many long disk flowers (features listed paren- thetically in Table 2). The new species do not occur close geo- graphically to H. hhomeri or H. suffruti- cosus, species with which they have been confused. Further, the new species are dis- tinct from the other two in their constant ab- sence of ray flowers. Ray flowers are almost always present in heads of H. bloomeri and H. suffruticosus, and, although occasionally in- dividual plants may be eradiate, some plants of the population will always have rays. Haplopappus compactus, once considered a subspecies of H. bloomeri, differs from that species in involucral length and width, disk flower number, corolla shape, and achene size as well as the absence of ray flowers and some aspects of leaf shape and size. Hap- lopappus compactus and H. zionis appear to be closely related. Both have dilated corolla tubes. They can be distinguished in habit, head size, and style length, and generally so in leaf shape and size. In addition, H. zionis usually has leafier involucres. The two also differ anatomically. Haplopappus crispus is intermediate geo- graphically between H. compactus and H. zi- onis, but it is closer to H. suffruticosus in relationship even though it is eradiate. In ad- dition to absence of rays, H. crispus can be further distinguished from H. suffruticosus by its habit, leaf size and thickness, degree of glandularity (and odor), narrower heads, few- er flowers, and sparsely strigose achenes. A key to the new species and related taxa is presented here. 1. Ray flowers present (at least in the population); widespread but not in S Nevada or S Utah 2 — Ray flowers always absent in each head; plants of S Nevada or S Utah 3 2(1). Achenes densely pubescent (sericeus); heads campanulate with more than 15 flowers H. suffruticosus — Achenes essentially glabrous; heads cylindric-turbinate with less than 15 flowers H. bloomeri 3(1). Achenes sparsely strigose with hairs distributed evenly; leaves crispate .. H. crispus — Achenes essentially glabrous but with few hairs near the pappus; leaves not crispate 4 Table 2 continued. Trichomes Corolla thickness Nectary length, jum Achene Corolla tube Index of Freq. Length, mm Type Freq. Length, jLtm A B C specialization 144 - 0.5 g + + 150 7 5 2 6.5 168 - 0.43 g -1- 168 8 5 2 5.6 115 + 0.53 g n -1- -1- 165 432 7 6 2 6.8 140 + + 0.8 g + + 160 7 5 2 6.9 240 - 0.52 g - 270 8 6 2 5.0 364 4(3). Great Basin Naturalist Vol. 43, No. 2 Heads mostly less than 14 mm long; styles less than 15 mm long; woody shrubs 3-5 dm tall H. compactus Heads mostly more than 14 mm long; styles over 17 mm long; low shrubs 1-3 dm tall H. zionis In the Asteraceae, amounts of secretory tis- sue and floral venation bear evolutionary sig- nificance with a reduction in both usually in- dicating advancement. In Chrysothamnus, these features (correlated with other charac- teristics such as karyotypes) were used to de- velop a phylogenetic index of specialization on a scale of 0-10 for the taxa (Anderson and Fisher 1970). Section Macronema is close to Chryso- thamnus with natural hybridization known to occur between H. macronema and C. nau- seosus (Anderson and Reveal 1966). There- fore, the index of specialization developed for Chrysothamnus should be applicable for showing relationships among the new species. Index values are listed in Table 2. Reduction in involucral bract vasculature is apparently correlated with that of the ovary wall; phyl- laries of H. zionis have three vascular bundles each; those of H. crispus have one. Of the taxa studied here and in Anderson (1970a), H. macronema (the most widespread species in the section) has the lowest index number of 4.5, and H. ophitidus (clearly a de- rived species that is specialized for serpen- tine) has the highest at 8.5. Hall (1928) con- sidered H. suffruticosus the most primitive member of section Macronema, but H. mac- ronema is probably a better candidate. Hap- hpappus compactus, and, especially, H. zion- is, though much more restricted in range than H. bloomeri and H. suffruticosus, seem- ingly represent more primitive stock phy logenetically . Acknowledgments Appreciation is expressed to Stanley L. Welsh for field trip support and to Wesley Niles for seed of H. compactus. Melanie Darst prepared the line drawings; Walter Forehand is thanked for assistance with the Latin diagnoses. This study was supported by National Science Foundation Grant DEB 76-10768. Literature Cited Anderson, L. C. 1964. Taxonomic notes on the Chryso- thamnus viscidiflorus complex (Astereae, Com- positae). Madrono 17:222-227. 1966. Cytotaxonomic studies in Chrysothamnus (Astereae, Compositae). Amer. J. Bot. 53: 204-212. 1970a. Floral anatomy of Chrysothamnus (As- tereae, Compositae). Sida 3:466-503. 1970b. Embryology of Chrysothamnus (Astereae, Compositae). Madroiio 20:337-342. 1980. Haplopappus alpinus (Asteraceae): a new endemic species from Nevada. Great Basin Nat. 40:73-77. Anderson, L. C, and J. B. Creech. 1975. Comparative leaf anatomy of Solidago and related Asteraceae. Amer. J. Bot. 62:486-493. Anderson, L. C, and P. S. Fisher. 1970. Phylogenetic indicators from floral anatomy in Chrysothamnus (Astereae, Compositae). Phytomorph. 20: 112-118. Anderson, L. C, and J. L. Reveal. 1966. Chryso- thamnus bolanderi, an intergeneric hybrid. Ma- drono 18:225-233. Anderson, L. C, D. W. Kynos, T. Mosquin, A. M. Powell, and P. H. Raven. 1974. Chromosome numbers in Compositae. IX. Haplopappus and other Astereae. Amer. J. Bot. 61:665-671. Clokey, I. W. 1951. Flora of the Charleston Mountains, Clark County, Nevada. Univ. California Publ. Bot. 24:1-274. Hall, H. M. 1928. The genus Haplopappus, a phyloge- netic study in the Compositae. Carnegie Inst. Publ. 389:1-391. Nelson, J. B. 1982. Floral anatomy of North American Haplopappus (Asteraceae): systematic consid- erations. Unpublished dissertation. Florida State Univ. Library, Tallahassee. A BOUQUET OF DAISIES (ERIGERON, COMPOSITAE) Stanley L. Welsh' Abstract. — Described as new are six critical species and one variety of Erigeron from Utah. They are E. awa- pensis Welsh, £. canaani Welsh, E. carringtonae Welsh, £. goodrichii Welsh, E. maguirei Cronq. var. harrisonii Welsh, E. tintemuinnii Welsh and Goodrich, and E. wahwahensis Welsh. Erigeron is perhaps the most complex of all genera within the Compositae of Utah. The genus is allied to both Aster and Conyza, but most of the problems lie within the genus it- self. To make matters more difficult Erigeron is one of the largest genera in Utah, with more than 60 taxa. Morphological features used as diagnostic characters often involve pubescence and its position on the plants. Other vegetative features have been relied upon also, because of the uniformity of flow- ers and bracts. Shape of leaves, whether mainly basal, mainly cauline, or distributed along the stem constitute important diagnos- tic characters. Pubescence type on the in- volucral bracts is considered as diagnostic in many cases. The technical nature of the dis- tinguishing characters have led to wide mis- understanding of the genus, and most bota- nists have avoided the members of the group, relying, where possible, on specialists for crit- ical determinations. Preparation of a manuscript of the Com- positae for the Utah flora project has led me to study Erigerons of Utah and the surround- ing states in some detail. That study has in- dicated the presence of several taxa whose descriptions appear to be beyond those pre- viously known to occur within Utah. The specimens have been compared to all other materials at BRY, and duplicates of many have been examined by Arthur Cronquist at NY. I wish to acknowledge his suggestions and consideration, but wish not to share blame for problems created by this author alone in the following interpretations. Erigeron awapensis Welsh, sp. nov. A E. abajoensis Cronq. in caulis erectis foliorum basalis non vaginatis et floribus plus numer- osis differt. Perennial herbs from a branching caudex, the caudex branches clothed with ragged brown marcescent leaf bases; stems erect or nearly so, 10-24 cm long, strigose, the hairs ascending; basal leaves 1.5-7 cm long, 2-8 mm wide, not especially sheathing; cauline leaves well developed, oblong to linear, most- ly 1-4 cm long, 2-4 mm wide; heads 2-4, rarely solitary; involucres 3-9 mm wide, 3.7-4.5 mm high, the bracts more or less im- bricate, thickened near the base dorsally, greenish, strigulose, the hairs multicellular; rays 35-45, pink purple to pink (or white?), 5-6 mm long, 0.9-1.8 mm wide; pappus ap- parently simple, of 15-20 slender bristles, and with a few inconspicuous shorter setae in some; achenes 2-nerved, hairy. Pinyon-juni- per and sagebrush communities at 2135 to 2260 m in Garfield and Wayne counties; endemic. Type.- USA. Utah. Garfield County, T32S, R2W, S23, igneous bouldery slope in canyon, in Artemisia community, at milepost 26.5 south of Antimony, 4 September 1969, S. L. & S. L. Welsh 9388 (Holotype BRY; Iso- types, 4 distributed previously as E. divergens T. & G.). Additional specimen: Utah. Wayne County, T29 S, R4E, S20, 1 mi SE of Teas- dale, sandy slope in pinyon-juniper commu- nity, 7 August 1980, M. E. Lewis 6657 (BRY). The Awapa daisy is most similar to E. aba- joensis, but stands apart from that taxon, which has become a catchall for all speci- mens that run to the end of the key. It seems apparent that the assemblage will be clarified only upon examination of much additional material not now available in collections. 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 365 366 Great Basin Naturalist Vol. 43, No. 2 Erigeron canaani Welsh, var. nov. Similis E. eatonii Gray in aspectum et capitulum amplitudem sed in foliis caulino et radi- calibus ambo linearibus et involutis folio basalibus ampliato et ciliato et radiis paucior- ibus differt. Perennial herbs from a simple (or branched?) caudex, this clothed with brown marcescent leaf bases, the taproot prominent; stems 7-20 cm tall, decumbent to ascending, sometimes pm-plish at the base, sparingly hairy with ascending hairs; leaves pubescent Uke the stem, the basal ones tufted, 1 -nerved, 1.4-9 cm long, 0.7-1 mm wide, linear, in- volute, sharply acute, conspicuously ex- panded and long-ciliate basally; cauline leaves numerous, reduced upwards; heads 1-3; involucres 9-13 mm wide, 5.3-6.5 mm high; bracts imbricate, conspicuously glandu- lar and sparingly to moderately villous-pilose with multicellular hairs, green or variously suffused with purple; rays 15-22, white or pinkish, 3.5-5 mm long, 1.8-2.1 mm wide; pappus single, of ca 20 slender bristles; achenes 2-nerved, hairy. Ponderosa pine community at 1585 to 2075 m in Washington County; endemic. Type.— USA. Utah. Washington County, with Petrophytum and Panicum, in crevices in Navajo Sandstone, Canaan Mountain, T43S, RlOW, S4, 11 June 1980, John Ander- son sn (Holotype BRY; Isotype NY). Addi- tional specimen: Utah. Washington County, Zion National Park, Clear Creek Canyon, T41S, RlOW, S24, ponderosa pine commu- nity, on Navajo Sandstone, 25 Sept. 1982, S. L. Welsh 21398 (BRY). The Canaan daisy is similar in general as- pect to E. eatonii. The involute linear glaucous leaves and few ray flowers appear to be diagnostic. Erigeron carringtonae Welsh, sp. nov. Af- finis E. untermannii in foliis crassis sed in pilis involucrorum longioribus et tenuioribus et radiis longioribus differt. Pulvinate perennial herbs with a plurici- pital caudex, the branches clothed with con- spicuous brown to straw colored or ashy mar- cescent leaf bases; leaves mainly basal, thickish, 0.6-3.5 cm long, 1-5 mm wide, spatulate to oblanceolate, strigose to pilosu- lose, obtuse to rounded apically; scapes 2.5-8 cm tall; heads solitary; involucres 9.8-15 mm wide, 5.8-7 mm high, the bracts imbricate, suffused with purple or green, the inner greenish with scarious margins, spreading-vil- lous with long multicellular hairs; rays 18-30, pink to pink purple, 6.8-8.2 mm long, 1.4-2.3 mm wide; pappus double, the inner of 25-35 barbellate bristles, the outer of short setae; achenes 2-nerved, pilose. Meadows and escarpment margins, commonly on Flagstaff Limestone at 3050 to 3355 m in Emery and Sanpete counties; endemic. Type.— USA. Utah. Sanpete County, T18S, R4W, S33, Manti-La Sal NF, 10 mi 115 de- grees from Manti, 3220 m, 3 July 1981, S. Goodrich 15780 (Holotype BRY; Isotypes 14, distributed as E. simplex). Additional speci- mens: Utah. Sanpete County. T20S, R4E, S33, ca 17 mi W of Ferron, 3355 m. Flagstaff Limestone barrens, 13 July 1977, S. L. Welsh and S. Clark 15393 (BRY); do, T18S, R4E, S22, 7 mi due NNW of Ferron Reservoir, 19 July 1977, E. Neese and S. White 3708 (BRY); do. Heliotrope Mountain, 13 July 1977, M. E. Lewis 4915 (BRY). Emery Coun- ty, Big East Mountain, head of Rilda Canyon, 13 July 1979, M. E. Lewis 6009 (BRY); do, Big East Mountain, above Upper Joes Valley, 8 August 1977, M. E. Lewis 5129 (BRY). The Carrington daisy has been identified with E. simplex by previous workers, but ap- pears to be more clearly allied to E. unter- mannii, at least superficially. It has long, spreading multicellular hairs similar to those of E. simplex, but the pulvinate caespitose habit and thick obtuse to rounded leaves of E. carringtonae appear to be diagnostic. The species is named in honor of the mem- ory of Jane Carrington, evidently the first woman to collect Utah plant materials desig- nated as types (Welsh 1982). Erigeron goodrichii Welsh, sp. nov. Similis et affinis E. clokeyi Cronquist sed in caudi- cibus tenuioribus et capitulis majoribus differt. Perennial herbs from a stout taproot and caudex, the caudex branches with dark brown marcescent leaf bases; stems 3-10 cm tall, de- cumbent-ascending to erect, spreading hairy; basal leaves 0.4-6 cm long, 1.2-5 mm wide, narrowly oblanceolate, the veins not appar- ent, pilosulose, obtuse apically; cauline leaves more or less developed, but much reduced upwards; heads solitary; involucres 10.5-18 April 1983 Welsh: Erigeron 367 mm wide, 6.4-7.8 mm high; bracts imbricate, spreading villous-pilose with multicellular hairs, thickened basally, green or the apices suffused purplish, the inner with scarious margins, the attenuate apices more or less glandular and sometimes spreading; rays 40-65, pink purple to pink or white, 6.8-10.4 mm long, 1.5-2 mm wide; pappus apparently single, of 20-30 minutely barbellate bristles; achenes 2-nerved, pilose. Engelmann spruce krummholz and meadow communities, often on rock outcrops or talus at 3050 to 3400 m in Duchesne, Summit, and Utah counties; endemic. Type.— USA. Utah. Duchesne County, T2N, R8W, S22, Ashley NF, Uinta Moun- tains, S rim of South Fork of Rock Creek, 14 mi n of Tabiona, 3264 m, with clumps of Engelmann spruce krummholz, on gravelly ground, 29 July 1981, S. Goodrich & R. Jep- son 15907 (Holotype BRY; Isotypes NY, CAS, POM, MO, US, UT, UTC, RM). Additional specimens: Utah. Duchesne County, divide between Log Hollow and Rock Creek, 12 mi 355 degrees from Tabiona, 31 July 1979, S. Goodrich 13550 (BRY); do, head of Log Hol- low, T2N, R7W, S30, 12 mi N of Tabiona, 3 July 1978, S. Goodrich and L. Hart 11696 (BRY); do, above Wedge Hollow, 11.5 mi and 354 degrees from Tabiona, 1 August 1979, S. Goodrich 13579 (BRY). Summit County, T2N, R12 E, S24, 11.5 mi NW of Kings Peak, East Fork of Blacks Fork, 31 August 1981, S. Goodrich 16203 (BRY). Utah County, Mt. Timpanogos, Emerald Lake, 7 August 1941, E. Castle 101c (BRY). This dwarf alpine species has been recog- nized as having affinities with E. asperu- gineus (D.C. Eaton) Gray and E. clokeyi Cronquist, each of them dwarf alpine species. The Goodrich daisy differs from the former by its more slender leaves and from the latter by the caudex which lacks the distinctive thatch of gray -brown marcescent leaf bases. The specific epithet honors the enthusias- tic collector of the type and other materials, Sherel Goodrich, 'student of western botany, whose energetic pursuit of Utah and Nevada plants is unsurpassed. Erigeron maguirei Cronq. var. harrisonii Welsh, var. nov. Similis var. maguirei sed in capitulis plus numerosis, radiis angustioribus, et disci brevioribus differt. Type.- USA. Utah. Wayne County, T29S, R6E, S14, ca 1 mi ENE of Fruita, ca 5700 ft elev., Navajo Sandstone, juniper community, 2 June 1982, S.L. & E.R. Welsh 21178 (Holo- type BRY; Isotypes NY; CAS). Additional specimens: Wayne County, canyon near Nat- ural Bridge, 6 April 1934, B.F. Harrison 7385 (BRY); do, Fruita (Hickman Bridge trail), 10 June 1938, D E. Beck s.n. (BRY). The harri- sonii phase of the E. maguirei differs in minor technical ways from materials of the type va- riety. Both phases are plants of sandy canyon bottoms, and perhaps they represent nothing more than ecological variants of a common theme. The variety is named in honor of its dis- coverer, Bertrand F. Harrison, collector, teacher, and student of Utah botany. Erigeron untermannii Welsh & Goodrich, sp. nov. Similis E. compactus sed in foliis la- tioribus pilis ascendentibus vel patentibus et radius brevioribus differt. Perennial pulvinate herbs with an in- tricately branched caudex, the caudex branches mainly basal, 0.8-3.3 cm long, 1-4 mm wide, narrowly oblanceolate to spatu- late, pilosulose with ascending, often curved, hairs; scapes 2-6 cm tall; heads solitary; in- volucres 7-11 mm wide, 5-5.7 mm high, the bracts more or less imbricate, green, or the inner somewhat chartaceous, the margins hyaline, the tips suffused with purple (some- times throughout), densely hispidulous with short spreading hairs; rays 14-26, white, 4-6.5 mm long, 1.5-2.1 mm wide; pappus apparently single, of ca 20 slender fragile- bristles; achenes 2-nerved, pilose. Pinyon- juniper community on calcareous shales and sandstones of the Uinta and Green River for- mations at 2135 to 2380 m in Duchesne and Uintah counties; endemic. Type.— USA. Utah. Duchesne Co., T5S, R6W, S22, Indian Canyon, 17 mi S Du- chesne, pinyon-juniper community, on Green River Shale, 4 June 1980, N. D. Atwood 7554 (Holotype BRY; Isotype NY). Additional specimens: Utah. Duchesne County, Tava- puts Plateau, Uinta Formation, junction of Right Fork and Left Fork of Indian Canyon, ca 10 mi SW of Duchesne, steep slopes and narrow ridge tops, shale and marly limestone, 26 May 1976, S. Goodrich 5317 (BRY); do, 13 mi 220 degrees from Duchesne, T5S, R6W, 368 Great Basin Naturalist Vol. 43, No. 2 S21, ridge between Right Fork and Left Fork of Indian Canyon, 31 May 1979, S. Goodrich 12402 (BRY). Uintah County, Vernal District, Ashley National Forest, dry ridge top, S rim of Red Pine Canyon, 5 June 1976, S. Good- rich 5652 (BRY). The Untermann daisy is compared to E. compactus Blake in the diagnosis, and it is probably allied to that taxon. However, the similarity to E. nematophyllus Rydb. cannot be ignored. It differs from that taxon in the broader leaves, generally harsher and more spreading hairs of the leaves, involucre, and achenes. The plant is named to honor the memory of the late G. E. and B. R. Unter- mann, an amazing husband and wife team who worked throughout their lives to under- stand the geology, natural history, and an- thropology of the Uinta Basin. They in- fluenced the lives of all persons whom they contacted. Erigeron wahwahensis Welsh, sp. no v. Herbis similis E. eatonii in aspectus sed ro- bustioribus caudicibus crassioribus pilis pat- entibus pro parte et bracteis crassioribus ad basim et dense vel sparse pilis patentibus differt. Perennial herbs, from a branching caudex, the caudex branches with conspicuous fibrous brown to ash-colored marcescent leaf bases; stems 15-40 cm long, decumbent to ascend- ing; basal leaves 3-18 cm long, 4-13 mm wide, linear-oblanceolate to oblanceolate or elliptic, 3-nerved, petiolate, appressed to spreading hairy with curved hairs; cauline leaves reduced, sessile, and bracteate above; heads solitary or 2 or 3; involucres 13-17 mm wide, 6-7 mm high, spreading-villous with multicellular hairs, glandular apically; bracts imbricate, green, the tips reddish, thickened basally; rays 30-40, pink or white, 5.5-7 mm long, 1.7-2.2 mm wide; disk corollas 3.5-4.2 mm long, the tube ca 2 mm long, the lobes 0.4 mm long; pappus of 15-20 bristles, with inconspicuous outer setae; achenes 2-nerved, short-hairy. Sagebrush, oak-maple, and pin- yon-juniper communities at 1670 to 2440 m in Beaver and Washington counties; endemic. Type.- USA. Utah. Beaver County, T28S, R15W, S31, Wah Wah Mountains, ca 12 mi SSW of Wah Wah Spring, Pine Grove Pass, 2450 m elev, sagebrush and pinyon-juniper community, 12 June 1982, S.L. Welsh 21229 (Holotype BRY; Isotypes NY, CAS, POM, RM, UT, UTC, US, GH, MO, and others to be distributed). Additional specimens: Utah. Beaver County, T29S, R16W, S12. Willow Creek, Wah Wah Mountains, 8 June 1978, K. Ostler & D. Anderson 1274 (BRY); do, T28S, R15W, S31, Wah Wah Mountains, divide be- tween Quartz Creek and Pine Grove Canyon, 22 May 1981, S.L. Welsh 20520 (BRY). The Wah Wah daisy is more or less inter- mediate between phases of E. eatonii and E. jonesii, but has features not shared by either. The pubescence is similar to E. jonesii, but the general aspect is more like E. eatonii. The plants tend to be larger than either, and the thick caudex appears to be diagnostic. Literature Cited Cronquist, a. 1947. Revision of the North American Species of Erigeron north of Mexico. Brittonia 6: 121-302. Welsh, S. L. 1982. Utah Plant Types— historical per- spective 1840 to 1981— annotated list, and bibli- ography. Great Basin Nat. 42:129-195. NEW TAXA IN THELESPERMA AND TOWNSENDIA (COMPOSITAE) FROM UTAH Stanley L. Welsh' Abstract. — Three new taxa are named and described: Thelesperma subnudum Gray var. alpinum Welsh, Towns- endia jonesii (Beaman) Reveal var. lutea Welsh, and Townsendia montana var. caelilinensis Welsh. Preparation of a manuscript of the sun- flower family for the Utah flora project dem- onstrated the existence of diversity within Thelesperma and Townsendia that seems best treated at taxonomic rank within existing species in those genera. In both genera the plants named are ecological specialists, with plants of each taxon occurring on calciferous or gypsiferous substrates of the Carmel, Flag- staff, Arapien, or other peculiar formations. Each of them belongs to the mound-forming group of substrate specialists of barren or semibarren habitats where water relations are controlled by the parent material. Generally the habitats are arid, despite the large amount of rainfall in the higher elevation sites. In all habitat examples cited there are other similarly restricted specialists, either species or varieties. Thelesperma subnudum Gray var. alpi- num Welsh, var. nov. Similis Thelesperma subnudo Gray var. subnudo sed in habito hu- milioribus et capitulis parvioribus differt. Perennial herbs from a taproot and less commonly with a caudex and creeping root- stock; stems 3-7 cm tall, subscapose; leaves mainly basal on the stem, 1.5-4 cm long, pin- nately lobed, or the upper entire; petioles ciliate and the blades puberulent; involucres 6.3-9 mm high, 9-14 mm wide; outer bracts oblong to lanceolate, with narrow scarious margins, to half as long as the inner; inner bracts united to below the middle; rays lack- ing; disk flowers yellow; pappus a toothed crown; achenes glabrous or hairy apically. Type.- USA. Utah. Wayne County, T28S, R4E, S13 (NEi/4;l. 3 mi due N of Bicknell, bristlecone pine forest on multicolored clay hills, 2745 m, 20 July 1980, D. Atwood and B. Thompson 7646a (Holotype BRY). Addi- tional specimen: Utah. Wayne County, S of Teasdale, 25 July 1978, D. Atwood 6924 (BRY). This dwarf phase of Thelesperma sub- nudum occurs about 500 m above the up- permost elevations known for the typical va- riety. Discoid plants are not uncommon for the widespread phase of the species, includ- ing at least a portion of the type (taken in Red Canyon, near Paragonah; isotype BRY), but the typical phase is ordinarily radiate. The dwarf alpine plants have involucres that are both shorter and narrower on the aver- age. The plants are apparently restricted to the Carmel Limestone, on the peculiar vari- colored phase of that formation as it occurs at the east margin of Rabbit Valley. Townsendia jonesii (Beaman) Reveal var. lutea Welsh, var. nov. Affinis T. jonesii var. jonesii sed in coroUis luteis ventralis differt et in substratis gypsiferis confinis. Subacaulescent to acaulescent caespitose herbs, the caudex commonly branched; stems not conspicuously white-strigose, mainly 2-3 cm tall, forming clumps to 5 cm wide; leaves 10-30 mm long, 1-4 mm wide, oblanceolate' to spatulate or almost linear, strigose; heads mostly solitary; involucres 9-12 mm high, 8-13 mm wide; bracts in 4 or 5 series, lan- ceolate, green or suffused purple, sparsely strigose; rays 13-21, yellow ventrally, often suffused reddish dorsally, glandular, 4-7 mm long; disk corollas yellow, ca 3.5 mm long; achenes 3-5 mm long, pubescent with glochi- diate hairs; ray pappus 2-4.5 mm long; disk pappus 5-8 mm long. Type.— USA. Utah. Sevier County, Ara- pien shale outcrop with scattered juniper, ca 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 369 370 Great Basin Naturalist Vol. 43, No. 2 4.2 km E of Sigurd, T23S, RIW, S8, at mile- post 14, Utah Hwy 24, 8 May 1975, S.L. Welsh 12700 (Holotype BRY; 2 isotypes dis- tributed previously as T. aprica Welsh & Reveal). Additional specimens: Utah. Sevier Coun- ty, Rainbow Hills area, 2 mi NE of Glen- wood, 7 May 1981, L. Greenwood 5013; do, 4 mi S of Salina, 24 Mar. 1981, L. Green- wood 5003; do, 2 mi E of Sigurd, 7 May 1981, L. Greenwood 5012. Piute County, NE side of Piute Reservoir, 26 Apr. 1979, E. Neese et al. 7149; do 30 Apr. 1981, D. At- wood and E. Neese 7794 (all BRY). The presence of yellow flowers in T. aprica was considered to be noteworthy for the genus, although corollas that dried yellowish were admitted for T. pnesii (Reveal 1970). The existence of populations with yellow co- rollas when fresh in Piute and western Sevier counties at first indicated placement of those plants with T. aprica, but the long pappus of the ray flowers indicates relationships with T. pnesii, in which the corollas sometimes fade yellowish. Most of the localities for this variety are situated on the Arapien Shale, which is noted for its deposits of commercial gypsum. The exception involves those plants from near the Piute Reservoir in Piute County, where the plants grow on gypsiferous or calciferous sub- strates amidst igneous gravels. Toumsendia montana Nutt. var. caelili- nensis Welsh, var. nov. A var. montana differt in foliis spatulatis late capitulis major- ibus et bracteis latioribus. Type:- USA. Utah. Sanpete Co., T20S, R4E, S33, ca 24 km W of Perron, 3050 m. Flagstaff Limestone, barrens, 13 July 1977, S. Welsh & S. Clark 15385 (Holotype BRY; 3 isotypes distributed previously as T. montana Nutt.). Additional specimens: Utah. Duchesne County, Argyle Canyon, 27 June 1978, E. Neese & L. England 5864; do, T6S, R6W, SW of Duchesne, 2 June 1978, E. Neese 5305. Sanpete County, 8 mi E of Spring City, 12 July 1977, E. Neese & S. White 3552; He- liotrope Mountain, 26 June 1977, M.E. Lewis 4254; do, 17 July 1981, D. Atwood 7998; do, head of Bacon Rind Canyon, J. W. Hum- phrey 52958, 22 July 1927; do, 11 mi 134 de- grees from Manti, 3 July 1981, 22 July 1927; South Tent Mountain, 27 July 1976, M. E. Lewis 4365; Skyline at head of Duck Fork, 22 July 1976, M. E. Lewis 4246; do, E of Manti, 28 July 1977, S. Clark 2877. Wasatch County, 27 mi 98 degrees east of Spanish Fork, 19 July 1982, S. Goodrich 17303 (all BRY). This is the large-headed phase of the spe- cies, with broadly rounded spatulate leaves, which lies intermediate with the typical montane materials and the dwarf plants of var. minima. The broad leaves and large heads are diagnostic in segregation of this en- tity from both the typical montane phase and the more southern var. minima. The substrate occupied by var. caelili- nensis on the Wasatch Plateau is composed of weathered Flagstaff Limestone. The Du- chesne County populations occur on calcif- erous members of the Green River Formation. Literature Cited Beaman, J. H. 1957. The systematics and evolution of Townsendia Compositae. Contr. Gray Herb. 183: 1-151. Reveal, J. L. 1970. A revision of the Utah species of Townsendia (Compositae). Great Basin Nat. 30:23-52. NEW HAPLOPAPPUS VARIETY IN UTAH (COMPOSITAE) Stanley L. Welsh' and Frank J. Smith- Abstract. — Reported as a new variety from the Green River Formation of Uintah and Duchesne counties, Utah, is Haplopappus armerioides (Nutt.) Gray var. gramineus Welsh & F. J. Smith. The arid Green River Formation, with its pecuhar soil relations, has been the spawning ground for numerous narrow endemics. The taxon described herein is another belonging to that unique group of mound-forming, low- growing taxa that grow on the peculiar shale substrates. Relatively little variation occurs within the typical variety of H. armerioides. Thus, the variation represented by the taxon proposed below falls well outside that of the typical variety and substantiates its recogni- tion. The main features that distinguish this variety from the main body of the species in- clude the short stature and slender leaves, but the heads are also smaller, on the average, and the involucral bracts tend to be narrow- er. This latter feature seems to indicate an af- finity with the related H. acaulis (Nutt.) Gray, but the bulk of the characteristics seem to represent an extension of the basic mor- phology of H. armerioides; hence, the place- ment with that taxon. The numerous speci- mens cited below have resulted from the laborious efforts of students of the Utah flora, whose contributions are gratefully ac- knowledged. Haplopappus armerioides (Nutt.) Gray var. gramineus Welsh & F. J. Smith, var. nov. Similis var. armerioides sed in staturis humilioribus et folius gracilioribus differt. Perennial caespitose herbs from a thick lig- neus pluricipital caudex and stout taproot, the caudex branches clothed with brown to ashy marcescent leaf bases and leaves; herb- age resinous-glandular, otherwise glabrous or with scabrous leaf margins; stems mainly 3-8 cm tall; basal leaves 1.5-4 cm long, 1-3 mm wide, rigid, linear, sharply mucronate, 1- or obscurely 3-nerved; cauline leaves few, re- duced upwards; heads solitary; involucres campanulate, 8-11 mm high, 10-14 mm wide; bracts in 3 or 4 series, imbricate, ob- long to oval or obovate, obtuse, sometimes lobed below the apex, greenish near the apex, glabrous; rays 8-12, 10-12 mm long, yellow, 3-4 mm wide; pappus white; achenes silky- villous. Type.- USA. Utah. Uintah County, T13S, R25E, S8, Atchee Ridge road, 1.4 km N of Boulevard Ridge, 2727 m elev, Green River Formation, pinyon-juniper, serviceberry, mountain mahogany, and sagebrush commu- nity, 29 May 1982, K. Thome & B. Neely 1836 (Holotype BRY; Isotypes NY; CAS). Additional specimens: Utah. Duchesne County, Dry Canyon, 40 km due NE of Price, 25 May 1978, E. Neese 4916. Uintah County, along Watson-Ouray rd, 8 km W of Bitter Creek, 26 May 1935, E. H. Graham 8988 (UTC); do. Big Pack Mtn., 16 May 1978, E. Neese & J. S. Peterson 4628; do, ca 45 km S of Ouray, 18 May 1978, E. Neese & J. S. Peterson 4729; do, Johnson Draw, be- tween Hill and Willow creeks, 27 Apr. 1978, ' E. Neese & J.L. England 4288; do. Long Draw, 2 km S of Ouray-Rainbow road, 18 May 1979, J. L. England 1777; do, Long Draw, ca 4.5 km SW of Rainbow, 16 May 1982, E. Neese & F. J. Smith 11388, 11389; do. Long Draw, ca 4 km W of Rainbow road, 26 May 1982, K. Thome and B. Neely 1793; do. Bitter Creek and West Fork of Asphalt Wash jet., 26 May 1982, K. Thome and B. Neely 1798; do, T13S, R25E, Sll, 30 May 1982, R. Kass & J. Trent 862; do, Atchee Ridge road, near East Seep Canyon, 7 June 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. '26 North 100 East, Smithfield, Utah 84335. 371 372 Great Basin Naturalist Vol. 43, No. 2 1982, F. J. Smith 1639, 1640; do, 4.5 km S of Rainbow, 15 May 1982, F. J. Smith 1596b, 1597; do. Asphalt Wash, 28 May 1982, R. Kass & J. Trent 842 (all BRY, except as noted). There is a tendency for some plants to have acutish bracts, and the broad subapical spot is only more or less developed. Despite the presence of these features, which are somewhat intermediate with phases of H. acaulis, the taxon stands with few inter- mediates to H. armerioides in a strict sense. Literature Cited Hall, H. M. 1928. The genus Haplopappiis. Carnegie Institution of Washington. 391 pp. NEW SPECIES OF HYMENOXYS AND PERITYLE (COMPOSITAE) FROM UTAH Stanley L. Welsh' and Elizabeth Neese^ Abstract. — Named as new species are Hymenoxys lapidicola Welsh & Neese, from Uintah County, Utah, and Perityle specuicola Welsh & Neese, from Grand County, Utah. Revision of the sunflower family, as part of the Utah Flora project, has led to the recog- nition of several taxa that were previously known but had been placed within other en- tities. Additionally, some of the taxa had ap- parently not been collected previously, but were similar to previously described taxa. Careful observation of each of the numerous specimens in the herbarium at Brigham Yoimg University has led to recognition of tiie great diversity in our flora. The two taxa described below each occur on sandstone, where they grow in crevices. They are part of a cadre of species with crevice habitats, and both are narrowly restricted endemics. Much more work remains to be done in the areas of the state occupied by massive sand- stones, limestones, and other geological strata that produce cliffs and escarpments. The fol- lowing species are named to indicate the pe- culiarities of their places of growth. Hymenoxys lapidicola Welsh & Neese, sp. nov. Ab H. torreyana (Nutt.) Parker in bracteis recuvatis et capitulis parvioribus differt et a H. depressa (T. & G.) Welsh & Reveal in bracteis recurvatis et marginem scariosis differt. Pulvinate caespitose herbs from a multi- cipital caudex, this densely clothed with brown marcescent leaf bases, acaulescent; leaves all basal, 0.3-1.2 cm long, 0.8-2 mm wide, narrowly oblanceolate, the inner con- spicuously glandular-punctate, the blades glabrous, the axils long-villous; heads solitary, immersed in the leaves; disks 5.5-9 mm wide; involucres 5-8 mm high; bracts distinct, in 2 or 3 subequal series, sparingly villous and suf- fused reddish, the margins scarious, the tips more or less squarrose-spreading and some- what thickened; rays 5 or 6, yellow, 5-6 mm long; pappus scales lance-acuminate, 2.3-3 mm long; achenes 2-2.5 mm long, pilose. Type.- USA. Uintah County, T5S, R25E, S20, Point of Pines camp area, S edge of Blue Mt. Plateau, at 2485 m, in ponderosa pine- manzanita community, sandy loam and standstone outcrop, 12 June 1982, E. Neese and C. Fullmer 11734 (Holotype BRY; Iso- type NY). Additional specimens: Utah. Uintah Coun- ty, Blue Mountain, cliff face, 11 June 1982, F. Smith & J. Trent 1653 (BRY); do. Point of Pines campground, Weber Sandstone, 11 June 1982, R. Cass & E. Neese 919 (BRY); do, 10 mi E of Jensen in draw at base of Blue Mountain, 14 May 1982, E. Neese et al, 11370 (BRY). The habit of H. lapidicola simulates that of H. depressa, but the features of the involucral bracts and the proportionally broader non- cuspidate leaves indicate a closer relationship with H. torreyana. From that entity it differs in the recurved outer involucral bracts with thickened reddish tips and much smaller heads. Perityle specuicola Welsh & Neese, sp.' nov. Similis Perityle congesta (Jones) Shinners sed in aspectis et pappo setarum 3 vel 4 (nee 1 vel 2), et in pappo palearum sigmoidis et expansis differt. Perennial suffruticose herbs, mainly 50-75 cm tall; stems sprawling or pendulous, much branched; herbage glandular-hispidulous; leaves mostly alternate, short-petiolate, the blades 3-6 mm long, 1.5-3 mm wide, ovate- elliptic, entire, hispidulous; heads few to many in a branching corymbose in- florescence; involucres 3.5-5 mm high, 5-6 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 'Life Science Museum, Brigham Young University, Provo, Utah 84602. 373 374 Great Basin Naturalist Vol. 43, No. 2 mm wide; bracts 11-16, oblong to elliptic, keeled; ray flowers lacking; disk flowers nu- merous, ca 2.5 mm long, whitish (?); pappus of 3 imequal scabrous bristles and often with 1 apically flattened and sigmoid scale; achenes 3-3.8 mm long, the faces flattened, glabrous, the margin thickened and with short ascending hairs. Type.- USA. Utah. Grand County, T25S, R21E, S24, 2 mi due N of Moab, 1220 m, hanging garden community, 29 Sept. 1977, S.L. Welsh 16283 (Holotype BRY; 6 isotypes distributed previously as Laphamia congesta Jones). Additional specimens: T25S, R22E, S29, Castle Valley, 2 mi due NNE of Moab, Negro Bill Canyon, 1220 m, Navajo sst, hanging gar- den community, 6 Oct. 1977, S.L. Welsh 16365 (BRY). The technical features of this taxon in- dicate an affinity with P. congesta, but the differences in the pappus features, inter alia, suggest the need for recognition of this dis- tinctive plant. It seems to be more than an etiolated shade form of its near ally, from which it is remote geographically by more than 300 km. Literature Cited Powell, A. M. 1973. Taxonomy of Perityle section Lap- hamia (Compositae-Helenieae-Peritylinae). Sida 5:61-128. NEW VARIETY OF STEPHANOMERIA TENUIFOLIA (COMPOSITAE) FROM UTAH Sherel Goodrich' and Stanley L. Welsh' Abstract. — Named as new is Stephanomeria tenuifolia (Torr.) Hall var. uintaensis Goodrich & Welsh. Identification of plants taken in prepara- tion of a flora of the Uinta Basin demon- strated the presence of a Stephanomeria whose description is beyond that of other taxa in the genus. Specimens were compared with all other taxa in the genus, and with published keys and descriptions of all con- temporary floras. The plants are situated at the margin of the range of S. tenuifolia, and are similar in general aspect to other phases of that species that are ecologically stressed. Plants taken on the West Tavaputs Plateau at 2745 m elevation (Welsh & Clark 15923 BRY) are similar to the aberrant materials from the Uinta Basin in general aspect and in leaf features, but the involucres are within the normal size range for typical S. tenuifolia. The material described below grows in one small isolated stand in a ponderosa pine com- munity and shows variation in size of heads and shape of the involucral bracts. The cau- dex is not well developed, producing solitary or few stems. Recognition at varietal level seems justified. More work in the genus is indicated. Stephanomeria tenuifolia (Torr.) Hall var. uintaensis Goodrich & Welsh, var. nov. A var. tenuifolia imprimis in involucris long- ioribus bracteis attenuatis radiis longioribus et foliis basalibus bipinnatifidis differt. Perennial herbs from a woody caudex; cau- dex sparingly branched or the branches lack- ing, with few marcescent leaf bases, not hairy; stems solitary or 2, 25-40 cm tall; herbage puberulent; leaves 1-11 cm long, 1-8 mm wide, the lower and sometimes the middle cauline ones runcinate-pinnatifid, the upper (at least) entire, linear, finally brac- teate; heads solitary at the ends of stems or on branches; involucres 10-16 mm high, 3-5 mm wide; main bracts lance-attenuate, green or suffused with purple, puberulent; outer bracts very short; rays 5, pink, 7-10 mm long; pappus white, plumose to the base; achenes 5-6 mm long, longitudinally ribbed, the angles smooth. Type.- USA. Utah. Uintah County, T2S, R19E, S14, NWi/4, 18 mi NW 319 degrees of Vernal, Ashley N.F., Uinta Mountains, Brownie Canyon, 2486 m; ponderosa pine, Douglas fir, Juniperus scopulorum, and sage- brush, south exposure, 20 Aug. 1982, Good- rich 17708 (Holotype BRY; isotypes NY, US, CAS, POM, MO, UT, UTC, RM, GH). Additional specimens: Utah. Uintah Coun- ty, Brownie Canyon, 13 Sept. 1982, S. Good- rich & D. Atwood 17968 (BRY). This variety approaches S. parryi in bract' size, but has the appearance of phases of S. tenuifolia at the edge of their ecological tol- erance. The tall involucres appear to be diag- nostic when taken with the other features mentioned in the diagnosis. 'USDA Forest Service, Intermountam Forest and Range Experiment Station, Ogden, Utah 84401, stationed in Provo, Utah, at the Shrub Sciences Laboratory. 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 375 NOTICE TO CONTRIBUTORS Manuscripts intended for publication in the Great Basin Naturalist or Great Basin Natural- ist Memoirs must meet the criteria outhned in paragraph one on the inside front cover. They should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Natu- ralist, 290 Life Science Museum, Provo, Utah 84602. Three copies of the manuscript are re- quired. 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 Council of Biology Editors Style Manual, Fourth Edition (AIBS 1978) 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 article. All manuscripts receive a critical peer review by specialists in the subject area of the manu- script under consideration. Authors may recommend one or two reviewers for their article. Accepted manuscripts less than 100 printed pages in length will automatically be assigned to the Great Basin Naturalist Those larger than 100 printed pages 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. The original illustrations or glossy photoprints of them should accompany the manuscript. Illustrations should be prepared for reduction by the printer to either single-column (2% inches) or double-column (5^ inches) width, with the length not exceeding 7^ inches. Costs Borne by Contributor. Contributors to the Great Basin Naturalist should be prepared to donate from $10 to $40 per printed page toward publication of their article (in addition to reprint costs). Excessive or complex tables requiring typesetting may be charged to the author at cost. Authors publishing in the Great Basin Naturalist Memoirs may be expected to contrib- ute $40 per printed page in addition to the cost of the printed copies they purchase. No re- prints are furnished free of charge. Reprint Schedule for the Great Basin Naturalist 100 copies, minimum cost for 2 pages, $26, Each additional 2 pages, $6. Each additional 100 copies, $4 for each 2 pages. Examples: 300 copies of 10 pages = $82; 200 copies of 13 pages = $86. 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 Intermoimtain biogeography: a symposium. By K. T. Harper, J. L. Reveal et al. $15. No. 3 The endangered species: a symposium. $6. No. 4 Soil-plant-animal relationships bearing on revegetation and land reclamation in Nevada deserts. $6. No. 5 Utah Lake monograph. $8. No. 6 The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. $60. TABLE OF CONTENTS Utah flora: Compositae (Asteraceae). Stanley L. Welsh 179 Haplopappus crispus and H. zionis (Asteraceae): new species from Utah. Loran C. Anderson 358 A bouquet of daisies (Erigeron, Compositae). Stanley L. Welsh 365 New taxa in Thelespetma and Townsendia (Compositae) from Utah. Stanley L. Welsh 369 New Haplopappus variety in Utah (Compositae). Stanley L. Welsh and Frank J. Smith 371 New species of Hymenoxys and Perityle (Compositae) from Utah. Stanley L. Welsh and Elizabeth Neese 373 New variety of Stephanomeria tenuifolia (Compositae) from Utah. Sherel Goodrich and Stanley L. Welsh 375 HE GREAT BASIN NATURALIST }lume 43 No. 3 July 31,1983 Brigham Young University LIPOAQY UNiVERSITV I'M; LIBRARY .„\V /.u/..^ . HARVARD GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Chairman, Botany; James R. Barnes, Zoology; Hal L. Black, Zoology; Stanley L. Welsh, Botany; Clayton M. White, Zoology. All are at Brig- ham Young University, Provo, Utah 84602. Ex Officio Editorial Board Members. Bruce N. Smith, Dean, College of Biological and Agricul- tural Sciences; Norman A. Darais, University Editor, University Publications. Subject Area Associate Editors. Dr. Noel H. Holmgren, New York Botanical Garden, Bronx, New York 10458 (Plant Taxonomy). Dr. James A. MacMahon, Utah State University, Department of Biology, UMC 53, Lo- gan, Utah 84322 (Vertebrate Zoology). Dr. G. Wayne Minshall, Department of Biology, Idaho State University, Pocatello, Idaho 83201 (Aquatic Biology). Dr. Ned K. Johnson, Museum of Vertebrate Zoology and Department of Zoology, Uni- versity of California, Berkeley, California 94720 (Ornithology). Dr. E. Philip Pister, Associate Fishery Biologist, California Department of Fish and Game, 407 West Line Street, Bishop, California 93514 (Fish Biology). Dr. Wayne N. Mathis, Chairman, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 (Entomology). Dr. Theodore W. Weaver III, Department of Botany, Montana State University, Boze- man, Montana 59715 (Plant Ecology). The Great Basin Naturalist was founded in 1939 and has been published from one to four times a year since then by Brigham Young University. Previously unpublished manuscripts in English of less than 100 printed pages in length and pertaining to the biological natural his- tory of western North America are accepted. Western North America is considered to be west of the Mississippi River from Alaska to Panama. The Great Basin Naturalist Memoirs was es- tablished in 1976 for scholarly works in biological natural history longer than can be accom- modated in the parent publication. The Memoirs appears irregularly and bears no geographi- cal restriction in subject matter. Manuscripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist for private individuals is $16.00; for institutions, $24.00 (outside the United States, $18.00 and $26.00); and for stu- dent subscriptions, $10.00. The price of single issues is $6.00 each. All back issues are in print and are available for sale. All matters pertaining to subscriptions, back issues, or other busi-* ness should be directed to Brigham Young University, Great Basin Naturalist, 290 Life Sci- ence Museum, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same office 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. See Notice to Contributors on the inside back cover. 9-83 650 66769 ISSN 017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 43 July 31, 1983 No. 3 A RE-EVALUATION OF THE POSTGLACIAL VEGETATION OF THE LARAMIE BASIN, WYOMING-COLORADO Deborah L. Elliott-Fisk', Betty S. Adkins-, and Jeanine L. Spaulding' Abstract.— Previous work by Wells in the Laramie Basin suggested that a coniferous forest /woodland covered the basin floor in the recent past (until the latest Holocene). We have found no evidence for this woodland and sug- gest instead that these scattered woodlands along sandstone outcrops and their immediate margins are outliers of the montane forest of the Medicine Bow Mountains, existing in their apparently anomalous locations due to favorable microenvironments set up by the topography and substrate. During the past two years, we worked at various sites in and surrounding the Laramie Basin in an attempt to collect sufficient infor- mation with which to reconstruct postglacial vegetation (and environment) change for the region. The basin's unique geography as one of the few high-altitude basins surrounded by the Rocky Mountains, and the knowledge that a periglacial climate existed here in pre- Holocene times (Mears 1981), encouraged us to seek postglacial paleoclimatic data from not only upper elevations in the mountains (as done traditionally in the Rocky Moun- tains), but for the basin itself. Faunal, soil, and geomorphic studies here (Hager 1972, Reider et al. 1974, Grasso 1979, Reider and Burgess, in prep.) point to late Pleistocene and Holocene environmental change. In ad- dition, a 116-year-old record of mete- orological data is available for the basin, which will aid any climatic reconstructions attempted. Upon visiting the Sand Creek region of the Laramie Basin (Fig. 1), we were delighted to find an abundance of fossiliferous material, including in situ dead trees (macrofossils) and Neotoma (woodrat) middens, as well as living trees that appeared to be at least several cen- turies old. Both living and dead trees were restricted to the sandstone outcrops and their immediate margins, these outcrops principal- ly lithified sand dunes of the Pennsylvanian Casper Formation (Fig. 2). It later surprised us when we read in the papers of Wells (1970a, 1970b) on the post- glacial vegetation of the Laramie Basin tRat he believed these trees had in recent times (the latter half of the Altithermal and per- haps part of the Neoglacial) covered the en- tire floor of the Laramie Basin. Knowing both the ecological requirements of these conifer species {Juniperus scoptilorum Sarg.— Rocky Mountain juniper; Pinus flexilis James— limber pine; and P. ponderosa Laws.— ponderosa pine) and the relatively minor de- gree of climatic change in the Holocene compared to the Pleistocene, we considered Wells's hypothesis not plausible. Instead, 'Department of Geography, University of California, Davis, California 95616. 'Department of Geography, University of Alberta, Edmonton, Alberta, Canada T6G 2H4. 'Department of Geography, University of Wyoming, Laramie, Wyoming 82071. 377 378 Great Basin Naturalist Vol. 43, No. 3 Fig. 1. Map of Sand Creek study area in the Laramie Basin of Wyoming-Colorado. topography and microenvironmental condi- tions could have favored a more vigorous woodland (as evidenced by population and individual sizes) on the outcrops sometime in the past. We therefore decided to critically reeval- uate the paleoenvironmental data for the Sand Creek area. This involved examining a larger area than apparently done by Wells (pers. comm. with local ranchers) and also an attempt to tie the botanical data in with other paleoenvironmental studies conducted in the region. The completion of this project awaits radiocarbon dates and further den- droclimatic reconstructions. It is our hypoth- esis that these scattered woodland stands are outliers of the montane forest of the Medi- cine Bow Mountains, existing in their appar- ently anomalous locations due to favorable microenvironments set up by the topography and substrate, and not relicts of a former for- est over the basin floor as hypothesized by Wells. Methods The methods we have used to solicit infor- mation as to the nature and degree of change of past environments at Sand Creek involve gathering various paleoecological materials. As we have yet neither found fossil soils in this immediate area, nor analyzed animal re- mains found in the Neotoma middens, we confine our discussion to the botany and pa- leobotany of the area. Extensive hiking and collecting was done in the area outlined in Figure 1 to determine the spatial and temporal extent of conifers in this region. Dead and living individuals were mapped, along with Neotoma middens (both July 1983 Elliott-Fisk et al.: Postglacial Vegetation 379 Fig. 2. Study site 1 at Sand Creek (Wells' primary site). The landscape here is characterized by lithified sand dunes of the Pennsylvanian Casper Formation. The tree in the middle of the photo is a Rocky Mountain juniper (Jiiniperiis scopuhrtim) approximately 1000 years of age. fossil and contemporary). Although collecting has been completed at many of these sites, materials are still being obtained. In addition, the vegetation, regenerative capacity, micro- climatic setting, geology, and soils of the sites were recorded. Initial tree-ring and midden analyses, fol- lowing the traditional methods of Stokes and Smiley (1968), Fritts (1976), and Wells (1976), have been conducted. Studies on the regenerative capacity of the conifer popu- lations follow the methods outlined by Elliott (1979). We present our initial findings here. We believe these findings are important for those attempting to reconstruct Holocene cli- matic change for the Western USA, because they offer a radically different conclusion than diat presented by Wells (1970a, 1970b). Also, for anyone undertaking recent pa- leoenvironmental studies, it becomes increas- ingly important to have a thorough under- standing of the life strategy and ecological requirements of the species investigated, and also of the local geography of a site, if one is to accurately reconstruct a climatic record. Results The present distribution of trees and tree species is shown in Figure 3. Multiple mono- specific and mixed species stands occur along this semicontinuous sandstone ridge extend- ing out from the southwestern Medicine Bow Mountains. The lower forest limit here is 2,500 m, with woodland patches reaching elevations of 2,300 m. It is important to note that the lowest trees surviving here are not on the floor of the ba- sin proper, but are restricted to the sandstone outcrops (Fig. 4) or their immediate margins. The dead trees (macrofossils) reported by Wells and found by us occupy the same topo- graphic position as the living conifers, where winter snowdrift and summer runoff can sup- ply them with the necessary moisture for sur- vival in this marginal environment (Fig. 5). It is interesting to note that two of the fall- en fossil junipers plus two standing dead stumps in the area are much larger than any living junipers in the area today (Fig. 6). As this species is apparently at its lower mois- 380 Great Basin Naturalist Vol. 43, No. 3 Juniperus scopulorum PInus tiexilis Pinus ponderosa Sites Outcrops CONIFER POPULATIONS: SAND CREEK Fig. 3. Distribution of living and fossil (in situ dead) trees in Sand Creek study area. Findings in woodrat middens are not included here. Sandstone outcrops ex- tend from the Medicine Bow Mountains to their south- west, decreasing in elevation toward the northeast. tiire, but not temperature, limit today (Po- wells 1965), it seems reasonable to speculate that more moisture (either due to increased precipitation or decreased temperatures/ evapotranspiration) was available during at least part of the past life spans of these trees. This hypothesis is also supported by the fact that no mixed stands of ponderosa pine and juniper (as evidenced in some of the fos- sil middens) exist here today. Stands of limber pine and juniper do exist, along with mixed stands of all three species and pure stands of juniper or ponderosa pine. Fossil midden equivalents of all stand types mentioned here except the monospecific stands are found. All three species are successfully sexually regenerating today in each stand in which they occur. This is witnessed by the presence of both viable seed (cones) and juveniles. It proves difficult to make any type of rig- orous statement in reference to the age struc- tures of the tree populations here. This is due not only to a lack of radiocarbon dates for each dead tree (macrofossil), but primarily because both dead and apparently also living trees have been removed from this area for use as fence posts, firewood, etc. Many of the cut junipers we have found were harvested for "cedar table tops" in the 1920s (pers. comm., F. Lilly to B. Mears 1980), a fact that is distressing to paleoecologists trying to re- construct the history of a site. Cross-dating of the tree rings in mature junipers is also ex- tremely difficult; however, limber and pon- derosa pine are easily cross-datable. Juveniles of all three species, ranging from a few to 50 years in age, are abundant. All age classes (standard 10-year groups) of pon- derosa pine are found for the last few hun- dred years, with no individuals over 350 years old (as yet found) surviving today. Pon- derosa pine macrofossils (both in situ dead trees and smaller macrofossils in middens) are infrequent, though we have found cones and seeds at site 1. Limber pine is found up to several hundred years old, with some individ- uals approaching 1,000 years in age. In situ limber pine macrofossils are common at sites 2 and 3; some of these dead trees were also much larger than those living today. Sam- pling of these macrofossils is not yet complete. Juniper appears to have had considerable difficulty through at least the last few thou- sand years (as documented by Wells's radio- carbon dates) with the establishment of an equilibrium population in this area. The liv- ing individuals today are primarily either several hundred to 1,000 years old or under 100 years in age. Juveniles tend to be clus- tered around singular dead or living mature individuals, which most likely served as the seed (mother) trees. Layering of juveniles from each other (but not from the larger trees) is also found. Occasionally, juveniles are found a great distance (over 1 km) from any possible seed trees. Few young adults are found, an age group in which the two pine species are abundant. Further dendroeco- logical work with response functions for a species may help explain this difference. One of the most intriguing facets of the Sand Creek sites is the tremendous variation in average tree-ring widths as one looks at living versus dead individuals. The ponderosa pines seem to have found a couple of sites at July 1983 Elliott-Fisk et al.: Postglacial Vegetation 381 Fig. 4. Trees at site 2. The majority ot trees in this stand are situated on top of the sandstone outcrop, with indi- viduals occasionally found along the margins. Junipcrus scopulontrn, Pinits flexilis, and Pinus ponderosa are all pres- ent here as both living individuals and fossils. which they can exist with only occasional stress (as evidenced by very narrow rings every 30 to 40 years which are cross-datable); these individuals show wide, symmetrical crowning with consistently good growth and low mean sensitivities. In contrast, limber pine and juniper appear to have been under considerable environmental (climatic) stress, though this has not precluded the estab- lishment of juveniles in recent times. Narrow rings and high mean sensitivities are common for both species here. For comparative purposes, we present av- erage ring-width data for several individuals of Juniperus scopulorum that we believe are representative of our samples. Whereas two measured, undated fossil trees have average ring widths of 1.09 and 2.38 mm, respective- ly, the largest living tree at Wells's site (our site 1) has an average ring width of 0.19 mm. One section of a large dead juniper cited by Wells with a radiocarbon date of 940 ± 105 BP (Gx-140F) on the outer wood has an aver- age ring width of 0.17 mm; however, this val- ue is derived from measurements on a branch cross-section and may therefore be expected to be smaller than those from the primary trunk. (All of the other measurements given are from main trunk cores and cross-sections.) The above data show almost a 25-fold dif- ference in ring-widths. As the living and fossil trees are found in the same habitats, this dif- ference in growth must be attributed to a change in climate, with other environmental factors (topography, etc.) remaining constant. It is not attributable to intrapopulation or age differences. Conclusion It appears that Holocene climatic change in the Laramie Basin has been sufficient to trigger the death of conifers in the most mar- ginal low-elevation sites. This is suggested by: (1) change in species composition of some of the lowest woodland stands, with only the most xeric tree species surviving this deterio- ration, (2) the death of individuals at the most marginal microclimatic/topographic loca- tions, and (3) the decrease in ring width 382 Great Basin Naturalist Vol. 43, No. 3 Fig. 5a. Remains of fossil (dead) Juniperus scopulorum at the margin of a sandstone outcrop. This individual, like many in the area, has been partly harvested for lumber. Site 1. Fig. 5b. Immature (not yet bearing seeds) Juniperus scopulorum along the margin of a sandstone outcrop between sites 1 and 2. This is a typical habitat for juniper regeneration. This individual is approximately 80 years old. July 1983 Elliott-Fisk et al.: Postglacial Vegetation 383 Fig. 6. One of two large, dead, fallen junipers at site 1. A cross-section through the midpart of this tree has been cut out, most likely for a table top. Presence of small branches on this individual attest to the fact that it has prob- ably not been dead for more than a few hundred years. (average growth rate) of the mature conifer populations. We have found no evidence that a con- iferous woodland or forest ever existed over the floor of the Laramie Basin in the Holo- cene (or even Pleistocene, though this latter time period is out of the scope of this study). It does appear, however, that more and occa- sionally larger individuals of the extant con- ifer species did exist on the sandstone out- crops and at their immediate margins in the early Neoglacial and perhaps late Altithermal. Acknowledgments This research was initiated while the au- thors were faculty (Elliott-Fisk) and students (Adkins and Spaulding) in the Department of Geography at the University of Wyoming; we appreciate the use of laboratory facilities there. We are also indebted to several local ranchers in the Laramie Basin for permission to work on their property and for various types of information and to Richard G. Rei- der, Brainerd Mears, Jr., and Thomas P. Har- lan for helpful suggestions. Tree-ring mea- surements were done by Elliott-Fisk at the Laboratory of Tree-Ring Research, Univer- sity of Arizona; the use of this facility is very much appreciated. We also thank the many students and faculty who have accompanied us in our field work at Sand Creek. » Literature Cited Elliott, D. L. 1979. The stability of the northern Cana- dian tree limit: current regenerative capacity. Unpublished dissertation. Univ. of Colorado, Boulder. 192 pp. FowELLS, H. A. 1965. Silvics of forest trees of the United States. USDA Agr. Handbk. 271. Washington, D.C. 762 pp. Fritts, H. C. 1976. Tree rings and climate. Academic Press, New York. 567 pp. Grasso, D. N. 1979. Paleoclimatic significance of fossil ice-wedge polygons in the Laramie Basin, Wyoming. Unpublished thesis. Univ. of Wyo- ming, Laramie, 105 pp. Hager, M. W. 1972. A late Wisconsin-Recent vertebrate fauna from the Chimney Rock animal trap, Lari- mer County, Colorado. Contrib. to Geol. 2:6.3-71. (Univ. of Wyoming, Laramie). Mears, B., Jr. 1981. Periglacial wedges and Late Pleisto- cene environment of Wyoming's intermontane basins. Quat. Res. 15:171-198. 384 Great Basin Naturalist Vol. 43, No. 3 Reider, R. G., N. J. KuNiANSKY, D. M. Stiller, and P. J. Uhl. 1974. Preliminary investigation of com- parative soil development on Pleistocene and Holocene geomorphic surfaces of the Laramie Basin, Wyoming. Pages 27-.33 in M. Wilson, ed.. Applied geology and archaeology: the Holocene history of Wyoming. Geol. Surv. of Wyoming, Rept. of Investigations 10. Stokes, M. A., and T. L. Smiley. 1968. An introduction to tree-ring dating. Univ. of Chicago Press, Chi- cago. 73 pp. Wells, P. V. 1970a. Postglacial vegetational history of the Great Plains. Science 167:1574-1582. 1970b. Vegetational history of the Great Plains: a post-glacial record of coniferous woodland in southeastern Wyoming. Pages 185-202 in W. Dort and K. J. Jones, eds.. Pleistocene and Recent environments of the Central Great Plains. Univ. of Kansas Press, Lawrence. 1976. Macrofossil analysis of woodrat (Neotoma) middens as a key to the Quaternary vegetational history of arid America. Quat. Res. 6:223-248. COMPARATIVE LIFE HISTORY AND FLORAL CHARACTERISTICS OF DESERT AND MOUNTAIN FLORAS IN UTAH Patrick D. Collins', Kimball T. Harper', and Burton K. Pendleton- Abstract.— Life forms and floral characteristics of plants at Arches National Park (desert communities), the Mt. Nebo complex, and a subalpine meadow in the Uinta Mountains (montane and subalpine communities) were com- pared. Characteristics observed were (1) life form, (2) longevity, (3) pollination system, (4) flower structure, (5) flower symmetry, and (6) flower color. Common families in each flora were also compared. Results showed that there is a significant overrepresentation of shnib species at Arches, and an underrepresentation of perennial forbs. Relative number of perennial forb species was significantly higher at Mt. Nebo and the subalpine meadow than at Arches Na- tional Park. Native annuals and wind-pollinated species were significantly overrepresented at Arches. Flowers with open stnicture that permit free access of most insects to nectar and pollen were overrepresented at Mt. Nebo and in the subalpine meadow. The distribution of flower colors also differs significantly among these ecologically con- trasting floras, with yellow being best represented in the desert and white in the mountains. Because of accelerating development of energy and other natural resources, plant communities of the world are constantly being altered. In the United States, law re- quires that such disturbed areas be restored to their natural condition (Public Law 95-87, 1977), but little is known of the relative pro- portion of life histories and floral character- istics that enhance coexistence and self- perpetuation of a variety of wild plant spe- cies on common sites. What controls the rela- tive success of pollen transfer by wind or ani- mal in various natural communities? Do the contrasting climatic conditions of certain en- vironments affect the success of species of various life forms, longevity, and/or floral characteristics? In this paper, we compare characteristics of three Utah floras: the floras are from Arches National Park (desert), the Mt. Nebo complex (midelevation montane vegetation), and a subalpine meadow at high elevation in the Uinta Mountains. Arches National Park is a semiarid, cold desert region in southeastern Utah with an average elevation of about 1,220 m above sea level. Its topography con- sists of rolling hills and sandstone outcrops. The mean annual precipitation at Moab, near Arches, was 21.7 cm with a standard devia- tion of 4.3 cm (Nat. Oceanic and Atmosph. Admin. 1971-1979). The average annual tem- perature was 13.5 C. Mt. Nebo and adjacent mountains form a montane habitat with ele- vations between 1,829 and 3,621 m, but aver- age elevation is in the neighborhood of 2,500 m. Average annual temperatures at Tim- panogos Cave (1,720) 73.3 km north of Mt. Nebo was 9.4 C. The mean annual precipi- tation at Timpanogos Cave was 55.3 cm for the 1971-1979 period and that at the Payson Guard Station (2,454 m) was 73.3 cm with a standard deviation of 11.9 cm (Whaley and Lytton 1978). The Payson Guard Station is 17.4 km north of the crest of Mt. Nebo. The series of subalpine meadows selected for this study are located just below Bald Mountain Pass at the 28-mile marker (45.1 km) on the Mirror Lake Highway (Utah Highway 150) in Summit County, Utah. Average elevation at these meadows is 3,216 m, whereas mean an- nual precipitation at Trial Lake (near the meadows) was 102.8 cm with a standard de- viation of 18.3 cm (Whaley and Lytton 1978). The mean annual temperature of this site is approximately -2.4 C (Callison and Harper, in review). Jaccard's community coefficient (1912) shows the Arches and Nebo floras to be 90 percent dissimilar, Arches and Bald Mountain 98 percent dissimilar, and the Nebo and Bald 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 'Department of Biological Sciences, Wayne State University, Detroit, Michigan 48202. 385 386 Great Basin Naturalist Vol. 43, No. 3 Mountain floras 90 percent dissimilar. Be- cause these floras are so dissimilar and oc- cupy such physically different ecological situ- ations, it was hypothesized that there would be statistically significant differences in re- productive strategies of the plant species of the three floras. This paper compares the dis- tribution of life form, longevity, and floral characteristics in the three floras. The com- parisons evaluate the relative success of vari- ous reproductive strategies of plant species native to these three contrasting envi- ronments. Methods Floristic checklists furnish the data on which this study is based. The list for Mt. Nebo and adjacent mountains, Utah County, was compiled from Collins (1979) and Allred (1975). Checklists for Arches National Park were prepared by Harrison et al. (1964) and Allan (1977). The Bald Mountain meadow checklist was taken from Pendleton (1981). Life history and floral characteristics were determined for all species from preserved specimens in the Brigham Young University Herbarium. There were 734 species (647 na- tive; 12.0 percent introduced) on the Mt. Nebo complex checkhst, 356 (322 native; 9.6 percent introduced) on the desert list (Arches National Park), and 134 (all native) on the high elevation meadow list (Bald Mountain). The following information was obtained for each of the native species: (1) longevity, (2) life form, (3) likely pollinating agent, (4) flower symmetry, (5) flower structure, and (6) flower color. All analyses reported in this pa- per are based on native species only. Longevity was simply recorded as annual or perennial. Species described in keys as biennials were treated here as annuals, except those species listed as "biennials to short- lived perennials" were considered perennials. Plant life form was noted as tree, shrub, forb, or grass. In respect to pollination system, plant species were classified as anemophilous or zoophilous. It is realized that some of the species may be self-pollinated, but this could not be determined without independent re- search on each species. Thus, no attempt was made to identify self-pollinated taxa. Flower symmetry was regarded as either zygomorphic or actinomorphic. Species were also classified according to flower structure. Structure of zoophilous flowers was de- scribed as restricted when access to nectar or pollen was difficult for unspecialized pollina- tors. Restricted access flowers had long co- rolla or calyx tubes or had nectaries at the base of long spurs, thus limiting access to pol- len or nectar. Flowers classified as open- structured were saucer or bowl shaped and appeared incapable of mechanically exclud- ing any pollinator. Moreover, some plants were considered to be only partially restric- tive, having short calyx tubes or deeply lobed and/or widely flaring sympetalous corollas: such flowers were listed as semi-restricted. Flower colors were listed as red, violet, blue, yellow, pink, white, or greenish. Important families for each location were summarized by the number of species found in each flora. Jaccard's similarity index was employed to test compositional similarity be- tween floras. The Chi-square statistic was used to identify departures from random ex- pectations. In the Chi-square analyses, ran- dom expectations are based on the propor- tion of the species in the pooled floras that share a particular trait (e.g., the proportion of the species in the combined floras from the desert and montane environments that have red flowers). If the trait is randomly dis- tributed between the two floras, the propor- tion of species having the trait in each flora should not differ significantly from the pro- portion having that trait in the pooled flora. The Chi-square statistic was used to test whether the observed and expected numbers of species (tests use absolute numbers, not the proportions) sharing a trait in the individual floras differed significantly. If there was a significant departure, the trait was consid- ered to be under- or overrepresented in a giv- en flora. Introduced species were omitted from all analyses on the assumption that they may not have achieved stable reproductive characteristics in their new home. Results and Discussion A total of 70 families, 307 genera, and 734 species occurred in the Mt. Nebo flora. There are 60 families, 203 genera, and 356 species July 1983 Collins et al.: Desert and Mountain Floras 387 reported for the Arches flora. Thirty-six fami- hes, 86 genera, and 134 species appear on the Bald Mountain meadows checklist. Jaccard's index (Jaccard 1912) shows the Arches and Nebo floras to be 10.0 percent similar on the basis of species, 32.5 percent similar on the basis of genera, and 75.5 percent similar on the basis of families. The Arches and Bald Mountain floras are 1.5 percent similar on the basis of species, 12.0 percent similar on the basis of genera, and 39.1 percent similar on the basis of families. The Mt. Nebo and Bald Mountain floras show 10.2 percent sim- ilarity by species, 23.2 percent by genera, and 45.2 percent by families. Because the families contributing species to these flora are so similar, most of the observed differ- ences in plant adaptations in the three areas can be attributed to ecological selection rather than to differences in basic phylogeny. Families contributing most of the species in the three floras are reported in Table 1. The families Asteraceae and Poaceae domi- nate the three floras: Fabaceae holds third place in the Arches flora, Cyperaceae claims that position at Bald Mountain, and Brassi- caceae takes that slot on Mt. Nebo. The fam- ily Chenopodiaceae contributes over three times as many species in relative terms in the desert as in the mountains. In contrast, spe- cies of Rosaceae, Caryophyllaceae, Polemo- niaceae, Saxifragaceae, and Scrophulariaceae are twice or more as common in our moun- tains as in the desert flora considered. Life Form.— In all three floras, the pre- dominant life form (as represented by num- ber of species) is the broad-leaved herb (forb). At Arches National Park, 64.3 percent of the total flora is contributed by forbs; on Mt. Nebo, 73.0 percent of the species are forbs; and at Bald Mountain, 64.9 percent of the species are forbs (Table 2). Analysis shows that forbs are significantly overrepresented in the midmontane flora, whereas they are un- derrepresented in the high-elevation mead- ows and in deserts (Table 3). The shrub life form contributes proportionally over twice as many species in the desert (14.3 percent) as in the mountain floras at Mt. Nebo (7.0 per- cent) and at Bald Mountain (6.8 percent). That difference is statistically highly signifi- cant (2X2 = 30.46, P < 0.005, Table 3A). The results support the hypothesis that un- der dry conditions, shrubs are more successful than forbs. Deserts are notorious for unpre- dictable climatic patterns, and many forbs do not tolerate moisture deficits for long periods (Hironaka 1963, Mueggler 1972, Harner and Harper 1973). Shrubs can tolerate such con- ditions. They exhibit a variety of adaptations to dry environments, such as deep root sys- tems and reduced reliance on turgor pressure to keep leaves expanded to collect light and carbon dioxide (Sharif and West 1968). Shrubs also have leathery or firm leaves that reduce breakage from heavy winds and are resistant to herbivory; and, finally, shrubs have well-developed secondary meristems Table 1. A comparison of important families showing the number of species and percent (in parenthesis) of thg total floras of Arches National Park, the Mt. Nebo complex, and Bald Mountain meadows, Utah. Family Arches No. (%) Mt. Nebo No. (%) Bald Mtn. No. (%) Asteraceae Poaceae Fabaceae Chenopodiaceae Brassicaceae Scrophulariaceae Boraginaceae Cyperaceae Rosaceae Liliaceae Caryophyllaceae Ranunculaceae Polemoniaceae Saxifragaceae Salicaceae Other Totals 80 (22.5) 51 (14.3) 25 (7.0) 22 (6.2) 17 (4.8) 10 (2.8) 10 (2.8) 9 (2.5) / (2.0) 7 (2.0) 2 (0.6) 4 (1.1) 5 (1.4) 1 (0.3) 6 (1.7) 100 (28.0) 115 (15.7) 105 (14.3) 30 (4.1) 14 (1.9) 50 (6.8) 36 (4.9) 19 (2.5) 16 (2.2) 34 (4.6) 14 (1.9) 18 (2.5) 28 (3.8) 19 (2.6) 18 (2.5) 15 (2.0) 203 (27.7) 356(100.0) 734 (100.0) 20 (14.9) 18 (13.4) 2 (1.5) 0 (0.0) 4 (3.0) 8 (6.0) 1 (0.7) 12 (9.0) 4 (3.0) 4 (3.0) 6 (4.5) 5 (3.7) 1 (0.7) 3 (2.2) 1 (0.7) 45 (33.6) 134 ( 100.0) 388 Great Basin Naturalist Vol. 43, No. 3 that probably permit individual roots to be longer lived than is possible for species that lack secondary meristems (as in grasses and sedges). During dry periods, shrubs persist and maintain root systems in both lateral and vertical space; when better moisture condi- tions do return, herbs attempting to colonize barren spaces between shrubs experience ex- treme competition from the already estab- lished root systems of shrubs. Even in moist years, however, the barren interspaces be- tween shrubs are only sparsely clothed with annual plants, but nearby areas that have been deprived of their shrub cover by abu- sive grazing or mechanical disturbance sup- port a nearly complete cover of annual plants (Hutchings and Stewart 1953). Graminoides tend to be better represented in the Arches flora (grasses) and at Bald Mountain (sedges) than at Mt. Nebo. Trees are best represented in the midelevation mountain flora (Table 2). Longevity.— Longevity of species in the desert and montane floras also show signifi- cant differences. The deserts have more an- nual species than one would expect by chance (2X2 = £1.27, P < 0.005, Table 3B). Native annuals contribute 18.9 percent of the 322 species at Arches National Park, 10.5 percent of the 647 species on the Mt. Nebo Table 2. Characteristics of the native floras of Arches National Park, the Mt. Nebo complex, and the Bald Moun- tain meadows. The table shows the number of species and the percent of the native flora (in parentheses). Arches Mt. Nebo Bald Mtn. No. Sp. (% flora) No. Sp. (% flora) No. Sp. (% flora) Life form Trees 13 (4.0) 39 (6.0) 3 (2.2) Shrubs 46 (14.3) 45 (7.0) 9 (6.7) Forbs 207 (64.3) 472 (73.0) 87 (64.9) Graminoides 56 (17.4) 91 (14.0) 35 (26.1) Total 322 (100.0) 647 (100.0) 134 (100.0) Longevity Annual 61 (18.9) 68 (10.5) 7 (5.2) Perennial 261 (81.1) 579 (89.5) 127 (94.8) Total .322 (100.0) 647 (100.0) 134 (100.0) Pollination system (annuals excluded) Anemophilous 101 (38.7) 162 (28.0) 39 (30.7) Zoophilous 160 (61.3) 417 (72.0) 88 (69.3) Total 261 (100.0) 579 (100.0) 127(100.0) Flower symmetry (zoophilous species) Radial 188 (88.7) 395 (84.4) 77 (83.7) Bilateral 24 (11. .3) 73 (15.6) 15 (16.3) Total 212 (100.0) 468 (100.0) 92 (100.0) Flower structure (zoophilous species) Open 55 (25.9) 179 (38.2) 42 (45.7) Restricted 59 (27.8) 128 (27.4) 14 (15.2) Semirestricted 98 (46.2) 161 (34.4) 36 (39.1) Total 212 (100.0) 468 (100.0) 92 (100.0) Flower color (zoophilous species) White 50 (23.6) 165 (35.3) 30 (32.6) Yellow 88 (41.5) 148 (31.6) 23 (25.0) Blue 20 (9.4) 57 (12.2) 8 (8.7) Violet 27 (12.7) 36 (7.7) 12 (13.0) Pink 8 (3.8) 34 (7.2) 12 (13.0) Red 14 (6.6) 15 (3.2) 1 (1.1) Green 4 (1.9) 8 (1.7) 5 (5.4) Other 1 (0.5) 212 (100.0) 5 (1.1) 1 (1.1) Total 468 (100.0) 92 (100.0) July 1983 Collins et al.: Desert and Mountain Floras 389 complex, and only 5.2 percent of the 134 spe- cies from the Bald Mountain flora. Climatic unpredictability enhances the success of annuals in deserts (Schaffer and Gadgil 1975), where precipitation is sporadic and scarce. The annual strategy seems well suited for such conditions, whereas perennial forbs consistently contain high levels of tissue moisture (Sharif and West 1968). Our results show that perennial forbs are under- represented in the desert (Table 3C). Pollination Systems.— Because shrubs have been shown to be overrepresented at Arches, and because anemophily is heavily favored among woody species (Ostler and Harper 1978, Freeman et al. 1980), we anticipated that anemophily would be most prevalent at Arches. At Arches National Park, 38.7 per- cent of the perennial flora is anemophilous; on Mt. Nebo only 28.0 percent and at Bald Mountain only 30.7 percent of the species are wind pollinated (annuals have been omitted from this analysis to minimize the possible confusing effect of self-pollinated species, which are believed to be especially common among annuals [Solbrig 1977]). The differences in modes of pollination in the three floras are statistically significant (SX^ = 9.64, P < 0.005, Table 3D). The reason that there are more anemo- philous species in the desert is not that there is more wind movement there. The Arches area receives only half as much wind (1,590.5 km at Moab) as the Mt. Nebo area (2,984.4 Table 3. Chi-square analyses comparing life histories and floral characteristics of the native plant species of Arches National Park, the Mt. Nebo complex, and the Bald Mountain complex of Utah. Observed and expected numbers of species (in parentheses) are shown. Asterisks indicate significance level: single < .05, double < .010, triple < .005 probability. Summation Chi-square Arches Mt. Nebo Bald Mtn. values Life form Trees 13 (16.1) 39 (32.3) I (6.7) Shrubs 46 (29.2) 45 (58.7) 9 (12.1) Forbs 207 (223.6) 472 (449.2) 87 (93.1) Grasses 56 (53.1) 91 (106.8) 35 (22.1) 30.46°°° Growth cycle Annuals 61 (39.7) 68 (79.8) 7 (16.5) Perrenials 261 (282.3) 579 (567.2) 127 (117.5) 21.27°°° Growth cycle/life form Perennial forbs 150 (184.5) 406 (370.7) 76 (76.8) Other native spec •ies 172 (137.5) 241 (276.3) 58 (57.2) 22.99°°° • Pollination system (annuals ( excluded) Anemophilous 101 (81.5) 162 (180.8) 39 (39.7) Zoophilous 160 (179.5) 417 (398.2) 88 (87.3) 9.64°°° Flower structure (for zoophilous species) All species considered ( annuals included) Open 55 (75.8) 179 (167.3) 42 (32.9) Restricted 59 (55.2) 128 (121.8) 14 (24.0) Semirestricted 98 (81.0) 161 (178.8) 36 (35.2) 19.11°°° Only perennial species considered (semirestricted taxa ignored) Open 46 (59.2) 160 (156.6) 38 (28.2) Restricted 61 (47.8) 123 (126.4) 13 (22.8) 14..35°°° Flower color (for zooph ilous species) Red 14 (8.2) 15 (18.2) 1 (3.6) Other 198 (203.8) 453 (449.8) 91 (88.4) 6.70° White 50 (67.3) 165 (148.5) 30 (29.2) Other 162 (144.7) 303 (319.5) 62 (62.8) 9.21°° Yellow 88 (71.1) 148 (157.0) 23 (30.9) Other 124 (140.9) 320(311.0) 69 (61.1) 9.82°° 390 Great Basin Naturalist Vol. 43, No. 3 km at Lehi) in the April-September period (Whaley and Lytton 1979). Conditions that may favor wind-pollinated species at Arches include dominance of most perennial covers by a few woody species that have large pop- ulations, low-growing, open vegetation, and severe, unpredictable periods of drought. Ac- curate wind movement readings were not available for the Bald Mountain area. When only perennial species are consid- ered, woody species are much better repre- sented in the Arches flora (22.6 percent) than at Mt. Nebo (14.5 percent) or Bald Mountain (9.4 percent). Furthermore, shrubs dominate all major communities at Arches (Allan 1977). Diversity of perennial species as mea- sured by number of species per 1.0 m^ (a var- iable known to reduce the success of wind- pollinated taxa [Ostler et al. 1982]) is shown by Allan (1977) to be 2.6 at Arches (10 com- munities considered) as compared with 4.3 in the Wasatch Moimtains (of which Mt. Nebo is a part [Ostler and Harper 1978, 25 com- munities reported]) and 6.9 at Bald Mountain (Pendleton 1981, 4 communities reported). Wind pollination is further facilitated at Arches by a plant cover that is more open than that at Nebo. Allan (1977) reported an average of 41.3 percent living cover at Arches, but, considering the fact that 70 per- cent of the Mt. Nebo study area is dominated by oak woodland or forests of aspen and/ or conifer, plant cover there undoubtedly aver- ages well over 65 percent (see Allan 1962, Crowther and Harper 1965, Kleiner 1966, and Harper 1981 for cover estimates for sim- ilar vegetations in northern Utah). Vegetative cover in subalpine meadow in the Uinta Moimtains averages about 76 percent (Ostler et al. 1982). Finally, annual precipitation is more likely to fall below a level sufficient to support flowering of many species at Arches than in the mountain study areas. Frequent years of sparse or no flowering distributed at random through time should reduce the like- lihood that insect pollinators can maintain large enough populations to pollinate all the flowers produced in years of adequate soil moisture. Wind-pollinated species should be favored in such situations provided individual plants are large enough to intercept a re- liable flow of air, foliage cover is not so dense that it seriously interferes with pollen move- ment in the wind, and conspecific individuals are close enough together to insure that most stigmas will receive pollen. On all counts, the Arches area is better suited for wind pollina- tion than the two mountain locations. Flower Structure.— If reproduction of animal-pollinated species is to be successful, floral structure should encourage the likeli- hood of sequential visits by specific pollina- tors. When flowers of coexisting species com- pete for pollinators, species having flowers that mechanically exclude many kinds of pol- linators should be able to conserve more nec- tar or pollen for adapted visitors than species whose flowers can be worked by any visitor. Thus visits by such adapted pollinators should be reinforced by more dependable nectar or pollen rewards, thus encouraging the pollinator to seek out other flowers of the same type. As a result, flowers with restricted access should be at a reproductive advantage in diverse assemblages of plants that flower simultaneously. Ostler and Harper (1978) show that flowers that have restricted access to the nectar and/ or pollen supply are positively corre- lated with the diversity of animal-pollinated species per unit area in the Wasatch Moun- tains of Utah and Idaho. Thus, one might be tempted to hypothesize that, because diver- sity is lower at Arches than in Utah moun- tains as noted above, one could expect an overrepresentation of open-structured flow- ers in the desert. The data show, however, that when all species are considered, the mountain floras have relatively more species with open flowers than the desert (ZX^ = 19.11, P < 0.005, Table 3E). Even among perennial species only, that pattern continues to hold (2X2 = 14.35, P < 0.005, Table 3E). The difference in flower structure among the floras of Arches, Bald Mountain, and Mt. Nebo may be attributable to differences in flowering phenology. Species must flower when moisture conditions are favorable. In the desert, moisture conditions for most spe- cies are apparently optimal in the spring, be- cause it is at that time that most desert spe- cies flower. Accordingly, although there are fewer species per imit area in the desert, more species may actually flower simultane- ously than in the mountain zone. Thus, at July 1983 Collins et al.: Desert and Mountain Floras 391 certain times, there may be greater com- petition among plant species for pollinating animals in the desert than in at least mid- elevation moimtains. In contrast to deserts, midelevation moun- tain commimities have favorable moisture conditions throughout much of the growing season. It is therefore possible for coexisting species to partition the available time by flowering out of synchrony. Such out-of- phase flowering should decrease competition for pollinators and allow for more open flow- ers (Mosquin 1971). This argument probably does not hold at high elevations where grow- ing seasons are short and flowering of all spe- cies is confined to that brief season. In such environments, simultaneous flowering of many species is undoubtedly commonplace. The profusion of open-flowered taxa in the subalpine meadows (Table 2) thus cannot be attributed to low flowering-plant diversity. It has become clear in recent years that pollinating insect faunas are larger and more diverse in warmer and lower elevation envi- ronments than in cold and/ or high elevation habitats (Arroyo et al. 1982, Warren et al. 1982). Hymenopteran insects especially ap- pear to decline in colder and higher elevation communities, and Dipteran and Lepidopte- ran pollinators become relatively more im- portant. It has long been recognized that Hymenopteran pollinators are the most ef- ficient of all insects at manipulation of com- plex flowers (Faegri and Pijl 1971). It is thus possible that the observed predominance of open flowers in our mountain floras is related to changes in composition of the pollinator fauna along the altitudinal gradient. Open flowers may be favored when there is com- petition among flowers for an impoverished and less efficient guild of pollinating insects. Flower Symmetry.— The native flora of Arches National Park consists of 88.7 percent radially symmetrical and 11.3 percent zygo- morphic-flowered species. The Mt. Nebo flora includes 84.4 percent radial and 15.6 percent zygomorphic flowers. Finally, the Bald Mountain Flora consists of 83.7 percent radial and 16.3 percent zygomorphic flower (Table 2). Although differences between the three floras are not statistically significant (2X2 = 2.42), there is a trend toward a great- er nmnber of zygomorphic flowers as eleva- tion and diversity increases. Zygomorphic structure is believed to force insects to ap- proach flowers in more stereotyped ways. Thus, in zygomorphic flowered taxa, muta- tions that result in loss of stamens and stigmas off the regular access route of the insect to the floral reward (nectar and/or pollen) could be accommodated without loss in fe- cundity. In harsh environments where car- bohydrate gains are marginal, zygomorphy and the energy economy associated with re- duced numbers of reproductive parts and lower pollen production could be expected to have selective advantages. Flower Color.— Is there any difference in the distribution of colors among these floras? Ostler and Harper (1978) showed that floral color diversity was positively correlated with species diversity of the communities studied. Our data demonstrate that red and yellow flowers are overrepresented at Arches Na- tional Park (Table 3F), with the differences being statistically significant for both (P < 0.05 or better). White flowers are significant- ly overrepresented at Mt. Nebo and Bald Mountain. White seems to be favored in moist and wooded environments (Ostler and Harper 1978, Del Moral and Standley 1979), whereas yellow flowers are consistently bet- ter represented in dry environments (Kevan 1972, Tikhomirov 1966, and Weevers 1951). The results for white flowers were expected in the Nebo flora (but not at Bald Mountain meadow), because white color had been shown to be more abundant in forest under- stories by both Ostler and Harper (1978) and Del Moral and Stanley (1979). The Mt. Nebo complex is largely dominated by forest or woodland commimities, but there is only a minor amount of woodland at Arches Nation- al Park and Bald Mountain. Ostler and Har- per (1978) speculated that white flowers re- flected more light and were more easily found by pollinators in forest understories. Baker and Hurd (1968) noted that moths tend to replace bees as major pollinators in shaded habitats, and moths show a preference for white flowers. Finally, it is possible that the insect eye, like that of the human, is unable to perceive color at low light intensities (Proctor and Proctor 1978). Thus, the insect may react to flower color in shaded environ- ments in terms of contrast alone rather than in terms of different hues per se. 392 Great Basin Naturalist Vol. 43, No. 3 The abundance of white flowers in the sub- alpine meadows cannot be explained by the foregoing argument. At this point, we can only hypothesize that the importance of white flowers and the diminished abundance of yellow-flowered species in high elevation meadows is somehow related to the cooler, more moist environment or to altered com- position of the pollinating insect community (or both) at high elevations. The proportionally greater number of red and, to a lesser extent, yellow flowers in the Arches flora seems attributable to the near- ness of the park to areas of high diversity of hummingbird species in Arizona (up to nine species, see Crosswhite and Crosswhite 1981). Hummingbirds are believed to have been the selective force responsible for the evolution of many orange- and red-flowered species in the American Southwest (Crosswhite and Crosswhite 1981). At the mountain sites, such "hot"-colored species are relatively less com- mon (Table 2). Only four species of hum- mingbirds occur in the Mt. Nebo area, and three hummingbird species occur in- frequently at the high subalpine meadows considered here. Conclusion The results demonstrate that there are def- inite differences in the distribution of plant longevity, life form, mode of pollination, flor- al structure, and flower colors in the desert and moujitain floras compared in this study. Annuals and shrubs are overrepresented in the Arches National Park flora, and zoophily is significantly more abundant in the moun- tain floras. Open-structured flowers are sig- nificantly overrepresented on the mountain floras. White flowers are most common in the mountain floras, whereas red and yellow flowers are best represented in the desert. Acknowledgments Sincere appreciation is extended to C. Da- vidson, Arthur Holmgren, and others who re- viewed this manuscript and made valuable suggestions. This paper was funded in part by the U.S. Forest Service, Uinta National For- est, Provo, Utah. Literature Cited Allan, J. S. 1977. The plant communities of Arches Na- tional Park. Unpublished dissertation. Brigham Young Univ., Provo, Utah. 1962. The plant communities of the Big Cotton- wood Canyon drainage. Unpublished thesis. Univ. of Utah. Salt Lake City. Allred, K. W. 1975. Timpanogos flora. Unpublished thesis. Brigham Young Univ., Provo, Utah. Arroyo, M. T. K., R. Primack, and J. Armesto. 1981. Community studies in pollination ecology in the high temperate Andes of central Chile, I. Pollina- tion mechanisms and altitudinal variation. Amer. J. Bot. 69:82-97. Baker, H. C, and P. H. Hurd, Jr. 1968. Intrafloral ecol- ogy. Ann. Rev. Entomol. 13:511-523. Callison, J., and K. T. Harper. In review. Temper- ature in relation to elevation in the Uinta Moun- tains, Utah. Brigham Young Univ., Provo, Utah. Collins, P. D. 1979. Checklist of the plants on Mt. Nebo and adjacent mountains. Unpublished re- port to the Uinta National Forest, Provo, Utah. Collins, P. D., and K. T. Harper. 1980. Vegetation map of the Mt. Nebo complex and San Pitch Mountains, Utah. BYU Printing Service, Provo, Utah. Crosswhite, F. S., and C. D. Crosswhite. 1982. Hum- mingbirds as pollinators of flowers in the red- yellow segment of the color spectrum, with spe- cial reference to Pensteinon and the "open habi- tat." Desert Plants 3:156-170. Crowther, E. C, and K. T. Harper. 1965. Vegetational and edaphic characteristics associated with aspen "strips" in Big Cottonwood Canyon. Proc. Utah Acad. Sci., Arts, Lett. 42(II):222-230. Del Moral, R., and L. A. Standley. 1979. Pollination of angiosperms in contrasting coniferous forests. Amer. J. Bot. 66:26-35. Faegri, K., and L. van der Pijl. 1971. The principles of pollination ecology, 2d ed. Pergamon Press, Oxford. Freeman, D., K. T. Harper, and W. K. Ostler. 1980. Ecology of plant dioecy in the intermountain re- gion of western North America and California. Oecologia 44:410-417. Harner, R. F., and K. T. Harper. 1973. Mineral compo- sition of grassland species in the eastern Great Basin in relation to stand productivity. Canadian J. Bot. 51:2037-2046. Harper, K. T. (ed.) 1981. Potential ecological impacts of snowpack augmentation in the Uinta Mountains, Utah. Final report to the Bureau of Reclamation, Engineering and Research Center, Federal Cen- ter, Denver, Colorado. Harrison, B. F., S. L. Welsh, and G. Moore. 1964. Plants of Arches National Monument. BYU Sci. Bull., Biol. Ser. 5(1): 1-23. HiRONAKA, M. 1963. Plant environment relations of ma- jor species in sagebrush-grass vegetation of south- ern Idaho. Unpublished dissertation. Univ. of Wisconsin, Madison. Hutchings, S. S., and G. Stewart. 1953. Increasing for- age yields and sheep production on inter- mountain winter ranges. USDA Circ. No. 925. July 1983 Collins et al.: Desert and Mountain Floras 393 Jaccard, p. 1912. The distribution of the flora of the al- pine zone. New Phytol. 11:37-50. Kevan, p. G. 1972. Floral colors in the high arctic with reference to insect-flower relations and pollina- tion. Canadian J. Bot. 50:2289-2316. Kleiner, E. F. 1966. A study of the vegetational com- munities of Red Butte Canyon, Salt Lake County, Utah. Unpublished thesis. Univ. of Utah, Salt Lake City. MosQUiN, T. 1971. Competition for pollinators as a stim- ulus for the evolution of flowering time. Oikos 22:398-402. MuEGGLER, W. F. 1972. Plant development and yield on mountain grasslands in southwestern Montana. U.S. Forest Serv. Paper INT-124. National Oceanic and Atmospheric Administration. 1971-79. Annual .summaries for Utah. Climatolo- gical Data 73-81(13):pp. variable. Ostler, W. K., and K. T. Harper. 1978. Floral ecology in relation to plant species diversity in the Wasatch Mountains of Utah and Idaho. Ecology 59:848-861. Ostler, W. K., K. T. Harper, K. B. McKnight, and D. C. Anderson. 1982. The effects of increasing snowpack on a subalpine meadow in the Uinta Mountains, Utah, USA. .\rctic and Alpine Res. 14:20.3-214. Pendleton, B. K. 1981. Aspects of the floral biology of a subalpine meadow of the Uinta Mountains, Utah. Unpublished thesis. Brigham Young Univ., Provo, Utah. Proctor, J., and S. Proctor. 1978. Color in plants and flowers. Everest House Publishers, New York. ScHAFFER, VV. M., and M. D. Gadgil. 1975. Selection for optimal life histories in plants. Pages 142-157 in M. L. Cody and J. M. Diamond, eds.. Ecology and evolution of communities. Belknap Press, Cambridge, Massachusetts. Sharif, C. M., and N. E. West. 1968. Forage moisture variations on mountain summer range. J. Range Mgt. 21:228-2.35. SoLBRiG, O. T. 1977. On the relative advantages of cross- and self-fertilization. Ann. Missouri Bot. Card. 63:262-276. TiKHOMiRov, B. A. 1966. The interrelationships of the animal life and vegetational cover of the tundra. Academy of Sciences of the USSR, Botanical In- stitute (Translated from Russian by the Israel Program for Scientific Translation, Jerusalem, 1966). U.S. Congress. Public Law 95-87. 91 Stat. 445. 95th Cong., 3 Aug. 1977. Warren, S. D., G. M. Booth, and K. T. Harper. 1982. Elevational distribution of insect pollinators. Un- published manuscript. Whaley, B. L., and L. Lytton. 1979. Summary of pre- cipitation measurements for Utah, 1930-1978. Report distributed by the Snow Survey Section, Soil Conservation Service, USDA, Salt Lake City, Utah. 168 pp. Weevers, T. H. 1952. Flower colours and their frequen- cy. Acta Bot. Neerl. 1:81-92. FLORA OF THE LOWER CRETACEOUS CEDAR MOUNTAIN FORMATION OF UTAH AND COLORADO, PART I. PARAPHYLLANTHOXYLON UTAHENSE G. F. Thayne', W. D. Tidwell", and W. L. Stokes' Abstract.— ParapItyUantJwxijIon utahense, sp. nov., is described from the Cedar Mountain Formation and com- pared with similar fossil and modern woods. Fossil angiosperm woods from the Early Cretaceous are of great interest because very few have been reported from strata of this age. This species demonstrates that the angiosperms had de- veloped many of their modern characteristics by Early Cretaceous time. The Lower Cretaceous Cedar Mountain Fonnation is fossiliferous at several localities. Fossils reported from this formation include the wood of conifers, Tempskya, and cyca- deoids, charophytes, pelecypods, gastropods, ostracods, and fish scales (Stokes 1952, Young 1960, Thayn et al. 1973, Tidwell et al. 1976), as well as dinosaur bones (Bodily 1969). A species of dicotyledonous wood assigned to the genus Paraphyllanthoxylon Bailey 1924, is described in this report from the Ce- dar Moimtain Formation. This is the first re- port of petrified dicotyledonous wood from the diverse flora in this formation. These angiosperm woods are of great interest in that very few Early Cretaceous angiosperm woods have been previously reported. Since the Cretaceous Period is the assumed time for the origin of the angiosperms, a tax- onomic study of Early Cretaceous angio- sperm wood is significant in that it expands oiu- knowledge of the early members of this division. The petrified wood described in this study was collected from two localities. Lo- cality 1 is 6 road miles (3.7 km) east of Castle Dale, Utah, and Locality 2 is 9 road miles (5.6 km) east of Ferron, Utah (Figs. 1, 5, 6, 7). The Cedar Mountain Formation at Local- ity 1 is composed of brown to grey shales. It contains at least one horizon of nearly coali- fied material from which Tempskya has been collected in growth position (Tidwell and Hebbert 1976). The dicotyledonous woods studied here were collected from a horizon between 10 (3.1 m) and 30 (9.2 m) feet below the overlying Dakota Sandstone, which is represented by 10 (3.1 m) to 20 (6.2 m) feet of coarse brown sandstone that forms a cap rock in the area. Fig. 1. Index map of collection sites. 'Bureau of Land Management, Salt Lake City, Utah 84112. 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 'Department of Geology, University of Utah, Salt Lake City, Utah 84112. 394 July 1983 Thayne et al.: Cretaceous Flora 395 Ulah 0 Prove Arizono Fig. 2. Geographic extent of the Cedar Mountain Formation (after Young 1960). Specimens were collected from seven dif- ferent logs at Locality 2, where the Cedar Mountain Formation consists of a bed of coarse white sandstone underlaid by channel fills of yellow conglomeritic sandstones alter- nating with grey-green shales. These are, in turn, underlaid by a dark green nodular weathering shale (Figs. 5-7). The Dakota Sandstone is missing at this particular site, but reappears in the section about 3 miles (1.9 km) to the southeast. Dicotyledonous woods were found associated with Tempskya and fossil conifer wood at this locality. Stratigraphy of the Cedar Mountain Formation Stokes (1944) defined the Cedar Mountain Formation as those sediments lying between the Brushy Basin Member of the Upper Juras- sic Morrison Formation and the Lower Cre- taceous Dakota Formation. These strata were formerly considered part of the Morrison Formation. Cedar Mountain deposits are present over much of eastern Utah, western Colorado, and northwestern New Mexico Fig. 3. ParaphijUanthoxijlon i/ta/iense— Ilhistration of the transverse section showing the relative abundance of ray tissue (dark areas) and the size, shape, and arrange- ment of the vessel elements (open circles). Note the radi- al pore multiples. (Fig. 2). Stokes (1952) defined two members of the formation, the Buckhorn Con- glomerate at the base and the Cedar Moun- tain Shale at the top. At the type locality near Castle Dale, Utah, the Buckhorn Con- glomerate is thick and massive, but it gener- ally thins to the east and is absent on the eastern side of the San Rafael Swell. East of the Colorado River a mudstone and con- glomeritic sandstone unit occupies the same Fig. 4. ParaphijUanthoxijlon lita/iense— Illustration of the tangential section showing the size, shape, and ar- rangement of the rays (dark lines) and a vessel element (center). 396 Great Basin Naturalist Vol. 43, No. 3 W" -* Fig. 5. Ferron collecting site, showing the lithology. A is white sandstone cap, B is the surface of the yellow conglomeritic channel fill from which the specimens were recovered, and C is the underlying grey-green shale. relative position as the Cedar Mountain For- mation on the west side of the river. These rocks were termed the "Post-McElmo" beds (Coffin 1921), but were later renamed the Burro Canyon Formation (Stokes and Phoe- nix 1948). Young (1960) proposed that the Burro Canyon Formation and the Cedar Mountain Formation are a physically contin- uous unit and should both be referred to as Cedar Mountain Formation. Based on the presence of the pelecypods Protoelliptio douglassi, Unio farri, and the conifer Frenelopsis varians, as well as the stratigraphic position of the Cedar Mountain Formation, Young (1960) as well as Stokes (1952), suggested that it is Lower Cretaceous in age. Another indication of its age is the presence of Tempskya, which Read and Ash (1961) considered to be an index fossil to the Lower Cretaceous (Albian). Fisher et al. Fig. 6. Overview of the Ferron collecting site. Snow- capped mountains in the background are in the Wasatch Plateau. Dicotyledonous logs along with Tempskya were collected from tlie uppermost layer of sediment shown in the foreground. Fig. 7. Petrified dicotyledonous wood shown as it is found weathered upon the surface of the Ferron site. (1960) listed the formation as Aptian, but it may be only Albian or, most probably, may include rocks of both ages. Paraphyllanthoxylon utahense sp. nov. Figs. 3-4, 8-18 Description.— This species is described from several pieces of black petrified second- ary wood. The preservation is excellent, and fine stRictural detail can be observed. Growth rings: Lacking. Vessels: Diffuse porous, approximately 12 /mm 2, solitary or more commonly in radial rows (pore multiples) of 2-3 up to 5 cells long; individual vessels range from 204 jum radial by 165 jum tangential diameter to 58 jum radial by 48 jum tangential, average 105 jum radial by 93 jum tangential diameter; per- forations exclusively simple, located on oblique end walls; thin-walled tyloses abun- dant, obscuring the vessel length; vessel walls 3 ium-5 jum thick; tangential pitting with nu- merous, often appressed, 6 jum-10 /xm diame- ter; alternate bordered pits with slitlike aper- tures and occasionally up to 12 |u,m long, slightly bordered pits with large elliptic aper- tures probably representing the vessel to pa- renchyma pitting; radial intervascular pitting similar to tangential; vessel to ray inter- vascular pitting similar to tangential; vessel to ray pitting consisting of small circular or large, up to 24 jum, scalariform, elliptic to an- gular slightly bordered pits; 3-6, occasionally more, pits per crossover field. Axial Parenchyma: Rare, apotracheal dif- fuse or scanty paratracheal. Rays: 12/mm2, heterogeneous with both uniseriate and multiseriate present; uniseriate July 1983 Thayne et al.: Cretaceous Flora 397 Fig. 8. Transverse section illustrating solitary vessels and vessel chains with tyloses. Note that the axial paren- chyma is scarce (65X). rays rare, many partially biseriate, with both procmnbent and upright cells, uniseriate rays range from 2 cells (80 jum) to 6 cells (300 jum) high (average 5 cells, 200 jam-220 jum); mul- tiseriate rays range from 9 cells (380 jum) to 33 cells (1360 [xm) high and 2 cells (30 jum) to 5 cells (100 jum), wide with 106 rows (com- monly 2) of upright border cells; procumbent cells range from 25 jam-40 jum vertical, 50 jum-80 jLim radial, and 25 jum-45 jUm tan- gential diameter; some cubodial cells present, approximately 40 jum in diameter; upright cells approximately same size as procumbent but radial and vertical dimensions reversed; ray cells' walls 2.5 jum thick, pitted and ap- pearing beaded in radial section. Fibretracheids: Septate, libriform, round to square in cross section, approximately 36 /xm in diameter, with approximately 2.4 jum thick walls. Repository: Brigham Young University, 2190 (Holotype) Horizon: Cedar Moimtain Formation Age: Early Cretaceous Discussion Twelve species of ParaphyUanthoxylon have been described in the past. The features generally constant in all of these reported species are as follows: Fig. 9. Transverse section illustrating distribution of vessels and multiseriate rays (30X). Diffuse porous wood; vessels in radial rows (pore multi- ples); exclusively simple perforations; alternate inter- vascular pitting; elongate vessel to ray pitting; rays of two sizes, 1-7 seriate, heterocellular with 107 rows of upright border cells, rays commonly over I mm high, ax- ial parenchyma lacking or scanty apotracheal diffuse, scanty paratracheal, or combination of both; septate fibretracheids; vessels commonly with tyloses. Paraphijllanthoxijlon utahense fits well with- in the boundaries of this genus. Comparison with Described North American Species Three species of ParaphyUanthoxylon have been described from Cretaceous strata in North America. ParaphyUanthoxylon arizonense Bailey 1924.— ParaphyUanthoxylon utahense differs from the upper Cretaceous P. arizonense in several ways. The most obvious variations are the size of the vessels and rays. Although Bailey (1924) gave no measurements and merely stated that the vessels of P. arizonense are large, it can be seen that the vessels shown in his figures at 35X are almost as large as those of P. utahense at 65X. Also, the rays shown at 35X in his figures are approx imately twice as high and wide and more parallel in outline than those of P. utahense at a comparable magnification. Another dif- ference is that P. arizonense has slitlike pits on the fiber walls that P. utahense lacks. On the basis of these differences, the Utah speci- mens have been determined to be distinct from P. arizonense. ParaphyUanthoxylon idahoense Spackman 1948.— ParaphyUanthoxylon utahense is closer to P. idahoense, which was reported 398 Great Basin Naturalist Vol. 43, No. 3 Fig. 10. Transverse section illustrating a closeup view of fibretracheids and a heterocellular multiseriate ray (495X). from the Lower Cretaceous Wayan Forma- tion of Idaho, than to other Para- phyllanthoxylon species. The diameter of the vessels in P. idahoense is 60 jum-160 jum. Fig. 11. Radial section with irregularly shaped vessel to ray cell crossover pits inked in. The beaded nature of the ray cell walls can also be seen (lOOX). Fig. 12. Radial section illustrating the relative size and distribution of the vessels and cells of the hetero- cellular rays (33X). However, in comparing the holotype slides of this species with those of P. utahense, it can be seen that the vessels of the former are dis- tinctly larger than those of the latter, which are 48 ]u,m-165 /xm in tangential diameter. The pitting is similar in both species, and the intervascular pits are also alternate, circular elliptical, and sometimes compacted and an- gular. The bordered pits of P. idahoense are 10 ju,m-12 jum in diameter, whereas, those in P. utahense vary from 6 /xm to 10 /xm in di- ameter. The major differences between these two species are the compaction of the vessels and the size of the rays. The number of ves- sels per square millimeter was not given for P. idahoense, but its vessels are more tightly compacted than those in P. utahense. The rays in P. idahoense are made up of smaller cells and are narrower than those of P. uta- hense, although both have multiseriate rays from two to five cells wide. Since P. utahense has smaller vessels that are fewer in number per square millimeter, and larger rays than P. idahoense, these two species are considered distinct from one another. Paraphyllanthoxylon alabamense Gaboon 1972.- As described by Gaboon (1972) from the Upper Gretaceous Tuscaloosa Formation, this species has a wide range of variation, which can be seen by comparing Figures 5, 11, and 14 of her paper. These photos are all July 1983 Thayne et al.: Cretaceous Flora 399 Fig. 13. Radial section illustrating the irregularly shaped, narrowly bordered vessel to parenchyma pits (495X). listed as being magnified 55X, but the rays in Figure 5 are approximately five times wider than those in Figure 14. She stated that this species was described from 11 different type specimens that are all similar but show some variation. The most obvious variation is in the size of the rays. Barghoorn (1941) has shown that such variation could conceivably be found within a species or even within the trunk of an individual tree, but, since pa- leobotanists are often restricted to working with fragments, they have traditionally de- scribed such fragments as form genera and hence form species. Spackman (1948) distin- guished P. idaJioense from P. arizonense be- cause P. idahoense has smaller vessels, less abundant pitting, and smaller rays and ray cells. He stated that: The magnitude and nature of these variations are well within the range of variability found in individuals of many living species, and thus the differences in the two fossils might be accounted for on the basis of the part of the tree from which the specimen was derived, differ- ences in growth rate, etc. In spite of this, however, it seems appropriate, because of these differences to de- scribe this new wood as a new species with the hope that the tnie relationship of these two fossils will be demonstrated in the hiture. (Spackman, 1948, p. 108). We agree with Spackman's reasoning, and therefore believe that P. alabamense as it now stands includes at least two or three Fig. 14. Radial section illustrating oblique simple per- foration plates and oppositely arranged bordered inter- vascular pits with slitlike apertures on the radial vessel wall (495X). form species. Therefore, P. utahense cannot be accurately compared to it at this time. The holotype specimen shown by Gaboon (1972) appears to differ from P. utahense by having larger rays. The other specimens re- ported by her appear similar to P. utahense, although one has larger rays and the other has smaller. Before any conclusions can be drawn as to the species boundaries and rela- tionship between P. utahense and the Ala- bama specimens, more detailed measure- ments and comparisons need to be made. Paraphyllanthoxylon pfefferi Platen 1908.— This species was collected from the Tertiary of California. It was originally de- scribed as Carpinoxylon pfefferi (Platen 1908), but was combined with Para- phyllanthoxylon by Madel (1962). Para- phyllanthoxylon utahense has larger vessels (up to 204 jum radial diameter as opposed to 137 jLim radial diameter), which are fewer per square millimeter (12 compared to 44), and broader rays (100 jum compared to 50 jum) than P. pfefferi. Comparison With Other Paraphyllantho- xylon Species.— Paraphyllanthoxylon uta- hense differs from species of Para- phyllanthoxylon described from geographical areas beyond the boundaries of North Ameri- ca (Table 1) in such features as the size and density of the vessels and dimensions of the 400 Great Basin Naturalist Vol. 43, No. 3 Fig. 15. Tangential section illustrating the relative size and distribution of multiseriate rays, fibretracheids, and vessels with tyloses (30X). rays. Paraphyllanthoxylon utahense is most similar to P. capense but differs in having fewer vessels per square millimeter, and ves- sels that are larger and fewer per pore multiple. Affinities of Paraphyllanthoxylon.— The original species, Paraphyllanthoxylon arizo- nense, was described by Bailey (1924) from silicified wood fragments of the Colorado Group in Arizona. He proposed the name to indicate a relationship to Bridelia and Phyl- lanthus in the section Phyllanthoidea of the Euphorbiaceae. Madel (1962) combined woods which had been described as Phyllan- thinium and Glochidioxylon into the genus Paraphyllanthoxylon and reserved the genus for woods with general structure of the Glo- chidion wood group of the Euphorbiaceae. Other authors have compared their species to a number of genera in several other families. Although Paraphyllanthoxylon alahamense may be an aggregation of species, further in- formation concerning the affinities of the genus may be inferred by the leaf com- pressions that occur along with it in the Tus- caloosa sediments. Gaboon (1972) reported Fig. 16. Tangential section ilkistrating the hetero- cellular rays and septa in the fibretracheids (lOOX). that of the families with wood similar to Par- aphyllanthoxylon only the Sapindaceae, Eu- phorbiaceae, and Lauraceae are represented by fossil leaves from the Tuscaloosa. Phylogenetic Considerations.— The pro- cesses of convergent and divergent evolution have obscured the genealogy of even modern genera and species. Pax and Hoffman (1931) considered the Euphorbiaceae to be poly- phyletic in origin, making it imlikely that the Lower Gretaceous Paraphyllanthoxylon spe- cies are ancestral to the various groups with- in the family. The possibility does exist that they are ancestral to at least some members of the Glochidion group. Considering the large number of genera that are similar to the genus, Paraphyllanthoxylon could be related to the taxon from which several genera in many families originated. By comparing the features of Para- phyllanthoxylon with Tippo's (1946) list of primitive and advanced wood characteristics (Table 2), it can be seen that the anatomy of the Lower Gretaceous members of the genus supports evidence from fossil leaf com- Table 1. Paraphyllanthoxylon species from outside of North America. Species Author Occurrence pseudohohash iraish i sahnii tertiunim bangalamodense ■ keriense capense yvardi teldense Ogura, 1932 Prakash, 1958 Ramanujam, 1956 Navale, 1960 Dayal, 1968 Madel, 1962 Koeniguer, 1967 Prive, 1975 Tertiary of Japan Tertiary of India Tertiary of India Tertiary of India Tertiary of India Upper Cretaceous of S. Africa Neogene of France Oligocene of France July 1983 Thayne et al.: Cretaceous Flora 401 Fig. 17. Tangential section illustrating various sizes and shapes of rays and dark cell contents in many of the ray cells (SOX). pressions that the angiosperms had developed many of their modern characteristics by Early Cretaceous times. Acknowledgments The authors express appreciation to S. R. Rushforth, J. D. Brotherson, and J. Keith Rig- by of Brigham Young University and S. R. Ash of the Department of Geology, Weber State College, Ogden, Utah, who graciously reviewed the manuscript. We also express appreciation to Naomi Hebbert who helped in preparation of the illustrations. Table 2. Comparison with primitive and advanced features. Primitive Advanced 4 ^ ci- 3 J- ^ « _ - T3 e Us ^ c 2 o So.^ o i; c/2 H &< < iZ 0; ._ 15 "3 ;s Sfi "5 5P § c/5 O c/2 £-4 c/i 3- Z Fig. 18. Tangential section showing appressed, op- positely arranged, bordered pits with slitlike apertures on the tangential vessel wall (495X). Literature Cited Bailey, I. W. 1924. The problem of identifying the wood of Cretaceous and later dicotyledons: Para- phifUanthoxylon arizonense. Ann. Bot. .38: 4.39-451. Barghoorn, E. S. 1941. The ontogenetic development and phylogenetic specialization of rays in the xy- lem of dicotyledons. II. Modification of the mul- tiseriate and uniseriate rays. Amer. J. Bot. 28:273-282. Bodily, N. M. 1969. An armored dinosaur from the Lower Cretaceous of Utah. Brigham Young Univ. Geol. Studies 16(.3):35-60. Cahoon, E. J. 1972. Paraphyllanthoxylon alabamense: a new species of fossil dicotyledonous wood. Amer. J. Bot. 59(1):5-11. Coffin, R. C. 1921. Radium, uranium and vanadium de- posits of southwestern Colorado. Colorado Geoft Surv. Bull. 16. Dayal, R. 1968. A new fossil wood of the Eu- phorbiaceae from the Deccan Intertrappean beds of Madhva Pradesh. Palaeobotanist 16(2): 148-150. Fisher, D. J., D. E. Erdmann, and J. B. Reeside. 1960. Cretaceous and Tertiary formations of the Book Cliffs, Carbon, Emery, and Grand counties, Utah, and Garfield and Mesa counties, Colorado. U.S. Geol. Surv. Prof. Paper .3.32. 1-80. KoENiGUER, J. C. 1967. Etude paleoxylologique de la Fouraine: Paraphyllanthoxylon yvardi, n. sp. C. R. 92c Congress Nat. Soc. Sav., Strasbourg, 3:71-75. Madel, E. 1962. Die fossilen Euphorbiaceen-Holzer niit besonderer Benicksichtigung neuer Funde aus der Oberkreide Sud-Afrikas. Senkenberg. Leth 43(4):283-321. Navale, G. K. B. 1960. Phyllanthinium bangalamo- dense, a new species of fossil Euphorbiaceous wood from the Cuddalore Series of India. Pa- laeobotanist 9:11-16. 402 Great Basin Naturalist Vol. 43, No. 3 Ogura, Y. 1932. On the stnicture of a silicified wood found near hobashiraishi at Najima near Fukuoka City. Japanese J. Bot. 6(2): 18.3-190. Pax, F., and K. Hoffman. 1931. Euphorbiaceae. In A. Engler and K. Prantl: die naturlichen pflanzenfa- milien, 2Aufl. 19(c): 11-233. Platen, P. 1908. Untersuchungen fossiler Holzer aus deni Westen der Vereinigten Staaten von Nor- dainerika. Diss: 1-55. Prakash, U. 1958. Studies in the Deccan Intertrappean Flora, 4: two silicified woods from Madhya Pra- desh. Palaeobotanist 7(1): 12-20. Prive, C. 1975. Sur la presence dun bois fossile D Eu- phorbiaceae dans loligocense du Cantal. Act. 95th Natl. Cong. Soc. Savantes 1970(,3):111-125. Ramanujam, C. K. G. 1956. Fossil woods from the Eu- phorbiaceae from the Tertiary rocks of south Ar- cot District, Madras. J. Indian Bot. Soc. 35(3):284-307. Read, C. B., and S. R. Ash. 1961. Stratigraphic signifi- cance of the Cretaceous fern Tempskya in the western conterminous United States. U.S. Geol. Surv. Res. 8.32:D-250-D-254. Spackman, W., Jr. 1948. A dicotyledonous wood found associated with the Idaho Tempskyas. Ann Mo. Bot. Card. .35:107-116. Stokes, W. L. 1944. Morrison and related deposits in and adjacent to the Colorado Plateau. Ceol. Soc. Amer. Bull. 55:951-992. 1952. Lower Cretaceous in Colorado Plateau. Amer. Assoc. Pet. Geol. Bull. .36(9): 1774- 1796. Stokes, W. L., and D. A. Phoenix. 1948. Geology of the Engas-Gypsum Valley Area, San Miguel and Montrose counties, Colorado. U.S. Geol. Surv. Prelim. Map 93, Oil and Gas Inv. Ser. Thayn, G. F., W. D. Tidwell, and W. L. Stokes. 1973. Petrified angiosperm wood from the Lower Cre- taceous Cedar Formation of Utah. Geol. Soc. Amer. Abstr. with programs 5(6):519. Tidwell, W. D., and N. E. Herbert. 1976. Tempskija from the Lower Cretaceous Cedar Mountain For- mation, Utah. Amer. J. Bot. Abstr. 63:33. Tidwell, W. D., G. F. Thayn, and J. L. Roth. 1976. Cretaceous and Early Tertiary floras of the Inter- mountain Area. Brigham Young Univ. Geol. Stud- ies 22:77-98. Tippo, O. 1946. The role of wood anatomy in phylogeny. Amer. Midi. Nat. 36:362-372. Young, R. G. 1960. Dakota Group of Colorado Plateau. Amer. Assoc. Pet. Geol. Bull. 44(1): 1,56-194. ACULEATA HYMENOPTERA OF SAND MOUNTAIN AND BLOW SAND MOUNTAINS, NEVADA R. W. Rust', L. M. Hanks,i2, and R. C. BechteP Abstract.— There were 198 species of aculeata Hymenoptera in 15 families collected from Sand Mountain and Blow Sand Mountains, Nevada. Four species are considered new to science and none are considered endemic to ei- ther dune area. Sand Mountain and Blow Sand Mountains were visited 19 times in a 13-month period for the purpose of surveying selected groups of arthropods. Here we report on the acu- leate Hymenoptera collected during the study. Over 2,000 specimens were obtained, representing 198 species in 15 families. Four species are considered new to science and 21 species were identified as "species" in un- studied genera. Most of the unknown species were bees (Apoidea) in the genera Perdita, Dialictus, Sphecodes, and Hesperapis. None of the species is considered sand obligate or endemic to either dune. Previous studies on Hymenoptera in Ne- vada include the faunistic inventory of the Nevada Test Site conducted by Brigham Young University from 1958 to 1966 that produced almost 8,000 adults and 1,100 im- matures (Beck and Allred 1968). Of these, the ants (Cole 1966), mutillid wasps (Ferguson 1967, Allred 1973), tiphiid wasps (Wasbauer 1973), and bees (G. E. Bohart, pers. comm.) have been identified. Wheeler and Wheeler (1978) studied the mountain ants of Nevada and they have produced a manuscript on the ants of Nevada (to be published by Los An- geles Coimty Museum). Pretruszka (1980) ob- tained almost 2,000 specimens, identified to family, from Fairview Valley, Nevada, from pitfall and malaise trapping. Study Sites Sand Mountain (SM) dunes and Blow Sand Mountains (BSM) dunes were sampled from June 1979 through July 1980. Sand Mountain is approximately 46 km ESE of Fallon, Churchill County, Nevada (39°20'N- 118°20'W) and is about 1,250 m in elevation. Blow Sand Mountains are approximately 52 km SE of Fallon, Nevada (39°10'N- 118°35'W) and are about 1,400 m in eleva- tion. The dunes are separated by 25 km air distance. Sand Mountain is a star dune of ap- proximately 3.2 km^ and Blow Sand Moun- tains are complex star and linear dunes of ap- proximately 9.2 km^. However, both dunes result from the same eolian sand deposited during the Turupah and Fallon formations of about 4,000 years B.P. (Morrison and Frye 1965). The floras of the two dunes were similar. The dominant vegetation was Atriplex con- fertifolia (Torr. & Frem.), Tetradymia tetra- meres (Blake), Chrysothamnus viscidiflorus (Hookl), Astragalus lentiginousus Dougl., and Psoralea lanceolata (Pursh.), and at Sand Mountain only Eriogonum kearneyi Tidestr. and Psorotharnnus polyadeniiis (Torr.). The* common grass was Oryzopsis hyrnenoides (R. &S.). Methods Several collecting techniques were used to obtain specimens (see Bechtel et al. 1981). Permanent pitfall traps were 0.95 L plastic cartons. They were operative for 30-day peri- ods. Temporary pitfall traps were 15 cm di- ameter cereal bowls placed level with the sand surface. Temporary pitfall traps were 'Biology Department, University of Nevada, Reno, Nevada 89557. ^Present address; Department of Entomology, University of Maryland, College Park, Maryland 20742. 'Nevada State Department of Agriculture, .350 Capitol Hill Avenue, Reno, Nevada 89502. 403 404 Great Basin Naturalist Vol. 43, No. 3 used for 12-18 hr during a survey period. Two UV light traps were operated from dusk to dawn. Hand-held UV lamps were used in searching the dimes for fluorescing arthro- pods. Sand was sifted through two screens 12 X 12 cm and 1.5 X 1.5 cm mesh to recov- er subsurface arthropods. General collecting involved the use of aerial nets, plant in- spection, and walking the dunes at night with lamps to obtain specimens. During a survey period, four or five different sites on the dune were visited and the sites were varied each survey period. Data are presented in the following man- ner: For each species its known geographic distribution, present location, numbers ob- tained, dates of occurrence, and, for the bees, their flower visitation as determined from pollen-load analysis (Hanks and Rust 1983). Once a genus and species of plant is given as a pollen source, subsequent references to it is by use of the first two letters of the genus and species, e.g. Astraglus lentiginosus is Asle. Geographic distributions were obtained from the Hymenoptera Catalog (Krombein and Hurd 1979) and are presented as North America (NA), western United States (WUS), southwestern United States (SWUS), Great Basin (GB), or by individual state. For the ants, Formicidae, the number given repre- sents collections and not individual ants. The following taxonomists identified the material being presented: R. C. Bechtel (Mu- tillidae), G. E. Bohart (Apoidea), R. M. Bo- hart (Chrysididae, Vespidae, Sphecidae), A. A. Grigarick (Megachilidae), L. Kimsey (Chrysididae), W. E. LaBerge (Apiodea), A. S. Menke (Sphecidae), F. D. Parker (Sphe- cidae), J. G. Rosen (Andrenidae), R. W. Rust (Apoidea), R. R. Snelling (Apiodea, Formi- cidae), R. W. Thorp (Andrenidae), M. S. Wasbauer (Tiphidae, Scoliidae, Pompilidae), and G. C. and J. Wheeler (Formicidae). All specimens, except new specimens, are in the collection of the University of Nevada, Reno. Results The most numerous species obtained was the California harvester ant, Pogonomyrmex californicus (Buckley), with 26 collections containing hundreds of specimens. One per- manent pitfall trap contained over 500 indi- viduals. It was present at both dunes from May through October. Two other ants were common at SM Vero7nessor lariversi M. R. Smith and Conomyrma insana (Buckley). Glyptacros new species and Xeroglypta egr- egia Mickel and Krombein were perhaps the most interesting wasps collected. They both have wingless females and were collected by sifting sand. Other wingless females obtained, mutillids, were found active on the surface or attracted to UV light traps. Next to the ants, the most numerous species were Parnopes ftilvicornis Cameron (Chrysididae), Para- nistrocerus toltecus (Saussure) (Vespidae), Acanthetropis aequalis (Fox) (Tiphiidae), Sphaerophthalma sp. (Mutillidae), Micro- benibex argyropletira Bohart and Eucerceris nevadensis (Dalla Torre) (Sphecidae), and Agapostemon melliventris Cresson and Dia- lictus sp. (Halictidae). Chrysididae Elampus nitidus Aaron (WUS) SM 3 Aug. Oct. Holopyga hora Aaron (WUS) SM 1 May. Hedychridium amahile Gockerell (WUS) SM 5 Aug. Sept. Hedychridium arietinum Bohart (CA) SM 1 Sept., BSM 1 Sept. Hedychridium mancopae Bohart (CA) SM 2 April May. Hedychridium mirum Bohart (CA) SM 14 June July, BSM 1 June. Hedychridium species (?) BSM 1 June. Chrysis inflata Aaron (WUS) SM 1 Sept. Spintharosoma trochilus (duBuysson) (WUS) SM 1 April. Parnopes fulvicornis ftilvicornis Cameron (WUS) SM 132 June July Aug. Sept. Tiphiidae Brachysistis lacustris lacustris Malloch (Mo- jave desert) SM 3 July Aug. Sept., BSM 9 July Aug. Brachycistis triangularis Fox (SWUS) SM 4 June, BSM 10 June July. Brachycistis agama (Dalla Torre) (NA) SM 12 July Aug. Sept., BSM 57 July Aug. Brachycistis species (?) BSM 9 June. Colocistis crassa (Bradley) (SWUS) BSM 3 June. July 1983 Rust et al.: Nevada Hymenoptera 405 Acanthetropis aequalis (Fox) (WUS) SM 30 July Aug. Sept., BSM 31 July Aug. Sept. Acanthetropis noctivaga (Bradley) (SWUS) SM 16 July Aug., BSM 6 July Aug. Glyptacros new species (?) SM 5 Sept. Oct., BSM 16 Jime Aug. Sept. Xeroglypta egregia Mickel and Krombein (SoCA) SM 1 Aug. Mutillidae Sphaeroptluilma species (?) SM 57 June July Aug., BSM 26 Jime July Aug. Dasymutilla gloriosa (Saussure) (SWUS) BSM I Aug. Dasymutilla satanas Mickel (SWUS) SM 2 June July. Scoliidae Crioscolia alcione (Banks) (SWUS) SM 3 July Aug. Formicidae Pogonomyrmex californicus (Buckley) (SWUS) SM 26 May to Oct. BSM 14. Veromessor lariversi M. R. Smith (WUS) SM II June July Aug. Sept., BSM 2 June Aug. Creniatogaster species (?) SM 1 Sept., BSM 1 Aug. Conornyrrna hicolor (W. M. Wheeler) (WUS) SM 2 July Aug. Conornyrrna insana (Buckley) (WUS) SM 15 June July Aug. Sept., BSM 2 Aug. Camponotus vicinus Mayr (US) SM 5 Aug. Sept. Myromecocystus kennedyi Cole (WUS) SM 4 Feb. Aug., BSM 2 Aug. Myromecocystus pyramicus M. R. Smith (WUS) SM 3 Aug. Sept., BSM 2 Aug. Myromecocystus new species (?) SM 4 April May. Vespidae Pterocheilus crispocornis Bohart (SWUS) SM 1 June, BSM 1 Aug. Pterocheilus diversicolor Rohwer (SWUS) BSM 1 Aug. Pterocheilus hurdi Bohart (CA) BSM 1 June. Pterocheilus hirsutipennis Bohart (SWUS) SM 1 May. Pterocheilus laticeps Cresson (WUS) BSM 1 Jime. Pterocheilus tricoloratus Bohart (SWUS) BSM 1 June. Leptochilus species (?) BSM 1 Aug. Maricopodynerus maricoporum (Viereck) (SWUS) SM 4 April July. Stenodynerus percampanulatus (Viereck) (WUS) SM 6 June July Aug. Sept., BSM 8 Aug. Sept. Parancistrocerus toltecus (Saussure) (WUS) BSM 95 June July Aug. Sept. Euodynerus annulatus sulphureus (Saussure) (WUS) SM 1 June. Ancistrocerus acatskill halophila Viereck (WUS) SM 2 Oct. Pompilidae Pepsis pallidolimbata pallidolimbata Lucas (WUS) SM 2 Sept., BSM 1 Aug. Hemipepsis ustulata ochroptera Stal. (SWUS) SM 1 June. Aporus hirsutus (Banks) (WUS) BSM 2 June Aug. Evagetes padrinus padrinus (Viereck) (WNA) BSM 9 June Aug. Sept. Agenioideus biedermani (Banks) (SWUS) SM 1 Aug. Episyron oregon Evans (WNA) SM 1 May, BSM 1 Aug. Anoplius relativus (Fox) (NA) SM 5 June, BSM 10 June July Aug. Sept. Oct. Anoplius tenebrosus (Cresson) (NA) SM 1 Sept. Pompilus orophilus Evans (NA) SM 3 May, BSM 3 Aug. Pompilus phoenix Evans (WUS) BSM 3 June. Aporinellus borregoensis Evans (SWUS) SM 1 June. Aporinellus fasciatus (Smith) (NA) SM 1 June, BSM 1 June. Aporinellus medianus Banks (NA) SM 12 June July Aug. Sept. Aporinellus taeniatus taeniatus (Kohl) (NA) Sm 1 Sept. Aporinellus yucatanensis (Cameron) (NA) SM 1 Sept., BSM 1 June. Sphecidae Prionyx canadensis (Provancher) (NA) SM 1 June, BSM 6 June July Aug. 406 Great Basin Naturalist Vol. 43, No. 3 Prionyx subatratus Bohart (WUS) SM 1 Aug. Podahnia communis (Cresson) (WUS) SM 12 June July, BSM 6 July. Ammophila aberti Haldeman (WUS) SM 2 May June. Ammophila polita Cresson (WUS) BSM 4 June. Ammophila pruinosa Cresson (WUS) SM 2 May Sept. Ammophila wrightii (Cresson) (WUS) BSM 1 June. Mimesa coquilletti (Rohwer) (CA NV) SM 13 May Sept., BSM 2 June. Astata bechteli Parker (SWUS) SM 3 June July Aug. Astata occidentalis Cresson (WUS) BSM 9 June July Aug. Larropsis waslioensis Bohart and Bohart (NV) SM 3 May June, BSM 11 June July Aug. Ancistromma granulosa (Bohart and Bohart) (WUS) SM 1 Aug., BSM 1 Aug. Tachytes ermineus Banks (SWUS) SM 2 June. Tachytes nevadensis Bohart (WUS) SM 3 Aug. Sept. Tachytes new species SM 7 June. Tachysphex apicalis fusiis Fox (NA) BSM 3 June. Tachysphex ashmeadii Fox (WUS) SM 7 Aug. Tachysphex spinosus Fox (WUS) SM 1 June. Tachysphex species (?) BSM 14 June. Plenoculus boregensis Williams (SoCA) SM 1 June. Pisonopsis species (?) SM 3 May June. Miscophus species (?) SM 2 May June. Oxybelus abdominalis Baker (WUS) SM 10 Aug. Sept., BSM 6 June. Oxybelus pitanta Pate (SWUS) SM 12 Aug. Sept., BSM 2 June. Crabro opalenscens Bohart (WUS) SM 2 May. Bicyrtes ventralis (Say) (NA) SM 3 Aug. Bicyrtes capnopteris (Handlirsch) (NA) SM 3 Aug. Microbembex argyropleura Bohart (SWUS) SM 39 June July Aug. Sept., BSM 15 July Aug. Microbembex californica Bohart (SWUS) SM 1 June, BSM 1 July. Bembix rugosa Parker (AZ) SM 18 Aug. Bembix stenobdoma Parker (AZ) SM 1 July. Bembix occidentalis Fox (SWUS) SM 2 June. Bembix americana comata Parker (SWUS) SM 2 Aug. Sept. Stictiella corniculata Mickel (WUS) SM 3 Sept. Stictiella nubilosa Gillaspy (SoCA) SM 2 June July. Stictiella speciosa (Cresson) (WNA) SM 1 Sept. Glenostictia argentata (Fox) (SoCA) SM 12 Aug. Glenostictia megacera (Parker) (WUS) SM 1 Aug. Glenostictia tenuicornis (Fox) (SWUS) SM 2 Aug. Philanthus crotoniphilus Viereck and Cock- erell (WUS) SM 19 Aug. Sept. Philanthus multimaculatus Cameron (WNA) SM 13 Aug. Sept. Philanthus pacificus pacificus Cresson (NA) SM 11 July Aug. Sept., BSM 1 June. Philanthus pulcher Dall Torre (SWUS) SM 1 May. Philanthus ventralis (Mickel) (PCS) SM 17 Sept. Philanthus zebratus Cresson (WUS) SM 1 Sept. Clypeadon evansi Bohart (SWUS) SM 4 July Aug. Clypeadon laticinctus (Cresson) (WUS) SM 8 Aug. Clypeadon utahensis (Baker) (SWUS) SM 7 Aug. Sept. Clypeadon species (?) SM 1 July. Cerceris bridwelli ScuUen (SoCA AZ) SM 8 Aug. Cerceris californica Cresson (WUS) SM 5 July Aug. Cerceris conifrons Mickel (WNA) SM 4 Aug. Sept. Cerceris crotonella Viereck and Cockerell (WUS) SM 5 June July Aug., BSM 1 June. Cerceris echo echo Mickel (WUS) SM 3 July Aug. Sept. Cerceris species (?) SM 1 Aug. Eucerceris arenaria Scullen (SWUS) SM 9 Aug. Sept. Eucerceris nevadensis (Dalla Torre) (WUS) SM 74 July Aug. Sept. Colletidae Colletes rnandibularis Smith (EUS) SM 3 Sept., Chrysothamnus viscidiflorus. July 1983 Rust et al.: Nevada Hymenoptera 407 CoUetes slevini Cockerell (WUS) SM 4 June July Sept., Chvi, Psorothamnus polijdeniiis. CoUetes stepJiani Timberlake (SoCA) SM 1 June. CoUetes species (?) SM 30 June Aug. Sept., BSM 4 June Sept., Chvi, Eriogonum keameyi, Pspo, Tetradymnia coniosa. Andrenidae Andrena (Diandrena) makwothricidis Thorp (SoCA) BSM 2 May, Malacothrix sonclioides. Andrena (Onagrandrena) chyUsmiae Linsley and MacSwain (ECA) SM 15 April, BSM 2 May. Andrena (Onagrandrena) Unsleyi Timberlake (SWUS) SM 5 April. Andrena (Thysandrena) vierecki Cockerell (GB) BSM 2 June, Mentzelia albicauUs. Nornadopsis (Nomadopsis) pueUae (Cockerell) (WUS) SM 7 May June, SM 32 May June, Maso. Nornadopsis (Micronomadopsis) phaceliae Timberlake (ECA) SM 2 May, BSM 1 June, PhaceUa sp. Nornadopsis new species (?) BSM 35 June July, Psorotliamnus kingii. Perdita (Cockerellia) utahensis Cockerell (SWUS) SM 16 July Aug., HeUanthus deserticola. Perdita (Perdita) lepidosparti Timberlake (GB) SM 11 July Aug. Sept., Cleome sparsifolia. Perdita (Perdita) hirticeps Timberlake (SWUS) SM 16 June July Aug., Chvi, BSM 5 July. Perdita (Perdita) phymatae Cockerell (SWUS) SM 2 Sept., Chvi. Perdita (Procockerellia) albonotata Tim- berlake (SoCA) SM 1 July. Perdita species 1 (?) SM 84 Aug. Sept., Chvi. Perdita species 2 (?) SM 1 Aug., Pspo. Perdita species 3 (?) SM 51 July, Tiguilia nuttaUii. Perdita species 4 (?) SM 15 July Aug., Pspo. Halictidae Nomia (Acunomia) howardi (SWUS) SM 2 Aug., Erke. Crawford Agapostemon femoratiis Crawford (WNA) BSM 1 June, Clsp, Sphaeralcea atnbigua. Agapostemon meUiventris Cresson (WUS) SM 73 June July Aug., BSM 2 Aug. Sept., Atriplex sp, Amaranthus sp, Camissonia clavaefonnis, Chvi, Clsp, Hede, Erke, Psoralea lanceolata Pspo. Lasioglosswn sisymbrii (Cockerell) (WUS) SM 16 April June July Aug. Sept., As- tragalus lentiginosus, Chvi. Evylaeus aberrans (Crawford) (WUS) SM 9 May June July, BSM 2 May June. DiaUctus species 1 (?) BSM 5 April. DiaUctus species 2 (?) SM 78 April May June July Aug. Sept., BSM 37 July Aug. Sept., Chvi, Erke, Hede, Psla, Step- hanomeria exigiia, Tetradymia tetrameres. DiaUctus species 3 (?) SM 9 Aug. Sept., Abr- ionia turbinata, Asle, Chvi, Erke, Pen- stemon acuniinatus, PhaceUa sp. Sphecodes species (?) SM 1 Sept. Melittidae Hesperapis species (?) SM 22 May, PhaceUa sp, Oenothera deUoides. Megachilidae Anthidium rodecki Schwartz (CO NV) SM 15 May June July Aug., BSM 44 July Aug., Psla, Pski, Pspo. Dianthidium subparviim Swenk (WNA) SM 4, Chaenactis xantiana, Chvi. AnthidieUiim notatum robertsoni (Cockerell) (WNA) SM 2 Aug., Psla. SteUs species (?) BSM 1 June. Proteriades (HolpUtina) bidUfacies (Michener) (ECA) SM 3 April May, BSM 5 May June, (Asle) PhaceUa. Anthocopa (Eremosmia) robustula (Cockerell) (SWUS) SM 4 April May, BSM 5 June, PhaceUa sp. Anthocopa (Eremosmia) timberlakei (Cock- erell) (SoCA) SM 28 May June, BSM 15 May June, Asle, Gael, Meal, Peac, Pha- ceUa sp, Pski, Pspo. AshmeadieUa (Ashmeadiella) aridula Cock- erell (WUS) SM 5 Aug., BSM 3 Aug, Clsp, PhaceUa sp, Psla, Pski, Pspo. 408 Great Basin Naturalist Vol. 43, No. 3 Ashmeadiella (Ashmeadiella) bticconis den- ticulata (Cresson) (WUS) SM 1 June, Pspo. Osmia (Nothosmia) titusi Cockerell (SoCA) SM 2 April June, Asle, Phacelia sp, Pspo. Megachile (Derotropis) xerophila Cockerell (SoCA AZ) SM 6 May, BSM 3 June, Chxa, Maso. Coelioxys (Coelioxys) mitchelli Baker (SUS) SM 2 May. Anthophoridae Diadasia australis (Cresson) (WUS) BSM 12 June, Opiintia pidchella, Mentzelia al- bicaidis, Hede. Diadasia vallicola Timberlake (AZ CA) SM 16 May. Synhalonia albescens Timberlake (SWUS) SM 3 May. Synhalonia primaveris Timberlake (SWUS) SM 30 April May, BSM 4 May June, Asle, Meal, Peac, Phacelia. Synhalonia speciosa (Cresson) (WUS) SM 1 May. Melissodes (Eumelissodes) biniatris LaBerge (WNA) BSM 2 Sept. Melissodes (Eumelissodes) lutulenta LaBerge (WNA) SM 14 June, BSM 2 June July, Chaetadelpha wheeleri, Hede, Pspo, Spam, Teco. Melissodes {Eumelissodes) montana Cresson (WUS) SM 39 Sept, BSM 1 Sept, Chvi, Erke. Anthophora (Anthophora) affabilis Cresson (WUS) SM 31 April May, BSM 3 May, Asle, Cacl, Peac, Phacelia. Anthopfiora (Anthophora) porterae Cockerell (WUS) SM 39 April May, BSB 9 May June, Asle, Peac. Anthcrphora (Anthoplwra) urbana Cresson (WUS) SM 35 Aug. Sept., BSM 7 Aug. Chvi, Clsp, Erke, Stex. Anthophora (Micranthophora) hololeuca Cockerell (SWUS) SM 18 June July Aug, BSM 38 June July Aug, Pski, Pspo. Anthophora (Micranthophora) petrophila Cockerell (SWUS) SM 11 Aug. Sept., BSM 3 Aug, Chvi. Centris (Xerocentris) californica Timberlake (CA AZ) SM 34 Aug. Sept., Clsp. Ceratina (Zadontomerus) neomexicana Cock- erell (SWUS) SM 10 May June July Aug., BSM 3 June July, Erke, Hede, Maso, Pspo, Stex, Teco. Apidae Apis mellifera Linnaeus (world wide) SM 7 April, Asel. Acknowledgments We thank G. E. Bohart, R. M. Bohart, A. A. Grigarick, L. S. Kimsey, W. E. LaBerge, A. S. Menke, F. D. Parker, J. G. Rozen, R. R. Snelling, R. W. Thorp, M. S. Wasbauer, and G. C. and J. Wheeler for the identification of specimens used in this study. Dave Goi- coechea, BLM State of Nevada, made the study possible. Literature Cited Allred, D. M. 1973. Additional records of mutillid wasps from the Nevada Test Site. Great Basin Nat. 33:156-162. Bechtel, R. C, L. M. Hanks, and R. W. Rust. 1981. Orthopteroids of Sand Mountain and Blow Sand Mountains, Nevada. Entoniol. News 92:125-129. Beck, D. E., and D. M. Allred. 1968. Faunistic inven- tory—BYU ecological studies at the Nevada Test Site. Great Basin Nat. 28:1.32-141. Cole, A. C. 1966. Ants of the Nevada Test Site. BYU Sci. Bull., Biol. Ser. 7(3): 1-26. Ferguson, W. E. 1967. Male Sphaeropthalmine wasps of the Nevada Test Site. BYU Sci. Bull., Biol. Ser. 8(4): 1-26. Hanks, L. M., and R. W. Rust. 1983. Bee pollinators in a sand dune community. Submitted to Ecology. Krombein, K v., and p. D. Hurd. 1979. Catalog of Hymenoptera in America north of Mexico. Smithsonian Institution Press, Washington, D.C. Morrison, R. B., and J. C. Frye. 1965. Correlation of the middle and late quaternary successions of the Lake Lahontan, Lake Bonneville, Rocky Moun- tains (Wasatch Range), southern Great Plains, and eastern midwest areas. Nevada Bureau Mines 9:1-45. Pietruszka, R. D. 1980. Observations on seasonal varia- tion in desert arthropods in central Nevada. Great Basin Nat. 40:292-297. Wasbauer, M. S. 1973. The male Brachycistidine wasps of the Nevada Test Site (Hymenoptera: Tiph- iidae). Great Basin Nat. 33:109-112. Wheeler, G. C, and J. Wheeler. 1978. Mountain ants of Nevada. Great Basin Nat. .38:379-396. STATUS AND LIFE HISTORY NOTES ON THE NATIVE FISHES OF THE ALVORD BASIN, OREGON AND NEVADA Jack E. Williams' and Carl E. Bond- .\bstract.— Three fishes, two species of Gila, and an undescribed subspecies of cutthroat trout, are endemic to the Alvord Basin. Historically, the Alvord cutthroat trout, Salmo clarki ssp., inhabited the larger creeks of the basin but has been extirpated in pure form because of introgression with introduced rainbow trout, Salmo gairdneri. Gila boraxobitis is restricted to the thermal waters of Borax Lake and its outflows in the northern part of the basin. This species is endangered because of alteration of its fragile habitat. The Alvord chub, G. alvordensis, is recorded from 16 localities throughout the basin, including springs, creeks, and reservoirs. Although G. alvordensis as a species is not in jeopardy, many populations are small and could be easily eliminated by habitat destruction or by the in- troduction of exotic fishes. Competition with exotic guppies, Poecilia reticulata, has extirpated the Thousand Creek Spring population of Alvord chubs. Both species of Gila are opportunistic omnivores, consuming primarily chironomids, microcrustaceans, and dia- toms. Tlie Borax Lake chub also consumed large numbers of terrestrial insects, but specialized feeding on molluscs was noted in the West Spring population of Alvord chubs. Borax Lake chubs spawn throughout the year; however, most spawning occurs in early spring. Borax Lake chubs mature at a small size, occasionally less than 30 mm stan- dard length, and seldom live more than one year. Alvord chubs are typically much larger than the Borax Lake spe- cies and live at least into their fifth year. The Alvord Basin of southeastern Oregon and northwestern Nevada is an endorheic part of the Great Basin province. Aquatic habitats are sparse and consist primarily of Trout Creek in Oregon, the Virgin-Thousand Creek system in Nevada, as well as several small streams and springs (Fig. 1). During the late Pleistocene, a lake of over 1,200 km^ covered much of the valley floor (Snyder et al. 1964). As pluvial waters dried, fishes were restricted to remaining permanent springs and creeks. Three native fishes are endemic to the Alvord Basin. Chubs, genus Gila, oc- cupy many of the isolated waters in the Al- vord Basin and have diverged into two spe- cies. The Alvord chub, Gila alvordensis, is the most common fish in the basin and occurs in a variety of springs and creeks. The Borax Lake chub, G. boraxobitis, is restricted to Bo- rax Lake and its outflows in Oregon. The Al- vord cutthroat trout, Salmo clarki ssp., is the third fish native to the basin. Historic habitat for the Alvord cutthroat trout consisted of the larger streams in the basin, such as Trout and Virgin creeks. Another undescribed sub- species of cutthroat trout occurs in Willow and Whitehorse creeks just east of the Trout Creek Mountains in a separate basin. Al- though exhibiting affinities for the Alvord cutthroat trout, the subspecies foimd in Wil- low and Whitehorse creeks will not be treated further in this report. Our knowledge of the native fishes of the Alvord Basin is limited. The monograph of Great Basin fishes by Hubbs and Miller (1948) provided the first detailed account of the Great Basin ichthyofauna and included a brief discussion of the native Alvord Basin fishes and their isolation. However, all the Alvord Basin fishes remained undescribed un- til 1972, when Hubbs and Miller (1972) diag- nosed the Trout Creek population of Gila as G. alvordensis. Our studies have resulted in the description of Gila boraxobitis (Williams and Bond 1980) and a further description of G. alvordensis with a taxonomic analysis of seven disjunct populations of the species (Williams 1980, Williams and Bond 1980). Characters of the Alvord cutthroat trout have been provided by Behnke (1979), but the sub- species remains undescribed. The only pub- lished life history information on Alvord 'Endangered Species Office, U.S. Fish and Wildlife Service, 1230 "N" Street, 14th Floor, Sacramento, California 95814. 'Department of Fisheries and Wildlife, Oregon State University, Corvallis, Oregon 97331. 409 410 119 mJk Great Basin Naturalist o Vol. 43, No. 3 ^N « 4 / ^Dufurrena Area ^og Hot Thou Cr . I T.- . Res /^ Borax Lake. ,^ :^J 50 I 1 I 1 km Fig. 1. Map of major aquatic habitats in the Alvord Basin, Oregon and Nevada. Basin Gila consists of a study of feeding ecol- of the native fishes of the Alvord Basin and ogy by Williams and Williams (1980). This presents additional information on their life paper documents the distribution and status history. July 1983 Williams, Bond: Native Fishes 411 Materials and Methods The distribution and status of fishes was determined by field surveys, museum records, and testimony of local residents. Many of the habitats in the Oregon part of the basin were known prior to this work. On the other hand, the Nevada part of the basin had received little attention by ichthyologists, and there- fore most of our survey efforts were focused in the southern one-half of the basin. Surveys were conducted from 12 June 1978 to 26 Au- gust 1979 and from 13 to 15 April 1982. Habitats were sampled with 3 m seines (9.5 mm mesh), dip nets, fish traps, backpack electroshocker, and 15 m gill nets (51 mm mesh). Fishes utilized in this study are depos- ited at Oregon State University (OS), The University of Michigan Museum of Zoology (UMMZ), Tulane University (TU), and the University of Nevada, Las Vegas (UNLV). Information concerning reproduction, lon- gevity, and adult sex ratio was determined for Borax Lake chubs collected monthly from March 1978 to January 1979. Fish were col- lected from the southwest one-quarter of Bo- rax Lake. Specimens were preserved in 10 percent formalin and transferred to 45 per- cent isopropanol after one week. Standard length (SL) of specimens was measured to the nearest 0.1 mm with dial calipers. After blotting fish dry on paper towels, wet weight was measured to the nearest 0.01 g. A gona- dosomatic index was calculated by weighing the left ovary or testis to the nearest 0.001 g, multiplying by two, thereby accounting for the right gonad, and dividing by fish weight. Three classes of ova were identified: class 1— mature ova, yellow color, 0.7 to 1.2 mm di- ameter; class II— immature ova, opaque white color, 0.4 to 0.6 mm diameter; and class III— immature ova, transparent, 0.1 to 0.3 mm diameter. The number of ova was enumerated in females that possessed only class I and/ or class II ova. By this method, accurate counts could be obtained and these numbers are probably more indicative of the actual number of eggs deposited during spawning. In females shorter than 35 mm SL, all ova were counted in both ovaries. In fe- males 35 to 50 mm SL, all ova were counted in the left ovary, then multiplied by two to derive the total number of ova. Ova were enumerated in larger females by multiple subsamples of the ovary. Age was determined by examining annuli of scales taken from the left side of the body above the lateral line. Monthly collections were grouped into sea- sons as follows: spring (March-N4ay), summer (June-August), fall (September-November), and winter (December-February). Population Accounts Alvord Cutthroat Trout, Salmo clarki ssp. The Alvord cutthroat trout, Salmo clarki ssp., is now extinct in pure form. This native trout was known from Virgin Creek in Ne- vada and Trout Creek in Oregon, but prob- ably existed in several of the larger Alvord Basin creeks during recent times (Hubbs and Miller 1948). Trout Creek (Harney County, Oregon).— The Alvord cutthroat trout occurred in the headwater canyon area of Trout Creek, where it flows through the Trout Creek Mountains. Introgression of introduced rain- bow trout, Salmo gairdneri, with the native cutthroat was already noticeable in 1934 col- lections of fish made in Trout Creek by Carl L. Hubbs, although trout from more head- water localities were quite similar in appear- ance to pure Alvord cutthroat trout (Behnke 1979). Cutthroat trout pigmentation was evi- dent in many specimens collected by one of us (CEB) in 1953 and 1957, but none had basibranchial teeth. Trout collected from* Trout Creek in 1972 (Behnke 1979) and 1978 to 1980 (our collections) exhibited only rain- bow trout characteristics and we conclude that the native trout has been extirpated from this creek. Collections made from Cot- tonwood Creek and other streams draining the Trout Creek Mountains yielded only rain- bow trout. Virgin Creek (Humboldt County, Ne- vada).—Thirty small (<15 cm) Alvord cut- throat trout were collected from Virgin Creek in or near Virgin Creek Gorge by Carl L. Hubbs in 1934 (UMMZ 130532). Behnke (1979) considers these specimens to be pure native cutthroat trout and provides their de- scription as follows: body with fewer than 50 relatively large round spots, spots concen- trated posteriorly and above lateral line; few 412 Great Basin Naturalist Vol. 43, No. 3 spots on caudal fin. Gill rakers 20 to 26. Lat- eral series scales 122 to 152. Scales above lat- eral line 33 to 37. Pelvic fin rays 8 or 9. Branchiostegal rays 8 or 9. Vertebrae 59 to 63. Trout collected in 1971 (OS 3832, OS 3834) from approximately the same region of Virgin Creek exhibited typical rainbow trout features (Behnke 1979). During 1978 surveys, we found only rainbow trout or introgressed trout in Virgin Creek at and upstream of the north end of Virgin Creek Gorge. No fish were found in Virgin Creek Gorge at the nexus of Hell and Virgin creeks. The up- stream sections of Virgin Creek, in Virgin Creek Gorge near Alkali Ranch, also are fish- less. Although we received reports of large trout from beaver ponds in Virgin Creek Gorge downstream of Wilson Ranch, none could be secured for examination. The large number of introduced rainbow trout in Vir- gin Creek Gorge would seem to preclude the survival of pure Alvord cutthroat trout in Virgin Creek. An extensive survey of Hell Creek, the only permanently flowing tribu- tary of upper Virgin Creek, revealed only a single fish, which was typically rainbow in character. This individual was apparently able to ascend the falls separating Hell and Virgin creeks during a flood. The negative survey of Hell Creek in 1978 and 1979 causes us to consider the Alvord cutthroat trout to be extinct. Borax Lake Chub, Gila boraxobius The Borax Lake chub, Gila boraxobius Williams and Bond, is restricted to the ther- mal waters of Borax Lake and its outflows. Gila boraxobius was described in 1980 and is considered a dwarf relative of G. alvordensis (Williams and Bond 1980). Borax Lake (T37S, R33E, Sec 14; Harney County, Oregon).— Borax Lake is a relatively shallow 4.1 ha natural lake that receives wa- ter from several thermal springs. These springs issue into the bottom of the southwest portion of the lake at approximately 35 to 40 C. Lake temperature is typically 29 to 32 C but can vary from 17 to 35 C depending on season, weather, and distance from the spring sources. The water is clear. Substrates range from rocky outcroppings in the southeast portion of the lake to gravels in the north and soft, easily roiled silt in the remainder of the lake. The lake shoreline consists of salt crusts, which have been deposited by the lake waters. These salt deposits have built up over hundreds or, more probably, thousands of years until the lake is now 10 m higher in elevation than the surrounding land. Histori- cally, the lake waters overflowed along the south and southwest shoreline, creating a marsh. These outflows also provided water for Lower Borax Lake, a reservoir southwest of Borax Lake. Adjacent to Borax Lake are two small pools, one about 25 m southwest of Borax Lake and an artificial pool about 75 m west of the lake. Borax Lake chubs occur throughout Borax Lake except in the hot spring inflows. Obser- vations at Borax Lake indicated that Borax Lake chubs avoided water with a temper- ature above 34 C. These observations are supported by unsuccessful attempts to chase the chubs into hot spring inflow areas. In aquaria. Borax Lake chubs lost equilibrium when water temperature was raised to 34.5 C, indicating a critical thermal maximum near this temperature. Borax Lake chubs also occurred in small numbers in the two small pools near Borax Lake. The pools are appar- ently formed from Borax Lake overflow waters. The Borax Lake chub is a dwarf species that typically reaches maturity at 30 to 35 mm SL. Males as small as 28.6 mm SL are highly tuberculate and females as small as 31.8 mm SL have been found with mature eggs. Typical adult size is 33 to 45 mm SL. The largest male collected from Borax Lake was 50.6 mm SL, whereas two exceptionally large females, 90.4 and 93.0 mm SL, have been collected from Borax Lake. Most spawning probably occurs in early spring, although some spawning can occur year around. The gonadosomatic index was highest in females during March and April, with mature, class I ova present during March, April, and January (Table 1). Ovaries were usually poorly developed during May through August. In males, the gonadosomatic index was highest in April and September, when testes averaged 0.97 and 1.11 percent body weight, respectively. A search of mu- seum specimens disclosed large females with mature ova collected on 17 June (OS 4137) July 1983 Williams, Bond: Native Fishes 413 and 11 September (OS 4106). Thus, spawning may occur at any time of the year. However, a major spawning in early spring is supported by observation of numerous larval chubs dur- ing April, May, and early June. Young fish, eight to 15 mm SL, typically inhabit the shal- low cove areas along the west and south mar- gins of Borax Lake. Water is cool, only a few cm deep, and vegetation is common in the coves. Ova number increases dramatically with fish length. The number of ova was deter- mined in eight females 32.7 to 93.0 mm SL, that contained only class I and/ or class II ova. The smallest females examined, 32.7 and 34.5 mm SL, contained 75 and 82 ova, re- spectively. Larger females, 39.0, 39.3, 44.6, and 49.4 mm SL contained 252, 246, 380, and 362 ova, respectively. The exceptionally large females, 90.4 and 93.0 mm SL con- tained 2,143 and 6,924 ova, respectively. Al- though females larger than 60 mm SL are very rare in Borax Lake, their contribution to recruitment may be substantial. Most Borax Lake chubs live one year, with few age I and II fish present. Annuli devel- opment, although difficult to discern, in- dicated that the 90.4 and 93.0 mm SL fe- males were probably age III. This appears to be the maximum age achieved by chubs in Borax Lake. A length frequency analysis of 113 individuals collected 5 August 1977 ap- pears in Figure 2. Because most spawning oc- curs in spring, with young of approximately 10 mm SL prominent in May and June, most fish in the August collection are probably young-of-the-year. Some age I fish, 33 mm to 51 mm SL are present, whereas the two larger individuals are probably age II (Fig. 2). Most older fish are females. Adults (>33 mm SL) typically comprised less than 25 percent of specimens collected during March, April, and May. The percentage of adults in month- ly collections then increased until reaching a peak of 82 percent during November. Based on monthly collections made throughout the year, the sex ratio of 190 indi- viduals greater than or equal to 30 mm SL was 1.0(5 :1.3 ? . Seasonally, the sex ratio was as follows: spring (n = 23) 1.0(5 :1.3 ? , sum- mer (n = 67) 1.0(5:1.9$, fall (n = 50) 1.0 5:11?, and winter (n = 50) 1 <5 :1 ? . The larger number of females may be in- dicative of higher survivorship following spawning. The feeding ecology of Borax Lake chubs has been reported by Williams and Williams (1980). The relative importance of foods var- ied seasonally, but diatoms, microcrustaceans, and chironomid larvae were often the pri- mary foods consumed. Terrestrial insects were important foods during summer and fall (Williams and Williams 1980). Lower Borax Lake (T37S, R33E, Sec 15; Harney County, Oregon).— Lower Borax Lake is a reservoir that receives water from the southwest outflow creek of Borax Lake. Water levels of Lower Borax Lake fluctuated seasonally, often holding little water during summer. Unfortunately, alteration to Borax Lake during 1979 diverted water away from the reservoir. Prior to the diversions, the res- ervoir occasionally harbored Borax Lake chubs that entered from the southwest out- flow creek. It is doubtful that Borax Lake chubs ever spawned in Lower Borax Lake* and the population was probably dependent on an influx of fish from Borax Lake via the outflow creek. Table 1. Monthly reproductive characteristics of female Borax Lake chubs longer than 30 mm SL. Classes Gonadosomatic index of ova Month n xSL Range X SD present March 3 38.0 0.91-5.01 2.89 2.05 LII,III April 5 33.1 0.78-10.56 3.45 4.13 i,n,ni May 4 34.8 0.45-0.55 0.51 0.05 III only June 7 35.9 0.53-2.38 1.01 0.66 II,III July 8 44.0 0.50-1.54 1.06 0.37 III only August 8 37.4 0.16-1.60 0.95 0.41 IIJII September 5 38.3 0.70-4.56 2.48 1.61 II,III November 5 39.3 1.28-2.13 1.61 0.34 IIJII December 8 42.5 1.20-4.45 2.02 1.03 II,III January 7 41.4 0.99-2.27 1.59 0.48 I,II,III 414 Great Basin Naturalist Vol. 43, No. 3 >« u c 0) 3 O" 0> 20 15 10 5- 15 21 27 33 39 45 51 n I n 57 63 6< 69 Standard Length in mm Fig. 2. Length frequency of 113 Borax Lake chubs, Gila boraxobius, collected 5 August 1977 from Borax Lake, Oregon. Alvord Chub, Gila alvordensis The Alvord chub, Gila alvordensis Hubbs and Miller, was found in 16 habitats located throughout much of the basin in Oregon and Nevada. The species was collected from a va- riety of habitats, including springs, creeks, and reservoirs. The species has been collected from Jimiper Lake, Oregon (Bond 1974), al- though its presence appears attributable to an introduction because the lake dries during drought years. Serrano Pond (T36S, R33E, Sec 1; Har- ney County, Oregon).— Serrano Pond is a 0.1 ha reservoir that receives water from a cool- water spring approximately 60 m distant. Water flows from the spring at approx- imately 17 C and water temperature in the pond is typically 16 to 21 C during the sum- mer. The substrate of the relatively shallow pond is primarily silt. The water is somewhat turbid and aquatic vegetation is abundant. Recent alteration to this area has resulted in a diversion canal draining part of the flow away from the pond. Alvord chubs are absent from the spring, but are abundant in remain- ing waters of the pond and in the diversion creek. More than 100 fish can be easily col- lected from the pond in a single seine haul during the summer. Adult males are typically about 50 mm SL and adult females average approximately 65 mm SL, but females great- er than 80 mm SL are occasionally collected. Alvord chubs from Serrano Pond are high- ly opportunistic feeders. Bottom in- vertebrates are grazed extensively, as are midwater crustaceans and diatoms. Very few foods are consumed from the water's surface. Chironomid larvae, diatoms and cladocerans were the principal foods during summer, and ostracods, harpacticoid copepods, and chi- ronomid pupae were of secondary impor- tance (Williams and Williams 1980). Eighty- nine percent of the intestines examined by Williams and Williams (1980) contained one food that accounted for more than 50 per- cent of intestinal volume. Thirty-nine per- cent of the intestines contained one food, chi- ronomid larvae, diatoms, or cladocerans, almost exclusively. Trout Creek and Alvord Lake (Harney County, Oregon).— Trout Creek is the largest stream in the Alvord Basin and discharges an average of 15 cfs, as measured in the canyon area 8 km east of Trout Creek Ranch (Libbey 1960). The creek heads in Trout Creek Mountains just north of the Nevada border. In the headwaters. Trout Creek flows July 1983 Williams, Bond: Native Fishes 415 through canyon areas where its waters are clear and fast-flowing. Water temperature in the canyon is cool during summer, near 15 C, and colder during winter months. As Trout Creek leaves the canyon and enters the val- ley floor, flows decrease and water temper- atures increase. Naturally lower summer flows and irrigation diversions often reduce the lower portions of Trout Creek to an in- termittent stream during late summer. Turbi- dity is often high (visibility 1 cm) in lower reaches during summer. Substrate type changes from mostly gravel in upstream areas to silt in downstream sections. Trout Creek eventually empties into Alvord Lake, a rem- nant of the large lake that covered the valley during pluvial times. Alvord Lake varies greatly in size and occasionally dries com- pletely during drought years. Alvord chubs are common, although not abundant, in upstream canyon areas, and abundant in downstream sections. Introduced rainbow trout, Salrno gairdneri, also occur in upstream regions of Trout Creek. This is the only habitat where Gila occur sympatrically with another fish in the Alvord Basin. Alvord chubs in the canyon area are large, the long- est measuring 122 mm SL. Three specimens 113 to 122 mm SL are all age class IV. Downstream areas also produce large Alvord chubs, although maximum length appears somewhat less. Color differences between Al- vord chubs in the canyon and downstream areas are striking. Alvord chubs from the can- yon are very dark, nearly black, dorsally, with golden sides possessing some black speckles, and a silver belly. In sections of Trout Creek in the valley floor, Alvord chubs are lighter in color, exhibiting a light green color on the dorsal part of the head and body, silver sides without speckles, and a white belly. All fins of Gila from Trout Creek are translucent red or orange in color except at the tips, which are white. Pueblo Slough (T40S, R35E and T41S, R35E; Harney County, Oregon).— Pueblo ( = Denio) Slough is a wetland area approx- imately 13 km long, extending from Tum Turn Lake in the north to just north of the Nevada border in the south. Various marsh, spring, and creek areas south of Tum Tum Lake provide most of the habitat in the slough. Water in Pueblo Slough is provided by Van Horn and Colony creeks, which drain the Pueblo Mountains, as well as at least 25 springs in the slough itself. These springs are mostly cool and shallow. Red Point School (T40S, R35E, Sec 14) is located in approx- imately the center of the slough. In August, Alvord chubs were collected from a shallow, clear pool at Red Point School where water and air temperatures were 15 and 17 C, re- spectively. Alvord chubs were abundant in the pool. Despite searches for chubs in streams draining the Pueblo Mountains, none could be found— although small rainbow trout were collected in lower Van Horn Creek. Bog Hot Reservoir (T46N, R28E, Sec 17; Humboldt County, Nevada).— Bog Hot Res- ervoir is a relatively small impoundment fed by thermal waters flowing from Bog Hot Springs. Water issues from Bog Hot Springs at approximately 44 C and flows for 1.2 km before entering Bog Hot Reservoir. Water from Bog Hot Springs enters the reservoir at about 30 C. Typical water temperatures in the reservoir are 20 to 21 C during early summer. The waters of Bog Hot Reservoir are slightly turbid (visibility 31 cm), and the substrate is mostly silt with some gravel. Al- vord chubs are abundant in Bog Hot Reser- voir but are absent in Bog Hot Springs and in practically all the inflow creek between the springs and the reservoir. No Alvord chubs were found upstream of the 31.1 C boundary, where water from the inflow creek enters Bog Hot Reservoir. Alvord chubs collected from Bog Hot Res- ervoir on 13 June 1978 were in spawning condition. On that date, water and air tem- peratures were 20.4 and 20.7 C, respectively. The fish were collected from open, slightly turbid water, 30 to 40 cm deep. Young-of- the-year fish 10 to 15 mm SL were abundant in the reservoir during June 1978, indicating a spawning season from at least April imtil July. Most young were observed in the shal- low northwest end of the reservoir near the warm inflow creek. Bog Hot Creek (T46N, R28E; Humboldt County, Nevada).— Bog Hot Creek flows for approximately 5.2 km below Bog Hot Reser- voir before entering Thousand Creek. Parts of Bog Hot Creek below the reservoir have 416 Great Basin Naturalist Vol. 43, No. 3 been diverted or otherwise modified by agri- cultural practices. Alvord chubs were not collected in Bog Hot Creek except in the lower reaches near Thousand Creek. One poeciliid, probably a guppy, was observed in the highly modified section of the creek about halfway between the reservoir and Thousand Creek. In the downstream part of Bog Hot Creek, Alvord chubs were rare to common during an April 1982 svirvey and oc- curred primarily in pools 20 to 46 cm in depth. Water and air temperatures on 15 April 1982 were 2.3 C and 1.5 C, respective- ly. The water was clear, but appeared brown and quite acidic. Unnamed spring (T46N, R26E, Sec 31; Humboldt Covmty, Nevada).— This spring, measuring 2.4 m wide and 5 m long at its maximiun extent, is by far the smallest habi- tat supporting fish in the Alvord Basin. Max- imum depth is 77 cm. The water is clear and the substrate is an easily roiled silt. Water temperature was 11.4 C (air 7.5 C) during April and 18.2 C (air 20.4 C) during June. The unnamed spring is well isolated from nearby Thousand Creek by approximately 100 m of greasewood flat. The closest waters of Thousand Creek do not support Alvord chubs. The nearest population occurs approx- imately 1 km away in Dufurrena Pond 19. A very small population of Alvord chubs, estimated at slightly less than 100 individuals, inhabits the spring. Many seine hauls, each encompassing the entire spring, yielded 52 fish during an April 1982 survey. The Alvord chubs ranged in size from 32 mm SL to, con- sidering the small size of the habitat, an ama- zingly large 87 mm SL individual. No juve- niles were seen during April, but young-of- the-year were observed on 14 June, when wa- ter temperature had risen to 18.2 C. Juveniles occurred among rushes, Juncus sp., in water only a few cm deep. Adults were occasionally observed darting across the open center of the spring, but spent most of the daylight hours under a narrow band of floating algal mats along the periphery of the spring. Thousand Creek Spring (T46N, R26E, Sec 31; Humboldt County, Nevada).— Thou- sand Creek Spring forms the headwaters of Thousand Creek. The clear spring waters achieve a maximum depth of approximately 31 cm. Substrates are mostly fine gravels with some silt. Water and air temperatures during June were 27.1 and 18.2 C, respec- tively. Currently, the spring is inhabited by swarms of exotic guppies, Poecilia reticulata. Guppies have become established here and in the spring pool at nearby Dufurrena Camp- ground. Competition from introduced gup- pies probably extirpated Alvord chubs from Thousand Creek Spring. Alvord chubs have not been recorded from Thousand Creek Spring, but their historic presence is in- dicated by the occurrence of Alvord chubs in downstream areas of Thousand Creek and in a nearby spring. Except for the presence of guppies. Thousand Creek Spring appears to provide a suitable habitat for Alvord chubs. Thousand Creek and Continental Lake (Humboldt County, Nevada).— Thousand Creek heads at Thousand Creek Spring, flows through the Dufurrena area, where it re- ceives Virgin Creek, and then enters Thou- sand Creek Gorge. Below the gorge. Thou- sand Creek becomes braided and receives water from Bog Hot Creek before eventually emptying into Continental Lake. In the Du- furrena area, Thousand Creek is dammed at several locations to create reservoirs. Thou- sand Creek is usually turbid (visibility 7 cm), shallow, and about 1 to 2 m wide. The sub- strate is mostly silt. In Thousand Creek Gorge, the creek is surprisingly deep (>300 cm) and cool, near 15 C during late summer. Below the gorge. Thousand Creek is often in- termittent during summer, when water tem- peratures can reach 27 C. Typical summer and fall water temperature is 16 to 18 C. Al- vord chubs are abundant in Thousand Creek except in some upstream areas where guppies have been introduced or habitat has been al- tered by reservoir construction. Guppies are abundant in Thousand Creek Spring and oc- cur sporadically in Thousand Creek between Thousand Creek Spring and Dufurrena Pond 19. None were found downstream of Dufur- rena Pond 19. Thousand Creek contains many large Alvord chubs; the largest mea- sured 104.9 mm SL. During summer, Alvord chubs concentrate in deep pools in down- stream areas of Thousand Creek. Continental Lake usually dries completely during sum- mer, but harbors Alvord chubs during winter months. July 1983 Williams, Bond: Native Fishes 417 The sex ratio of 23 adults greater than 35 mm SL was 1.1 5 :1.0 ? . Food habits of Al- vord chubs collected during Jime from Thou- sand Creek were reported by Williams and Williams (1980). They found 10 foods in the intestines, of which chironomid larvae, cla- docerans, copepods, and ostracods were of greatest importance. Chironomid larvae oc- curred in all intestines examined and ac- counted for approximately 26 percent mean volume (Williams and Williams 1980). Mi- crocrustaceans comprised almost 45 percent mean volimie of intestines, whereas diatoms accoimted for only 5 percent mean volume. No terrestrial insects were observed in the in- testines. Alvord chubs in Thousand Creek ap- pear to feed primarily on bottom in- vertebrates and midwater crustaceans, avoiding surface foods. DuFURRENA PoND 19 (T46N, R26E, Sec 32; Humboldt County, Nevada).— Dufurrena Pond 19, approximately 1 km downstream of Thousand Creek Spring, is the first reservoir on Thousand Creek. The reservoir is shallow (typically < 50 cm) and moderately turbid (visibility 14 cm). Alvord chubs occur in the reservoir but are not abundant. Young-of-the- year (<20 mm SL) were abundant during June in the inflow diversion creek feeding the reservoir. At this time, the inflow creek was shallow (< 8 cm) and clear, with water and air temperatures of 14.5 and 13.0 C, respectively. Dufurrena Pond 22 (T45N, R26E, Sec 2; Humboldt County, Nevada).— Dufurrena Pond 22 is a reservoir fed by waters of Thou- sand and Virgin creeks. The water is very turbid (visibility 2 cm) and shallow (typically 15 to 20 cm deep). Water level fluctuates greatly with season. During late summer, the reservoir is reduced to a small pool. Water and air temperatures during June were 17.6 and 13.2 C, respectively. Alvord chubs are abundant and achieve a large size in the reservoir. Virgin Creek (Humboldt County, Ne- vada).— Virgin Creek heads near the southern extent of the Alvord Basin, flows north through the more than 300-m-deep Virgin Creek Gorge, and then flows east until reach- ing Thousand Creek. Springs scattered along most of the length of Virgin Creek provide its flow. Alvord chubs are abundant in Virgin Creek from the north end of Virgin Creek Gorge to the nexus of Virgin and Thousand creeks. Alvord chubs are absent in Virgin Creek Gorge, where introduced rainbow trout are common. Virgin Creek below the gorge is cool, relatively shallow, and moder- ately turbid (visibility 10 cm). Below the gorge, water temperature is typically 15 to 18 C during summer months and maximum depth is usually less than 75 cm. The sub- strate is mostly silt. Warm Spring (T45N, R25E; Humboldt County, Nevada).— Warm Spring and its out- flow are tributary to Virgin Creek approx- imately 2.5 km downstream from Virgin Creek Gorge. The outflow creek is a small, clear-water stream with an easily roiled silt bottom. Summer water and air temperatures of the creek just below the spring were 26.0 and 26.4 C, respectively. Alvord chubs are abundant in the outflow creek. The spring it- self was not sampled, but Carl Hubbs col- lected Alvord chubs from the spring in 1934 ("Italian Camp Spring," UMMZ 130533). The steep gradient and low flow of the Warm Spring crenon as it approaches Virgin Creek inhibits mixing between the Virgin Creek and Warm Spring populations of Al- vord chubs. Although Warm Spring flows through Virgin Valley Ranch, the spring and outflow creek have been only slightly altered and no exotic fish were present during a 1978 survey of the spring system. Dufurrena Pond 13 (T45N, R26E, Sec 17; Humboldt County, Nevada).— Dufurrena Pond 13 is the only reservoir on Virgin Creek between Virgin Creek Gorge and Thousand Creek. Water characteristics are typically those of Virgin Creek except that maximum depth is greater (almost 2 m) and aquatic vegetation is abundant. Alvord chubs are common to abundant just upstream and downstream of the reservoir but are rare in the pond itself. Gridley Springs (T44N, R27E, Sec 22; Humboldt County, Nevada).— Gridley Springs is a series of approximately 17 cool- water springs located on an alkali flat just south of the Gridley Lake playa. Many of the springs are little more than seeps, but a few have spring pools nearly 2 m deep with out- flow creeks 30 or 40 m in length. During April 1982, most of the spring waters were 418 Great Basin Naturalist Vol. 43, No. 3 10- 5- ■R U4 27 33 39 45 51 57 63 69 Standard Length in mm 75 81 ^ 87 93 Fig. 3. Length frequency of 44 Alvord chubs, Gila alvordensis, collected 14 April 1982 from Gridley Springs, Nevada. clear, with a temperature of 12 C (air 10 C). Rushes, Juncus sp., were the dominant plants around the springs, with some larger springs also harboring pondweed, Potamogeton sp., and cattails, Tijpha sp. Only one of the 17 springs examined contained Alvord chubs. This spring is located near the northwestern margin of the Gridley Springs series. Alvord chubs were found in the outflow creek, which extended approximately 40 m and was 3 m wide at its greatest extent. Only a trace of current could be detected in the creek. The water was imusually turbid, visibility 4 cm, with a maximimi depth of 30 cm. Water temperature was 11.5 C (air 6.8 C) on 14 April 1982. The bottom was silt. Unlike most of the larger springs in the area, this spring contained only rushes along its margin. The Gridley Springs area is overgrazed by cattle and horses, but it is not known to what extent this is detrimental to the Alvord chub population. Alvord chubs were not abundant in the outflow creek, but were common enough to collect 50 fish in three short seine hauls. Forty-four individuals collected in April 1982 ranged in size from 27 to 91 mm SL, but were mostly 30 to 38 mm SL (Fig. 3). The sex ratio of 32 individuals greater than 35 mm SL was 1 5 :3 ? . West Spring (T44N, R27E, Sec 20; Hum- boldt County, Nevada).— West Spring issues from the base of Big Mountain and forms West Creek, which flows into the alkali flat south of Gridley Springs. The waters of West Spring are clear and shallow. Maximum depth is 12 cm over a gravel and sand sub- strate. Water and air temperatures recorded during summer were 21.8 and 20.4 C, respec- tively. Alvord chubs are common in the spring and its outflow creek. The largest of 33 individuals collected on 18 August 1978 was 62.8 mm SL. The sex ratio of 30 adults greater than 40 mm SL was 1^:1$. Alvord chubs collected during August from West Spring fed exclusively or almost exclusively on small hydrobiid snails. This snail, which occurs in great abundance, apparently repre- sents an undescribed species endemic to West Spring (Jerry Landye, pers. comm.). West Creek (T44N, R27E, Sec 20, 28, 29; Humboldt County, Nevada).— West Creek flows for nearly 3 km before emptying into the alkali flat approximately 2 km south of Gridley Springs. Because West Creek is formed by West Spring, water characteristics are similar for both. The creek is quite small, often 1 to 2 m in width and less than 15 cm deep. Current is moderate in the upper reaches but slows considerably upon reaching the flat. Alvord chubs occur throughout the creek, but are somewhat smaller than those in West Spring. Discussion The Alvord cutthroat trout, Alvord chub, and Borax Lake chub are all restricted in dis- tribution to waters of the Alvord Basin. The Alvord cutthroat trout is now extinct but for- merly occurred in larger creeks of the basin. Hybridization with introduced trout caused the demise of the native form. The Borax Lake chub has the most restricted natural dis- tribution of the three fishes, occurring only in Borax Lake and adjacent lake outflows. The Alvord chub is relatively widespread in the basin and was recorded from 16 localities, in- cluding Bog Hot Reservoir, Bot Hot Creek, Thousand Creek Spring, an unnamed spring. July 1983 Williams, Bond: Native Fishes 419 Dufurrena Pond 19, Dufiirrena Pond 22, Du- furrena Pond 13, West Spring, and West Creek as new locality records. Because of the fragility of the small aquat- ic habitats and the overall paucity of water in the basin, the two extant native fishes are easily threatened by the activities of man. The naturally restricted range of the Borax Lake chub and threats from geothermal energy development prompted the American Fisheries Society to list the species as threat- ened in 1979 (Deacon et al. 1979). Also dur- ing 1979, several portions of the north and east shoreline of Borax Lake were altered so that overflow waters exited the lake to the north and east rather than to the south and west, as was the historical condition. This al- teration caused Lower Borax Lake as well as the marsh and pools to the south and west to dry, thus eliminating Borax Lake chubs from these waters. Leasing of surrounding lands for geothermal exploration and alteration of the shoreline caused the U.S. Fish and Wild- life Service temporarily to list the Borax Lake chub as an endangered species on 28 May 1980. As a result of the listing, geother- mal exploration was prohibited from a one- mile buffer zone around Borax Lake. The emergency listing has since been supple- mented by a final rulemaking that designated the species as endangered pursuant to the Endangered Species Act. The Alvord chub has fared better than the Borax Lake species because of its wider distribution. Never- theless, competition with exotic guppies has extirpated the Thousand Creek Spring popu- lation of Alvord chubs, and other populations are threatened by habitat alteration. The Al- vord chub appears easily eliminated by the presence of exotic fishes. Thousand Creek reservoirs stocked with game fish, such as Dufurrena Ponds 20 and 21, lack Alvord chubs. White crappie, Pomoxis annularis, pumpkinseed, Lepomis gibbosiis, and large- mouth bass, Micropterus sahnoides, were col- lected from Dufurrena Ponds 20 and 21. Borax Lake chubs are dwarf and typically mature at 30 mm SL. Adults are usually 33 mm to 45 mm SL and typically live for one year. A few Borax Lake chubs, mostly fe- males, live more than one year. Adult Alvord chubs are larger, achieving more than 100 mm SL in Trout and Virgin creeks. Even in very small springs, such as the unnamed spring and Gridley Springs, Alvord chubs achieve 90 mm SL. The presence of large chubs in the cool springs and creeks indicates a longer life span for the Alvord chub than typically occurs for the Borax Lake species. Borax Lake chubs spawn year around in their thermal lake habitat, but a spring spawning peak is indicated. Alvord chubs appear to spawn only once a year in their thermally fluctuating habitats. Both species of Gila are opportunistic omnivores, consuming primari- ly chironomids, microcrustaceans, and dia- toms. The Borax Lake species also consumed large quantities of terrestrial insects during summer and fall. The Alvord chubs in West Spring are unusual in that they are greatly dependent on the endemic hydrobiid snail for food. Acknowledgments Funds to study the Alvord Basin fishes were provided by U.S. Fish and Wildlife Ser- vice contract 14-16-0001-78025 to inventory the fishes of the Sheldon National Wildlife Refuge and by the Department of Fisheries and Wildlife at Oregon State University. The study greatly benefited from the field and editorial assistance of Cynthia D. Williams. Ray S. Taylor, J. J. Long, K. M. Howe, G. DeMott, B. Boccard, A. Tiehm, J. E. Deacon, M. S. Deacon, D. E. Deacon, and E. M. Lo- rentzen assisted with field collections. Robert R. Miller lent museum specimens and collec- tion notes of our late mentor, Carl Leavitt Hubbs. Reviews of this paper were provided by C. D. Williams, J. E. Deacon, E. P. Pister, and S. V. Gregory. This contribution is Tech- nical Paper 6510 of the Oregon Agricultural Experiment Station. Literature Cited Behnke, R. J. 1979. Monograph of the native troiits of the genus Salmo of western North America. Re- port to U.S. Fish and Wildlife Service. 215 pp. Bond, C. E. 1974. Endangered plants and animals of Oregon I. Fishes. Agricultural Exp. Station, Ore- gon State Univ. Spec. Rept. 205. 9 pp. Deacon, J. E., G. Kobetich, J. D. Williams, S. Co.NTRERAS, et al. 1979. Fishes of North America endangered, threatened, or of special concern: 1979. Fisheries 4:29-44. Hubbs, C. L., and R. R. Miller. 1948. The zoological evidence: correlation between fish distribution 420 Great Basin Naturalist Vol. 43, No. 3 and hydrographic history in the desert basins of western United States. Pages 17-166 in The Great Basin, with emphasis on glacial and post- glacial times. Bull. Univ. Utah, Vol. .38. HuBBS, C. L., AND R. R. Miller. 1972. Diagnoses of new cyprinid fishes of isolated waters in the Great Ba- sin of western North America. Trans. San Diego See. Nat. Hist. 17:101-106. LiBBEY, F. W. 1960. Boron in Alvord Valley, Harney County, Oregon. Ore-Bin. 22:97-105. Snyder, C. T., G. Hardman, and F. F. Zdenek. 1964. Pleistocene lakes in the Great Basin. U.S. Geol. Surv., Misc. Geol. Investigations, Map 1-416. Williams, J. E. 1980. Systematics and ecology of chubs {Gila: Cyprinidae) of the .\lvord Basin, Oregon and Nevada. Unpublished dissertation. Oregon State Univ. 175 pp. Williams, J. E., and C. E. Bond. 1980. Gila boraxohius, a new species of cyprinid fish from southeastern Oregon with a comparison to G. alvordensis Hubbs and Miller. Proc. Biol. Soc. Washington 93:293-298. Williams, J. E., and C. D. Williams. 1980. Feeding ecology of Gila boraxohius (Osteichthyes: Cypr- inidae) endemic to a thermal lake in southeastern Oregon. Great Basin Nat. 40:101-114. KRAMER PALOUSE NATURAL AREA Del W. Despain^'^ and Grant A. Harris^ Abstract.— The 27-acre Kramer Palouse Natural Area located in southeastern Washington State represents the best remaining example of what was once the most productive portion of the original Palouse Prairie. This area is being maintained in its pristine condition as a key to the past and as a memory to a unique and once extensive prairie land by the Department of Forestry and Range Management at Washington State University. Many of the most productive rangelands of the past are now agricultural lands. The Pa- louse Prairie of the inland Pacific Northwest is no exception. Once part of a vast prairie- land that extended throughout eastern Wash- ington and Oregon, as well as adjacent Idaho, most of this region has since been turned un- der by the plow, to become some of the most productive unirrigated farmland in the world. The Palouse Prairie occupies a region of relatively gentle topography, with the princi- pal relief being low hills having the general appearance of dunes. These wind-deposited loesal materials originated in the arid lands and volcanoes to the west, and were depos- ited on a basalt rock foundation. Fertile xe- rolls of silty and clay loam texture have de- veloped under the influence of a semiarid climate. Average armual precipitation is from about 400 to 600 mm (16-24 inches), coming mostly as rain or snow during fall, winter, and spring. Prefarming era vegetation was characteristic of a true grassland region, and was composed of dense stands of caespitose perennial grass species (Daubenmire 1970). The climax vegetation, though palatable and nutritious, apparently developed without significant grazing use. Dominant species are easily injured by close cropping, and under poor grazing management are replaced by introduced annual grasses (chiefly Bromiis tectorum). Archeologists estimate that the small bison poulation of the region became extinct about 2,000 years ago, and large her- bivore grazing was practically nil from that time until horses from early Spanish missions of the southwest were introduced in about 1730 (Osborne 1953). The region is so re- markably adapted to intensive wheat and pea cropping that today livestock grazing never has become an important land use, except in waste places. With the development of a side-hill com- bine and other technologically advanced farm equipment, only the very steepest "eye- brows" and slopes of the loesal deposits of the Palouse have not been tilled. Many of these small islands of native prairie have been used for other purposes and have changed dramatically over the past. Most re- maining uncultivated segments are so small as to have been changed to dense stands of brush or weeds through activities on adjacent land including tillage, herbicide applications, and fertilization. Probably the best remaining example of the more productive portions of this region is the Kramer Palouse Natural Area. Ownership has been obtained by Washington State Uni- versity in an effort to retain this unique spec- imen in its natural condition as a reference point to the past for demonstration and research. Historical Background The 27-acre Kramer Palouse Natural Area was once part of a producing wheat farm, but it was held out of production by virtue of steep topography and shape of the ownership boundaries of the farm. It reaches over the crest of a high ridge, too steep for even the intrepid Palouse area farmers to cultivate. 'Department of Forestry and Range Management, Washington State University, Pullman, Washington 99163. 'Present address: School of Renewable Natural Resources, University of Arizona, Tucson, Arizona 85721. 421 422 Great Basin Naturalist Vol. 43, No. 3 The ownership boundaries included the steepest part of the ridge top, with neighbors on three sides, and no access to the back side without crossing their land. Consequently, the area was not plowed until 1961 when the owner decided to cultivate the lower north- em extent of the parcel. However, after hav- ing made one pass with the plow, the farmer says he didn't have the heart to continue and left the tract untouched by further dis- tubance. The path of that one pass, half- circle in shape along the base of the ridge, can still be faintly seen today, but the vegeta- tion is now similar to adjacent sites. The area was occasionally used as a pas- ture for farm milk cows, but only a small part was noticeably changed where the cattle con- gregated for resting. A road that once tra- versed the area for movement of farm equip- ment and animals is the most evident scar of disturbance from the past, but even this has healed over with native vegetation similar to the undisturbed portions. Dr. Rexford F. Daubenmire is reported to have found the area in about 1955 during his search for bench mark natural areas in sup- port of his ecological studies. He established permanent study plots there at that time and continued to make observations as needed. He later showed the area to Dr. Grant A. Harris, who pursued the possibility of pur- chasing the tract through John P. Nagle, then chairman of the Department of Forestry and Range Management at Washington State University. The parcel, previously owned by Caroline Kramer, was not immediately avail- able for purchase. However, following pro- bate settlements for the Kramer estate, the University was able to purchase the 27-acre tract. The deed was filed in the imiversity's name on 28 March 1962. The Department of Forestry and Range Management, with spe- cific assignment to Dr. Harris, was given the responsibility for administration and mainte- nance of the Natural Area, and this assign- ment continues at present. Description The Kramer Palouse Natural Area, 27 acres in size, is located about 5 miles west of the farming community of Colton, Whitman County, in the southeastern portion of the state of Washington (N %, SE V4, NE V4 of Section 25, Township 13 North, Range 44 East, WPM). Aspects are generally steep on the south and north, with elevations of 805 to 869 m. Precipitation at the site averages ap- proximately 550 mm (22 inches) annually. Zonal vegetation in this environment is ex- pressed in the Festuca idahoensis/ Symphoricarpos alhus association (Dauben- mire 1970). A major part of the north slopes of the Natural Area supports this habitat type. Major species include the caespitose grasses, Idaho fescue {Festuca idahoensis), bluebunch wheatgrass {Agropijron spicatum), and June grass (Koelaria cristata), in associ- ation with shrubs dominated by snowberry {Symphoricarpos alhus) and wild rose {Rosa nutkana and Rosa woodsii). The type has a rich diversity of perennial forbs, creating a virtual sea of flowers across the Natural Area during much of the growing season. The snow-catching boundary fences along the north slope have sufficiently altered the mi- croclimate in places to increase the normal coverage of the tall shrub phase {Sym- phoricarpus), creating a border of dense shrubs including chokecherry {Frunus virgi- nana) and occasionally bittercherry {Frunus emarginata). There are also small patches of the black hawthorn/cow parsnip {Crataegus douglassii/ Heracleum kinatum) habitat type at the base of the steep north slopes, typical of bottom lands that are more moist than lo- cal zonal soils. The south face of the Natural Area supports a topographic climax cover of the bluebunch wheatgrass/ Sandberg's blue- grass {Agropyron spicatum /Foa sandhergii) habitat type typical of the more droughty vegetation zone to the west. Detailed soil and vegetation studies by Al- ler et al. (1981) on the site have revealed a "perched water table" situation in the solid steep north exposures. On this site, as well as two other similar locations, they found an un- described topoedaphic climax plant as- sociation dominated by Carex geyeri, as well as a Festuca phase and Symphorocarpos phase of Daubenmire's Festuca idahoensis/ Symphorocarpos alhus association that had not previously been reported. Soils on the site have been classified into the Palouse and Calouse series, which are rel- atively deep, well-drained soils that formed July 1983 Despain, Harris: Kramer Palouse Area 423 in loess deposits containing varying amounts of volcanic ash (Donaldson 1980). Approx- imately 80 percent of the surface is classified as Palouse-Thatuna silt loam, 10 percent Pa- louse silt loam (SE comer), and 10 percent Calouse silt loam (SW corner). The Palouse- Thatima series is a complex of approximately 50 percent Palouse and 50 percent Thatuna silt loam, mapped together because of their intermingled occurrence in the field. The Pa- louse silt loams are generally found on con- vex slopes and Thatuna on concave slopes, examples of which are found on south and north exposures of the natural area, respec- tively. All the soils found there are deep, well-drained, and fertile, and, in cultivation, produce excellent crops of wheat, lentils, peas, barley, and alfalfa. An abimdance of wildlife inhabits the area, including coyotes, badgers, and occasionally whitetail deer. Management philosophy encourages non- destructive scientific study of entire ecosys- tems found in the Natural Area. Scientists from several imiversity departments, includ- ing the Universities of Idaho and Washing- ton, as well as from state and federal agencies, have established short-term biologi- cal studies there. In addition to vegetation, studies have investigated natural status of soils, rodents, insects, birds, and atmosphere, all in considerable detail. Interest in the Pa- louse Natural Area as a bridge to the past continues to increase as research interest in biological subjects increases, and the area will become ever more valuable as time passes. Access to the Natural Area is somewhat hampered because it is isolated from the lo- cal road system and is completely surrounded by cultivated land. The lack of easy access, however, simplifies maintenance of the un- disturbed characteristics of the area. Ar- rangements have been made with the neigh- bor on the west to walk one-half mile along his fence line southward from Rim Road near its junction with the Colton— Wawawai Road. This provides the principle access for observation and study. Management Problems Maintaining a natural area in its natural state, especially one as small as the Kramer Area, is not easy. Most of the problems stem from the "unnatural" interface of the area with adjacent cultivated farmland. There is some indication that the dense cover of tall shrubs that has developed along some of the fence lines is migrating inward and increasing its extent over the area. The potential loss of the grass-forb- and low- shrub-dominated communities may become a concern in the future. Because of good cover and protection pro- vided by the shrubs along the fence line, and because of availability of adjacent cultivated crops as forage, the population of Columbian ground squirrels (Citellus columhianus) has increased dramatically around the perimeter of the track. In the past, these rodents have stripped adjacent field crops as far as 100 feet out from the boundary. Following com- plaints from neighboring farmers, attempts were made to control excessive populations of the squirrels, but most methods considered or tried proved to be either largely unsuc- cessful or involved equipment that was diffi- cult to get to the inaccessible area or that would cause undue disturbance. Recently, however, a very successful control effort has been implemented using a product with the trade name "Ramik Green," made by Velsi- col Chemical Corporation. This rodenticide was placed in bait stations located around the perimeter of the area during early spring. The immediate question that has been raised is whether or not this control of rodents changes the natural balance of the ecosystem. It is our opinion that rodent populations pri- or to control were artificially high due to the adjacent crops as a forage source. No attempt has been made to completely destroy rodent populations, with control efforts directed only at critical locations around the perime- ter of the Natural Area. The alternative, legal action taken by neighboring farmers due to obvious crop damage, would clearly endan- ger the status of the Natural Area. Another problem of the intensive agricul- ture interface is herbicide drift from adjacent fields, primarily 2-4-D. The most obvious im- pact is damage to shrubs along the bound- aries. What changes in species composition herbicide drift causes is unknown at this time, but it does not appear to be a serious problem as far as maintaining natural condi- tions over most of the area. 424 Great Basin Naturalist Vol. 43, No. 3 Except for cheatgrass [Bromus tectorum) along the ridge top, noxious, introduced weeds now common to the region have largely been excluded from the unit by the well-established stands of native vegetation. Canada thistle {Cirsium arvense) has ap- peared in a couple of places and could possi- bly become a problem to deal with in the future. Research use is encouraged, but care is taken to limit the kinds of uses to those that can be made without significant disturbance to its natural character. A system of approval has developed in which the applicant submits a copy of a detailed proposal for review by the administrator. If destructive procedures are found, negotiations usually modify these so that the work can be completed. Visits by undergraduate classes are not encouraged. Due to limited access to the Natural Area, disturbance by the general public has not been a problem. Public use has been limited primarily to upland game-bird hunters. Literature Cited Aller, a. R., M. a. Fosberg, M. C. LaZelle, and A. L. Falen. 1981. Plant communities and soils of north slopes in the Palouse region of eastern Washington and northern Idaho. Northwest Sci. 55(4):248-262. Daubenmire, R. F. 1970. Steppe vegetation of Washing- ton. Washington Agr. Res. Center Tech. Bull. 62, Pullman, Washington 99164. Do.NALDSON, N. C. 1980. Soil survey of Whitman Coun- ty, Washington. Soil Conser. Serv. U.S. Dept. Agr. in cooperation with Washington State Univ. Osborne, D. 1953. Archaeological occurrence of prong- horn antelope, bison, and horses in the Columbia Plateau. Sci. Monogr. 77:260-269. WINTER FOOD HABITS OF COUGARS FROM NORTHEASTERN OREGON Chris Maser' and Ronald S. Rohweder- Abstract.— Sixty-four cougar {Felis concolor) stomachs and 41 intestinal tracts were examined for food items in northeastern Oregon from 1976 through 1979. Food items, in order of decreasing frequency, were mule deer {Odo- coileus hemionus). North American elk (Cervus elaphus), porcupine (Erethizon dorsattim), snowshoe hare [Lepus americanus), and deer mouse {Peromijsciis maniciilatus). Bounties existed on cougars in Oregon from 1843, when the Oregon Territorial Gov- ernment offered them on most "predators," until the boimty system was repealed by the 1961 Oregon Legislature (Ebert 1971, Kebbe 1961). Once found throughout most of Ore- gon, the decline of the cougar during the 1950s and early 1960s aroused concern for its continued existence within the state. The cougar was classified as a game animal in September 1967 (Oregon State Game Com- mission 1967). The hunting season was imme- diately closed and remained closed until De- cember 1971, when the first controlled hunting season was opened. Twenty-two cou- gars were killed by hunters during the De- cember 1971 and December 1972 himting seasons. Little is generally known about food habits of the cougar (Young and Goldman 1964), particularly in Oregon, where indiscriminate boimty hunting kept cougar populations too low for such studies. Maser et al. (1981) and Toweill and Meslow (1977) discussed cougar food habits in general; Toweill and Meslow (1977) also discussed the food habits of those cougars killed during the 1971 and 1972 hunting seasons. The purpose of this paper is to present data on the winter food habits of cougars from northeastern Oregon and to of- fer some tentative interpretations of these data. Methods Sixty cougars were obtained from hunters during four one-month hunting seasons (De- cember 1976, 1977, 1978, 1979) in north- eastern Oregon counties: Baker (5), Union (15), and Wallowa (40). An additional four cats were obtained from Umatilla (1) and Wallowa (3) counties; one was killed illegally and three were killed because of their prox- imity to livestock. Cats, killed by himters, were brought into an Oregon Department of Fish and Wildlife office within 48 hours of being killed. Most cats were received intact, but a few had been field dressed. Each individual was sexed, weighed, mea- sured, and, if intact, eviscerated. Each cat's heart, lungs, liver, stomach and intestinal tract, and reproductive organs were placed in separate plastic bags, labeled, and quick- frozen for later analysis. The present food habit study was done in conjunction with a study of endoparasites, which necessitated separately examining the stomach, small in- testine, and large intestine. Each cougar thus had three separate analyses for food items. This procedure worked well because we could determine what appeared to be tlie contents of two meals for each cat that con- tained food in its alimentary canal— one meal in the stomach and a different meal in the co- lon. The two meals "mixed" in the small in- testine. Thus, by identifying the stomach con- tents first, the colon contents second, and the small intestine contents third, we had a cross- check on the content determinations. Mate- rials from the small intestine usually con- tained elements of both stomach and colon contents and have not been included in the discussion. Further, by identifying the plant 'U.S. Department of the Interior, Bureau of Land Management, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon 973.3L ■Oregon Department of Fish and WildUfe, Route 2, Box 2283, La Grande, Oregon 97850. 425 426 Great Basin Naturalist Vol. 43, No. 3 material in the alimentary canal, we could determine, by tree associations and by infer- ence, the general habitat within which the cougar had taken its meal. Results and Discussion Results of the food habit study are given in Tables 1 and 2. The sexes of cougars analyzed during this study probably approximates a random sample (Table 3). Although cougars are, in some sense, opportunistic predators (Robinette et al. 1959, Sitton 1977) their main diet was mule deer. North American elk, and porcupine (Table 1), which concurs with studies reviewed in Toweill and Meslow (1977). Stomachs ranged from being empty to containing 3.5 kilograms of mule deer. Mule deer was the most frequently con- sumed prey; North American elk was second (Table 1). In eight cases, the consumed elk could be classified as adult or calf. From the limited sample, the five male cougars that had eaten, and presumably killed, adult elk were in the upper size-limit of the overall sample: 68 kg (150 lbs)— the largest cougar— 66 kg (146 lbs), 64 kg (140 lbs), 64 kg (140 lbs), and 62 kg (137 lbs). The average weight of the five male cats was 64.8 kg (142.7 lbs). On the other hand, the three cats that had ea- ten known calf elk were a female (32 kg— 70 lbs), a male (41 kg— 91 lbs), and another fe- male (50 kg— 110 lbs). The average weight of these three cats was 41 kg (91 lbs). The 50-kg female that killed a calf elk was 12.3 kg (27 lbs) lighter than the smallest of the males that killed an adult elk. Thus it seems that the larger a cougar, the larger a prey animal it can kill, and the more energy efficient such a kill will be. Porcupine would seem to be an energy- efficient meal as soon as a young cougar is old enough to kill because these large rodents are slow, easily caught, and seem to be read- ily dispatched by cougars. Although porcu- pines occurred in the diet with only 10.6 per- cent frequency in stomachs and 5.2 percent frequency in colons in our study, Robinette et al. (1959) found them to account for 19 per- cent of the cougar's diet, based on scat analy- sis, in Utah and Nevada. Evidence— in the form of quills embedded in and around the gum lines, the skinned shoulders and feet, and embedded in stomach walls— indicated that most cougars encounter porcupines at some time during their life. Such quills, repre- sented by their embedded tips, appear as dark streaks. Apparently, a cougar's body readily absorbs the softer, light shaft of a quill but not the harder, dark tip. Cougars seem to be variously adept at eat- ing porcupines. For example, some cougars appear to avoid the quills as much as possible and have only a few hairs mixed with the porcupine flesh in their digestive tract, whereas others eat almost everything. In ad- dition, a cougar killed in 1973 had eaten a porcupine about an hour prior to being shot. It had consumed the entire porcupine, except the head and digestive tract. The quills had already begun to soften in the cat's stomach. The proportion of a cougar's diet that is composed of porcupine is probably related to the availability of the prey. Connolly (1949 cited in Robinette et al. 1959), for example. Table 2. Miscellaneous associated items consumed by 60 cougars killed in December. Content Stomach % frequency Colon % frequency Table 1. Prey species consumed by b 0 cougars killed Grass 20.0 23.8 in December. Douglas-fir needles 17.1 4.8 Grand fir needles 14.3 19.0 Stomach Colon Ponderosa pine needles Engelmann spruce needles 14.3 14.3 Content % frequency % frequency 11.4 Mule deer 55.3 42.1 9.5 Elk 21.3 15.8 Twigs 5.7 4.8 Porcupine 10.6 5.2 Soil 5.7 23.8 Unidentified hair 6.4 10.5 Alder leaf 2.9 — Snowshoe hare 4.3 _ Lichen (Alectoria Bird 2.1 _ fremonti) 2.9 — Lagomorph — 5.2 Larch needles 2.9 - Cougar hair - 21.1 99.9 Pebbles Total 2.9 — Total 100.0 100.1 100.0 July 1983 Maser, Rohweder: Winter Food of Cougars 427 indicated that, in his Utah study area, cougars killed one porcupine per week in winter. Of the 64 cougars examined in this study, plus 97 cougars examined prior to this study, none showed ill effects from encounters with por- cupines, even when quills remained em- bedded in a cat's tissues. Such lack of serious damage or infection from porcupine quills has also been noted in the fisher {Maries pen- nati) and spotted skunk {Spilogale piitorius) (Maser et al. 1981). The snowshoe hare occurred fourth (of the identifiable items) in the cougars' stomach contents (Table 1). Because these hares were relatively abundant in the coniferous forests of northeastern Oregon during our study, their low frequency (4.3 percent) indicates that they were taken incidentally by the cougars. The lagomorph remains in the colon (Table 1) were either snowshoe hare or mountain cottontail (Sylvilagus nuttalli), but they could not be identified to species once they reached the colon. The bird (Table 1) was probably a grouse. Miscellaneous items associated with food are given in Table 2. Other than grasses, some of which were intentionally eaten, iden- tifiable vegetation gave clues to the habitats in which the cats presumably had been hunt- ing and had consumed their prey. Of the five stomachs that contained elk and vegetation, Engelmarm spruce {Picea engelmannii) oc- curred in 20 percent, western larch (Larix oc- cidentalis) in 20 percent, grand fir {Abies grandis) in 40 percent, and ponderosa pine {Pinus ponderosa) in 20 percent. Of the 13 stomachs that contained deer and vegetation, Engelmann spruce occurred in 15 percent, grand fir in 8 percent, ponderosa pine in 38 percent, and Douglas-fir {Pseudotsuga men- ziesii) in 38 percent. From the conifer nee- dles in the stomachs, it seems that elk were killed primarily in denser, moister forests be- cause the Engelmann spruce, western larch, and grand fir accoimted for 80 percent of the needles, whereas ponderosa pine, character- istic of more open habitat, accounted for only 20 percent. With respect to mule deer, on the other hand, ponderosa pine-Douglas- fir, which occiu-s as a drier, more open forest, accounted for 76 percent of the conifer nee- dles, as opposed to the moister, denser forests of grand fir and Engelmann spruce, which represented 23 percent of the needles. Thus, it seems that the elk were usually killed in dense forest where the advantage would lie with the stalking cougar and the smaller, more easily subdued mule deer was most of- ten hunted in more open habitats. In addition to vegetation, several cats had eaten much soil— evidence of having cleaned up a kill. Conclusions Of the 60 cougars killed during the De- cember hunting season, 31.6 percent had vir- tually or totally empty stomachs, and 30 per- cent had empty colons. The 4 cougars killed because of their proximity to livestock also had virtually empty stomachs. Thus, if the stomachs or the colons are used independent- ly as the sole source of food habit data, a large sample is needed. If, on the other hand, both the stomach and colon contents are used independently but together as dietary sam- ples, the chances of getting adequate food habit data are good because it is unlikely that both stomach and colon are simultaneously empty. Acknowledgments Murray L. Johnson (Puget Sound Museum of Natural History, University of Puget Sound, Tacoma, Washington), Donald K. Grayson (Department of Anthropology, Uni- versity of Washington, Seattle, Washington), Maurice Hornocker (USDI, Fish and Wildlife Service Cooperative Wildlife Research Unit, University of Idaho, Moscow, Idaho), and Paul E. Ebert (Oregon Department of Fish and Wildlife, Portland, Oregon) read and im- proved the paper. Phyllis Taylor-Hill (USDI Table 3. Year of capture, number, and sex of cougars studied in northeastern Oregon. No. of No. of No. of Year cougars males females 1976 4 3 1 1977 21 13 8 1978 22 10 12 1979 17 7 10 Total 64 33 31 428 Great Basin Naturalist Vol. 43, No. 3 Bureau of Land Management, Forestry Sci- ences Laboratory, Corvallis, Oregon) typed the various drafts of the manuscript. The following Oregon Department of Fish and Wildlife personnel helped collect the cougar viscera: Ronald Bartels, Vic Coggins, Paul Ebert, Mark Henjum, Dick Humphreys, Mike Kemp, and Walt Van Dyke. Laboratory space and partial financing for this study were provided by the USDA Forest Service, Pacific Northwest Forest and Range Experi- ment Station, Project 1701, Range and Wild- life Habitat Laboratory, La Grande, Oregon. We sincerely appreciate their help. Literature Cited Connolly, E. J., Jr. 1949. Food habits and life history of the mountain Hon, Felis concolor hippolestes. Un- pubUshed thesis, Univ. of Utah, Salt Lake City. 176 pp. (Cited in Robinette et al, 1959). Ebert, P. W. 1971. The status and management of the felids of Oregon. Pages 69-71 in S. E. Jorgensen and L. D. Mech, eds., Proceedings of a sym- posium on the native cats of North America. U.S. Department of the Interior, Fish and Wildl. Serv., Bur. Sport Fish and Wildl, Region 3. Kebbe, C. E. 1961. Bounties. Oregon State Game Comm. Bull. 16:3, 6-7. Maser, C, B. R. Mate, J. F. Franklin, and C. T. Dyrness. 1981. Natural history of Oregon Coast mammals. U.S. Department of Agricutlure, For- est Service Gen. Tech. Rept. PNW-1.33. Pacific Northwest Forest and Range Expt. Sta., Portland, Oregon. 496 pp. Oregon State Game Commission. 1967. Cougar. Ore- gon State Game Comm. Bull. 22. Robinette, W. L., J. S. Gashwiler, and O. W. Morris. 1959. Food habits of the cougar in Utah and Ne- vada. J. Wildl. Mgt. 23:261-272. SiTTON, L. W. 1977. California mountain lion in- vestigations with recommendations for manage- ment. State of California Resour. Agency, Dept. Fish and Game Wildl. Restoration Proj. W-51-R, Big Game Invegations. 35 pp. Toweill, D. E., and E. C. Meslow. 1977. Food habits of cougars in Oregon. J. Wildl. Mgt. 41:576-578. Young, S. P., and E. A. Goldman. 1964. The puma, mysterious American cat. Dover Publ., Inc., New York. 358 pp. A NEW SPECIES OF PENSTEMON (SCROPHULARIACEAE) FROM THE UINTA BASIN, UTAH Elizabeth Neese' and Stanley L. Welsh' Abstract.— Described as a new species is Penstemon flowersii Neese & Welsh from Utah. An illustration is pro- vided. Tlie species is named in honor of the late Seville Flowers, professor of botany at the University of Utah. The genus Penstemon is large and complex within the state of Utah. The flowers of plants of this genus are among the most beau- tiful within the state. Several taxa have been described as new from the intermountain re- gion during recent years. It is not surprising that another such novelty should occur in the region. The species was discovered during in- vestigations of the rare plants in the Uinta Basin, a region noted for its narrowly dis- tributed endemics. Penstemon flowersii Neese & Welsh, sp. nov. Species haec ab P. immanifesto N. Holm- gren in staminodiorum barbis multo breviore, et ab P. carnoso Pennell in floribus roseo non violaceo-caeruleis, ab uterque foliis basalibus nullis differt. Perennial glabrous glaucous herbs, with simple ascending stems arising from a branching woody caudex, 8-25 (32) cm tall, the basal rosette lacking; cauline leaves all entire, fleshy-thickened, (1.5) 2-5.5 cm long, (4) 10-25 mm broad, the lower shortly petio- late, spatulate, the middle ones larger, sessile, lanceolate or elliptic, obtuse, the upper re- duced, broadly ovate, acute; thyrsus cylindric (not secund), with 4-9 verticils, the cymes many flowered; calyx glabrous, 5-6.5 mm long, the lobes broadly lanceolate, acumi- nate, the margin scarious, suffused with rose; corolla 15-18 mm long, rose within, the striae dark rose-pink, the limb ampliate, 10-12 mm in diameter; staminode equaling the tube, not exserted, the apex shortly bar- bellate (to 0.1 mm long); stamens included, the anthers glabrous, dehiscent throughout and in the connective, not explanate, the sacs opposite, 1-1.2 mm long; capsules 7-10 mm long. Type: USA. Utah: Uintah Co., T3S RIE S9-10, 5.6 km W of Randlett, 12 May 1980, Neese & White 8609 (Hoiotype: BRY; Iso- types: NY, US, RM, CAS, UTC, MINN). Paratypes: Utah. Uintah Co., T3S RIE SIO, 4.8 km W of Randlett, 12 May 1898, Neese & White 8600 (BRY, UT, NY, MO); do T3S RIW S3, 5.8 km S of U.S. Hwy 40, 14 km W of Randlett, 12 May 1980, Neese & White 8606 (BRY, NY, CAS); do T3S RIE SIO, 4.8 km W of Randlett, 16 May 1979, E. Neese & B. Welsh 7212 (BRY, NY, GH, MO). Duchesne Co., T3S R2W S21, 3.2 km WNW of Myton, 16 May 1979, Neese & B. Welsh 7218 (BRY, NY); do T3S R2W S12, 4 km due N of Myton, 15 May 1980, Neese & White 8662 (BRY, NY, UC). The Flowers beardtongue grows in shad scale communities on pale-colored clay slopes and benches between 1,500 and 1,600 m, where old terraces of the Uinta Formation in the Duchesne River drainage are mantled with Pliocene or Pleistocene pedimen,tal gravels. It is common on such habitat in an area of about 8 X 25 km between Randlett and Myton. The species is remarkable in its uniformity in regard to both morphology and habitat. The plants, with their dusty pink flowers and pale gray-green foliage are in- conspicuous against the gray clay on which they grow. The near congener, P. immani- festus, of central eastern Nevada and western Utah possesses a more prominently bearded staminode. Penstemon carnosus Pennell is similar in diagnostic characteristics, but the Flowers beardtongue is quite distinctive in its smaller stature, tufted, usually numerous 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Prove, Utah 84602. 429 430 Great Basin Naturalist Vol. 43, No. 3 mm Fig. 1. Penstemon floiversii Neese & Welsh: A, Habit. B, Flower. C, Anther-sacs. D, Staminode. July 1983 Neese, Welsh: A New Penstemon 431 stems, absence of a basal rosette, and pink (not lavender-blue) flowers. Penstemon car- nosus is a species of the western Colorado Plateau, from the San Rafael Swell and the Henry Mountains westward to Aquarius Plateau (Holmgren 1978). The plant is named to honor the memory of Dr. Seville Flowers, late professor of bot- any at the University of Utah. Dr. Flowers was a student of lichens, mosses, and higher plants, and his imtimely passing has left a void in the understanding of the plants of Utah and the West. Acknowledgments We express thanks to Noel H. Holmgren for his examination of the type materials, and for his comments regarding relationships of this taxon. Kaye Thome provided the illustra- tions, and for this we are grateful. Literature Cited Holmgren, N. H. 1978. An overlooked new species of Penstemon (Scrophulariaceae) from the Great Ba- sin. Brittonia 30:334-339. A COMPARATIVE STUDY OF COYOTE FOOD HABITS ON TWO UTAH DEER HERDS Jordan C. Pederson' and R. Gary Tiickfield' Abstract.— Coyote {C-anis latrans) scats from two southern Utah deer herd units were collected and analyzed to establish diet selection. The category showing the most consistent frequency of occurrence was mule deer Odo- coileus hemionus; lagomorphs were next. Formal statistical analysis revealed that the only significant difference in coyote food habits between herd units was in the frequency of rabbits eaten. These data suggest that coyotes in this region of southern Utah show a comparatively higher preference for mule deer but, at the same time, do not eat deer in proportion to the frequency of their occurrence. Documented reductions in deer popu- lations in most southern Utah mule deer (Odocoileus hemionus) herds have led to speculation concerning the cause or causes for these declines (Workman and Low 1976). This paper investigates the hypothesis that coyote {Canis latrans) predation may reflect differential selection for deer. This was done by assessing coyote food habits in two adja- cent deer herd units in southern Utah's San Juan Coimty. Areas studied included the Blue Moimtain (31 A) and Elk Ridge (3 IB) herd units. Since the deer population is known to be larger within the Blue Mountain unit (Jense 1981), an examination of coyote scats from both areas could indicate whether deer occur in coyote diets in relationship to herd size. If this relationship was positive at a high level of significance, it would lend some cre- dence to the coyote predation hypothesis. Study Area The San Juan-Blue Mountain deer herd unit (31 A) is, for the most part, that portion of San Juan County east of the North and Table 1. Relative frequency of occurrence of food items in coyote diets as determined from 460 scats collected from September 1977 to December 1979. (n) Blue Mountain Perioc? Vegetation Rodent Deer Lagoniorph Cattle Bird Carrion 1 (18) 11.1 11.1 44.4 48.9 0.0 5.6 0.0 2 (15) 26.7 20.0 46.7 33.3 0.0 0.0 0.0 3 (105) 23.8 3.8 61.0 15.2 1.0 1.0 6.7 4 (41) 4.9 0.0 14.6 87.8 0.0 2.4 0.0 5 (12) 16.7 16.7 16.7 91.7 0.0 0.0 0.0 6 (62) 21.0 16.1 58.1 33.9 1.6 1.6 .30.6 7 (37) 59.5 40.5 37.8 13.5 10.8 10.8 21.6 8 (12) 14.3 50.0 46.4 25.0 17.0 10.7 0.0 Totalis (318) mean'^ 26.8 19.8 40.7 42.4 3.9 4.0 7.4 Standard deviation*^ 16.60 17.27 17.17 30.56 4.81 4.52 12.04 ^Period Dates 1 Sep-Dec 1977 2 Jaii-Jiin 1978 3 Jul- Sep 1978 4 Oct-Dec 1978 5 Jan-Mar 1979 6 Apr-Jun 1979 7 Jul-Sep 1979 8 Oct-Dec 1979 'Utah Wildlife Resources, 115 North Main Street, Springville, Utah 84663. ^Department of Statistics, Brigham Young University, Provo, Utah 84602. Present address: Department of Biology Indiana University, Bloomington, In- diana 47405. 432 July 1983 Pederson, Tuckfield: Coyote Food Habits 433 South Cottonwood drainages. Its highest point is Abajo Peak at 11,360 ft (3,463 m), and it ranges to a low elevation at Bluff City of 4,473 ft (1,363 m). The summer range area of this unit is 153 mi^ (396 km^), and the area of the winter range is 1,394 mi 2 (3,610 km 2). Major vegetational types within this unit are conifer, aspen, mountain brush, sagebrush, pinyon-jimiper, and blackbrush (Coles and Pederson 1968, 1969). The San Juan-Elk Ridge deer herd unit (3 IB) is that area of San Juan County west of the North and South Cottonwood Wash drainages. Horse Mountain, at 9,320 ft (2,840 m) elevation, is the highest point; and the lowest is also at Bluff City, which divides these two herd units. The area of the summer range is 195 mi^ (505 km^), and that of the winter range is 1,132 mi^ (2,932 km2). Major vegetational complexes include conifer, as- pen, mountain brush, sagebrush, pinyon- juniper, and salt desert shrub (Coles and Pe- dersen 1968, 1969). During the period from 1976 to 1979, the number of deer harvested per 1000 ha of summer range was 2.9 for the Blue Mountain unit and 1.10 for the Elk Ridge unit. The number of deer harvested per hunter day (ef- fort) for the same time period was 0.061 and 0.049, respectively. Materials and Methods Data on dietary selection were obtained from analyses of coyote scats collected along established roads. Scat analysis was chosen over stomach content analysis because a larger sample size could be collected during specific time periods and at specified local- ities without diminishing the predator popu- lation (Knowlton 1964, Meinzer et al. 1975). Scats were collected every three months dur- ing a 27-month period from 1 September 1977 to 31 December 1979, with the excep- tion of a 6-month lapse during period 2. Scats were air dried for a minimum of 30 days and then analyzed after thoroughly crumbling. All remains were identified with the aid of a binocular dissecting microscope, hair (Moore et al. 1974), and feather keys, as well as a ref- erence collection of skeletons and vegetation. Table 1 continued. Elk Ridge (n) Vegetation Rodent Deer Lagomorph Cattle Bird Carrion (0) _ _ — — — — - (4) 0.0 0.0 50.0 25.0 50.0 0.0 0.0 (26) 19.2 15.4 57.7 19.2 3.8 0.0 3.8 (5) 0.0 0.0 80.0 20.0 0.0 0.0 0.0 (22) 9.1 0.0 18.2 81.8 4.5 0.0 0.0 (18) 16.7 16.7 72.2 33.3 0.0 5.6 5.6 (31) 77.4 58.1 25.8 9.7 6.5 0.0 3.2 (36) 8.3 9.3 27.8 86.1 5.6 2.8 0.0 (142) 18.7 14.1 47.4 39.3 10.1 1.2 26.92 20.71 24.13 31.33 17.80 2.20 2.36 ''63.2 and 74.6 percent of all scats contained unidentifiable material from the Blue Mountain and Elk Ridge herd units, respectively. 'Computed as the average overtime periods 434 Great Basin Naturalist Vol. 43, No. 3 Food habits are reported as relative frequen- cy of occurrence. Comparisons between the two coyote pop- ulations were made using three statistical procedures, viz., normal approximation to two sample binomial data (Snedecor and Cochran 1967), stepwise logistic regression (Fienberg 1980), and stepwise discriminant analysis (Morrison 1976). The statistical com- puting programs PIF, PLR, and P7M, re- spectively, were employed from the BMDP series (Brown 1977). In the first statistical procedure, each scat was considered to represent a bernoulli trial for each category of remains identified. Hence the total number of scats from each herd unit was treated as a binomial random sample, of which a certain proportion con- tained remains but the complement did not. In the second procedure, we treated the lo- cation (herd unit) category as a "response" variable and all other dichotomous categories Table 2. Summary of coyote dietary studies of identified remains as "design" or explana- tory variables. The logic of the response vari- able was then regressed on the explanatory variables. In the final procedure, each scat was con- sidered to be a multivariate observation, i.e., a vector of remains categories. Discriminant analysis was then used to determine which variables (categories) best discriminated be- tween the two groups (herd units). Results and Discussion We collected and analyzed 460 coyote scats: 318 from the Blue Mountain imit and 142 from the Elk Ridge unit. Equal search ef- fort was not expended on both areas, and scat numbers are not indicative of coyote num- bers. The major food items found in the scats from both areas were mule deer, birds, car- rion, lagomorphs (black-tailed jackrabbit [Lepus calif ornicus], mountain cottontail Authority Study area Sample size Source Bond 1939 California 282 S and S Murie 1940 Yellowstone National Park 5,086 Scats Sperry 1941 8,339 Stomachs Murie 1945 British Columbia 311 Scats 1945 Montana 286 Scats 1945 Montana 67 Scats Fitch 1948 California 1,173 Scats Ferrel at al. 1953 California 2,222 Scats Fichter et al. 1955 Nebraska 747 Stomachs 1955 Nebraska 2,500 Scats Korschgen 1957 Missouri 770 Stomachs Korschgen 1957 Missouri 326 Scats Ozoga** et al. 1966 Michigan 92 Scats Gier 1968 Kansas 1,451 Stomachs Clark 1972 Utah and Idaho 186 Stomachs Hawthorne 1972 California 384 Scat Mathwig 1973 Iowa 151 Stomachs Richens et al. 1974 Maine 51 Stomachs Gipson 1974 Arkansas 168 Stomachs Meinzer et al. 1975 Texas 514 Scats 1975 Texas 55 Stomachs Niebauer et al. 1975 Wisconsin 3,353 S and S Nellis et al. 1976 Alberta, Canada 344 Stomachs Johnson et al. 1977 Arizona 224 Scats Ribic^ 1978 Colorado 54 Scats Neffetal. 1979 Arizona 65 102 Scats Scats Litvaitis et al. 1980 Oklahoma 361 Scats Springer and Smith'^ 1981 Wyoming 404 Scats ^Percent could not be determined from data presented. "Winter study only. '^Summer study only. "Largely carrion; irmards. heads, and feet. July 1983 Pederson, Tuckfield: Coyote Food Habits 435 [Sylvilagus nuttallii]), rodents (rock squirrel [Spertnophilus variegatus], least chipmunk [Eutomius 7Jiinimiis], Apache pocket mouse [Perognathtis opoche], and deer mouse [Per- omyscus riianiculatus]), and vegetation (Table 1). When results of our study are compared to data collected in 23 previous studies of coy- ote diets (Table 2) dating from 1939 through 1981, only two show deer occurring in the diets with greater relative frequency (Ozoga and Harger 1966, Hawthorne 1972). Coyote diets from both our study areas also showed a higher relative frequency of carrion than most other studies reported (Table 2). How- ever, since it was difficult to postively identi- fy carrion during the winter months, this cat- egory was not included in the statistical analyses reported hereafter. The greatest amoimt of fluctuation from one time period to another occurred in the category of la- gomorph remains. Mule deer were the diet- Table 2 continued. ary item showing the most consistent use (highest relative frequency) across collection periods occurring in four out of eight and four out of seven collection periods for the Blue Mountain and Elk Ridge herd units, re- spectively. Lagomorphs were the second most consistently used food item identified in scats, occurring in two of eight and two of seven collection periods, respectively. Analy- sis suggests coyotes could be a factor in the fluctuations of deer populations in these southeastern Utah herd imits. These results do not constitute evidence for a cause and ef- fect relationship. Mule deer may be killed and eaten by coyotes or they may be eaten as carrion. Deer carrion could occur as a result of winter stress, other predators, disease, par- asites, or other factors, but the reason for these mortality factors warrants further investigation. Table 3 contains the single category com- parisons of binomial proportions between Percentage of specimens in which item occurred Lagomorphs Rodents Carrion Livestock Birds Deer Vegetation 38.8 62.5 4.2 8.8 2.5 26.1 16.9 4.0 59.9 0.0^ 0.1 3.1 1.0 2.0 4.3.0 32.0 25.0 20.0 13.0 6.0 4.0 69.4 6.1 9.2 6.6 7.4 4.6 1.8 3L8 40.1 12.6 0.0 3.5 1.1 2.4 52.7 1.7 8.9 6.4 12.9 1.1 3.2 45.4 43.7 1.0 1.0 2.0 0.0 3.0 29.3 49.1 0.0' 23.2 18.1 18.5 0.0* 58.2 0.0» 0.0^ 26.1 44.1 0.4 3.6 23.0 0.0» 0.0» 30.5 33.7 7.6 16.0 55.3 36.3 8.6 13.8 22.0 2.9 7.9* 80.4 33.3 o.oa 2.8 14.7 0.0 23.5 17.0 69.8 83.0 2.1 .5.1 91.4 19.8 54.3 41.5 37.7 0.0* 24.8 0.0* 3.1 84.0 15.0 0.0» 10.3 2.1 2.0 1.6 5.7 74.2 0.0* 1.5 3.7 35.2 45.3 6L0 37.7 0.0* 31.1 21.2 0.0 64.2 19.6 19.6 0.0* 11.8 19.6 15.9 78.4 7.0 9.0 30.0 13.0 44.0 5.0 36.0 10.5 24.5 6.0 0.0 1.1 0.0 48.5 10.8 20.2 21.1 0.0 4.5 0.3 20.5 28.0 21.0 0.0* 0.0 12.1 26.9 36.2 3.0 22.0 0.0* 44.0^ 11.0 0.0* 7.0 27.0 19.4 0.0* 0.0 18.6 0.0 4.5 24.0 45.0 0.0 0.0 30.0 13.0 78.0 26.2 43.0 0.0 15.4 1.5 12.3 32.8 0.0 38.2 0.0 6.9 34.3 2.0 39.5 11.0 53.0 0.0 6.0 19.0 20.0 32.0 63.0 45.0 0.0* 0.0* 0.0* 5.0 42.0 436 Great Basin Naturalist Vol. 43, No. 3 herd units. Normally we would conclude a significantly higher (p = 0.27) proportion of coyote scats from the Elk Ridge herd unit contained lagomorph remains than did that of the Blue Mountain unit, but such a con- clusion would be somewhat misleading. All tests reported in Table 3 are not independent of one another since the information in each came from the same sample of coyote scats. One generally acknowledged and conserva- tive interpretation of these kinds of results uses a Bonferroni procedure (Neter and Was- serman 1974) in which the level of accept- able Type I error is divided by the number of simultaneous tests (six in this investigation). Hence, the "appropriate" significance level for the results in Table 3 (assuming P = .05) is .008, in which case none of the test results are significant. It is interesting to note that the only other category approaching the point of demonstrating even weak evidence in favor of a difference in coyote diets be- tween herd units was deer. The results of the stepwise discriminant analysis indicated the most important variable (category) to signifi- cantly discriminate between groups was lago- morphs (approx. F-statistic at 1st step = 4.941, p = .027). Similarly, the results of the stepwise logistic regression analysis indicated lagomorph remains were the only variable to account for a significant (x^ = 4.859 at 1st step, p = .028) amount of variability in the logit (response) variable. These results suggest coyotes do not in- clude deer in their diets based on the poten- tial frequency of occurrence of this food item. However, we did not conduct any si- multaneous census of deer numbers in either of the areas where scats were collected. Fur- ther investigation is warranted. Acknowledgments We thank Marlene and Robert Hasenya- ger, A. Ray Johnson, John C. Kimball, and Aurelia, Mary Ann, and Linda Pederson for Table 3. Cell frequencies and .statistical test results when each category of coyote scat material is considered to be a normal approximation to a two-sample binomial problem. 1 Lagomorph; s Deer Present Absent Total Present Absent Total Deer herd 111 207 318 151 167 318 2 65 77 142 56 86 142 Total 176 284 460 207 253 460 Pi" = .349 P.'' = .458 P," = .475 P,b = .394 z = 2.216 Prob. = .027 z = 1.603 Prob. = .109 1 Birds Cattle Present Absent Total Present Absent Total Deer herd 10 308 318 12 306 318 2 2 140 142 8 134 142 Total 12 448 460 20 440 460 P," = .031 P,b = .014 Pi" = .038 P,b = .056 z = 1.079 Prob. = .280 z = .904 Prob. = .366 1 Rodents Vegetation Present Absent Total Present Absent Total Deer herd 52 266 318 77 241 318 2 28 114 142 36 106 142 Total 80 380 460 113 347 460 Pi" = .164 P,b = .197 p," = .242 P,b = .254 z = .880 Prob. = .379 z = .263 Prob. = .793 1 = Blue Mountain 2 = Elk Ridge "Proportion of scats collected from the Blue Mountain Range containing the indicated remains "Proportion of scats collected from the Elk Ridge Mountain Range containing the indicated remains July 1983 Pederson, Tuckfield: Coyote Food Habits 437 their help with this study. We also acknowl- edge the help of Leonard Newlin, Albert W. Heggen and Norman V. Hancock of the Utah Division of Wildlife Resources. This study was supported by the Utah Division of Wild- life Resources and the Brigham Young Uni- versity Department of Statistics. Critical re- view of this manuscript was provided by Dwight Bimnell, Jerran T. Flinders, K. T. Harper, Clyde L. Pritchett, Alvin C. Ren- cher, and Bnice L. Welsh. Literature Cited Bond, R. M. 1939. Coyote food habits on the Lava Beds Monument. J. Wildl. Mgt. .3:180-198. Brown, M. B. 1977. BMDP-77 Biomedical Computer Programs, P-Series. Univ. of California Press, Berkeley. 880 pp. Clark, F. W. 1972. Influence of jackrabbit density on coyote population change. J. Wildl. Mgt. 36:343-356. Coles, F. H., and J. C. Pederson. 1968. Utah big game range inventory. Utah Div. Fish Game Pubi. 68- 2. 120 pp. 1969. Utah big game range inventory. Utali Div. Fish Game Publ., 69-2. 164 pp. Ferrel, C. M., H. R. Leach, and D. Tillotson. 1953. Food habits of the coyote in California. Califor- nia Fish and Game .39:301-341. Fighter, E., G. Schildman, and J. H. Sather. 1955. Some feeding patterns of coyotes in Nebraska. Ecol. Monogr. 25(l):l-.37. Fienberg, S. E. 1980. The analysis of cross-classified categorical data. MIT Press, Cambridge, Eng- land. 198 pp. Fitch, H. S. 1948. A study of coyote relationships on cattle range. J. Wildl. Mgt. 12:7.3-78. Gier, H. T. 1968. Coyotes in Kansas. Kansas State Univ. Agr. Exp. Stn.Bull. 393. 118 pp. GiPsoN, p. S. 1974. Food habits of coyotes in Arkansas. J. Wildl. Mgt. 38:848-853. Hawthorne, V. M. 1972. Coyote food habits in Sagehen Creek Basin, northeastern California. California Fish and Game 58:4-12. Jense, G. K. 1981. Utah big game investigations and management recommendations 1980-81. Utah State Div. of Wildl. Publ. No. 81-4. Johnson, M. K., and R. M. Hansen. 1977. Foods of coy- otes in the lower Grand Canyon, Arizona. J. Ari- zona Acad. Sci. 12:81-83. Knowlton, F. F. 1964. Aspects of coyote predation in south Texas with special reference to white- tailed deer. Unpublished dissertation. Purdue Univ. 189 pp. Korsghgen, L. J. 1957. Food habits of the coyote in Mis- souri. J. Wildl. Mgt. 21:424-435. Litvaitis, J. A., and J. H. Shaw. 1980. Coyote move- ments, habitat use and food habits in south- western Oklahoma. J. Wildl. Mgt. 44:62-68. Mathwig, H. J. 1973. Food and population character- istics of Iowa coyotes. Iowa State J. Research 47:167-189. Meinzer, W. p., O. N. Clegkert, and J. T. Flinders. 1975. Food niche of coyotes in the rolling plains of Texas. J. Range Mgt. 28:22-27. Moore, T. D., L. E. Spenge, and C. E. Dugnolle. 1974. Identification of the dorsal guard hairs of some mammals of Wyoming. Wyoming: Game and Fish Dept. Bull. 14. 177 pp. Morrison, D. F. 1976. Multivariate statistical methods. 2d ed. McGraw-Hill, New York. Murie, a. 1940. Ecology of the coyote in Yellowstone, Fauna Series No.' 4, 1940. U.S. Govt. Print. Off., Washington, D.C. 206 pp. Murie, O. J. 1945. Notes on coyote food habits in Mon- tana and British Columbia. J. Mammal. 26:33-40. Neff, D. J., AND N. WooLSEY. 1979. Effects of predation by coyotes on antelope fawn survival on Ander- son Mesa. Spec. Rept. No. 8. Fed. Aid in Wildl. Restoration Proj. W078-R. Phoenix, Arizona. Ari- zona Game and Fish Dept. Nellis, C. H., and L. B. Keith. 1976. Population dy- namics of coyotes in central Alberta, 1964-68. J. Wildl. Mgt. 40:389-399. Neter, J., and W. M. Wasserman. 1974. Applied linear statistical models. Richard D. Irwin, Inc., Home- wood, Illinois. Niebauer, T. J., and O. J. Rongstad. 1975. Coyote food habits in northwestern Wisconsin. Proc. 1975 Predator Symp. Montana Forest and Conser. Exp. Stn., University of Montana, Missoula. OzoGA, J. J., AND E. M. Harger. 1966. Winter activities and feeding habits of northern Michigan coyotes. J. Wildl. Mgt. 30:809-818. Ribic, C. a. 1978. Summer foods of coyotes at Rocky Flats, Colorado. Southwestern Nat. 23:152-153. RiCHENS, V. B., AND R. D. HuGiE. 1974. Distribution, taxonomic status, and characteristics of coyotes in Maine. J. Wildl. Mgt. 38:447-454. Snedecor, G. W., and W. G. Cochran. 1967. Statistical methods. Iowa State Univ. Press, Ames. 593 pp. Sperry, C. C. 1941. Food habits of the coyote. U.S. Fish Wildl. Ser. Res. Bull. 4. 70 pp. Springer, J. T., and J. S. Smith. 1981. Summer food habits of coyotes in central Wyoming. Great Ba- sin Nat. 41:449-456. Workman, G. W., and J. B. Low, eds. 1976. Mule deer decline in the West: a symposium. Utah State Univ., Logan. 134 pp. A SMALL CARNIVORE SURVEY TECHNIQUE Tim W. Clark' and Thomas M. Campbell III- Abstract.— a track station survey method for small, nocturnal mammalian carnivores is described. The method has been field tested under a wide varietv of conditions. Stations were visited by rodents, mustelids, and canids, as well as insects and birds. Small carnivores, because they often range relatively widely, occur at low densities, and tend to be solitary and nocturnal, are one of the more difficult mammal groups to study. Five basic categories of carnivore survey and census methods were described by Linhart and Knowlton (1975): (1) direct counts or capture-recapture, (2) counts of sign (dens, tracks, or droppings), (3) questionnaires and boimty payments, (4) catch per imit of effort (trap-nights), and (5) elicited responses to man-made stimuli (frequency of visitations to scent stations, howl responses to sirens). These methods vary in their effectiveness by species and habitat type. We used a track recording method em- ploying two varieties of lures (olfactory and acoustic) to elicit small carnivore responses between 1978 and 1982. Field-use deter- mined: (1) efficacy, (2) ease and thrift of con- struction and setup, and (3) durability and amount of maintenance required. Elicited re- sponses sought included: (1) deposition of tracks and scats at stations and (2) drawing animals to these stations to make them more visible during spotlight surveys. Our track- recording technique was a combination and modification of methods used by Mayer (1957) and Justice (1961) to determine small mammal presence and by Linhart and Knowlton (1975) to index coyote populations. We also compared our track stations accord- ing to the above objectives with Linhart and Knowlton's (1975) station design. We were particularly interested in small carnivores on prairie dog colonies, and therefore placed track stations of both types on white-tailed {Cynomys leucurus), Gunnison's (C. gun- nisoni), and black-tailed prairie dog (C. hi- dovicianus) colonies, although they could be placed anywhere. Track Stations Our track station consisted of track- sensitive smoked kymograph paper covering a base of 4 mm thick plywood (0.6 m^) (Fig. 1). The kymograph paper was smoked in the field by burning kerosene-soaked cotton in- side an aluminum can which had a long, 1 cm wide slit cut in one side. Track impressions were preserved by spraying with quick drying shellac. Scent and bait attractants were placed on a stake in the center of each track station (Fig. 1). Scents consisted of a variety of com- mercial mustelid and other lures (i.e., weasels Mustela frenata, mink M. visoji, marten Mar- ies americana, fisher M. pennanti, badger Taxidea taxus, wolverine Gulo giilo, and black bear Ursus americanus. Baits consisted of an aged liver and sardine mixture. The acoustic attractant, a modified elec- tronic bird call producing a "chirping" sovmd, was used in association with track sta- tions. This call was developed by the U.S. Fish and Wildlife Service (electronic sche- matic available from the authors). The call was capable of fimctioning for a couple of weeks on a single 6-volt battery. The call was placed inside a camouflaged paint can (one- pint, 0.5 liters) with holes in the lid to emit the sounds. Location of track stations was determined in the field to minimize wind damage and to 'Department of Biology, Idaho State University, Pocatello, Idaho 8.3201 and Biota Research and Con.sulting, Inc., Box 2705, Jackson, Wyoming 83001. •Biota Research and Consulting, Inc., Box 2705, Jackson, Wyoming 83001. 438 July 1983 ■<'*!fjc^'' Clark, Campbell: Survey Techni9ue 439 ^Ca-'T*!*'--" ^"'Mii^f^^^^^ Fig. 1. Photograph of kymograph-smoked track station used by a prairie dog. Scent stake is coated with hver/sardines. maximize dispersal of odors and sounds. Each station was checked early each morning for nocturnal tracks and again in evenings for diurnal tracks for several consecutive days. Baits, scents, and kymograph paper were re- placed and soil resifted as needed. Linhart and Knowlton's (1975) track sta- tions consisted of a 1-m diameter circle of freshly sifted fine dirt 1 cm deep and scented and baited with specially prepared tabs or our lures. Results We accumulated 264 kymograph and 112 sifted dirt track stations days (1 track station day was 1 track station in operation for 24 hrs) on 24 different prairie dog colonies. A variety of animals left track impressions on both kinds of stations; they were: long-tailed weasels, skunks {Mephitis mephitis), kit foxes {Viilpes velox), coyotes {Canis latrans), badg- ers, mice {Peromyscus sp., Dipodomys sp., OnycJiomys sp.), ground squirrels (Sperrno- phihis sp.), prairie dogs, unidentified passe- rine birds, and numerous insects. As a substitute for the rarest carnivore on prairie dog towns, the black-footed ferret {Mustela nigripes), we presented kymograph track stations to Steppe ferrets (M. evers- manni) in laboratory conditions and they locomoted on them. Discussion The utility of the kymograph and sifted dirt track stations depended on the target species sampled, the quality and permanence of visitation record sought, and field condi- tions encountered. Kymograph stations re- quired about 4 minutes each to prepare, in a sheltered location, and about 5 minutes to set out. This type of station was most effective in low humidity and low to moderate winds— where stations might be effective 3-5 days before needing new smoked paper. Concerns that the kerosene odor and unfamiliar sub- strate texture might deter all wild species was not substantiated. The sifted dirt stations required less time and equipment to establish and were easier 440 Great Basin Naturalist Vol. 43, No. 3 to replace, but tracks were sometimes in- definite and harder to positively identify than with kymograph tracks. Photography was the only method to permanently record these tracks. No scats or urinations were found within the track station areas. We never observed a carnivore at a track station during nearly continuous nighttime surveys, even though tracks indicated their visitations. We think our track station method has utility in surveying site-specific areas, espe- cially if camera monitoring of animal visits as described by Dodge and Synder (1960) and Loveless et al. (1963) were added to our sys- tem. Our technique, as suggested by Clark (1977, 1978) and Clark and Campbell (1980), may offer a valuable addition to traditional survey methods for some rare species like the black-footed ferret. Since its efficacy has been demonstrated, needed now are follow- up quantitative experimental studies on the value of the kymograph track station. Acknowledgments We thank those organizations who sup- ported our field studies during which our track station method was tried— National Audubon Society, National Geographic So- ciety, Defenders of Wildlife, U.S. Bureau of Reclamation, Cleveland Cliffs Iron Co., and Mid-American Pipeline Co. Denise Casey and John Weaver provided critical advice on the manuscript. Wildlife Preservation Trust International and the New York Zoological Society Animal Research and Conservation Center provided support for manuscript preparation. Literature Cited Clark, T. W. 1977. A research proposal to develop sur- vey techniques and search for the black-footed ferret in Wyoming. Submitted to U.S. Fish and Wildlife Service— Office of Endangered Species, Washington, D.C., and Denver Regional Office. 23 pp. 1978. Current status of the black-footed ferret in Wyoming. J. Wildl. Mgt. 42:128-1.34. Clark, T. W., and T. M. Campbell. 1980. Suggested guidelines for black-footed ferret surveys. Avail- able from: Box 2705, Jackson, Wyoming 8.3001. 7 pp. Dodge, W. E., and P. Snyder. 1960. An automatic cam- era device for recording wildlife activity. J. Wildl. Mgt. 24:341-344. Justice, K. E. 1961. A new method for measiuing home ranges of small mammals. J. Mammal. 42:462-470. Llnhart, S. B., and F. F. Knowlton. 1975. Determin- ing relative abundance of coyotes by scent station lines. Wildl. Soc. Bull. 3:119-124. Loveless, C. M., J. D. Coffelt, D. E. Medin, and L. E. Yeager. 1963. A photoelectric-cell device for use in wildlife research. .'Vmer. Instit. Biol. Sci. Bull. 13:55-57. Mayer, W. V. 1957. A method for determining the ac- tivity of burrowing mammals. J. Mammal. .38.5.31. EVALUATION OF DRABA OLIGOSPERMA, D. PECTINIPILA, AND D. JUNIPERINA COMPLEX (CRUCIFERAE) Robert W. Lichvar' Abstract.— Since Dniha pectinipihi Rollins was described in 1953, it has been assigned to several different tax- onomic categories. It has be n recognized at the species and variety level and has also been placed in synonymy un- der D. oUgospemui Hook. Then Dorn (1978) described D. junipenna and contrasted it to D. pectinipila and D. oligos- perma. To clarify the status of these three taxa, the evaluation included field and herbarium observations and scanning electron microscope studies. A proposal to give Draba pectinipila Rol- lins protection under the Endangered Species Act prompted extensive field studies on this and two closely related taxa. Rollins (1953) described D. pectinipila from the alpine habi- tat of Clay Butte, Park County, Wyoming. It is now known from two other alpine loca- tions in British Columbia and Colorado. Since this taxon was described, it has been as- signed to several different taxonomic cate- gories. Before further action could be taken to protect the plants, the taxonomic status had to be reevaluated. Hitchcock (1964) gave Draba pectinipila varietal status imder D. oligospenna Hook. His only comment was that this variety was the only fairly distinct variant for the species. Mulligan (1972) placed it in synonomy under D. oligospenna without making field observa- tions. Then Dorn (1978) described plants from the low elevation areas near the Utah- Wyoming border, mentioned by Rollins (1953) under D. pectinipila, as D. juniperina. To clarify the taxonomy of this species com- plex, D. pectinipila and D. juniperina are compared to one another and to D. oligospenna. Methods This analysis of Draba oligosperma, D. pec- tinipila, and D. juniperina included field and herbarium observations and scanning elec- tron microscope (SEM) studies. Field obser- vations included habitat, plant aspects, and flowering dates. Herbarium specimens were studied for shape, and scape characteristics. The herbarium analysis was done at the Table 1. Character differences between the three Draba species. Character D. oligosperma D. pectinipila D. juniperina Habitat Scape pubescence Scape height Mature fruit pedicel length Petal color Silique tip Silique base Style length Flowering dates Basal leaf trichomes Valve trichomes Distribution exposed rocky slopes and ridges glabrous 1-4 (9) cm 0.1-0.5 (1.0) cm yellow mostly rounded rounded 0.1-1 mm May-July fine simple (90% + ) Western U.S. and Canada alpine slopes pubescent (4) 5-9 (11) cm 0.5-1.2 (1.4) cm yellow tapered rounded 0.3-0.8 mm July-August medium doubly pectinate (90% + ' British Columbia, Wyoming, Colorado pinyon-juniper woodlands pubescent (5) 7-15 cm 0.5-1.0 cm yellow tapered tapered 0.6-1.5 mm April-May coarse doubly pectinate (90% + ) Wyoming, Colorado, Utah 'Wyoming Natural Heritage Program, Nature Conservancy. 1603 Capitol .Avenue. Room 325, Cheyenne, Wyoming 82001. 441 442 Great Basin Naturalist Vol. 43, No. 3 Fig. 1. Leaf trichome of Draha pectinipila X200 (Lichvar 2066). Rocky Mountain Herbarium, Laramie, Wyoming; the Gray Herbarium, Cambridge, Massachusetts; and the University of Colo- rado Museum, Boulder, Colorado. The fol- lowing specimens were most pertinent to this study: At RM: Dawson, s.n.; McCoun, s.n.; Dorn, 897; Nelson, 1223; Lichvar, 2066; Johnston, 1424, 1442A, 1434; at GH: Rollins and Porter 51269; Williams, 476; at COLO: Beaman and Erbisch, 1208, 1276; Johnston, 1273, 2314; Calder, Savile, and Ferguson, 13780. Results and Discussion Comparison of field observations, herba- rium, and SEM analysis showed considerable differences between taxa in this complex. These taxa have overlapping ranges with areas of sympatry. Of the three taxa, Draha oligospemia has the widest range and is the most variable in habitat specificity. Draha oligospemia occurs from lower basin areas to high alpine ridges on either sandstone or limestone formations. Draha pectinipila is al- ways found on limestone in an alpine habitat, and D. jiiniperina is found on sandstone for- mations in association with a pinyon-juniper woodland at lower elevations. When D. oli- gospemia and D. pectinipila are syinpatric, D. pectinipila is readily identifiable in the field by its taller, more erect stature and the high percentage of pectinate hairs on the fruit valves. Near the Flaming Gorge area of Wyoming, Utah, and Colorado, D. juniperina is not found in the same habitat as D. oligo- spemia. Draha oligospemia occurs on sand- stone or gravelly outwash ridges and out- crops, but D. juniperina is almost always found in association with pinyon-juniper woodlands or adjacent sagebrush that overlies a sandstone formation. Certain characteristics separate Draha pec- tinipila and D. juniperina from D. oligos- pemia (Table 1). The correlation of charac- ters that are similar between D. pectinipila July 1983 Lichvar: Draba 443 Fig. 2. Leaf trichonie of Draba oligosperma X200 {Lichvar 1981). and D. jiiniperina are the tapered fruit tips with doubly pectinate hairs on the valves of the fruits and the taller scapes that are pu- bescent. Draba oligosperma has rounded fruit tips with mostly simple hairs and shorter scapes that are glabrous. To distinguish be- tween the two taxa with a more limited dis- tribution, D. pectinipila and D. jiiniperina, habitat, style length, and leaf trichomes may be used. Draba pectinipila is an alpine taxon found on limestone and has tapered fruits above witli styles 0.3-0.8 mm long, but D. jii- niperina is found at lower elevations in the basins on sandstone in a pinyon-juniper woodland and has tapered fruits above and below, and styles 0.6-1.5 mm long. Rollins (1953) noted that the lower eleva- tion taxon, Draba juniperina, had not only ta- pered fruits above and below but also coarse hairs on the basal leaves. Scanning electron micrographs (Figs. 1, 2, 3) show that D. ju- niperina (Fig. 3) has doubly pectinate leaf hairs twice the diameter of D. pectinipila (Fig. 1). Draba oligosperrna (Fig. 2) is inter- mediate in leaf trichome diameter. Recognition of each of these species can be justified at the species level for three reasons. First, specimens or plants of each of these taxa can consistently be separated in either the field or herbarium. Second, when any combination of these species occurs sympa- trically in the field they are easily and con- sistently identifiable. Finally, based upon tri- chome characters, those used to distinguish these species are consistent with those used throughout the genus of Draba. Key to the Species 1. Scapes glabrous; 1-4 (9) cm long; fruits with 90 percent or more simple hairs ... D. oligosperma 444 Great Basin Naturalist Vol. 43, No. 3 Fig. 3. Leaf trichome of Draba jtiniperina X200 {Lichvar 2821). — Scapes pubescent, 4-12 cm long; fruits with 90 percent or more doubly pectinate hairs 2 2(1). Plants alpine with fine hairs on basal leaves; siliques tapered above, rounded below; styles 0.3-0.8 mm long D. pectinipila — Plants of pinyon-juniper woodlands with coarse hairs on basal leaves; siliques tapered above and below; styles 0.6-1.5 mm long D. juniperina Acknowledgments Thanks are extended to Robert Dorn for reviewing the manuscript, to Robert Bow- men for doing the SEM work, and to the curators of the collections mentioned above for allowing me to study specimens in their herbaria. Literature Cited Dorn, R. D. 1978. A new species of Draba (Cruciferae) from Wyoming and Utali. Madrono 25:101-103. Hitchcock, C. L., A. Cronquist, M. Ownby, and J. W. Thompson. 1964. Vascnlar plants of the Pacific Northwest. Part 2. Univ. of Washington Press, Seattle. 496 pp. Mulligan, G. A. 1972. Cytotaxonomic studies of Draba species in Canada and Alaska: D. oligosperma and D. incerta. Canadian J. Bot. 50(8): 1763-1766. Rollins. R. C. 1953. Draba on Clay Butte, Wyoming. Rhodora 55:229-235. PRESENCE OF MAXILLARY CANINE TEETH IN MULE DEER IN UTAH Jordan C. Pederson' Abstract.— A hunter-harvested adult female mule deer {Ock>coiletis hemionus) was found to have canine teeth present along both rows of teeth of the maxillae. The reported occurrence of erupted canine teeth in mule deer {Odocoileus hemionus) has been observed in CaHfornia (Nordquist 1941), Utah (Robinette 1958), and Colorado (Robin- ette et al. 1977). The Utah incident occurred in a three-year-old male having a single erupted canine in the right maxilla (Robin- ette 1958). On 27 November 1982, a hunter-harvested 4-year-old female mule deer was checked and collected by me. She was killed near In- dianola in the Little Clear Creek drainage in Utah County. The presence of upper max- illary canine teeth on both sides was observed when the jaw was cut for field aging (Fig. 1). The head was removed from the carcass and, after cleaning and photographing, donated to the Brigham Young University Life Science Museum, Provo, Utah, Catalog no. 7225. The following skull measurements were taken: total length 291 mm, condylobasal length 272 mm, zygomatic breadth 128 mm. Both maxillary canines are found posterior to the nasomaxillary suture identical to the posi- tion of canines found in elk {Cervus cana- densis). Further measurements showed each Fig. 1. Palatal view of a mule deer skull showing canine teeth on maxillae. 'Utah Division of Wildlife Resources, 1115 North Main, Springville, Utah 84663. 445 446 Great Basin Naturalist Vol. 43, No. 3 canine is positioned 46 mm posterior of the anterior point of the premaxilla and 43 mm anterior of the premolar. Measurements of the left and right canine, respectively, are: length of outside (labial), exposed part, 9.2, 9.9 mm; anteroposterior diameter of crown, 4.9, 5.1 mm; labial lingual diameter of crown, 3.8, 4.0 mm. Acknowledgments Thanks are extended to Dr. Jerran T. Flin- ders, Dr. Kent M. Van De Graaff, Dennis L. Shirley, and Laura John for their help in the preparation of this manuscript. Literature Cited NoRDQuiST, G. 1941. Elk teeth in California deer. Cali- fornia Fish and Game 27:39-40. RoBiNETTE, W. L. 1958. Unusual dentition in mule deer. J. Mammal. .39:1,56-157. RoBiNETTE, W. L., N. V. Hancock, and D. A. Jones. 1977. The Oak Creek mule deer herd in Utah. Publ. No. 77-15. Utah Wildlife Resources, Salt Lake City. 148 pp. COMPARATIVE SUCCESSIONAL ROLES OF TREMBLING ASPEN AND LODGEPOLE PINE IN THE SOUTHERN ROCKY MOUNTAINS Albert J. Parker' and Kathleen C. Parker' Abstract.— A review of the ecological distribution and siiccessional roles of lodgepole pine and trembling aspen in the Southern Rocky Mountains suggests that the two species have different strategies for occupying disturbed sites. Lodgepole pine's easily dispersed seeds and faster growth from seed in unsuppressed conditions allows it to col- onize severe bums, even from remote seed sources. Aspen appears to compensate for ineffective development from seed by vegetative reproduction from durable root stocks, which promotes geographic persi.stence. Such persistence is achieved by the maintenance of a forest structure conducive to light surface fires, which stimulate suckering and retard conifer invasion, and by the accumulation of soil organic matter, which improves site nutrient retention and water availabilitv. Empirical studies of the dynamics of trembling aspen {Popiihis tremiiloides) and lodgepole pine (Pinus contorta ssp. latifolia) forests show that both tree species commonly colonize open sites following disturbance (Clements 1910, Ives 1941, Stahelin 1943, Marr 1961, Langenheim 1962). In portions of the upper montane and subalpine zone (2,400-3,000 m) of the Southern Rocky Mountain region, the geographic and habitat ranges of these two important colonizers overlap, so that either species (or both) might be encountered on a disturbed site. Within this zone of cooccurrence, neither the site preferences of nor the successional relation- ship between these two species is satisfac- torily detailed (Marr 1961, Peet 1981). Re- garding habitat range, early workers thought that aspen more frequently occurred on mes- ic sites, and lodgepole more commonly occu- pied drier settings (Bates 1924, Daubenmire 1943). More recently, Marr (1961) and Peet (1978) have questioned the simplicity of this arrangement. Peet (1978) asserts that both species possess a comparable ecological opti- mum on mesic sites in the lower/ middle sub- alpine zone, as evidenced by the distribution of aspen in mountainous regions where lodgepole is absent. He observed that, in re- gions where both species occur, lodgepole is a better competitor than aspen on prime sites and therefore tends to preempt aspen from optimal settings. Aspen maintains popu- lations in this region of cooccurrence by pos- sessing a broader environmental tolerance range, often being restricted to a variety of both wetter and drier sites at higher and lower elevations than lodgepole. The successional relationship of the two species in this region of cooccurrence is com- plex (Moir 1969, Reed 1971, 1976, Whipple and Dix 1979, Peet 1981). Differences in their respective patterns of colonization are likely related to a number of factors, chief among which is the fundamental dissimilarity in their reproductive strategies. Lodgepole is a prolific seeder, depending on widespread wind dispersal of its light seeds to facilitate invasion of disturbed sites. Aspen, although it is capable of reproduction by seed, more of- ten reproduces by vegetative suckering. Marr (1961) observed that aspen roots often sur- vive fire, thus providing a stock for vegeta- tive propagation on burned sites. Further- more, both Marr (1961) and Peet (1981) noted that aspen is often found in the under- story of a variety of different forest covers, including dense, mature conifer forests. Thus, aspen is able to maintain a suppressed but vi- able population on a site through long peri- ods of time, and is capable of colonizing burned sites by the release of the persi.stent rock stock. Horton and Hopkins (1965), in an examination of fire ecology in aspen groves, found that light burns (i.e., low temperatures) stimulate aspen suckering (probably through 'Department of Geography, University of Georgia, Athens, Georgia 30602. 447 448 Great Basin Naturalist Vol. 43, No. 3 both the reduction of competition with the thick ground layer and mobilization of the nutrient supply tied up in the ground layer vegetation), but heavy burns (i.e., high tem- peratures) inhibit aspen suckering (presum- ably through damage to perennating organs in the root stock). Heavy burns are likely to enhance the establishment of lodgepole pine on disturbed sites, because they create a min- eral seedbed and eliminate much of the ground layer vegetation that might normally inhibit development from seed of lodgepole pine. Hence, postburn colonization of sites by either aspen or lodgepole in their zone of cooccurrence is influenced by their respec- tive reproductive modes and ecological toler- ances of environmental factors, by burn in- tensity, and by a chance element (Marr 1961) associated with the probability/proximity of a lodgepole seed source or an aspen rootstock. The present study presents a review and interpretation of both the habitat ranges and successional relationship of lodgepole pine and trembling aspen in the Colorado Front Range. This discussion is accompanied by a data set examining tlie age/size structure and community characteristics of an abrupt aspen /lodgepole ecotone on the south flank of Bierstadt Moraine in Rocky Moimtain Na- tional Park, Colorado. Bierstadt Moraine is a lateral moraine of approximately 200 m relief, deposited by al- pine glaciers during the late Wisconsin gla- cial maximum (Pinedale stage, Richmond 1960). Bierstadt Moraine trends slightly north of east, extending for approximately 6 km along the northern margin of the Glacier Creek valley from Bear Lake to near Glacier Basin Campgroimd. The dominant particle size in the till is sand (exceeding 70 percent of the total fine earth fraction); soils devel- oped under both aspen and lodgepole forests are immature (typic Cryorthents), although there are distinct differences in the A-horizon under each cover type. A transect down the south-facing slope of Bierstadt Moraine from top to bottom reveals the following sequence of plant communities: lodgepole pine forest on the gently rolling upland, sagebrush {Arte- misia ssp.) scrub on the steep upper slope, as- pen forest on the middle slope, and lodgepole pine forest on the lower slope and through- out the adjacent valley bottom. The ecotone studied is between the aspen forest and the lower lodgepole forest, at an elevation of 2,700 m. The study area is located within a much larger region (perhaps 10 km^) which was burned by the Bear Lake fire of 1900 (Peet 1981). None of the trees cored on the study site is older than this burn, so that the mod- ern forest is representative of postburn recov- ery dating three-quarters of a century from this extensive fire. Methods Seven 4 X 60 m belt transects were placed with their long-axis oriented normal to eleva- tion contours and the aspen /lodgepole eco- tone. Each transect was subdivided into six 4 X 10 m quadrats and placed so that three of these quadrats were under aspen cover and three were under lodgepole cover. Although precise location of the "boundary" between types is subjective, in this case abrupt differ ences in both litter type and ground cover were used to determine the midpoint of the belt transect. Belt transects were spaced along the flank of the moraine at intervals of 60 m. In each quadrat, all living trees (stem DBH > 6.25 cm) were identified to species and their diameter recorded. All saplings (0 < stem DBH < 6.25 cm) and seedlings (stems less than breast height) in each quad- rat were counted by species. All standing dead stems in each quadrat were counted by species. Along the central long axis of each belt transect, the coverage of all herbs and shrubs was determined in 10-m intervals by the line-intercept method (Canfield 1941). In two of the seven belt transects a radial core was extracted from each tree (at 0.3 m height) with an increment borer, and the tree's age determined. Measurements of slope aspect, steepness, and configuration were taken for each 4 X 10 m quadrat. In two of the belt transects, the type and depth of litter was measured, using a point-frame, at the center of each 4 X 10 m quadrat. The point- frame was 1 m wide, with a 5 cm recording interval; hence, there are 21 litter measure- ments per quadrat. In addition, two soil pits were dug, one under each cover type, and the soil profiles were described. July 1983 Parker, Parker: Succession 449 Table 1. Vegetation data by forest type. Lodgepole cover .odgepole Aspen Aspen coyer Characteristic L Lodgepole Aspen Tree density (stems-ha"l) 2420 430 100 2800 Stand basal area dm2-ha-l) 2280 170 140 1910 Mean diameter (cm'stem"^) 15.5 10.2 19.5 13.2 Sapling density (sfha-i) 290 190 80 540 Seedling density (sfha-l) 10 210 20 3870 Standing dead density (st-ha-i) 250 920 10 2510 Results The aspen forest on the south flank of Bierstadt Moraine is on a slightly steeper slope than the lodgepole forest (15° vs. 12°), fingering to lower elevations along local con- vexities of the slope face. The principal difference in the soil profiles under each cover type is the presence of a dark brown (10 YR 3/2) surface mineral hori- zon, approximately 10 cm thick, under the aspen forest. This darker topsoil, which is ab- sent under lodgepole, is indicative of higher organic matter content associated with humi- fication of the deciduous aspen leaves and lit- terfall from the thick ground layer vegetation (cf Hoff 1957, Tew 1968). Hydrometer analy- sis (Bouyoucos 1962) of soil material col- lected from varying depths in both profiles revealed no meaningful differences in texture with depth or between profiles, all samples being coarse sandy loams. This texture, which normally would permit only low water reten- tion and rapid leaching of cations, imparts in- creased significance to the organic buildup in the A-horizon under aspen (Morgan 1969), in that organic colloids increase the water re- tention capacity and the cation (or nutrient) exchange capacity of the aspen soil relative to the lodgepole soil. The ground layer of the lodgepole forest was poorly developed, with 3 percent ground coverage divided evenly between herbs and woody shrubs. The aspen forest ground layer was well developed (58 percent ground cov- er) and almost exclusively herbaceous. Such differences between aspen and conifer ground layer development have been re- ported elsewhere (Hoff 1957, Marr 1961, Peet 1981) and apparently result from a com- bination of the high acidity of conifer litter (Daubenmire 1953) and increased light pene- tration to the forest floor under aspen (par- ticularly in the spring prior to aspen leaf flush). Under the lodgepole cover, tree density was 2,850 stems-ha-^; lodgepole pine ac- counted for 84.9 percent of this total (Table 1). Under the aspen cover, tree density was 2,900 stems'ha'; aspen accounted for 96.6 percent of this total. Similarly, under lodge- pole cover, stand basal area .was 2,450 dm^-ha' (93.1 percent lodgepole); under as- pen cover, stand basal area was 2,050dm^ha-i (93.2 percent aspen). Little evidence of invasion by more toler- ant tree species over the last 75 years exists. The only other tree species encountered were Table 2. Cross transect patterns of mean tree basal area and understory representation by species. Cover/Quadrat/Species Mean tree Basal area (dm^'stem"^) Number of saplings and seedlings Number of standing dead . Lodgepole Aspen Lodgepole _^ Lodgepole cover Aspen ^Lodgepole — Ecotone- Aspen ■ Lodgepole Aspen Aspen -Lodgepole cover Aspen ^Lodgepole Aspen 1.74 0.47 1.68 0.37 2.34 0.89 10 9 5 10 10 15 9 12 56 2.52 1.21 1.47 1.45 4.91 1.47 3 121 2 125 4 124 0 92 0 60 1 59 450 Great Basin Naturalist Vol. 43, No. 3 Lodgepole Forest Cover Lodgepole Pine UJ CD Aspen 20 10 fc IV d cvj id f^' O CO lo K d ro C£)CD 10 Aspen Forest Cover Lodgepole Pine 72—1—^2 I r / J / /P^ 1^" d c\J in h-' d oJ in h-^ ' ^.— ■— ■— CJOJC\JCM ro I I I I I I I I i^'dcJidi^doJid '"•—'"'" CJ CVJ CVJ DIAMETER -CLASS (cm) Fig. 1. Diameter-class distribution by forest cover. Histograms of stem number by diameter class for both lodge- pole pine and aspen under each forest cover are depicted (diameter class interval = 2.54 cm). The number of stems in the smallest diameter class (6.3-7.5 cm) has been doubled to adjust for its half-interval width. willow {Salix spp.), with two saplings present in a quadrat adjacent to the valley bottom, and Douglas-fir {Psendotsiiga menziesii), with a single seedling found under aspen. Cross transect trends in the number and size of aspen and lodgepole (Table 2) demon- strate that lodgepole pine is infrequently en- countered under aspen cover, but, where July 1983 Parker, Parker: Succession 451 Lodgepole Pine Aspen CO < 15 > 10 ^ 5 I tiii < 9 > or UJ CO w I I ' I I I I I I ^ OinOLOQi^omo w I I I I I I I I I ^ oioOinoioOinO AGE-CLASS (yr.) Fig. 2. Age-class distribution of lodgepole pine and aspen in cored transects. Histograms of the stem number by five-year age class intervals are shown for each species. present, trees are generally large. Conversely, aspen is commonly encountered beneath lodgepole, although these trees are generally small. This pattern is more emphatically ex- pressed in the understory; aspen saplings, seedlings, and standing dead stems (chiefly aborted suckers) are much more common un- der lodgepole cover than lodgepole is under aspen cover. The high number of aspen seed- lings and standing dead stems under lodge- pole cover is indicative of a successional strategy that relies on maintenance and grad- ual spread of the aspen root stock into the understory of adjacent conifer forests. The trend in mean tree diameter and in the num- ber of both understory and dead stems across the transect clearly demonstrates the progres- sive spread of aspen across the ecotone (Table 2). The mean tree diameter of aspen and the total number of understory and dead aspen stems steadily decline away from the aspen grove. The collection of 72 lodgepole pine and 51 aspen tree cores was used to correlate age with stem diameter. Both species exhibited a comparable age-diameter relationship, the coefficient of variation of age (lodgepole = 0.22, aspen = 0.25) being less than the coefficient of variation of diameter (lodge- pole = 0.38, aspen = 0.41) in each case. Furthermore, age was significantly correlated with diameter for each species (for aspen r = 0.654, p < 0.001; for lodgepole r = 0.665, p < 0.001). Lodgepole pine displayed a some- what stronger tendency toward even- agedness than aspen, although both species exhibited unbroken representation in age ranges between 30 and 75 years. Fifty per- cent of the lodgepole stems were in the 60-to 75-year age range, suggesting colonization 452 Great Basin Naturalist Vol. 43, No. 3 (1) Aspen Cover Trees Soil/Ground layer Annual nutrient- rich litter fall Open canopy branctiing pattern Nonflammable plant parts Reproduction by suckering Pronounced biocycling (good nutrient reserve) Thick herbaceous layer Low-temperature surface fires Fig. 3. Diagrammatic representation of the interactions among the dominant tree species, soil, ground layer vege- tation, and fire characteristics under aspen and lodgepole pine forest cover. immediately following the Bear Lake burn, and the maximum concentration of aspen stems (42 percent) occurred in the 45- to 60- year range (Fig. 1). The unsuppressed diame- ter growth rate of each species was deter- mined using the 10 largest lodgepole and as- pen trees cored. Our data reveal that following successful establishment unsup- pressed lodgepole pine grows more rapidly than aspen (0.37 cm yr' vs. 0.32 cm yr') on the study site. It should be cautioned that the period of establishment is generally several years longer for lodgepole pine produced from seed, which may require from 3 to 20 years to reach 0.2 m height (Romme and Knight 1981), than for aspen suckers, which may reach 3 m or greater height in 6 to 8 years (Jones and Trujillo 1975). The composite diameter class diagram (Fig. 2), which is based on all seven belt transects and presented by cover type, does not show a tendency for concentration of lodgepole pine in larger size-classes even though many lodgepole trees are relatively old, suggesting that older lodgepole stems may persist as suppressed individuals follow- ing postburn colonization for a lengthy peri- od. Examination of these diameter-class dia- grams reiterates that aspen is a fairly common understory tree beneath lodgepole forests, but only a few generally large-sized lodgepole individuals are scattered through- out the aspen canopy. Discussion The soil profile and age structure differ- ences between aspen and lodgepole pine stands suggest that each species, where domi- nant, reinforces a distinct group of vegeta- tion-soil-fire interactions (Fig. 3). Further- more, the persistence of these cover types appears to be more closely tied to stand his- tory than to direct environmental gradients. Under aspen cover, the deciduous, nutri- ent-rich foliage of aspen (Daubenmire 1953) and the dense herbaceous understory com- bine to enhance nutrient cycling and humifi- July 1983 Parker, Parker: Succession 453 (2) Lodgepole Cover Trees Soil/Ground layer Nutrient-poor litter fall Dense canopy branching pattern Flammable plant parts Reproduction by seeds a) requires high light regime and mineral seedbed. b) serotinous cones. c) widely dispersed seeds Slow biocycling (low nutrient reserve) sparse ground cover, shrub dominated High-temperature crown fires Fig. 3 continued. cation, resulting in an increased cation ex- change capacity and nutrient concentration in surface mineral horizons (Hoff 1957). The increased soil water retention conferred by the humus accumulation acts in concert with the ground cover (which buffers soil surface temperature and decreases windspeed, thus reducing evaporation) to increase the avail- ability and effective use of soil moisture. Hence, aspen maintains a broad habitat range by direct enhancement of soil nutrient and moisture status (Lutz and Chandler 1946). On the contrary, the acidic lodgepole pine nee- dles promote leaching, and the paucity of ground cover under lodgepole pine limits biocycling of nutrients; consequently, soils under lodgepole pine are often impoverished relative to adjacent aspen substrates (Hoff 1957). In addition, the mechanical resistance of pine needles to decomposition results in a decrease in himiification and the buildup of pine needle litter. Fire plays a prominent role in the perpetu- ation of discrete populations of both aspen and lodgepole pine (Fig. 3). Aspen domin- ance is maintained on a site through stimu- lation of vegetative propagation following low-temperature surface fires, presumably through reduction of apical dominance (Dan- iel 1980). The buildup of surface fuels by the thick herbaceous layer, the mesicness of the ground layer, and the relative openness of the aspen canopy favor light-burning surface over crown fires (Horton and Hopkins 1965). Because of its suckering habit, aspen can sus tain and is capable of slowly expanding local populations vegetatively into adjacent favor- able sites. The ability of aspen to sucker in relatively dense shade facilitates this spread. Continued aspen dominance on a site re- quires the perpetuation of a surface fire re- gime that releases advance regeneration (Marks 1974, Oliver 1981) and stimulates vig- orous reestablishment of aspen suckers. In the absence of fire, eventual ascendance of more tolerant conifer species often does not pre- clude the persistence of the aspen root stock in a suppressed condition (Marr 1961). De- pendence on the maintenance of a "parental" aspen root stock is necessary to offset the competitive superiority of lodgepole pine (and other conifers) when both species are es- 454 Great Basin Naturalist Vol. 43, No. 3 tablished from seed, and favors persistent re- colonization by aspen following light surface fires. In lodgepole forests a denser canopy branching network, greater leaf area index, and a high resin production combine with a paucity of imdergrowth to favor hot crown fires. By creating high surface light levels and mineralizing the litter layer, crown fires cre- ate a favorable seedbed for lodgepole pine establishment and trigger a wave of lodge- pole pine regeneration that manifests itself in a tendency toward even-agedness in Rocky Mountain lodgepole forests. This contagious postbum colonization pattern is facilitated by lodgepole pine's lightweight, easily wind- dispersed seeds and rapid growth rate follow- ing seedling establishment on disturbed sites (allowing colonization from a remote seed source). Furthermore, colonization of severe- ly burned sites by lodgepole pine is rein- forced locally by partial core serotiny (Po- wells 1965). In addition to favoring lodgepole establishment, severe burns inhibit aspen suckering, because most suckers develop from roots which are within 5 cm of the soil surface, and hence are easily killed in a hot fire (Daniel 1980). The persistence of both aspen and lodge- pole pine populations on sites with little evi- dence of successional alteration suggests that both species can be expected to maintain do- minance for extended periods, in accordance with Egler's (1954) view of vegetation devel- opment. Only with prolonged fire exclusion are stands likely to be invaded and replaced by more tolerant conifers. Moreover, changes in dominance on a site appear to be related to the character of initiating disturbances (Henry and Swan 1974, Anderson and Holte 1981) and the differential reproductive habits of each species. A low-temperature surface fire regime favors the maintenance and grad- ual spread of aspen dominance by aggressive suckering. Stand-destroying crown fires open sites to rapid colonization by lodgepole pine, and repeated crown fires reinforce lodgepole pine dominance. In summary, both trembling aspen and lodgepole pine are successful colonizer spe- cies in the southern Rocky Mountain region, although they accomplish colonization and persistence in different ways. Aspen perpetu- ates itself through time on suboptimal sites by its suckering habit that rapidly recolonizes lightly burned sites, by promotion of a favor- able surface fire regime, and by enhancement of site quality through improved soil water and nutrient retention capacity. Lodgepole pine, which develops more rapidly from seed than aspen and has easily dispersed seeds, is able to preempt aspen on optimal sites, and exhibits a relatively even-aged, contagious colonization pattern following stand- destroying crown fires. Literature Cited Anderson, J. E., and K. E. Holte. 1981. Vegetation de- velopment over 25 years vvithont grazing on sage- brush-dominated rangeland in southeastern Idaho. J. Range Mgt. 34:25-29. B.\TEs, C. G. 1924. Forest types in the central Rocky Mountains as affected bv climate and soil. USDA Bull. 1233. 152 pp. BouYoucos, G. J. 1962. Hydrometer method improved for making particle size analysis of soils. Agron. J. 54:464-465. Canfield, R. H. 1941. Application of the line intercept method in sampling range vegetation. J. For. 39:388-.394. Clements, F. E. 1910. The life history of lodgepole burn forests. USDA Forest Service Bull. 79:7-56. Daniel, T. W. 1980. The middle and southern Rocky Mountain region. Pages 277-.340 in J. W. Barrett, ed.. Regional silviculture of the United States. Wiley-Interscience, New York. Daubenmire, R. F. 1943. Vegetation zonation in the Rocky Mountains. Bot. Rev. 9:325-393. 1953. Nutrient content of leaf litter of trees in the northern Rocky Mountains. Ecology 34:786-793. Egler, F. E. 1954. Vegetation science concepts I. Initial floristic composition, a factor in old-field vegeta- tion development. Vegetatio 14:412-417. Fowells, H. a. 1965. Silvics of forest trees in the United States. USDA Agr. Handbook 271. 762 pp. Henry, J. D., and J. M. A. Swan. 1974. Reconstnict- uring forest history from live and dead plant ma- terial—an approach to the study of forest succes- sion in southwest New Hampshire. Ecology 55:772-783. HoFF, C. C. 1957. A comparison of soil, climate, and biota of conifer and aspen communities in the central Rocky Mountains. Amer. Midi. Nat. 58:115-140. HoRTON, K. W., AND E. J. HoPKiNs. 1965. Influence of fire on aspen suckering. Canadian Dept. Forestry Publ. 1095:1-19. Ives, R. L. 1941. Forest replacement rates in the Colo- rado headwaters area. Bull. Torrev Bot. Club. 68:407-408. Jones, J. R., and D. P. Trujillo. 1975. Development of some young aspen stands in Arizona. USDA For- est Service Research Paper RM-151. 11 pp. July 1983 Parker, Parker: Succession 455 Langenheim, J. H. 1962. Vegetation and environmental patterns in the Crested Butte area, Gunnison County, Colorado. Ecol. Monogr. 32:249-285. LuTZ, H. J., AND R. F. Chandler. 1946. Forest soils. Wiley & Sons, New York. Marks, P. L. 1974. The role of pin cherry (Pruniis pen- stjlvanica L.) in the maintenance of stability in northern hardwood ecosystems. Ecol. Monogr. 44:7.3-88. Marr, J. W. 1961. Ecosystems of the east slope of the Front Range in Colorado. Univ. Colo. Stud., Biol. 8. 134 pp. MoiR, W. H. 1969. The lodgepole pine zone in Colo- rado. Amer. Midi. Nat. 81:87-98. Morgan, M. D. 1969. Ecologv of aspen in Gunnison County, Colorado. Amer. Midi. Nat. 82:204-228. Oliver, C. D. 1981. Forest development in North Amer- ica following major disturbances. Forest Ecol. Mgt. 3:153-168. Peet, R. K. 1978. Latitudinal variation in southern Rocky Mountain forests. J. Biogeogr. 5:275-289. 1981. Forest vegetation of the Colorado Front Range: composition and dvnaniics. Vegetatio 45:3-75. Reed, R. M. 1971. Aspen forests of the Wind River Mountains, Wyoming. Amer. Midi. Nat. 86:327-343. 1976. Coniferous forest habitat types of the Wind River Mountains, Wyoming. Amer. Midi. Nat. 95:159-173. Richmond, G. M. 1960. Glaciation of the east slope of Rocky Mountain National Park, Colorado. Geol. Soc. Amer. Bull. 71:1.371-1382. Rom.me, W. H., and D. H. Knight. 1981. Fire frequency and subalpine forest succession along a topo- graphic gradient in Wyoming. Ecology 62:319-326. Stahelin, R. 1943. Factors influencing the natural re- stocking of high altitude burns by coniferous trees in the central Rocky Mountains. Ecology 24:19-30. Tew, R. K. 1968. Properties of soil under aspen and herb-shrub cover. USDA Forest Service Res. Note INT-78. 4 pp. Whipple, S. A., and R. L. Dix. 1979. Age structure and successional dynamics of a Colorado subalpine forest. Amer. Midi. Nat. 101:142-158. DIFFERENTIAL UTILIZATION OF BAT BOXES BY HOUSE WRENS {TROGLODYTES AEDON) Hal L. Black' Abstract.— Pine boxes nailed to trees in four habitat types in southeastern Utah as roosting sites for bats proved ineffective for bats but were utilized bv house wrens in all but one habitat. Boxes were most often utilized in the as- pen habitat. Hypotheses to account for differential utilization are presented. In late Fall of 1977, 35 rough-sawn, pine- wood, day-roosting boxes for bats, con- structed after the design of Stebbings (Quar- terly J. Deven Trust for Nature 6:114-119, 1974) were placed in each of four habitat types on the Abajo Mountains of southeastern Utah in the hope of attracting several mon- tane species of vespertilionid bats. The aspen forest habitat was within a watershed rela- tively ungrazed by livestock and had a rich understory of low-growing shrubs, forbs, and grasses. The oak-pine habitat, which is grazed annually by cattle, had, relative to the aspen, a poor understory and more hetero- geneous structure. The cottonwood habitat was within a rather broad stream bed and was not grazed by livestock. The spruce-fir forest was also ungrazed and consisted of large mature trees, which formed a fairly closed canopy with essentially no understory vegetation. Permanent water was available within each habitat. Boxes were attached to tree tnmks with nails at heights of 3-4 m and arranged in a zig-zag fashion within each habitat at 12-15 m intervals. Unlike tradi- tional bird boxes, these had a slit entrance on the ventral side that measured about 25 X 175 mm. As roosting sites for insectivorous bats the boxes were disappointing, but as nesting sites for house wrens {Troglodytes aedon) the boxes were immediately successful. Table 1 in- dicates by habitat and year the number of boxes from which young wrens were fledged. Hypotheses to explain the differential utiliza- tion of these boxes include: (1) differences in productivity of the understory vegetation and, therefore, insect biomass on which wrens feed and (2) differences in the avail- ability of natural cavities for nesting. The as- pen habitat appears to have the most produc- tive understory and the most homogeneous forest structure. The importance of nest boxes in the aspen habitat suggests that re- moval of understory vegetation by grazing or through competitive processes may have a rather severe effect on the abundance and distribution of foliage gleaning species like the house wren. These data suggest that the addition of nest boxes to habitats may selecti- vely enhance population densities. Table 1. Summary of patterns of utilization of bat-roosting boxes by house wrens. The open number under each habitat indicates the number of boxes out of .35 that were used. The number in parenthesis represents percent of total. Habitat Type Year Aspen (El. 2700m) Oak-Pine (El. 2400m) Cottonwood (El. 2400m) Spruce-Fir (El. 3200m) 1977 1978 10 (29) 12 (34) 2(6) 2(6) 0 1(3) 0 0 'Department of Multicultural Education and Department of Zoology, Brigham Young University, Provo, L'tah 84602. 456 PLANT AND SOIL RELATIONSHIPS IN TWO HYDROTHERMALLY ALTERED AREAS OF THE GREAT BASIN N. M. Milton' and T. L. Purely' Abstract.— In two areas of hydrothermally altered rocks in the Great Basin, the native vegetation differs in com- position and areal cover from imaltered to altered sites on the same geologic formations. Analysis suggests that phys- ical rather than chemical factors may be the cause of the vegetation differences, especially permeability of bedrock, depth and texture of soils, and, possibly, amounts and types of clay minerals present. These characteristics influence the ability of soils to absorb and retain water. In the East Tintic Moimtains, Utali, the soils from argillized or mixed argillized and silicified parent materials have more characteristics associated with dryness and support sparser vegetation and more species especially adapted to dry conditions than do soils from unaltered or silicified parent materials. In Battle Mountain, Nevada, unaltered areas have greater vegetation cover and have soil depth and texture that are more favorable for plants than do altered areas. Soil pH is higher in altered areas than in unaltered areas. Where geology is obscured by vegetation, as in the humid regions of the world, vegeta- tion can be used as a clue to the underlying rocks and minerals. Changes in the vegeta- tion along zones of mineralization have been recognized since ancient times and have been well docimiented in several recent reviews (Malyuga 1964, Rommel 1968, Nesvetailova 1970, Cannon 1971, Brooks 1972). The rea- sons for the vegetation differences should be found in the physical and chemical proper- ties of the soils that develop on hydro- thermally altered, locally mineralized rocks which differ from those of soils that develop on unaltered rocks. Two previous workers have studied the vegetation in hydrothermally altered areas in the Great Basin of the western United States. Billings (1950) concluded from greenhouse experiments that acid conditions and defi- ciencies in nitrogen and phosphorus were re- sponsible for the lack of sagebrush on altered sites in the Virginia Range near Virginia City, Nevada. Salisbury (1954, 1964) per- formed similar experiments at Big Rock Can- dy Mountain, Utah, and concluded that nutri- ents in acid soils (pH 3.3) were chemically bound by iron and aluminum and therefore were unavailable to plants. These studies de- scribe extreme states, in which the soil pH is very low and the vegetation is drastically dif- ferent from that on nearby unaltered rocks. In many other hydrothermally altered areas, altered soils have pH ranges comparable with those of unaltered soils, and vegetation differ- ences are more subtle. The most striking* vegetation differences in the study areas described in this report are on sites of intermediate alteration intensity, whereas the unaltered and the most intensely altered sites have more similar vegetation. Since intensely altered sites have been sub- jected to more leaching than intermediate sites, nutrient deficiencies and toxicities would seem to be ruled out as likely causes of the vegetation differences. The most common limiting factor of plant growth in arid and semiarid areas is water. Infiltration and water retention are closely related to soil depth, soil texture, i.e., the distribution of different-sized particles in the soil, and the type of clay min- erals in the soil (Black 1968, Foth and Turk 1972). For the present study, the hypothesis of drier soil conditions in areas of inter- mediate alteration intensity was tested by ex- amining soil characteristics on altered and imaltered sites and by comparing the vegeta- tion distribution with that found by other workers in similar terrain. Study Areas Two study areas in Utah and Nevada were chosen to coincide with areas of ongoing re- 'Mailstop927, U.S. Geological Survey, Reston, Virginia 22092. 457 458 Great Basin Naturalist Vol. 43, No. 3 search in the use of remote sensing tech- niques to identify hydrothermally altered rocks. The Utah study area has been used as a test site for mapping hydrothermally altered rocks from high altitude and satellite imagery (Rowan and Abrams 1978a, b, Rowan and Kahle 1982). Krohn et al. (1978) used Landsat imagery to detect hydrothermally altered rocks at the Nevada study area. Both areas were found to be at the limit beyond which limonitic hydrothermally altered rocks could not be detected through the plant cover by the remote sensing techniques used. For this reason they were considered ideal sites for devising a method to detect altered rocks by using differences in vegetation. Emplacement of intrusive bodies during the Tertiary resulted in alteration of host rocks to form several types of altered rocks. Argillic and silicic alteration are the most common types in the study areas, and are the only types included in this study. In general, the acidic hydrothermal fluids followed faults and fractures in the rocks. The mineralizing fluids changed in composition away from the source, which resulted in a gradual decrease in alteration along the path of the fluid. An- other change that took place outward from the path resulted in a gradation of alteration away from the conduit. These changes pro- duced a zonation of altered rocks. Silicified rocks are closest to the conduit and the source and are surrounded by argillized rocks, beyond which is an area of gradual transition to unaltered rock. Widths of zones vary and can range from a few meters to sev- eral hundred meters (Lovering 1949, 1960, Lovering and Shepard 1959). Degrees of alteration are reflected in the different clay minerals produced. Moderate alteration and supergene weathering under mild conditions favored the formation of montmorillonite. More intense alteration and a more acidic weathering environment re- sulted in the formation of kaolinite. In addi- tion, supergene weathering resulted in the conversion of ferrous iron to limonite, caus- ing the weathering environment to be more acidic in altered areas than in unaltered areas (Lovering 1949). The study includes unaltered, argillized, and silicified sites. The argillized rocks are bleached, limonite stained, and friable. They are formed by cation leaching, addition of water, and formation of clays. Kaolinite, mixed-layer clays, and montmorillonite are present. Kaolinite decreases away from the conduit. Initial porosity is greater in the al- tered rock than in the host rock. In the Utah study area, however, gravity compacts the argillized rocks, causing the pores to close, and a nearly impermeable rock results (H. T. Morris, oral comm. 1979). Silicified rocks are limonite stained and very hard, composed largely of silica, and containing some kaoli- nite and mixed-layer clays. Porosity remains higher than in the fresh and argillized rock because of the rigidity of the matrix sur- rounding the pores (H. T. Morris, oral comm., 1979). The SiOa content of silicified rocks can be as high as 90-95 percent, mak- ing the original texture of the rock difficult or impossible to determine. East Tintic Mountains, Utah The East Tintic Mountains are located in Juab and Utah counties, west central Utah, near the eastern edge of the Great Basin (Fig. 1). The area is classified as semiarid desert, having an average annual precipitation of about 30 cm. The range is made up of Paleo- zoic sedimentary rocks partly overlain by Tertiary volcanic rocks. The Tertiary rocks include quartz latite and latite tuffs and flows. Intrusive bodies associated with the tuffs and flows are numerous, though not large, and range from monzonite to quartz monzonite. The Paleozoic rocks were exten- sively folded and faulted prior to the Tertiary volcanism (Morris 1957, 1964a, Morris and Lovering 1961, 1979, Lovering 1960). The study was confined to two Tertiary units, the Packard Quartz Latite and the overlying Laguna Springs Volcanic Group. The Packard Quartz Latite consists of quartz latite tuffs and flows that contain phenocrysts of andesine, sanidine, quartz, and biotite in a fine-grained to glassy groundmass (Morris and Lovering 1961). The Laguna Springs Volcanic Group con- sists of latite tuffs and flows. The tuffaceous member ranges from fine to coarse grained and in some areas is agglomeratic. The flow is a medium- to coarse-grained latite that contains phenocrysts of orthoclase, plagio- July 1983 Milton, Purdy: Plant-Soil Relationships 459 120^ 115^ + WYOMING 0 100 200 Kilometers L I I ■ J ' Fig. 1. Location of study areas in the Great Basin. clase, hornblende, biotite, augite, magnetite, and quartz (Morris and Lovering 1961). Battle Mountain, Nevada Battle Mountain is located in Humboldt and Lander counties, north central Nevada (Fig. 1). The climate is arid, having about 15 cm average annual precipitation. The area is made up of Cambrian to Tertiary sedimen- tary and volcanic rocks. The pre-Tertiary rocks have been intruded by early Tertiary stocks, sills, and dikes. The geology of Battle Mountain is further complicated by its posi- 460 Great Basin Naturalist Vol. 43, No. 3 tion along the Roberts Mountain and Gol- conda thrust zones. Alteration at Battle Mountain took place at various times throughout the Paleozoic and Mesozoic, and most recently during the Tertiary. Potassic, argillic, and silicic altera- tion are present but are difficult to dis- tinguish from each other in the field; for this report, rocks are categorized only as unal- tered or altered (Roberts and Arnold 1965, Roberts et al. 1971, Theodore and Roberts 1971, Shawe and Stewart 1976, Silberman et al. 1976). Three formations were chosen for the study. The Devonian Scott Canyon Forma- tion is composed predominantly of chert, ar- gillite, and greenstone with minor shale, limestone, and orthoquartzite. As the clastic content increases, the chert grades into argil- lite. Recrystallization of Scott Canyon chert by hydrothermal alteration resulted in an al- tered chert difficult to distinguish in the field from unaltered chert except for the presence of small amounts of altered argillite (Theo- dore and Blake 1975). The Late Cambrian Harmony Formation is composed of feldspathic sandstone inter- bedded with shale and limestone. The sand- stone is medium grained, subangular to sub- rounded, and poorly sorted. The formation is highly susceptible to alteration due to its tex- ture and fine fracture patterns. Many of the mineralized areas were also enriched by su- pergene alteration (Theodore and Blake 1975, Suczek 1977). The Pumpernickel Formation is Early Pennsylvanian to Early Permian in age. It consists of chert and argillite with minor shale, greenstone, limestone, sandstone, and conglomerate. Alteration resulted in recrys- tallization to quartzose hornfels. In addition, some silica was added and most oxides were decreased (Roberts 1964, Theodore and Blake 1975). Vegetation The vegetation of the study areas is in- cluded in the sagebrush and pinyon-juniper zones described by Billings (1951), Blackburn et al. (1968, 1969), Cronquist et al. (1972), Tueller (1975), Young et al. (1976, 1977), and MacMahon (1979). Artemisia tridentata (big sagebrush), commonly with Chrysothamnus nauseosus (rabbitbmsh), Purshia tridentata (antelopebrush), grasses, and forbs, inhabits wide valleys and lower slopes, and occurs more sparsely within the pinyon-juniper woodland. Juniperus osteosperma (Utah juni- per) is the most prevalent tree, found in stands having little undergrowth or scattered among the shrub communities. Pinus mon- ophylla (single-leaf pinyon pine) is a common associate. Stream valleys and moist north- facing slopes contain a dense growth of Acer grandidentatum (bigtooth maple), Prunus vir- giniana (choke cherry), Symphoricarpos oreo- philus (snowberry), and Amelanchier tita- hensis (shadbush). Many altered areas support a sparse and low flora, often including Arte- misia nova (black sagebrush), Petradoria pu- mila (rock goldenrod), and other low matted shrubs and herbs. In addition, several halo- phytic species from the surrounding bajada slopes are found in altered areas in Battle Mountain. Nomenclature for Utah follows Welsh and Moore (1973); that for Nevada follows Munz (1968). Methods Sites for sampling of vegetation and soils were chosen from geologic and alteration maps. To minimize variables, slopes chosen ranged from 1,500 to 2,100 m in altitude, were south facing (azimuth 135° to 225°), and inclined between 12° and 20°. Side slopes and spurs were chosen rather than coves to minimize drainage-catchment differ- ences, and slopes containing springs were avoided. In the field, the sampling sites were further limited to those near roads and rela- tively undisturbed (e.g., not recently chained or burned). Vegetation Sampling A floristic list for each site was made, in- cluding forbs and grasses. Unfortunately, the lists are not complete, because each study area could not be visited during each part of the growing season, and the vegetation data presented here do not include forbs. How- ever, shrubs and trees can be used to indicate soil and water conditions, as well as lithologic variations (e.g., Chikishev 1965). July 1983 Milton, Purdy: Plant-Soil Relationships 461 Areal cover of vegetation was measured by using a modification of the line-interception method (Canfield 1941). Two 15 m tapes were stretched at right angles, one along the contour. The intercept of each species along the tapes was measured to the nearest cen- timeter, and the percent cover was calcu- lated. Dead organic matter (standing dead and litter) was measured as "mulch." For each slope, fifteen 30-meter transects were measured and averaged to give a representa- tive vegetation sample for each site. Total vegetation cover is the percentage of ground covered by trees, shrubs, grasses, or mulch. Soil Sampling Soil samples were taken from the same slopes on which vegetation was measured. Four sites on each alteration type in each geologic formation were sampled at depths of 20 to 60 cm. Rocks larger than about 1.0 cm were removed from samples at the time of collection. Because of the difficulty of an- gering such dry, rocky soils, holes were dug with a small shovel as deeply as possible. As a result, measurements of pore space or bulk density were not possible. Silicification of flow and tuff units ob- scures the original texture so that these units are difficult to distinguish from one another in the field; also they would be expected to form identical soils because of their similar initial chemical composition. For this reason, the silicified Laguna Springs latite soil sam- ples used in the summaries are the same for flow and tuff examples. Soil pH was measured by a glass electrode pH meter on a 1:1 soil-water suspension. Ca- tion exchange capacity (CEC) was measured by the modified barium chloride-triethanola- mine procedure (Chapman and Pratt 1961), using a flame photometer. Clay minerals were identified by X-ray dif- fraction analysis. Oriented slides were run imtreated, after heating to 350 C and 500 C, and after treatment with ethylene glycol. Minerals having (001) peaks at 7.1 A that dis- appeared after hearing to 500 C were identi- fied as kaolinite-group. Illite-group minerals were identified as those having (001) peaks at 10.2 A. Minerals having (001) reflections that expanded from 14 A on untreated samples to 15 A to 17 A on samples treated with eth- ylene glycol were called mixed-layer illite- montmorillonites, and the percentage of ex- pandable layers was estimated from the rela- tive peak intensities. In the absence of other important peaks, relative proportions of the three clay groups could be estimated from the peak intensities (M. Hess, oral comm., 1977). Particle size analysis was done by the hy- drometer method and a series of sieves, using methods adapted from ASTM (1978) and Lambe (1951). Organic matter was not re- moved. Weighed samples were soaked over- night in sodium metaphosphate and sonified to aid dispersion. Soils were separated into the following fractions: clay (<2 jn), silt (2-50 ju), fine sand (50-250 ju) and coarse sand (250-1,000/1). Statistical Analysis The Kruskal-Wallis test for central ten- dency (Gibbons 1976) was used to test the null hypothesis that there were no significant differences in the areal cover of vegetation in the East Tintic Mountains. Areal cover data for Battle Mountain were analyzed by using the Mann-Whitney-Wilcoxon test (Gibbons 1976). Different tests were used because of the different numbers of independent varia- bles in the two study areas. Vegetation and soil differences were tested only within, and not between, the two study areas. Non- parametric tests were chosen in preference to the corresponding parametric tests because of small sample sizes and unknown distributions. A binary discriminant analysis (Strahler 1978a, b) was used to compare affinity of plant species for rock formation and altera- tion type, resulting in a list of plant species for each type that best describes its differ- ence from the other types. This is not a floris- tic list nor a list of dominant species; only those species strongly correlated with rock and alteration type (p = 0.01) are listed. In this test, each vegetation sample is entered separately rather than averaged to give a composite sample for each rock and altera- tion type. Frequency, rather than areal cov- er, is the variable used in this analysis. 462 Great Basin Naturalist Vol. 43, No. 3 Results East Tintic Mountains, Utah The two most common species on the Packard Quartz Latite areas, Artemisia tri- dentata and Purshia tridentata, have higher areal cover measurements on unaltered and silicified sites than on argillized sites (Table 1). Chrysotliamnus nauseosus and Ephedra viridis are found only in imaltered and silici- fied areas. Grasses and mulch also have high- er areal cover on unaltered and silicified sites than on argillized sites. In contrast, the cover of Juniperns osteosperma is higher on the ar- gillized sites. This results in a high total vege- tation measurement, even though there is ac- tually substantial bare ground under and between the trees in argillized areas. With the exception of /. osteosperma, each species has lower areal cover on argillized than on unaltered and silicified sites. The samples from the latite flow sites of the Laguna Springs Volcanic Group also show that samples from unaltered and silici- fied areas have higher shrub and nontree sub- totals than do the samples from argillized sites (Table 1). Grasses and mulch have lower areal cover in argillized areas. An important difference is the presence of Artemisia nova only on the argillized sites. The argillized samples from the Laguna Springs latite tuff (Table 1) have the lowest shrub and nontree subtotals and the lowest Artemisia tridentata, Purshia tridentata, grasses, and mulch. Artemisia nova occurs only on argillized sites. A greater variety of herbaceous species was observed in argillized areas than in unaltered and silicified areas on both tuff and flow, although forbs, as stated above, are not included in the vegetation summaries. The statistically significant differences in shrub cover, nontree cover, and tree cover in Table 1. Vegetation cover Silicified. (in percent) for the East Tintic Mountains study areas. U, Unaltered. A, Argillized. S, Packard Quartz Latite Laguna Springs Volcanic Group Flows U A Tuffs U A Tuffs and flows undifferentiated S Artemisia tridentata 8.23 .81 14.68 16.20 9.19 12.86 7.09 9.28 A. nova 1.42 1.71 Purshia tridentata L61 .49 6.22 .21 .26 5.74 .63 2.64 Chrysothamntis nauseosus .08 .11 .78 .98 .17 C. viscidiflorus Ephedra viridis Opuntia sp. Gutierrezia sarothrae L50 .08 .14 .08 .01 .17 .12 .07 .05 .03 .50 Tetradijynia sp. Arenaria kingii .02 .04 1.18 .02 .10 Petradoria pumila .07 1.01 1.25 .05 Cercocarptis montanus Subtotal n.42 L45 .,31 22.58 16.70 11.72 19.58 11.98 12.76 Grasses Mulch Subtotal 1.96 8.18 10.14 .06 3.30 3.36 1.17 9.58 10.75 2.02 11.74 13.76 1.28 7.85 9.13 4.03 14.62 18.65 1.60 6.85 8.45 3.71 10.85 14.56 Nontree subtotal 21.56 4.81 ,33.,33 30.46 20.85 38.23 20.43 27.32 Juniperus osteosperma Pinus monophylla Tree subtotal 6.90 2.47 9.37 29.34 .29 29.63 5.18 2.99 8.17 5.76 2.75 8.51 6.25 1.53 7.78 4.32 1.99 6.31 1.64 1.64 Total vegetation 30.93 ,34.44 41.50 38.97 28.63 38.23 26.74 28.96 Standard deviation Number of samples ±10.5 15 ±15.9 15 ±12.7 15 ± 9.1 15 ±13.5 15 ± 8.4 15 ±11.5 15 ±11.6 30 July 1983 Milton, Purdy: Plant-Soil Relationships 463 Table 2. Significant differences in vegetation on different alteration types using the Kniskal-Wallis test on the East Tintic Mountains data. Shrubs Total Shrub grasses Tree vegetation cover mulch cover Packard Quartz Latite Unaltered vs. argillized Unaltered vs. silicified Argillized vs. silicified Laguna Springs Volcanic Group: Flows Unaltered vs. argillized Unaltered vs. silicified Argillized vs. silicified Laguna Springs Volcanic Group: Tuffs Unaltered vs. argillized Unaltered vs. silicified Argillized vs. silicified "Significantly different at p < 0.15 samples from the Packard Quartz Latite sites occur between unaltered and argillized sam- ples and between silicified and argillized samples (Table 2). That is, the argillized sam- ples are significantly different from both unaltered and silicified samples, whereas unaltered and silicified samples differ from each other only in total vegetation cover. The Lagvina Springs flow data show signifi- cant differences between unaltered and argil- lized samples in total vegetation, shrub cov- er, and nontree cover (Table 2). The argillized and silicified samples are signifi- cantly different only in the nontree measurements. In the Laguna Springs tuff results, the unaltered and argillized samples differ signif- icantly in total cover, shrub cover, and non- tree cover, and the argillized and silicified samples differ in nontree cover (Table 2). In addition unaltered and silicified samples differ significantly in all but total vegetation cover. Thus, the vegetation is similar in unaltered and silicified areas but different in argillized areas; on the Laguna Springs Volcanic Group, silicified areas are more like argil- lized than like unaltered areas. The binary discriminant analysis results for the Packard Quartz Latite data (Table 3) show that unaltered areas are characterized by Artemisia tridentata and Ephedra viridis, argillized areas by Jtiniperus osteosperma, and silicified areas by Purshia tridentata. The correlation of A. nova with argillized flow and tuff is notable in the Laguna Springs data. Soils in unaltered areas were dug to depths of 25-35 cm. A thin organic layer alternates with pebble- to cobble-sized angular float. Table .3. Binary discriminant analysis results for East Tintic Mountains. Species listed are significantly correlated with rock and alteration type at p = 0.01. U, Unaltered. A, Argillized. S, Silicified. + , d > 2.0. -, d < -2.0. Packard Quartz Latite U A 5 Laguna Springs Volcanic Group U U A S Artemisia tridentata Ephedra viridis Juniperus osteosperma A. nova Purshia tridentata Chrysothamnus nauseosus C. viscidiflorus Gutierrezia sarothrae Arenaria kingii Petradoria pumila Haplopappus acauUs 464 Great Basin Naturalist Vol. 43, No. 3 The rock fragments increase in size as depth increases. The soil is light brown (Munsell color 7.5YR5/4) (Munsell Color Company 1969) to grayish brown (7.5YR4/2). Roots de- crease in frequency as depth increases and penetrate crevices in the rock beyond the depth possible to dig by hand. Soils in argillized areas are shallower (about 15 cm) and range in color from mod- erate brown (7 5YR4/4) to light yellowish or rusty brown (10YR7/4). The organic layer is absent or thinner than in unaltered areas. Pebble- to boulder-sized bleached and limo- nite coated rock fragments cover the top lay- er of soil and are profuse at all levels. At the deepest level dug, rocks are friable and roots are few. Soils in silicified areas are nearly as deep as those in unaltered areas, and color is similar. Bleached rock fragments occur at the surface. Finer particles are found deeper than in soils on unaltered areas, and roots are profuse in lower parts of the soil profile. Soil pH ranges from 6.6 to 7.7 (Table 4). No significant differences were found be- tween soils on different rock or alteration type. Cation exchange capacity (CEC) ranged from 17 to 43 Me/lOOg (Table 4). In general, soils on Packard Quartz Latite areas have higher CEC than soils on the Laguna Springs Volcanic Group, except in silicified areas. Using the Spearman coefficient of cor- relation (Gibbons 1976), the CEC was found to be weakly positively correlated with the total clay content (r = 0.413). CEC is strong- ly negatively correlated (r = -0.780) with the amount of kaolinite in the soil and posi- tively correlated (r = 0.630) with the amount of montmorillonite. However, the lack of significant differences between soils of different alteration types seems to indicate that, although the tests were done with rea- sonable accuracy, the CEC and clay content are too variable within soils of a single altera- tion type to be diagnostic of it. Physical analysis of the soil samples in- cludes particle size analysis for texture and X-ray diffraction analysis for identification of clay minerals. In the Packard Quartz Latite samples, clay content is about the same on all three alteration types (Table 4). Silt content, however, is higher and coarse sand content is lower in silicified areas than in unaltered and argillized areas. In the samples from the latite flow in the Laguna Springs Volcanic Group, the unal- tered soils have the most silt and the least coarse sand, argillized soils have the least silt and the most coarse sand, and silicified soils have intermediate amounts of each. In the sites on the tuffs, the argillized and silicified samples are similar, and the unaltered soils have less silt and more coarse sand. Because clay is formed in the alteration process, a higher clay content would be ex- pected in altered, particularly in argillized, areas. The small clay differences recorded in the samples suggest that dispersion was not complete in all tests. The predominant types of clay minerals in the samples vary (Table 4). The standard de- Table 4. pH, cation exchange capacity (CEC), particle size distribution and relative clay content for East Tintic Mountains soils. U, Unaltered. A, Argillized. S, Silicified. Packard Quartz Latite Laguna Springs Volcanic Group U A S 1 U A S pH 6.8-7.7 6.9-7.6 6.7-7.6 6.7- 7.2 6.6-7.5 6.7-7.4 CEC (Me/lOOg) 36 ±5 37 ±6 25 ±8 28 ±4 Flows 27 ±3 Tuffs 31 ±3 Particle size (percent) 30 ±12 29 ±3 U A U A S Coarse sand 8 ±6 9 ±1 18 ±3 23 ±3 12 ±4 11 ±6 Fine sand 27 ±5 24 ±3 28 ±5 31 ±5 27 ±8 27 ±3 31 ±6 30 ±4 Silt 16 ±5 19 ±6 36 ±5 ,35 ±3 27 ±3 23 ±7 31 ±3 30 ±6 Clay 27 ±5 28 ±3 28 ±2 25 ±3 28 ±7 27 ±2 26 ±4 29 ±5 Relative clay content (percent) Kaolinite 14 25 40 27 21 42 41 33 Ulite 47 49 44 44 67 55 56 61 Mixed-la ver 50 33 24 24 24 9 7 11 Montmorillonite 39 26 16 14 12 4 3 6 July 1983 Milton, Purdy: Plant-Soil Relationships 465 viations of the X-ray data averages are large, so that these data can only be used to make rough comparisons from one alteration type to another. In the samples from the Packard Quartz Latite, the unaltered samples have the least kaolinite and the most mixed-layer clay and montmorillonite. Soils of silicified areas have the most kaolinite and the least mixed-layer clay and montmorillonite. Soils of argillized areas are intermediate between the two but have the greatest amount of illite. The Laguna Springs latite samples show fewer differences in clay type (Table 4). All the soils contain large amounts of illite, less kaolinite, and small amounts of montmorillonite. The soil characteristics of silicified areas resemble those of argillized latite areas in the Lagima Springs Volcanic Group. Following a suggestion of R. P. Ashley (oral comm., 1978), the silicified areas were examined and found to contain large amounts of argillized float around the silicified outcrops. This in- dicates that the soils from silicified areas are mixed with argillized material, resulting in smaller differences in soils and vegetation on the two alteration types than would be ex- pected. The amount of argillized float on sili- cified areas of the Packard Quartz Latite is small, so that soils and vegetation differences are large. Battle Mountain, Nevada The vegetation patterns in the Battle Mountain study area show differences in composition and areal cover from unaltered to altered sites (Table 5). The altered areas have lower total vegetation cover on all three formations, and, except on the Harmo- ny Formation, more variety of species is found in altered areas. Shrub cover is lower on altered sites of the Pumpernickel and Scott Canyon formations than on unaltered sites but remains nearly the same on the Har- mony Formation; areal cover of grasses and mulch is higher on imaltered than on altered sites. Areal cover of Artemisia tridentata is higher on unaltered than on altered sites on all three formations and is absent on altered Harmony Formation. Chrysothamnus nau- seosus is more likely to be found on unaltered sites, and C. viscidiflorus on altered sites. Ar- temisia nova ra'ther than A. tridentata is found on the altered Harmony Formation. Shrub cover and total vegetation cover are significantly different on unaltered and al- tered areas of the Pumpernickel and Scott Canyon formations (Table 6). On the Harmo- ny Formation, the shrub cover is similar on unaltered and altered sites, but the greater amount of mulch on unaltered sites makes the total vegetation cover significantly differ- ent. Differences in vegetation cover between Table 5. Vegetation cover (in percent) for the Battle Mountain study areas. U, Unaltered. A, Altered. Pumpernickel Formation Scott Canyon Formation Harmony Formation Artemisia tridentata A. nova Purshia tridentata Chrysothamnus nauseosus C. viscidiflorus Tetradymia sp. Atriplex confertifolia Ephedra nevadensis PeucephyUum schottii Subtotal 15.9 4.4 .4 3.5 1.9 1.8 1.6 .3 trace 15.2 .2 .2 1.5 trace 11.1 .5 1.4 .4 0.9 .3 19.1 .5 .3 18.8 1.0 20.7 9.1 17.1 14.6 19.9 19.9 Grasses Mulch Subtotal 2.6 10.9 13.5 4.5 3.8 8.3 2.0 9.1 11.1 1.7 9.1 10.8 2.2 1.1 10.0 2.6 12.2 3.7 Total Vegetation 34.2 17.4 28.2 25.4 32.1 23.6 Standard deviation ±7.8 ±4.4 ±6.9 ± 4.3 ±4.6 ±6.7 Number of samples 15 15 15 15 15 15 466 Great Basin Naturalist Vol. 43, No. 3 Table 6. Significant differences in vegetation on dif- ferent alteration types using the Mann-Whitney- Wilcoxon test on the Battle Mountain data. Total Shrub vegetation cover Pumpernickel Formation Unaltered vs. altered • " Scott Canyon Formation Unaltered vs. altered • " Harmony Formation Unaltered vs. altered ' - 'Significantly different at p < 0.15 unaltered and altered sites are greater than those on different rock types. Binary discriminant analysis results (Table 7) show that unaltered Pumpernickel Forma- tion sites are characterized by Artemisia tri- dentata and Purshia tridentata, altered sites by Atriplex confertifolia, Chrysothamnus vis- cidiflorus, and Tetradymia sp. (Table 3). The altered Scott Canyon Formation sites contain Peucephylhini schottii, Chrysothamnus vis- cidiflorus, and Ephedra nevadensis, whereas only Artemisia tridentata is significantly cor- related with the unaltered sites. Arteynisia tri- dentata is characteristic of unaltered Harmo- ny Formation and A. nova, of altered sites. The pH of unaltered Battle Mountain soils was lower than that of altered soils in all three formations (Table 8). Soil depths were greatest (35 cm) in unaltered Pumpernickel chert and Scott Canyon chert soils. Depths of altered soils and unaltered Harmony sand- stone soil averaged 20-25 cm. The particle size distribution analysis re- sults show similar patterns for all three for- mations (Table 8). Soils of unaltered areas are coarser than those of altered areas. The silt difference is greatest in the Harmony Forma- tion, and the coarse sand difference is great- est in the Scott Canyon Formation. As in the Utah study area, the absence of the expected higher clay content in altered areas suggests that the clays may not have dispersed completely. The soils in altered areas of all three for- mations have more kaolinite and less illite than those of unaltered areas (Table 8). In ad- dition, the unaltered Scott Canyon soils con- tain montmorillonite. Again, differences are not large, particularly in the Pumpernickel soils, and the standard deviations are high. Consequently, only general comparisons can be made between soils of unaltered and al- tered areas. The altered sites on all three formations are lower in altitude than the corresponding unaltered sites. This factor, plus the presence of Atriplex confertifolia and the higher pH on the altered sites, suggests that the vegetation may be influenced by increased salinity as well as by a factor in the alteration process. Discussion In the East Tintic Mountains study area, total vegetation cover is lower in argillized areas than in unaltered and silicified areas, except on the argillized Packard Quartz La- tite, where large numbers of Junipenis os- teosperma are found. All other species have lower areal cover on the argillized Packard areas, and the ground is relatively bare under and around the trees. In addition, the compo- sition of the plant communities varies with rock type and alteration history. On unal- tered and silicified areas are large amounts of mulch, Artemisia tridentata, Purshia triden- tata, and Ephedra viridis. Argillized areas, in contrast, contain mostly Juniperus os- teosperma and a very few shrubs. Table 7. Binary discriminant analysis results for Battle Mountain vegetation data. Species listed are significantly correlated with rock and alteration type at p = 0.01. U, Unaltered. A, Altered. +, d > 2.0. -, d < -2.0. Pumpernickel Scott Canyon Harmony U U A U A A Artemisia tridentata + -1- -1- -1- - A. nova 4- Purshia tridentata + Ch njsotham mis natiseosus - - -1- C. viscidiflorus - -t- -1- - - Atriplex confertifolia + Tetradymia sp. - + + - Peucephyllum schottii -1- Ephedra nevadensis -t- July 1983 Milton, Purdy: Plant-Soil Relationships 467 Table 8. pH, particle size distribution and relative clay content for Battle Mountain soils. U, Unaltered. A, Altered. Pumpemi( ckel Formation A Scott Canyon Formation Harmony U Formation U U A A pH 6.8-7.3 7.6-7.9 6.8-7.3 7.2-7.9 6.7-7.3 6.9-7.8 Particle size (percent) Coarse sand 7 ± 1 4 ±2 15 ±2 5 ±3 11 ±3 8 ±2 Fine sand 17 ±2 18 ±1 24 ±2 24 ±4 23 ±4 23 ±4 sat 51 ±2 54 ±3 34 ±2 38 ±6 38 ±5 49 ±3 Clay 25 ±3 24 ±3 27 ±6 31 ±1 28 ±6 20 ±6 Relative clay content (percent) Kaolinite 2.3 28 20 49 17 29 Illite 77 72 70 51 83 71 Montmorillonite 5 0 On the latite of the Laguna Springs Vol- canic Group, vegetation differences are large between unaltered and argillized sites, but small between silicified sites and argillized sites. The vegetation on unaltered sites is composed largely of Arteinisia tridentata and Purshia tridentata, whereas the argillized areas contain A. nova and a wide variety of minor shrubs. In areal cover and in composition of vege- tation the silicified areas of the Laguna Springs Volcanic Group fall between argil- lized areas and unaltered areas. The areas la- beled 'silicified' contain a mixture of silicified and argillized float, which accounts for the similarity of soils and vegetation on argillized and silicified sites. In the Battle Mountain study area, shrub cover and total vegetation cover are greater in unaltered areas than in altered areas; Arte- misia tridentata is the most common shrub in all areas except on altered Harmony Forma- tion, where it is replaced by A. nova. Atriplex conferiifolia occurs on altered sites. Several soil parameters were measured in an attempt to explain the causes of the vege- tation patterns. The hypothesis of low nutri- ent levels to explain the low vegetation cover in some altered areas was discarded because of high vegetation cover measurements in the most highly altered, leached areas. Physical and chemical analyses of the soils include measurements of pH, cation exchange capac- ity, particle size distribution, and X-ray dif- fraction for identification of clay minerals. Although not quantitatively significant, some relationships seem to exist between areal cov- er of vegetation and soil characteristics. The hypothesis that water is the most important limiting factor in plant growth is supported by the following reasons and comparisons: 1. Junipertis osteospemia and Ari:emisia nova, which are found on argillized Packard Quartz Latite and on the argillized Laguna Springs Volcanic Group and altered Har- mony Formation, respectively, are known to occur in the drier habitats in the Great Basin (Blackburn et al. 1968, 1969, Cronquist et al. 1972, Zamora and Tueller 1973, Vasek and Thorne 1977). 2. The high percentage of bare ground, around and under the trees on the argillized Packard Quartz Latite and between the low shrubs on the argillized Laguna Springs Vol- canic Group and altered Scott Canyon and Pumpernickel Formations, results in high runoff and low infiltration of rainfall. 3. Although soil was not dug to bedrock due to the difficulty of digging, the soil ap- peared to be shallower in argillized areas and mixed argillized and silicified areas than in unaltered and silicified areas in the East Tintic Mountains, and shallower in the al- tered Battle Mountain areas than in the unal- tered areas, except on the Harmony Formation. 4. Fractured unaltered and silicified bed- rock allows greater infiltration of rainfall than does the highly compacted argillized bedrock in the East Tintic Mountains. 5. The argillized areas are low in mixed- layer clays and montmorillonite, which would retain more moisture. The presence of montmorillonite on the unaltered Scott Can- yon Formation, Packard Quartz Latite, and Laguna Springs Volcanic Group may in- 468 Great Basin Naturalist Vol. 43, No. 3 crease water capacity. However, total clay content is probably more important than type of clay in areas that have a mixture of clay types. 6. The other major difference between unaltered and altered soils in Battle Moun- tain, the higher pH of altered soils, can ac- coimt for the presence of salt-loving plants such as Atriplex confertifolia but does not ex- plain the decrease in vegetation cover on al- tered sites, because nearby areas that contain halophytic communities have quite dense vegetation. Conclusions Vegetation patterns of areal cover and dis- tribution of species are related to the distri- bution of hydrothermally altered and unal- tered rocks in two areas within the Great Basin. Several factors, including bare ground, shallow soil, impermeable rock, soil texture, and, possibly, clay composition in some areas appear related to low vegetation cover in ar- gillized areas, and suggest that water may be limiting in these areas. The results are con- sistent with those of other workers in the Great Basin. This type of information is needed for the development of techniques for using vegetation as an aid to prospecting in vegetated regions. Acknowledgments The authors are grateful to Drs. J. T. Hack and F. A. Branson of the U.S. Geological Sur- vey for critical reviews of the manuscript. Literature Cited American Society for Testing and Materials. 1978. Standard method for particle-size analysis of soils. Pages 71-81 in ASTM Standards, Part 19. Billings, W. D. 1950. Vegetation and plant growth as affected by chemically altered rocks in the west- em Great Basin. Ecology 31:62-74. 1951. Vegetational zonation in the Great Basin of western North America. Pages 101-122 in Les bases ecologiques de la regeneration de la vegeta- tion des zones arides. International Union of Biol. Sci., Series B, No. 9. Black, C. A. 1968. Soil-plant relationships. Wiley & Sons, New York. 792 pp. Blackburn, W. H. 1973. Infiltration rate and sediment production of selected plant communities and soils in five rangelands in Nevada. College of Ag- riculture, Univ. of Nevada, Reno, Rept. No. R-92. 99 pp. Blackburn, W. H., R. E. Eckert, Jr., and P. T. Tueller. 1969. Vegetation and soils of the Cow Creek Watershed. College of Agriculture, Univ. of Nevada, Reno, Rept. No. R-49. 77 pp. Blackburn, W. H., P. T. Tueller, and R. E. Eckert, Jr. 1968. Vegetation and soils of the Duckwater watershed. College of Agriculture, Univ. of Ne- vada, Reno, Rept. No. R-40. 81 pp. Brooks, R. R. 1972. Geobotany and biogeochemistry in mineral exploration. Harper & Row, New York. 290 pp. Canfield, R. H. 1941. Application of the line inter- ception method in sampling range vegetation. J. of Forestry 39:388-394. Cannon, H. L. 1971. The use of plant indicators in ground water surveys, geologic mapping, and mineral prospecting. Taxon 20:227-256. Chapman, H. D., and P. F. Pratt. 1961. Methods of analysis for soils, plants, and waters. Univ. of Cal- ifornia, Div. of Agr. Sci., Santa Barbara, Califor- nia. 309 pp. Chikishev, a. G. 1965. Plant indicators of soils, rocks and subsurface waters. New York Consultants Bureau, trans.. New York. 210 pp. Cronquist, a., a. H. Holmgren, N. H. Holmgren, and J. L. Reveal. 1972. Intermountain flora. Vol. I. Hafner, New York. 270 pp. FoTH, H. D., and L. M. Turk. 1972. Fundamentals of soil science. Wiley & Sons, New York. 454 pp. Gibbo.ns, J. D. 1976. Nonparametric methods for quan- titative analysis. Holt, Rinehart & Winston, New York. 463 pp. Krohn, M. D., M. J. Abrams, and L. C. Rowan. 1978. Discrimination of hydrothermally altered rocks along the Battle Mountain-Eureka, Nevada, Min- eral Belt using Landsat images. U.S. Geol. Surv. Open-File Rept. 78-585. 66 pp. Lambe, T. W. 1951. Soil testing for engineers. Wiley & Sons, New York. 165 pp. Lovering, T. S. 1949. Rock alteration as a guide to ore- East Tintic district, Utali. Econ. Geol., Monogr. 1. 65 pp. 1960. Geologic and alteration maps of the East Tintic district, Utah. U.S. Geol. Surv. Mineral In- vestigation Field Studies Map MF-230. Scale 1:9600. Lovering, T. S., and A. O. Shepard. 1959. Hydro- thermal argillic alteration on the Helen claim. East Tintic district, Utah. Pages 193-220 in A. Swineford, ed., Clays and clay minerals. Vol. 8, 8th National Conference on Clays and Clay Min- erals, Proc. MacMahon, J. A. 1979. North American deserts: their floral and faunal components. Pages 21-82 in D. W. Goodall and R. A. Perry, eds.. Arid-land eco- systems: structure, hmctioning and management. Cambridge Univ. Press, New York. 881 pp. Malyuga, D. p. 1964. Biogeochemical methods of pros- pecting. New York Consultants Bureau, trans.. New York. 205 p. Morris, H. T. 1957. General geology of the East Tintic Moimtains, Utah. Pages 1-56 in Geology of the East Tintic Mountains and Ore Deposits of the Tintic Mining Districts. Utah Geological Society Guidebook to the Geology of Utah, No. 12. July 1983 Milton, Purdy: Plant-Soil Relationships 469 1964a. Geology of the Eureka quadrangle, Utah and Juab counties, Utah. U.S. Geol. Surv. Bull. 1142-K. 29 pp. 1964b. Geology of the Tintic Junction quad- rangle, Tooele, Juab and Utah Counties, Utah. U.S. Geol. Surv. Bull. 1142-L. 23 pp. Morris, H. T., and T. S. Lovering 1961. Stratigraphy of the East Tintic Mountains, Utah, with a section on Quaternary deposits by H. D. Goode. U.S. Geol. Surv. Prof. Paper 361. 145 pp. 1979. General geology and mines of the East Tintic Mining District, Utah and Juab counties, Utah. U.S. Geol. Surv. Prof. Paper 1024. 203 pp. MuNSELL Color Company. 1969. Munsell book of color. Neighboring hues edition, Matte finish collection, (looseleaf, unnumbered pages). Baltimore, Maryland. MuNZ, P. A. 1968. A California flora. Univ. of California Press, Berkeley. 1681 pp. Nesvetailova, N. G. 1970. Botanical prospecting for ores. Office of Secretary of State of Canada, trans., Ottawa, Canada. 97 pp. OsBORN, B. 1955. How rainfall and nmoff erode soil. Pages 126-135 in Water, USD A Yearbook of Ag- riculture. U.S. Govt. Print. Off., Washington, D.C. Roberts, R. J. 1964. Stratigraphy and structure of the Antler Peak quadrangle, Humboldt and Lander counties, Nevada. U.S. Geol. Surv. Prof. Paper 459-A. 93 pp. Roberts, R. J., and D. C. Arnold. 1965. Ore deposits of the Antler Peak Quadrangle, Humboldt and Lander counties, Nevada. U.S. Geol. Surv. Prof. Paper 459-B. 94 pp. Roberts, R. J., A. S. Radtke, and R. R. Coats. 1971. Gold-bearing deposits in north-central Nevada and southwestern Idaho. Econ. Geol. 66:14-33. Rommel, M. A. 1968. Application of biogeochemistry to mineral prospecting, a survey. NASA SP-5056. 134 pp. Rowan, L. C, and M. J. Abrams. 1978a. Evaluation of Landsat multispectral scanner images for map- ping altered rocks in the East Tintic Mountains, Utah. U.S. Geol. Surv. Open-File Rept. 78-736. 81pp. 1978b. Mapping hydrothermally altered rocks in the East Tintic Mountains using 0.4-2.38 jum multispectral scanner aircraft images, [abs]. Page 156 in International Association on the Genesis of Ore Deposits, Snowbird, Alta, Utah. Program with abstracts. Rowan, L. C, and A. B. Kahle. 1982. Evaluation of 0.46 to 2.36 jum multispectral scanner images of the East Tintic Mining District, Utah, for map- ping hydrothermally altered rocks. Econ. Geol. 77:441-452. Salisbury, F. B. 1954. Some chemical and biological in- vestigations of materials derived from hydro- thermally altered rock in Utah. Soil Sci. 78:277-294. 1964. Soil formation and vegetation on hydro- thermally altered rock material in Utah. Ecology 45:1-9. Sayers, R. W., M. C. Tippett, and E. D. Fields. 1968. Duval's new copper mines show complex geolog- ic history. Pages 55-62 in Mining Engineering, March 1968. Shawe, D. R., and J. H. Stewart. 1976. Ore deposits as related to tectonics and magnetism, Nevada and Utah. American Institute of Mining, Metallur- gical and Petroleum Engineers, Society of Mining Engineers, Trans. 260:225-231. Silberman, M. L., J. H. Stewart, and E. H. McKee. 1976 Igneous activity, tectonics and hydro- thermal precious-metal mineralization in the Great Basin during Cenozoic Time. American In- stitute of Mining, Metallurgical and Petroleum Engineers, Society of Mining Engineers, Trans. 260:253-263. Str.\hler, a. H. 1978a. Binary discriminant analysis: a new method for investigating species- environment relationships. Ecology 59:108-116. 1978b. Response of woody species to site factors of slope angle, rock type and topographic posi- tion in Maryland as evaluated by binary discrimi- nant analysis. J. of Biogeography 5:403-423. SuczEK, C. A. 1977. Tectonic relations of the Harmony Formation, northern Nevada. Unpubli.shed dis- sertation. Stanford Univ., Stanford, California. Theodore, T. G., and D. W. Blake. 1975. Geology and geochemistry of the Copper Canyon porphyry copper deposit and surrounding area. Lander County, Nevada: U.S. Geol. Surv. Prof. Paper 798-B,'86pp. Theodore, T. G., and R. J. Roberts. 1971. Geochem- istry and geology of deep drill holes at Iron Can- von. Lander County, Nevada. U.S. Geol. Surv. Bull. 1318. 32 pp. TuELLER, P. T. 1975. The natural vegetation of Nevada. Mentzelia 1:3-28. Vasek, F. C, and R. F. Thorne. 1977. Transmontane coniferous vegetation. Pages 797-832 in M. G. Barbour and J. Major, eds.. Terrestrial vegetation of California. Wiley & Sons, New York. Welsh, S. L., and G. Moore. 1973. Utah plants: tracheophyta. Brigham Young Univ. Press, Provo, Utah. 474 pp. Young, J. A., R. A. Evans, and P. T. Tueller. 1976. Great Basin plant communities— pristine and grazed. Nevada Archaeological Surv. 6: 186-214. Young, J. A., R. A. Evans, and J. Major. 1977. Sage- brush steppe. Pages 763-796 in M. G. Barbour and J. Major, eds.. Terrestrial vegetation of Cali- fornia. Wiley & Sons, New York. Zamora, B., and P. T. Tueller. 1973. Artemisia arbtis- cida, A. longiloba, and A. nova habitat types in northern Nevada. Great Basin Nat. 33:225-242. PLASTICITY AND POLYMORPHISM IN SEED GERMINATION OF MIMULUS GUTTATUS (SCROPHULARIACEAE) Robert K. Vickery, Jr.' Abstract.— Seeds of 12 populations of Mimulus guttatus representative of the Wasatch Mountain ecotype were incubated for 17 months (one natural season plus a year) in five artificial climates found in phytotron studies to be important to the growth of the plants of that form of monkey flower. In all but the coldest climate, germination oc- curred promptly (3-8 days, on average), peaked during the first three weeks, and then tapered off gradually well into the second season. Generally, the amount and timing of germination was plastic, showing much the same range of responses in widely different climates both overall and for individual populations. However, in some cases, there were significant differences between populations indicative of polymorphism within the species. For example, ger- mination was significantly slower, more variable, and less in amount the higher the elevation of origin of the popu- lations. The responses of the population suggest the presence of both nuich plasticity and much polymorphism for germination characteristics in this form of M. guttatus. The purpose of this investigation is to study seed germination in the yellow Monkey flower, Mimulus guttatus Fischer ex DC, in greater depth than was possible in the earlier surveys (Vickery 1963, 1967). In those sur- veys small samples of a series of species and varieties of Mimulus were studied in a broad range of artificial climates. The present study concentrates on the Wasatch ecotype of Mimulus guttatus (Vickery 1978) and on five artificial climates found in phytotron studies to be important for the growth of M. guttatus (Vickery 1972, 1974). Materials and Methods Seeds from 12 populations of M. guttatus were collected for the study from the Wasatch mountain area of northern Utah and southern Idaho (Table 1). The experiments were carried out in four laboratory artificial climates and one greenhouse climate (Table 2). The seeds were germinated on moist blot- ters in petri dishes. Samples of 500 seeds, 125 per petri dish, were used for each population in each climate. The climates included ex- tremes of the earlier studies (1 and 5), optim- al and suboptimal growth conditions (3 and 4, respectively), and the contrasts of fluctuat- ing and steady temperatures (1 and 2 vs. 3, 4, and 5). Germination was scored for 17 months, that is, through the 5-month germi- nation season normal for M. guttatus in the Wasatch Mountains plus an additional year. Results and Discussion Overall, germination in the four warmer climates (2-5), started as early as the third Table 1. Origins of the populations of Mimulus gut- tatus studied, arranged by culture number, locality, and elevation. M. guttatus Fischer ex DC, n = 14 5839 Spruces, Big Cottonwood Canyon, Salt Lake Co., Utah, 2350 m. 7273 Draper, Salt Lake Valley, Salt Lake Co., Utah, 1390 m. 7274 Gorgoza Ranch, Parley's Summit, Summit Co., Utah, 1910 m. 7311 Fish Haven, Bear Lake, Bear Lake Co., Idaho, 2030 m. 7312 Rick's Springs, Logan Canyon, Cache Co., Utah, 2000 m. 7314- East Canyon, Salt Lake Co., Utah, 2060 m. 7315 Thousand Springs, Mill Creek Canyon, Salt Lake Co., Utah, 2215 m. 7316 Mill F East, Big Cottonwood Canyon, Salt Lake Co., Utah, 2670 m. 7317 Brighton, Big Cottonwood Canyon, Salt Lake Co., Utah, 2645 m. 7318 Homestead, Heber Valley, Wasatch Co., Utah, 1570 m. 7319 Snow Pine, Alta, Little Cottonwood Canyon, Salt Lake Co., Utah, 2710 m. 11157 Mill D North, Big Cottonwood Canyon, Salt Lake Co., Utah, 2520 m. Note: The experiments were carried out at the University of Utah, elev. 1500 m, near the center of the study area. 'Department of Biology, University of Utah, Salt Lake City, Utah 84112. 470 July 1983 Vickery: Mimulus Germination 471 Table 2. Experimental climates used for the seed ger- mination study. 1 Steady 4 C day and night, no photoperiod 2 Steady 25 C day and night, no photoperiod 3 Gradually changing from 4 C night to 17 C day, 16- hour photoperiod 4 Gradually changing from 14 C night to 17C day, 16- hour photoperiod 5 Standard greenhouse, changing from 10 G nights on average to 30 G days on average, 16-hour photoperiod. NoTE:Artificial climates 1 and 2 employed incubators, whereas climates 3 and 4 employed growth chambers programmed to rise and fall like natural July climates in the Wasatch Mountains (Dept. of Commerce, Climatolo- gical Data, 1971-1980). day, peaked during the following week, ta- pered off to a low level by the end of the third week, but continued to occur occasion- ally well into the next year, forming a typical (Went 1957, Vegis 1963) leptokurtic curve (Fig. 1). Despite the overall pattern, germina- tion varied noticeably from climate to cli- mate in both speed and amount (Table 3) as Stakanov (1976) observed in similar studies on beans. For example, in the 17/14 C cli- mate (4), the monkey flower seeds were sig- nificantly slower than in the other cliamtes both in starting to germinate and in achiev- ing 50 percent of the ultimate, total germina- tion for the 17-month trial period (Table 4). The slowing effect on germination of the sub- optimal, 17/14 C climate parallels the strik- ing reduction in plant growth observed in that climate in the phytotron (Vickery 1972, 1974) and suggests that the posited cause, too similar day and night temperatures, acts on speed of germination as well as plant growth. In fact, if the temperature is constant as in climate 2, total germination is significantly less than in the fluctuating climates, 3, 4, 5 (Table 4). Overall, the variable, but generally similar ranges of germination results in the four diverse, warmer climates suggest wide plasticity of response in M. guttatus. In the cold, steady 4 C climate (1), in sharp contrast to tiie pattern of early germination in the four warmer climates, no germination occurred at all during the first four weeks (Fig. 1). After that, apparently the cumula- tive effect of the time spent at room temper- ature while the seeds were being watered and scored triggered a little germination— 2 or 3 seedlings per petri dish— followed by a spurt of germination when the watering and scor- ing time was inadvertently prolonged. Thus, 4^ -U Days Fig. 1. The average number of seeds of all 12 popu- lations of M. guttatus that germinated per day (Table 3) for each of the five climates studied (full data available on request). Peak germination occurred on day 7 in cli- mate 3 with an average of close to 50 of the 500 seeds studied per population per climate germinating that day. The germination rate dropped off to approximately one seed germinating per 25,000 seeds per day by day 255 and to one-tenth of that rate by day 512, the end of the experiment. the latter data for climate 1 are ambiguous and were not analyzed. However, the early results are clear and are consonant with the extremely slow growth of the young plants in the steady 4 C climate of the phytotron (Vickery 1972, 1974). In general, the 12 individual populations of M. guttatus exhibit statistically similar ranges of germination responses to the test climates (Table 5) with some apparent differences (Table 2, 5), much as Wright (1978, 1980) ob- served in Panicwn. The differences range from slight and insignificant to moderate to three cases in which they are so pronounced as to be statistically significant (Table 5). On one hand, the overall similarities suggest a wide plasticity of the populations of re- sponse. On the other hand, the differences appear to reflect underlying genetic differ- ences—polymorphisms—of the populations. The germination results of the 12 popu- lations correlate with the elevation of origin of the populations (Table 6). The populations show significantly longer times to first germi- nation and to 50 percent germination as well as less total germination with increasing ele- vation. This is true not only overall but in most of the individual climates as well (Table 6). The variances of the overall times to first germination are also significantly greater the higher the elevation of origin (Table 6). The 472 Great Basin Naturalist Vol. 43, No. 3 Table 3. Germination results for four replicates of each of the 12 populations of M. guttatus in each of the four warmer climates (Table 2). Climate 1, steady 4 C, was omitted due to the ambiguity of the later results. Population Climate Days to first germination Days to 50% germination Total germination 5839 Spruces 7273 Drape 7274 Gorgoza 7311 Bear Lake 7312 Logan Canyon 7314 East Canyon 7315 1000 Springs 7316 Mill Creek East 7317 Brighton 7318 Homestead 7319 Alta 11,157 MillD North All populations 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 Average 2 3 4 5 5.25 ± 2.50 5.00 ± 1.41 8.00 ± 1.41 5.00 ± 1.82 5.81 ± 2.10 4.00 ± 0.00 5.00 ± 0.00 6.00 ± 0.82 4.50 ± 0.58 4.88 ± 0.89 4.00 ± 0.00 4.25 ± 0.50 5.75 ± 0.96 4.00 ± 0.00 4.50 ± 0.89 6.33 ± 0.00 5.25 ± 0.00 7.75 ± 0.58 5.00 ± 0.50 6.08 ± 0.73 6.33 ± 2.31 5.25 ± 0.50 7.75 ± 0.96 5.00 ± 1.41 6.08 ± 1.67 4.00 ± 0.00 6.00 ± 1.41 5.25 ± 0.50 4.75 ± 0.50 5.00 ± 1.03 4.25 ± 0.50 6.00 ± 0.00 7.00 ± 0.82 5.00 ± 0.00 5.56 ± 1.15 8.00 ± 2.16 6.00 ± 0.82 9.50 ± 1.73 8.50 ± 2.52 8.00 ± 2.16 7.25 ± 2.63 5.25 ± 0.50 7.50 ± 0.58 6.50 ± 1.00 6.63 ± 1.59 3.25 ± 0.50 4.25 ± 0.50 6.75 ± 0.50 5.00 ± 0.00 4.81 ± 1..38 16.00 ± 10.74 7.25 ± 0.96 11.75 ± 1.26 8.50 ± 1.91 10.88 ± 6.04 3.75 ± 0.50 5.25 ± 0.50 7.50 ± 1.73 4.75 ± 0.50 5.31 ± 1.66 5.71 ± 4.60 5.29 ± 1.11 7.35 ± 2.02 5.52 ± 1.80 15.00 ± 9.90 34.75 ± 16.38 8.00 ± 1.83 81.50 ± 51.45 13.50 ± 2.08 73.25 ± 11.38 11.00 ± 0.82 52.25 ± 15.84 11.90 ± 5.37 60.44 ± 31.86 5.75 ± 0.96 56.25 ± 13.77 6.50 ± 0.58 63.50 ± 16.09 8.50 ± 0.58 68.00 ± 41.29 6.50 ± 1.00 86.25 ± 34.60 6.81 ± 1.28 68.50 ± 28.30 14.75 ± 13.10 24.75 ± 6.95 6.50 ± 1.00 109.00 ± 36.21 10.25 ± 2.50 98.75 ± 20.37 6.00 ± 1.15 95.75 ± 23.16 9.38 ± 7.03 82.06 ± 40.69 6.25 ± 0.50 65.50 ± 15.20 6.25 ± 0.50 102.75 ± 22.88 8.75 ± 1.26 78.00 ± 23.85 6.50 ± 1.00 90.25 ± 9.18 6.94 ± 1.34 84.13 ± 22.06 6.50 ± 3.69 2.67 ± 1.41 10.00 ± 0.82 79.25 ± 6.45 12.75 ± 0.50 21.25 ± 16.19 14.00 ± 4.24 67.25 ± 6.85 10.81 ± 4.47 55.11 ± 32.97 10.50 ± 2.65 31.50 ± 7.85 8.00 ± 2.71 88.75 ± 35.88 9.50 ± 1.29 98.00 ± 10.86 8.00 ± 2.16 97.75 ± 11.35 9.00 ± 2.31 79.00 ± 33.21 8.25 ± 2.22 15.25 ± 2.36 9.50 ± 0.58 68.72 ± 7.50 16.50 ± 7.59 77.75 ± 23.21 10.50 ± 1.00 59.00 ± 17.15 11.19 ± 4.85 55.19 ± 28.24 17.25 ± 8.34 17.50 ± 9.61 22.50 ± 12.40 60.50 ± 28.18 32.00 ± 4.32 49.25 ± 7.89 13.50 ± 2.38 34.00 ± 10.95 21.31 ± 10.10 40.31 ± 22.19 10.50 ± 2..38 8.75 ± 7.59 10.50 ± 1.73 47.75 ± 8.66 12.50 ± 1.00 61.50 ± 6.66 9.25 ± 2.06 35.75 ± 9.54 10.69 ± 2.06 38.43 ± 21.34 5.00 ± 0.00 65.25 ± 9.43 6.25 ± 0.50 85.00 ± 7.44 9.75 ± 0.50 74.50 ± 19.16 5.50 ± 1.00 80.25 ± 14.22 6.63 ± 2.00 76.25 ± 14.16 16.75 ± 10.20 3.25 ± 1.21 15.75 ± 12.30 8.50 ± 4.65 15.00 ± 5.35 8.50 ± 3.70 11.50 ± 0.58 9.00 ± 3.74 14.75 ± 7.82 7.31 ± 4.05 5.00 ± 0.00 82.00 ± 26.99 6.00 ± 0.00 97.25 ± 10.34 11.75 ± 1.71 75.25 ± 9.00 5.00 ± 0.00 77.00 ± 22.69 6.94 ± 3.00 83.00 ± 19.09 9.98 ± 7.25 33.90 ± 28.32 9.65 ± 6.56 74.38 ± 34.14 13.40 ± 6.77 69.54 ± 28.18 8.94 ± 3.42 65.38 ± 30.94 July 1983 Vickery: Mimulus Germination 473 Table 4. Comparison of germination in the different artificial climates. Climates that are underlined together are statistically inseparable according to the Student, Newman, Keuls Multiple Range Test (Woolf 1968). aver- age values are given below the lines. Days to 1st germination Days to 50% germination Total germination (Av. per petri dish) 3 5 2 4 5.3 5.5 5.8 7.4 5 3 2 4 8.9 9.6 10.2 13.4 2 5 4 3 34.6 65.4 69.5 74.4 slower and more varied germination re- sponses seem a reasonable adaptation to the increasing unpredictability of the climate with increasing elevation in the Wasatch area (Department of Commerce, Climatolo- gical Data, 1971-1980). This finding of de- creased germination is compatible, also, with my field observations that the populations appear to rely more on rhizomes than on seeds at higher elevations, i.e., there appears to be a cline of decreasing r-selection and in- creasing K-selection with increasing elevation. Although the great majority of seeds ger- minated promptly during the first season in the four warmer artificial climates, a few, about 1 percent on average, in each of the populations did not germinate until the fol- lowing season (Fig. 1). This result parallels the germination response of M. guttatus in the Wasatch Mountains. There, most of the seeds germinate soon after shedding, i.e., dur- ing the later part of the summer. They do not require an after-ripening period as many spe- cies do (Mayer and Poljakoff-Mayber 1975). They continue to germinate in decreasing numbers well into the fall. The seedlings overwinter as small (1-2 cm) rosette stage plants. A few new seedlings appear in the spring along the streams as new habitats are exposed with receding water levels. The late germinating seeds appear to constitute small but important seed banks (Harper 1977), both for springtime germination and for the survi- val of the populations in unfavorable years. Table 5. Comparison of germination of the various populations. Populations within underlined groups are statis- tically inseparable, whereas populations in nonoverlapping groups are statistically distinct according to the Student, Newman, Keuls Multiple Range Test (Woolf 1968). Population culture numbers (Table 1) are given above the lines and average values for each population below the line. Days to first germination 7274 7311 7318 7273 7314 11157 7315 5839 73127317 4.5 4.6 4.8 4.9 5.0 5.3 5.6 5.8 6.1 6.6 7317 7316 6.6 8.0 7319 10.9 Days to 50% germination 7318 7273 7311 11157 7314 7274 7317 7312 7315 6.6 6.8 6.9 6.9 9.0 9.4 10.7 10.8 11.2 7274 7317 7312 7315 5839 7319 9.4 10.7 10.8 11.2 11.9 14.8 7316 21.3 Average total germination 7319 7.3 7317 7316 7315 7312 5839 38.4 40.3 55.2 .58.6 60.4 7315 7312 5839 7273 7318 7314 7274 11157 7311 55.2 58.6 60.4 68.5 76.3 79.0 82.1 83.0 84.1 474 Great Basin Naturalist Vol. 43, No. 3 Table 6. Regression analysis of seed germination in the artificial climates versus elevation of origin of the popu- lation, given as F-ratio and p value. Significant values are underlined. Climates First germination F= p = 50% germination F= p = Total germination F= p = 2 3 4 5 All together Standard deviation (all) 11.462, .007 7.667, .019 16.586, .002 16.599, .002 11.096, .007 6.168, .032 5.520, .041 7.942, .018 4.434, .061 2.767, .127 7.649, .019 4.265, .066 2.444, .149 4.946, .050 6.894, .025 19.650, .001 7.112, .024 2.578, .139 The Wasatch ecotype of Mimulus guttatus exhibits both much plasticity in the overall similarity of its range of seed germination re- sponses to widely different climates and some apparent genetic polymorphisms for speed and amount of germination during the first season, for the speed and amount of germina- tion in populations with different elevations of origin and for the ability to delay germina- tion until the second season. Thus, the eco- type appear to be well adapted for survival in its climatically unpredictable area. Acknowledgments I thank Michael Nellestein for his technical assistance with the seeds and Harold Hurst and Dennis Phillips for their assistance with the statistical checks. Literature Cited Harper, J. L. 1977. Population biology of plants. Aca- demic Press, London. 892 pp. Mayer, A. M., and A. Poljakoff-Mayber. 1975. The germination of seeds. 2d ed. Pergamon Press, Ox- ford, 178 pp. Stakanov, F. S. 1976. Changes in the rate and simulta- neity of bean seed germination in relation to ger- mination temperature. Krishnev Inst. Transac- tions 153:27-29. U.S. Department of Commerce. 1971-1980. Climatolo- gical data. Utah. Vols. 73-82. Vegis, a. 1963. Climatic control of germination, bud break, and dormancy. In L. T. Evans, ed.. Envi- ronmental control of plant growth. Academic Press, New York. 449 pp. VicKERY, R. K., Jr. 1963. The evolutionary potential, as measured by seed germination, of chromosome races of Mimulus (Scrophulariaceae). Proc. XI In- ter. Cong. Cenet. 1:46. 1967. Ranges of temperature tolerance for germi- nation of Mimulus seeds from diverse popu- lations. Ecology 48:647-651. 1972. Range of climatic tolerance as an in- dication of evolutionary potential in Mimulus (Scrophulariaceae). Symp. Biol. Hung. 12:31-42. 1974. Growth in artificial climates— an indication of Mijuulus' ability to invade new habitats. Ecol- ogy 55(4):796-807.' 1978. Case studies in the evolution of species complexes in Mimulus. Evol. Biol. 11:405-506. Went, F. W. 1957. The experimental control of plant growth. Chronica Botanica, Waltham, Massachu- setts. 343 pp. WooLF, C. M. 1968. Principles of biometry: statistics for biologists. Van Nostrand, Princeton, New Jersey. 359 pp. Wright, L. N. 1978. Recurrent selection for changing gene frequency of germination rate in Blue Panic grass. Crop Sci. 18(5):789-791. Wright, L. N. 1980. Germination rate and growth char- acteristics of Blue Panic grass, Panicum anti- dotale. Crop Sci. 20(l):42-44. PREDATORY BEHAVIOR OF LARVAL AMBYSTOMA TIGRINUM NEBULOSUM ON LIMNEPHILUS (TRICHOPTERA) LARVAE Joseph R. Holomuzki' Abstract.— Examination of stomach contents indicated that second-year Ambystoma tigrintim nebulostiin larvae consumed Limneptnlus sp. larvae but rarely ingested the case. Feeding observations of captive salamanders on cad- disfly larvae supported this finding. Extraction of caddisfly larvae from their case was accomplished only when larval .salamanders quickly seized the anterior portion of ambulatorv Limnephihis sp. extended from their case and vigor- ously shook the trichopteran from side to side. Interest in the predatory behavior of Am- hystoina tigrinum nebtiloswn on Limnephihis sp. was prompted by the examination of stomach contents of 29 second-year larvae (^-13 mos. old) from east central Arizona. Fourteen salamanders had eaten a total of 71 Limnephilus sp. larvae, yet remains of only 7 cases were evident. This indicated that A. tigrinum were extracting caddisfly larvae from their case. Such feeding behavior appar- ently contrasts with some A. tigrinum in Utah where individuals ingested caddisfly larvae with cases (Tanner 1931). Moreover, extrac- tion of larva from the case is discordant with the notion that larval A. tigrinum exploit aquatic resources in a manner nearly identi- cal to freshwater fish (Zaret 1980), since fish typically consume both caddisfly case and larva (Elliot 1967, Tippets and Moyle 1978, W. L. Minckley, pers. comm.). This paper describes the ability of captive A. tigrinum nebulosum larvae to extract Limnephilus sp. larvae from their cases. Salamanders used for feeding observations and stomach analyses were collected in June 1981 from Big Meadows Tank 1, a per- manent pond located 1.0 km NNW of the western edge of Sunrise Lake, Apache Co., Arizona (elev. 2,774 m). Eight second-year larvae varying from 78 to 98 mm from tip of snout to posterior margin of vent were indi- vidually kept in 36 X 22 X 26 cm aquaria partially filled with 50 percent Holtfreter's solution. Animals were acclimated for 24 hours before feeding observations were in- itiated. Limnephilus sp. collected from Big Meadows Tank 1 also were kept in 50 per- cent Holtfreter's solution. Each salamander was provided six caddisflies during feeding runs. The number of strikes and successful captures were counted in each 1-3 hour run. Salamanders were not fed between observations. Movement by Limnephilus sp. seemed to provide a visual stimulus for a strike response by these salamanders. Previous studies also noted A. tigrinum larvae typically striking moving prey (Dodson and Dodson 1971, Rose and Armentrout 1976). My observations, however, suggest tactility may also play a role in stimulating an attack on prey. Attrac- tion of a salamander to a caddisfly case was apparently frequently caused by any move- ment of the case. The salamander usually halted and placed its snout or chin against a case that had moved. Further movement by the caddisfly stimulated a strike. Salamanders withdrew from the case if cessation of move- ment was protracted. During 26 hours of observations, only 2 (3.4 percent) Limnephilus sp. larvae were eaten in 58 strikes. Unsuccessful attempts at prey capture consisted of a salamander tak- ing the entire case into its mouth. The animal then manipulated the case and discarded it after about 14 seconds (N = 12, range: 3-85 seconds). On no occasion was the case con- sumed. Caddisfly larvae were successfully at- tacked and eaten only when a salamander slowly approached an ambulatory Limne- philus sp. extended from its case and quickly seized the anterior portion of the larva. The salamander then vigorously shook the tri- chopteran from side to side until extracted. 'Department of Zoology, Arizona State University, Tempe, Arizona 85287. 475 476 Great Basin Naturalist Vol. 43, No. 3 Similar head-whipping behavior by metamor- phosed A. trigrinum on elongate prey was de- scribed by Larsen and Guthrie (1975) and Lindquist and Bachmann (1980). This method of Limnephilus sp. capture seemed to be sup- ported by the stomach analyses, in which 10 of 71 consumed larvae were severed 0-2 mm behind the metanotum. The relatively poor capture success by salamanders in the laboratory may also be characteristic of the natural habitat. For ex- ample, Dodson and Dodson (1971) found rel- atively few tichopteran larvae in the diet of A. tigrinum larvae from Colorado, even though the insects were abundant in the sam- led pond. Limnephilus sp., however, com- prised about 16 percent of the total volume of prey in the diet of larvae in June from Big Meadows Tank 1. This suggests salamander larvae of this population frequently attacked trichopterans. In sum, stomach contents showed that A. tigrinum larvae from this population in- frequently ingested caddisfly cases. Absence of case consumption in the laboratory sup- ported the finding. Successful attacks on Lim- nephilus sp. were few and occurred only when A. tigrinum quickly seized the anterior portion of a caddisfly extended from its case. These observations suggest caddisfly cases are an effective means of deterring predation by larval salamanders of this population. Acknowledgments Thanks are extended to J. P. Collins and W. L. Minckley for commenting on an ear- lier draft of the manuscript. Literature Cited Dodson, S. I., and V. E. Dodson. 1971. The diet of Ajji- bijstoma tigrinum larvae from western Colorado. Copeia 1971:614-624. Elliot, J. M. 1967. The food of trout (Salmo tnitta) in a Dartmoor stream. J. Appl. Ecol. 4:59-71. Larsen, J. H., Jr., and D. J. Guthrie. 1975. The feeding system of terrestrial tiger salamanders {Ambtjs- toma tigrinum mekmostictum Baird). J. Morph. 147:137-154. Lindquist, S. B., and M. D. Bachmann. 1980. Feeding behavior of the tiger salamander, Ambystoma tigrinum. Herpetologica .36:144-158. Rose, F. L., and D. Armentrout. 1976. Adaptive strate- gies oi Ambystoma tigrinum Green inhabiting the Llano Estacado of west Texas. J. Anim. Ecol. 45:71.3-729. Tanner, V. M. 1931. A synoptical study of Utali am- phibia. Utah Acad. Sci. 8:159-198.' Tippets, W. E., and P. B. Moyle. 1978. Epibenthic feeding by rainbow trout. J. Anim. Ecol. 47:549-559. Zaret, T. M. 1980. Predation and freshwater commu- nities. Yale Univ. Press, New Haven. NOTES ON REPRODUCTION OF THE SIDE-BLOTCHED LIZARD UTA STANSBURIANA STANSBURIANA IN SOUTHWEST IDAHO George Bakewell', Joseph M. Chopek', and Gary L. Burkholder' Abstract.— In 1968 and 1973 in southwestern Idaho, 275 Ufa stanshuriana were collected. Uta occupies low hills with rocky outcrops and flat sparse vegetation consisting of the following shrubs: Arternisia tridentata, Grcnjla spin- osa, Atriplex confertifolia, Chrysothamnus nauseosus, and Atriplex canescens. Emergence from hibernation occurs from mid-March to early April and the first yolked follicles appear in early April, with oviducal eggs present in late April. Testicular cycle begins with the emergence of males, and spermatozoa are produced from April through July. Uta reaches sexual maturity in one year at a SVL of 40.0 mm in males and 43 mm in females. Overall clutch size is 3.75 eggs (1-2 clutches per year). Fat body size at emergence is not known, but what is present shows a decline in males and females until July, when a substantial increase occurs. Sex ratios are about 1:1 in all months and seasons except June 1968, according to chi-square analysis. This study has hmited scope for two rea- sons: (1) the years of study are five years apart; (2) in 1968 there were only six trips to the field (which was about 10 miles from the 1973 area) in May and June, whereas in 1973 field trips were made once each week from 4 April through 25 July. In view of this, the 1968 data are minimized to some extent in drawing our conclusion and emphasis is on the 1973 data. Despite these limitations, the study sheds some light on the reproductive cycle of Uta stanshuriana in the northern portion of its range. Uta has been studied in considerable detail in Texas (Tinkle 1961, 1967a, Hahn and Tinkle 1965), Colorado (Tinkle 1967b), south- em Nevada (Hoddenbach and Turner 1968, Turner, Hoddenbach, Medica, and Lannom 1970, Medica and Turner 1976, Tanner 1972), and Oregon (Nussbaum and Diller 1976). Our results show that latitudinal (and therefore climatic) differences exert observ- able modifications of the reproductive cycle as compared to that of southern populations in Colorado, Nevada, and Texas, which are 800-1200 km south of our study, respective- ly. In the case of the northern population studied by Nussbaum and Diller (1976), our results are similar in some aspects, as would be expected, since their study area is only 225 km further north. Physical and Biotic Environment These counties included in the study area are part of the Snake River Valley (Fig. 1). Elevations of study sites and collection areas range from 530 to 750 m. Dominant topo- graphical features consist of small canyons, which are a part of the Snake River Drain- age, low rolling hills, boulder-strewn areas, rocky outcroppings, and intermittent streams that form sandy washes and/ or alluvial fans Fig. 1. Geographic location for capture sites (hollow circles) of Uta stanshuriana in southwestern Idaho. In- sert shows the portion of Idaho where collections were made. 'Mount Vemon Nazarene College, Martinsburg Road, Mount Vemon, Ohio 43050. 477 478 Great Basin Naturalist Vol. 43, No. 3 where they reach the desert floor. Annual rainfall varies from 10.1 to 30.4 mm; most of this occurs as snow in winter (Shreve 1942). Summers are typically hot and dry, with few overcast days. Frost free days average 150 per year from April to September. The dominant floral description for the study areas is that of "cold desert formation" (Shreve 1942). The dominant floral associ- ations consist of several different com- binations of the following desert shrubs: Grayia spinosa (Hook.), Atriplex canescens (Pursh, Ex. Nutt.), Atriplex confertifolia (Torr.), Artemisia tridentata (Nutt.), and Chrysothamniis nauseosus (Poll.). These shrubs reach heights of 1 to 1.8 m and grow in a clumped pattern that creates open areas (Burkholder and Walker 1973). Many species of annuals are present in the open areas be- tween the shrubs, the majority of which flow- er during the early and midspring rainy peri- od. Others flower during the occasional rainy periods of late spring and summer. Methods and Materials Specimens for this study were collected in 1968 and 1973 in Canyon, Ada, and Owyhee counties of southwestern Idaho. All lizards were weighed, measured (SVL), and autop- sied (except hatchlings). The ovaries and ovi- ducts of the females were removed and counts were made of yolked ovarian follicles, oviducal eggs, and corpora lutea. Yolked fol- licles and oviducal eggs were measured to the Table 1. Sex ratios for six samples of Vta stansbu- ruina in 1968 and 1973. Each year total was tested for goodness of fit by the chi-square statistic as well as the month of Jime, where the largest differences occurred. All fit the 95 percent confidence limit except June 1968. nearest 0.1 mm and weighed to the nearest .01 g. In males, one testis was removed and used to determine sexual maturity by a squash preparation using aceto-orcein stain. Clutch size data followed the procedure out- lined by Tinkle (1961). Fat bodies were also removed and weighed to the nearest .01 g. Results Sex ratio.— Sex was determined in 275 specimens by the examination of gonads. In May and June 1968, 34 females and 50 males were collected. The April through July 1973 sample consisted of 90 females and 101 males. Both years combined resulted in 124 (45.1 percent) females and 151 (54.9 percent) males. Data for sex ratios are summarized in Table 1. Statistical analysis for fitting the expected ratio of 1:1 are also included in Table 1. Size at maturity and at hatching.— Size of individuals when reaching sexual maturity was determined by analyzing the gonads. In females, yolked ovarian follicles, oviducal eggs, and corpora lutea were used as criteria for maturity. From this analysis, it was deter- mined that the smallest sexually mature fe- male was 43 mm SVL for the 1973 sample and 44 mm SVL for the 1968 sample. Size at sexual maturity in males was deter- mined by analysis of a single testis removed from each male. An aceto-orcein squash preparation was performed to check for the presence of mature spermatozoa. All male specimens, even the smallest at 40 mm SVL, yielded a positive test for spermatozoa. The Table 2. Data for hatchling Vta stansbiiriana cap- tured in July 1973. Two lizards not listed were observed on 26 June but not captured. Month Year Females 14 Males 13 SVL (mm) Date Captured May 1968 26.0 3 July June 1968 20 37 25.5 18 July Total 34 50 25.0 23 July Chi-square test .95 = 5.06 for Tune 25.0 23 July Chi-square test .95 = 3.04 overall 26.0 31.0 23 July 23 July April 1973 20 16 31.0 23 July May 1973 29 33 31.5 23 July June 1973 23 32 36.0 23 July July 1973 18 20 28.0 25 July Total 90 101 29.0 25 July Chi-square test .95 = 1.472 for June 31.0 25 July Chi-square test .95 = .632 overall 33.0 25 July July 1983 Bakewell et al.: Side-Blotched Lizard 479 6 - X=6.00 5 - 4 - 5a?. Q? X=4.13 11 II c cc Y— •3 £1 3 •» ^^ VO 1 i-l C\J 11 II c OS A-J.O 1 A-J.O/ un 1 CJ CO II 11 ^ . X=3.00 c<^ 1 II 11 in O 1 11 1! 2 - 1 - 0 1 11 II C PC 43-44 45-46 47-48 49-50 51-52 53-54 Size class of females (SVL) Fig. 2. The mean number of yolked ovarian follicles, oviducal eggs, and corpora lutea for six arbitrarily sized class- es of females. N = number of individuals, R = range, and X = mean 1968 and 1973 females combined. 12 hatchlings were not sexed but were as- sumed not to be reproductively mature. The first hatchlings were observed, but not collected, on 26 June 1973. Twelve speci- mens were collected from 3 July 1973 to 25 July 1973. The data for hatchlings is pre- sented in Table 2. Mean size of mature individuals.— The largest male specimen was 57.5 mm SVL and the smallest 40.0 mm SVL. The largest fe- male was 53.0 mm SVL and the smallest was 43 mm SVL. Table 3 summarizes the remain- ing data related to mean size of mature individuals. Reproduction in females.— Counts were made of oviducal eggs, corpora lutea, and yolked ovarian follicles in the ovaries of 124 females. Figure 2 shows a comparison of six arbitrary size classes that indicates that clutch size increases with size of the female, with the two smallest categories (43-44 and 45-46 SVL) presumably being first-year re- producive females, and those 47 and over second-year females. Using 1973 female data alone, because it covers the entire reproduc- tive season, there was an attempt to deter- mine the number of clutches per year, which is difficult in multiple clutch species. The 1973 data appears to indicate a two- clutch ability by some individuals. The first clutch group is clustered between 23 April and 22 May (based on shelled oviducal eggs). The second clutch group is clustered between 2 June and 10 July. Between the period of 22 May and 2 June, there are females with ovi- ducal eggs but these are late individuals re- productively, which will probably produce only one clutch because of time constraints. To substantiate the double clutch, we began with the correlation of the first appearance of hatchlings, assuming a 45-50 day in- cubation period, which is based on Bur- kholder and Tanner's (1974) work on Scelo- porus graciosus, which has a mean incubation period of 49 days (from laboratory and field incubation data). If the females of the first group laid eggs from 23 April through 22 May, with a 50-day incubation period, the hatchlings would appear as early as 11 June and as late as 10 July. The first hatchlings Table 3. The mean SVL in millimeters of sexually mature males and females for 1968 and 1973, based on sperma- tozoa in testis and epididymus and yolked ovarian follicles, oviducal eggs, or corpora lutea, respectively. Males Females Year X N R X N R 1968 1973 49.27 49.66 51 100 44-57 40-55 47.70 48.06 34 99 44.5-52 43-53 480 Great Basin Naturalist Vol. 43, No. 3 were observed on 26 June and 3 July (Table 2), which falls within the predicted time. The second clutch hatchlings would appear 21 July through 28 August. The smallest hatch- lings captured between 23 July and 25 July (Table 2) fit into this category. The second point that supports two clutches is that of the time span between the first and second onset of vitellogenesis. Turn- er, et al. (1970) state that 31 days are suf- ficient for production of a second clutch. If the days are counted between the proposed first and second clutch (clustering of females with shelled oviducal eggs), there are 42 days, which would appear to be ample time. The only nonsupportive evidence is the ab- sence of corpora lutea when the second pro- duction of yolked ovarian follicles occurs. However, it is our opinion, based on observa- tions of autopsied individuals, that corpora lutea in Uta disappear very quickly (1-5 days), which would explain the lack of overlap. Along with the double clutch phenomena is the feature of clutch size fluctuation. The mean size of the first clutch (based on ovi- ducal eggs only) was 4.40 (N = 14) and the second was 3.81 (N = 16). This corresponds to that reported by Tinkle (1967b) for Texas populations, Turner et al. (1970), and Medica and Turner (1976) for Nevada, and Nussbaum and Diller (1976) for Oregon, though the dif- ference is not as significant. The earliest onset of vitellogenesis is 14 April (1973). This is based on yolked ovarian follicles of 2 mm diameter or larger and a definite yellow color. Table 4 summarizes the numbers of yolked ovarian follicles, oviducal eggs, and corpora lutea for the determination of clutch size (1968 and 1973 data were Table 4. Data for yolked ovarian follicles, oviducal eggs, corpora lutea, and mean clutch size per female for the years 1968 and 1973. Ovarian Mean yolked Oviducal Corpora clutch Year follicles eggs lutea size 1968 45(N=12) 65(N=17) 0 3.79 1973 112(N = 33) 135(N = 33) 7(N = 2) 3.74 Total 157(N = 45) 200(N = 50) 7(N = 2) 3.64 Mean 3.49 4.00 3.50 3.75 Clutch size Clutches per year pooled). Reproductive potential was not cal- culated; the actual numbers of females laying a second clutch was not determined. Fat bodies.— The fat bodies decrease in weight from the time of emergence of adults through the end of the reproductive period in both males and females (Table 5) Discussion Although the volume of data and length of time are much less than studies which we will use for comparison and contrast, this study provides some additional information concerning the total knowledge of natural history of Uta stansburiana. In essence, the comparisons made with regard to Uta con- cern that of northern vs. southern popu- lations and the changes that occur along that continuum. The first feature is that of sex-ratio. On a yearly basis, all (1968 and 1973) fit the Table 5. Fat body weight changes in 1973 male and female Uta stansburiana April-July in grams. Single as- terisk means that all had yolked follicles or oviducal eggs. Double asterisk means the two at < .01 still had oviducal eggs and corpora lutea and the one at .08 cor- pora lutea only, and the rest had no reproductive activi- ty. Triple asterisk means the .25 specimen had just be- giui vitellogenesis; the two at < .01, two at .02, and two at .03 had not started vitellogenesis, whereas all others had. April May Jv July Males 16=<.01 32=<.01 26=<.01 3=<.01 1 = .08 1 = .01 2 = .01 1 = .03 2=.04 1 = .05 2 = .06 2 =.07 1 = .08 2 =.09 1 = .10 1 = .15 N 16 33 32 18 Females 11 = < .01 18= < .01 14= < .01 1 = <.01 4 =.03 1 = .05 3 =.02 1 = .04 1 = .25 2 =.04 2 =.03 2 = .02 1 = .05 2 =.03 1 = .02 3.75 1-2 1 = .02 2 = .05 1 = .06 1 = .07 1 = .08 1 = .09 2=.10 2 = .ll 1 = .12 1 = .15 21.0. 24° 18° 15° July 1983 Bakewell et al.: Side-Blotched Lizard 481 expected ratio of 1:1. Monthly fluctuations are not as consistent, especially in June for both years. These data indicate a 3:2 ratio of males to females, which is similar to what Tinkle (1961) recorded in Texas. Tinkle (1961) attributed this to territorial vigilance in males, and this appears to be true for those in Idaho. In addition to this, the difference may be further enhanced by female oviposi- tion in June. By July these activities (male and female) cease, and the ratio returns to 1:1. Idaho Uta attain sexual maturity in one year; they hatch, over winter, and emerge as adults (Tinkle 1961, Tanner 1972, Medica and Turner 1976). Nussbaum and Diller (1976), however, observed that in north cen- tral Oregon some Uta that hatched late in the year (late August) would not have suf- ficient time for growth and therefore would not be sexually mature by the time the next Table 6. Comparison of data for six Uta populations. reproductive season arrived. We think that this is rare in Idaho, based on size of earliest individuals collected, because they had yolked ovarian follicles, spermatozoa, and larger SVL than the minimum SVL as deter- mined for sexual maturity of Oregon Uta. The longer growth period of Idaho vs. Ore- gon is most likely the basis for the differences in the two populations. The remainder of our findings concerning Idaho Uta have been added to Table 6, which is a modification from Nussbaum and Diller (1976). Table 6 calls attention to sever- al areas of interest. The elevation is as low or lower than all the previous studies, though further north than all but the one in Oregon. Length of growing season and therefore length of reproductive season are longer than that found in Oregon but less than the four southern studies. These physical aspects put the Idaho population in a somewhat inter- mediate position between that of Oregon and Texas Colorado Nevada (Rock Valley) Nevada (Rainier Mesa) Oregon Idaho (1973) Elevation (ft) Length of growing season (days) Length of reproductive season (davs) Habit' Sexual dichromatism Aggressiveness Social stnicture Density (individuals /acre) Male home range (acres) Female home range (acres) Hatchling size (nun(SVL) Size range mature males (mm) Size range mature females (mm) Average size adult males (mm) Average size adult females (mm) Clutch size Clutch frequency Percent males fail to breed first season Percent females fail to breed first season Percent males two years old and older Percent females two years old and older Date hatchlings appear 2900 ^4250 3400 7840 2400 1722-2437 215 175 225 200 110-140 150-180 121-141 120 135 _ 70 90-100 arenicolous saxicolous — — saxicolous saxicolous high 103 — — high high high 103 — — low territorial dominance — — dominance (?) dominance (?) 36.109 17.5 24 10 71 _ 0.50 0.27 — — 0.54 — 0.17 0.23 — — 0.43 — 22 ^22 s22 ^22 22 26' 40-60 42- - 40-56 40-53 40-55 40-60 37- 40 + 41-49 43-53 - - - 49 48.4 49.6 48.9 42.8 45.4 48.0 ^4.0 3.20 .6- .4 4.85 3.33 3.75 3.5 3 .3-5 - 1-2 1-2 0.0 <25 - - = 19.0 - 0.0 <25 - - = 47.0 I 7 33 18-28 36-65 57.6 - 7 33 18-28 36-65 69.4 _ 20 June 25 June 25 June 17 July 17 July 26 June Texas and Colorado (Tinkle 1961, 1967a, b, 1969a, Tinkle and Woodward 1967) Oregon (Nussbaum and Diller 1976) Nevada-Rock Valley (Turner et al., 1970, Medica and Turner 1976) Nevada— Rainier Mesa (Tanner 1972) Idaho— (this study) "Smallest individual caught 3 July 1973 482 Great Basin Naturalist Vol. 43, No. 3 the four southern populations, even though it is not located halfway between from the standpoint of miles. Because of these factors and others (i.e., precipitation and primary production), there should be some influence on reproductive cycle, average adult size, date of hatching, appearance, etc. In reference to the above-mentioned pa- rameters and the information in Table 6, it becomes apparent that Idaho Uta do lay more than one clutch per year. This is based on the broad period of time when females have oviducal eggs present (the same is true for yolked ovarian follicles) and the appear- ance and size (SVL) of hatchlings from 26 June through late July. The percentage of fe- males laying a second clutch is speculative because of incomplete data, but it would ap- pear to occur in the majority of the popu- lation as compared to a small percentage in the Oregon population as reported by Nuss- baum and Diller (1976). It is doubtful if any could produce a third clutch, as is the case in Colorado Uta (Tinkle 1976), due to length of reproductive season, though the possibility does exist if favorable conditions prevail. Coupled with this is the fairly high overall clutch size of 3.75 or, if based on oviducal eggs only (first and second clutch), 4.40 and 3.81/female, which approaches the size if not equals that of the Texas, Colorado, and Ne- vada (Rock Valley) populations. The only dif- ference is the number of clutches per year. The size is significantly higher than that of Oregon. The explanation for the similarity to populations much farther south and the dif- ference from the more northern Oregon pop- ulation is complicated but again probably is within the realm of the response of Uta to changes in the elevation and/or latitude, length of growing and reproductive seasons, annual precipitation as it affects primary production, and general habitat. Because of the limitation of this study, we feel in- adequate to speculate on these inter- relationships at this time. Average size of males and females (SVL), as well as size range for mature males and fe- males, does not appear to deviate dramatical- ly from other populations. Fat body cycle follows that reported first by Hahn and Tinkle (1965) and by many subsequent au- thors with regard to various lizard species. Literature Cited BuRKHOLDER, G. L., AND W. W. Tanner. 1974. Life his- tory and ecology of the Great Basin sagebrush swift, Sceloporus graciosits graciosus Baird & Gi- rard, 1972. BYU Sci. Bull., Biol. Ser. 19(5): 1-42. BuRKHOLDER, G. L., AND J. M Walker. 1973. Habitat and reproduction of the desert whiptail lizard, Cnemidophorns tigris Baird and Girard at the northern part of its range. Herpetologica 29:76-83. Hahn, W. E., and D. W. Tinkle. 1965. Fat body cycling and experimental evidence for its adaptive signif- icance to ovarian follicle development in the liz- ard Uta stansbuhana. J. Exp. Zool. 158(l):79-86. Hoddenbach, G. a., and F. B. Turner. 1968. Clutch size of the lizard Uta stanshuriana in southern Nevada. Amer. Midi. Nat. 80(l):262-265. Medica, p. a., and F. B. Turner. 1976. Reproduction by Uta stanshuriana (Reptilia, Lacertilia, Igua- nidae) in southern Nevada. J. of Herpetology 10(2): 123-128. NussBAUM, R. a., and L. v. Diller. 1976. The life his- tory of the side-blotched lizard, Uta stanshuriana Baird and Girard, in north-central Oregon. Northwest Sci. 50(4):243-260. Shreve, F. C. 1942. The desert vegetation of North America. Bot. Rev. 8:195-246. Tanner, W. W. 1972. Notes on the life history of Uta stanshuriana. BYU Sci. Bull., Biol. Ser. 15(4):31-39. Tinkle, D. W. 1961. Population structure and reproduc- tion in the lizard Uta stanshuriana stejnegeri. Amer. Midi. Nat. 66:206-234. 1967a. Home range, density, dynamics, and struc- ture of a Texas population of lizard Uta stanshu- riana. Pages 5-59 in W. W. Milstead, ed.. Lizard ecology: a symposium. Univ. of Missouri Press. 1967b. Life and demongraphy of the side- blotched lizard. Univ. Michigan Misc. Publ. Mus. Zool. 1.32:1-182. Turner, F. B., G. D. Hoddenbach, P. A. Medica, and J. R. Lannon. 1970. The demography of the liz- ard Uta stanshuriana Baird and Girard in south- ern Nevada. J. Anim. Ecol. 39:505-519. OBSERVATIONS ON ALPINE VEGETATION NEAR SCHOOLROOM GLACIER, TETON RANGE, WYOMING John R. Spence' and Richard J. Shaw' Abstract.— Quadrat and propagiile trapping studies were made on the moraine of the Schoolroom Glacier and in adjacent dry alpine meadow vegetation in the Teton Range in 1978-1979. Forty-six species of vascular plants were collected. Distributionally, three groups of species exist. One of these is concentrated primarily on the moraine, a second in the meadow, and the third in a narrow ecotonal band at the base of the distal slope of the moraine. The moraine slopes are steep and unstable, with vegetation cover ranging from 1 to 9 percent, dominated by Cirsium tweedyi (Rydb.) Petr. Along the more stable moraine crest the vegetation cover is heavier, and is similar to that in the meadow. The meadow vegetation cover is about 50 percent, dominated by Astragalus kentrophyta Gray. Using a combination of cover and frequency as a measure of importance, dominance-diversity curves were constructed for the moraine and meadow. Both approach geometric series, which are suggested as indicating harsh environments. Abiotically pollinated species are more successful on the moraine than biotically pollinated species, but the reverse is true for the meadow. Propagule trapping studies suggest that dispersal of anemochorous propagules onto the mo- raine is very low compared with dispersal in the meadow. Relatively little is known about the struc- ture and dynamics of alpine vegetation in the Teton Range, Grand Teton National Park, Wyoming. In this paper we report some pre- liminary quadrat and propagule trapping studies from alpine vegetation in the south- em fork of Cascade Canyon in the center of the range. The selected study area is a complex of al- pine meadow and morainal deposits at the head of the south fork of Cascade Canyon, about 4 km southwest of the Grand Teton (Fig. 1). An east-facing cliff called The Wall bounds the study area on the west; this for- mation forms part of the hydrographic divide of the Teton Range. A shallow cirque has been carved into this cliff by the Schoolroom Glacier, so called because of the almost per- fect end moraine fronting it (Fig. 2). The age of this moraine is unknown, but similar de- posits elsewhere in the range are of Neo- glacial age, which places the moraine age from about 100 to 3,000 years (Mahaney 1975, Mahaney and Spence 1983). Between this moraine and the glacier lies a small melt- water lake about 50 m across, which is drained by a stream that has cut through the center of the moraine. The two ends of the moraine merge into extensive talus derived from The Wall to either side of the glacier. Northeast of the glacier is a large outcrop of gneiss and schist of Precambrian age, which is vegetated by a mosaic of fen and tim- berline krummholz stands. To the east and southeast lies an extensive, slightly undulat- ing dry alpine meadow underlaid by glacial and talus deposits of Pinedale or older age. The Wall and Schoolroom Glacier moraine are composed primarily of Death Canyon Limestone of the Gros Ventre Formation (Cambrian), with some debris of the Wolsey Shale member of the Gros Ventre Formation, in addition to Flathead Sandstone (Cambrian, Love, and Reed 1968, Reed 1973). Climate data from the Teton Range and park are summarized elsewhere (Spence 1981). Mean annual temperature from Jack- son Hole to the east of the range, at an eleva- tion of 2,040 m, is about 1.3 C. Using the ele- vation of the study area, 3,060 m, and the adiabatic lapse rate (Cole 1975), average an- nual temperature in front of the Schoolroom Glacier would be about -4.8 C. Snow depth during the winter is unknown, but it is usual- ly gone from the area by late June to early July during an average year. Common ani- mals at the site include marmots {Marmota flaviventris), pikas {Ochotona princeps), and Rosy Finches {Leucosticte atrata). The purposes of this paper are (1) to char- acterize and contrast the vegetation on the 'Department of Botany, University of British Columbia, Vancouver, B.C., Canada V6T 1W5. 'Department of Biology, Utah State University, UMC45, Logan, Utah 84322. 483 484 Great Basin Naturalist Vol. 43, No. 3 3 N 42' rpn glacial moraine I'^I^I glacier/snowbank lake '' fen it^ Legend stream geologic unit major topographic barrier g-mo gneiss-quartz monzonite qt glacial/periglacial deposits qa alluvial deposits ewf Wolsey Shale edc Death Canyon Limestone A peak Fig. 1. A sketch map of the study area at the head of the south fork of Cascade Canyon. Only major features are shown on the map. The boundaries for the geologic units are only approximate. The map is derived from 7..5 minute uses topographic maps. Love and Reed (1968), Reed (1973), and personal observations of the senior author (JRS). See the legend for map details. Schoolroom Glacier moraine and in the dry alpine meadow adjacent to it, and (2) to sug- gest possible dynamics between the two communities. Methods and Materials Initial site observations and collections were made in the summer of 1978. In 1979, July 1983 Spence, Shaw: Schoolroom Glacier Plants 485 Fig. 2. Schoolroom Glacier at the head of the south fork of Cascade Canyon, t.rand ieton i\ational Park, Wyoming. the quadrat data were gathered on 19 Sep- tember. Two transects, each 90 m long, were run from the base of the proximal slope of the moraine adjacent to the meltwater lake, up to and over the moraine crest, down the distal slope, and out into the meadow. At 3 m intervals, a 0.5 X 0.5 m quadrat was placed down on alternating sides of the transect line. Cover and presence of all species was noted visually, using a modified Braun-Blanquet scale, as follows: + = 0-1 percent, 1 = 1-5 percent, 2 = 5-25 percent, 3 = 25-50 per- cent, 4 = 50-75 percent, 5 = 75-100 per- cent. For calculating the total and average cover values, the midpoints of the ranges were used. Prominence Values (PV) were cal- culated using the formula PV = percent cov- er X the square root of percent frequency. Along the transect lines, slope was measured using an Abney level. Propagule trapping was studied in two ways, by water-filled plastic trays 40 X 15 X 5 cm in size, and 15 X 15 cm wooden plates coated with petroleum jelly. The wa- ter trays were used in 1978, with six of them placed in a line from the edge of the melt- water lake up and over the moraine to the meadow. In 1979, nine wooden plates were used, and were placed in a line 10 m apart from the edge of the lake up and over the moraine and out into the meadow. Once in July and once in September the wooden plates were in operation, for a total of 38 hours in July and 24 hours in September. This amounts to a total of 558 trap-hours in oper- ation. Trapped propagules were placed in glass vials for later identification. A reference set of propagules from species at the site was made to aid in identification. Specimens col- lected are on deposit at the Moose Herba- rium in Grand Teton National Park and the Intermountain Herbarium at Utah State Uni- versity (UTC). Nomenclature follows Shaw (1976). Results and Discussion Forty-six species of vascular plants were collected from the moraine and adjacent meadow in 1978 and 1979. Details on the Great Basin Naturalist Vol. 43, No. 3 floristics and comparisons with other glacial moraine sites in the Tetons will be published elsewhere (Spence 1983). One species, Ta- raxacum lyratum Ledeb., is a new report for the park. The distribution of life forms of the species is: 1 shrub, 1 fern ally, 1 annual dicot, 11 graminoids, and 32 biennial/perennial forbs. Of these species, 31 were encountered along the two transects (Table 1). The re- maining species are quite rare at the site, many of them consisting of only a few indi- vidual plants. Along the first transect, average cover was 0.6 percent for the moraine proximal slope, 8.9 percent for the distal slope, and 47.4 per- cent for the meadow. For the second tran- sect, the respective values were 2.8 percent, 9.2 percent, and 63.3 percent. Cover values per quadrat ranged from 0 percent (7 times; 6 on the moraine proximal slope, 1 on the moraine distal slope, and 0 in the meadow) to 106.5 percent (once, in the meadow). Pooling the results of the two transects, total cover on the moraine is 1.7 percent for the proximal slope and 9.1 percent for the distal slope. Av- erage cover for the meadow is 54.6 percent. These values are all significantly different from one another at a = 0.05. Data from the two transects are summarized in Table 1 for the moraine and the meadow. Although most of the species encountered in the transects are found on both the mo- raine and in the meadow, they tend to be much more common on one or the other (see Table 1). General observations elsewhere on the moraine and in the meadow tend to sup- Table 1. The quadrat data from the two Hne transects in summarized form. Each transect was 90 m long, with a 0.5 X 0.5 m quadrat placed at every 3 m interval, for a total of 60 quadrats and 15 m^. The species are arranged alphabetically, and three numbers are listed for each species; percent frequency, average percent cover, and promi- nence value, which is calculated as average percent cover X square root of percent frequency. For details on tran- sect placement and method of measuring species, see Methods and Materials. Moraine (N = 38) Meadow (N = 22) Species (percent frequency/average percent cover/prominence value) Achillea millefolium Agmpyron caninuvi A. scribneri Antennaria umbrinella Arabis h/allii Arenaria nutiallii Arnica longifolia Astragalus kentrophyta Carex species Cirsium tweedyi Cymoptertts hendersonii Epilobium alpiniim Erigeron compositus E. leiomerus Erysimum asperum Festuca ovina Hymenoxys grandiflora Oxyria digymt Phacelia sericea Poa alpina P. pattersonii Polemonitim viscosurn Salix arctica Selaginelki densa Senecio fremontii Silene acaulis Solidago multiradiata Taraxacum lyratum T. officinale Townsendia montana Trisetum spicatum Unknown grasses Unknown herbs 10.5/0.54/1.75 2.6/0.07/0.11 7.9/0.09/0.25 2.6/0.01/0.02 2.6/0.45/0.73 26.3/0.92/4.72 7.9/0.04/0.11 18.4/0.63/2.70 7.9/0.14/0.39 2.6/0.01/0.02 2.6/0.01/0.02 7.9/0.14/0.39 15.8/0.24/0.95 18.4/0.58/2.49 23.7/0.12/0.58 2.6/0.39/0.63 5.3/0.08/0.18 5.3/0.03/0.07 10.5/0.05/0.16 7.9/0.14/0.39 7.9/0.14/0.39 42.1/0.70/4.54 18.4/0.09/0.39 4.5/0.68/1.44 22.7/4.25/20.25 45.5/4.82/32.51 13.6/0.07/0.26 13.6/0.07/0.26 68.2/28.20/232.S 9.1/0.14/0.42 4.5/0.68/1.44 4.5/0.02/0.04 13.6/0.34/1.25 4.5/0.68/1.44 18.2/0.09/0.38 4.5/0.02/0.04 72.7/4.20/35.81 9.1/0.14/0.42 40.9/3.59/22.96 4.5/0.68/1.44 9.1/1.82/5.49 4.5/0.68/1.44 4.5/1.70/3.61 31.8/0.43/2.42 22.7/0.11/0.52 9.1/0.14/0.42 27.3/1.64/8.57 50.0/4.36/30.83 July 1983 Spence, Shaw: Schoolroom Glacier Plants 487 port the conclusions drawn from the transect data. Only a few species, such as Poa patter- sonii, Agropyron caninum, and Taraxacum officinale, appear to be equally common in both areas. Furthermore, a group of species appears to be restricted to an area at the base of the distal moraine slope. They are found in a band ranging from 1 m up to 10 m wide be- tween the moraine slope and the meadow proper. This band is formed primarily of debris derived from slumping and sliding off the distal moraine slope. Some of the species that were found in this ecotonal region in- clude Anemone multifida, Draba loncho- carpa, Oxytropis deflexa. Taraxacum lyratum, Androsace septentrionalis, Castilleja sul- phurea, and Eritrichium nanum. Few of these species were found on the moraine, and they were all either rare or absent from the mead- ow. The transect data show that this ecotonal region is richer in species than either the mo- raine or meadow. Average number of species per quadrat ranged from 4 on the moraine, to 5.1 in the meadow, to 7.3 in the ecotone be- tween the two. Using prominence values (PV), the most important species on the moraine are Cirsium tweedyi, Epilobium alpinum, Polemonium viscosum, Agropyron caninum, and Poa pat- tersonii. In the meadow the most important species are Astragalus kentrophyta, Hyme- noxys grandiflora, Antennaria umbrinella, Poa pattersonii, and Agropyron caninum. Us- ing the contribution of each species PV to the total summed PV for the moraine and the meadow, dominance diversity curves can be constructed (Fig. 3). Both curves approach geometric series, which have been suggested to indicate harsh environments in which do- minance by one or a few species is strong (Whittaker 1975). On the moraine, several species share dominance; these are Cirsium tweedyi (28 percent of the total summed PV's), Epilobium alpinum (16 percent), Pole- monium viscosum (15 percent), and Agropy- ron caninum (10 percent). In the meadow. Astragalus kentrophyta dominates (63 per- cent). Other species include Hymenoxys grandiflora (10 percent), Antennaria umbri- nella (9 percent), and Poa pattersonii (6 per- cent). Of the eight species listed above, the majority are either western North American alpine species {P. viscosum, A. kentrophyta. Moraine Meadow 0.1% Species Rank Fig. 3. Dominance-diversity curves for the moraine and the meadow. The measure of importance used was each species contribution to the total summed promi- nence values in percent. There are 21 species in the mo- raine curve and 23 species in the meadow curve. All species with values less than 0.1 percent had 0.1 percent added to their value to include them on the graph. This tended to slightly flatten the two curves at the bottom. 488 Great Basin Naturalist Vol. 43, No. 3 P. pattersonii, and A. umbrinella) or south/central Rocky Mountain alpine species (C. tweedyi and H. grandi flora). Of the other two, Epilobium alpinum (sensu lato) is a widespread circumpolar arctic-alpine species and Agropyron caninum is a North American boreal-montane species. Using information derived from Fryxell (1957), Swales (1979), and Ostler and Harper (1978), the species from the transects can be divided into two groups, biotically and abiotically pollinated species. Wind-polli- nated, suspected autogamous and apomictic species (i.e., Arabis lyallii. Taraxacum offici- nale, and Epilobium alpinum), and the fern ally Selaginella densa, which requires water for fertilization, are included in the abioti- cally pollinated group. Biotically pollinated (primarily entomophilous) dicot species com- prise the biotically pollinated group. Average number of abiotically pollinated species per quadrat on the moraine is 1.0, in the meadow 1.4. These are not significantly different at a = 0.05. Average number of biotically polli- nated species per quadrat on the moraine is 1.2, in the meadow 3.1. These are significant- ly different at a = 0.05 (using a t-test for un- equal sample sizes). These and other com- parisons between these two groups of species are summarized in Table 2. The total Table 2. Comparisons between the moraine and the meadow using the quadrat data derived from the hne transects. Two groups of species, abiotically pollinated (the abiotic group) and biotically pollinated (biotic group) are compared for the two areas. See the text for details on the two groups of species. The sample size (N) is the number of quadrats. Moraine (N = 38) Meadow (N = 22) Average percent cover Abiotic 2.5 10.7 Biotic 3.0 44.5 Average number of species per quadrat Abiotic 1.0 1.4 Biotic 1.2 3.1 2) Prominance values (PV) Abiotic 11.0 54.9 Biotic 6.4 343.1 Number of species Abiotic 8 9 Biotic 13 14 Total number of species Abiotic 12 Biotic 19 summed PV for all the moraine species in the abiotically and biotically pollinated groups are 11.0 and 6.4, respectively. For the mead- ow they are 54.9 and 343.9, respectively. The ratio of PV for the abiotically pollinated spe- cies in the meadow and the moraine is 5:1, and that for the biotically pollinated species is 53:1. The total number of species on the moraine and in the meadow for the abioti- cally pollinated group are 8 and 9, respec- tively. For the biotically pollinated group the values for the moraine and meadow are 13 and 14, respectively. All these comparisons suggest that the species in the abiotically pol- linated group are relatively more succesful at colonizing and establishing on the moraine deposits than the species in the biotically pol- linated group. The reverse is true for the meadow, where the biotically pollinated spe- cies dominate. There could be several reasons for these differences. Perhaps the open na- ture of the vegetation on the moraine makes it more difficult for biotically pollinated spe- cies to attract pollinators. Thus species that are autogamous, apomictic, or anemophilous may be at a reproductive advantage. It is also possible that such breeding systems are linked with other traits that confer greater colonizing abilities than is found in the bioti- cally pollinated species (Jain 1976). Differen- tial dispersion of propagules onto the mo- raine by species in the two categories does not appear to be the reason (see below). Turning to the propagule trapping results, the water trays caught only five propagules during several weeks of operation. The water in the trays tended to evaporate quickly, and many propagules were probably blown out of the trays once they had dried out. These five propagules and those trapped by the petro- leum-jelly-coated wooden plates used in 1979 are listed in Table 3. No propagules were trapped by the wooden plates in July, pre- sumably because the plants were still flower- ing and had yet to set seed. In September, the trap furthest out into the meadow (30 m) trapped 23 propagules over a 24-hour period. The only other trap that caught anything was placed on the distal moraine slope in about midslope position. This trap caught a single unidentifiable composite achene with a pap- pus. Extrapolating from the data on number of propagules trapped and the size of the July 1983 Spence, Shaw: Schoolroom Glacier Plants 489 plates, the 23 propagules trapped in the meadow represent about 1022 propagules dispersing into and through every 1 m^ every 24 hours, at a time in September when most species had finished flowering and were dis- persing propagules. The calculated value for the trap on the moraine is 45 propagules per 1 m^ per 24 hours, assuming that the single propagule trapped is representative of the propagule rain on the deposits. Most of the propagules (90 percent) have some mor- phological feature that might aid in more ef- ficient wind dispersal. Such features include the coma of the Salix and Epilobium seeds and the pappus of the composite achenes. Two propagules, the Carex achene and the Astragalus kentrophyta legume, have no ap- parent features that could enhance wind dis- persal ability. The propagule of Geum rossii represents a special case. This species is the only one that did not occur in the vicinity of the moraine or meadow. The propagule con- sists of a persistent calyx with numerous stiff hairs, enclosing many small achene fruits. It was found in one of the water trays in 1978, which raises the possibility that the prop- agule could have accidently dropped into the water from the fur of an investigating pika. Pikas are known to collect plants of Geum rossii for their hay piles (Johnson 1967). All the propagules trapped consist of dicot spe- cies except for the achene of the Carex species. The moraine deposits appear to be in an active state of collapse. Fresh slumps and old slump scars can be found on both slopes, and are especially common on the proximal slope. The deposits are very loose and tend to shift easily underfoot. Numerous small ero- sion channels (rills) exist, attesting to the ef- fects of snow meltwater erosion. The steep- ness of the slopes, which are from 36° to 38° on the distal slope and 35° to 41° on the pro- ximal, contributes to the instability of the moraine. The combination of continual dis- turbance and apparently low rates of dis- persal of propagules from the meadow is the probable explanation for the low average vegetation cover on the moraine slopes. The only portions of the moraine where the plant cover is as dense as in the meadow is along the crest, which in places is flat and presum- ably more stable than the steep slopes. The patches of vegetation on the flat portions of the crest are very similar to the meadow veg- etation, including the presence of the three most common meadow species. Astragalus kentrophyta, Hymenoxys grandiflora, and An- tennaria umbrinella. This suggests that, as the moraine deposits stabilize, they will become vegetated by the species that dominate and characterize the meadow. A perusal of alpine vegetation literature from the south central Rocky Mountains failed to reveal any reports of vegetation sim- ilar to that found in the meadow (Rydberg 1914, Cox 1933, Cain 1943, Hayward 1952, Ellison 1954, Langenheim 1962, Johnson and Billings 1962, Bamberg and Major 1968, Ha- beck 1969, Bonham and Ward 1970, Lewis 1970, Anderson et al. 1979, Komarkova 1979), although many of the species in the meadow are common and widespread in the Rocky Mountains. On the other hand, several reports list vegetation that is strongly similar to that of the moraine (Buttars 1914, Ma- haney 1974, Given and Soper 1975). In a de- tailed study from the Colorado Front Range, Komarkova (1979) listed several species that are characteristic of scree, talus, and glacial deposits (her Order Aquilegio-Cirsietalia sco- pulorum). Many of these species are also found at the Schoolroom Glacier moraine. These include Senecio fremontii, Poa patter- sonii, Oxyria digyna, Draba lonchocarpa, Epilobiujn alpinum. Taraxacum officinale, and Trisetum spicatum. The thistle Cirsium Table 3. The identity of the propagules trapped dur- ing 1978 and 1979 are hsted, along with the number caught, and the presence of any morphological feature that might aid in more efficient wind dispersal. Number Morphological Species trapped feature Epilobium alpinum (seeds) 8 coma Salix arctica (seeds) 9 coma Arnica longifolia (achenes) 3 pappus Geum rossii (achenes enclosed in calyx) 1 hairy calyx Solidago multiradiata (achene) 1 pappus Astragalus kentrophyta (legume) 1 — Carex species (achene) 1 — Unknown composites (achenes) 5 pappus 490 Great Basin Naturalist Vol. 43, No. 3 scopulonim appears to play an ecological role similar to Cirsium tweedyi in the Tetons. In summary, the vegetation on the School- room Glacier moraine is very open, with av- erage cover ranging from less than 1 percent up to 9 percent, compared with over 50 per- cent in the alpine dry meadow adjacent to it. Dominance-diversity curves using promi- nence values as the measure of importance were constructed for the meadow and the moraine. Both curves approach geometric series, suggesting harsh environments in which dominance by one or a few species is strong. The legume Astragalus kentrophyta dominates the meadow, and the thistle Cir- sium tweedyi dominates the moraine. Using cover and frequency data, abiotically polli- nated species are relatively much more suc- cessful on the moraine than biotically polli- nated species. The reverse is true in the meadow. Propagule trapping studies suggest that wind dispersal onto the moraine is very low compared with wind dispersal within the meadow. Distributionally, three groups of species can be discerned. One group is con- centrated primarily on the moraine, only rarely straying into the meadow. The second group is found primarily in the meadow. The third group consists of species that are found in a narrow band between the base of the dis- tal moraine slope and the meadow. This band of vegetation has many of the characteristic of an ecotone. The moraine deposits are highly unstable, and, in the few places along the crest of the moraine that tend to be the most stable, the vegetation is strongly similar to the vegetation in the meadow. Acknowledgments This paper represents partial requirements for a M.Sc. thesis by JRS, under the super- vision of RJS. We thank the National Park Service, particularly Linda Olson and Bob Wood, for collecting permits and for other help. Literature Cited Anderson, D. C, R. S. Hoffman, and K. B. Armitage. 1979. Aboveground productivity and floristic structure of a high subalpine herbaceous mead- ow. Arctic and Alpine Res. 11:467-476. Bamberg, S. A., and J. Major. 1968. Ecology of the veg- etation and soils associated with calcareous par- ent materials in three alpine regions of Montana. Ecol. Monogr. .38:127-167. BoNHAM, C. D., AND R. T. Ward. 1970. Phytosociolog- ical relationships in alpine tufted hairgrass (Des- champsia caespitosa [L.] Beauv.) meadows. Arctic and Alpine Res. 2:267-275. BuTTARs, F. K. 1914. Some peculiar cases of plant distri- bution in the Selkirk Mountains, British Colum- bia. Minnesota Bot. Studies 4:313-331. Cain, S. A. 1943. Sample-plot technique applied to al- pine vegetation in Wyoming. Amer. J. Bot. 30:240-247. Cole, F. 1975. Introduction to meterology. 2d ed. Wiley & Sons, New York. Cox, C. F. 1933. Alpine plant succession on James Peak, Colorado. Ecol. Monogr. 3:300-372. Ellison, L. 1954. Subalpine vegetation of the Wasatch Plateau, Utah. Ecol. Monogr. 24:89-184. Fryxell, p. a. 1957. Mode of reproduction of higher plants. Botanical Rev. 23:135-233. Given, D. R., and J. H. Soper. 1975. Pioneer vegetation on moraines near Clachnacudainn Snowfield, British Columbia. Syesis 8:349-354. Habeck, J. R. 1969. A gradient analysis of a timberline zone at Logan Pass, Glacier Park, Montana. Northwest Sci. 43:65-73. Hayward, C. L. 1952. Alpine biotic communities of the Uinta Mountains, Utah. Ecol. Monogr. 22:93-120. Jain, S. K. 1976. The evolution of inbreeding in plants. Ann. Rev. Ecol. Syst. 7:469-495. Johnson, D. R. 1967. Diet and reproduction of Colorado pikas. J. Mammol. 48:311-315. Johnson, P. L., and W. D. Billings. 1962. The alpine vegetation of the Beartooth Plateau in relation to cryopedogenic processes and patterns. Ecol. Monogr. 32:105-135. Komarkova, v. 1979. Alpine vegetation of the Indian Peaks Area, Front Range, Colorado Rocky Moun- tains. Flora et Vegetatio Mundi, 7. R. Tiixen, ed. Cramer, Vaduz. Langenheim, J. H. 1962. Vegetation and environmental patterns in the Crested Butte area, Gunnison County, Colorado. Ecol. Monogr. 32:249-285. Lewis, M. E. 1970. Alpine rangelands of the Uinta Mountains. U.S. Forest Service— Region 4. Love, J. D., and J. C. Reed. 1968. Creation of the Teton Landscape. Grand Teton Natural Hist. Assoc. Press. Mahaney, W. C. 1974. Soil stratigraphy and genesis of Neoglacial deposits in the Arapaho and Hender- son cirques, central Colorado Front Range. Geogr. Monogr. 5:197-240. 1975. Soils of post-Audubon age, Teton Glacier area, Wyoming. Arctic and Alpine Res. 7:141-153. Mahaney, W. C, and J. R. Spence. 1983. Late Qua- ternary deposits, soils, chronology, and floristics. Jaw Cirque area, Central Teton Range, Wyom- ing. In preparation. Ostler, W. K., and K. T. Harper. 1978. Floral ecology in relation to plant species diversity in the Wasatch Mountains of Utah. Ecology 59:848-861. July 1983 Spence, Shaw: Schoolroom Glacier Plants 491 Reed, J. C. 1973. Geological Map of Grand Teton Na- tional Park. U.S. Geol. Surv. Rydberg, p. a. 1914. Phytogeographical notes on the Rocky Mountain region III. Formations in the al- pine zone. Bull. Torrey Bot. Club 41:459-474. Shaw, R. J. 1976. Field guide to the vascular plants of Grand Teton National Park and Teton County. Utah State Univ. Press, Logan. Spence, J. R. 1981. Comments on the cryptogam vegeta- tion in front of glaciers in the Teton Range. Bryo- logist 84:564-568. 1983. A floristic analysis of the vegetation on Neoglacial deposits in the Teton Range, Wyom- ing. Submitted to Arctic and Alpine Res. Swales, D. E. 1979. Nectaries of certain arctic and sub- arctic plants with notes on pollination. Rhodora 81:367-407. Whittaker, R. H. 1975. Communities and Ecosystems. 2d ed. Macmillan, New York. WINTER STOMACH CONTENTS OF SOUTH DAKOTA BADGERS E. Blake Hart' and Michael Truinbo- Abstract.— Stomach contents of 33 hunter and trapper killed badgers, Taxidea taxtis, from northeastern South Dakota were examined during the 1980-81 fur season. Mammal prey species made up the bulk of the ingested food, followed by plant materials. Approximately 40 percent of stomach contents were mammals, 35 percent were of plant origin, 10 percent were birds, 10 percent were insects, while the remainder was mostly inorganic materials. Little significant difference was found in variety of food items consumed by each sex. South Dakota badgers are opportu- nistic foragers in the wintertime when food is scarce. Individual food items are usually taken in quantity when en- countered by badgers. The severity of typical northern plains winters fixes food as a priority item in the lives of nonhibernating carnivorous mam- mals. Answers to the question as to varieties of prey consumed was sought by examination of a ubiquitous carnivore, the American badger, Taxidea taxus. Stomachs of 33 badgers (15 male, 18 fe- male) from the northeastern South Dakota counties of Faulk, Brown, Spink, and Ed- munds were obtained from a local fur broker during the 1980-1981 season. Several keys were used to identify prey animal hairs, par- ticularly Moore, Spence, and Dugnole (1974). Cuticular scale patterns of hairs were pressed into a film of nail polish and observed (Wein- gart 1973). A variety of materials were found in stom- achs, including plants, inorganic materials (soil, stones), insects, and prey animals. Stom- achs contained an average of 98 grams of food materials each. From the standpoint of relative volume and variety of identifiable re- mains, 40 percent were mammal prey ani- mals, 35 percent were of plant origin, 10 per- cent were birds, 10 percent were insects, and the remainder were mostly undigested in- organic materials. Similar foods were found in stomachs of both sexes with the exception that straw and small stones were not found in female stomachs. At least one-half of all badger carcasses purchased by the fur dealer had empty stom- achs. This suggests that either stomach con- tents had largely been digested during long trap stays or that many badgers spend consid- erable time with empty stomachs. Most of the food matter contained within the stomachs had been thoroughly crushed and mascerated; the largest bone fragment was 1.7 X 0.8 cm. In fact, skull remnants of mammal prey were difficult to identify. In several instances portions of hair-covered epidermis of larger prey were present; these were the largest pieces of food material seen, although occasional intact mouse viscera and other assorted prey extremities were observed. Sunflower seeds, corn, and millet were present in several stomachs in considerable quantities, though these too had mostly been chewed into a paste, with few kernels intact. We observed what possibly may have been several masses of feces. The majority of stom- achs contained from 5 to 20 Ascaris-\ike parasites. We found, as did Jense (1968), who studied badger food habits and energy utilization in east central South Dakota, that badgers are opportunistic foragers of edible plant and an- imal materials. Inasmuch as they feed on whatever is at hand, variety is often lacking, especially if the prey is a large animal or is plant material. One markedly distended stomach contained 375 grams of plant and animal materials. Full stomachs contained little variety; rather, they were often replete with a single food type. 'Department of Mathematics, Natural Sciences and Health Professions, Northern State College, Aberdeen, South Dakota 57401. 'Tulare, South Dakota 57476. 492 July 1983 Hart, Trumbo: Badger Diet 493 Jense (1968) suggested that grains were im- portant fall foods, that birds and eggs were eaten only during the spring and summer. We found that squirrels, mice, rabbits, and other small mammals formed much of the typical winter badger diet, but also evidence was foimd of birds, insects, and grains. Snead and Hendrickson (1942) found many of the same food items in diets of Iowa badgers; they found that percentages of kinds of foods taken changed as availability changed with the seasons. Hibernating prey animals appear to be at- tractive food sources throughout the winter, especially ground squirrels. During the mid- winter 1978-1979 fur season, 17 toads were found in a single badger stomach. Rosen- weig's (1966) statement that there is a clear trend for larger predators to seek larger prey appears true enough, although, judging from the quantities of mice in stomachs, badgers obviously do not ignore such small mammals as important auxiliary food sources. In summary, winter diets of South Dakota badgers vary with the prey species, which represent most nonaquatic vertebrate groups, as well as insects. Opportunistic feeding ap- pears to be common among badgers, which is not unexpected considering sparse food re- sources during the severe cold of typical northern plains winters. Literature Cited Jense, G. K. 1968. Food habits and energy utilization of badgers. Unpublished thesis. South Dakota State Univ., Brookings. 39 pp. Moore, T. D., L. E. Spence, and C. E. Dugnole. 1974. Identification of the dorsal gviard hairs of some mammals of Wyoming. Wyoming Fish and Game Dept. Bull. 14. 177 pp. Rosenweig, M. L. 1966. Community structure in sym- patric carnivora. J. Mammal. 47:602-612. Snead, F., and G. O. Hendrickson. 1942. Food habits of the badgers fn Iowa. J. Mammal. 23:380-391. Weingart, E. L. 1973. A simple technique for revealing hair scale patterns. Amer. Midi. Nat. 90(2):508-509. A LIST OF UTAH SPIDERS, WITH THEIR LOCALITIES Dorald M. Allred' and B. J. Kaston- Abstract. — The 621 species of spiders known to occnr in Utah as recorded in the Hterature or Utah universities' collections are listed with their junior synonyms and collection localities. Two-fifths (265 species) are known from onlv one locality each, and only one-fifth (123 species) from five or more localities in the state. Little is known of the distribution or eco- logical relationships of Utah spiders. Each of 265 species of the 621 recorded for the State is known from only one locality. Even the ubiquitous black widow, Latrodectus hes- perus, has been recorded from only 24 sites in Utah. Most collections from the 249 local- ities, listed primarily from the literature and some unpublished data, are from a few places that seem to have been favorite or conven- ient collecting areas for early naturalists and students of arachnology. For example, 166 species are recorded from Box Elder County in the northwest comer of Utah, principally in the Raft River Mountains, which apparent- ly was a favorite place of Wilton Ivie, a con- temporary and associate of Ralph Chamber- lin and Willis Gertsch. All three men were associated with the University of Utah in Salt Lake City, and 166 species are listed from that environs. Two naturalists and avid col- lectors who were contemporaries and associ- ates of Chamberlin were Vasco Tanner and Angus Woodbury. Both had family ties in St. George in Washington County in the south- west comer of the state and lived and fre- quently revisited there for many years. One hundred species are listed for St. George, and 80 from nearby Zion National Park, where Woodbury was employed as a naturalist for several years. Chamberlin frequently visited Tanner in St. George, and collected in that area. Such large numbers of species recorded from these localities is indicative of the lack of study done in other areas of the state, fur- ther exemplified by the fact that each of only 123 species is known from five or more localities. Much of our knowledge of Utah spiders was contributed by Ralph Chamberlin, who authored or coauthored the naming of 220 of the species listed for Utah. Wilton Ivie and Willis Gertsch authored 106 and 73 species, respectively, some in coauthorship with Chamberlin. Stanley Mulaik, arthropodolo- gist, naturalist, and avid collector, was also contemporary with these men and coau- thored several species with Gertsch. We are indebted to Willis Gertsch, who provided some unpublished records and valu- able criticism of the manuscript. James Mac- Mahon of Utah State University also provid- ed unpublished records of specimens collected by him and his students. Anne Bond, research assistant, initially helped with much of the literature search. Sources of collection records are desig- nated by initials and dates in parentheses im- mediately following specific, or a group of, localities. A key to these is given below. Where more than one publication is repre- sented by an author's initials, the specific source is indicated by a date. Key to Locality Sources (Published articles except as indicated) A = Allred Ac = Archer AG = Allred & Gertsch AU = Allred (unpublished) BU = Beck (unpublished) BA = Beedlow & Abraham Br = Brady BS = Bowling & Sauer C = Chamberlin CG = Chamberlin & Gertsch CI = Chamberlin & Ivie 'Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. 'Department of Zoology, San Diego State University, San Diego, California 92182. 494 April 1983 Allred, Kaston: Utah Spiders 495 cw = Chamberlin & Woodbury D = Dondale DR = Dondale & Redner E = Exline G = Gertsch GI = Gertsch & Ivie GR = Gertsch & Russell GU = Gertsch (unpublished) GW = Gertsch & Wallace H = Hatley I = Ivie lU = Ivie (unpublished) K = Knowlton Ks = Kaston KsU = Kaston (unpublished) Ky = Keyserling L = Levi LD = Lowrie & Dondale Le = Leech LL = Levi & Levi LR = Levi & Randolph M = Millidge McU = MacMahon (unpublished) MG = Munia & Gertsch P = Platnick PS = Platnick & Shadab R = Roewer S = Schick SP = Sauer & Platnick V = Van Helsdingen W = Waagen WE = Wallace & Exline List of Utah Collection Localities with County Designation Allen Cvn (Rich) Alta (Salt Lake) American Fork Cyn (Utah) Aquarius Plateau (Garfield) Aspen Grove (Utah) Bear Lake (Rich) Beaver (Beaver) Beaver Crk (in 12 counties) Beaver Cyn (Beaver) Beaver Dam Mts (Washington) Beaver Dam Wash (Washington) Benson (Cache) Bicknell (Wayne) Big Cottonwood Cyn (Salt Lake) Big Indian Rock (San Juan) Bills Cyn (Salt Lake) Black Rock (Salt Lake) Blanding (San Juan) Blue Sprvice Cyn (Garfield) Bluff (San Juan) Boulder Mt (Garfield) Bountiful (Davis) Brigham (Box Elder) Brigham Cyn (Box Elder) Brigham Plains Flat (Kane) Bryce Cyn Nat Park (Garfield) Butterfield Cyn (Salt Lake) Caineville (Wayne) Calf Crk (Garfield) Carter Crk (Daggett) Carter & Deep Crk Jet (Daggett) Castle Cliffs (Washington) Castle Dale (Emery) Castle Park (?) Cedar Cyn (Iron) Cedar Hills (Box Elder) Cedar Jet (Box Elder) Cedar Mt (Juab) Cedar Mts (in Uinta Mts — Summit?) Chalk Crk (Summit) City Crk Cyn (Salt Lake) Clarkston (Cache) Clear Crk (Box Elder) Clear Crk Cyn (Box Elder) Clear Lake (Millard) Cobble Rest (Wasatch) Cove Fort (Millard) Coyote Gulch (Garfield) Curlew Valley (Box Elder) Cyclone Lake (Garfield) Daniels (Wasatch) Deep Crk (Daggett) Delta (Millard) * Devils Cyn (San Juan) Diamond Valley (Washington) Dinosaur Nat Mon (Uintah) Dixie Nat Forest (Iron) Dove Crk (Box Elder) Dry Cyn (Salt Lake) Duchesne (Duchesne) East Cyn (Salt Lake) Egg Island (Tooele) Elk Ridge (San Juan) Elsinore (Sevier) Emery (Emery) Emigration Cyn (Salt Lake) Emory (Summit) Ensign Peak (Salt Lake) Escalante (Garfield) Eureka Cyn (Utah) Farmington (Davis) Farmington Cyn (Davis) Ferron (Emery) Ferron Res (Sanpete) Fillmore (Millard) Fillmore Cyn (Millard) Fish Lake (Sevier) Fisher Pass (Tooele) Four-mile Bench (Kane) Franklin Basin (Cache) Fremont River (Wayne) Fruita (Wayne) Ft Douglas (Salt Lake) Garden City (Rich) Glen Cyn City (Kane) Glenwood (Sevier) Government Crk (Tooele) Grand Gulch (San Juan) Grandaddy Lake (Duchesne) Granite (Salt Lake) Grantsville (Tooele) Green Cyn (Cache) Green Lake (Daggett) 496 Great Basin Naturalist Vol. 43, No. 3 Greenriver (Emery) Grouse Crk (Box Elder) Gunnison Butte (Emery) Hanksville (Wayne) Hat Island (Tooele) Hatch (Garfield) Heber (Wasatch) Helper (Carbon) Henry Mts (Garfield) Hidden Lake (Kane & Summit) Holliday (Salt Lake) Horse Valley (Wayne) Hughes Cyn (Salt Lake) Hurricane (Washington) Junction (Piute) Kaibab Forest (Kane) Kanab (Kane) Kems (Salt Lake) Kelton (Box Elder) Lake Powell (Kane) Laketown (Rich) Lambs Cyn (Salt Lake) La Sal Jet (San Juan) La Sal Mts (Grand) La Sal Pass (San Juan) Layton (Davis) Lehi (Utah) Leidy Peak (Uintah) Levan (Juab) Liberty (Weber) Little Cottonwood Cyn (Salt Lake) Loa (Wayne) Locomotive Spngs (Box Elder) Logan (Cache) Logan Cyn (Cache) Lynn (Box Elder) Lynndyl (Millard) Manila (Daggett) Marysvale (Piute) Marysvale Cyn (Piute) Mill Crk (Summit) Mill Crk Cyn (Salt Lake) Mirror Lake (Duchesne) Moab (Grand) Monroe Cyn (Sevier) Monticello (San Juan) Moroni (Sanpete) Motauqua (Washington) Mounds (Emery) Mt Agassiz (Summit) Mt Ellen (Garfield) Mt Nebo (Juab) Mud Spngs (Emery) Navajo Mt (San Juan) Nipple Bench (Kane) Noton (Wayne) Ogden (Weber) Ogden Cyn (Weber) Ophir (Tooele) Oquirrh Mts (Tooele) Orton (Garfield) Ouray (Uintah) Pangiiitch (Garfield) Paradise (Cache) Park Valley (Box Elder) Parleys Cyn (Salt Lake) Parowan (Iron) Payson (Utah) Pickleville (Rich) Pinecrest (Salt Lake) Pine Cyn (Millard) Pine Spngs (Garfield) Pine Valley (Washington) Pine Valley Mts (Washington) Pink Sand Dunes (Kane) Pintura (Washington) Plain City (Weber) Posey Lake (Garfield) Price (Carbon) Promontory Point (Box Elder) Provo (Utaii) Provo River, Upper (Duchesne) Puffer Lake (Beaver) Raft River (Box Elder) Raft River Mts (Box Elder) Red Butte Cyn (Salt lake) Red Cyn Camp (Garfield) Richardson (Grand) Richfield (Sevier) Rock Island (Utah) Rotary Park (Salt Lake) Salina (Sevier) Saltair Beach (Salt Lake) Salt Lake Airport (Salt Lake) Salt Lake City (Salt Lake) San Rafael (Emery) San Rafael River (Emery) Santa Clara (Washington) Santaquin (Utah) Santaquin Res (Utah) Scipio (Millard) Silver Lake (Salt Lake & Utah) Smith & Morehouse Cyn (Summit) Smokey Mt (Kane) Snow Crk (Sevier) Snow Crk Cyn (Sevier) Snowville (Box Elder) Spring Crk (Carbon) Spring Cyn (Carbon) Spring Lake (Utah) St John (Tooele) St George (Washington) Standardville (Carbon) Steep Crk (?) Straight Wash (Emery) Stockton (Tooele) Strawberry (Wasatch) Swan Lake (Rich) Table Cliff Pass (Garfield) Table Cliff Plateau (Garfield) Teapot Lake (Summit) Terrys Ranch (Washington) Thompsons (Grand) Three Lakes (Garfield & Kane) Tibbet Spng (Kane) Timpanogos Cave Nat Mon (Utah) Timpanogos Park (Utali) Tooele Cyn (Tooele) Torrey (Wayne) Tremonton (Box Elder) April 1983 Allred, Kaston: Utah Spiders 497 Tropic (Garfield) Tropic Res (Garfield) Trout Crk City (Juab) Uinta Mts (in 5 counties) Utah Lake (Utah) Utah State Univ School Forest (Rich) Valley City (San Juan) Verdure (San Juan) Vermillion Castle (Iron) Vernal (Uintah) Vernon (Tooele) Wah Wah Mts (Beaver) Wanship (Summit) Wasatch (Salt Lake & Summit) Wasatch Mts (Salt Lake) Wasatch Plateau (in 6 counties) White River (Uintah) Widtsoe (Garfield) WiUard (Box Elder) Willow Crk (Tooele) Willow Tank Spngs (Kane) Yost (Box Elder) Zion Nat Park (Washington) Species and Localities If the name associated with a specimen in a collection or a record in the literature is not foimd in this list of species, consult the list of "Synonymies of Utah Records" in the latter part of this report. Adiaearanea ambera Levi 1963 (Bull. Mus. Comp. Zool. 128:204). Mill Crk Cyn (L63). Achaearanea canionis (Chamberlin & Gertsch) 1928 (J. Ent. Zool. Pomona Coll. 21:103). American Fk Cyn, Pinecrest (lU), Beaver Cyn, Cobble Rest, Dinosaur Nat Mon, Dry Cyn, Richfield, Salt Lake City, St George (L55), Zion Nat Park (CG28). Achaearanea tepidarioritm (C.L. Koch) 1841 (Die Arachn. 8:75). HoUiday, Provo, Salt Lake City (L55). Actinoxia sp. Glen Cyn City (AU). Aculepeira carbonaria (L. Koch) 1869 (Zeits. Ferd. Tirol. Voral. 15:58). St George (CW). Aculepeira packardi (Thorell) 1875 (Kongl. Svenska Vet. Akad. Handl. 13:3-203). Green Cyn (H), Kelton Pass (K), USU School Forest (W). Agelenopsis aperta (Gertsch) 1934 (Amer. Mus. Novitates 726:24). Aspen Grove (BU), Motauqua (CI41), Salt Lake City, Trout Crk, Zion Nat Park (G34c), St George (CW). Agelenopsis californica (Banks) 1896 (J. New York Ent. Soc. 4:88). Dove Crk, Grouse Crk (City), Lynn, Raft River S fk, Yost (CI33), St George (CW). Agelenopsis Oklahoma (Gertsch) 1936 (Amer. Mus. Novitates 852:12). Duchesne (CI41). Agelenopsis iitahana (Chamberlin & Ivie) 1933 (Bull. Univ. Utah Biol. Ser. 2(2):43). Clear Crk (CI33a), Fish Lake, La Sal Mts, Wasatch Mts (CI41). Agelenopsis sp. Nipple Bench (AU). Agroeca ornata Banks 1892 (Proc. Acad. Nat. Sci. Philadelphia 44:23). Clear Crk, Raft River S fk (CI33). Agroeca pratensis Emerton 1890 (Trans. Conn. Acad. Sci. 8:155). Raft River S fk(CI33). Agroeca trivittata (Keyserling) 1887 (Verb, zool. bot. Ges. Wien 37:444). Fillmore (C19), Moab (CG28), St George (CW). Alopecosa gulosa (Walckenaer) 1837 (Hist. Nat. Ins. Apt., 1:38). Clear Crk, Yost (CI33), La Sal Jet (CG28), St George, Zion Nat Park (CW). Alopecosa kochi (Keyserling) 1877 (Verb, zool. bot. Ges. Wien 26:636). Green Cyn (H), Posey Lake, Steep Crk, Table Cliff Pass, Three Lakes (N Kanab) (BU), USU School Forest (W). Amaurobius americanus (Emerton) 1888 (Trans. Conn. Acad. Sci. 7:443). Bluff, Fruita, Moab, Monticello, Mounds, San Rafael River, Verdure (CI28), Clear Crk (in Raft River Mts), Dove Crk, Grouse Crk, Lynn (CI33), Lake Powell (CW). Anacornia microps Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2):29). Clear Crk, Raft River S fk, Uinta Mts (CI33). Anacornia proceps Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:483). Chalk Crk, Cobble Rest, Mirror Lake (C48a), USU School Forest (W). Antrodiaetns hageni (Chamberlin) 1917 (Bull. Mus. Comp. Zool. 61:74). Raft River S fk (CI33). Antrodiaetns montanus (Chamberlin & Ivie) 1935 (Bull. Univ. Utah Biol. Ser. 2(8):4). Green Cyn (H), USU School Forest (W). Anyphaena californica (Banks) 1904 (Proc. Calif. Acad. Sci. 3:338). Mill Crk (C20b). Anyphaena pacifica (Banks) 1896 (Trans. Amer. Ent. Soc. 23:63). Green Cyn (H), Ver- dure (CG28). Anyphaena sp. Clear Crk, Dove Crk, Raft River S fk (CI33), Escalante, Three Lakes (N Kanab) (BU). 498 Great Basin Naturalist Vol. 43, No. 3 Aphonopelma angusi Chamberlin 1940 (Bull. Univ. Utah Biol. Ser. 5(8):21). W of Beaver Dam Mts (C40). Aplwnopelma simulatiim (Chamberlin & Ivie) 1939 (Bull. Univ. Utah Biol. Ser. 5(1):8). Fruita (CI39a). Aphonopelma zionis Chamberlin 1940 (Bull. Univ. Utah Biol. Ser. 5(8) :24). Zion Nat Park (C40). ApollopJianes texanus Banks 1904 (J. New York Ent. Soc. 12:113). Blanding, Bluff, Moab, Verdure (CG28), Glen Cyn City 5 km W (AG). Aranetis gemma (MacCook) 1888 (Proc. Acad. Nat. Sci. Philadelphia 50:193). Clear Crk, Lynn, Park Valley (in Raft River Mts) (CI33), Ferron, Salt Lake City (CI35b), Green Cyn (H), Locomotive Spngs (K), Pro- vo, Three Lakes (nr Escalante) (BU), Zion Nat Park (CW). Araneus mammatus (Archer) 1951 (Amer. Mus. Novitates 1487:17). Mill Crk Cyn (L81a). Araneus marmoreus Clerck 1757 (Aranei Suecici, p. 29). Chalk Crk (in Uinta Mts) (C19). Araneus nordmanni (Thorell) 1870 (Rem. Syn. Europ. Spid., p. 4). USU School Forest (W). Araneus pima Levi 1971 (Bull. Mus. Comp. Zool. 141:176). "Utah" (L71). Araneus saevus (L. Koch) 1872 (Zeits. Ferd. Tirol Voralberg (3) 17:323). Salt Lake City (Ac). Araneus trifolium (Hentz) 1847 (J. Boston Soc. Nat. Hist. 5:471). Provo (BU). Araneus varians Thorell 1899 (Bih. Svenska Vet. Akad. Handl. 25:49). Zion Nat Park (CW). Araneus sp. Coyote Gulch (BU). Araniella displicata (Hentz) 1847 (J. Bos- ton Soc. Nat. Hist. 5:476). Aspen Grove (BU), Clear Crk (CI33), Green Cyn (H), USU School Forest (W). Araniella octopunctata Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1) :76). Emigration Cyn, Pine Vallev, Zion Nat Park (CI42a), Steep Crk (BU). Arctosa alpigena (Doleschall) 1852 (Sitz.- ber. Akad. Wiss. Wien, 9:643). USU School Forest (W). Arctosa chamberlini Gertsch 1934 (Amer. Mus. Novitates 693:4). Vernal (G34). Arctosa littoralis (Hentz) 1844 (J. Boston Soc. Nat. Hist. 4:388). Calf Crk, Coyote Gulch (in Escalante Basin), La Sal Mts, Rock Island (in Utah Lake), Steep Crk, Three Lakes (nr Kanab), Willow Tank Spngs (BU), Grand Gulch, Utah Lake (G34), Greenriver, Moab (CG28), St George (CW). Arctosa parva (Banks) 1894 (J. New York Ent. Soc. 2:52). "Utah" (C8). Arctosa rubicunda (Keyserling) 1877 (Verh. zool. bot. Ges. Wien 26:663). Raft River S fk (CI33). Arcuphantes decoratus Chamberlin & Ivie 1943 (Bull. Univ. Utah Biol. Ser. 7(6): 17). Zion Cyn (Nat Park) (CI43). Arcuphantes fragilis Chamberlin & Ivie 1943 (Bull. Univ. Utah Biol. Ser. 7(6): 17). American Fork Cyn, Timpanogos Nat Mon (CI43). Argenna obesa Emerton 1911 (Trans. Conn. Acad. Sci. 16:399). Utah Lake W shore (CI35b), Zion Nat Park (CG58). Argenna saphes Chamberlin 1948 (Bull. Univ. Utah Biol. Ser. 10(6):6). Mirror Lake (C48). Argenna sp. Raft River S fk (CI33). Argennina reclusa Gertsch & Ivie 1936 (Amer. Mus. Novitates 858:1). Cove Fort 10 mi N (GI36). Argiope trifasciata (Forskal) 1775 (Descr. Anim., p. 86). Green Cyn (H), Kelton, Kelton Pass, Snowville (K), Salt Lake City (BU), St George (CW). Ariadna bicolor (Hentz) 1842 (J. Boston Soc. Nat. Hist. 4:225). Lake Powell (CW), Verdure (CG28). Aysha gracilis (Hentz) 1847 (J. Boston Soc. Nat. Hist. 5:452). Zion Nat Park (CW). Aysha incursa (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:239). St. George (CW). Bathyphantes concolor (Wider) 1834 (In: Reuss, Zool. Misc. Mus. Senck. 1:267). Provo, Salt Lake City (169). Bathyphantes latescens (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:248). Aspen Grove, Logan Cyn, Mill Crk Cyn, Ogden River Cyn (169), Chalk Crk (in Uinta Mts) (C19), Clear Crk, Grouse Crk, Raft River S fk (CI33). Bathyphantes pullatus (O. Pick. -Cam- bridge) 1863 (Zoologist 21:8580) Cobble Rest Camp, Uinta Mts, Salt Lake City (169). Bathyphantes sp. USU School Forest (W). Batroceps sp. Kelton (K). April 1983 Allred, Kaston: Utah Spiders 499 Brachybothritim montaniim Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8):4). Provo River N fk (in Uinta Mts), Raft River Mts, Salt Lake City (CI35). Calilena gertschi Chamberlin & Ivie 1941 (Ann. Ent. Soc. Amer. 34:612). Monticello (CI41). Calilena multiformis dixiana Chamberlin & Ivie 1941 (Ann. Ent. Soc. Amer. 34:608). Diamond Valley, Pintura, St George (CI41). Calilena restricta Chamberlin & Ivie 1941 (Ann. Ent. Soc. Amer. 34:606). Aquarius Plateau, Cedar Mt, Escalante, Eureka Cyn, Ferron, Loa, Mt Ellen, Noton, Panguitch, Price, Raft River Mts, Richfield, Standard- ville, Utah Lake W side, Zion Nat Park (CI41), Locomotive Spngs (K), Tibbet Spng 2 km NE (AG), Widtsoe (BU). Calilena sp. Calf Crk, Cyclone Lake (nr Escalante), Steep Crk, Three Lakes (nr Ka- nab) (BU), Smokey Mt (AU). Callilepis eremellus Chamberlin 1928 (Proc. Biol. Soc. Washington 41:77). Caine- viUe (CG28). Callilepis zionis Chamberlin & Woodbury 1929 (Proc. Biol. Soc. Washington 43:133). Zion Nat Park (CW). Callilepis sp. Kelton, Kelton Pass (K), Four- mile Bench, Smokey Mt, Tibbet Spng 2 km NE (AU). Callobiiis nevadensis (Simon) 1884 (Bull. Soc. Zool. France 9:318). American Fk Cyn, City Crk Cyn, Dry Cyn, Logan, Logan Cyn, Mill Crk Cyn, Timpanogos Park (Le), Fill- more (C19), Green Cyn (McU), Hughes Cyn, Ogden, Ogden Cyn, Wasatch Mts (C47). Callobius nomeus (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:240). Cedar Mts, Chalk Crk (in Uinta Mts), Ferron Res, Mirror Lake, Upper Provo River (C47), Cobble Rest, Grand Daddy Lake, La Sal Pass, Mill Crk Cyn, Ouray (Le), USU School Forest (W), Wasatch Plateau (CI47a). Castianeira aurata (Hentz) 1847 (J. Boston Soc. Nat. Hist. 5:459). Raft River S fk (CI33). Castianeira descripta (Hentz) 1847 (J. Bos- ton Soc. Nat. Hist. 5:456). Zion Nat Park (CW). Castianeira longipalpa (Hentz) 1847 (J. Boston Soc. Nat. Hist. 5:457). "Utah"(DR82). Castianeira occidens Reiskind 1969 (Bull. Mus. Comp. Zool. 138:211). Brigham Plains Flat, Four-mile Bench (8 km SE cow camp at head Wesses Cyn) (AG), Green Cyn (H), USU School Forest (W). Castianeira zionis Chamberlin & Wood- bury 1929 (Proc. Biol. Soc. Washington 42:139). Zion Nat Park (CW). Catabrithorax clypiellus Chamberlin 1920 (Canad. Ent. 52:199). Bear Lake, Logan Cyn (C20), Clear Crk, Dove Crk, Raft River S fk. Raft River Mts (CI33). Catabrithorax plwnosus (Emerton) 1882 (Trans. Conn. Acad. Sci. 6:53). Fish Lake, Moab, Price (C48). Catabrithorax stylifer Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:486). Clear Crk, Dove Crk, Lynn 8 mi S, Raft River S fk (C48). Catabrithorax iitus (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:253). Clear Lake (C19), Cyclone Lake (nr Escalante), Posey Lake (BU), Dove Crk, Raft River S fk, Yost (CI33). Ceraticelus crassiceps Chamberlin & Ivie 1939 (Verb. 7 intern. Kongr. Ent. 1:68). Mir- ror Lake (CI39). Ceraticelus subniger Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:491). Salt Lake City 10 mi W (C48). Ceraticelus sp. Curlew Valley (K). Ceratinella acerea Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2):26). Raft Riv- er S fk, Wasatch Mts (CI33). Ceratinella brunnea Emerton 1882 (Trans. Conn. Acad. Sci. 6:36). Clear Crk, Dove Crk (CI33). Ceratinops uintana Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:508). Cobble Rest, Mirror Lake (C48). Ceratinops watsinga Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:516). Salt Lake City (C48). Ceratinops sp. Clear Crk (CI33). Cesonia gertschi Platnick & Shadab 1980 (Bull. Amer. Mus. Nat. Hist. 165:352). Zion Nat Park (PS80). Cesonia sincera Gertsch & Mulaik 1936 (Amer. Mus. Novitates 851:10). Smokey Mt (AG). Cheiracanthium inclusum (Hentz) 1847 (J. Boston Soc. Nat. Hist. 5:451). Calf Crk, Ka- nab, Kanab Cyn, Three Lakes (N Kanab) (BU), Green Cyn (H), St George, Zion Nat Park (CW). 500 Great Basin Naturalist Vol. 43, No. 3 Cheiracanthitim rnildei L. Koch 1864 (Abh. naturh. Ges. Nurnberg 1864:144). Provo (A80), Brigham (McU). Chrosiothes chirica (Levi) 1954 (Trans. Amer. Microscop. Soc. 73:184). Utah (LR). Chrysso nordica (ChamberHn & Ivie) 1947 (Ann. Ent. Soc. Amer. 40:29). Utah (LR). Chrysso pelyx (Levi) 1958 (Psyche 64:104). Salt Lake City (L58). Cicurina deserticola Chamberlin & Ivie 1940 (Bull. Univ. Utah Biol. Ser. 5(8) :65). Bicknell, Bluff, Perron, Henry Mts, Pintura, S Price, Santa Clara, Thompsons, Tropic (CI40), Glen Cyn City 5 km SW, Nipple Bench, Smokey Mt (14 km from Last Chance Jet) (AG). Cicurina intermedia Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2) :46). Clear Crk, Henry Mts, Raft River S fk, Uinta Mts, Wasatch Mts (CI33), Fish Lake, Salt Lake City (E), Green Cyn (H). Cicurina parma Chamberlin & Ivie 1940 (Bull. Univ. Utah Biol. Ser. 5(9):67). Bryce Cyn Nat Park (CI40). Cicurina rohusta Simon 1886 (Ann. Ent. Soc. Belg. 30:40). Bluff (CG28), Chalk Crk (in Uinta Mts) (C19), Clear Crk, Henry Mts, Raft River S fk (CI33), Perron, Pish Lake, La Sal Mts, Liberty, Mirror Lake, Provo River N fk. Raft River Mts, Smith & Morehouse Cyn, Verdure, Wasatch Mts (CI40), Pine Cyn (C13), Steep Crk, Widtsoe (BU), USU School Porest (W). Cluhiona abbottii Koch 1866 (Arach. Fam. Drassiden., p. 303). City Crk Cyn, Utah Lake W side (G41), Clear Crk (CI33), Moab (CG28). Cluhiona canadensis Emerton 1890 (Trans. Conn. Acad. Sci. 8:181). Logan (McU). Cluhiona mimula Chamberlin 1928 (Proc. Biol. Soc. Washington 41:184). Clear Crk (CI33), Pruita (CG28). Cluhiona maesta Banks 1896 (Trans. Amer. Ent. Soc. 23:64). Chalk Crk (in Uinta Mts) (C19), Clear Crk, Dove Crk, Park Valley (City), Raft River S fk. Raft River Mts (CI33). Cluhiona mutata Gertsch 1941 (Amer. Mus. Novitates 1184:14). Salt Lake City (G41). Cluhiona norvegica Strand 1900 (Kong. Norske Vidensk. Selsk. Skr., p. 30). "Utah" (DR82). Cluhiona pacifica Banks 1896 (Trans. Amer. Ent. Soc. 23:65). Zion Nat Park (CW). Cochlemholus provo Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:522). Cobble Rest (C48). Coreogonal hicornis (Emerton) 1923 (Ca- nad. Ent. 55:242). USU School Porest (W). Coriarachne hrunneipes Banks 1893 (J. New York Ent. Soc. 1:133). Raft River S fk (CI33), NE Utah (BS). Coriarachne versicolor Keyserling 1880 (Spinn. Amer. Lat. 1:53). Benson (G53), Bluff, Salt Lake City, Zion Nat Park (G32), Clear Crk (CI33), Greenriver, Moab (CG28), Heber (McU). Crustulina sticta (O. Pick. -Cambridge) 1861 (Ann. Mag. Nat. Hist., ser. 3, 7:432). Moab, Price, San Rafael River (CG28), Salt Lake City, St George (lU), Uinta Mts (L57). Ctenium eremophilus (Chamberlin) 1928 (Proc. Biol. Soc. Washington 41:180). Devils Cyn (CG28), Verdure (Ks46). Ctenium fusca (Emerton) 1894 (Trans. Conn. Acad. Sci. 9:407). Locomotive Spngs (K). Ctenium vigerens (Chamberlin & Ivie) 1933 (Bull. Univ. Utah Biol. Ser. 2(2):9). Dove Crk, Raft River S fk, Yost (CI33), Raft River Mts, Salt Lake City, Verdure (Ks46). Cybaeota concolor Chamberlin & Ivie 1937 (Ann. Ent. Soc. Amer. 30:211). Mill Crk Cyn (CI37). Cybaeota wasatchensis Chamberlin & Ivie 1937 (Ann. Ent. Soc. Amer. 30:211). Hughes Cyn, Mill Crk Cyn (CI37). Cyclosa conica (Pallas) 1772 (Spicil. Zool. 1(9):48). Clear Crk (CI33), Logan SE (McU), Zion Nat Park (CW). Delopelma iodius Chamberlin & Ivie 1939 (Bull. Univ. Utah Biol. Ser. 5(1):6). Castle Cliffs 2 mi W, Zion Nat Park (CI39a). Delopelma marxi (Simon) 1891 (Acta Bull. Soc. Linn. Bordeaux 44:321). Pruita (G35). Delopelma melanius Chamberlin & Ivie 1939 (Bull. Univ. Utah Biol. Ser. 5(1):5). Brig- ham, Salt Lake City, Stockton, and Davis and Utah counties (CI38a). Delopelma steindachneri (Ausserer) 1875 (Verb. zool. bot. Ges. Wien 25:199). St George (C21). Dendryphantes (prob is Metaphidippus) mylothrus Chamberlin 1925 (Proc. Calif. Acad. Sci. (4)14:134). Mill Crk Cyn (C25). April 1983 Allred, Kaston: Utah Spiders 501 DendrypJiantes (prob is Metaphidippus) uteanus Chamberlin & Gertsch 1929 (J. Ent. Zool. Pomona Coll. 21:110). Clear Crk, Raft River S fk (CI33), Lambs Cyn, Zion Nat Park (CG29). Dictyna abundans Chamberlin & Ivie 1941 (Bull. Univ. Utah Biol. Ser. 6(3):5). St George (CI41a), Zion Nat Park (CG58). Dictyna apacheca Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8):28). Ameri- can Fk Cyn (CG58). Dictyna artemisia Ivie 1947 (New York: priv. publ.). Provo River N fk, Scipio, Wasatch Mts (nr Salt Lake City) (CG58). Dictyna bellans Chamberlin 1919 (Ann. Ent. Soc. Amer. 12:242). Moab (CG58). Dictyna bicornis Emerton 1915 (Trans. Conn. Acad. Sci. 20:141). Salt Lake City (G46). Dictyna borealis cavernosa Jones 1947 (Field & Lab. 15:12). Salt Lake City (CG58). Dictyna brevitarsus Emerton 1915 (Trans. Conn. Acad. Sci. 20:140). USU School Forest (W). Dictyna calcarata Banks 1904 (Proc. Calif. Acad. Sci. 3:342). Calf Crk (BU), St George, Zion Nat Park (CG58). Dictyna cholla Gertsch & Davis 1942 (Amer. Mus. Novitates 1158:12). Salt Lake City (CG58). Dictyna completa Chamberlin & Gertsch 1929 (J. Ent. Zool. Pomona Coll. 21:101). Green Cyn (H), Moab (CG28). Dictyna coloradensis Chamberlin 1919 (Ann. Ent. Soc. Amer. 12:241). Utah (C58). Dictyna cornupeta Bishop & Ruderman 1946 (Proc. Biol. Soc. Washington 59:1). Salt Lake City (CG58). Dictyna horta Gertsch & Ivie 1936 (Amer. Mus. Novitates 858:4). City Crk Cyn, Layton (CG58). Dictyna idahoana Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2) :4). Cedar Hills, Kelton, Kelton Pass, Snowville (K), Green Cyn (H), Grouse Crk (CI33), Mt Nebo (CG58). Dictyna littoricolens Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8):30). Black Rock, Salt Lake City (CG58), Locomo- tive Spngs 3 mi E (K), Utah Lake W side (CI35). Dictyna major Menge 1869 (Schr. naturf. Ges. Danzig 2:247). Farmington Cyn (LL). Dictyna minuta Emerton 1888 (Trans. Conn. Acad. Sci. 7:447). Price (CG58), Salt Lake City (GI36). Dictyna moaba Ivie 1947 (New York, priv. publ.). Moab (C48). Dictyna pallida Keyserling 1887 (Verb, zool. bot. Ges. Wien 37:472). Richfield (CG58). Dictyna personata Gertsch & Mulaik 1936 (Amer. Mus. Novitates 851:9). East Cyn, Hur- ricane (CG58), Glen Cyn City (AG), Glen- wood 2 mi E (G46). Dictyna pictella Chamberlin & Gertsch 1958 (Bull. Amer. Mus. Nat. Hist. 116:97). Pintura, Scipio (CG58). Dictyna piratica Ivie 1947 (New York, priv. publ.). Dry Cyn (CG58), East Cyn, Pro- vo River N fk, Wanship (C48). Dictyna reticulata Gertsch & Ivie 1936 (Amer. Mus. JVovitates 858:7). Curlew Valley (K), Richfield, Santa Clara 4 mi SW (GI36). Dictyna secuta Chamberlin 1924 (Proc. Calif. Adad. Sci. 12:583). St George (CG58). Dictyna stulta Gertsch & Mulaik 1936 (Amer. Mus. Novitates 851:7). Aspen Grove, Ogden Cyn (G46). Dictyna terrestris Emerton 1911 (Trans. Conn. Acad. Sci. 16:399). Bill's Cyn, Delta, Dry Cyn, Mill Crk Cyn (CG58). Dictyna tertanea Ivie 1947 (New York, priv. publ.). American Fork Cyn, Bluff, Castle Park, Dinosaur Nat Mon, Greenriver, Henry Mts, Richfield, Salt Lake City (CG58). Dictyna tridentata Bishop & Ruderman 1946 (Proc. Biol. Soc. Washington 58:2). Elk Ridge (CG58). Dictyna tucsona Chamberlin 1948 ( Bull. Univ. Utah Biol. Ser. 10(6):8). Kanab (CG58). Dictyna uintana Chamberlin 1919 (Ann. Ent. Soc. Amer. 12:240). Bluff (CG28), Chalk Crk (in Uinta Mts) (C19), City Crk Cyn (C48), Clear Crk, Grouse Crk, Raft River Mts, Raft River S fk, Yost (CI33), Elsinore, Fish Lake, Richfield, Salt Lake City (G46). Dictyna variana Chamberlin & Gertsch 1958 (Bull. Amer. Mus. Nat. Hist. 116:57). St George (CG58). Dictyna vincens Chamberlin 1919 (Ann. Ent. Soc. Amer. 12:243). Clear Crk, Raft Riv- er S fk. Raft River Mts (CI33). Dictyna volucripes Keyserling 1882 (Verb, zool. bot. Ges. Wien 31:286). Central & northern Utah, St George (C21), La Sal Mts (C28). 502 Great Basin Naturalist Vol. 43, No. 3 Dictyna zaha Barrows & Ivie 1942 (Ohio J. Sci. 42:21). City Crk Cyn, Layton, Richfield (CG58). Dictyna sp. Coyote Gulch, Posey Lake, Steep Crk, Table Cliff Plateau, Willow Tank Spngs (BU). Dictynina eutypa (Chamberlin & Ivie) 1929 (J. Ent. Zool. Pomona Coll. 21:101). Bluff (CG28). Dictynoides sp. Zion Nat Park (CW). Digiietia canities (McCook) 1890 (Amer. Spiders 2:136). St George (G35), Zion Nat Park (CW). Dipoena atopa (Chamberlin) 1948 (Ann. Ent. Soc. Amer. 41:541). City Crk Cyn, Dry Cyn, Timpanogos Nat Mon (L53), Grantsville (L63), Red Butte Cyn (C48a). Dipoena malkini Levi 1953 (Amer. Mus. Novitates 1647:33). Government Crk, Mill Crk Cyn (L53). Dipoena nigra (Emerton) 1882 (Trans. Conn. Acad. Sci. 6:21). Beaver Cyn, City Crk Cyn, Government Crk, Wanship (L53), Pick- leville (W). Dipoena prona Menge 1868 (Schr. naturf. Ges. Danzig 2:177). Moab (L53). Dipoena provalis Levi 1953 (Amer. Mus. Novitates 1639:34). Cobble Rest, Hughes Cyn (L53), Provo River N fk (GU). Dipoena tibialis Banks 1906 (Proc. Ent. Soc. Washington 7:96). Green Cyn (H), USU School Forest (W). Dipoena sp. Clear Crk, Raft River Mts (CI53). Disembolus alpha (Chamberlin) 1948 (Ann. Ent. Soc. Amer. 41:550). Dry Crk Cyn (C48a). Disemholus anguineus Millidge 1981 (J. Arach. 9:276). Utah (M81a). Disemholus beta Millidge 1981 (J. Arach. 9:274). Dry Cyn (M81a). Disemholus galeatus Millidge 1981 (J. Arach. 9:268). Utah (M81a). Disemholus implexus Millidge 1981 (J. Arach. 9:279). Fish Lake (M81a). Disemholus implicatus Millidge 1981 (J. Arach. 9:277). Cobble Rest, Upper Provo River (M81a). Disemholus kesimbus (Chamberlin) 1948 (Ann. Ent. Soc. Amer. 41:552). Fish Lake, Henry Mts, Strawberry Res (C48a). Disemholus stridulans Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2) :20). Fillmore Cyn, Lynn 8 mi S, Mirror lake, Snow Crk Cyn, Wasatch Mts (nr Salt Lake City) (CI45), Henry Mts, Raft River S fk, (CI33). Disemholus vicinus Millidge 1981 (J. Arach. 9:281). Grantsville (M81a). Dolomedes triton (Walckenaer) 1837 (Hist. Nat. Ins. Apt. 1:340). Raft River Mts (CI33). Drassodes gosiutus Chamberlin 1919 (Ann. Ent. Soc. Amer. 12:245). City Crk Cyn, Dry Cyn, Ensign Peak, Greenriver, Helper, E Monticello, Mud Spngs, Rotary Park, Salt Lake Airport 5 mi SW, Spring Crk, Utah Lake W side. Willow Crk (PS76), Fillmore (C19), Nipple Bench (AG), Zion Nat Park (CW). Drassodes neglectus (Keyserling) 1887 (Verb. zool. bot. Ges. Wien 37:434). Bluff, Marysvale, Verdure (CG28), Fish Lake (C36), La Sal Mts (C28), also counties of Garfield, Iron, Salt Lake, Summit, Tooele, and Utah (PS76). Drassodes saccatus (Emerton) 1890 (Trans. Conn. Acad. Sci. 8:178). Fillmore (C19), Glen Cyn City 5 km SW, Nipple Bench (AG), Green Cyn (H), Raft River Mts (CI33), also counties of Carbon, Emery, Garfield, Grand, Salt Lake, San Juan, Sevier, and Washington (PS76). Drassodes sp. Curlew Valley (K), Glen Cyn City (AU), Steep Crk, Widtsoe (BU). Drassyllus conformans Chamberlin 1936 (Amer. Mus. Novitates 841:22). Richfield, Salt Lake City (C36). Drassyllus dromeus Chamberlin 1922 (Proc. Biol. Soc. Washington 35:169). Dry Cyn, Pinecrest (C36). Drasyllus inanus Chamberlin & Gertsch 1940 (Amer. Mus. Novitates 1068:17). Bluff (CG40). Drassyllus insularis (Banks) 1900 (Canad. Ent. 32:97). Bluff, Salt Lake County, Straight Wash, Valley City (CG28), Green Cyn (H), Grouse Crk (CI33), Lake Powell (C58), St George (CW). Drassyllus lamprus (Chamberlin) 1920 (Ca- nad. Ent. 52:193). Mill Crk (C22), USU School Forest (W). Drassyllus lepidus (Banks) 1899 (Proc. Ent. Soc. Washington 4:190). Pine Valley Mts (C36). Drassyllus mexicanus (Banks) 1898 (Proc. Calif. Acad. Sci (3)1:217). La Sal Mts (CG40). April 1983 Allred, Kaston: Utah Spiders 503 Drassyllus mormon Chamberlin 1936 (Amer. Mus. Novitates 841:27). St George (C36). Drassyllus nannelliis Chamberlin & Gertsch 1940 (Amer. Mus. Novitates 1068:11). Tremonton 10 mi W (CG40), Green Cyn (H). Drassyllus notonus Chamberlin 1928 (Proc. Biol. Soc. Washington 41:179). Noton (CG28). Drassyllus tonaquintus Chamberlin & Gertsch 1940 (Amer. Mus. Novitates 1068:1). Farmington, St George (CG40). Drassyllus sp. Curlew Valley (K). Ebo evansae Sauer & Platnick 1972 (Ca- nad. Ent. 104:41). Butler, Logan, Salt Lake City (SP72), Green Cyn (H). Ebo sp. Locomotive Spngs (K), Raft River S fk (CI33). Enoplognatlm Joshua Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1) :44). Brigham Plains Flat (AG), Hurricane, Pin- tura, Helper, Spring Cyn (CI42a), counties of Emery, Grand, Salt Lake, and Sevier (L57). EnoplognatJia marmorata (Hentz) 1850 (J. Boston Soc. Nat. Hist. 6:273). Dry Cyn, Emi- gration Cyn (lU), Oquirrh Mts, Santaquin (CI42), Park Valley, Raft River Mts, Raft Riv- er S fk (CI33), Sevier County (L57). Enoplognatha ovata (Clerck) 1757 (Aranei Suecici, p. 58). Green Cyn (H). Enoplognatha wyuta Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):43). American Fork Cyn, Hughes Cyn (CI42a), and Emery County (L57). Erigone canthognatha Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 13). Colorado River 5 mi up from Moab (CI35). Erigone denticulata Chamberlin & Ivie 1939 (Verh. 7 intern. Kongr. Ent. Berlin 1:57). Mirror Lake (CI39). Erigone dentosa O. Pick. -Cambridge 1894 (Biol. Centr. Amer. Arach. Ar. 1:128). Clear Crk, Grouse Crk, Raft River Mts, Raft River S fk, Yost (CI33), Green Cyn (H), Posey Lake (BU), USU School Forest (W). Erigone uintana Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 14). Mirror Lake (CI35). Erigone viabilis Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2): 11). Mirror Lake, Raft River S fk (CI33). Eris marginata (Walckenaer) 1837 (Hist. Nat. Ins. Apt. 1:466) Mill Crk Cyn (C25), St George (CW). Eris nigromaculatus (Keyserling) 1884 (Verh. zool. bot. Ges. Wein 34:500) Lambs Cyn (CG29). Ero canionis Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8):23 ). City Crk Cyn (CI35). Eularia chelata Chamberlin & Ivie 1939 (Verh. 7 intern. Kongr. Ent. Berlin 1:61). Mir- ror Lake (CI45). Eularia dela Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2): 15). Clear Crk, Raft River S fk, Raft River Mts (CI33), Lynn 8 mi 5 (CI45). Eularia kaiba Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:530). Mirror Lake (C48). Eularia mana Chamberlin & Ivie 1935 (Bull. Univ. Ufah Biol. Ser. 2(8): 16). Mill Crk Cyn (C45). Eularia schediana Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2): 17). Cedar Mt, City Crk Cyn, Ferron, Fish lake, Henry Mts, Horse Valley, Kaibab Forest, Mirror Lake, Upper Provo River (CI45), Dove Crk (CI33), Raft River Mts (CI45). Euryopis coki Levi 1954 (Amer. Mus. Novi- tates 1666:33). Salt Lake City (L54). Euryopis formosa Banks 1908 (Canad. Ent. 40:206). American Fork, Salt Lake City (CI41), Dry Cyn, Smith & Morehouse Cyn (L54). Euryopis scriptipes Banks 1908 (Canad. Ent. 40:206). Clear Crk, Grouse Crk, Raft River Mts (CI33), Glen Cyn City (AG), Mt Agassiz, Bryce Cyn Nat Park, Jet Deep Crk 6 Carter Crk (in Uinta Mts), Fish Lake, Fruita, Hughes Cyn, Mill Crk Cyn, Trout Crk, Vermillion Castle (nr Parowan), Zion Nat Park (L54), Green Cyn (H). Euryopis spinigera O. Pick. -Cambridge 1895 (Biol. Centr. Amer. Arachn. Ar. 1:146). City Crk Cyn, Junction, Parleys Cyn, Pintura 10 mi N (L54). Euryopis taczanowskii Keyserling 1886 (Spinn. Amer., Theridiidae 2:47). Orton (L54). Euryopis texana Banks 1908 (Canad. Ent. 40:207). Hurricane (L54). Euryopis sp. Brigham Plains (AU), Snow- ville (K), Coyote Gulch, Escalante, Steep Crk (BU). 504 Great Basin Naturalist Vol. 43, No. 3 Evarcha leucophaea (Koch) 1846 (Die Arachn. 13:216). Dove Crk, Raft River S fk (CI33). Filistata hurca Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):3). Hurricane (CI42a). Filistata utahana Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8) :5). Brigham Plains Flat, Glen Cyn City 7 km SW, Tibbit Spng 2 km NE (AG), Marysvale, St George (CI35b). Floricottius littoralis Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 16). Utah Lake W shore (CI35b). Frontinella pyramitela (Walckenaer) 1841 (Hist. Nat. Ins. Apt. 2:261). Green Cyn (H), Washington County, Willow Tank Spngs (BU). Gayenna dixiana Chamberlin & Wood- bury 1929 (Proc. Biol. Soc. Washington 42:138). St George (CW). Geolycosa fatifera (Hentz) 1842 (J. Boston Soc. Nat. His. 4:229). Utah (C9). Geolycosa rafaelana (Chamberlin) 1928 (Proc. Biol. Soc' Washington 41:186). Big In- dian Rock, San Rafael Desert (CG28), Glen Cyn City 5 km SW (AG). Gnaphosa antipola Chamberlin 1933 (Amer. Mus. Novitates 631:4). Farmington, Moab, Salt Lake City (PS75), Utah Lake W shore (C33). Gnapliosa brwnalis Thorell 1875 (J. Boston Soc. Nat. Hist. 17:497). La Sal Mts, Verdure (CG28), Salt Lake City (PS75). Gnapliosa calif ornica Banks 1904 (Proc. Calif. Acad. Sci. (Zool.) 3:335). Four-mile Bench (5 km SW cow camp at head Wesses Cyn) (AG), Ft Douglas, Glenwood, Grant- sville. Hat Island, Manilla, Pintura, Raft Riv- er Mts, Richfield, Salt Lake City, St George, Stockton, Straight Wash, Tooele Cyn, Tre- monton (PS75), Grouse Crk (CI33). Gnaphosa clara (Keyserling) 1887 (Verh. zool. bot. Ges. Wien 37:429). Bridger Basin (C22), Greenriver, Grouse Crk, Moab, Mud Spngs, Wah Wah Mts (PS75). Gnaphosa dentata Platnick & Shadab 1975 (Bull. Amer. Mus. Nat. Hist. 155:18). Utah Lake and Washington County (PS75). Gnapliosa gosoga Chamberlin 1928 (Proc. Biol. Soc. Washington 41:178). Marysvale, Straight Wash (CG28), Tooele Cyn (C36). Gnaphosa hirsutipes Banks 1901 (Proc. Acad. Nat. Sci. Philadelphia 53:573). Fruita, Salt Lake City, San Rafael River, Verdure (CG28), Grouse Crk (CI33), St George (CW). Gnaphosa musco^'uni (L. Koch) 1866 (Arachn. Fam. Drassiden., p. 14). Blue Spruce Camp, Castle Dale, Emigration Cyn, Fish Lake, Fillmore, Hatch, Leidy Peak, Mill Crk, Provo, Richfield, Salt Lake City (PS75), Clear Crk, Raft River Mts, Raft River S fk, Yost (CI33), Curlew Valley, Kosmo (K), La Sal Mts (C28a), Monroe Cyn (C36), Oquirrh Mts (CG28), Rock Island (in Utah Lake) (BU), St George (CW). Gnaphosa salsa Platnick & Shadab 1975 (Bull. Amer. Mus. Nat. Hist. 155:22). Terrys Ranch (Beaver Dam Wash) (PS75). Gnaphosa saxosa Platnick & Shadab 1975 (Bull. Amer. Mus. Nat. Hist. 155:17). Salt Lake City (PS75). Gnaphosa sericata (L. Koch) 1866 (Arach. Fam. Drassiden., p. 31). Green Cyn (H), Lo- comotive Spngs (K), Moab, Price, Salt Lake City (PS75). Gnaphosa synthetica Chamberlin 1924 (Proc. Calif. Acad. Sci. 12:620). St George (PS75). Gnaphosa utahana Banks 1904 (J. New York Ent. Soc. 12:110). Egg Island, Farm- ington NW, Hat Island, Plain City, San Ra- fael River, Silver Lake (PS75). Gnaphosa sp. Brigham Plains (AU), Posey Lake, Provo, Widtsoe (BU). Gnathantes ferosa Chamberlin & Ivie 1943 (Bull. Univ. Utah Biol. Ser. 7(6):5). City Crk Cyn (CI43). Gosiphrurus unicolor Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8):40). Ferron (CI35b). Grammonota salicicola Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:532). Salt Lake City (C48a). Habrocestrum sp. Raft River S fk (CI33). Habronattus sp. Calf Crk, Kanab, Steep Crk (BU). Hahnia cinerea Emerton 1890 (Trans. Conn. Acad. Sci. 8:197). Butterfield Cyn, Mill Crk, Logan Cyn (G34b), Zion Nat Park (CW). Hahnia ononidiwn Simon 1875 (Arach. de France 2:135). Fish Lake, La Sal Mts, Mt El- len, Pine Spngs (in Henry Mts) (CI42a). April 1983 Allred, Kaston: Utah Spiders 505 Haplodrassus hicornis (Emerton) 1909 (Trans. Conn. Acad. Sci. 14:218). Blanding, City Crk Cyn (CG28), Marysvale Cyn (C36a). Haplodrassus dixiensis Chamberlin & Woodbury 1929 (Proc. Biol. Soc. Washington 42:134). St George (CW). Haplodrassus eunis Chamberlin 1922 (Proc. Biol. Soc. Washington 35:162). Cedar Cyn, East Cyn, Emory, Provo River N fk, Smith & Morehouse Cyn, Tropic Res (CI46), Nipple Bench, Smokey Mt (14 & 23 km from Last Chance Jet) (AG), USU School Forest (W). Haplodrassus signifer (Koch) 1839 (Die Arachn. 6:31). Blanding, Marysvale, Moab, Noton, Salt Lake County, Straight Wash (CG28), Cyclone Lake (nr Escalante) (BU), Green Cyn (H), Gunnison Butte (nr Green- river), Moab, Richardson (CG40), La Sal Mts (C28a). Haplodrassus sp. Posey Lake (BU). Helopfiora orinoma (Chamberlin) 1920 (Ca- nad. Ent. 52:195). Bear Lake (C20). Helophora reducta (Keyserling) 1886 (Spinn. Amer. Theridiidae 2:54). Chalk Crk (in Uinta Mts), Clear Lake (C19), Clear Crk, Raft River S fk, Yost (CI33). Helopliora hinagijna Chamberlin & Ivie 1943 (Bull. Univ. Utah Biol. Ser. 7(6):6). USU School Forest (W). Herpyllus cockerelli (Banks) 1901 (Proc. Acad. Nat. Sci. Philadelphia 53:571). Red Cyn Camp (11 mi SE Panguitch) (P). Herpyllus convallis Chamberlin 1936 (Amer. Mus. Novitates 841:2). St George, Zion Nat Park (P). Herpyllus ecclesiastica Hentz 1832 (Amer. J. Sci. 21:102). Bluff, HanksviUe, San Rafael River, Valley City (CG28). Herpyllus hesperolus Chamb^lin 1928 (Proc. Biol. Soc. Washington 42:176). Big In- dian Rock, Bluff, Fruita, HanksviUe, Mary- svale, Moab, San Rafael River, Valley City (CG28), Locomotive Spngs (K), Richfield (C36a), St George, Zion Nat Park (CW), and counties of Carbon, Davis, Duchesne, Mor- gan, Salt Lake, San Juan, Tooele, Utah, and Wayne (P). Herpyllus propinquus (Keyserling) 1887 (Verb. zool. bot. Ges. Wien 37:430). Four- mile Bench (8 km SE cow camp at head Wesses Cyn) (AG), La Sal Mts (CG28), Lynn, Raft River Mts (CI33), St George (CW) and counties of Beaver, Carbon, Duchesne, Mil- lard, Uintah, Utah, and Wayne (P) and San Juan (PS77). Herpyllus sp. Curlew Valley, Kosmo (K), Glen Cyn City (AU), Green Cyn (H). Hillhousia microtarsus (Emerton) 1882 (Trans. Conn. Acad. Sci. 6:57). Mirror Lake, upper Provo River (CI45). Hololena hola (Chamberlin & Gertsch) 1928 (Proc. Biol. Soc. Washington 41:183). Blanding, Devils Cyn, Moab, Noton, Verdure (CG28), Monticello (CI42). Hololena mimoides (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:256). Devils Cyn, Moab, Pine Valley, Pintura, St George, Zion Nat Park (CI42), Fillmore Cyn (C19), Posey Lake (BU). Hololena nevada (Chamberlin & Gertsch) 1929 (J. Ent. Zool. Pomona Coll. 21:107). Beaver Cyn, Moab 5 mi NE, Price, Richfield (CI42). Hololena oquirrhensis (Chamberlin & Gertsch) 1930 (Proc. Biol. Soc. Washington 43:142). Butterfield Cyn (CG30), Oquirrh Mts, Wasatch Mts (CI42). Hybocoptus dentipalpis (Emerton) 1915 (Trans. Conn. Acad. Sci. 20:149). Chalk Crk (in Uinta Mts) (C19). Hypselistes florens (O. Pick. -Cambridge) 1875 (Proc. Zool. Soc. London, p. 403). Smith & Morehouse Cyn (CI35b). Hypselistes reducens Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 17). Mirror Lake (CI35b). Hypsosinga funebris (Keyserling) 1893 (Spinn. Amer. Epeiridae 4:37). Alta (L75). Hypsosinga groenlandica (Thorell) 1872 (Ofvers Kongl. Vet. Akad. Forh. 29:157). Franklin Basin (L75). Hypsosinga pygmaea (Sundevall) 1831 (Kongl. Svenska Vet. Akad. Handl., p. 121). Kanab Cyn (BU), Salt Lake City (L75). Hypsosinga singaefomiis (Scheffer) 1904 (Ent. News 15:259). Green Cyn (H). Hijptiotes gertschi Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 12). Cedar Cyn (nr Cedar City), Henry Mts, Salt Lake City (CI35b). Hyptiotes puebla Muma & Gertsch 1964 (Amer. Mus. Novitates 2196:14). N central Utah (MG). Icius annectans Chamberlin & Gertsch 1929 (J. Ent. Zool. Pomona Coll. 21:110). Zion Nat Park (CG29). 506 Great Basin Naturalist Vol. 43, No. 3 Idionella anomala (Chamberlin & Ivie) 1939 (Verb. 7 intern. Kongr. Ent. Berlin 1:69). Dry Cyn(CI39). Idionella tuganus (Chamberlin) 1948 (Ann. Ent. Soc. Amer. 41:494). Salt Lake City 10 mi W (C48). Lahuella prosaica Chamberlin & Ivie 1943 (Bull. Univ. Utah Biol. Ser. 7(6): 10). Smith & Morehouse Cyn (CI43). Larinia borealis Banks 1894 (Ent. News 5:8). Green Cyn (H), nr Salt Lake City (L75a). Larinia famitlatoria (Keyserling) 1883 (Ver. zool. bot. Ges. Wien 32:201). Central and southern Utah (L75a). Latrodectus hesperus Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 3(1): 15). Ahlstrom Point, Glen Cyn City 6 km SW, Tibbet Spng 2 km NE (AG), Bluff, Moab, Price (CG28), Clear Crk, Grouse Crk, Raft River Mts (CI33), Curlew Valley, Kosmo (K), Dry Cyn, Mill Crk, Logan Cyn, Weber Coimty, Millard County (GU), Green Cyn (H), Provo (A74), St George, Zion Nat Park (CW), Salt Lake City (CI35a), Tooele (L59), Utah Lake, Three Lakes (N Kanab) (BU). Lathys deliculata Gertsch 1946 (Amer. Mus. Novitates 1319:3). Moab, Pintura (CG58). Lathys hesperus Chamberlin 1948 (Bull. Univ. Utah Biol. Ser. 10(6): 14). Utah (C48). Lepthypliantes agressus Chamberlin & Ivie 1943 (Bull. Univ. Utah Biol. Ser. 7(6): 15). Lambs Cyn head. Mirror Lake, Provo River (in Uinta Mts) (CI43). Lepthypliantes arhorea (Emerton) 1915 (Trans. Conn. Acad. Sci. 20:151). Posey Lake (BU), USU School Forest (W). Lepthypliantes calcarata (Emerton) 1909 (Trans. Comi. Acad. Sci. 14:197). USU School Forest (W). Lepthypliantes concolor (Wider) 1834 (In: Reuss, A. Zool. Misc. Arachn., Mus. Senck. 1:267). Provo, Salt Lake City (169). Lepthypliantes furcillifer Chamberlin & Ivie 1933 (Amer. Mus. Novitates 631:32). Clear Crk, Raft River S fk (CI33). Lepthypluintes lamprus Chamberlin 1920 (Canad. Ent. 52:195). Clear Crk, Raft River S fk (CI33), Lake Powell (C58), La Sal Mts (C28), Logan Cyn (C20b), Posey Lake (BU). Lepthypliantes nehulosus (Sundevall) 1829 (Kongl. Svenska Vet. Akad. Handl, p. 218). Lynn (CI33). Lepthypliantes pollicaris Zorsch 1937 (Amer. Midi. Nat. 18:897). USU School For- est (W). Lepthijphantes ranieri Emerton 1926 (Ca- nad. Ent. 58:118). USU School Forest (W). Lepthypliantes sp. Cyclone Lake (nr Esca- lante) (BU). Linyphantes ephedrus (Chamberlin & Ivie) 1933 (Bull. Univ. Utah Biol. Ser. 2(2):31). Dry Cyn, Fish Lake, St George, Yost, Zion Nat Park (CI42a). Linyphia tattphora Chamberlin 1928 (Proc. Biol. Soc. Washington 41:180). Bluff, Zion Nat Park (CG28). Loxosceles deserta Gertsch 1975 (Toxicon 13:203). Southern Utah (GR). Loxosceles rufipes (Lucas) 1834 (Mag. Zool. Guerin 4:354). Clear Crk, Raft River Mts (CI33), San Juan County (CG28), Zion Nat Park (CW). Loxosceles sp. Lake Powell (C58). Lycosa antelucana Montgomery 1904 (Proc. Acad. Nat. Sci. Philadelphia 56:282). St George (C21a). Lycosa carolinensis Hentz 1842 (J. Boston Soc. Nat. Hist. 4:230). Utah (COS). Lycosa frondicola Emerton 1885 (Trans. Conn. Acad. Sci. 6:484). Clear Crk (CI33). Lycosa helluo Walckenaer 1837 (Hist. Nat. Ins. Apt. 1:337). Utah (C8). Lycosa uinticolens Chamberlin 1936 (Proc. Biol. Soc. Washington 49:15). Green Lake (in Uinta Mts) (C36). Lycosa sp. Green Cyn (H), Provo (A78). Mallos alpheus Chamberlin 1948 (Bull. Univ. Utah Biol. Ser. 10(6): 14). American Fork Cyn (C48). Mallos eutyptis (Chamberlin & Gertsch) 1929 (J. Ent. Zool. Pomona Coll. 21:101). Bluff, Monroe Cyn, Richfield, St George (G46). Mallos niveus O. Pick.-Cambridge 1902 (Biol. Centr. Amer., Arachn. Ar. 1:308). Calf Crk, Posey Lake, Three Lakes (nr Kanab), Willow Tank Spngs (BU), Hughes Cyn, Oph- ir, Richfield, Salt Lake City, Zion Nat Park (G46), Hurricane (C48). Mallos trivittatiis (Banks) 1901 (Proc. Acad. Nat. Sci. Philadelphia 53:577). Calf Crk, Ka- nab, Posey Lake, Steep Crk, Three Lakes (N Kanab) (BU), Richfield (G46), USU School Forest (W), Zion Nat Park (C48). April 1983 Allred, Kaston: Utah Spiders 507 Marpissa californica (Peckham) 1888 (Trans. Wisconsin Acad. Sci. 7:81). San Ra- fael (CG28). Mmo perplexa Chamberlin & Ivie 1939 (Verb. 7 intern. Kongr. Ent. Berlin 1:64). Cobble Rest (€145), Provo River N fk (in Uinta Mts) (C139). Meioneta fiUmomna (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:249). Fillmore Cyn (C19). Meioneta sp. Green Cyn (H). Metacyrba taeniola (Hentz) 1846 (J. Boston Soc. Nat. Hist. 5:353). Moab (CG28), Nipple Bench (AG). Metaphidippus aeneolus (Curtis) 1892 (Zoe 3:332). Kelton Pass, Snowville (K), USU School Forest (W). Metaphidippus diplacis Chamberlin 1924 (Proc. Calif. Acad. Sci. (4)12:616). Zion Nat Park (CW). Metaphidippus galathea (Walckenaer) 1837 (Hist. Nat. Ins. Apt. 1:456). La Sal Mts (C28), Raft River S fk (CI33), Zion Nat Park (CW). Metaphidippus helenae (Banks) 1921 (Proc. Calif. Acad. Sci. 11:101). Fish Lake, Richfield (G34c). Metaphidippus paiutus Gertsch 1934 (Amer. Mus. Novitates 726:1). Richfield, St George (G34c). Metaphidippus unicus (Chamberlin & Gertsch) 1930 (Proc. Biol. Soc. Washington 43:143). Uintah County (CG30). Metaphidippus verecundus (Chamberlin & Gertsch) 1930 (Proc. Biol. Soc. Washington 43:144). Clear Crk, Raft River S fk (CI33), Dry Cyn (CG30), Green Cyn (H). Metaphidippus vitis (Cockerell) 1894 (En- tomologist 27:207). Kanab Cyn, Three Lakes (N Kanab) (BU). Metaphidippus sp. Coyote Gulch, Esca- lante, Steep Crk, Table Cliff Plateau, Three Lakes (N Kanab), Widtsoe, Willow Tank Spngs (BU), Green Cyn (H), Nipple Bench (AU). MeteUina mimetoides Chamberlin & Ivie 1941 (Bull. Univ. Utah Biol. Ser. 6(3): 15). Utah (L80). Metepeira foxi Gertsch & Ivie 1936 (Amer. Mus. Novitates 858:20). Curlew Valley (K), Fish Lake, Richfield (GI36), Green Cyn (H). Metepeira grandiose alpina Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):74). Fish Lake (CI42a). Metepeira gosoga Chamerlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8):21). Esca- lante, Steep Crk (BU). Metepeira labyrinthea (Hentz) 1847 (J. Bos- ton Soc. Nat. Hist. 5:471). Clear Crk, Dove Crk, Raft River S fk (CI33). Metepeira nanella Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):71). Fillmore (CI42a). Metepeira sp. Smokey Mt (AU). Micaria ahana Gertsch 1933 (Amer. Mus. Novitates 637:6). Utah (G35). Micaria formicoides Chamberlin & Wood- bury 1929 (Proc. Biol. Soc. Washington 42:139). St George (CW). Micaria gosiuta Gertsch 1942 (Amer. Mus. Novitates 1195:1). City Crk Cyn (G42). Micaria jeanae Gertsch 1942 (Amer. Mus. Novitates 1195:4). Glenwood (G42). Micaria rjiontana Emerton 1890 (Trans. Conn. Acad. Sci. 8:168). Clear Crk, Dove Crk, Lynn (CI33). Micaria mormon Gertsch 1935 (Amer. Mus. Novitates 805:17). City Crk Cyn (G35). Micaria saUna Gertsch 1942 (Amer. Mus. Novitates 1195:5). Salina (G42). Micaria sp. Ahlstrom Point, Four-mile Bench (AU), Green Cyn (H), Kelton, Lo- comotive Spngs (K), USU School Forest (W). MicroHnyphia mandibulata (Chamberlin & Ivie) 1943 (Bull. Univ. Utah Biol. Ser. 7(6):24). Emery, Fillmore, Provo River mth, Scipio, Verdure, Wasatch Mts, Zion Nat Park (CI43). Microneta anopla Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2) :35). Clear Crk, Dove Crk, Grouse Crk, Raft River S fk (CI33). Microneta cornupalpis (O. Pick. -Cam- bridge) 1875 (Proc. Zool. Soc. London, p. 401). Clear Crk (CI33). Microneta fratrella (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:250). Clear Crk (CI33), Uinta Mts (C19). Microneta lophophor Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2) :35). Raft River S fk (CI33). Microneta orines Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2):35). Clear Crk, Raft River S fk (CI33). Microneta protrudens Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2) :37). Clear Crk, Uinta Mts (CI33). 508 Great Basin Naturalist Vol. 43, No. 3 Microneta tumoa Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2):37). Clear Crk (CI33). Microneta uta Chamberlin 1920 (Canad. Ent. 52:196). Logan Cyn (C20b). Microneta viaria (Blackwall) 1841 (Trans. Linn. Soc. London 18:645). Chalk Crk (in Uinta Mts) (C19), Clear Crk, Dove Crk (CI33). Microneta sp. Zion Nat Park (CW). Minietus aktius Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8) :23). Green Cyn (H), Utah Lake W side (CI35b). Mimetus hesperus Chamberlin 1923 (J. Ent. Zool. Pomona Coll. 15:5). Grouse Crk (CI33). Minyriolus plesius Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:538). Cobble Rest (C48a). Misumena vatia (Clerck) 1757 (Aranei Suecici, p. 128). Aspen Grove (BU), Clear Crk (CI33), St George, Zion Nat Park (CW). Misumenoides aleatoria (Hentz) 1847 (J. Boston Soc. Nat. Hist. 6:444). La Sal Mts (C28a). Misumenops asperatus (Hentz) 1847 (J. Boston Soc. Nat. Hist. 5:447). Green Cyn (H). Misumenops californicus (Banks) 1896 (J. New York Ent. Soc. 4:91). Coyote Gulch (BU). Misumenops celer (Hentz) 1847 (J. Boston Soc. Nat. Hist. 5:446). Clear Crk, Raft River S fk (CI33), Curlew Valley (K), Green Cyn (H). Misumenops coloradensis Gertsch 1933 (Amer. Mus. Novitates 635:17). Upper Esca- lante Basin at 10-mile Crk (BU). Misumenops importunus (Keyserling) 1881 (Verb. zool. bot. Ges. Wien 31:307). Cedar Jet, Kelton Pass (K). Misumenops oblongus (Keyserling) 1880 (Spirm. Amer. Lat. 1:79). Three Lakes (N Ka- nab) (BU). Misumenops utanus Chamberlin 1929 (Proc. Biol. Soc. Washington 2:137). Zion Nat Park (CW). Misumenops varia (Keyserling) 1880 (Spinn. Amer. Lat. 1:94). USU School Forest (W). Misumenops sp. Calf Crk, Coyote Gulch, Escalante, Kanab Cyn, Steep Crk, 10-mile Crk (in Escalante Basin), Three Lakes (N Ka- nab), Widtsoe, Willow Tank Spngs (BU). Nanavia monticola Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2):27). Clear Crk (CI33). Neoanagraphis pearci Gertsch 1941 (Amer. Mus. Novitates 1147:19). Ahlstrom Point, Brigham Plains Flat, Four-Mile Bench (5 km SE cow camp head Wesses Cyn), Nipple Bench, Smokey Mt (14 & 23 km from Last Chance Jet) (AG). Neoantistea gosiuta Gertsch 1934 (Amer. Mus. Novitates 712:24). Lynn 8 mi S, Raft River S fk, Yost (G34b), Table Cliff Pass (BU). Neoantistea magna (Keyserling) 1887 (Verh. zool. bot. Ges. Wien 37:463). Raft Riv- er S fk, Yost (CI33a), Spring Lake (G34b). Neoantistea sp. Glen Cyn City, Nipple Bench (AU), USU School Forest (W). Neon nellii G. & E. Peckham 1888 (Trans. Wisconsin Acad. Sci. 7:88). City Crk, Jordan River (nr Salt Lake City) (GI55), Clear Crk (CI33), Verdure (CG28). Neon pixii Gertsch & Ivie 1955 (Amer. Mus. Novitates 1743:15). Salt Lake City (GI55). Neoscona arabesca (Walckenaer) 1841 (Hist. Nat. Ins. Apt. 2:74). Curlew Valley (K), Green Cyn (H), St George (CW). Neoscona benjamina (Walckenaer) 1841 (Hist. Nat. Ins. Apt. 2:42). St George (CW). Neoscona oaxacensis (Keyserling) 1864 (Sitz.-ber. naturw. Ges. Isis, Dresden., p. 121). Saltair Beach (C20a), Washington County (CW), West Mt (A73). Neoscona utahana (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:254). Fillmore (C19). Neriene montana (Clerck) 1757 (Aranei Suecici, p. 64). Clear Crk (CI33), St George, Zion Nat Park (CW). Nodocion eclecticus Chamberlin 1924 (Proc. Calif Acad. Sci. 12:613). Ophir (CG40), Price, Promontory Point, Utah Lake W side, Zion Nat Park (PS80a), Valley City (CG28). Nodocion moronius Chamberlin 1936 (Amer. Mus. Novitates 853:5). Moroni (C36b). Nodocion rufithoracicus Worley 1928 (Ann. Ent. Soc. Amer. 21:620). American Fork Cyn mth, Mill Crk Cyn, Willard (PS80a), Green Cyn (H). Nodocion utus (Chamberlin) 1936 (Amer. Mus. Novitates 841:7). Glen Cyn City 7 km April 1983 Allred, Kaston: Utah Spiders 509 SW (AG), Richfield (C36a), Santaquin 3 mi E, Utah Lake W side (PS80a). Nodocion voluntarius (Chamberlin) 1919 (J. Ent. Zool. Pomona Coll. 12:5). Panguitch 11 mi SE, Red Cyn Camp (PSSOa). Novalena idahoana (Gertsch) 1934 (Amer. Mus. ovitates 726:25). Price, Wasatch (CI42). Novalena lutzi (Gertsch) 1933 (Amer. Mus. Novitates 637:12). Cedar Cyn, Horse Valley (in Henry Mts) (CI42). Nuctenea patagiata (Clerck) 1757 (Aranei Suecici, p. 38). Aspen Grove, St George (BU), Benson (McU), Dove Crk, Grouse Crk, Raft River S fk (CI33), Lake Powell (C58), La Sal Mts (C28a). Oedothorax lasalanus Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 18). La Sal Mts (CI35b). Oedothorax sp. Dove Crk (CI33). Olios fasiculatiis Simon 1880 (Acta Soc. Linn. Bordeaux 34:307). St George (C21), Zion Nat Park (CW). Orchestina moaba Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 10). Bluff, Moab (CI35b). Orchestina utaliana Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 10). Utah Lake W shore (CI35b). Oreonetides vaginatus (Thorell) 1872 (Of- vers. Kongl. Vet. Akad. Forh. 29:153). Utah (V). Orodrassus coloradensis (Emerton) 1877 (Bull. U.S. Geol. Surv. 3:528). Posey Lake, Steep Crk, Widtsoe (BU), Raft River S fk (CI33), USU School Forest (W), Uinta Mts (C19). Orthonops gertschi Chamberlin 1928 (Psyche 35:235). Straight Wash (in San Rafael Desert) (C28). Oxyopes rufipes Banks 1893 (J. New York Ent. Soc. 1:133). lear Crk, Raft River S fk (CI33), Kanab, Kanab Cyn, Escalante (BU), St George (CW). Oxyopes salticus (Hentz) 1845 (J. Boston Soc. Nat. Hist. 5:196). St George (C21). Oxyopes scalaris (Hentz) 1845 (J. Boston Soc. Nat. Hist. 5:196). Green Cyn (H), Lo- comotive Spngs (K). Oxyopes tanneri Chamberlin 1928 (Canad. Ent. 60:95). La Sal Mts (C28a). Oxyopes tridens Brady 1964 (Bull. Mus. Comp. Zool. 131:472). Glen Cyn City 5 km W (AG). Oxyopes sp. Escalante (BU), USU School Forest (W). Ozyptila heaufortensis Strand 1916 (Arch. Naturgesch. 81A(9):124). Smith & Morehouse Cyn (DR74). Ozyptila conspurcata Thorell 1877 (Bull. U.S. Geol. Surv. 3:496). Dry Cyn, Fish Lake, Fruita, Salt Lake City, White River, Zion Nat Park (DR74). Ozyptila sp. Cyclone Lake (nr Escalante) (BU). Pachygnatha dorothea McCook 1894 (Amer. Spiders 3:270). nr Salt Lake City (L80). Pachygnatha mccooki Banks 1916 (Proc. Acad. Nat. Sci. Philadelphia 68:79). Kanab (BU). Pachygnatha xanthostoma C. L. Koch 1845 (Die Arachn. 12:148). Grouse Crk (CI33), & other localities not designated (L80). Pagomys monticola (Gertsch & Mulaik) 1936 (Amer. Mus. Novitates 851:2). Mirror Lake (CG58). Pagomys uinta Chamberlin 1948 (Bull. Univ. Utah Biol. Ser. 10(6): 15). Mirror Lake (C48). Paidisca camano Levi 1957 (Bull. Amer. Mus. Nat. Hist. 112:105). Mill Crk Cyn (L57). Paidisca pallida (Emerton) 1913 (Trans. Conn. Acad. Sci. 18:213). Counties of Salt Lake & Utah (L57). Pardosa altamontis Chamberlin & Ivie 1946 (Bull. Univ. Utah Biol. Ser. 9(5):7). East Cyn (CI46). Pardosa atra Banks 1894 (J. New York Ent. Soc. 2:52). Blanding, Devils Cyn (CG28), Clear Crk (CI33), Salt Lake City, Zion Nat Park (G34), Green Cyn (H), Steep Crk (BU), USU School Forest (W). Pardosa bellona Banks 1898 (Proc. Calif. Acad. Sci. 1:275). Monroe Cyn (G34). Pardosa californica Keyserling 1887 (Verh. zool. bot. Ges. Wien 37:483). Utah (C8). Pardosa coloradensis Banks 1894 (J. New York Ent. Soc. 2:51). Raft River S fk (CI33). Pardosa concinna (Thorell) 1877 (Bull. U.S. Geol. Surv. 3:506). Sevier County (G34). Pardosa distincta (Blackwall) 1846 (Ann. Mag. Nat. Hist. London (1)17:32). Aspen Grove (BU), Beaver Crk (G34), Clear Crk, Dove Crk, Lynn, Raft River S fk, Yost (CI33). Pardosa dorsalis Banks 1894 (J. New York Ent. Soc. 2:51). Several localities not desig- nated (LD81). 510 Great Basin Naturalist Vol. 43, No. 3 Pardosa dorsuncata Lowrie & Dondale 1981 (Bull. Amer. Mus. Nat. Hist. 170:130). Several localities not designated (LD81). Pardosa falcifera F. Pick.-Cambridge 1902 (Biol. Centr. Amer., Arachn. Aran. 2:318). Hurricane, Salt Lake City, Zion Nat Park (G34). Pardosa giebeli (Pavesi) 1873 (Atti Soc. Ital. Sci. Nat. 16:30). Navajo Mt (AG), Steep Crk (BU), Uinta Mts (G34). Pardosa groenlandica (Thorell) 1872 (Oefv. Vet. Akad. Forh. 29:157). Devils Cyn, Green- river, Moab, Monticello, Price, San Rafael River, Valley City, Verdure (CG28), Dove Crk, Grouse Crk, Raft River S fk, Yost (CI33). Pardosa lapidicina Emerton 1885 (Trans. Conn. Acad. Sci. 6:494). Blanding, Bluff, Fniita, Greenriver, Hanksville, Moab, San Rafael River, Straight Wash (CG28), La Sal Mts (C28a), St George, Zion Nat Park (CW). Pardosa mackenziana (Keyserling) 1877 (Verb. zool. bot. Ges. Wien 26:621). Aspen Grove, Washington County, Steep Crk (BU), Clear Crk, Raft River S fk (CI33), La Sal Mts (C28a). Pardosa modica (Blackwall) 1846 (Ann. Mag. Nat. Hist. London (1)17:33). Fish Lake (G34), Utah Lake (C8). Pardosa saxatilis (Hentz) 1844 (J. Boston Soc. Nat. Hist. 4:392). Marysvale Cyn (G34). Pardosa sierra Banks 1898 (Proc. Calif. Acad. Sci. 1:274). Coyote Gulch (in Escalante Basin) (BU). Pardosa solituda Levi & Levi 1951 (Zoo- logica 36:225). Hidden Lake, Mt Timpanogos (LL). Pardosa sternalis (Thorell) 1877 (Bull. U.S. Geol. Surv. 3:504). Blanding, Fruita, Moab, Price, Valley City, Verdure (CW), Clear Crk, Dove Crk, Grouse Crk, Raft River S fk, Yost (CI33), Rock Island (in Utah Lake), Steep Crk, Three Lakes (N Kanab), Kanab (BU), St George (C21), USU School Forest (W). Pardosa tristis (Thorell) 1877 (Bull. U.S. Geol. Surv. 3:510). Cyclone Lake (nr Esca- lante) (BU). Pardosa iiintana Gertsch 1933 (Amer. Mus. Novitates 636:27). Uinta Mts (G33), SW Utah (LD81). Pardosa uncata (Thorell) 1877 (Bull. U.S. Geol. Surv. 3:508). Undesignated localities in Utah (LD). Pardosa titahensis Chamberlin 1919 (Ann. Ent. Soc. Amer. 12:258). Chalk Crk (in Uinta Mts) (C19), Navajo Mt (AG), Steep Crk (BU). Pardosa wasatchensis Gertsch 1933 (Amer. Mus. Novitates 636:25). Utah (R). Pardosa xerampelina (Keyserling) 1876 (Verb. zool. bot. Ges. Wien 26:622). Utah (C8). Pardosa yavapa Chamberlin 1925 (Bull. Mus. Comp. Zool. 68:231). Fish Lake, Henry Mts, Richfield, Weber Cyn (G34), La Sal Mts (C28a), Boulder Mt, Steep Crk, Table Cliff Pass (BU). Pardosa zionis Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):33). Coyote Gulch (BU), Zion Nat Park (CI42a). Pardosa sp. Calf Crk Crossing (nr Esca- lante), Kanab Cyn, Steep Crk (BU), Four- mile Bench (AU). Pelecopsis digna Chamberlin & Ivie 1939 (Verb. 7 Kongr. intern. Ent. Berlin, p. 70). Clear Crk, Uinta Mts (CI33), Provo River (in Uinta Mts), Raft River Mts (CI39). Pelecopsis sculptum (Emerton) 1917 (Ca- nad. Ent. 49:261). Clear Crk, Uinta Mts (CI33). Pellenes americanus (Keyserling) 1884 (Verb. zool. bot. Ges. Wien 34:506). USU School Forest (W). Pellenes candidus G. & E. Peckham 1901 (Bull. Wisconsin Nat. Hist Soc. (N.S.) 1:206). Rock Island (in Utah Lake) (BU), Verdure (CG28). Pellenes clypeatus (Banks) 1895 (Canad. Ent. 27:102). Moab, Verdure (CG28). Pellenes decorus (Blackwall) 1846 (Ann. Mag. Nat. Hist. (1)17:34). Utah (C58). Pellenes griseus G. & E. Peckham 1901 (Bull. Wisconsin Nat. Hist. Soc. (N.S.) 1:206). Price (CG28). Pellenes hirsutus (G. & E. Peckham) 1888 (Trans. Wisconsin Acad. Sci. 7:64). Green Cyn (H). Pellenes laggani G. & E. Peckham 1909 (Trans. Wisconsin Acad. Sci. 16:532). USU School Forest (W). Pellenes mimtda Chamberlin & Gertsch 1929 (J. Ent. Zool. Pomona Coll. 21:111). St George (CG29). Pellenes oregonense (G. & E. Peckham) 1888 (Trans. Wisconsin Acad. Sci. 7:66). Val- ley City CG28). April 1983 Allred, Kaston: Utah Spiders 511 Pellenes viridipes (Hentz) 1846 (J. Boston See. Nat. Hist. 5:362). Verdure (CG28). Pellenes sp. Curlew Valley (K), Four-mile Bench, Glen Cyn City, Nipple Bench (AU). Phidipptis apacheanns Chamberlin & Gertsch 1929 (J. Ent. Zool. Pomona Coll. 21:109). Black Rock (W Salt Lake City), City Crk Cyn, Fisher Pass, Strawberry (CG29), Green Cyii (McU), Locomotive Spngs (K). Phidippus asottis Chamberlin & Ivie 1933 (Amer. Mus. Novitates 631:50). Grouse Crk (CI33). Phidippus johnsoni (G. & E. Peckham) 1883 (Desc. Spiders Fam. Attidae, p. 22). As- pen Grove, Provo, Rock Island (in Utah Lake) (BU), Clear Crk, Dove Crk, Raft River S fk (CI33), Green Cyn (H), Mounds, San Ra- fael, Straight Wash (CG28), St George (C21). Phidippus octopunctatus (G. & E. Peck- ham) 1883 (Descr. Spiders Fam. Attidae, p. 6). Green Cyn (H). Phidippus pogonopus Chamberlin 1925 (Proc. Calif. Acad. Sci. 14:132). Greenriver (C25). Phidippus pruinosus G. & E. Peckham 1909 (Trans. Wisconsin Acad. Sci. 16:388). Blanding, Bluff (CG38). Phidippus purpuratus Keyserling 1884 (Verh. zool. bot. Ges. Wien 34:489). USU School Forest (W). Phidippus workmanii (G. & E. Peckham) 1901 (Trans. Wisconsin Acad. Sci. 13:287). St George (C21). Phidippus sp. Glen Cyn City (AU), Posey Lake, Provo (BU). Philodromus alascensis Keyserling 1884 (Verh. zool. bot. Ges. Wien 33:674). Farm- ington Cyn (LL), Salina (CG28), USU School Forest (W). Philodronius californicus Keyserling 1884 (Verh. zool. bot. Ges. Wien 33:676). Big Cot- tonwood Cyn, Fruita, Mill Crk, Ogden River Cyn, Payson, Salt Lake City (D), Bluff, Moab, San Rafael River, Zion Nat Park (CG28), Green Cyn (H), St George (CW). Philodromus cespitum (Walckenaer) 1802 (Faune Parisienne, Insecta, 2:230). Curlew Valley (K), Richfield, W Utah Lake (D). Philodromus histrio (Latreille) 1819 (N. Die. hist. nat. Nouv. edit., Paris, p. 36). Green Cyn (H). Philodromus imbecillus Keyserling 1880 (Spinn. Amer., Lat., p. 224). Paradise (DR). Philodromus infuscatus Keyserling 1880 (Spinn. Amer., Lat., p. 222). St George (C21). Philodromus insperatus Schick 1965 (Bull. Amer. Mus. Nat. Hist. 129:66). East Cyn, Salt Lake City, Silver Lake (DR68). Philodromus pernix Blackwall 1846 (Ann. Mag. Nat. Hist (1)17:38). Clear Crk, Raft River S fk, Yost (CI33), Mill Crk (D). Philodromus praelustris Keyserling 1880 (Spinn. Amer., Lat., p. 209). Allen Cyn (D). Philodromus quercicola Schick 1965 (Bull. Amer. Mus. Nat. Hist. 129:56). Kelton Pass (K). Philodromus rodecki Gertsch & Jellison 1939 (Amer. Mus. Novitates 1032:27). Red Cyn Camp (DR68). Philodromus rufus Walckenaer 1826 (Faune frangaise, Paris, p. 91). Green Cyn (H), USU School Forest (W). Phildromits satullus Keyserling 1880 (Spinn. Amer., Lat., p. 211). Green Cyn (H), Little Cottonwood Cyn, Salt Lake City, Zion Nat Park (DR68). Philodromus speciosus Gertsch 1934 (Amer. Mus. Novitates 707:22). Green Cyn (H), Salt Lake City (G34a), Vermillion Castle (nr Parowan) (LL). Philodromus spectabilis Keyserling 1884 (Verh. zool. bot. Ges. Wien 33:676). SW Utah (DR76). Philodromus sp. Escalante, Kanab Cyn, Posey Lake, Provo, Rock Island (in Utah Lake), Three Lakes (N Kanab), Widtsoe, Wil- low Tank Spngs (BU). Pholcophora americana Banks 1893 (Trans. Amer. Ent. Soc. 23:57). Four-mile Bench (5 km SE cow camp at head Wesses Cyn) (AG), Lehi (G35), Lynn, Raft River Mts (CI33). Phruronellus pictus Chamberlin & Gertsch 1930 (Proc. Biol. Soc. Washington 43:138). Bountiful (CG30). Phruronellus similis (Banks) 1895 (J. New York Ent. Soc. 3:81). Utah (R). Phrurotimpus alarius (Hentz) 1847 (J. Bos- ton Soc. Nat. Hist. 5:461). Caineville, Moab, San Rafael River (CG28), Four-mile Bench (5 km SE cow camp at head Wesses Cyn) (AG), Green Cyn (H). Phrurotimpus borealis (Emerton) 1911 (Trans. Conn. Acad. Sci. 16:404). Clear Crk (CI33). Phrurotimpus mormon (Chamberlin & Gertsch) 1930 (Proc. Biol. Soc. Washington 512 Great Basin Naturalist Vol. 43, No. 3 43:140). Ferron (CI35b), Salt Lake City (CG30). Phrurotimpus woodburyi (Chamberlin & Woodbury) 1929 (J. Ent. Zool. Pomona Coll. 21:18). Dry Cyn, Smith and Morehouse Cyn (CI35b). Physocyclus tanneri Chamberlin 1921 (Ca- nad. Ent. 53:245). Lake Powell (C58), St George (C21), Zion Nat Park (CW) Piabuna nanna Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2) :41). Dove Crk, Grouse Crk (CI33). Piabuna xerophila Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8):38). Moab (CI35b). Pimoa utahana (Gertsch & Ivie) 1936 (Amer. Mus. Novitates 858:16). Salt Lake City (CI43). Pirata minutus Emerton 1885 (Trans. Conn. Acad. Sci. 6:493). Ferron, Logan 1 mi S (WE). Pirata montanus Emerton 1885 (Trans. Conn. Acad. Sci. 6:493). Fruita (CG28). Pirata piraticus (Clerck) 1757 (Aranei Suecici, p. 102). Coyote Gulch (in Escalante Basin), Rock Island (in Utah Lake) (BU), counties of Davis, Emery, Grand, Millard, Rich, and Salt Lake (WE). Pirata sedentarius Montgomery 1904 (Proc. Acad. Nat. Sci. Philadelphia 56:312). Counties of Cache, Grand, Millard, Salt Lake, and Utah (WE). Pirata sp. Kanab (BU). Pityohypliantes costatus (Hentz) 1850 (J. Boston Soc. Nat. Hist. 6:31). Bear Lake, Chalk Crk (in Uinta Mts), Logan Cyn (C20b), Clear Crk, Raft river S fk (CI33). Pityohypliantes cristatus Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):38) USU School Forest (W). Pityohypthantes navajo Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):60). Henry Mts (CI42a). Pityohyphantes sp. Posey Lake (BU). Plectreurys tristis Simon 1893 (Ann. Ent. Soc. France 62:300). Blanding, Bluff, Devils Cyn, Straight Wash (CG28), Nipple Bench (AG), Zion Nat Park (CW). Pocadicnemis pumila (Blackwall) 1841 (Trans. Linn. Soc. London 18:639). Dove Crk, Raft River S fk (CI33), St John, Verdure (CG28), Zion Nat Park (CW). Poeciloneta bellona Chamberlin & Ivie 1943 (Bull. Univ. Utah Biol. Ser. 7(6): 13). Mirror Lake (CI43). Poeciloneta canionis Chamberlin & Ivie 1943 (Bull. Univ. Utah Biol. Ser. 7(6): 13). Cobble Rest (CI43). Porrhomma ocella Chamberlin & Ivie 1943 (Bull. Univ. Utah Biol. Ser. 7(6):4). City Crk Cyn, Smith & Morehouse Cyn (CI43). Prolinyphia litigiosa (Keyserling) 1886 (Spinn. Amer., Theridiidae 2:62). Aspen Grove (BU), Clear Crk (CI33), Zion Nat Park (CW). Psilochorus imitatus Gertsch & Mulaik 1940 (Bull. Amer. Mus. Nat. Hist. 77:321). Four-mile Bench (8 km SE cow camp at head Wesses Cyn), Nipple Bench (AG), Salt Lake City (GU). Psilochorus utahensis Chamberlin 1919 (Ann. Ent. Soc. Amer. 12:247). Clear Crk, Fillmore, Fillmore Cyn, Pine Cyn (C19), Coyote Gulch, Cyclone Lake (nr Escalante), Steep Crk (BU), Devils Cyn, Moab, Straight Wash, Verdure (CG28), Ahlstrom Pt, Brig- ham Plains Flat, Four-mile Bench (5 km SE cow camp at head Wesses Cyn), Glen Cyn City 5 km W, Nipple Bench, Smokey Mt (14 and 23 km from Last Chance Jet), Tibbet Spng 2 km NE (AG), Green Cyn (H), Lake Powell (C58), Grouse Crk, Lynn, Raft River Mts (CI33), St George (C21). Rhysodromus alascensis dondalei Schick 1965 (Bull. Amer. Mus. Nat. Hist. 129:73). Utah (S). Rhysodromus virescens (Thorell) 1877 (Bull. U.S. Geol. Surv. 3:500). Calf Crk, Pink Dunes (N Kanab) (BU), Clear Crk, Raft River S fk (CI33), Curlew Valley (K), Lake Powell (C58), St George (C21). Salticus austinensis Gertsch 1936 (Amer. Mus. Novitates 852:20). St George (C21). Salticus scenicus (Clerck) 1757 (Aranei Suecici, p. 117). Benson (McU). Salticus peckhamae (Cockerell) 1897 (Ca- nad. Ent. 29:223). Lynn (CI33), Snowville (K). Sassacus papenhoei G. & E. Peckham 1895 (Occ. Pap. Nat. Hist. Soc. Wisconsin 2:177). City Crk Cyn (CG29), Curlew Valley (K), Green Cyn (H), Steep Crk (BU). Satilatlas gentilis Millidge 1981 (Bull. Amer. Mus. Nat. Hist. 170:251). Smith & Morehouse Cyn (M81b). Schizocosa avida (Walckenaer) 1837 (Hist. Nat. Inst. Apt. 1:322). Bluff, Caineville, Fre- mont River, Fruita, Moab, Straight Wash, April 1983 Allred, Kaston: Utah Spiders 513 Valley City (CG28), Clear Crk, Raft River S fk (CI33), Four-mile Bench (8 km SE cow camp at head Wesses Cyn), Glen Cyn City 5 km W & 6.5 km S, Smokey Mt (23 km from Last Chance Jet) (AG), Widtsoe (BU). Schizocosa (?) celerior Chamberlin 1910 (Ent. News 21:2). (Not a Schizocosa accord- ing to Dondale & Redner 1978). St. George (GW). Schizocosa crassipalpata Roewer 1951(Abh. naturw. Ver. Bremen 32:440). nr Salt Lake City (DR78). Schizocosa mccooki (Montgomery) 1904 (Proc. Acad. Nat. Sci. Philadelphia 56:283). Green Cyn (H), Hughes Cyn (CI42a), USU School Forest (W), other localities not desig- nated (DR78). Schizocosa mimula (Gertsch) 1934 (Amer. Mus. Novitates 726:5). Salt Lake City (DR78). Schizocosa minnesotensis (Gertsch) 1934 (Amer. Mus. Novitates 726:4). Utah (DR78). Schizocosa saltatrix (Hentz) 1844 (J. Boston Soc. Nat. Hist. 4:387). Fish Lake (GW). Sciastes simplex (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:250). Bear Lake, Logan Cyn, Mirror Lake, upper Provo River, Swan Crk Cyn, Teapot Lake (in Uinta Mts), Wasatch Mts (CI45), Chalk Crk (in Uinta Mts) (C19). Scotinella formidabilis (Chamberlin & Gertsch) 1930 (Proc. Biol. Soc. Washington 43:137). Fish Lake (CG30), Posey Lake (BU), Raft River S fk (CI33). Scotinella pelvicolens (Chamberlin & Gertsch) 1930 (Proc. Biol. Soc. Washington 43:138). Clear Crk (CI33), Mt Ellen (in Hen- ry Mts) (CG30), USU School Forest (W). Scotinella pugnata (Emerton) 1890 (Trans. Conn. Acad. Sci. 8:188). "Utah" (DR82). Scotinotylus castorus (Chamberlin) 1948 (Ann. Ent. Soc. Amer. 41:520). Beaver Cyn (C48). Scotinotylus diibiosus Millidge 1981 (J. Arach. 9:205). Logan (M81). Scotinotylus kenus (Chamberlin) 1948 (Ann. Ent. Soc. Amer. 41:519). Ferron Res, Mirror Lake (C48). Scotinotylus pallidus (Emerton) 1882 (Trans. Conn. Acad. Sci. 6:1). Utah (M81). Scotinotylus sacratus Millidge 1981 (J. Arach. 9:181). Mirror Lake (M81). Scotinotylus sanctus (Crosby) 1929 (Ent. News 40:81). Clear Crk, Raft River S fk (CI33). Scotoussa bidentata (Emerton) 1882 (Trans. Conn. Acad. Sci. 6:56). Carter Crk & Deep Crk jet, Smith & Morehouse Cyn (CI45). Scylaceus sp. Four-mile Bench (AU). Sergiolus angustus (Banks) 1904 (Proc. Ca- lif. Acad. Sci. 3:337). City Crk Cyn, Richfield (C36b), Clear Crk Cyn, Hughes Cyn, Salt Lake City (PS81), Fruita (CG28). Sergiolus columbianus (Emerton) 1917 (Canad. Ent. 49:269). Salt Lake County (PS81). Sergiolus iviei Platnick & Shadab 1981 (Amer. Mus. Novitates 2717:34). Lynn, Moab, Salt Lake City, West Jordan (PS81). Sergiolus lowelli Chamberlin & Woodbury 1929 (Procf Biol. Soc. Washington 41:177). Moab (PS81), St George (CW). Sergiolus montanus (Emerton) 1890 (Trans. Conn. Acad. Sci. 8:175). Clear Crk, Lynn, Raft River Mts (CI33), Green Cyn (H), Rich- field, Salt Lake City (C36b), Zion Nat Park (CW), and counties of Garfield, Rich, San Juan, Uintah, Utah, Wayne and Weber (PS81). Sisicottus montanus (Emerton) 1882 (Trans. Conn. Acad. Sci. 6:55). Chalk Crk (in Uinta Mts) (C19), Clear Crk, Raft River S fk (CI33). Sisicottus uintanus Chamberlin & Ivie 1939 (Verb. 7 intern. Kongr. Ent. Berlin 1:65). Mirror Lake (CI39). Sitticus finschi (L. Koch) 187,9 (Verb, zool bot. Ges. Wien 28:483). USU School Forest (W). Sosticus loricatus (L. Koch) 1866 (Arach. Earn. Drassiden, p. 131). City Crk Cyn, Green River, Price (PS76a), Logan (CG40). Spirembolus humilis Millidge 1980 (J. Arach. 8:12). Mirror Lake, Tooele County (M80). Spirembolus monticolens (Chamberlin) 1919 (Ann. Ent. Soc. Amer. 12:251). Chalk Crk (in Uinta Mts) (C19), Grouse Crk, Raft River S fk (CI33), Logan Cyn, Lynn 8 mi S, Smith & Morehouse Cyn (CI45a). Spirembolus mundus Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2): 18). City Crk Cyn, Raft River Mts (CI45a), Clear Crk (CI33), Green Cyn (H). 514 Great Basin Naturalist Vol. 43, No. 3 Spirembolus pachygnathus Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 18). City Crk Cyn, Fish Lake, Mill Crk Cyn, Smith & Morehouse Cyn (CI45a). Spirembolus pallidus Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser 2(8): 19). Pine Spngs (on Mt Ellen in Henry Mts) (CI45a). Spirembolus spirotubus (Banks) 1895 (Ann. New York Acad. Sci. 8:424). Mt Ellen (in Henry Mts), and other localities not desig- nated (M80). Spirembolus vallicolens (Chamberlin) 1920 (Canad. Ent. 52:198). Fillmore, Fish Lake, Mt Ellen (in Henry Mts), Provo River mth, Richfield, Salt Lake City 10 mi W, Santaquin Res (CI45a), Mill Crk (C20b), Verdure (CG28a). (Millidge 1980 states that many of Chamberlin and Ivies 1945a records of this species actually are S. spirotubus.) Steatoda albomaculata (DeGeer) 1778 (Mem. pour, servir a I'hist. des Ins. 7:257). Clear Crk, Grouse Crk, Lynn, Raft River Mts, Raft River S fk (CI33), La Sal Mts (C28a), Boulder Mt, Cyclone Lake, Posey Lake, Steep Crk (BU), Bryce Cyn Nat Park, comities of Kane, Millard, San Juan (L57), and Wayne (lU). Steatoda americona (Emerton) 1882 (Trans. Comi. Acad. Sci. 6:23). Fort Douglas, Salt Lake City, Washington County (C28), Fruita, Verdure (CG28), Green Cyn (H), Oquirrh Mts (lU). Steatoda castanea (Clerck) 1757 (Aranei Suecici, p. 49). Logan (McU). Steatoda fidva (Keyserling) 1882 (Spinn. Amer. Theridiidae 2:142). Dry Cyn, Fort Douglas, Great Salt Lake shore (lU), Ahlst- rom Point, Brigham Plains, Glen Cyn City, Nipple Bench, Tibbet Spng 2 km NE (AG), Spring Lake (Ky), and counties of Carbon, Sevier, Tooele and Weber (L57). Steatoda grandis Banks 1901 (Proc. Acad. Nat. Sci. Philadelphia 53:578). Beaver Dam Wash, Utah Lake W shore (lU), Cedar Cyn nr Cedar Breaks Nat Mon, Ferron, Henry Mts, Levan, Salt Lake City, Zion Nat Park (CI35b), Escalante, Fish Lake, Fruita, Help- er, Noton, Parowan, Price, Red Cyn, Rich- field, Scipio, Vernal, Watson, White River (nr Evacuation Crk) (L57), St George (CW), Steep Crk, Torrey 15 mi S (BU). (According to Gertsh (pers. comm.), this species and S. mexicana need clarification, and the records above are tentatively assigned to these two species. Steatoda mexicana actually is south- ern Mexico in distribution, and the Utah specimens need other names, possible only after revision of the group.) Steatoda hespera Chamberlin & Ivie 1933 (Bull. Univ. Utah Biol. Ser. 2(2):9). Aspen Grove, Cyclone Lake (nr Escalante) (BU), Clear Crk, Raft River Mts, Raft River S fk (CI33), St George (CW), USU School Forest (W). Steatoda medialis (Banks) 1898 (Proc. Ca- hf. Acad. Sci. (3)1:239). Several localities not designated (L59). Steatoda mexicana Levi 1957 (Bull. Mus. Comp. Zool. 117:415). Beaver Dam Wash, Boulder 15 mi N, Bryce Cyn Nat Park, Fish Lake, Henry Mts, Lehi, Panguitch 10 mi SE, Salt Lake City, St George, Utah Lake W shore (L57). (See comments under S. grandis). Steatoda triangulosa (Walckenaer) 1802 (Faune parisienne, Paris 2:207). Utah (LR). Steatoda variata Gertsch 1960 (Amer. Mus. Novitates 1982:24). Ferron, Fruita, Green- river, Grouse Crk, Moab, Raft River Mts, Salt Lake City, White River (on Evacuation Crk) (G60), Glen Cyn City (AG). Steatoda washona Gertsch 1960 (Amer. Mus. Novitates 1982:21). Ferron, Fish Lake, Glenwood, Loa, Moab, Richfield, Salt Lake City (G60). Steatoda sp. Three Lakes (N Kanab) (BU). Stemonyphantes blauveltae Gertsch 1951 (Amer. Mus. Novitates 1514:1). Strawberry Res (G51a). Stylophora puUata (O. Pick. -Cambridge) 1863 (Zoologist 21:8580). Cobble Rest (169). Synagales sp. Green Cyn (H), Cedar Hills, Curlew Valley (K). Tachygyna haydeni Chamberlin & Ivie 1939 (Verh. 7 intern. Kongr. Ent. Berlin, p. 63). Mirror Lake (CI39). Tachygyna paita Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:548). Mirror Lake (C48a). Tachygyna pallida Chamberlin & Ivie 1939 (Verh. 7 intern. Kongr. Ent. Berlin, p. 63). Provo River N fk (in Uinta Mts) (CI39). Tachygyna tuoba (Chamberlin & Ivie) 1933 (Bull. Univ. Utah Biol. Ser. 2(2):23). Raft River S fk (CI33), Raft River Mts (CI39). Tachygyna watona Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:549). Mirror Lake (C48a). April 1983 Allred, Kaston: Utah Spiders 515 Talavera mintita (Banks) 1895 (Canad. Ent. 27:99). Green Cyn (H), USU School Forest (W). Tapinocyba gamma Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:550). Mirror Lake (C48a). Tetragnatha extensa (Linnaeus) 1758 (Syst. Nat. 10th ed., p. 621). Aspen Grove, Calf Crk Crossing (nr Escalante), Boulder (BU), Clear Crk, Dove Crk, Raft River S fk (CI33), Cur- lew Valley (K), Zion Nat Park (CW). Tetragnatha laboriosa Hentz 1850 (J. Bos- ton Soc. Nat. Hist. 6:27). Curlew Valley (K), Dove Crk, Raft River S fk (CI33), Green Cyn (H), Moab (CG28), Calf Crk, Kanab Cyn, Posey Lake, Steep Crk, Three Lakes (N Ka- nab), Utah Lake (BU), Zion Nat Park (CW). Tetragnatha marginata (Thorell) 1890 (Ann. Mus. Civ. Stor. Nat. Genova 28:230). Lake Powell (C58). Tetragnatha pallescens F. Pick. — Cam- bridge 1903. (Biol. Centr. Amer., Arachn., Ar. 2:436). NE Utah (L81). Tetragnatha shoshone Levi 1981 (Bull. Mus. Comp. Zool. 149:312). Bear Lake SE shore. Granite, Laketown, Provo River mth, Richfield, Salt Lake City, Utah Lake W side (L81). Tetragnatha straminea Emerton 1884 (Trans. Conn. Acad. Sci. 6:335). N central Utah (L81). Tetragnatha versicolor Walckenaer 1841 (Hist. Nat. Ins. Apt. 2:215). Many localities not designated (L81). Tetragnatha sp. Willow Tank Spngs (BU). Thanatus altimontis Gertsch 1933 (Amer. Mus. Novitates 636:6). Glen Cyn City (AG). Thanatus coloradensis Keyserling 1880 (Spinn. Amer. Lat. 1:206). Clear Crk, Raft River S fk (CI33), St George (BU). Tlianatus formicinus (Clerck) 1757 (Aranei Suecici, p. 134). Green Cyn (H), USU School Forest (W). Theridion albidum Banks 1895 (J. New York Ent. Soc. 3:84). Green Cyn (H). TJieridion australe Banks 1899 (Proc. Ent. Soc. Washington 4:191). Richfield (L57). Ttieridion berkeleyi Emerton 1924 (Pan Pa- cif. Ent. 1:30). Counties of Salt Lake & Utah (L57). TJieridion denticulatum (Walckenaer) 1802 (Fauna parisienne, Paris 2:208). Utah (LR). Theridion differens Emerton 1882 (Trans. Conn. Acad. Sci. 6:9). Counties of Box Elder, Grand, Rich, Salt Lake and Sevier (L57). Theridion dihititm Levi 1957 (Bull. Amer. Mus. Nat. Hist. 112:37). Richfield, Washing- ton County (L57). Theridion goodnightorum Levi 1957 (Bull. Amer. Mus. Nat. Hist. 112:41). Counties of Sevier and Utah (L57). TJieridion Jiaivea Levi 1957 (Bull. Amer. Mus. Nat. Hist. 112:48). Zion Nat Park (L57). TJieridion leecJii Gertsch & Archer 1942 (Amer. Mus. Novitates 1171:8). Counties of Morgan, Salt Lake and Utah (L57). Theridion montanum Emerton 1882 (Trans. Conn. Acad. Sci. 6:10). Pickleville (Wa), USU School Forest (W), and counties of Beaver, Emery (L47) and Rich (L57). TJieridion murarium Emerton 1882 (Trans. Conn. Acad.* Sci. 6:11). Counties of Davis, Tooele and Utah (L57). TJieridion neomexicanum Banks 1901 (Proc. Acad. Nat. Sci. Philadelphia 53:577). Beaver, Bryce Cyn Nat Park, counties of Car- bon, Grand, Juab, Kane, Sevier, Wayne and Weber (L57), Green Cyn (H), Hughes Cyn, Logan Cyn (lU), Pickleville (Wa), St George (CW), Straight Wash (CG28), USU School Forest (W). TJieridion ohlerti Thorell 1870 (Remarks Syn. European Spiders, p. 85). Pickleville (Wa), USU School Forest (W), counties of Tooele and Utah (L57). TJieridion ornatum Hahn 1831 (Monogr. der Spinnen, H.6, T.3, 7.c). Richfield, Salt Lake County (L57). TJieridion petraeiim L. Koch 1872 (Zeits. Ferd. Tirol Voral. 17:246). Green Cyn (H), counties of Grand, Millard, Morgan, Salt Lake, Summit, Utah and Washington (L57). TJieridion pictum (Walckenaer) 1802 (Favine parisienne, Paris 2:207). Utah (LR). TJieridion rabuni Chamberlin & Ivie 1944 (Bull. Univ. Utah Biol. Ser. 8(5):53). Green Cyn (BA), Pickleville (Wa), USU School For- est (W), counties of Morgan, Salt Lake, San Juan and Utah (L57). TJieridion sexpunctatum Emerton 1882 (Trans. Conn. Acad. Sci. 6:12). Clear Crk, Raft River Mts (CI33), Zion Nat Park, coun- ties of Salt Lake, Summit, Utah and Wasatch (L57). 516 Great Basin Naturalist Vol. 43, No. 3 Theridion timpanogos Levi 1957 (Bull. Amer. Mus. Nat. Hist. 112:31). American Fk Cyn, Mt Timpanogos (L57). Theridion transgressum Petrunckevitch 1911 (Bull. Amer. Mus. Nat. Hist. 29:208). Ogden Cyn, Weber River, counties of Gar- field, Washington and Utah (L57). Theridion sp. Steep Crk (BU). Theridula opidenta (Walckenaer) 1837 (Hist. Nat. Ins. Apt. 1:322). Utah (LR). TInodina sylvana (Hentz) 1846 (J. Boston Soc. Nat. Hist. 5:364). Utah (C58). TInodina sp. Willow Tank Spngs (BU). TJiomisiis sp. La Sal Mts (C28a). Thymoites camano Levi 1957 (Bull. Amer. Mus. Nat. Hist. 112:105). Mill Crk Cyn (L57). Tfiymoites edinburgensis (Gertsch & Mu- laik) 1936 (Amer. Mus. Novitates 863:9). Hughes Cyn (L57). Thymoites palUdns (Emerton) 1913 (Bull. Amer. Mus. Nat. Hist. 32:255). Utah (LR). Tibelhis chamberlini Gertsch 1933 (Amer. Mus. Novitates 593:10). Elsinore, Monroe, Zion Nat Park (G33), Green Cyn (H), Kelton Pass (K). TibeUus diittoni (Hentz) 1847 (J. Boston Soc. Nat. Hist. 5:448). Lake Powell (C58), St George (C21). Tibelhis gertschi Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):81). Smith & Morehouse Cyn (CI42a). TibeUus maritimus (Menge) 1875 (Schr. na- turf. Ges., Danzig, N.F. 3:398). Zion Nat Park (G33). TibeUus oblongus (Walckenaer) 1802 (Faune parisienne, Paris 2:228). Clear Crk, Raft River S fk (CI33), Curlew Valley (K), Green Cyn (H), USU School Forest (W), Utah Lake (BU). Tibelhis sp. Moab (CG28), Kanab Cyn, Three Lakes (N Kanab) (BU). Titanebo magnificus Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):81). Salt Lake City (CI42a). Titanebo sp. Escalante, Posey Lakes, Three Lakes (N Kanab) (BU). Titanoeca nigrella (Chamberlin) 1919 (J. Ent. Zool. Pomona Coll. 12:2). Beaver 10 mi E, Beaver Cyn, Beaver Mt, Bountiful, City Crk Cyn, Cobble Rest, Daniel, Dry Cyn, Eu- reka, Evacuation Crk, Farmington, Fruita, Garden City, Glenwood, Junction, Moab, Parleys Cyn, Richfield, Utah Lake (Le), Green Cyn (H). Titanoeca silvicola Chamberlin & Ivie 1947 (Bull. Univ. Utah Biol. Ser. 10(3): 15). City Crk Cyn (C47), Fish Lake, Glendale (at Virgin River), La Sal Forest, Snow Crk (Le). Tmarus angulatiis (Walckenaer) 1837 (Hist. Nat. Ins. Apt. 1:537). Zion Nat Park (CW). Trachelas deceptus (Banks) 1895 (J. New York Ent. Soc. 3:81). Moab, Verdure, Wayne County (CG28), St George (CW). Trachelas mexicanus Banks 1898 (Proc. Calif. Acad. Sci. 1:226). St George, Zion Nat Park (CI35b). Trachelas tranquillus (Hentz) 1847 (J. Bos- ton Soc. Nat. Hist 5:450). Bluff (CG28), St George, Zion Nat Park (CW). Trachelas sp. Three Lakes (BU). Tricholathys spiralis Chamberlin & Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8):28). Kelton (K), Salt Lake City, Utah Lake W side (CI35b). Trochosa avara (Keyserling) 1877 (Verb, zool. bot. Ges. Wien 27:661). La Sal Mts (BU). Trochosa gosiuta (Chamberlin) 1908 (Proc. Acad. Nat. Sci. Philadelphia 60:281). Devils Cyn, Fruita, San Rafael, Verdure (CG28), St George, Zion Nat Park (CW), Utah Lake (BU), counties of Carbon, Duchesne, Grand, Salt Lake, Sevier, Tooele, and Wasatch (Br). Trochosa terricola Thorell 1856 (Nova Acta Reg. Soc. Sci. Upsala (3)2:171). Aspen Grove (BU), City Crk Cyn, Lambs Cyn (CG29), Clear Crk, Dove Crk, Grouse Crk (CI33), Salt Lake City, Tooele (G34c), counties of Daggett, Rich, Sevier and Summit (Br). Trogloneta paradoxuin Gertsch 1960 (Amer. Mus. Novitates 1981:12). Timpanogos Cave Nat Mon (G60). Tutelina similis (Banks) 1895 (Canad. Ent. 27:100). Clear Crk (CI33), Green Canyon (H). Uloborus diversus Marx 1898 (Proc. Calif. Acad. Sci. 1:234). Fillmore (C19), N, S and central Utah (MG). Usofila flava Chamberlin & Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):8). Provo Riv- er (in Uinta Mts), Raft River Mts, Wasatch Mts (CI42a). Usofila gracilis Marx 1891 (Proc. Ent. Soc. Washington 2:9). Mill Crk (G35). Walckenaeria communis (Emerton) 1882 (Trans. Conn. Acad. Sci. 6:41). Dove Crk (CI33), Steep Crk (BU). April 1983 Allred, Kaston: Utah Spiders 517 Walckenaeria perditits (Chamberlin) 1948 (Ann. Ent. Soc. Amer. 41:555). Salt Lake City (C48a). Walckenaeria spiralis (Emerton) 1882 (Trans. Conn. Acad. Sci. 6:39). Raft River S fk (CI33). Walckenaeria thrinax (Chamberlin & Ivie) 1933 (Bull. Univ. Utah Biol. Ser. 2(2) :24). Dove Crk (CI33). Walckenaeria weber (Chamberlin) 1948 (Ann. Ent. Soc. Amer. 41:557). Smith & Morehouse Cyn (C48a). Walckenaeria sp. Fniita (CG28). Wubana drassoides (Emerton) 1882 (Trans. Conn. Acad. Sci. 6:72). Chalk Crk (in Uinta Mts), Fillmore, La Sal Mts (CI36). Wubana reminescens Chamberlin 1948 (Ann. Ent. Soc. Amer. 41:560). USU School Forest (W). Wubana utahana Chamberlin & Ivie 1936 (Ann. Ent. Soc. Amer. 29:89). Chalk Crk (in Uinta Mts), Fish Lake, Mirror Lake, Smith & Morehouse Cyn (CI36). Xysticus apachecus Gertsch 1933 (Amer. Mus. Novitates 593:22). Blanding (G3), Glen- wood (G53). Xysticus benefactor Keyserling 1880 (Spirm. Amer., Lat. 1:22). Bridger Basin, Brig- ham Cyn, Logan (G53). Xysticus californicus Keyserling 1880 (Spinn. Amer., Lat. 1:37). Brigham, Clarks- ton, Escalante 22 mi N, Fish Lake, Logan, Ogden, Parowan, Salina, Willard, Zion Nat Park (G53), Grouse Crk, Raft River S fk, Lynn (CI33), St George (CG29), Straight Wash, Verdure (CG28), Green Cyn (H), Utah Lake (BU). Xysticus coloradensis Bryant 1930 (Psyche 37:133). Tooele County (G53). Xysticus facetus O. Pick. -Cambridge 1896 (Biol. Centr. Amer., Arach. Ar. 1:179). Moab (G53). Xysticus ferox (Hentz) 1847 (J. Boston Soc. Nat. Hist 5:445). Bluff, Fruita, Moab, Price, San Rafael River (CG28). Xysticus gertschi Schick 1965 (Bull. Amer. Mus. Nat. Hist. 129:159). Clear Crk Cyn, El- sinore, Monroe Cyn (G34a). Xysticus gosiutus Gertsch 1933 (Amer. Mus. Novitates 593:20). Little Cottonwood Cyn, Zion Nat Park (G33). Xysticus gulosus Keyserling 1880 (Spinn. Amer., Lat. 1:43). Blanding (CG28), Glen Cyn City (AG), Kelton Pass (K), Lake Powell (C58), Provo (BU), St George (CW). Xysticus imitarius Gertsch 1953 (Bull. Amer. Mus. Nat. Hist. 102:442). Little Cot- tonwood Cyn (G53). Xysticus knowltoni Gertsch 1939 (Bull. Amer. Mus. Nat. Hist. 76:399). Vernon (G53). Xysticus lassanus Chamberlin 1925 (Bull. Mus. Comp. Zool. 67:218). Glen Cyn City, Smokey Mt (AG), Hanksville, St George (G53). Xysticus locuples Keyserling 1880 (Spinn. Amer., Lat. 1:24). Brigham (G53), Green Cyn (McU), Grouse Crk (CI33). Xysticus lutulentus Gertsch 1939 (Bull. Amer. Mus. Nat. Hist. 76:396). Mill Crk Cyn (CI42a). Xysticus lutzi Gertsch 1935 (Amer. Mus. Novitates 792:27). Glen Cyn City (AG). Xysticus '^ montanensis Keyserling 1887 (Verb. zool. bot. Ges. Wien 37:479). Clear Crk (CI33), Ferron, Salt Lake City (G34a), Green Cyn (H), Kelton (K), USU School For- est (W). Xysticus Orizaba Banks 1898 (Proc. Calif. Acad. Sci. 1:260). Beaver Dam Wash (G53), Little Cottonwood Cyn, St George (G33). Xysticus pellax O. Pick.-Cambridge 1894 (Bio. Centr. Amer., Arach. Ar. 1:138). Kearns (G53). Xysticus sp. Coyote Gulch, Steep Crk, Three Lakes (N Kanab), 10-mi Crk (in Esca- lante Basin) (BU), USU School Forest (W). Zanomys kaiba Chamberlin 1948 (Bull. Univ. Utah Biol. Ser. 10(6):18). Dry Cyn (C48). Zanomys ochra Leech 1972 (Mem. Canad. Ent. Soc. 84:90). Lynndyl 10 mi N (Le). Zelotes fratris Chamberlin 1920 (Canad. Ent. 2:193). Blanding, Bluff, Grantsville, Ver- dure (CG28), Clear Crk, Raft River Mts, Raft River S fk, Yost (CI33), Curlew Valley (K), Fish Lake, Monroe Cyn, Richfield (C36a), Green Cyn (H), Logan Cyn (C20b), St George (CW). Zelotes latnpra Chamberlin 1920 (Canad. Ent. 52:193). Mill Crk (C20b). Zelotes lasalanus Chamberlin 1928 (Ca- nad. Ent. 60:93). La Sal Mts (C28a). Zelotes nannodes Chamberlin 1936 (Amer. Mus. Novitates 853:10). Tremonton 10 mi W (C36b). 518 Great Basin Naturalist Vol. 43, No. 3 Zelotes nanniis Chamberlin & Gertsch 1940 (Amer. Mus. Novitates 1068:18). Bluff, Richardson (CG40). Zelotes puritanus Chamberlin 1922 (Proc. Biol. Soc. Washington 35:164). Puffer Lake, Tooele Cyn (C36a). Zelotes tuohus Chamberlin 1919 (Ann. Ent. Soc. Amer. 12:247). Ahlstrom Point, Brigham Plains (AG), Dove Crk, Raft River Mts (CI33), Fillmore (C19), USU School Forest (W). Zelotes sp. Brighams Plains, Glen Cyn City (AU), Kelton, Locomotive Spngs (K), St George, Tliree Lakes (N Kanab) (BU). Zornella ctiltrigera (L. Koch) 1879 (Kongl. Svenska Vet. Akad. Handl. 16:11). USU School Forest (W). Synonymies of Utah Records In the list below, the names on the left are as recorded for some specimens in collections of Utah universities, or as listed in some pub- lished articles, and are considered as junior synonyms of the names on the right, which are the only ones included in the main body of this report. Aculepeira verae = A. packardi Agelena californica = Agelenopsis califomica Agelena hola = Hololena hola Agelena mimoides = Hololena mimoides Agelena oquirrhensis = Hololena oquirrhensis Agelenopsis mimoides = Hololena mimoides Agelenopsis naevia = A. aperta Agroeca oaba = A. trivittata Alopecosa helluo = Lycosa helluo Amaurobius nevadensis = Callobius nevadensis Amaurobius nomeus = Callobius nomeus Amaurobius utahensis = Callobius nevadensis Aranea carbonaria = Aculepeira carbonaria Aranea cucurbitina = Araniella displicata Aranea displicata = Araniella displicata Aranea ocellatula = Nuctenea patagiata Areanea ocellatus = Nuctenea patagiata Aranea solitaria = Araneus saevus Aranea tusigia = Araneus marmoreus Aranea utahana = Neoscona utahana Arctachaea pelyx = Chrysso pelyx Arctosa cinerea = A. littoralis Aysha nigrifrons = A. incursa Bathyphantes fillmoranus = Meioneta fillmorana Bathyphantes phylax = Helophora orinoma Bathyphantes spatulifer = Leptyphantes lamprus Ceraticelus guttatus = Idionella anomala Ceraticelus tuganus = Idionella tugana Cheraira castoris = Scotinotylus castoris Cheraira kena = Scotinotylus kenus Cicurina garrina = C. robusta Clubiona orinoma = C. moesta Cochembolus sanctus = Scotinotylus sanctus Cornicularia communis = Walckenaeria communis Cornicularia thrinax = Walckenaeria thrinax Cylphosa gosoga = Gnaphosa gosoga Cylphosa sericata = Gnaphosa sericata Delopelma simulatum = Aphonopelma Simula tum Dendryphantes diplacis = Metaphidippus diplacis Dendryphantes mylothrus = prob. Metaphidippus mylothrus Dendryphantes nigromaculatus = Kris nigromaculatus Dendryphantes pruinosus = Phidippus pniinosus Dendryphantes unicus = Metaphidippus unicus Dendryphantes uteanus = prob. Metaphidippus uteanus Dendryphantes verecundus = Metaphidippus verecundus Dendryphantes workmanii = Phidippus workmanii Dictyna dactylata = D. calcarata Dictyna eutypa = Mallos eutypus Dictyna hoples = D. calcarata Dictyna socarina = D. uintana Diplocentria bidentata = Scotoussa bidentata Diplocentria perplexa = Maso perplexus Dipoena daltoni = D. atopa Dipoena hamata = D. prona Drassodes celes = D. saccatus Drassodes robinsoni = D. saccatus Drassyllus apacheus = D. insularis Drassyllus devexus = D. dromeus Drassyllus gertschi = D. conformans Drassyllus lasalus = D. mexicanus Drassyllus mephisto = D. lepidus Emblyna completa = Dictyna completa April 1983 Allred, Kaston: Utah Spiders 519 Emblyna rena = Dictyna completa Emblyna reticulata = Dictyna reticulata Emblyna urica = Mallos niveus Emblyna utesca = Dictyna piratica Epeira labyrinthea = Metapeira labyrinthea Eperigone taibo = Tachygyna tuobo Eularia simplex = Sciastes simplex Euryopis nigripes = E. taczanowskii Evarcha leucophaea = E. hoyi Frontinella communis = F. pyramitela Fuentes taeniola = Metacyrba taeniola Garritus vigerens = Ctenium vigerens Gayenna saniuana = Anyphaena pacifica Geodrassus gosiutus = Drassodes gosiutus Geolycosus carolinensis = Lycosa carolinensis Gertschia sp. = Synagales sp. Gnaphosa gigantea = G. muscorum Hahnia inomata = H. ononidium Haplodrassus dystactus = H. signifer Haplodrassus uncifer = H. bicornis Haplodrassus utus = H. eunis Herpyllus atopophysis = Nodocion eclecticus Herpyllus piedicus = H. propinquus Herpyllus validus = H. hesperolus Herpyllus vasifer = H. ecclesiasticus Icius similis = Tutelina similis Labuella utahana = Pimoa utahana Lathys moabana = Dictyna moaba Latrodectus curacaviensis = L. hesperus Latrodectus geometricus = L. hesperus Latrodectus mactans = L. hesperus Latrodectus variolus = L. hesperus Linyphia communis = Frontinella communis Linyphia ephedra = Linyphantes ephedrus Linyphia litigiosa = Prolinyphia litigiosa Linyphia montana = Neriene montana Linyphia phrygianus = Pityohyphantes costatus Liodrassus metalleus = Nodocion eclecticus Liodrassus utus = Nodocion utus Lithyphantes albomaculatus = Steatoda albomaculata Lycosa avara = Trochosa avara Lycosa gosiuta = Trochosa gosiuta Lycosa orophila = Trochosa terricola Lycosa piraticus = Pirata piraticus Lycosa pratensis = Trochosa terricola Lycosa rafaelana = Geolycosa rafaelana Meriola sp. = Trachelas sp. Metaphidippus nigromaculatus = Eris nigromaculatus Metaphidippus verecundus = M. diplacis Metastinus oblongus = Tibellus oblongus Metepeira alpina = M. grandiosa alpina Misumenops lepidus = M. celer Neoantistea riparia = N. magna Neoscona naiba = N. arabesca Paraphidippus marginatus = Eris marginatus Pardosa atromedia = P. sierra Pardosa wyuta = P. atra Peocilochroa montana = Sergiolus montanus Peponocranium pumila = Pocadicnemis pumila Phidippus borealis = P. purpuratus Phidippus capitatus = Metaphidippus galathea Phidippus formosus = P. johnsoni Philodromus cespiticolis = P. cespitum Philodromus hoples = P. californicus Philodromus virescens = Rhysodromus virescens Pirata sylvestris = P. piraticus (part) & P. insularis (part) Pityohyphantes phrygianus = P. costatus Platyxysticus utahensis = Coriarachne versicolor (part) & C. utahensis (part) Prosopotheca sp. = Walckenaeria sp. Pselothorax atopus = Dipoena atopus Pterotrichia clara = Gnaphosa clara Pusillia mandibulata = Microlinyphia mandibulata Robertus fuscus = Ctenium fusca Robertus eremphilus = Ctenium eremophilus Sassacus uteanus = S. papenhoei Schizocosa wasatchensis = S. mccooki Sergiolus clarus = S. angustus Sergiolus fruitanus = S. angustus Singa variabilis = Hyposinga pygmaea Sostogeus zygethus = Sosticus loricatus Spirembolus chera = Scotinotylus sanctus Steatoda punctulata = S. medialis Tapinocyba alpha = Disembolus alpha Tapinocyba kesimba = Disembolus kesimbus Teutana castanea = Steatoda castanea Theridion placens = T. neomexicanum Theridion pygmaea = Hyposinga pygmaea Theridium canione = Achaearanea canione Tigellinus weber = Walckenaeria weber Tigellinus perditus = Walckenaeria perditus Tosyna calcarata = Dictyna calcarata Tosyna cholla = Dictyna cholla Tosyna terrestris = Dictyna terrestris Trachelas utahanus = T. mexicanus Tricholathys reclusa = Argennina reclusa Trochosa frondicola = Lycosa frondicola 520 Great Basin Naturalist Vol. 43, No. 3 Trochosa pratensis = T. terricola Uloborus utahensis = U. diversus Xysticiis ciinctator = X. californicus Xysticus quinquepunctatus = X. gertschi Zelotes pananus = Sergiolus angustus Zelotes subterraneus = Z. fratris Literature Cited Allred, D. M. 1973. An unusual population of spiders in Utah. Great Basin Nat. 33:51-2. 1974. An unusual spider bite. Great Basin Nat. 34:82. 1975. Arachnids as ecological indicators. Great Basin Nat. .35:405-6. 1978. Bite of wolf spider. Great Basin Nat. 38:478. 1980. A Chiracanthitim spider bite. Great Basin Nat. 40:116. Allred, D. M. and W. J. Gertsch. 1976. Spiders and scorpions from northern Arizona and southern Utah. J. Arach. 3:87-99. Archer, A. F. 1951. Studies in the orb-weaving spiders. Part 1. American Mus. Novitates 1487. 52 pp. Bowling, T. A., and R. J. Sauer. 1975. A taxononiic re- vision of the crab spider genus Coriarachne for North America north of Mexico. J. Arach. 2:183-93. Brady, A. R. 1979. Nearctic species of the wolf spider genus Trochosa. Psyche 86:167-212. Chamberlin, R. V. 1908. Revision of North American spiders of the family Lvcosidae. Proc. Acad. Nat. Sci. Phila. 60: 158-318. ' 1919. New western spiders. Ann. Ent. Soc. Amer- ica 12:239-60. 1920. A new American Amaurobius. Ent. News 31:293. 1920a. The spider of Saltair Beach. Ent. News 31:165-9. 1920b. New spiders from Utah. Candadian Ent. 52:193-201. 1921. A new genus and a new species of spiders in the group Phrurolitheae. Canadian Ent. 53:69-70. 1921a. On some arachnids from southern Utah. Canadian Ent. 53:245-7. 1922. The North American spiders of the family Gnaphosidae. Proc. Biol. Soc. Washington 35:145-72. 1925. Diagnosis of new American Arachnida. Harvard Univ. Bull. Mus. Comp. Zool. 67:221-48. 1925a. New North American spiders. Proc. Calif. Acad. Sci. 14:105-42. 1928. A two-eyed spider from Utah. Psvche 35:235-6. 1928a. Notes on the spiders from the La Sal Mountains of Utah. Canadian Ent. 60:93-5. 1933. Four new spiders of the family Gnapho- sidae. American Mus. Novitates 631. 8 pp. 1936. A new burrowing spider of the genus Ly- cosa from the Uinta Mountains, Utah. Proc. Biol. Soc. Wash. 49:15-16. 1936a. Records of North American Gnaphosidae with descriptions of new species. American .Mus. Novitates 841. .30 pp. 1936b. Further records and descriptions of North American Gnaphosidae. American Mus. Novi- tates 853. 21 pp. 1940. New American tarantulas of the family Aviculariidae. Bull. Univ. Utah Biol. Ser. 5(8). 39 pp. 1947. A summary of the known North American Amaurobiidae. Bull. Univ. Utah Biol. Ser. 10(5). 31pp. 1948. The genera of North American Dictynidae. Bull. Univ. Utah Biol. Ser. 10(6). 31 pp. 1948a. On some spiders of the family Erigonidae. Ann. Ent. Soc. America 41:483-562. 1958. Araneida. In Preliminary report on biologi- cal resources of the Glen Canyon Reservoir. Univ. Utah Anthropological Papers 31:167-71. Chamberlin, R. V., and W. J. Gertsch. 1928. New spi- ders from Utah and California. Pomona College J. Ent. and Zool. 21:101-12. 1928a. Notes on spiders from southeastern Utah. Proc. Biol. Soc. America 41:175-88. 19.30. On fifteen new North American spiders. Proc. Biol. Soc. Washington 43:137-44. 1940. Descriptions of new Gnaphosidae from the United States. American Mus. Novitates 1068. 19 pp. 1958. The spider family Dictvnidae in America north of Mexico. Bull. American Mus. Nat. Hist. 116:1-152. Chamberlin, R. V., and W. Ivie. 1933. Spiders of the Raft River Mountains of Utah. Bull. Univ. Utah Biol. Ser. 2(2). 79 pp. 1933a. A new genus in the family Agelenidae. Bull. Univ. Utah Biol. Ser. 2(3). 7 pp. 1935. Nearctic spiders of the family Urocteidae. Ann. Ent. Soc. America 28:265-79. 1935a. The black widow spider and its varieties in the United States. Bull. Univ. Utali Biol. Ser. 3(1). 29 pp. 1935b. Miscellaneous new American spiders. Bull. Univ. Utah Biol. Ser. 2(8). 79 pp. 1936. Nearctic spiders of the genus Wubana. Ann. Ent. Soc. America 29:85-98. 1937. New spiders of the family Agelenidae from western North America. Ann. Ent. Soc. America .30:211-41. 1939. Studies on North American spiders of the family Micrvphantidae. Verb. VII Int. Kongr. Ent. 1:56-73.' 1939a. New tarantulas from the southwestern states. Bull. Univ. Utah Biol. Ser. 5(1). 17 pp. 1940. Agelenid spiders of the genus Ciciirma Bull. Univ. Utah Biol. Ser. 5(9). 108 pp. 1941. North American Agelenidae of the genera Agelenopsis, Calilena, Ritalena, and Tortolena. Ann. Ent. Soc. America 34:585-628. 1941a. Spiders collected by L. W. Saylor and others, mostly in California. Bull. Univ. Utah Biol. Ser. 6(3). 49 pp. _ 1942. Agelenidae of the genera Hololena, Nova- fl lena, Rualena, and Melpomene. Ann. Ent. Soc. ^ America 35:203-41. April 1983 Allred, Kaston: Utah Spiders 521 1942a. A hundred new species of American spi- ders. Bull. Univ. Utah Biol. Ser. 7(1). 117 pp. 1943. New genera and species of North American linvphiid spiders. Bull. Univ. Utah Biol. Ser. 7(6). 39 pp. 1945. Some erigonid spiders of the genera Eularki and Diploccntrki. Bull. Univ. Utah Biol. Ser. 9(4). 19 pp. 1945a. Erigonid spiders of the genera Spirein- bohis, Disembohis. and Bactroceps. Trans. Conn. Acad. Arts and Sci. 36:215-35. 1946. On .several new American spiders. Bull. Univ. Utah Biol. Ser. 9(5). 15 pp. 1947. North American dictynid spiders: the ben- netti group of Amaurobius. Ann. Ent. Soc. Amer- ica 40:29-55. Chamberlin, R. v., and A. M. Woodbury. 1929. Notes on the spiders of Washington County, Utah. Proc. Biol. Soc. Washington 42:131-42. DoNDALE, C. D. 1961. Revision of the aureolus group of the genus Philodromus in North America. Cana- dian Ent. 93:199-222. DoNDALE, C. D., AND J. H. Redner. 1968. The imbe- cillus and nifus group of the spider genus Philo- dromus in North America. Ent. Soc. Canada, Memoir 55. 78 pp. 1975. The genus Ozyptila in North America. J. Arach. 2:129-81. 1976. A review of the spider genus Philodromus in the Americas. Canadian Ent. 108:127-57. 1978. Revision of the Nearctic Wolf Spider genus Schizocosa. Canadian Ent. 110:143-81. 1979. Revision of the wolf spider genus Alopecosa Simon in North America. Canadian Ent. 111:1033-55. 1982. The sac spiders of Canada and Alaska. Can- ada Dept. Agr. Bull. 1724. 194 pp. ExLiNE, H. 1936. Nearctic spiders of the genus Ciciirina Menge. American Mus. Novitates 850. 25 pp. Gertsch, W. J. 1932. A new generic name for Coria- rachne versicolor Keyserling, with new species. American Mus. Novitates 563. 7 pp. 1933. Notes on American spiders of the family Thomisidae. American Mus. Novitates .593. 22 pp. 19.34. Notes on American Lycosidae. American Mus. Novitates 693. 25 pp. 19.34a. Notes on American crab spiders. Ameri- can Mus. Novitates 707. 25 pp. 19.34b. Some American spiders of the family Hah- niidae. American Mus. Novitates 712. 32 pp. 1934c. Further notes on American spiders. Amer- ican Mus. Novitates 726. 26 pp. 1935. Spiders from the southwestern United States, with descriptions of new species. Ameri- can Mus. Novitates 792. 29 pp. 1941. New American spiders of the familv Club- ionidae. Part 1. American Mus. Novitates 1147. 20 pp. 1942. New American spiders of the family Club- ionidae. Part III. American Mus. Novitates 1195. 18 pp. 1946. Notes on American spiders of the familv Dictynidae. American Mus. Novitates 1319. 21 pp. 1953. The spider genera Xijsticus, Coriarachne, and Oxtjptila (Thomisidae, Misumeninae) in North America. Bull. American Mus. Nat. Hist. 102:417-82. 1960. Descriptions of American spiders of the family Smyphytognathidae. American Mus. Novi- tates 198l' 40 pp. Gertsch, W. J., and W. Ivie. 1936. Descriptions of new American spiders. American Mus. Novitates 858. 21pp. 1955. The spider genus Neon in North America. American Mus. Novitates 1743. 17 pp. Gertsh, W. J., and F. E. Russell. 1975. Loxosceles deserta Gertsch. Toxicon 13:203-4. Gertsch, W. J., and H. K. Wallace. 1937. New Ameri- can Lycosidae with notes on other species. Amer- ican Mus. Novitates 919. 22 pp. Hatley, C. L. 1978. The role of vegetation architecture in determining spider community organization. Unpubublished thesis, Utah State Univ., Logan. Helsdingen, p. J. VAN 1981. The Nearctic species of Oreonetides. Bull. Amer. Mus. Nat. Hist. 170:229-41. IviE, W. 1967.' Some synonyms in American spiders. J. New York Ent. Soc. 75:126-31. 1969. North American spiders of the genus Batli- ijphantes. American Mus. Novitates 2364. 70 pp. Kaston, B. J. 1946. North American spiders of the genus Ctenium. American Mus. Novitates 1.306. 19 pp. Knowlton, G. F. 1971. Some Curlew Valley spiders. Utah State Univ. Ecol. Center, Terrestrial Ar- thropods, Mimeo. Ser. 2. 5 pp. Leech, R. 1972. A revision of the Nearctic Amauro- biidae. Canadian Ent. Soc. Memoir 84. 182 pp. Levi, H. W. 1953. New and rare Dipoena from Mexico and Central America. American Mus. Novitates 1639. 11 pp. 1954. Spiders of the genus Eurijopis from North and Central America. American Mus. Novitates 1666. 48 pp. 1955. The spider genera Coressa and Achaea- ranea in America north of Mexico. American Mus. Novitates 1718. 33 pp. 1957. The spider genera Enoplognatha, Theri- dion, and Paedisca in America north of Mexico. Bull. Amer. Mus. Nat. Hist. 112:1-12. 1958. Spiders of the genus Arctachaea. Psyche 64:102-6. 1959. Problems in the spider genus Steatoda. Syst. Zool. 8:107-16. 1963. American spiders of the genus Achaearanea and the new genus Echinotheridion. Harvard Univ. Bull. Mus. Comp. Zool. 129:187-240. 1975. Additional notes on the orb-weaver genera Araneus, Hypsosinga, and Singa north of Mexico. Psyche 82:265-74. 1975a. The American orb-weaver genera Larinia, Cercidia, and Mangora north of Mexico. Harvard Univ. Bull. Mus. Comp. Zool. 147:101-35. 1980. The orb-weaver genus Mecijnogea, the sub- familv Metinae and the genera Pachijgmitha, Gle- nognatlia, and Azilia of the subfamily Tetragna- thinae north of Mexico. Harvard Univ. Bull. Mus. Comp. Zool. 149:1-75. 522 Great Basin Naturalist Vol. 43, No. 3 1981. The American orb-weaver genera Dolichog- natha and Tetragnatha north of Mexico. Harvard Univ. Bull. Mus. Comp. Zool. 149:271-318. . 1981a. More on the genus Araneus from North America. Bull. Amer. Mus. Nat. Hist. 170:254-6. Levi, H. W., and L. R. Levi. 1951. Report on a collec- tion of spiders and harvestmen from Wyoming and neighboring states. Zoologica .36:219-37. Levi, H. W., and D. E. Randolph. 1975. A key and checklist of American spiders of the family Theri- diidae north of Mexico. J. Arach. 3:31-51. LowRiE, D. C, and C. D. Dondale. 1981. A revision of the nigra group of the genus Pardosa in North America. Bull. Amer. Mus. Nat. Hist. 170:125-39. MiLLiDGE, A. F. 1980. The erigonine spiders of North America. Part 2. The genus Spirembolus Cham- berlin. J. Arach. 8:109-58. 1981. The erigonine spiders of North America. Part 3. The genus Scotinotyhis Simon. J. Arach. 9:167-213. 1981a. The erigonine spiders of North America. Part 4. The genus Disembolus Chamberlin and Ivie. J. Arach. 9:259-84. 1981b. The erigonine spiders of North America. Part 5. The genus Satilathis. Bull. Amer. Mus. Nat. Hi.st. 170:243-53. (In press) The erigonine spiders of North Ameri- ca. Part 6. The genus Wolckenaeha Blackwall. J. Arach. MuMA, M. H., and W. J. Gertsch. 1964. The spider family Uloboridae in North America north of Mexico. American Mus. Novitates 2196. 43 pp. Platnick, N. L and M. U. Shadab. 1975. A revision of the spider genus Gnaphosa in America. Bull. Amer. Mus. Nat. Hist. 155:1-66. 1976. A revision of the spider genera Drassodes and Tivodrassus in North America. Amer. Mus. Novitates 2593. 29 pp. 1976a. A revision of the spider genera Rach- odrassus, Sosticiis, and Scopodes. Amer. Mus. Novitates 2594. 33 pp. 1977. A revision of the spider genera Herpi/Uiis and Scotophaeits in North America. Bull. Amer. Mus. Nat. Hist. 159:1-44. 1980. A revision of the spider genus Cesonia. Bull. Amer. Mus. Nat. Hist. 165:337-85. 1980a. A revision of the North American spider genera Nodocion Litopyhis, and Synaphosus. Amer. Mus. Novitates 2691. 26 pp. 1981. A revision of the spider genus Sergiolus. Amer. Mus. Novitates 2717. 41 pp. 1982. A revision of the American spiders of the genus Drassyllus. Bull. Amer. Mus. Nat. Hist. 173:1-97. (In press) A revision of the American species of the genus Zetotes. Bidl. Amer. Mus. Nat. Hist. Sauer, R. J., and N. I. Platnick. 1972. The crab spider genus Ebo in the United States and Canada. Ca- nadian Ent. 104:.35-60. Schick, R. X. 1965. The crab spiders of California. Bull. Amer. Mus. Nat. Hist. 129:1-180. Waagen, G. N. 1979. Changes in spider community at- tributes along a subalpine successional gradient. Unpubublished dissertation, Utah State Univ.^ Logan. Wallace, H. K., and H. Exline. 1977. Spiders of the genus Pirata in North America, Central America, and the West Indies. J. Arach. 5:1-112. NOTICE TO CONTRIBUTORS Manuscripts intended for publication in the Great Basin Naturalist or Great Basin Natural- ist Memoirs must meet the criteria outhned in paragraph one on the inside front cover. 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See Notice to Contributors on the inside back cover. 1-84 650 68551 ISSN 017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 43 October 31, 1983 No. 4 ALPINE AND SUBALPINE WETLAND PLANT COMMUNITIES OF THE UINTA MOUNTAINS, UTAH George M. Briggs' and James A. MacMahon- Abstract.— Seven wetland areas in the subalpine and alpine region^ of the Uinta Mountains are described. Most areas have a mosaic vegetation structure, comprised of several patches that are usually sharply demarcated from one another. Each patch has a distinct species composition, usually with only one or two species. Water level is associ- ated with some of the vegetation patterns found in these sites. Standing crop varied from 28 to 360 g/m^. Sites show considerable variation in standing crop that could not be explained by elevation differences between sites. Sites with water flowing over them have substantially greater standing crops than sites where water stagnates. Seasonal pat- terns in shoot density and standing crop indicate one late summer peak in standing crop and little recruitment of shoots over the summer. On stagnant sites, the average stem weight shows a strong relationship to stem density. This pattern did not appear to be caused by thinning mortality and did not follow a "3/2 power law" pattern. A conspicuous aspect of many of the west- em North American mountain ranges is their wetlands. The Uinta Mountains of north- eastern Utah have an abundance of wetland areas, from the lower elevations (around streams in sagebrush slopes) to areas near springs at high elevations in the alpine zone. The majority of wetlands are found at the heads of glaciated valleys in a zone just above and below treeline. The vegetation in these areas is dominated by members of the Cyperaceae (sedges), as is much of the upland region in the alpine zone of the Uintas. Al- though both wetland and upland regions are dominated by Carex spp., the structure of the two regions is markedly different (Briggs and MacMahon 1982). In this study we describe a variety of sedge-dominated wetlands, detail some of their structural attributes, and dis- cuss some of the factors that we think are im- portant in determining their patterns of occurrence. Study Sites and Methods All sites are in the Uinta Mountains of northeastern Utah (40°45'N, 110°-111°W), the largest east-west trending mountain range in North America. Bedrock throughout the upper elevations of the range is quartzite. The range was heavily glaciated in Pleisto- cene time (Hansen 1975) and is dominated by numerous large U-shaped valleys ending abruptly in steep-walled cirques. No attempt was made to describe or cata- log all the wetland areas of the Uintas. In- stead, we chose to study a variety of wetland regions that varied in their species composi- tion and standing crop. Most of these regions had a patchwork appearance because they were composed of regions (2-100 m^) of dis- tinct species composition, often monotypic, which abruptly gave way to regions of a dif- ferent species mixture. In sampling these areas, we first made a rough map that demar- cated the "patches" found in them. Each 'University of Michigan Biological Station, Pellston, Michigan 49769. 'Department of Biology, Utah State University, Logan, Utah 84322. 523 524 Great Basin Naturalist Vol. 43, No. 4 patch was then sampled by the methods out- lined below. We refer to each patch as a site and present figures representing the species composition and standing crop of each of these. The larger units of vegetation (the en- tire meadow composed of several patches) are not quantitatively described in this paper. Each site was given a two-letter code that represented the area from which it came and a number that indicated the specific patch sampled (e.g., SC-1 is patch 1 in area SC). The boundaries to each site were sub- jectively demarcated and a point was picked within the stand. A 20 X 50 cm frame was placed at that point and all vegetation (ex- cept bryophytes) was clipped at ground level. Nine additional samples were taken at regu- lar intervals from the first point. These sam- ples were sorted to species in the field, re- turned to the lab, air dried for at least two months, and oven dried at 40 C for at least 24 h. Sorting to species was quite easy, even when based on vegetative characters, since there were rarely more than two species per sample. For larger species, the number of stems was directly counted and the average weight per stem calculated. For the smaller species, the average weight per stem was de- termined by weighing three replicates of 100 stems. The total number of stems was obtain- ed by dividing the total weight of that spe- cies in the sample by the weight/ stem. All sites were sampled in August, when the fruits of the dominant species were mature. Sea- sonal changes in aboveground standing crop, number of shoots, and number of fruiting shoots were determined in three of the sites by sampling three or four times during the summer. In addition, several individual plants were observed. The length of each leaf and the length and condition of the fruiting culm (if present) were measured at each sampling time. At each site, soil samples were dug and pH of the soil was determined. All these sites were submerged at least part of the summer, and notes were made of both the length of time the site was submerged and the degree of water movement over the site. Site Descriptions Seven wetland areas were mapped and 21 sites were sampled within these areas. Each wetland area contains between one and five sites. Table 1 lists the dominant species, ele- vation, and standing crop of all 21 sites. Fig- ure 1 maps the vegetational patterns found in the more complex areas. The highest wetland found was at an ele- vation of 3,768 m, in an area adjacent to the steep, rocky talus pile that makes up the northeast side of King's Peak in the east cen- tral section of the Uinta range. Springs moist- en the site all summer and feed a small stream that traverses the site. Large rocks are scattered throughout. The wetlands of this re- gion consist of a series of small irregularly shaped pools containing 1-10 cm of water at the end of the summer. The emergent vege- tation consists solely of Eriophorum scheiich- zeri Hoppe. These pools were sampled as site ER-1. To the north of King's Peak lies Henry's Fork Basin. The head of this basin has large expanses of sedge meadows and willow thick- ets interrupted by upland regions. In spite of a homogeneity in species composition (much of the area is dominated by Carex aquatilis), there is considerable variation in stature and standing crop of wet meadows within upper Henry's Fork Basin. We sampled two sites that represent extremes in Carex aquatilis meadows. HF-1 is a border to a small (20 m diameter) pond at an elevation of 3,278 m. Vegetation consisted of robust individuals of Carex aquatilis and Caltha leptosepala. Wa- ter stands at least 10 cm deep throughout the year, draining off the site to the north. HF-2 is a very different Carex aquatilis region, with a more dense stand of much smaller plants. There is no proximate open water and the ground surface was dry at the end of the summer although the ground was saturated with water at a depth of 2-5 cm. The remaining five areas are within four miles of Mirror Lake on the west end of the Uintas. Area TL is part of a circular meadow approximately 200 m in diameter. The mead- ow has a small stream meandering through it and has numerous small "oxbow lakes" and "kettleholes." The vegetation displays a pat- tern commonly found in moist subalpine meadows of the Uintas: monotypic stands of Carex aquatilis and Eleocharis pauciflora, areas of Carex aquatilis and Eleocharis pau- ciflora mixed, and sections of bare ground, all October 1983 Briggs, MacMahon: Uinta Plant Communities 525 surrounded by an upland that has soil with less organic matter and a vegetation of Carex illota L. H. Bailey, Deschampsia cespitosa, and Ligusticwn filicinum var. tenuifolium (S. Wats.) Mathias & Constance. A large portion of this area has standing water year around. This area was sampled in five sites (Fig. 1). Areas FT and SE are small meadows in Picea enge/mannii-dominated forests. The vegetation in them is similar to that in TL and is mapped in Figure 1. Both areas were sampled as four sites. Area SC is a meadow in a Picea engelmannii-Pinns contorta-domi- nated forest. The meadow has a stream flow- ing through it and is kept moist by two large seepage areas. The vegetation presents a complex patterning, with part of the area ex- hibiting the mix described at the TL site. Table 1. Elevation, standing crop, and species found on the sites. Site Elevation (m) Standing crop (g/m2) Dominant species (> 10 percent standing crop) Other species ER 3,768 28 Eriophorum scheuchzeri Hoppe. None HF-1 3,278 360 Carex aquatilis Wahl. Caltha leptosepala DC. None HF-2 3,278 103 Carex aquatilis Eleoeharis pauciflora (Lightf.) Link, Pedicularis groenlandica Retz. SE-1 3,260 172 Carex aquatilis Carex eanescens L. None SE-2 3,260 234 Carex aquatilis None SE-3 3,260 264 Carex aquatilis None SE-4 3,260 106 Eleoeharis pauciflora Carex aquatilis Pedicularis groenlandica FT-1 3,256 152 Eleoeharis pauciflora Carex aquatilis Caltha leptosepala, Pedicularis ■-2 3,256 ■-3 3,256 -4 3,256 104 114 Carex aquatilis Eleoeharis pauciflora Carex illota L. H. Bailey Ligusticwn filicinum (S. Wats.) Mathias and Constance groenlandica, Dodecatheon pulchellum (Ref.) Merrill, Epilobium alpinum L., Agrostis alba L. Eleoeharis pauciflora Carex aquatilis Carex aquatilis, Eleoeharis pauciflora, Caltha leptosepala, Pedicularis groenlandica, Veronica wormskjoldii Roem. and Schult. TL-1 3,110 86 Carex aquatilis Deschampsia cespitosa (L.) Beauv. TL-2 3,110 96 Carex illota Deschampsia cesp itosa Eleoeharis pauciflora, Caltha leptosepala, Ligusticwn filicinum, Pedicularis groenlandica, Veronica wormskjoldii TL-3 3,110 139 Eleoeharis pauciflora None TL-4 3,110 97 Carex aquatilis Eleoeharis pauciflora None TL-5 3,110 92 Carex aquatilis None AQ 3,085 141 Carex aquatilis None SC-1 3,073 194 Scii-pus cespitosus L. Caltha leptosepala Carex nigricans A. Meyer, Pedicularis groenlandica, Veronica wormskjoldii SC-2 3,073 109 Scirpus cespitosus Carex aquatilis Eleoeharis pauciflora Caltha leptosepala, Ligusticum filicinum, Pedicularis groenlandica, Veronica wormskjoldii SC-3 3,073 83 Carex aquatilis Eleoeharis pauciflora None SC-4 3,073 251 Carex illota Carex rostrata Stokes Carex aquatilis Caltha leptosepala Agrostis alba, Carex eanescens, Ligusticum filicin um 526 Great Basin Naturalist Vol. 43, No. 4 Other parts of the meadow have a dense turf contains a mix of Carex illota and C aaua- of Scirpus cespuosus L. associated with Carex tilis. The whole meadow has standing wa"er aqtuitths Eleochans pauciflora, and Caltha throughout the year. SC was diviL fnto leptosepala. A final portion of the meadow four sites (Fig. 1). u^^ CAAO -"r"" °^ '^"r'n^t '''"^ ^' ^'"'^ ^^- ^' ^'^^ ^L. C, Area SC. D, Area FT. ARMO = Arnica t^ieocnans pauciflora, JUPA - /uncti* parryi, POPA = Po« pra/en^., SCCE = Scirpus cespitosa. October 1983 Briggs, MacMahon: Uinta Plant Communities 527 Area AQ is on the margin of a small pond that narrows into a stream at one end. It is surrounded by a Picea engelmannii-Pinus contorffl-dominated forest. This area is at least 30 cm imder water in the spring. Stand- ing water is present to a depth of 10 cm in the fall. One site was sampled in this area, a region of nearly pure Carex aquatilis. Results and Discussion Vegetation The number of species sampled on each of the 21 sites (Table 1) varied from one to sev- en. Most of the species present in the wet- land sites are common species of the western United States. The Carex species collected can be separated, phytogeographically, into two groups: western cordilleran species {Carex nigricans, C. illota) and circumboreal species (C. rostrata, C. aquatilis, and C canescens). One of the striking features of these wet- lands is the dominance by plants that are ca- pable of extensive spread by rhizomes. The mosaic structure (Fig. 1) of these commu- nities is a function of the strongly rhizo- matous nature of the dominant plants and some, perhaps most, of the "patches" seen represent single plants (genets. Harper 1977) formed by growth from a single propagule. The patterns seen in these wetlands are prob- ably dependent on both the history of prop- agule arrival and on variations in environ- mental conditions affecting the success of particular species. A complicating factor re- sults from the "inertia" of these sites— a re- sistance to vegetational change resulting from both the harsh nature of the climate and the vigorous mode of wetland plant growth. Both these factors could make established vegetation patterns difficult to disrupt. Al- though seedling establishment is common in some wetlands (Lieffers and Shay 1982), both Costello (1936) and Bernard (1975) noted that seedling establishment was rare in some Carex wetlands. Our sites included areas where the dominant was in very poor condi- tion (e.g., HF-2) or had died off completely, leaving areas of bare ground (e.g., part of TL). Both biotic and abiotic factors at these sites may change with time. These changes will sometimes be to the detriment of the species that has been dominant. But, because of the difficulty in establishing additional species, the dominant may continue to exist in a depauperate condition and may even die back completely before another species is able to invade the area. Thus, the species present today may not reflect present condi- tions but may represent conditions of an ear- lier time. One environmental variable that we found associated with community structure is water level. In TL and FT the Carex illota section was always upland relative to the Carex aquatilis-Eleocharis pauciflora sections. In SC the part of the plot containing Scirpus cespitosus was on higher ground than those regions containing C. aquatilis and Eleocharis pauciflora. Some species distributions could not* be associated consistently with water lev- el. Carex aquatilis could be found in areas with both more and less standing water than the often adjacent E. pauciflora areas (areas TL, FT). Carex illota, although usually re- stricted to areas with no standing water (FT- 4, TL-2), sometimes was found in areas 5 cm deep in running water (SC-4). It is generally thought that the influence of water level in wetland regions is based on variations in waterlogging and aeration. Soil aeration may influence plants either by af- fecting root respiration (Mendelssohn et al. 1981), which could affect water and nutrient flow through roots, or by affecting soil nutri- ent status. The influence on soil nutrition can occur directly, by influencing the chemical forms available and their solubilities (Jones 1971, 1972), or indirectly, by affecting soil microbial activity. The importance of aeration on species dis- tribution is represented in our observations on Carex illota, which is intolerant of water- logged soils if the water is stagnant (and pre- sumably poorly aerated), yet grows quite well under 5 cm of water when it is flowing over the site (and probably better aerated). Further evidence for the importance of aera- tion in these sites is shown in standing crop data presented below. If aeration does in- fluence the structure of these communities, the complex nature of some of these sub- alpine sites could be due to underground wa- ter flow patterns and resultant aeration pat- terns. These flow patterns may be due to 528 Great Basin Naturalist Vol. 43, No. 4 300- 200 100- HF-I • SE-3 • SC-4 • SE-2 • SC-I • SE-I • FT-I • AQ SC-2 TL-4 SC-3 I TL-3 FT-2 TL-5 FT-3 o^ .• PT-4 SE.4«#HF.2 TL-2: TL-I • ER I Running water year-round Running water in spring, standing water all summer Standing water all summer Standing water half of summer Standing water only in early summer MOISTURE CLASS Fig. 2. Graph showing the relationship between standing crop on a site and a measure of water movement over that site. springs and may not be readily discemable without measurement of soil conditions. The peak aboveground standing crop val- ues (Table 1) varied considerably, both be- tvi'een areas and also between adjacent sites in one area (e.g., area SC). The very low standing crop value for site ER was undoubt- edly due to the short growing season and low temperatures associated with this, the highest site. However, aside from site ER, there is very little relationship between elevation and peak aboveground standing crop. Gorham (1974) derived a regression equation relating standing crop of Carex meadows to the high- est monthly mean temperature of the year. His data on 11 pure Carex stands (on both rich and poor soils) fit the regression line well (r = 0.84). The data of Auclair et al. (1976) on wetland stands that were not pure Carex also fit this regression. Using weather data for areas in Colorado that are comparable to ours, Gorham 's equation predicts a standing crop of 292 g/m- for a site at 3,109 m (10,200 ft) and 235 g/m^ for a site at an ele- vation of 3,566 m (11,200 ft). The standing crop values that we found are both above and below these values, with most sites be- low. Gorham 's equation is most appropriate for those sites where water is not stagnant (HF-1, SE-2, SE-3, SC-4). The sites with less water movement have less standing crop than predicted by Gorham. The reduced growth on sites where water stagnates is shown in Figvire 2, where standing crop is plotted against a gradient in the amount of water movement through the site. Reduced growth on the stagnant sites is probably due to low oxygen levels that may be disturbing root res- piration, mineral uptake, or soil nutrient status. An oxygen-limited situation in Carex meadows might explain the timing of nutri- ent uptake in wetlands. Boyd (1970) and Ber- nard and Solsky (1977) note considerable October 1983 Briggs, MacMahon: Uinta Plant Communities 529 300H CM E Qi O c I lOOH (0 1 — r 1 I' 15 5 2 3 JULY AUG. SEPT. OCT. Fig. 3. Seasonal trends in aboveground standing crop for two of the pure Carex aqtiatilis sites, SE-2 and SE-3. mineral uptake by wetland plants in the early spring. Boyd hypothesized that this early mineral uptake (preceding the growth peri- od) was an adaptation to allow certain spe- cies to procure nutrients before other species. Another possible reason for mineral uptake early in the spring could be related to oxygen concentrations. In the early spring, oxygen in wetland soils may be at its highest levels be- cause of well-oxygenated runoff waters and because low temperatures increase the amount of oxygen that water can hold. Thus, plants may take up nutrients in the early spring because that is the most favorable time for root respiration. In several of the Carex aquatilis-domina.ied sites, we observed the variation in standing crop during the summer. The typical pattern in these subalpine meadows (Fig. 3) includes a single peak in aboveground standing crop which occurs in late summer. Gorham and Somers (1973) and Bernard and MacDonald (1974) also found such a pattern for wetlands composed of C. aqimtilis and C. lacustris, re- spectively. On our sites there was relatively little new shoot production over the course of the summer (Fig. 4), and the changes in aboveground standing crop were being caused by the growth of individual shoots. Some Carex species (e.g., C. rostrata) have biannual periods of shoot production and Fig, Carex 4. Seasonal trends in stem density (stems/m^) for oquatilis on SE-2 and SE-3. these may be responsible for bimodal pat- terns in biomass (Gorham and Somers 1973), although other factors may cause bimodal patterns. The pattern in shoot production that we observed for C. aquatilis had been found previously for this species (Gorham and Somers 1973), as well as for C. lacustris (Bernard and MacDonald 1974), and would be expected in regions with short growing seasons such as the one we studied. Based on our observations, the life history of C. aquatilis is initiated by growth of a new shoot sometime during the winter or spring. The shoots grow throughout the summer and then overwinter. The following spring the old shoots are distinguishable from shoots of the current year by their large size and old leaves. One or two of these old leaves may undergo additional growth in the spring. At least some and possibly all of the shoots un- dergo two or possibly more seasons of growth before they flower or die. Many shoots may not flower. Floral initiation is evident very early in the spring; floral parts are visible less than two weeks after growth initiation. Dur- ing the growing season leaves are initiated, grow quickly to a maximum length, and re- main green until early September. There was considerable variation in plant density in the C. aquatilis stands. On the stagnant sites (those where water was not flowing), the variation in density was related to stem weight. If a plot is made of the natu- ral logarithm of plant density vs. the natural logarithm of plant weight (Fig. 5), it can be seen that the stagnant sites fall on a line 530 Great Basin Naturalist Vol. 43, No. 4 n (density) 6.3 JU -0.86 Fig. 5. Graph showing the relationship between stem density and average stem weight for five Carex aquatilis stands where water was stagnant. (r^ = 0.98). Sites with flowing water do not fall on this line and have substantially greater plant weights at a particular density than those on stagnant sites. The regression line of Figure 5 describes the following equation: [1] w = 5.53p-o86 where w = individual weight; p = plant density plant The form of this equation is characteristic of monotypic stands and an exponential value of -3/2 has been found for a wide variety of species of differing life forms and habitats (Gorham 1979, White 1981). The equation has been labeled the "3/2's thinning law" be- cause it describes the changes in density and plant weight that occur in stands during self- thinning. On our plots it is likely that the pattern of density and plant weight is not being caused by mortality (thinning) but rather by recruitment (see Fig. 4). Gorham (1979) cites studies with imcrowded condi- tions, and therefore little self-thinning, where the exponent value in equation (1) is closer to -1. Our stagnant sites appear to be similar areas. With an exponent of -1, standing crop (w o p) remains constant regardless of density. Such sites may represent areas where stand- ing crop is limited by factors (such as fer- tility?) that prevent densities to reach levels where self-thinning in the typical 3/2's fash- ion can occur. The precise factors that dic- tate the 3/2's thinning law are still not eluci- dated (White 1981). Perhaps studies on such areas as these, where the "law" does not hold, may help to clear up the problem. Literature Cited AUCLAIR, A. N. D., A. BuCHARD, AND J. PaJACKOWSKI. 1976. Productivity relations in a Corex-dominated ecosystem. Oecoiogia 26:9-31. Bernard, J. M. 1975. The life history of shoots of Carex lacustris. Canadian J. Bot. 53:256-260. Bernard, J. M., and J. G. MacDonald. 1974. Primary production and life history of Carex lacustris. Ca- nadian J. Bot. 52:117-124. Bernard, J. M., and B. A. Solsky. 1977. Nutrient cy- cling in a Carex lacustris wetland. Canadian J. Bot. 55:630-638. Boyd, C. E. 1970. Production, mineral accumulation and pigment concentrations in Typha kitifolia and Scirpiis americanus. Ecology 51:285-290. Briggs, G. M., and J. A. MacMahon. 1982. The struc- ture of alpine plant communities near King's Peak, Uinta Mountains, Utah. Great Basin Nat. 42:50-59. CosTELLO, D. F. 1936. Tussock meadows in southeastern Wisconsin. Bot. Gaz. 97:610-648. Gorham, E. 1974. The relationship between standing crop in sedge meadows and summer temper- ature. J. Ecol. 62:487-491. 1979. Shoot height, weight and standing crop in relation to density in monotypic stands. Nature 279:148-150. Gorham, E., and M. G. Somers. 1973. Seasonal changes in the standing crop of two montane sedges. Ca- nadian J. Bot. 51:1097-1108. Hansen, W. R. 1975. The geologic story of the Uinta Mountains. Geological Survey Bulletin 1291. U.S. Government Printing Office, Washington, D.C. 144 pp. Harper, J. L. 1977. The population biology of plants. Academic Press, London. 891 pp. Jones, H. E. 1971. Comparative studies of plant growth and distribution in relation to waterlogging. III. The response of Erica cinerea L. to waterlogging in peat soils of differing iron content. J. Ecol. 59: 583-591. 1972. Comparative studies of plant growth and distribution in relation to waterlogging. VII. The influence of water table fluctuations on iron and manganese availability in dune slack soils. J. Ecol. 61:107-116. Lieffers, V. J., AND J. M. Shay. 1981. The effects of wa- ter level on the growth and reproduction of Scirptis maritiimis var. paludosus. Canadian J. Bot. 59:118-121. Mendelssohn, I. A., K. L. McKee, and W. H. Patrick. 1981. Oxygen deficiency in Spartina alterniflora roots: Metabolic adaption to anoxia. Science 214:439-441. White, J. 1981. The allometric interpretation of the self- thinning rule. J. Theor. Biol. 89:475-500. UTAH FLORA: SALICACEAE Sherel Goodrich' Abstract.— A revision of the willow family, Salicaceae, is presented for the state of Utah. Included are 31 species and 5 subspecific taxa of indigenous and introduced plants. Keys to genera and species are provided, along with de- tailed descriptions, distributional data, and comments. No new taxa or combinations are proposed. This paper is another in a series of works leading to a definitive treatment of the flora of Utah. The willow family as represented in Utah is rather small when compared to sever- al other families, but its taxa cover the state, and it is complex. Herbarium specimens are frequently misidentified. Unisexual plants, ex- treme variation in leaves of fertile and vege- tative or short and long twigs, and early de- ciduous flowers all contribute to the difficulty in identification of taxa. Hybridiza- tion especially in Populus further complicates identification. Several members of the family are culti- vated for ornamental plants or shade trees. Not all of these are included in this treat- ment. Among those not treated are Populus candicans Ait. (Balm of Gilead), P. simonii Carr., and Salix viminalis L. (Golden Osier). Members of the family are important to many kinds of wildlife. For example, in Utah, beaver are almost totally dependent on the family. They utilize aspen, cottonwoods, and willows and avoid most other woody plants. Among the very few exceptions are probably Alnus and Betula. The arabic numerals following the dis- cussion of each taxon indicate the number of specimens examined in the preparation of this treatment. The roman numerals indicate the number of specimens collected by me. Acknowledgments Appreciation is expressed to Dr. Arthur Cronquist for permitting me to preview his Salix manuscript of the intermountain flora. Appreciation is also expressed to the direc- tors and curators of the following herbaria of Utah: Brigham Young University, Provo; For- est Service Herbarium, Ogden; Garret Her- barium, University of Utah, Salt Lake City; Intermountain Herbarium, Utah State Uni- versity, Logan. I appreciate the loan of speci- mens from each of these herbaria. These specimens are the basis of this work. Salicaceae Mirbel. Willow Family Dioecious dwarf shrubs to large trees; leaves alternate, simple, entire, serrate, cre- nate, rarely lobed, usually stipulate, but the stipules often readily deciduous; flowers borne in aments (catkins), without a perianth, each subtended by a small, scalelike bract (commonly referred to as a scale); staminate flowers of (l)2-many stamens; pistillate flow- ers of a single pistil with 2-4 carpels and as many stigmas; placentation parietal or basal; fruit a sessile or stipitate capsule with 2-4 valves; seeds numerous, small, covered with long white hairs, dispersed easily by wind. 1. Trees with pendulous aments; leaf buds covered by several, usually resinous scales; each flower subtended by a cup-shaped disk, without obvious glands; stamens 6 to many; scalelike bracts subtending the flowers laciniate or fimbriate (except in P. alba), otherwise glabrous or ciliate Populus 'Forest Service, U. S. Department of Agriculture, Intermountain Forest and Range Experiment Station, Ogden, Utah 84401, stationed in Provo, Utah, at the Shrub Sciences Laboratory. 531 532 Great Basin Naturalist Vol. 43, No. 4 — Trees, shrubs, or dwarf shrubs with mostly ascending to erect aments; leaf buds covered by a single nonresinous scale; each flower subtended by 1 or 2 basal glands, but without a disk; stamens (1)2-8, rarely more; scalelike bracts sub- tending the flowers entire or occasionally shallowiy toothed, usually densely pubescent Salix PopuLus L. valves, glabrous in our taxa except in P. hahamifera. Small to large trees; leaf buds covered by several overlapping scales, resinous in most References taxa; aments pendulous, mostly appearing be- Eckenwalder, J. E. 1977. North American cottonwoods fore the leaves, and often soon deciduous, the {Populm, Salicaceae) of sections Abaso and Aieg- i,.,, . -iij-j J iros. T. Arnold Arboretum 58(3): 194-208. scalelike bracts very quickly deciduous, deep- hjtchcock, C. L., and a. Cronc^u.st. 1964. Vascular ly lobed to laciniate, often dilated (entire or plants of the Pacific Northwest. Part 2: Salicaceae to nearly so and not dilated in P. alha); each Saxifragaceae. Univ. Washington Publ. Biol. Vol. 17. flower subtended by a cuplike disk; stamens „ ^'^' ,0=1 \n i r u ^ w j u 1 1 f I r Kehder, a. 1951. Manual ot cultivated trees and shrubs 6-60 or more, the filaments free; inserted on hardy in North America. Macmillan Co., New the disk; capsules pedicellate, with 2-4 York. 996 pp. 1. At least some of the mature leaves deeply 3-5 lobed and aceriform, often densely tomentose beneath; bracts of flowers entire or shallowiy toothed, long pilose-ciliate; twigs of the season and winter buds often white-woolly; stigma lobes slender; plants introduced, cultivated, and escaping P. alha — Leaves not deeply lobed, not aceriform, merely toothed, glabrous or nearly so; scales of flowers deeply lobed to lacerate 2 2(1). Bark white and smooth except blackened and rough where scarred, covered with a whitish powdery bloom; bracts of flowers more or less persistent, deeply lobed or cleft, ciliate with long white hairs; leaves orbicular to reniform- cordate; bud scales shiny but hardy resinous; stamens 6-14; capsules 4-6 mm long, with 2 carpels; stigmas slenderly lobed; plants not confined to water courses P. tremuloides — Bark turning gray or brown and roughly furrowed on older trunks; bracts of flowers laciniate-fringed, otherwise glabrous or inconspicuously short hairy; stamens 12-60 or more; capsules mostly longer, with 2-4 carpels; stigmas broadly dilated; plants mostly cultivated or growing along water courses or edges of lakes 3 3(2). Leaves 0.67-1.3 times longer than wide, deltoid to rhombic or ovate; petioles compressed laterally 4 — Leaves (1)1.2-7(10) times longer than wide, ovate to lanceolate; petioles terete or dorsiventrally compressed 6 4(3). Bud scales and twigs of the season pubescent; leaf blades commonly with 4-10(15) fine to coarse teeth on each side; branches widely spreading and the crown often as broad or broader than the tree is tall; plants native, sometimes cultivated, most common along the drainages of the Colorado River system, but sporadic along the Wasatch Front and elsewhere P. fremontii — Bud scales and twigs mostly glabrous; leaf blades commonly with 15-25(30) fine teeth on each side; branches ascending to erect and the crown mostly longer than wide; plants introduced, cultivated, sometimes persisting 5 5(4). Leaf blades rhombic-ovate, cuneate at the base, seldom over 7 cm long, capsules 2 valved; branches often comparatively small, strongly ascending to erect and the crowna narrow and columnar (in the trees planted in our area) P. nigra October 1983 Goodrich: Utah Flora, Salicaceae 533 — Leaf blades more or less deltoid or broadly ovate, broadly cuneate at the base, some regularly over 7 cm long; capsules 2 or more valved; branches large, spreading-ascending, the crown not columnar P. canadensis 6(3). Leaf blades distinctly darker above than beneath, very strongly resinous espe- cially when young, the petiole terete or nearly so; ovary and young fruit hairy or glabrous; stamens 30-60 P. balsamifera — Leaf blades about equally yellow-green on both sides; ovary and young fruit glabrous; stamens mostly 12-30 7 7(6). Leaf blades (1.8) 2.5-6 (9.5) times longer than wide; petioles 1/5-1/3 (2/5) as long as the blades, dorsiventrally compressed; carpels 2 P. angustifolia — Leaf blades 1-2.4 times as long as wide; petioles 1/5-3/4 as long as the blades, subterete or somewhat flattened; carpels 2 or 3; plants hybrids, intergrading into P. angustifolia on one hand and into P. fre7nontii and other broad-leaved poplars on the other P. acuminata Popuhis acuminata Rydb. Lanceleaf Cot- tonwood. A series of hybrids between P. an- gustifolia and P. fremontii and other taxa with broad leaves, with features intermediate between the parents and intergrading into P. angustifolia on one hand and into the broad- leaved parent on the other; petioles com- monly (1.5) 2.5-5.5 (6.5) cm long, 1/5-3/4 as long as the blade; leaf blades 1-2.4 times longer than wide. Along streams and rivers, edges of ponds and lakes, often in mouths of canyons where the parental types come to- gether, probably cultivated, from (1370) 1525-1920 m, in Box Elder, Cache, Du- chesne, Emery, Garfield, Iron, Kane, Salt Lake, San Juan, Sevier, Uintah, Utah, Wasatch, Washington, Wayne counties; throughout the range of P. angustifolia. The name P. acuminata in the strict sense is ap- plied to crosses of P. angustifolia and P. del- toides Marsh, var. occidentalis Rydb. It is used here in a broad sense to include crosses with other broad-leaved taxa, including P. balsamifera and P. fremontii; 29(0). Populus alba L. White poplar. Trees spreading by root sprouts, to about 30 m tall, the tnmk to 1 m or more in diameter, the branches usually spreading, the crown more or less roimded; bark gray-green to whitish and smooth on upper parts of the trunk and branches, rough and furrowed and turning blackish on lower parts of old trunks; twigs tomentose or glabrous; buds tomentose; pet- ioles terete 1-5 cm long, 0.2-0.6 times as long as the blade; leaf blades longer than wide, deltoid-ovate in outline, undulate toothed to deeply palmately 3-5 lobed and a-ceriform, the lobes serrate or crenate, the two primary lateral lobes sometimes has- tately lobed, dark green above, silvery white- tomentose beneath or glabrous; aments ap- pearing before or with the leaves, the rachis pilose-tomentose, the bracts entire to toothed, not laciniate, ciliate-fringed with long-pilose hairs, very quickly deciduous; staminate aments 8 cm long or more, the flowers with 6-10 stamens; pistillate aments 4-9 cm long; capsules 2-5 mm long, glabrous, 2-3 valved, the pedicels about 1(2) mm long; stigmas 2, each 2 lobed, the lobes linear, not dilated. Introduced from Eurasia, cultivated, escaping, and more or less natu- ralized, in populated areas, along fencelines, ditchbanks, and abandoned homesteads and fields, up to about 1980 m, to be expected in all counties of the state. Trees with leaves densely white-tomentose beneath are refe- rable to var. alba. Those with leaves and twigs glabrous or glabrate and fastigiate crowns are referable to var. bolleana Lauche. These may be hybrids between P. alba and some other species; 21 (ii). Populus angustifolia James Narrowleaf Cottonwood. Trees about 7-15(20) m tall, the trunk 30-60(80) cm in diameter, the branches erect-ascending, the crown more or less pyra- midal; bark pale green to whitish when young, furrowed and grayish on old trunks, twigs glabrous or pubescent; buds ovoid-con- ic, pointed, strongly resinous, reddish brown, glabrous or pubescent; petioles semiterete or horizontally flattened and channeled above. 534 Great Basin Naturalist Vol. 43, No. 4 especially near the blade, 3-25 mm long, up to 0.3 (rarely 0.4) times as long as the blade; leaf blades 4-14 cm long, 0.7-2.5 (4.0) cm wide, (1.8) 2.5-6 (9.5) times longer than wide, lanceolate or occasionally narrow elliptical or ovate, glabrous or nearly so, usually acute at the apex, roimded at the base, the margins finely to coarsely serrate; aments often devel- oping with the leaves, the rachis glabrous or nearly so, the bracts broadly obovate, deeply and irregularly lacrate; staminate aments 2-6 cm long, the flowers with 12-20 stamens; pistillate aments 6-10 cm long; capsules 3-6(7) mm long, 2 valved, glabrous, the pedi- cels about 2-10 mm long; stigmas 2, dilated, irregularly lobed. Along water courses, often in canyons, from about 1525-2135 (2440) m, in all counties of the state. Rather freely crossing with the broad-leaved species of the genus; 79 (i). Populus balsamifera L. Balsam poplar, Black Cottonwood. [P. trichocarpa T. & G.]. Tree 15-30(50) m tall; the trunk mostly 0.6-1 (1.5) m in diameter, bark furrowed and gray- ish on older trunks; buds large, the scales very resinous, glabrous or inconspicuously puberulent; petioles more or less terete, 2-6.2 cm long, 1/4-3/4 as long as the blade; leaf blades 4.3-11 cm long, 3.2-8 cm wide, 1.3-2.6 times longer than wide, ovate- accuminate, cuneate to cordate at the base, the margins crenulate, sometimes short ci- liate, strongly resinous, glabrous at maturity on both sides, the upper side dark green, the lower side distinctly paler and often rufous tinged in dried specimens; bracts of aments lacerate-fringed, otherwise glabrous or some- times with minute hairs, these not over 0.5 mm long; staminate aments 2-3(5) cm long, readily deciduous; stamens commonly 30-60; pistillate aments 8-20 cm long; cap- sules 5-8 mm long, glabrous or pubescent, subsessile; stigmas broadly dilated. Along streams, mostly in canyons and cultivated, 1370-2350 m, in Cache, Juab, Salt Lake, Se- vier, Utah, Wasatch, and Wayne counties; widespread in North America from New- foundland south to New York and west to Alaska (ssp. balsamifera), and from Alaska south to Baja California in the western part of the continent (ssp. trichocarpa). The native trees of our area are expected to be ssp. trichocarpa (T. & G.) Brayshaw with mostly pubescent and 3 (rarely 2-4) carpellate cap- sules. Some of the cultivated trees might be ssp. babamifera with mostly glabrous and 2 (rarely 3-4) carpellate capsules; 9 (0). Populus X canadensis Moeneh. Carolina poplar. Gray poplar. Cultivated and per- sisting, rarely escaping, to 40(50) m tall, the trunk 0.75-1.5(2) m in diameter; bark deeply furrowed and grayish on old trunks; buds large, the scales glabrous, but resinous; pet- ioles laterally flattened 3.5-8.5 cm long, 1/3 to as long as the blade; leaf blades mostly 3.5-11.5 cm long, 3.5-11 cm wide, or much larger on stump sprouts, 0.9-1.3 (rarely to 1.5) times as long as wide, deltoid-ovate, acuminate at the apex, mostly broadly cu- neate or trvmcate at the base, the margin cre- nate-serrate; glabrous and equally green on both sides; staminate aments about 7 cm long; stamens 15-25; pistillate aments un- known. Cultivated for shade trees, probably originated in France as a cross between P. deltoides Marsh, and P. nigra (Rehder, 1951), to be expected in nearly all counties of the state. Populus deltoides might also be ex- pected in the state as an introduced tree from the Plains and eastward, but no specimens were seen that were clearly assignable to that taxon. The original Carolina poplar was P. deltoides, but for many years the nursery stock distributed under that name has been P. X canadensis (Hitchcock and Cronquist, 1964); 15 (ii). Populus fremontii Wats. Fremont cotton- wood. Trees 10-25 m tall with broad rounded crowns, the crown often as broad or broader than the tree is high, the trunk 0.5-1 (1.5) m in diameter; bark smooth and whitish on young trees and on twigs and young branches, deeply furrowed and grayish or brownish on old trunks; petioles (0.8) 3-9.5 cm long, one half to as long as the blade, flattened; rarely with two glands at the summit; leaf blades (2) 4-10 cm long, (15) 4.5-12.5 cm wide, or much larger on sterile sprouts, 0.67-1.2 times as long as wide, del- toid, ovate, rarely nearly rhombic, with trun- cate, cuneate, or occasionally cordate base, acuminate at the apex, coarsely to finely cre- nate or serrate with about 8-11 (15) glandu- lar teeth, glabrous, greenish or yellow-green on both sides, turning yellow in autumn; staminate aments 4-10 cm long, the flowers October 1983 Goodrich: Utah Flora, Salic ace ae 535 with a broad oblique disk and 50-80 stamens with dark red anthers; pistillate aments 5-15 cm long, the flowers with a cup-shaped disk, this to 5 mm wide in fruit; capsules 7-10 (12) mm long, to 8 mm wide, ovoid to subglobose, 3 to 4 valved, glabrous, the stipes 2-6 (10) mm long; stigmas strongly di- lated and irregularly lobed. Along flood plains of rivers and along washes, irrigation ditches, and occasionally cultivated, from 762 to about 1860 m, in Cache, Duchesne, Gar- field, Grand, Iron, Kane, Salt Lake, San Juan, Sevier, Tooele, Uintah, Utah, Washington, Wayne, and Weber counties. The Fremont Cottonwood is abundant along the Colorado, Green, San Juan, and Virgin rivers and their tributaries within the Colorado Drainage, to be expected anywhere in the state as it has been cultivated for a shade tree. This tree is part of a transcontinental complex, of which P. arizonica Sarg., P. deltoides, P. sargentii Dode, and P. wislizeni (Wats.) Sarg. are a part. Popuhis arizonica and P. wislizeni have generally been considered closely allied to P. fremontii and they have by some authors been included as varieties of or as synony- mous with P. fremontii. Specimens that have capsules with stipes up to 6 or even 10 mm long are found in Emery County and other points along the Colorado River system. These trees have been referred to as P. fre- montii var. wislizeni Wats. Based on the long stipes, these trees have recently been as- signed to P. deltoides var. wislizenii (Wats.) Eckenwalder (Eckenwalder, 1977). However, these trees are like P. fremontii in the lack of glands at the jimction of petiole and blade and with few, broad, and coarse teeth on leaf margins. Based on my provincial study, I am not well prepared to make a judgment as to the specific assignment of these trees, but I prefer the traditional approach. If P. fre- montii is to be kept separate at all from P. deltoides, I feel these plants are best kept as a part of P. fremontii., 98 (ii). Populus nigra L. Black poplar. Tree to 30 m tall; bark deeply furrowed and grayish on old trunks; bud scales glabrous, resinous; petioles flattened laterally, slender 1-4.5 cm long, 0.4-0.8 times as long as the blade; leaf blades 2.2-6.5 cm long, 1.8-8 cm wide, occa- sionally larger 0.8-1.2 (rarely 1.4) times as long as wide, very often as wide or wider than long, rhombic ovate, or orbicular, usual- ly strongly accuminate at the apex, cuneate at the base, glabrous, equally green on both sides or a little darker above, the margin crenate-serrate, not ciliate; bracts of aments laciniate; staminate aments 4-6 cm long; sta- mens 20-30; pistillate aments not seen. In- troduced, cultivated for shade and wind breaks, specimens seen from Beaver, Salt Lake, and Utah counties, but to be expected throughout the state. Most of the trees in our area are from a staminate clone with strongly ascending branches that produced a narrow, often nearly cylindrical crown. Trees of this clone have been assigned to var. italica Duroi, Lombardy poplar; 6 (0). . Populus tremuloides Michx. Aspen, quak- ing aspen, quakey. Colonial tree 10-15 (20) m tall, seldom taller; the trunk seldom over 40 cm in diameter; bark white and smooth, covered with a powdery white bloom, turning black and rough where scarred and at the base of very old trunks; branches usually spreading, the crown usually rounded; bud scales shiny but hardly re- sinous; petioles laterally flattened, 2-5.5 cm long (1/2) 3/4 to nearly as long as the blade; leaf blades 2-6.5 cm long, 1.8-6.5 cm wide, or much larger on stump sprouts, 3/4-1 1/3 times longer than wide, ovate to reniform- cordate, the margin subentire to serrate or undulate, ciliate, glabrous on the surfaces at maturity; bracts of the aments more or less persistent, especially the staminate ones, 3-7 lobed or cleft, silky-pilose ciliate, the hairs up to 2 mm long; staminate aments 2-4 cm long, readily deciduous; stamens 6-14; pistillate aments 4-12 cm long, to 13 mm wide; cap- sules 4-6 mm long, the stipes 1-2 mm long, subtended by a cuplike disk about 2 mm across; carpels 2; stigmas 2, each deeply cleft into 2 or more slender lobes. Along water courses and forming clones and aggregates of clones on canyon walls and mountain sides, from (1400) 1830-3050 (3200) m, in all coun- ties of the state; widespread in North Ameri- ca from Labrador to Alaska and south to Tennessee and northern Mexico. Aspen is cul- tivated as a shade or ornamental tree. In re- cent years, nursery stock has become readily available from commercial nurseries; 96 (i). 536 Great Basin Naturalist Vol. 43, No. 4 Salix L. Depressed, mat-forming dwarf shrubs to large trees; buds covered with one non- resinous scale; aments erect to spreading, rarely drooping, developing before (pre- cocious), with (coetaneous) or after (sero- tinous) the leaves, the bracts mostly entire, occasionally with a slightly toothed apex; flowers with 1, occasionally 2 minute glands near the base; stamens (1) 2-8 (12), the fila- ments free or imited toward the base, in- serted on the base of the bract; capsules ses- sile or stipitate, glabrous or pubescent. A large genus of about 300 species, mostly of the Northern Hemisphere, most common in arctic and temperate regions. Identification of the willows is com- pounded by unisexual plants, aments that are sometimes precocious and mostly early de- ciduous, and variation among the usually smaller leaves of the flowering branches which often lack or have inconspicuous stip- ules and the usually much larger leaves and stipules of vegetative branches and particu- larly of vigorous young shoots. Thus, her- barium specimens of each species present specimens of 3 or 4 phases (pistillate, stami- nate, flowering twigs with or without the de- ciduous aments, and vegetative twigs). Vigor- ous young shoots sometimes add a fifth dimension. At times whole plants in the field present only one or two of the various phases. To facilitate identification of plants of the different phases, pistillate, staminate, and vegetative features have been included in many of the leads in the key. Thus, some of the leads are rather long, and features not ap- plicable to a particular specimen will need to be skipped. An alternative approach to lengthy leads is separate keys for the differ- ent sexual and vegetative phases. Many such keys have been written, but these sometimes also contain a mixing of vegetative and sexual features. To establish an adequate basis for a staminate key, I feel that many more stami- nate specimens are needed in the herbaria of the state. References Archer, W. A., and E. E. Little. 1965. "Salicaceae of Nevada." Contr. Flora Nevada 50: 10-59. Argus, G. W. 1965. The taxonomy of the Salix glauca complex in North America. Contr. Gray Herb. 196: 1-142. 197.3. The genus Salix in Alaska and the Yukon. Nat. Mus. Canad. Publ. Bot. 2: 1-279. 1980. The typification and identity of S. erioce- phala Michx. (Salicaceae). Brittonia 32(2): 170-177. Arnow, L., B. Albee, and A. Wycoff. 1980. Flora of the central Wasatch Front, Utah. Univ. of Utah Printing Service, Salt Lake City. 663 pp. DoRN, R. D. 1975. A systematic study of Salix section Cordatac in North America. Can. J. Bot. 53: 1491-1522. 1976. A svnopsis of American Salix. Can. J. Bot. 54:2769-2789. 1977. Willows of the Rocky .Mountain States. Rhodora 79:390-429. Hitchcock, C. L., and A. CRONguisT. 1964. Vascular plants of the Pacific Northwest. Part 2: Sali- caceae to Saxifragaceae. Univ. Washington Puhl. Biol. Vol. 17. ,597 pp. Rehder, a. 1951. Manual of cultivated trees and shrubs hardv in North America. Macmillan Co., New York. 996 pp. 1. Plants shrubs or dwarf shrubs not over 1 (1.5) m tall, subalpine to alpine 2 — Plants shrubs or trees, mostly over 1.5 m tall, of valleys to montane 3 2(1). Plants depressed dwarf shrubs 1-10 (20) cm tall mostly alpine, often forming mats, the stems creeping on or below the ground surface KEY I — Plants (10) 20-100 cm tall or taller, subalpine or alpine, not forming mats on the ground, the stems ascending to erect KEY II 3(1). Leaves (8) 10-20 (32) times longer than wide; plants often strongly colonial, spreading underground and forming patches and occasionally thickets, our most common and widespread lowland willow S. exigua — Leaves less than 8 times as long as wide 4 4(3). Bracts persistent, dark brown to blackish or if pale green or pale brown in age then silky pilose with the hairs exceeding the bract by 1-2 mm and the cap- sules pubescent (rarely glabrous in unusual specimens); stamens 2 per flower, October 1983 Goodrich: Utah Flora, Salicaceae 537 the filaments glabrous or pilose in a few species; plants shrubs or occasionally treelike, mostly native KEY III — Bracts of at least the pistillate aments quickly deciduous, pale green or yellow- ish tan in age, short pubescent, the hairs hardly if at all exceeding the bract by more than 1 mm; capsules glabrous; stamens more than 2 per flower, or if only 2 then plants introduced trees, the filaments pilose; plants mostly trees or treehke except in S. lasiandra, mostly of valleys and lower montane 5 5(4). Plants native; stamens 3-9 per flower; stipes of capsules mostly 1-2 mm long, obviously longer than the gland KEY IV — Plants introduced trees; stamens 2 except in S. pentandra; capsules sessile or the stipes mostly less than 1 mm long and hardly longer than the gland KEY V KEY I. Depressed, mat-forming dwarf shrubs, 1-10 (20) cm tall, at or above timberline 1. Bracts of aments pale green or yellowish, glabrous dorsally; filaments 1.5-2 mm long; style obsolete or to 0.2 mm long, shorter than the stigma; leaves elliptic to orbicular, 1.4-2.6 times longer than wide, glaucous and strongly reticulate-veined beneath, the tips mostly rounded or obtuse S. reticulata — Bracts of aments blackish, pilose dorsally; filaments over 2 mm long; styles 0.5 mm long or longer, longer than the stigmas; leaves elliptic or narrow ellip- tic, (1.25)2.3-4.7 times longer than wide, glaucous or not, not strongly reticulate veined beneath, the tips mostly pointed 2 2(1). Leaves 2-5 (7) mm wide, 2-4.7 times longer than wide, sessile or the petiole to 3 mm long; plants seldom over 3 cm tall, aments 0.5-2.2 cm long S. cascadensis — Leaves 5-20 mm wide, mostly 2-3 times longer than wide, with petiole 3-13 mm long; plants mostly 5-10 (20) cm tall; aments (1) 2-4 cm long S. arctica KEY II. Low shrubs (10) 20-100 (300) cm tall, mat forming, subalpine or alpine 1. Capsules glabrous, the style and stigma together less than 1 mm long; leaves permanently pubescent on both sides, the lower surface not glaucous but often more densely pubescent and thus lighter than the upper surface; twigs of the season glabrous or thinly villous-puberulent S. wolfii — Capsules pubescent at least until mature or style and stigma together over 1 mm long; leaves often glaucous beneath, glabrous or pubescent 2 2(1). Mature leaves glabrous, dark green and shiny above, strongly glaucous and glabrous or with a few hairs beneath; twigs of the season glabrous or very scat- tered pubescent, dark chestnut to lustrous purplish black; aments precocious or coetaneous, sessile or nearly so or rarely on a stalk to 0.5(1) cm long, this nei- ther bearing nor subtended by bractlike leaves; style and stigmas collectively 1.5 mm long or longer; filaments of stamens glabrous S. planifolia — Mature leaves pubescent on both sides, but sometimes glabrate or glabrous in age; twigs of the current season densely pubescent; aments coetaneous or sub- serotinous, born on stalks to 2 (4) cm long, these usually bearing and subtended by bractlike leaves; style and stigmas collectively up to 1.5 mm long; filaments of stamens sometimes pilose 3 538 Great Basin Naturalist Vol. 43, No. 4 3(2). Bracts of aments pale green when young, tan in age; capsules 3-5 mm long, pubescent even in age, crowded and nearly sessile so as to mostly conceal the rachis at the center of the aments, the stipes seldom over 0.5 mm long; pistil- late aments 0.8-2 (2.5) cm long, 8-10 mm wide; staminate aments about 0.8-1 (1.2) cm long, 5-6 mm wide, the filaments densely pilose at the base and for 1/2 to 3/4 their length, the pilose portion often equaling or exceeding the scale, the anthers usually less than 0.5 mm long; petioles 1-4 mm long, seldom exceeding the bud even on vegetative twigs S. brachycarpa — Bracts of aments brown to blackish, sometimes light brown to whitish tan but not green even when young; capsules (4) 5-7 (8) mm long, sometimes glabrate in age, dense but often not so crowded as to conceal the rachis at the center of the ament, the stipes 0.5-2 mm long; pistillate aments (1.8) 2.5-5 cm long, 11-15 mm wide; staminate aments 0.8-2(4) cm long, sometimes over 6 mm wide, the filaments glabrous or pilose but usually not so conspicuously pilose as above, the anthers mostly over 0.5 mm long; petioles (1) 2-6 (10) mm long, equaling or often exceeding the bud, especially on vegetative twigs S. glauca KEY III. Mostly native shrubs or small trees; aments mostly with dark bracts; stamens 2; capsules glabrous or pubescent 1. Capsules glabrous; leaves not both glaucous and pubescent on the lower sur- face when fully expanded; hairs of aments mostly crisped-villous and more or less tangled except in S. wolfii with aments 0.8-2(3) cm long or in S. planifolia and then plants keyed both ways 2 — Capsules mostly pubescent except in S. lasiolepis; leaves glaucous and pu- bescent on the lower surface when fully expanded; hairs of aments straight or slightly wavy but hardly crisped-villous or tangled; aments sometimes longer than in S. wolfii 6 2(1). Leaves glaucous beneath, not or scarcely pubescent when fully expanded 3 — Leaves not glaucous beneath, although sometimes lighter colored from pu- bescence; pubescent at least in part on both sides when fully expanded, but sometimes glabrate in age 5 3(2). Aments sessile or on a stalk, the stalk to 0.5(1) cm long neither bearing nor sub- tended by bractlike leaves; pubescence of aments straight or nearly so; leaves mostly entire, often slightly revolute; twigs dark chestnut to lustrous purplish black, essentially glabrous; plants often less than 1.5 m tall and keyed also in Key II S. planifolia — Aments usually stalked, the stalk usually subtended by or bearing 1-4 bractlike leaves; pubescence of aments crisped-villous; leaves serrate, serrulate, or en- tire, not at all revolute; twigs variously colored, glabrous or those of the current season more often pubescent; plants often over 1.5 m tall 4 4(3). Styles 0.7-1.5(1.8) mm long; leaves of fertile and vegetative twigs often less than 3 times longer than wide, evidently crenulate-serrate or subentire; bark of older twigs not ashy gray or whitish; plants apparently uncommon, in the eastern and central part of the state, mostly montane S. monticola — Styles 0.2-0.7 mm long; leaves of vegetative twigs 2-5 times longer than wide, serrulate or entire; bark of older twigs usually ashy gray or white; plants widespread, mostly of valleys and lower montane S. lutea October 1983 Goodrich: Utah Flora, Salic ace ae 539 5(2). Aments precocious or coetaneous (1.5) 2-5 cm long, with dense crisped-villous, tangled hairs; leaves subglabrate in age, with inconspicuous hairs, entire or sometimes serrulate; plants sometimes over 2 m tall S. boothii — Aments coetaneous, 0.8-1.5 (3) cm long, with hairs straight or nearly so; leaves permanently pubescent throughout on both sides even in age, the hairs readily conspicuous with a 10- power lens, entire; plants 0.6-1.5(2)m tall, also keyed in Key II S. wolfii 6(1). Twigs strongly blue glaucous, the bloom sometimes deciduous, but then the twigs glabrous or sometimes puberulent; larger leaves mostly 3-5 times longer than wide, sericeous beneath; capsules densely pubescent 7 — Twigs not glaucous or those of the current season often pubescent, or leaves not sericeous; the larger leaves various but sometimes wider than above; capsules pubescent or glabrous 8 7(6). Pistillate aments 2-5 cm long; capsules sessile or the stipes to 1 mm long, the style and stigmas together 0.8-1.3 mm long; staminate aments about 2 cm long, the filaments glabrous; aments sessile or nearly so with or more often without subtending bractlike leaves, precocious or subcoetaneous; bracts of the aments blackish; leaves permanently silvery, silky-sericeous to subtomentose beneath, dark green and glabrous above in age S. drummondiana — Pistillate aments 1-2 cm long; capsules stipitate, the stipes 2-3 mm long, the style and stigmas together about 0.5 mm long; staminate aments 8-15 mm long, the filaments pilose on the lower 1/2; aments borne on 2-10 mm long, bracteate-leafy stalks; coetaneous or subprecocious; bracts of the aments dark at the tip and pale below; leaves sericeous when unfolding, sparsely or moder- ately sericeous, especially beneath when fully expanded, glabrate in age especially above S. geyeriana 8(6). Plants shrubs 0.6-3 m tall, midmontane to above timberline, the stems less than 4 cm thick; leaves mostly less than 2 cm wide, occasionally wider on vegetative twigs, elliptic to narrowly lanceolate KEY II — Plants shrubs or small trees, commonly 3-4 m tall or taller, but sometimes shorter, of valleys or montane, the stems of mature plants often 4-10 cm thick or thicker; leaves sometimes over 2 cm wide, oblong, obovate, oblanceolate, or elliptic 9 9(8). Capsules glabrous; filaments about 3-5 mm long, bracts of aments blackish or purplish black, about as wide as long and rounded at the apex, densely pilose- tomentose, the hairs exceeding the bracts by about 1 mm; leaves oblong to ob- lanceolate, less than 15 mm wide except on vigorous young shoots; plants of Great Basin and Virgin River drainages S. lasiolepis — Capsules pubescent; filaments longer or bracts not as dark; bracts of aments of lighter color or if blackish then with hairs exceeding the bracts by about 2 mm, pointed or somewhat rounded; leaves elliptic, obovate, to oblanceolate, sometimes over 15 mm wide; plants of various distribution 10 10(9). Twigs of the second and current year and the dark red bud scales velvety vil- lous; lower surface of leaves densely velvety villous throughout the season, twigs with longitudinal ridges beneath the bark; aments precocious; plants introduced, cultivated S. cinerea L. — Twigs of the second year glabrous, those of the current season villous or with appressed hairs; lower surface of leaves villous at first but usually rather scattered-villous to glabrate in age; aments various; plants native 11 540 Great Basin Naturalist Vol. 43, No. 4 11(10). Bracts of aments pale green or tan to very light brown in age, silky pilose, the hairs exceeding the bract by about 1 mm; aments coetaneous; capsules long beaked, loosely arranged so as to expose much of the rachis; filaments of sta- mens 3-6 mm long; leaves mostly elliptic, occasionally lanceolate or obovate; twigs of the season with mostly appressed or ascending hairs or occasionally glabrous S. hehbiana — Bracts of aments black or purplish black, reddish or pale only at the very base, pilose, the hairs exceeding the bract by about 2 mm; pistillate aments pre- cocious, or subprecocious, the capsules not long beaked, densely arranged and mostly concealing the rachis; staminate aments strictly precocious, the fila- ments about 10 mm long; leaves obovate or oblanceolate; twigs of the season with mostly widely spreading hairs S. scouleriana KEY IV. Plants native, tall shnibs or small trees; bracts of aments pale green or yellow, at least the pis- tillate ones, deciduous; stamens 3-8(12); capsules glabrous with a 1-2 mm long stipe 1. Bracts of aments 3-4 mm long; bud scales fused, without free overlapping mar- gins, blunt to rounded at the tip; staminate aments 2-3.5 times longer than wide, 1-3.5 cm long; styles 0.5-1 mm long; petioles of larger leaves often with wartlike glands near the base of the blade; plants mostly multistemmed, large shrubs from large root crowns, rarely trees, widespread in the northern half of the state S. lasiandra — Bracts of aments 1-2 mm long; bud scales with free overlapping margins, usu- ally pointed at the tip; staminate aments often narrower and longer than above; styles 0.1-0.2 mm long; petioles without wartlike glands or occasionally with glands in S. nigra; plants mostly trees with solitary or few trunks, of various distribution 2 2(1). Leaf blades not glaucous beneath, (2.5)4-7 times longer than wide; twigs whitish or grayish yellow; plants of the southern half of the state S. nigra — Leaf blades glaucous beneath 3 3(2). Twigs reddish or reddish brown, often pubescent at least near the nodes, hori- zontal or spreading; some of the leaf blades usually 4-5 times longer than wide, shiny dark green above; plants of San Juan and Washington counties S. laevigata — Twigs ashy gray or yellowish when fresh, glabrous, tending to droop; leaf blades usually not over 3 times longer than wide, not shiny dark green above; plants widespread in the state, mostly north of the counties listed above S. amygdalioides KEYV. Plants small or rather large trees, introduced, cultivated, sometimes escaping and persisting; bracts of aments pale green or yellowish, at least the pistillate ones deciduous; capsules glabrous, sessile or nearly so (Note: The cultivated species of this key, except S. fragilis, are not described due to lack of adequate specimens in herbaria.) 1. Stamens 3-12; leaves with wartlike glands on upper part of petiole and lower margins of blade, the blade seldom over 3 times longer than wide, usually glabrous except above along the midrib, lighter beneath than above but not glaucous (bay willow) S. pentandra L. October 1983 Goodrich: Utah Flora, Salicaceae 541 — Stamens 2; leaf blades 3-5 times longer than wide, usually glaucous beneath, glabrous or variously pubescent 2 2(1). Pistillate aments 1-2.5 (3) cm long, the capsules 1-2.5 mm long; staminate aments to 4 cm long; petioles glandless; trees weeping, with very slender, greatly elongate, pendulous branches, or if not weeping then the twigs more or less contorted 3 — Pistillate aments mostly over 3 cm long, the capsules 3-6 mm long; petioles sometimes with small glands near the base of the blade; trees not weeping, with upright branches; twigs spreading, not contorted 5 3(2). Trees not weeping; twigs not pendulous, more or less contorted, aments 1-1.5 cm long; (all specimens seen from the state were referrable to f. tortuosa Rehd. with the branches twisted and contorted — corkscrew willow) S. matsudana Koidz. — Trees weeping; twigs pendulous, very straight; aments sometimes longer than above 4 4(3). Leaves mostly 3-15 mm wide, mostly deciduous in October; twigs often bright yellow; capsules sessile (weeping willow) S. babylonica L. — Leaves 15-22 mm wide, often persisting into December; twigs greenish or yel- low-green; capsules with stipe exceeding the gland; plants hybrids of S. babylonica x S. fragilis (Niobe or Wisconsin weeping willow) .... S. x blanda Anderss. 5(2). Leaves glabrous when unfolded, the margin of mature leaves usually serrate with 4-8 teeth per cm; twigs glabrous, or nearly so; stipe of capsules 0.5-0.8(1) mm long; plants common, cultivated, and escaping S. fragilis — Leaves sericeous, or glabrous when unfolded, the margin of mature leaves fine- ly serrulate with 9-10 teeth per cm; twigs sometimes pubescent; capsules sessile or subsessile; plants not known outside of cultivation (white willow) S. alba L. Salix amygdaloides Anderss. Peach-leaf half; pistillate aments (1.5) 2.5-8 cm long, willow. Plants mostly small trees, rarely 13-20 mm wide; capsules 4-7 mm long, shrublike, mostly 4-10 (12) m tall, often with glabrous, the stipe 1.2-3 mm long, the style 2-4 leaning tnmks; twigs whitish, yellowish, about 0.2 mm long, not longer than the or ashy gray, rarely reddish, glabrous except stigmas. Lake and pond margins and along when very young; stipules usually minute and ditches, streams, and rivers, and in neglected soon deciduous; petioles (3) 5-15 (25) mm fields and pastures, from about 1070-1710 m long; leaf blades, (1.8) 2.3-6 (7.5) cm long, (7) in Box Elder, Davis, Duchesne, Emery, Juab, 12-19 (23) mm wide, or up to 10.5 cm and Salt Lake, Tooele, Uintah, Utah, and Wash- 3.2 cm wide on vigorous young shoots, ellip- ington counties; southern Canada and wide- tical to lanceolate, entire or serrulate, spread in the United States except the south- glabrous except when very young, glaucous ern part; 63 (vi). beneath, green above; aments coetaneous, Salix arctica Pall. Arctic willow. [S. anglo- rarely subprecocious, on leafy or bracteate rum Cham. var. antiplasta Schneid.]. De- twigs of the season, 1.5-4 cm long; bracts of pressed shrubs with stems creeping on or un- the aments 1-2 mm long, at least the pistil- der the ground, seldom rising more than 10 late ones soon deciduous, pale green, orbicu- (20) cm above ground level, tending to form lar, the dorsal side woolly-pilose below and mats, but not so much as in S. cascadensis or along the margins, but mostly glabrous to- S. reticulata; stipules minute or lacking; pet- ward the apex, the ventral surface woolly-vil- ioles 2-12 mm long; leaf blades (5) 11-47 mm lous throughout, the hairs seldom exceeding long, (4) 6-16 mm wide, elliptical, narrow el- the bract by more than 0.5 mm; staminate liptical, obovate, or oblanceolate, entire, aments 2-10 cm long, 7-11 mm wide; sta- slightly paler beneath than above but not mens 4-7, the filaments pilose on the lower strongly glaucous; pilose-sericeous when 542 Great Basin Naturalist Vol. 43, No. 4 young, sparingly pubescent or glabrous when mature; aments coetaneous, borne on glabrous or pubescent 7-35 mm long leafy- bracteate or barren twigs of the season; bracts of the aments persistent, dark brown, pinkish purple at the base, pilose-sericeus on both sides, sometimes less so dorsally than ventrally, the hairs exceeding the bract by about 1 mm; staminate aments 15-25 mm long, about 7-9 mm wide; stamens 2, the fila- ments glabrous, to about 7 mm long; pistil- late aments 1.5-7 cm long, about 10-12 mm wide, with 25-75 fruits; capsules 4-7 mm long, pubescent, the stipe about 1 mm long, the style and stigmas together about 1-2 mm long. About snowbanks, meadows, shores of lakes, and rocky slopes near or a little above timberline, 2775-3600 m on the Bear River (Mt. Naomi), Tushar (Delano Peak), western Uinta, and the Wasatch mountains in Cache, Piute, Salt Lake, Summit, and Utah counties; circumboreal and south in mountains of west- ern North America to California and New Mexico. Our plants are var. petraea Anderss. They more or less intergrade into S. casca- densis in the Uinta Mountains; 14 (0). Salix bebbiana Sarg. Bebb willow. Plants shrubs, occasionally treelike, (2) 4-6 (8) m tall, with 1 to several stems, young twigs glabrous, puberulent or densely pubescent; stipules usually inconspicuous and soon de- ciduous; petioles (2) 3-8 (10) mm long, red- dish or pale; leaf blades 1-4 cm long, 1.2-2 cm wide or to 7 cm long and 3 cm wide on vigorous young shoots, 2.2-2.8 times longer than wide, mostly elliptical, occasion- ally obovate or oblanceolate, entire to slightly undulate-crenate, dark green above, glaucous beneath, pubescent when young on both sides; fully expanded leaves glabrous above, usually with a few hairs beneath near the midrib; aments coetaneous, on a brac- teate 3-15 mm long peduncle; bracts of the aments persistent, pale green to very light brown in age, sometimes reddish at the apex, particularly in staminate aments, silky pu- bescent, the hairs exceeding the bract by about 1 mm; staminate aments 1.5-2 cm long, to 13 mm wide; stamens 2, the fila- ments 3-6 mm long, glabrous or sparingly pi- lose at tlie base; pistillate aments 1.5-4(5) cm long, to 2 cm wide; capsules 6-8(10) mm long, rostrate with a rounded basal portion 1-2 mm wide and a long slender beak, pu- bescent, rather loosely arranged and not con- cealing the rachis, the stipe 2-3.5 mm long, the style about 0.1-0.2 mm long; stigmas 0.3-0.5 mm long, bilobed to the base. Ripa- rian communities on canyon bottoms and along streams in mountains, occasionally along irrigation ditches, from (1370) 1830-2710 m in Box Elder, Cache, Daggett, Davis, Garfield, Grand, Juab, Kane, Rich, Salt Lake, San Juan, Sevier, Summit, Uintah, Utah, Wasatch, Washington, and Wayne counties; across much of Canada and north- ern United States. Our plants with leaves sparsely appressed pubescent and soon glabrous beneath and rather weakly raised reticulate-veiny are often referred to as var. perrostrato (Rydb.) Schneid., but the separa- tion probably merits no recognition. 77(x). Salix boothii Dorn Booths willow. [S. pseudocordata (Anderss.) Rydb., misapplied]. Shrubs (1.5) 2-4 m tall; young twigs finely hairy, stipules small, inconspicuous and soon deciduous or larger and leaflike on vigorous young shoots; petioles mostly 2-5 mm long; leaf blades (0.8) 2.5-6 cm long, (4) 8-22 mm wide, or to 11.2 cm long and 4 cm wide, with petiole to 2 cm long on vigorous shoots, ellip- tical, lanceolate, occasionally nearly linear, rarely oval, entire or serrulate, not glaucous beneath, sparingly to moderately pubescent at least in part on both sides, or glabrate to- ward the end of the season, about as pu- bescent at the apex as at the base, coriaceous in age; aments subprecocious or coetaneous, sessile or on a barren or 1-3 bracteate pe- duncle to 8 mm long; bracts of the aments persistent, dark brown to purplish black at the apex, often with a lighter base; pu- bescence of aments sericeus-pilose at first but soon becoming crisped-villous and somewhat entangled, the hairs usually exceeding the bracts by 1-2 mm, sometimes deciduous; staminate aments 1-2.5 cm long; stamens 2, the filaments about 5 mm long, glabrous; pis- tillate aments (1) 2-4 (6) cm long; capsules 3-6 mm long, glabrous, the stipe 1.5-2 mm long; styles 0.3-1 (1.5) mm long. Riparian and wet meadow communities from about 2075-3050 m, particularly common on the plateaus of central Utah, but from all coun- ties of the state except Millard, Morgan, October 1983 Goodrich: Utah Flora, Salicaceae 543 Rich, Tooele, Washington, Wayne, and We- ber and to be expected in some of these; Col- orado Rockies west to northern California and north to southern Alberta and British Co- lumbia. Our plants are closely related to S. myrtillifolia Anderss. of Alaska and Canada. They vary from those of Alaska and Canada by either taller stature or pubescent leaves or both, and they have longer stipules that are more sharply acute at the apex. They might be treated as a variety of S. myrtillifolia, but no new combination is proposed here. Some- times referred to as S. pseudocordata, but this name is synonymous with S. myrtillifolia (Dom 1975). Occasionally grading toward S. wolfii in pubescence of leaves and sometimes difficult to distinquish from that species veg- etatively. Like S. lutea in color and pu- bescence of scales and rachis of aments, and sometimes confused with that species, but with leaves coriaceous in age and more and persistently pubescent and not glaucous be- neath, and generally of higher elevations, but sometimes nearly impossible to distinguish from S. lutea in leafless or very young-leaved specimens with precocious aments. However, older twigs of S. boothii are not whitish as they often are in S. lutea, and specimens with older twigs are more easily distinguished; 139 (xh). Salix brachycarpa Nutt. Barrenground wil- low. Short-fruited willow. Shrubs (0.25) 0.6-1.5 m tall, rarely taller; twigs below the leaves with epidermis breaking in translucent flakes, twigs of the season dark or reddish un- der the dense pubescence; stipules inconspic- uous, deciduous; petioles 1-4 mm long, usual- ly not longer than the bud, often reddish, the reddish color sometimes extending up the midrib of the blade; leaf blades (0.6) 1.5-4 cm long, (3) 5-18 mm wide, or to 7 cm long and 3 cm wide on sterile branches, 2-4 (5) times longer than wide, elliptical, broadly lanceolate, occasionally nearly linear, entire, thinly to moderately sericeous to nearly glabrous on both sides, strongly glaucous be- neath; aments coetaneous or serotinous, nearly sessile or more often on bracteate pe- dimcles at the ends of leafy twigs; bracts of the aments pale green, tan, or light brown in age, rarely pink or pale reddish at apex, scat- tered to densely pilose on both sides, the hairs exceeding the bract by about 1 mm or less; staminate aments (6) 8-10 (12) mm long, 5-6 mm wide; stamens 2, the filaments 2.5-5 mm long, densely pilose at base and scattered pilose to 1/3 to 3/4 the entire length, the pubescent portion sometimes ex- ceeding the scale, anthers 0.3-0.5 (0.6) mm long, orbicular, yellowish; pistillate aments 8-25 mm long, 3-10 mm wide; capsules 3-5 mm long, densely arranged and mostly concealing the rachis, sessile or on stipes up to 0.5 (1) mm long, sometimes persisting over winter, pubescent, the hairs persistent even on over-wintering capsules, the style 0.5-1 mm long, the stigmas about 0.5 mm long, bilobed to the base. Along streams, in wet meadows, dry rocky and talus slopes, and rocky, open ground in mountains from 2070-3230 m, mostly on ground with basic substrate in Cache, Duchesne, Emery, Grand, Iron, Juab, Kane, Salt Lake, Sanpete, Sevier, Summit, Utah, and Wasatch counties; wide- spread in Alaska, Canada, and south in west- ern United States from Oregon south and east to Colorado. Our plants are assignable to var. brachycarpa with bracts greenish at anthesis and subspherical or short cylindrical, densely flowered pistillate aments, leaves coarsely pubescent on both sides and with com- paratively tall stature. Closely related to and often confused with S. glauca, but distinct in the state by small but numerous features. In addition to the features given in the key, S. brachycarpa more or less differs from S. glauca in having twigs with more numerous aments, distal leaves of fertile twigs often considerably larger than the 3 or 4 proximal ones, and reddish as well as yellowish petioles with the reddish color sometimes extending up the midrib of the leaf blade. Although most of our plants seem quite distinct, appar- ently there is widespread introgression with S. glauca in the Rocky Mountain Region and particularly southward in Colorado (Argus 1965); see discussion under S. glauca; 74 (xviii). Salix cascadensis Cockerell. Cascades wil- low. Depressed, mat-forming subshrubs, 1-3 cm tall, from tap root and rhizomatously much-branched caudex; petiole lacking or to 3 mm long; leaf blades 6-18 mm long, 1.5-4 mm wide, 2-4.7 times longer than wide, linear or narrow elliptical, entire, pilose-sericeous when young, soon glabrous 544 Great Basin Naturalist Vol. 43, No. 4 and green on both sides or slightly paler be- low, some marcescent for 1 or more years; aments coetaneous, terminal on short leafy lateral branches, these about 8-22 mm long; bracts of the aments persistent, black or purplish black, reddish-purplish at the very base, about 1-2 mm long, 1 mm wide, pilose on both sides, but less so to nearly glabrous at the base ventrally, the hairs about 1 mm long; staminate aments 3-12 mm long, 5-8 mm wide; stamens 2, separate to the base, the filaments about 3-4 mm long, glabrous, the anthers reddish or purplish; pis- tillate aments 5-22 mm long, 5-11 mm wide; capsules 3-4 mm long, pubescent, sessile or the stipe less than 1 mm long, the style and stigmas together about 1.5 mm long. Alpine tundra on the Uinta Mountains, 3350-3932 m, in Daggett, Duchesne, Sum- mit, and Uintah counties; southwestern Brit- ish Columbia, east to Montana south through Wyoming to Colorado and Utah; 15 (iv). Salix drummondiana Barratt in Hook. Drummond willow. [S. subcoerulea Piper]. Shmbs (1) 2-3(4) m tall; twigs glabrous or puberulent when very young, heavily glaucous, the bloom persisting into the sec- ond year, yellow-brown to blackish purple beneath the bloom; stipules narrow, small and deciduous, or larger and more persistent on vigorous young shoots; petioles 4-12 mm long; leaf blades 2.2-8 cm long, (5) 13-20 mm wide, or to 14 cm long and 3 cm wide on vigorous young shoots, lanceolate or narrowly elliptical, rarely oblanceolate, en- tire, sometimes with slightly revolute mar- gins, dark green and glabrous or thinly pu- bescent above, densely silvery white pubescence beneath with short appressed or spreading and slighty tangled hairs, pale glaucous beneath the pubescence; aments precocious or sub-coetaneous; bracts of the aments persistent, purplish black or purplish brown, pilose on both sides, the longest hairs exceeding the bract by 1.5-2 mm; staminate aments 19-22 mm long, 3-10 mm wide, ses- sile or on a peduncle to 3 mm long; stamens 2, the filaments, 4-9 mm long, glabrous; pis- tillate aments 2-4.5 cm long, 3-12 mm wide; capsules 3-6 mm long, pubescent, sessile or the stipe to 1 mm long, the style 0.5-0.7 mm long, the stigmas 0.3-0.6 mm long. Along streams and rivers, wet meadows, and other wet places from 2135-3140 (3290) m in Bea- ver, Box Elder, Cache, Daggett, Davis, Du- chesne, Emery, Grand, Piute, Salt Lake, San- pete, Sevier, Summit, Uintah, Utah, and Wasatch counties; British Columbia and Al- berta south to California and New Mexico; 84 (xxv). Salix exigua Nutt. Coyote willow. Dusky willow. Narrow-leaf willow. Colonial shrub (1) 2-3 m tall or rarely treelike and to 8 m tall; stems ashy gray, branches often reddish, twigs of the season greenish, pubescent; leaves (1) 2-11 cm long, (0.1) 0.2-1 cm wide, sessile or with 1-3 mm long petiole, or to 17.5 cm long and 1.6 cm wide with petiole up to 12 mm long on vigorous yoving shoots, linear, entire or serrulate-dentate with glandular teeth, glabrate to densely white se- riceous; aments coetaneous or serotinous on slender leafy peduncles or twigs of the sea- son, these 0.5-14 cm long; bracts of the aments about 2 mm long, about 1 mm wide, pale green or yellowish, deciduous, pu- bescent on both sides but often glabrate or glabrous dorsally especially toward the apex, occasionally only ciliate ventrally; staminate aments 1.5-4.5 cm long, 0.5-1 cm wide; sta- mens 2, the filaments pilose on the lower half; pistillate aments 1.5-6 cm long, 8-16 mm wide; capsules 4-7 mm long, most- ly glabrous, sometimes pubescent, sessile or the stipe up to 0.8 mm long, the style obso- lete. Along rivers and streams, irrigation dit- ches, washes, in neglected fields and pastures, around ponds and reservoirs, tolerant of alka- line soils, from 825-2315 (2590) m in all counties of the state. Our plants are part of a complex that extends from the Atlantic to the Pacific in the northern United States and southern Canada and extends from Alaska to northern Mexico in the western part of North America. Most of our specimens of ssp. exigua have glabrous capsules and can be as- signed to var. stenophyUa (Rydb.) Schneid. Some specimens from the northern part of the state have somewhat pubescent capsules and these may be var. exigua. The closely re- lated S. melanopsis Nutt. has been reported for the area, but I have not seen any speci- men that clearlv belongs to that taxon. 170 (ii). Salix fragilis L. Crack willow. Large trees to 20 m tall, the trunks to 1.3 m in diameter, solitary or few, erect or strongly leaning, October 1983 Goodrich: Utah Flora, Salicaceae 545 with thick fiirrowed gray or blackish gray bark; branches ascending, often large; twigs spreading, not pendulous, very brittle and easily broken at the base; leaf blades lanceo- late to narrow elliptic, (2.5)3-17 cm long, (7)10-32 mm wide, acute or accuminate, ser- rate, glaucous or glaucescent beneath, glabrous or sericeous when young, glabrous when mature; aments coetaneous, on twigs of the current season, these twigs about 1-2.5 cm long, with (1)2-3(4) reduced leaves, the leaves like the larger leaves of the non- floriferous twigs but sometimes oblanceolate; bracts of the aments pale green, pale yellow- green, or greenish white, tan or very pale brown upon drying, the pistillate ones de- ciduoiLS by the time the capsules mature, se- riceous with the hairs exceeding the bract by about 1-1.5 mm; staminate aments 3.5-7(9) cm long, 9-12 mm wide; stamens 2; filaments about 3-6 mm long, yellow, pilose toward the base, the pilose portion about equal or shorter than the subtending bract; pistillate aments (2.5)4-7 cm long, 10-13 mm wide; capsules 4-6 mm long, crowded but usually not so dense as to conceal the rachis, glabrous, the stipes about 1 mm long, the styles 0.5-1 mm long, the stigmas 0.2-0.3 mm long. Introduced from Eurasia, cultivated at homes and along streets, persisting, escaping, and naturalized along irrigation and natural waterways and lake margins, solitary to form- ing groves, from 1370-2075 m, in Beaver, Box Elder, Cache, Duchesne, Juab, Rich, Salt Lake, Sanpete, Sevier, Summit, Tooele, Uin- tah, Utah, Wasatch, and probably most other counties of the state. Hybrids of S. fragilis x S. alba (S. x rubens Schrank.) have been de- veloped (Rehder 1951). If such hybrids have been cultivated in the state, they could add considerable complication to the taxonomic separation of the two species; 30 (xiv). Salix geyeriana Anderss. Geyer willow. Shnibs 1.5-4.5 m tall, twigs glabrous or scat- tered puberulent, strongly glaucous, the bloom sometimes deciduous; stipules minute and deciduous; petioles 3-10 mm long; leaves (1) 2-4.5 cm long (4) 8-12 mm wide, ellipti- cal, narrow elliptical to narrow lanceolate, entire or nearly so, glaucus beneath, sericeous when unfolding, sparsely to moderately se- riceous at maturity, especially below, the hairs white or a few pale reddish; aments subprecocious to coetaneous; peduncles of aments leafy or bracteate, the staminate 2-5 mm long, the pistillate 3-10 mm long; bracts of the aments persistent, sericeous-pi- lose on both sides, or glabrate or glabrous ventrally especially in age, the hairs exceed- ing the bract by 0.5-1 mm, the staminate ones light brown when very young, turning reddish to purplish black at the tips, those at the tips of aments turning first, the pistillate ones greenish brown to brown; staminate aments 7-15 mm long, 5-9 mm wide; sta- mens 2, the filaments about 4 mm long, pi- lose to about midlength, the pilose portion about equaling or exceeding the bract; pistil- late aments 1-2 cm long, 6-15 mm wide; capsules 4-7 mm long, pubescent, the stipe (1) 2-3 mm long, the style 0.2-0.3 mm long; stigmas about 0.2 mm long. Along streams and rivers and in other wet places from 2195-2895 m, most common in the Uinta Mountains to Strawberry Valley, occasional elsewhere in the state, in Beaver, Cache, Daggett, Duchesne, Emery, Kane, Rich, Salt Lake, Sevier, Summit, Uintah, Utah, Wasatch, Washington, and Wayne counties; southern British Columbia south to California and east to Montana and Colorado. With glaucous twigs and whitish pubescence, our plants are assignable to var. geyeriana; 67 (xix). Salix glauca L. Glaucous willow, Grayleaf willow. [S. pseudolapponum Seem, in En- gler]. Plants mostly low shrubs (0.1) 0.3-1 (3) m tall; twigs sometimes glaucous but mostly not, those below the leaves with epi- dermis exfoliating in translucent flakes, those of the season reddish under whitish pu- bescence, occasionally glabrate, often with a tuft of pilose hairs at the node; stipules most- ly small and soon deciduous; petioles (1) 2-6 (18) mm long, mostly yellowish or greenish, the color often extending up the midrib of the blade; leaf blades 2-55 mm long, 7-22 mm wide, or to 9 cm long and to 5 cm wide on ends of vegetative twigs, elliptical, pu- bescent when young to glabrate or glabrous in age, mostly entire or rarely serrate; aments coetaneous, nearly sessile on old twigs or more often on bracteate peduncles or leafy, current twigs; bracts of the aments persistent, pale brown to blackish, pilose; staminate aments 1.5-4 cm long; stamens 2, the fila- ments free or united in imusual specimens, glabrous or sparsely pilose at the base, the 546 Great Basin Naturalist Vol. 43, No. 4 anthers 0.5-0.8 mm long; pistillate aments 1.5-5 cm long, 11-15 mm wide; capsules (4) 5-7 (9) mm long, densely pubescent to gla- brate or glabrous in age, crowded but usually not so dense as to conceal the rachis, the stipes 0.5-2 mm long, the style 0.6-1 mm long, the stigmas about 0.5 mm long. Along streams, around springs, on talus slopes, snowflush areas, and dry alpine timdra but then usually in or near krummholz, from 2775-3660 m, on the Bear River, Uinta, and Wasatch mountains and Wasatch Plateau in Cache, Daggett, Duchesne, Salt Lake, San- pete, Summit, Uintah and Wasatch counties; circumboreal, south in western North Ameri- ca in the Rocky Mountains to New Mexico. Highly variable plants with geographic phases. The Uinta Mountain plants from high elevations are more or less comparable to plants that have passed under the name of S. pseiidolapponum, and they generally repre- sent a rather low-statured phase. On wind- swept alpine summits, these plants approach the stature of S. arctica, but the stems are still ascending to erect. These Uinta Mountain plants tend more toward glabrescence in the capsules and have darker scales than is typi- cal of those in the Bear River Range. The twigs are quite persistently pubescent. Leaves are seldom over 5 cm long or over 2 cm wide. Plants from the Bear River and Wasatch ranges have densely and persistently pubescent capsules, pale brown to dark brown to occasionally pinkish tan or rarely whitish tan scales, the twigs are sometimes early glabrate and some of the leaves are fre- quently over 5 cm long and over 2 cm wide. Specimens from Horseshoe Flat area of the Wasatch Plateau have glabrous or pubescent capsules, mostly dark scales, and glabrate and unusual, distinctly serrate leaves. The varia- bility in S. glauca nearly encompasses S. brachycarpa. However, I prefer to follow Argus (1965) and keep the two separate; 57 (xii). Salix laevigata Bebb. Red willow. Shrub or tree 2-15 m tall; twigs reddish brown or dull brown, ashy red or ashy gray during exfolia- tion, stipules inconspicuous or to 6 mm long on vigorous vegetative twigs; usually de- ciduous, petioles stout, 4-14 mm long; leaf blades (1) 1.8-4 (6) cm long, 5-20 cm wide, or to 19 cm long and 4 cm wide on vigorous young shoots, narrowly to broadly lanceolate. glandular-serrulate, somewhat revolute, usu- ally thick and firm, dark green and glabrous above, glaucous, and glabrous or pubescent toward the base and along the midrib; aments subprecocious to coetaneous, on leafy or bracteate twigs of the season; bracts of aments 1-2 mm long, at least the pistillate ones deciduous, pale yellow, crinkly pilose on both sides or often glabrous dorsally, entire or erose to dentate at apex; staminate aments 3-6 cm long, about 1 cm wide; stamens 3-7, pilose on lower half; pistillate aments 4-8 (11) cm long, to 1.5 mm wide; capsules 4-5 (6) mm long, glabrous, the stipes 1.5-2.5 mm long, styles 0.1-0.2 mm long, equaling the bi- lobed stigmas. Along drainages from 701-1370 m, in Washington, San Juan, and probably Kane counties; Arizona, California, Nevada, Utah, and northern Baja California. Perhaps not distinct from S. bonplandiana H.B.K., and treated as synonymous with that taxon by Dorn (1977); 16 (0). Salix lasiandra Benth. Whiplash willow; Caudate willow. Shrub or small tree (2) 3-6 (12) m tall; twigs glabrous or finely hairy when young; stipules often well developed, broadly rounded, gland toothed, 2-10 mm long, eventually deciduous; petioles 3-15 (25) mm long, often bearing 2 or more wart- like glands on the upper side at or near the base of the blade; leaf blades (2.2) 5.5-11.5 cm long, (5) 12-21 mm wide, or to 26 cm long and 5.5 cm wide on vigorous young shoots, lanceolate, elliptical or narrow elliptical, gradually long accuminate, closely serrulate, glabrous except when very young; aments coetaneous, on 1-3.5 cm long leafy- bracteate twigs, the leaves or bracts of the ament-bearing twigs 3-5 in number to 6.5 cm long and 1.2 mm wide, deciduous after the fruit matures; bracts of the aments deciduous (at least the pistillate ones) by the time the capsules start to open, 3-4 mm long, glabrous or nearly on the upper half, pubescent to- ward the base usually more so ventrally than dorsally, entire or minutely toothed at the apex with a few rounded teeth, the staminate yellow, the pistillate pale greenish; staminate aments 1.8-4.5 cm long, 3-12 mm wide; sta- mens 3-8, usually 5, the filaments pilose; pis- tillate aments 2-7 cm long, 11-18 mm wide; capsules 4-8 mm long, glabrous, the stipe 1-2 mm long, the style 0.5-1 mm long, the October 1983 Goodrich: Utah Flora, Salicaceae 547 stigmas to 0.5 mm long. Along streams and rivers, on flood plains, occasionally along irri- gation canals, or around ponds, and reser- voirs, from 1525-2440 (262 l)m in Beaver, Box Elder, Cache, Carbon, Daggett, Davis, Duchesne, Emery, Garfield, Juab, Piute, Rich, Salt Lake, Sanpete, Sevier, Summit, Uintah, Utah, and Wasatch counties; to be expected elseu'here; Alaska and Yukon to California and New Mexico. Our plants are var. caudata (Nutt.) Sudw, with leaves about equally colored on both sides. Var. lasiandra with leaves glaucous beneath has been re- ported for the state, but I have seen no speci- men; 98 (xxi). Salix lasiolepis Benth. Arroyo willow. Shrubs or small trees mostly 4-6 m tall in our range; twigs yellowish olive to reddish, usual- ly soft puberulent when young; stipules mi- nute, soon deciduous or lacking, occasionally well developed on vigorous young shoots; petioles 3-15 mm long; leaf blades 1.5-4.2 cm long, 6-13 mm wide, or to 11 cm long and 2.5 cm wide on vigorous young shoots, usually oblanceolate or oblong, occasionally elliptical, entire, rarely minutely toothed, somewhat revolute margined, dark green and glabrous above, at maturity glaucous be- neath, more or less coriaceous, rather densely soft pubescent on both sides when unfolding, less so above than beneath, few to many of the hairs persisting beneath at maturity; aments precocious to subcoetaneous on 3-6 mm long bracteate or bare peduncles; bracts of the aments persistent purple-black, obovate with broad rounded apex, densely villous, almost hidden in the hairs; staminate aments 2.2-4.5 cm long; stamens 2, the fila- ments glabrous; pistillate aments (1.8) 2.2-4.5 cm long (to 7 cm long outside of our area), 10-12 mm wide; capsules 3-4 (5) mm long, glabrous, the stipe 1-2 cm long, the style about 0.5 mm long, the stigmas 0.2-0.3 mm long. Along streams, ditches, and washes from about 1463-2328 m, in western Utah, Great Basin and Virgin River drainages, in Beaver, Iron, Juab, Millard, Sevier, Tooele, Utah, and Washington counties; southern British Columbia south to Baja California and east to Idaho, Utah, Texas and northern Mex- ico; 32 (xvii). Salix lutea Nutt. Yellow willow. [S. I. var. platyphylla Ball; S. /. var. watsonii (Bebb) Jeps.]. Shrubs or rarely small trees but then still generally several stemmed at the base; (2) 3-5 (9) m tall; young twigs slender, yel- lowish to reddish at first, often pale on one side and red-purple on the other, glabrous; older twigs and smaller branches often gray- ish white; stipules small and inconspicuous or to 1 cm long or more and leaflike in texture on vigorous young shoots, usually deciduous; petioles 1-11 (20) mm long; leaf blades (1) 2-5.5 cm long, (4) 9-21 mm wide or to 10.7 cm long and 3 cm wide on vigorous young shoots, elliptical or lanceolate, rarely linear, entire or occasionally serrulate, glaucous beneath but hardly so when very young, usually glabrous at maturity, the lower surface glabrous from the first or less pubescent than above, the upper surface sometimes pubescent toward the base while th'e leaves are unfolding; aments precocious or subprecocious, on 1-7 mm long barren or 1-3 bracteate stalks; rachis and usually the stalk of the aments covered with a tangle of crisped-villous white hairs; bracts of the aments persistent, pubescent with crisped- villous, soon-entangled hairs, sometimes only moderately pilose-woolly toward the base or near the apex ventrally, the dorsal side usual- ly glabrous toward the apex and often throughout as the crinkly hairs are readily deciduous; staminate aments 2-5 cm long, about 1 cm wide; stamens 2, the filaments glabrous, the anthers yellowish or turning purple; pistillate aments 2-7 cm long, to 2 cm wide; capsules 3-6 mm long, glabrous, mostly densely arranged on the rachis, occa- sionally a little scattered, the stipe (1) 1.3-3 (4) mm long, the style 0.2-0.7 mm long, the stigmas often scarcely bilobed. Along streams and ditches in valleys and canyons and occa- sionally on mountains from 1340-2255 (2350) m, in all counties of the state except Beaver, Carbon, Davis, Iron, Morgan, and Rich, and to be expected in some or all of these; New Mexico to California and north to Alberta. Our plants are closely related to and possibly a part of the S. eriocephala Michx. complex. They have been referred to as S. rigida Muhl., but Argus (1980) has placed S. rigida in synonomy under S. eriocephala. He did not place S. lutea in synonomy, but suggested that more study is needed. Until such a study is made, I believe it best to re- tain the traditional name of S. lutea for our 548 Great Basin Naturalist Vol. 43, No. 4 plants. Salix ligulifolia Ball has been reported for southern Utah. Tliis has been separated from S. lutea by pedicels of capsules 1-2 mm long versus (1)2-4.5 mm long in S. lutea, and by having mostly entire rather than mostly semilate leaves. At the varietal level such plants are referrable to S. lutea var. liguli- folia Ball. See discussion under S. boothii; 144 (xxv). Salix monticola Bebb ex Coult. Shrubs 1.5-4 m tall; twigs yellowish when fresh, drying blackish, pubenilent at first; stipules small and inconspicuous or leaflike on vigor- ous young shoots; petioles 5-10 (15) mm long; leaf blades 2-5 cm long, 0.7-1.5 mm wide or up to 1 1 cm long and 4 cm wide on vigorous young shoots, mostly elliptical or el- liptic-obovate, crenate-serrate or subentire, slighty pubescent when very young, more so above than beneath, usually glabrous when fully expanded, glaucous beneath when ma- ture; aments precocious or coetaneous, sub- sessile or on short stalks to 1 cm long, often subtended by bractlike leaves; bracts of the aments persistent, dark brown to blackish, pi- lose, or soon crisped-villous, the hairs exceed- ing the bract by about 2 mm, more or less tangled; staminate aments 2-3.5 cm long, about 1-1.5 cm wide; filaments 2, glabrous; pistillate aments 2-6 cm long, 1-1.5 cm wide; capsules 4-7 mm long, glabrous, sub- sessile, the stipe less than 1 mm long; style 0.7-1.8 mm long, longer than the stigmas. Along streams and other wet places from 2195-3200 m, on mountains of eastern and central Utah in Beaver, Garfield, Piute, San Juan, Sanpete, Sevier, Uintah, and Wasatch coimties, and to be expected elsewhere in eastern Utah, but apparently uncommon in the state; Rocky Mountains of southern Wyoming, Colorado, Utah, Arizona, and New Mexico. Closely allied to S. boothii and S. lutea and rather easily confused with them. Separation from S. boothii is often com- pounded by the lack of glaucescence on yoimg leaves; I am indebted to Dr. Robert Dom for his annotations of specimens of this taxon. More specimens are needed to gain a better understanding of this plant in the state; 12 (0). Salix nigra Marsh. Black willow. [S. good- dingii Ball]. Trees or occasionally shrubs (2) 6-10 (24) m tall; twigs of the season yellow- ish, glabrous, or finely pubescent at first; stip- ules to 8 mm long, more or less glandular, usually quickly deciduous; petioles 3-7 mm long; leaf blades 2-7.5 cm long, 6-16 mm wide or to 10.2 cm long and 18 mm wide with petiole to 15 mm long on vigorous vege- tative twigs, narrowly to broadly lanceolate, apex short to long acuminate, entire or more often glandular-serrulate, greenish on both sides, pubescent when unfolding but becom- ing glabrous or glabrate; aments coetaneous, on lateral 1-6 cm long twigs of the season with 3-6 leaves or bracts; bracts of aments pale green or pale yellow, soon fading to tan, and at least the pistillate ones deciduous, pubescent on both sides or glabrous toward the apex, entire or with 1-3 minute, rounded teeth; staminate aments 2.5-6.5 cm long, 5-10 mm wide; stamens 3-6, the filaments pilose to about midlength; pistillate aments 1.5-6 cm long, 10-17 mm wide; capsules 4-7 mm long, glabrous, not so densely ar- ranged as to conceal the rachis, the stipe 1-2 mm long, the style 0.1-0.3 mm long. Along the Virgin and San Juan rivers and other drainages in southern Utah, and up the Green River to near Moab from 825-1585 m, in Garfield, Grand, Kane, San Juan, and Washington counties; widespread in the con- tinental United States, southern New Bruns- wick, and southern Quebec and Ontario, and northeastern Mexico. Utah specimens quite consistently have light-colored twigs and have been called S. gooddingii. I feel as did Archer (1965) that S. gooddingii is not clearly distinct from S. nigra. Arthur Cronquist (un- publ. ms.) has placed S. gooddingii in syn- onomy under S. tiigra var. venulosa (An- derss.) Bebb, and he recognized our plants as being different in having smaller stature and usually having some pubescence on the cap- sules or stipes, or both, as well as having light-colored twigs. However, he further states that these features are not consistent. I prefer to follow Cronquist's approach and recognize the differences in our plants at the varietal rather than at the specific level. Salix nigra and S. amygaloides come together near Moab on the Green River, and notes on specimens from that area by Arthur Cronquist indicate that the two hybridize at that location; 23 (0). October 1983 Goodrich: Utah Flora, Salic ace ae 549 Salix planifolia Pursh. Plainleaf willow. [S. plnjlicifolia L. ssp. planifolia (Pursh) Hiito- nen]. Shnibs 0.5-1.5 (4) m tall; twigs below the leaves often with epidermis exfoliating in translucent flakes or strips, younger twigs typically glabrous and lustrous black or purplish black, rarely glaucous in part; stip- ules small and asually deciduous; petioles 2-10 mm long; leaf blades 1.2-3.8 (8) cm long, 4-13 (30) mm wide, or to 5 (13) cm long and 2 (5) cm wide on vigorous sterile twigs, elliptical or narrow elliptical, soon glabrous and dark green above, glaucous and glabrous to sparingly pubescent below, entire or rarely with minute teeth; aments pre- cocious (at least the staminate) to coetaneous, nearly sessile or rarely on a short, mostly bar- ren pedvmcle to 0.5-1 cm long; bracts of the aments persistent, blackish, scattered to densely villose to pilose, the hairs usually ex- ceeding the bract by about 2 mm; staminate aments 10-25 mm long; stamens 2, the fila- ments glabrous, about 6 mm long; pistillate aments 2-4 cm long, 1-1.5 cm wide; capsules 3-7 mm long, typically pubescent at least near the base, occasionally glabrous or nearly so in age, the stipe mostly less than 1 mm long, the style and stigmas together mostly over 1.5 mm long. Streamside meadows, around lakes and ponds and other wet places, most abundant and sometimes forming wil- low fields in the Uinta Mountains, scattered on high points of the plateaus and mountains of the central and southern part of the state, from (2255) 2895-3660 m in Daggett, Du- chesne, Garfield, Iron, Salt Lake, Sanpete, Sevier, Summit, and Uintah counties; circum- boreal, south to California and New England. I have followed Argus (1973) in listing our plants under S. planifolia rather than S. phij- licifolia. Our plants mostly fall well within the concept of var. monica (Bebb) Jeps., though a few taller plants with larger leaves from moderate elevations of the major drain- ages in the Uinta Mountains are apparently var. planifolia. However, the differences are merely of stature and of leaf size and the two varieties are hardly worthy of separation; 39 (ix). Salix reticulata L. [S. nivalis Hook.; S. n. var. saximontana (Rydb.) Schneid.]. Caespi- tose dwarf shrubs, stems creeping at or just below the ground surface, the slender aerial twigs rarely more than 2-3 cm long, usually prostrate; stipules minute and deciduous or none; petioles 1-8 (15) mm long; leaf blades 0.5-3 cm long, 0.3-2 cm wide, ovate, ob- ovate, orbicular or occasionally broadly ellip- tical, entire, glabrous, green above, glaucous beneath, strongly reticulate veined; aments subcoetaneous, but mostly serotinous on the ends of shoots of the season; bracts of the aments persistent, pale green or yellowish, sometimes with reddish tops, spatulate or obovate, glabrous or sparsely pubescent ven- trally, especially toward the margin, with short hairs that extend less than 1 mm beyond the bract; staminate aments 0.5-2 cm long, slender, the flowers loose and not concealing the puberulent rachis, on a slender glabrous peduncle about 10-12 mm long; stamens 2; fikments 1.5-2 mm long, glabrous or pilose toward the base; anthers soon reddish or purple; pistillate aments 5-15 mm long, 5-8 mm wide, on a slender 1-2 mm long pe- duncle; capsules 1.5-3 mm long, pubescent or glabrous in age, sessile or the stipe to 0.5 mm long, the style obsolete or to 0.2 mm long, the stigmas about 0.1-0.2 mm long. Open rocky slopes and ridges and alpine tun- dra from 2987-3965 m, on the LaSal, Uinta, and Wasatch Mountains in Duchesne, Grand, Salt Lake, San Juan, Summit, and Utah coun- ties; circumboreal, south in the mountains of western North America to California, New Mexico, Utah, and Colorado. Most of our plants are referable to var. saximontana (L.) Kelso, which may not be distinct from var. reticulata. A few specimens seem to be like var. nivalis (Hook.) Anderss. The features used for separation seem to be poorly corre- lated in our plants. Some plants with pistil- late aments less than 1 cm long (that should be var. nivalis) have leaves well over 15 mm long, which is indicative of the other variety; 32 (0). Salix scouleriana Barratt in Hook. Scouler willow. Shrubs or small trees 3-7 m tall; stip- ules small and inconspicuous or large and leaflike on vigorous young shoots, eventually deciduous; petioles 2-11 mm long; leaf blades 2-6 cm long, (0.8) 1-3 cm wide or to 11.5 cm long and 4 cm wide on vigorous young shoots, obovate to oblanceolate, rounded to acute or occasionally accuminate at the apex, entire or finely serrate, or occa- 550 Great Basin Naturalist Vol. 43, No. 4 sionally coarsely crenate or serrate on larger leaves of vegetative twigs, densely crisp-hairy or sericeous, especially beneath as they un- fold, the mature ones dark green and glabrous above except sometimes puberulent along the midrib, the lower side strongly glaucous, sparsely puberulent with trans- lucent whitish or rusty minute hairs, or occa- sionally densely felty-villous; bracts of aments blackish or purplish black nearly throughout, reddish or pale at the very base, sericeous-pilose on both sides, the hairs at the apex usually exceeding the bract by 1.5-2 mm; staminate aments 15-35 mm long, nearly as wide as long, strictly precocious, nearly sessile or on thickened bracteate pe- duncles to 7 mm long, the bracts 3-4 mm long, about 2 mm wide, pale green to whit- ish, sericeous; stamens 2, the filaments to 11mm long at maturity, glabrous; pistillate aments 2-6 cm long, 13-17 mm wide, pre- cocious or subcoetaneous, nearly sessile or on thickened bracteate peduncles to 17 mm long, the bracts to 7 mm long and 2 mm wide, not at all leaflike; capsules (5) 6-9 mm long, pubescent, rarely nearly sessile, usually on a 1-3 mm long stipe, the style 0.3-0.4 mm long, rarely shorter, the stigmas 0.5-1 mm long. Around springs, along streams, and on well-drained slopes in aspen and conifer woods, from (1400) 2377-2835 (3355) m, in Box Elder, Cache, Carbon, Daggett, Davis, Duchesne, Garfield, Grand, Juab, Millard, Rich, Salt Lake, Sanpete, San Juan, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, Washington, and Weber counties; Alaska and Yukon to California, Arizona, and New Mexi- co. Salix scouleriana is most closely allied to S. humilus Marshall and to S. discolor Muhl. of eastern United States and Canada. Salix discolor (pussy willow) may be cultivated in our area. It is generally distinguished from S. scouleriana by: looser aments with longer stipes (1.5-3 mm), more elliptic, pointed, and toothed leaves that are usually more quickly and fully glabrate, but none of these features is wholly consistent (Hitchcock and Cronquist, 1964). Occasionally specimens have leaves densely pubescent beneath. Ar- now et al. (1980) attributed this to hybridiza- tion with S. drummondiana; 69 (xi). Salix tvolfii Bebb in Rothr. Wolf's willow. Shrubs 0.6-1.5 (2) m tall; twigs yellow to or- ange when young, chestnut brown in age, those of the season thinly villous-puberulent; stipules 1-5 mm long, often glandular-serru- late, eventually deciduous; petioles 2-10 mm long; leaf blades 1.2-4.2 cm long, 5-13 mm wide or to 5.3 cm long and 16 mm wide to- ward the ends of vigorous vegetative twigs, narrow elliptical, linear-lanceolate, or occa- sionally oblanceolate, entire, sparsely to densely sericeous-tomentose on both sides even in age or glabrate beneath very late in the season; aments coetaneous or sub- serotinous, nearly sessile or on bracteate pe- duncles to 1 cm long; bracts of the aments persistent, blackish or pale at the very base, pilose-sericeous on both sides, the hairs ex- ceeding the bract by about 1 mm; staminate catkins 10-15 mm long, about 8-10 mm wide; stamens 2, the filaments about 3-4 mm long, glabrous; pistillate aments 8-20 (30) mm long, 6-10 mm wide; capsules 3-5 mm long, glabrous or rarely pubescent, the stipe less than 1 mm long, the style about 0.5 mm long, the stigmas about 0.2 mm long. Along streams and around the margins of lakes and ponds, occasionally forming willow fields, in the Bear River and Uinta mountains and West Tavaputs and Wasatch plateaus from 2470-3290 m, in Cache, Daggett, Duchesne, Emery, Summit, Uintah, and Wasatch coun- ties. Oregon to Montana, south to Nevada, Utah, and Colorado. Our plants are var. tvolf- ii with mostly glabrous capsules. One speci- men (B. Maguire, D. Hobson, & R. Maguire 14104) from White Pine Lake, Cache Coun- ty, has pubescent capsules and leaves that are larger than others from the state. This speci- men is like S. wolfii var. idahoensis Ball, which is known from well north and west of Utah. Other specimens from the vicinity of White Pine Lake and other points in the Bear River Range have glabrous capsules, and I prefer not to list var. idahoensis for the state based on this one specimen. The plants from the Bear River Range with pistillate aments 15-30 mm long do, however, seem in- termediate toward var. idahoensis when com- pared to those of the Uinta Mountains with pistillate aments 8-15 mm long. The speci- men with pubescent capsules and somewhat large leaves is probably the basis of reports of S. commutata Bebb for Utah; 44 (ix). A MAMMALIAN HUMERUS FROM THE UPPER JURASSIC OF COLORADO Donald R. Prothero' and James A. Jensen- Abstract.— The first reported mammal fossil from Dry Mesa Quarry (Upper Jurassic Morrison Formation, Mesa Countv, Colorado) is the distal end of a right humerus. It is very similar to humeri described by Jenkins (1973) from the Morrison Formation at Como Bluff, Wyoming. It has a distinct ulnar condyle and a spiral humero-ulnar joint, both features found in prototherian mammals but not in therians. Postcranial remains of Jurassic mammals are extremely rare. An articulated skeleton of a dryolestid therian mammal has been report- ed from the Jurassic of Portugal (Henkel and Krebs 1977), but is still undescribed. Early Jurassic mammalian postcranial fossils are also known from India (Datta et al. 1978), but are undescribed. A few fragmentary post- cranial remains of mammals have been de- scribed from the Upper Jurassic of England (Seeley 1879, Simpson 1928, Haines 1946) and from the Upper Jurassic Morrison For- mation of Wyoming (Jenkins 1973). Of the five important mammal-producing localities in the Morrison Formation (listed in Clemens et al. 1979:23-26), two have produced mam- malian postcranial fossils prior to this paper: Como Bluff, Wyoming (Jenkins 1973), and the Fruita Paleontological Area, Mesa Coun- ty, Colorado (Rasmussen and Callison 1981). In 1977, the distal portion of a right hu- merus of a mammal was found in Dry Mesa Quarry, Mesa County, Colorado. This speci- men (BYU 2026) was first mentioned by Cle- mens et al. (1979:24), and is described below. Locality and Associated Fauna Dry Mesa Quarry is located in the lower section of the Brushy Basin Member of the Upper Jurassic Morrison Formation, 135 feet below its contact with the overlying Cre- taceous Cedar Mountain Formation. The quarry sediments include very fine to coarse sands, grits, and fine gravels containing angu- lar to well-rounded clay and bone pebbles. Stream gradient was sufficient to move very large bones, with the long axes of all large bones usually oriented at right angles to the stream flow. Sorting was biased by shape rather than by size. Sediments overlying the bone layer are predominantly light-colored, cross-bedded sands with occasional lenses of clay and fine gravel, the latter often containing clay peb- bles. Sediments underlying the bone layer are principally a light, blue-green clay with oc- casional traces of bright yellow zones of oxidation. The bone layer consists of an imusual vari- ety of disarticulated bones of all sizes, includ- ing specimens representing crocodilians, fish, turtles, pterosaurs, four new theropods, an unknown variety of sauropods, some ornitho- pods, and the mammal described herein. Due to the great variety of disarticulated bones in the deposit, and the generic novelty of the fauna, descriptive work has been postponed until enough material is available. Field work has been carried out for the last 10 years. The following have been identified so far: Lungfish tooth plate (probably Ceratodus— K. Thomson, pers. comm.) Pterodactyloid phalanx (Jensen and Ostrom 1977) Torvosaurus tanneri, a megalosaur (Galton and Jensen 1979) Prototherian mammal humerus (this paper) Description BYU 2026 (Fig. 1) is the distal portion of a right humerus of a mammal. It has been bro- ken at midshaft, but is otherwise well pre- served. There is relatively little evidence of crushing or distortion. The shaft cross-section 'Department of Geology, Knox College, Galesburg, Illinois 61401. ■Earth Science Museum, Brigham Young University, Prove, Utah 84602. 551 552 Great Basin Naturalist Vol. 43, No. 4 is a mediolaterally compressed triangle with the apex pointing anteriorly. Distally the shaft expands transversely to form the large medial and lateral epicondyles. The distal end of the hiunerus is naturally flattened an- teroposteriorly. The long axis of the distal end is not perpendicular to the major axis of the shaft cross-section, but is rotated about 20 degrees clockwise (viewed from the distal end). The shaft of the humerus has a strong anterior crest that is deflected anterolat- erally. This crest is probably the distal end of the deltopectoral crest. A faint posterolateral crest merges with the lateral epicondyle. The medial epicondyle is considerably more prominent than the lateral epicondyle. It flares medially and is anteroposteriorly compressed. The entepicondylar foramen is visible in anterior view. It is broken at the anterior end, where it passes anteromedially. The lateral epicondyle merges with the radial condyle. It is connected to the shaft of the humerus by a thin posterolaterally arched crest. The radial and olecranon fossae are in- terconnected, forming an apparent supra- trochlear foramen. This feature may be an ar- tifact of breakage, however. In anterior view, the main body of the shaft bifurcates to form crests joining the radial and ulnar condyles. These crests sur- round the radial fossa. In posterior view, the olecranon fossa is broadly concave and ex- tends partially up the shaft. From this view the apparent supratrochlear foramen has an irregular margin that is clearly enlarged by breakage. In distal view, three main features are seen: the medial epicondyle, the ulnar con- dyle, and the radial condyle-lateral epicon- dyle. The latter two features are confluent and separated only by a shallow groove. The ulnar and radial condyles, on the other hand, are separated by a narrow, deep inter- condylar groove. The radial condyle is broad and bulbous in anterior view. The spiral ul- nar condyle is very similar to that shown by Jenkins (1973, Fig. 13). It is wrapped around the distal end of the humerus, with a prox- imolaterally oriented extensor surface and a proximodistally oriented flexor surface. How- ever, the anterior portion of the ulnar con- dyle is more bulbous than the same feature in the humerus figured by Jenkins (1973, Fig. Fig. 1. BYU 2026, distal end of a right humerus. A, Anterior view. B, Posterior view. C, Distal view. Abbre- viations: dpc, deltopectoral crest; enf, entepicondylar foramen; le, lateral epicondyle; me, medial epicondyle; of, olecranon fossa; re, radial condyle; rf, radial fossa; uc, ulnar condvle. 18). The axis of the ulnar condyle as it crosses over the distal end of the humerus is at an approximately 60 degree angle to the trans- verse (interepicondylar) axis of the humerus (seen in distal view). This compares with an- gles of 58-65 degrees reported by Jenkins (1973:286) for several humeri from Como Bluff, Wyoming. Discussion The Dry Mesa Quarry mammal very close- ly resembles the Como Bluff humeri de- scribed by Jenkins (1973). It differs from them in having a more bulbous and broader ulnar condyle. In this respect, it is more like the humeri referred to the multituberculate Catopsolis by Jenkins (1973, Fig. 19). The multituberculates Tugribataar (Kielan-Jawo- rowska and Dashzeveg 1978) and Ptilodus (Gidley 1909), the triconodont Eozostrodon (Jenkins and Parrington 1976), and the mon- otremes (Howell 1937, Haines 1946) also have prominent bulbous ulnar condyles. BYU 2026 clearly does not have a trochlear con- dyle, which Jenkins (1973) considers charac- teristic of therian mammals. The only other feature that distinguishes the Dry Mesa Quarry mammal from the Como Bluff humeri is the apparent supra- trochlear foramen. As noted above, this fea- ture may be an artifact of breakage. October 1983 Prothero, Jensen: Jurassic Mammalian Humerus 553 The affinities of BYU 2026 are difficult to assess based on such hmited evidence. The presence of a distinct ulnar condyle with a spiral humero-ulnar joint is characteristic of prototherian mammals (Jenkins 1973). The advanced therian trochlear condyle is known from rocks as old as the Lower Cretaceous (Jenkins 1973, footnote 3). The Dry Mesa Quarry mammal humerus could have be- longed to a number of prototherian mammal taxa presently known from the Morrison For- mation (Prothero 1981, Clemens et al. 1979). It could also have come from some of the primitive Morrison therian mammals that may or may not have had a trochlear con- dyle. Until the Portuguese dryolestid therian skeleton (Henkel and Krebs 1977) is fully de- scribed, we cannot rule out the possibility tliat the Dry Mesa Quarry humerus belonged to a very primitive therian mammal. Of the possible candidates among nontherian mam- mals, BYU 2026 resembles the known humeri of multituberculates and triconodonts. The skeleton of docodonts is presently unknown. The systematic affinities of the Dry Mesa Quarry mammal cannot be determined more precisely at present than Mammalia incertae sedis. Acknowledgments The Dry Mesa Quarry has been worked for 10 years by the Earth Science Museum of the Brigham Young University under permits from the U.S. Forest Service, as authorized by the Antiquities Act of 1906. Funds for this work have been provided by research sup- port grants from Brigham Young University, with additional support from the National Geographic Society, Kenneth R. Thomson, and several other private donors. Measurements of angles were made with an E.P.O.I. Shopscope. D.R. Prothero was supported by a Columbia Faculty Fellowship diu-ing preparation of this paper. We thank Henry Galiano for advice on the illustration. Literature Cited Clemens, W. A., J. A. Lillec;raven, E. H. Lindsay, and G. G. Simpson. 1979. Where, when and what— a survey of known Mesozoic mammal distribution. Pages 7-58 in J. A. Lillegraven, Z. Kielan-Jawo- rowska, and W. A. Clemens, eds., Mesozoic mammals: the first two-thirds of mammalian his- tory. Univ. of California Press, Berkeley. Datta, p. M., p. Yadagiri, and B. R. J. Rao. 1978. Dis- covery of Early Jurassic micromammals from up- per Gondwana sequence of Pranhita (iodavari Valley, India. Geol. Soc. India J. 19:64-68. Galton, p. M., and J. A. Jensen. 1979. A new large theropod dinosaur from the Upper Jurassic of Colorado. Brigham Young Univ. Geology Studies. 26(2):1-12. Gidley, J. W. 1909. Notes on the fossil mammalian genus Ptilodus, with descriptions of new species: Proc. U.S. National Museum .36:611-626. Haines, R. VV. 1946. A revision of the movements of the forearm in tetrapods. J. Anatomy 80:1-11. Henkel, S., and B. Krebs. 1977. Der erste Fund eines Saugetier-Skelettes aus der Jura-Zeit. Umschau in Wissen.schaft und Technik 77:217-218. Howell, A. B. 1937. The swimming mechanism of the platypus. J. Mammal. 18:217-222. Jenkins, F. A., Jr. 1973. The fimctional anatomy and ev- olution of the mammalian humero-ulnar articu- lation. Amer. J. Anat. 137:281-298. Jenkins, F. A., and F. R. Parrington. 1976. The post- cranial skeletons of the Triassic mammals Eo- zostrodon, Megazostrodon, and Enjthrotherium. Roy. Soc. London Philos. Trans., B (Biol. Sci.) 27.3:387-431. Jense.n, J. A., and J. H. Ostrom. 1977. A second Jurassic pterosaur from North .-America. J. Paleont. 51:867-70. Kielan-Jaworowska, Z., and D. Dashzeveg. 1978. New Late Cretaceous mammal locality in Mongolia and a description of a new multituberculate. Acta Palaeont. Polonica 23:115-130. Prothero, D. R. 1981. New Jurassic mammals from Como Bluff, Wyoming, and the interrelationships of the non-tribosphenic Theria. Bull. Amer. Mus. Nat. Hist. 167 (art ,5):277-326. Rasmussen, T. E., and G. Callison. 1981. A new spe- cies of triconodont mammal from the upper Ju- rassic of Colorado. J. Paleont. 55:628-6.34. Seeley, H. G. 1879. Note on a femur and humerus of a small mammal from the Stonesfield Slate. Quar- terly Journal (Teological Society of London 35:456-463. Simpson, G. G. 1928. A catalogue of the Mesozoic mam- malia in the Geological Department of the Brit- ish Museum. British Museum (Natural History), London. BATS OF THE COLORADO OIL SHALE REGION Robert B. Finley, Jr.,' William Caire,' and Dallas E. Wilhelm' Abstract.— New records for Myotis californicus, M. evotis, M. leibii, M. lucifugus, M. thysanodes, M. volans, M. yunuinensis, Lasionycteris noctivagans, Pipistrelhis hespertis, Eptesicus fiiscus, Lasiuriis cinereus, Plecotus town- sendii, and Antrozous pallidus and their habitat occurrence in northwestern Colorado are reported. Mortality of 27 bats of six species trapped in an oil sludge pit is described. In 1974 the National Fish and Wildlife Laboratory began field work in the Piceance Basin as part of a survey of the vertebrates of the coal and oil shale regions of northwestern Colorado. The information was needed as baseline data in preparation for energy devel- opment and to better define the poorly known distribution of mammals in this area. From the oil shale region of Colorado, roughly defined as Rio Blanco and Garfield counties west of a line between Meeker and Rifle, records of eight species of bats were summarized by Armstrong (1972), seven of which had been reported only from the vi- cinities of Meeker or Rifle. Since then speci- mens of five additional species have been col- lected and numerous locality and habitat records obtained. This information is report- ed herein to make it available for manage- ment decisions, and to facilitate and stimu- late further work on the bats of northwestern Colorado. Study Area and Methods The elevation of the Oil Shale Region ranges approximately from 1,585 to 2,805 m, falling within the Upper Sonoran, Transition, and Canadian life zones of Gary (1911). The Roan Plateau extends east-west, roughly along the Rio Blanco-Garfield county line, di- viding the two main drainages, the Colorado River to the south and the White River to the north. The region is semiarid with pre- dominantly shaly alkaline soils and has only a few long permanent creeks, with many short intermittent tributaries. The few permanent creek bottoms are occupied by ranches and irrigated hay meadows. Low cliffs and rock ledges border some of the creeks, and high cliffs mark the southern rim of the Roan Plateau and the western rim of the Cathedral Bluffs east of Douglas Creek. Specimens were obtained primarily by mist-netting and were preserved as study skins and skulls or in liquid. All specimens ex- amined are in the Biological Surveys/ Fort Collins collection (BS/FC) of the U.S. Fish and Wildlife Service, unless otherwise in- dicated: (CU) Colorado University Museum or (KU) Kansas University Museum of Natu- ral History. Results The known kinds of bats from the Colo- rado oil shale region include seven species of Myotis and one species in each of six other genera {Lasionycteris, Pipistrellus, Eptesicus, Lasiurus, Plecotus, and Antrozous). Species found to be most common were the hoary bat {Lasiurus cinereus cinereus), long-legged myotis {Myotis volans interior), long-eared myotis {Myotis evotis evotis), and small-foot- ed myotis {Myotis leibii melanorhinus). The spotted bat {Euderma maculatum) has been reported in some environmental impact re- ports as "potentially" or "possibly" present; yet to our knowledge no specimen has been taken in the oil shale region. One was picked up in Browns Park, 65 km north of the oil shale region, in 1981 by J. Creasy (Finley and Creasy 1982). 'U.S. Fish and Wildlife Service, 1300 Blue Spruce Drive, Fort Collins, Colorado 80524-2098. 'Department of Biology, Central State University, 100 North University Drive, Edmond, Oklahoma 73034. 'Department of Biology, Hastings College, P.O. Box 269, Hastings, Nebraska 68901. 554 October 1983 FiNLEY ET AL.: COLORADO BaTS 555 Accounts of Species Myotis californicus stephensi Dalquest California Myotis Specimens examined (5).— Garfield Co.: 5'/2 mi N, 2 mi W Rifle, 5,900 ft, 5. Five California myotis, four unsexed and one female, were found partially decomposed in an oil sludge pit on the east side of the val- ley of Government Creek north of Rifle. Four specimens were picked up on 17 Sep- tember 1974 and one on 17 May 1975. They are referred to stephensi on geographic grounds. The circumstances of these and 22 other bat casualties at the same sludge pit are described in this paper under "Effects of Energy Development on Bats." The National Museum of Natural History has a specimen taken 14 August 1907 by Merritt Cary (1911) in a house 7 mi W Rifle, 5,300 feet. Myotis evotis evotis (H. Allen) Long-eared Myotis Specimens examined (31).— Garfield Co.: 5'/2 mi N, 2 mi W Rifle, 5,900 ft, 7; Rio Blanco Co.: Cascade Gulch, T3S, R95W, Sec 8, 1; Duck Cr tank, TIS, R98W, Sec 7, 1; Little Duck Cr, TIS, R98W, Sec 10, 6,200 ft, 10; Ryan Gulch, 24 mi W, 10 mi S Meeker, 6,500 ft, 5; 25 mi'w, 10 mi S Meeker, 6,400 ft, 2; 1 mi N, 4 mi W Rio Blanco, 6,900 ft, 5. Seven long-eared myotis carcasses (1 male, 3 females, and 3 unsexed) were picked up on the bank of the sludge pit north of Rifle on 17 September 1974. This species was the most abundant bat taken at pools adjacent to big sage (Artemisia tridentata), greasewood (Sarcobatus vermiculatus), and pinyon-juni- per habitats. Sandstone ledges or low cliffs were also present within a few hundred yards at all localities. The most productive mist netting site was on Little Duck Creek, at 2,000 m elevation. There on 7 August 1975 Caire and Finley collected 10 M. evotis, 2 M. leibii, 4 M. vol- ans, 3 Eptesicus fuscus, and 2 Lasiurus cine- rus. The valley bottom was covered with tall, dense big sagebrush and greasewood on deep alluvial soil. Little Duck Creek was en- trenched in an arroyo about 3-4V2 m deep. The steep upper slopes of the valley were of nearly bare shaly soil. Higher rock outcrops along the rim supported old mature junipers (Juniperus osteosperma). The water in the arroyo was barely enough to provide a flow, but a deposit of silt at the mouth of a side gully had dammed the flow and formed a long pool about 4 m wide and 20 cm deep. Nets were set over this pool and another about 40 m downstream, where we dammed the stream with a shovel, creating a pool. One long-eared myotis caught at Little Duck Creek had a large laceration in the left wing membrane, which had fully healed. The study skin (BS/FC 2119), pinned with wings fully extended, has the posterior margin of the membrane between the tibia and the 5th metacarpal deeply concave so that the width (chord) of the left wing measures only 22 njm, compared with 37 mm at the same place on the right wing. A lobe of smooth scar tissue extends anteriorly from the de- formed posterior margin, leaving only an 8 mm band of normal membrane between the scar tissue and the radius. In Ryan Gulch Wilhelm netted bats at a steel stock tank by a windmill on 15 August 1974. He took 5 Af. evotis, 5 M. volans, and 2 M. leihii. The terrain and habitat there were almost the same as on Little Duck Creek, ex- cept that Ryan Gulch was dry at the time and not so deeply gullied. Less than a mile away in June two macerated skeletons of M. evotis that had apparently drowned in a small watering trough were found. Three of four female M. evotis taken on 4 August 1977 on Piceance Creek, 1 mi N, 4 mi W Rio Blanco, were lactating or recently lactating. The habitat there is described in the M. lucifugus account. Of the 26 total specimens of long-eared myotis that were sexed, 14 were females and 12 were males. Myotis leibii melanorhinus (Merriam) Small-footed Myotis Specimens examined (14).— Garfield Co.: 3 mi N Douglas Pass, 7,000 ft, 1; 4.6 mi W Rifle (by rd), T6S, R94W, Sec 14, 1; 5'/2 mi N, 2 mi W Rifle, 5,900 ft, 5; Rio Blanco Co.: Cascade Gulch, T3S, R95W, Sec 8, 2; Little Duck Cr, TIS, R98W, Sec 10, 2; Ryan Gulch stock tank, 24 mi W, 10 mi S Meeker, 6,500 ft, 2; Ryan Gulch, 25 mi W, 10 mi S Meeker, 1. Five M. leibii were picked up (4 on 17 September 1974 and 1 on 29 April 1975) on 556 Great Basin Naturalist Vol. 43, No. 4 the bank of the same oil sludge pit north of Rifle where 5 M. califomicus and 7 M. evotis were found. There were 2 females, 1 male, and 2 imsexed carcasses. This species seems to be found mainly at lower and intermediate elevations in sage- bnish, greasewood, and pinyon-juniper habi- tats, but is less abundant there than M. evotis. Our highest record of occurrence was at 2,130 m, 3 mi N Douglas Pass, where one was netted along with Lasiunis cinereus on 2 August 1977 over an earthen stock pond. The adjacent hillsides supported mixed chaparral, sagebrush, and grassland with scattered Douglas-fir (Pseudotsuga menziesii) and juni- pers (Junipenis). A female taken on 22 July 1975 at a sheep stock tank 4.6 mi W Rifle contained an em- bryo 20 mm long. Of the 12 individuals col- lected that were sexed, half were females. The eight skins at hand show a wide varia- tion in color of the back and sides, from light buffy or reddish brown to dull medium brown, but have uniformly blackish ears and wings. Myotis lucifugus carissima Thomas Little Brown Bat Specitnens examined (3).— Garfield Co.: 4 mi W, 1 mi S Rifle, 5,300 ft, 1; 1 mi N, 4 mi W Rio Blanco, 6,900 ft 2. A female was taken on 28 August 1975 over a sheep watering pond in the Colorado River valley west of Rifle. The pond had been bulldozed in alkali soil on a greasewood flat south of cliffs and a steep rock slope with pinyon-juniper. Two males were taken in a net across up- per Piceance Creek west of Rio Blanco on 4 August 1977. At that point the valley is nar- row between steep canyon sides covered with pinyon-juniper and mountain shrub, with Douglas-fir in the tributary gulches. The val- ley bottom was grazed by sheep, and the creek was barely flowing between long shal- low pools. Other bats taken there were four M. evotis and one Lasionycteris. Cary (1911:206) collected two M. luci- fugus on the White River meadows a few miles east of Meeker in August 1905. Myotis thysanodes thysanodes Miller Fringed Mvotis Specimens examined (1).— Rio Blanco Co.; 11 mi W Meeker, '/2 mi S Hwy 64, Hav Gulch Rd, 1 (CL'). A male fringed myotis was netted by Jerry Freeman on 17 August 1978 at a stock tank where Hay Gulch opens into the White River Valley. Myotis volans interior Miller Long-legged Myotis Specimens examined (14).— Garfield Co.: 2 mi E Rio Blanco, 1; Rio Blanco Co.: Little Duck Cr, TIS, R98W, Sec 10, 4; Little Hills Game Research Station, 15 mi W, 1.9 mi S Meeker, 4 (CU); Ryan Gulch, 24 mi VV, 10 mi S Meeker, 5,600 ft, 5. Myotis volans occurs in the sagebrush, greasewood, and pinyon-juniper habitats along with M. evotis and M. leihii. Specimens were taken at sites described under the ac- counts of these species, and also at a more wooded site described under Lasiurus cinereus. Two males and two females were taken in the barn at the headquarters of the Little Hills Game Research Station by Bissell, Olivas, and Webb of the Colorado Division of Wildlife on 29 June and 7 July 1977. The barn is in a gulch flanked by rocky slopes with pinyon-juniper. One M. ywnanensis and one Eptesicus fuscus were also obtained there. Eight of the 14 M. volans examined were males and 6 were females. Myotis yumanensis yumanensis (H. Allen) Yuma Myotis Specimens examined (4).— Rio Blanco Co.: Little Hills Game Research Station, 15 mi W, 1.9 mi S Meeker, 1 (CU); Rio Blanco Lake, 3 (CU). Three female Yuma myotis were taken on 6 and 7 July 1977 by Ribic and Olivas in the attic of a deserted house by Rio Blanco Lake, a small reservoir on the White River near the mouth of Piceance Creek. One contained three embryos 4 mm in length. Lasionycteris noctivagans (Le Conte) Silver-haired Bat Specimens examined (8).— Garfield Co.: 5'/2 mi N, 2 mi W Rifle, 5,900 ft, 6; Rio Blanco Co.; 31/2 mi W Rio Blanco, 1; 1 mi N, 4 mi W Rio Blanco, 6,900 ft, 1. October 1983 FiNLEY ET AL.: COLORADO BaTS 557 Six silver-haired bats (5 males and 1 fe- male) were foimd at the sludge pit north of Rifle. One each day was picked up in a fairly good state of preservation on 29 April, 14 May, and 17 May, two on 4 June 1975, and one still alive on 17 September 1974. Only two Lasionycteris were taken in mist nets, one each at two sites on Piceance Creek west of Rio Blanco, where they were associ- ated with M. lucifugns at one site and La- siurus cinereus at the other. The sites are de- scribed under those accounts. Pipistrelltts hesperus hesperus (H. Allen) Western Pipistrelle Specimens examined (2).— Garfield Co.: 5V2 mi N, 2 mi W Rifle. 5,9(X) ft, 1; 4 mi W, 1 mi S Rifle, 5,300 ft, 1. One P. hesperus was picked up on 17 Sep- tember 1974 at the sludge pit north of Rifle. The only pipistrelle netted was in the Colo- rado River valley at the locality described under M. lucifugus. These records and the absence of specimens from our other sites at higher elevations bear out Cary's (1911:209) statement that pipistrelles "inhabit the Upper Sonoran zone in the western and south- western valleys . . . [and] hve only about cliffs and in rock- walled canyons." Eptesicus fuscus pallidus Young Big Brown Bat Specimens examined (4).— Rio Blanco Co.: Little Duck Cr, TIS, R98W, Sec 10, 6,200 ft, 3; Little HilLs Game Research Station, 15 mi W, 1.9 mi S Meeker, 1 (CU). One big brown bat was shot flying over tall sagebrush and two were netted over a pool in a gully at Little Duck Creek on 7 Au- gust 1975. The habitat and other captures are described under the account of Myotis evotis. Cary (1911:209) obtained a big-brown bat from the White River, a few kilometers east of Meeker in August 1905. The four skins vary in color, as discussed by Armstrong (1972:70), and fall within the range of a series of nine pallidus in the BS/FC collection from 5 mi E, 1 mi N Fort Collins, except that one is more light reddish than any in the Fort Collins series. Lasiurus cinereus cinereus Palisot de Beauvois Hoary Bat Specimens examined (16).— Garfield Co.: 3 mi N Douglas Pass, 7,000 ft, 1; 3.4 mi N Rifle, on Government Creek, 1; 5'/2 mi N, 2 mi W Rifle, 5,900 ft, 3; 2 mi E Rio Blanco, 7,500 ft, 3; Rio Blanco Co.: Little Duck Cr, TIS, R98W, Sec 10, 2; TIN, R102W, Sec 4, W of Rangely [5,300 ft], 1; 3'/2 mi W Rio Blanco, 5. The hoary bat was the most abundant spe- cies in mist nets set over pools of Piceance Creek adjacent to cottonwood (Populus) and Douglas-fir stands between 2,000 and 2,300 m elevation. One was also netted over an ephemeral pond a few miles west of Rangely in salt desert shrub at 1,615 m elevation. Three hoary bats were picked up at the sludge pit north of Rifle. A female was found OH 15 July 1974 and a male and a female on 17 September 1979. Two male hoary bats were netted on Little Duck Creek on 7 Au- gust 1975 at the site described under the ac- count of Myotis evotis. Five male hoary bats were caught in two nets over Piceance Creek, SVz mi W Rio Blanco, on the night of 10-11 July 1975. One net was set over a narrow beaver pond in a grove of cottonwoods and the other in a grassy area just below the junction of Cow Creek and Piceance Creek. The Piceance valley was narrow between steep hillsides wooded with small Douglas-fir and mixed mountain shrubs on the shady side and mostly pinyon-juniper on the sunny slope. One female and two male hoary bats, as well as one female M. volans, were netted on 10-11 July 1975 over a beaver pond on up- per Piceance Creek, 2 mi E Rio Blanco, 7,300 ft. The site was more wooded than the one just described (3V2 mi W Rio Blanco). The north-facing slope above the beaver pond was covered with Douglas-fir, and the opposite slope had a heavy stand of pinyon- juniper and mountain mahogany (Cerco- carpus montanus). Other localities where hoary bats were taken are described under the accounts of Myotis leihii and Antrozous pallidus. A pregnant female (BSC/FC 5109, alco- holic) was taken on 11 July 1975, 2 mi E Rio Blanco. She weighed 30 grams and carried two embryos, 22 mm. All 11 of the other 558 Great Basin Naturalist Vol. 43, No. 4 hoary bats taken by mist nets were males. The 11 skins available vary considerably in appearance. The three taken the latest in the summer (7 August) have longer dorsal fur with a heavier overlay of silver tips. Some taken in July have short pelage and some are moulting. Plecotus townsendii pallescens (Miller) Townsend's Big-eared Bat Speci7nens examined (5).— Rio Blanco Co.: Spring Cave, 7,850 ft, 9 mi S, 4 mi E Biiford, 4; 5 mi N, 10 mi W Rangely, 5,800 ft, 1 (KU). On 19 February 1977 Finley visited Spring Cave to investigate reports of two kinds of bats there in winter, one with "long" ears and one with "short" ears. This limestone cave is located high on a forested mountain- side in the White River National Forest. A detailed description, photos, and maps of Spring Cave were published by Parris (1973:221). The mountainside was snow cov- ered, but temperatures were mild. There were a few scattered dormant bats on the wall of the entry passage between the two main entrances, some even in the twilight zone. There were about 90 bats in all high on the wall of the "Long Room" extending southwest from the entry passage. About 25 were hanging singly and about 65 in 3 clus- ters, mostly 2 to 3.7 m above the floor. All appeared to be Plecotus, some with ears ex- tended and some with ears folded, giving a first impression of two kinds of bats present. No bats were seen beyond the bend in the Long Room or on the walls of the "Tunnel" or "Pirates' Den." Wilhelm visited Spring Cave on 15 August 1974 and explored it without finding any bats or guano deposits. Armstrong (1972) reported one specimen at the University of Colorado Museum from Spring Cave. A single male big-eared bat was taken by Finley on 29 August 1948 in a sandstone cave 5 mi N, 10 mi W Rangely, 5,800 ft., Rio Blanco County. It was found in an overhead vertical cleft through the roof of an arched rock shelter that was also occupied by a bushy-tailed wood rat (Neotoma cinerea arizonae). Antrozous pallidus pallidus (Le Conte) Pallid Bat Specimens examined (4).— Garfield Co.: .3.4 mi N Rifle, on Government Cr, 3; Rio Blanco Co.: TIN, R102W, Sec 4, W of Rangely, 1. Three pallid bats, lactating females, were taken by Caire on 25 July 1975 in a net set under cottonwoods over a trickle of water in Government Creek north of Rifle. The creek was nearly dry and lined with large cotton- woods and a few tamarisks {Tamarix gallica). On 7 August 1975 another lactating female was taken west of Rangely on the dry, low sage-saltbush bench south of the White Riv- er. The net was set across the lower end of a shallow, mud-flat pool in a small arroyo. A single hoary bat was the only bat taken in the evening, and the pallid bat was found in the net the following morning. Gary (1911:206) reported seeing a pallid bat "among the cliffs along the Grand [Colo- rado] River, 7 miles west of Rifle, August 14, 1907." Effects of Energy Development ON Bats Man's increasing activities for devel- opment of energy resources in western Colo- rado can entail several kinds of adverse ef- fects on bat populations, such as disturbance of hibernating bats in caves, mine tunnels, and other sites; destruction of daytime roost sites; loss or contamination of watering sites; and contamination of the food chain by in- secticides and pollutants. Bats are extremely vulnerable because of their specializations as volant, nocturnal in- sectivores. For protection against ground predators they require secure daytime roosts such as cave walls, cliff crevices, or hollow tree snags. Suitable shelters in the oil shale region are usually located in narrow canyons and ravines or on juniper-covered rimrock. Such terrain features occupy a low percent- age of total land area and should be avoided as waste disposal sites for spent shale from retorts. Both natural and man-made water surfaces are widely available to bats for drinking in northwestern Colorado. The main permanent streams such as the White River and Pi- ceance Creek are important and likely to October 1983 FiNLEY ET AL.: COLORADO BaTS 559 remain dependable drinking sites. But the small intermittent streams and springs are very susceptible to loss by lowering water ta- bles when wells are drilled, and may become contaminated by waste dumping or spillage. Loss of springs and small stream pools on the upland areas may be compensated for by presence or addition of stock-watering tanks. As long as cattle raising is maintained as a vi- able industry in the areas of energy devel- opment, the adverse effects on bats of spring and creek flow loss can be at least partially mitigated. However, production of aquatic insect food would still be reduced. Drilling for oil and gas in Colorado results in a certain amount of drilling fluids and oil at well sites being spilled and impounded in pools or sludge pits. These are recognized hazards to birds and other wildlife, but bats have not previously been reported as casu- alties. A source of bat mortality was called to our attention by Richard E. Fillmore, who picked up a mummified hoary bat carcass on 15 July 1974 on the bank of a sludge pit 5V2 mi N, 2 mi W Rifle at 1,800 m elevation. Pio- neer Drilling, Inc., had drilled a test well and dug two pits for impounding spilled oil. The sludge pool was situated in the bottom of a pit with steep banks rising about 3 m above the surface, which was about 23 by 30 m in size. Oil in the pool formed a thin slick over the water, which was a few feet deep. This lower pool was formed by drainage from an adjacent higher pit that contained much heavier oil sludge and was nearly dried up. The oil well and pits were on a bulldozed pad of alkaline clay soil on the east side of the valley. The pad was surrounded by a gentle slope covered with greasewood. To the east was a steep rocky slope of pinyon- juniper, leading up to high cliffs bounding the east side of the valley of Government Creek. The lower of the two sludge pits proved to be a lucrative source of bats. They were found 3 to 8 m from the edge of the lower pool, but none were found around the upper, nearly dry pit. In the following 16 months re- peated visits to this site by Finley and others yielded 27 poor-quality but identifiable spec- imens. The kinds and numbers of individuals collected are as follows: hoary bat (3), long- eared myotis (7), small-footed myotis (5), Cal- ifornia myotis (5), western pipistrelle (1), sil- ver-haired bat (6). A live, oil-soaked Lasionycteris was seen on 17 September 1974 hanging on the side of a rock about 1 m from the oil-slick surface of the pond. It responded to touch but did not seem to be much affected by the oil. In comparison with collections from other localities, there were relatively high numbers of silver-haired bats, only two of which have been taken by mist net in the region, and of California myotis, none of which has been mist-netted; but the pallid bat, three of which were netted only 5 km away, is absent from the sludge pit sample. A more common species not far away, Myotis volans, was also absent from the sludge pit. Perhaps various species of bats differ in their vulnerability to entrapment. On 19 September 1974 Finley visited the Rangely oil field and inspected sludge pits without finding any bats. The six pits that contained oil were all much smaller than the pit north of Rifle and had heavy oil, appar- ently without water, in the bottom. They had wires with colored streamers stretched over them, apparently to keep out birds. It does not seem likely that bats would mistake such pits for water ponds. Four sludge pits con- taining oil on water were inspected on 1 May 1981 between Craig, Colorado, and Rock Springs, Wyoming, without finding any bats. Three instances of entrapment of bats in oil are known to us in the literature: Krutzsch (1948) described finding 3 bats as well as insects, lizards, and birds in three small pools of oil formed by drainage into a gully from a recently oiled road in the Bo- rego Desert, California. Barbour and Davis (1969) cited a report by E. J. Koestner of 15 red bats {Lasiurus borealis) entrapped in oil on a 500-foot stretch of road in Illinois. Gil- lette and Kimbrough (1970) reported a bat seen in a "tar pit" of undescribed origin at Fort Sill, Lawton, Oklahoma. We thank Jerry Freeman for permission to report a specimen of Myotis thysanodes taken by him, Michael Bogan for identifying several bats, and the Colorado Division of Wildlife for generous information and assistance. 560 Great Basin Naturalist Vol. 43, No. 4 Literature Cited Armstrong, D. M. 1972. Distribution of mammals in Colorado. Mono. Mus. Nat. Hist., Univ. of Kansas 3:1-415. Barbour, R. W., and W. H. Davis. 1969. Bats of Ameri- ca. Univ. Press of Kentucky, Lexington. 286 pp. Gary, M. 1911. A biological survey of Colorado. N. Amer. Fauna .3.3:1-255. FiNLEY, R. B., Jr., and J. Creasy. 1982. First specimen of the spotted bat {Euderma macukitmti) from Colorado. Great Basin Nat. 42:360. Gillette, D. D., and J. D. Kimbrough. 1970. Chiropte- ran mortality. In Slaughter and Walton, eds.. About bats. Southern Methodist Univ. Press, Dallas. Krutzsch, p. H. 1948. Water-like surfaces attract volant desert animals. Ecology 29:391-392. Parris, L. E. 1973. Caves of Colorado. Pruett Publ. Co., Boulder. 247 pp. NEW GENERIC CONCEPTS IN THE TRITICEAE OF THE INTERMOUNTAIN REGION: KEY AND COMMENTS Mary E. Barkvvorth', Douglas R. Dewey% and Riley J. Atkins' Abstract.— Revision of the perennial genera of North American Triticeae requires recognition of seven new gen- era in the Intermonntain Region: Eltjtrigia, Letjmiis, Psatlnjwstachys, Pseudoroegneria, Thinopijrum, and the hybrid genera X Ehjleijmus and X Pseudelymus. One previously recognized genus, Sitanion, is included in Ehjmiis. Several new combinations are presented to accommodate the taxonomic changes. Ehjmits tmcln/caulus is treated as a wide- spread, polymorphic species with three subspecies in the region: subsp. trachycauhts, subsecundus, and latigliimis. Agropyron dd.systuchyum and A. albicans are treated as conspecific subspecies of Elymus lanceolatus. A key to the genera of the Triticeae occurring in the Intermonntain Region is presented as well as keys and brief descriptions for those genera not included in, or substantially modified from, other regional treatments. Agrostologists have been aware for a long time that traditional North American treat- ments of the Triticeae (e.g., A. Hitchcock 1951, Gould 1968, C. Hitchcock 1969, Hol- mgren and Holmgren 1977) do not reflect the evolutionary relationships within the tribe. Nevertheless, in the absence of any well- documented revision that included a high proportion of the North American species, most North American taxonomists have adopted A. Hitchcock's (1951) treatment, with relatively minor modifications, as the best available. Recently, however, Dewey (1982, 1983a, 1983b) has published a revision of the perennial genera that better reflects the genomic and ecological data available and is consistent with the morphological data. Although written in terms of North American taxa, Dewey's treatment is based on data from the full geographic and tax- onomic range of the tribe. This paper is designed to assist those who wish to use Dewey's generic concepts for plants from the Intermonntain Region. It in- cludes a key to the genera of the tribe in the region and, for those genera not included in, or substantially modified from Holmgren and Holmgren (1977), brief generic descriptions and keys to the infrageneric taxa that we rec- ognize. Readers are referred to the Holm- grens' article for illustrations, detailed de- scriptions of the species, and the complete synonymy. Table 1 summarizes the differ- ences between the treatment presented here and that found in their article. Taxonomic Treatment The genera that are most affected by the revised generic boundaries are Agropyron Gaertn., Elymus L., and Sitanion Raf. Agro- pyron has been restricted to the crested wheatgrasses, the remaining species being as- signed to Elymus, Elytrigia, Pseudoroeg- neria, or Thinopyrum. Several species of Elymus have been placed in the segregate genus Leymus Hochst., but all species of Sita- nion are now included in Elymus. The rea- sons for these and other changes are given in the discussion of individual genera. To assist those not familiar with the subtribal classifi- cation of the Triticeae, the genera and spe- cies within genera are treated alphabetically after the generic key. The intergeneric hybrids are treated after the nonhybrid genera. Readers are advised that such hybrids are relatively rare in na- ture. We include them because they do exist but, in our experience, most plants thought to be hybrids are aberrant forms of good spe- cies. Interspecific hybrids are more common, particularly in disturbed areas. Part of the problem in identifying hybrids in the Tri- ticeae, particularly interspecific hybrids, is that most can backcross to their parents. 'Department of Biology. UMC 45, Utah State University, Logan, Utah 84.322. -Crops Research Laboratory, UMC 63, U.S. Department of .■\griculture. Logan, Utah 84322. 561 562 Great Basin Naturalist Vol. 43, No. 4 Table 1. Synopsis of our revised treatment of the Trititeae occurring in the Interniountain Region compared with that presented in the Intermountain Flora (Hohngren and Hohngren 1977). Revised treatment Intermountain Flora Aegilops cylindrica L. Agropyron cristatwn L. Agropyron clesertortim J. A. Shultes Agrpyron fragile Roth X Elyhordeiiin macounii (Vasey) Barkworth & D. R. Dewey X Ehjleymus aristatits (Merrill) Barkworth & D. R. Dewey Elymus canadensis L. Elymiis elymoides (Raf.) Sweezy Elymus glaucus Buckley Elymus X lumsenii Scribner, pro sp. Elymus lanceolatus (Scribner & J. G. Smith) Gould subsp. lanceolatus subsp. albicans (Scribner & J. G. Smith) Barkworth & D. R. Dewev Elymus multisetiis (J. G. Smith) M. E. Jones Elymus X pseudorepens (Scribner & J. G. Smith) Barkworth & D. R. Dewey Elymus X saundersii Vasey Elymus trachycaulus (Link) Gould ex Shinners subsp. trachycaulus subsp. subsecundus (Link) Barkworth & D. R. Dewey, pro parte subsp. latiglumis (Scribner & J. G. Smith) Barkworth & D. R. Dewey subsp. subsecundus (Link) Barkworth & D. R. Dewey, pro parte Elymus virginicus var. submuticus Hooker Elytrigia intermedia (Host) Nevski subsp. intermedia subsp. barbulata (Schur) A. Love Elytrigia repens (L.) Nevski Elytrigia spicata (Pur.sh) D. R. Dewey Eremopyrum triticeum (Gaertner) Nevski Hordeum brachyantherum Nevski Hordeum depressum (Scribner & Smith) Rydb. Hordeum marinum Hudson subsp. gussonianuni (Pari.) Thell. Hordeum murinum L. subsp. murinum subsp. glaucum (Steudel) Tsvelev .subsp. leporinuni (Link) Arcang. Hordeum pusillum Nutt. Hordeum vulgare L. Leymus cinereus (Scribner & Merrill) A. Love Leymus flavescens (Scribner & Smith) Pilge Leymus salinus (M. E. Jones) A. Love subsp. salinus subsp. stdnionis (C. L. Hitchc.) R. J. Atkins Leymus simplex (Scribner & Williams) D. R. Dewey Leymus triticoides (Buckley) Pilger Pascopyrum smithii (Rydb.) A. Love Aegilops cylindrica L. Agropyron cristatum L. included in A. cristatum L. included in A. cristatum L. X Agrohordeum macounii (Vasey) Lepage X Elysitanion aristatum (Merril) Bowden Elymus canadensis L. Sitanion hystrix (Nutt.) J. G. Smith Elymus glaucus Buckley X Elysitanion hansenii (Scribner) Bowden Agropyron dasystachyum (Hooker) Scribner var. dasystachyum Var. riparium (Scribner & J. G. Smith) Bowden Agropyron albicans Scribner & J. G. Smith var. albicans var. griffithsii (Scribner & J. G. Smith) A. A. Beetle Sitanion jubatum J. G. Smith Agropyron X pseudorepens Scribner & J. G. Smith X Agrositanion saundersii (Vasey) Bowden Agropyron trachycaulum (Link) Malte var. trachycaulum var. glaucum (Pease & Moore) Malte var. latiglumis (Scribner & J. G. Smith) A. A. Beetle var. unilaterale (Cassidy) Malte Elymus virginicus var. submuticus Hooker Agropyron intermedium (Host) Beauv. var. intermedium var. trichophora (Link) Halac Agropyron repens (L.) Beau--'. Agropyron spicatum Pursh Eremopyrum triticeum (Gaertner) Nevski Hordeum brachyantherum Nevski Hordeum depressum (Scribner & Smith) Rydb. Hordeum geniculatum All. Hordeum murinum L. Hordeum glaucum Steudel Hordeum leporinum Link Hordeum pusdlum Nutt. Hordeum vulgare L. Elymus cinereus (Scribner & Merrill) Elymus flavescens Scribner & Smith Elymus salina M. E. Jones Elyvtus ambiguus var. salmonis C. L. Hitchc. Elymus simplex Scribner & Williams Elymus triticoides Buckley Agropyron smithii Rydb. October 1983 Barkworth et al.: Intermountain Triticeae 563 Table 1 continued. Revised treatment Intermountain Flora Psetidoroegneria spicata (Pursh) A. Love X Psi'udehjnnis saxkolus (Scribner & J. C Smith) Barkwortli & D. R. Dewey Psdthtjrostaclnjs jtincea (Fischer) Nevski Secale cereale L. Taeniatheriim caput-medusae (L.) Nevski Thinopyrum ponticum (Podp.) Barkworth & D. R. Dewey Triticwn aestivttm L. Agwpyron spicatum Pursh X Agrositanion saxicohi (Scribner & ]. G. Smith) Bowden Elymus juncetis Fischer Secale cereale L. Taeniatheriim capitt-mediisae (L.) Nevski Agropyron elongatum (Host) Beauv. Triticwn aestivitm L. resulting in a morphological continuum as well as partial restoration of fertility. This problem is not unique to the Triticeae but it is exacerbated by the relatively small number of diagnostic characters available for these grasses. Key to the Genera 1. Spikelets three at a node, each with only one floret; only the central floret fertile (exc. in Hordeum vulgare in which all three florets are fertile) Hordeum — Spikelets not three at a node OR with more than one floret per spikelet (n.b watch out for some forms of Elymus elymoides [ = Sitanion hystrix] in which the second floret of the central spikelet is reduced and the lateral spikelets have only a single, sterile, floret) 2 2(1). Annual or biennial, introduced cereals or weeds 3 — Perennial; native or introduced 7 3(2). Glumes ovate, with three or more (often many) nerves at midlength 4 — Glumes subulate to lanceolate, only one vein evident at midlength 5 4(3). Spikelets sunk in the rachis, the spike therefore very slender, less than 5 mm in diameter; rachis disarticulating at maturity Aegilops — Spikelets not sunk in the rachis, the spike therefore with a larger diameter; rachis not disarticulating at maturity Triticum 5(4). Spikes less than 2.5 cm long; lemmas 5—7.5 mm long Eremopyrum — Spikes more than 4 cm long; lemmas more than 8 mm long 6 6(5). Glumes more than 15 mm long, subulate, united at the base and tapering into a long slender awn; spikelets with only one fertile floret; lemmas glabrous Taeniatherum — Glumes 6-15 mm long, narrowly lanceolate to linear, free to the base, gradu- ally acuminate but not awned; spikelets with two fertile florets; lemmas conspicuously scabrous on the keel Secale 7(2). Spikelets solitary at each node, closely imbricate, often pectinate; internodes short, less than one third the length of the spikelets Agropyron — Spikelets two or more at some nodes OR, if single at all nodes, neither closely imbricate nor pectinate and with internodes about half as long as the spikelets or longer 8 8(7). Glumes 3-10 mm long, very narrow, 1-nerved at midbody length, their keels lying over the sides of the lowest lemmas rather than the midvein; lemmas awnless or with awns up to 7 mm long 9 564 Great Basin Naturalist Vol. 43, No. 4 — Glumes 5-90 mm long, with 2-5 nerves evident at midbody length; keels of the glumes lying opposite the midveins of the lowest lemmas; lemmas often truncate or with awns more than 10 mm long 10 9(8). Rachis disarticulating at maturity; truly cespitose, branching intravaginal, rhizomes never present; old leaf sheaths becoming fibrous Psathyrostachys — Rachis not disarticulating at maturity; often rhizomatous, sometimes shortly so, branching extravaginal; old leaf sheaths not fibrous Leymus 10(8). Plants fertile; anthers well-filled prior to anthesis, dehiscent, usually bleached and falling off after anthesis 11 — Plants sterile; anthers poorly filled prior to anthesis, nondehiscent, retaining their color and usually retained on the plant 13 11(10). Glumes 6-12 mm long; linear-lanceolate to lanceolate, tapering from below midlength into an awn-tip; only one spikelet at most nodes Pascopynim — Glumes varied but if 6-12 mm long either obtuse or tapering only in the distal third; number of spikelets per node 1-4, varying between species 12 12(11). Glumes acute to long awned, never truncate or obtuse; anthers 2-3.5 mm long and plants cespitose or anthers more than 3.5 mm long, plants rhizomatous, and leaf blades with subequal ribs Elymus — Glumes varied, often truncate or obtuse; anthers 4-7 mm long, plants rhizo- matous or cespitose, if both long anthered and rhizomatous [E. repens], leaf blades with 2-3 minor ribs alternating with the major ribs 13 13(12). Plants cespitose; glumes acute-tipped; spikelets only slightly longer than the internodes Pseudoroegneria — Plants rhizomatous or cespitose, if cespitose the glumes truncate to obtuse; spikelets almost twice as long as the internodes 14 14(13). Plants cespitose Thinopyrum — Plants rhizomatous Elytrigia 15(10). Lemmas with divergent awns more than 15 mm long; nodes with only one spikelet X Pseudelymus — Lemmas awnless or with nondiverging awns; nodes with one or two spikelets 15 16(15). Internodes less than 3 mm long; lowest lemmas usually less than 8.5 mm long ... X Elyhordeiim — Internodes more than 3 mm long; lowest lemmas usually 9 mm or longer 16 17(16). Glumes 12-24 mm long X Ehjleymus — Glumes awnless or 25-85 mm long Elymus hybrids A „.; T ploid wheats. His treatment has since been adopted by Morris and Sears (1967) and Bowden (1959) argued that Aegi/ops should ^°"J*^ ^J??^^', . , ^. ^. ^ ., be included in Triticum, primarily because The difficulty with this approach is that, if species of both genera have been involved in ^PP^^^^^ consistently, the tribe has to be re- the evolution of many of such polyploid ^"^'^^ *« ^ ^^"g^^ g^""^ ^^f^"f '^' members wheat species at T. durum Desf. and T. aesti- '^'^ connected by a network of introgressants vum L. He noted also that the International ^nd hybrids. Krause (1898) advocated recog- Code of Botanical Nomenclature requires i^ition of a single genus, but most taxonomists that intergeneric hybrids must have a differ- have rejected his position because it ignores ent generic name from their parents. By in- the differentiation, both morphological and eluding Aegilops in Triticum he obviated the physiological, that has occurred within the need for a new generic name for the poly- tribe. MacKey (1975) pointed out that the October 1983 Barkworth et al.: Intermountain Triticeae 565 combination of annual growth habit and self- fertihzation, such as occurs in Aegilops and Triticum, "stimulates morphological and physiological discontinuity in connection with ecological specialization without the ne- cessity for a simultaneous construction of sterility barriers based on karyological differ- entiation." Of the two genera in question, Aegilops has remained a weedy genus with a relatively narrow ecological amplitude and is generally restricted to poor soils. Triticum, on the other hand, has a much wider ecological amplitude and greater ability to occupy fer- tile land. This, combined with its tendency to produce a larger grain, has led to rapid evo- lution in response to selection pressures exerted in part by human cultivation. Thus, we prefer to treat the two as separate genera both because of their morphological dis- continuity and their different evolutionary potentials. The nomenclatural code requires that a hybrid genus be given a different name from any of its parents, but it does not state what groups of species are to be treated as hybrid genera. A group that has become sufficiently well established that its origins are "ancient history" can be treated as a "normal" genus even if it is known to have originated through hybridization. The species of Triti- cum are such a group. Agropyron Gaertner This genus is now restricted to members of the crested wheatgrass group. Its members can be recognized by the very short inter- nodes of the inflorescence and, in most in- stances, the pectinate arrangement of the spikelets. All our species are more or less ces- pitose, although forms that produce short rhi- zomes exist. Only one genome, the C gen- ome, has been found in Agropyron s.str. Both diploids and polyploids are known. Agropijron s.str. includes about 10 species, all of which are native to Eurasia. Consid- erable controversy exists concerning the ap- propriate taxonomic treatment for the plants found in North America (cf, e.g., Hitchcock 1951, Sarkar 1956, Schulz-Schaeffer et al. 1963, Dewey 1969a, Taylor and McCoy 1973). The species exhibit considerable mor- phological intergradation (cf. Tsvelev 1976), and the problems of identification are exacer- bated by their ability to hybridize when brought into contact (Knowles 1955, Dewey 1969a), as has happened in North America. The genus needs detailed biosystematic study, based on wild populations, a project beyond the scope of this paper. The treat- ment presented here is based in part on Dew- ey's examination of specimens in the Koma- rov Institute (the National Herbarium of the Soviet Union) and discussions with Tsvelev. 2(1). Key to the Species of Agropyron Spikelets diverging from the rachis at an angle of more than 40 degrees; glumes widespread, forming an angle of more than 120 degrees, giving the spike a bristly appearance; spikes at least 8 mm wide A. cristatum Spikelets diverging from the rachis at an angle of less than 350 degrees; glumes appressed; spikes 5-10 mm wide 2 Lemma with an awn 1-2(4) mm long; glumes forming an angle of approximately 60 degrees A. desertorum Lemma without an awn, sometimes mucronate; glumes forming an angle of approximately 45 degrees (not common) A. fragile (= A. sihiricum) Elymus L. Elymus is the largest genus in the Tri- ticeae, but genomically it is very uniform. All of its members are allopolyploids in which two genomes are present, one derived from Pseudoroegneria spicata or a relative thereof. and the other from Hordeum. Almost all plants examined, including all those from the Intermountain Region, are tetraploids (2n = 28). Despite their genomic similarity, species of Elymus fall into two distinct morphological groups. The largest group consists of self- 566 Great Basin Naturalist Vol. 43, No. 4 fertilizing, cespitose species with small an- thers; the other of rhizomatous, outcrossing species, with long anthers. Dewey (1983a) earlier included the latter group in Elytrigia with other rhizomatous, long-anthered, but genomically distinct species; but he now (1983b) includes them in Elymiis, a treatment that better reflects their phylogenetic affi- nities. These rhizomatous species of Ehjmus differ from those of Elytrigia in having glumes that are acute or shortly awned, rather than truncate or long-awned, and leaf blades with no evident alteration of major and minor ribs on the adaxial surface. As in- terpreted here, there is only one such species in the Intermountain Region, Elymus lan- ceolatus [= Agropyron dasystachyum and Agropyron albicans, cf. Table 1.]. The change in epithet is necessary because the com- bination Elymus dasystachys has been used for a European species. Elymus includes two other species that used to be included in Agropyron (£. scribneri and E. trachycaulus), because they have only one spikelet per node. We maintain that the morphological, reproductive, and genomic similarity of these two species to others with a similar genomic composition is more signif- icant than the number of spikelets per node. Elymus elymoides [ = Sitanion hystrix] and E. multisetus [ = S. jubatum] have previously been included in Sitanion (A. Hitchcock 1951, C. Hitchcock 1969, Holmgren and Holmgren 1977), a genus characterized by a readily disarticulating rachis and subulate, long-awned glumes. Genomic studies have shown, however, that the species included in Sitanion are just as closely related to the SH species previously included in Elymus or Agropyron as these species are to each other (Stebbins and Snyder 1956, Stebbins et al. 1946, Stebbins and Vaarama 1954, Brown and Pratt 1960, Dewey 1967, 1969b, Church 1967a, b). The disarticulating rachis, long subulate glumes, and reduced sterile florets constitute a set of adaptations for dispersal in open en- vironments because the segments of the spike are easily blown over the ground. Similar fea- tures are found in one of the forms of the di- morphic species Aegilops speltoides. The other form consists of plants with a non-dis- articulating rachis, short glumes, and more fertile florets. Zohary and Imber (1963) showed that the differences between the two forms are determined by a group of closely linked genes that are normally inherited as a block. No studies have been conducted to de- termine whether the same is true of the char- acteristics used to delimit Sitanion, but Zo- hary and Imber's study lends credence to our conviction that Sitanion does not merit rec- ognition at the generic level. Three hybrid species are included in our interpretation of Elymus, E. X hansenii, E. X pseudorepens, and E. X saundersii. These were previously referred to X Elysitanion, Agropyron, and Agrositanion, respectively. The change in their generic position results from changes in the treatment of their paren- tal taxa. Our treatment of Elymus trachycaulus differs somewhat from that endorsed by Holmgren and Holmgren (1977). The tax- onomy of the slender wheatgrass complex, of which Elymus trachycaulus is a part, is ex- tremely difficult to elucidate. Jozwik (1966) recognized four groups in North America, primarily on the basis of field, hybridization, and herbarium studies. He suggested that many of the members of two of his groups may have been derived by hybridization, one of them comprising plants derived from a va- riety of different hybrid combinations. He described the largest of the other two groups (which corresponds to subsp. trachycaulus in our treatment) as morphologically diverse, probably as a result both of innate genetic plasticity and introgression from other taxa. It has a wide ecological amplitude, growing along stream banks and in forests, meadows, and moist prairies. Geographically it is ex- tremely widespread, extending from Mexico to Alaska and to both the west and east coasts of North America. Subspecies latiglumis corresponds to Joz- wik's other, primarily nonhybrid, group. Its members are more or less restricted to sub- alpine, alpine, and far northern locations, but at lower elevations they tend to intergrade with subsp. trachycaulus, probably in part because of hybridization. Our third subspecies, subsp. suhsecundus, corresponds to Jozwik's second group. This is the group that he believed consisted almost entirely of hybrids. His data indicated that October 1983 Barkworth et al.: Intermountain Triticeae 567 the second parent could be one of several available, it would be impossible to design a taxa, e.g. E. elymoides, E. multisetus, E. completely satisfying treatment for such a glaucus, and H. jubatum. Intermediates be- group because the formal requirements of the tvt'een the members of this subspecies and nomenclatural code cannot perfectly reflect subsp. trachycaulus were numerous. He also the dynamic interactions occurring in a found intermediates with subsp. latiglumis, group such as the slender wheatgrass but these were much less frequent. complex. We admit that our treatment of this com- Our treatment of E. elymoides also differs plex is not altogether satisfying, but it seems from that in Holmgren and Holmgren (1977) the most appropriate treatment considering in that we are not recognizing any in- the data available. Even if more data were fraspecific taxa. Key to Species and Hybrids of Elymus 1. Spikelets 2-7 at a node 2 — Spikelets solitary at each node 7 2(1). Glumes subulate, 1-2-nerved at midlength and with awns more than 20 mm long; rachis disarticulating at maturity 3 — Glumes lanceolate, 2-5-nerved at midlength, if 2-nerved the awns less than 5 mm long; rachis not disarticulating at maturity 5 3(2). Awns not diverging, even at maturity E. X hansenii — Awns widely divergent at maturity 4 4(3). Glumes longitudinally divided into 3 or more narrow sections E. multisetus — Glumes entire or bifid E. elym,oides 5(2). Rachis flexible, spike nodding; glumes with an awn 10-30 mm long E. canadensis — Rachis stiff, spike erect; glumes unawned or short awned 6 6(5). Glumes bowed outward and indurate at the base, the nerves not evident in the indurate portion E. virginicus var. submuticus — Glumes not bowed out, membranous at the base, nerves evident throughout E. glaucus 7(1). Plants rhizomatous; anthers 3-5 mm long 8 — Plants cespitose; anthers 1-3 mm long 10 8(7). Plants sterile, anthers not well filled at anthesis, not dehiscent E. X pseudorepens — Plants fertile, anthers well filled at anthesis, dehiscent (£. lanceolatus) 9 9(8). Lemmas awnless or with an awn-tip less than 5 mm long E. lanceolatus subsp. lanceolatus — Lemmas with a divergent awn 5-12 mm long E. lanceolatus subsp. albicans 10(7). Lemmas awned, the awns widely divergent; culms decumbent, usually less than 35 cm tall E. scribneri — Lemmas unawned or, if awned, then erect or only slightly divergent; culms erect, usually more than 50 cm tall 11 11(10). Glumes l-2(3)-nerved; rachis tending to disarticulate at maturity; plants sterile E. X saundersii — Glumes (3) 5-nerved; rachis not disarticulating at maturity; plants fertile (£. trachycaulus) 12 12(11). Lemma awns 8-24 mm long, erect to divergent . E. trachycaulus subsp. subsecundus — Lemmas awnless or with short, erect, awns less than 5 mm long 13 568 Great Basin Naturalist Vol. 43, No. 4 13(12). Culms erect, 30-130 cm tall; glumes with a narrow hyaline margin E. trachycaulus subsp. trachycaulus — Culms often geniculate or decumbent, less than 55 cm tall; glumes with a broad hyaline margin E. trachycaulus subsp. latiglumis Elymus lanceolatus subsp. albicans (Scribner & J. G. Smith) Barkworth & D. R. Dewey, cotnb. nov.— Basionym: Agro- pyron albicans Scribner & J. G. Smith, USDA Div. Agrost. Bull. 4:32, 1897. Elymus X pseudorepens (Scribner & J. G. Smith) Barkworth & D. R. Dewey, comb, nov.— Basionym: Agropyron pseudorepens Scribner & J. G. Smith, USDA Div. Agrost. Bull. 4:34, 1897, pro sp. Elytrigia Desv. All species of Elytrigia are outcrossing, but in their other characteristics, including their genomic composition, they are very diverse. It is undoubtedly the least satisfactory genus as presently constituted and the one that most needs further study. There are only two species in the Intermountain Region, both of which are introduced. 2(1). Key to the Species of Elytrigia Glumes acute to awn tipped, membranous; rachis only slightly concave adjacent to spikelet E. repens Glumes truncate or mucronate, thick; rachis markedly concave adjacent to the spikelet (£. intermedia) 2 Lemmas glabrous; spikelets 3-8-flowered E. intermedia subsp. intermedia Lemmas hirsute; spikelets 2-3(-6)-flowered E. intermedia subsp. barbulata Hordeum L. The limits of this genus have not been changed but we have adopted the in- frageneric treatment recommended by von Bothmer (pers. comm.), since he has studied the genus in both North and South America as well as Europe. This seems particularly appropriate since the taxa for which von Bothmer's treatment differs from that in Holmgren and Holmgren (1977) are all in- troduced Mediterranean weeds. Moreover, although Holmgren and Holmgren treated the subspecies of H. murinum at the specific level, they noted that the taxa were very closely related and often difficult to dis- tinguish. Thus, the differences between the two treatments are not as great as it may ap- pear. No key is presented since the Holm- gren's key can be used, the only changes needed being nomenclatural. These are in- dicated in Table 1. Leymus Hochst. In our region, the species of this genus can be recognized by their short, subulate glumes that lie over the sides rather than the mid- veins of the lemmas, and by the absence of long awns. The genus includes both rhizo- matous and cespitose, but extravaginally branching, species. Species of Leymus, both here and else- where, tend to grow in alkaline or saline soils. Some are coastal in distribution; others are inland species. The two groups are mor- phologically distinct. Our species, not sur- prisingly, belong to the inland group. Despite the morphological discontinuity between its coastal and inland members, spe- cies of Leymus are genomically similar. They are all allopolyploids based on the J genome, from Psathyrostachys, and the X genome whose origin is unknown. Tetraploids (2n = 28), hexaploids (2n = 42), and octoploids (2n = 56) are known. Thus species of Leymus differ from species of Elymus both in their genomic composition and their tendency to form higher polyploids. In traditional treatments of the tribe, Leymus is included in Elymus since most of its members have more than one spikelet at a October 1983 Barkworth et al.: Intermountain Triticeae 569 node. As indicated above, however, the two genera differ from each other in a number of other morphological characteristics. More- over, L. salinits and L. simplex usually have only one spikelet at most, if not all, nodes. The treatment of L. salinus presented here is based on work by Atkins (1983; Atkins et al., in press). Key to Species of Leymiis 1. Plants strongly rhizomatous, the rhizomes long and slender 2 — Plants cespitose, sometimes with short rhizomes 5 2(1). Lemmas conspicuously hirsute, the hairs 1-2 mm long, not closely appressed to the lemma L. flavescens — Lemmas glabrous to, at most, inconspicuously hirsute with hairs less than 1 mm long 3 3(2). Leaf blades with more than 7 veins, not densely hirsute above the ligule; most nodes with two or more spikelets L. triticoides — Leaf blades with 5-7 prominent veins, densely short-hirsute above the ligule; most nodes with only one spikelet f. L. simplex 5(1). Leaf blades 4-15 mm wide, flat, many nerved; ligules 2-5 mm long; culms more than 1 m tall L. cinereus — Leaf blades 2-4 mm wide when flat, strongly involute, 5-7-nerved; ligules less than 2 mm long; culms less than 1 m tall (L. salinus) 6 6(5). Basal leaf sheaths glabrous; most nodes with only one spikelet L. salinus subsp. salinus — Basal leaf sheaths conspicuously hirsute; most nodes with two or more spikelets L. salinus subsp. salmonis Leym.us salinus subsp. salmonis (C. Hitchc.) Atkins, comb, nov.— Basionym: Elymiis amhiguus var. salmonis C. Hitchc. Univ. Wash. Publ. Biol. 17(1):558, 1969. Holotype: WTU! Pascopyrum A. Love Pascopyrum is a monotypic genus com- prising only P. smithii. This species is an oc- toploid, its probable parents being Elymus lanceokitus and Leymus triticoides (Dewey 1975). Morphologically it is intermediate be- tween its parents. This is particularly evident in the glumes, which are membranous and flat at the base, as it typical for Elymus, but then taper gradually into an acuminate tip resembling the linear lanceolate glumes char- acteristic of Leymus. Holmgren and Holm- gren (1977) recognized two varieties within the species, but we do not consider either merits formal recognition. Psathyrostachys Nevski This genus is comprised of eight species that are native to the steppes and arid re- gions of southeastern Europe. They are all strictly cespitose and have disarticulating rachises and two spikelets at a node. All the species studied so far are diploids based on the J genome. Psathrostachys juncea (Russian wild rye) is the only species to have become established in North America. Pseudoreogneria A. Love All species of Pseudoroegneria are based on a single genome, the S genome. The genus consists of several Eurasian species but only one North American species, P. spicata. Most of its members can be recognized by their rather slender habit and the single spikelets that are only slightly longer than the internodes. 570 Great Basin Naturalist Vol. 43, No. 4 In previous discussions of the tribe, Dewey (1982, 1983a, b) has included these species in Elytrigia in conformity with Tsvelev's (1976) treatment. It is clear, however, that they are genomically distinct and consequently, in keeping with the philosophy guiding this re- vised treatment, should be recognized at the generic level. No new combinations are nec- essary for the Intermountain Region. Thinopyrum A. Love This is a Eurasian genus but one of its members, T. ponticum, has been introduced into North America and occurs along high- ways in the Intermountain Region. We have followed Holub (1973) and Melderis (1980) in adopting the epithet pontica for the plants that Holmgren and Holmgren (1977) referred to Agropyron elongatum. The epithet ehn- gata is now interpreted as referring to a west- em Mediterranean species of relatively small, slender plants, all of which are diploids. Thinopyrum ponticum (Tall Wheatgrass) con- sists of robust decaploid plants that are wide- spread in Eurasia. It has been seeded at scat- tered locations in the Intermountain Region. The new combination is presented here: Thinopyrum, ponticum (Podp.) Barkworth & D. R. Dewey, comb, nov.— Basionym: Triticum elongatum (Host), Gram. Austr. 2:18, 1802). X Elyhordeum Mansf. ex Zizin & Petr. One hybrid between Elymus and Hordeum is established in the Intermoiantain Region, X E. macounii. Its parents are Elymus trachycaulus and Hordeum jubatum (Boyle and Holmgren 1955). In previous treatments it was included in X Agrohordeum macounii. The transfer to X Elyhordeum is made neces- sary by the transfer of Agropyron trachycau- lum to Elymus. X Elyhordeum macounii (Vasey) Barkworth & D. R. Dewey, comb, nov.— Elymus trachycaulus (Link) Gould ex Shinners X Hordeum jubatum L.— Basionym: Elymus macounii Vasey, Grasses U.S. 46, 1883. Macoun, Great Plains of B.C. X Elyleymus Baum One hybrid between Elymus and Leymus occurs in the Intermountain Region, X Ely- leymus aristatus. Dewey and Holmgren (1962) have shown that its parents are prob- ably Elymus elymoides and L. triticoides. Holmgren and Holmgren (1977) referred it to X Elysitanion aristatum. X Elyleymus aristatus (Merrill) Barkworth & D. R. Dewey, comb, nov.— Elymus elymoides (Raf.) Barkworth & Dewey X Leymus triticoides (Buckley) Pil- ger.— Basionym: Elymus aristatus Mer- rill, Rhodora 4:147, 1902. Cusick 2712, "in large clumps, Silver Creek, Harney Co., Oregon." X Pseudelymus Barkworth & D. R. Dewey, gen. hybr. nov. X Pseudelymus Barkworth & D. R. Dewey, gen. hybr. nov.— Pseudoroegneria A. Love X Elymus L. One hybrid between Pseudoroegneria and Elymus has become established in west- ern North America, X P. saxicola. The ge- neric name X Pseudelymus is presented here to accommodate it and other such hybrids that may occur elsewhere. The parents of X P. saxicola are Pseudo- roegneria spicata and Elymus elymoides (Dewey 1964). The plants are usually com- pletely sterile but, being perennial, once they are established at a location, they will persist there. The change in generic name is made necessary by changes in generic boundaries affecting its parents. X Pseudelymus saxicola (Scribner & J. G. Smith) Barkworth & D. R. Dewey, comb, nov.— Pseudoroegneria spicata (Pursh) A. Love X Elymus elymoides (Raf.) Sweezy.— Basionym: Elymus saxicolus Scribner & J. G. Smith, USDA Div. Agrostol. Bull. 18:20, 1899, pro. sp. Discussion Selection of the most appropriate tax- onomic treatment of a polyploid complex is always difficult. This is particularly true October 1983 Barkworth et al.: Intermountain Triticeae 571 when the members of the complex hybridize as readily as do members of the Triticeae. The problem is compounded by the great morphological reduction that characterizes all grasses and the prevalence of convergent evolution. The treatment presented here seeks to reflect as completely as possible all available data. It is therefore a compromise between a strictly genomic interpretation and one based entirely on morphological data. The main advantage of this treatment is that it reflects a higher proportion of the genomic and morphological information available than does the traditional treatment. It is also in closer accord with the systems adopted by Tsvelev (1976) and Tutin et al. (1980). Since both the Soviet Union and Eu- rope have more species of Triticeae than the United States and Canada combined, it is ap- propriate to consider seriously the treatments advocated by taxonomists in those regions. Some of the new genera are not, perhaps, as easy to recognize as the old interpretation of Agropyron and Elymus. On the other hand, numerous herbarium specimens indicate that Leymus salinus and Leymus simplex, which have only one spikelet at most of their nodes, have often been misidentified as species of Agropyron rather than Elymus. Thus, even the traditional treatment was sometimes diffi- cult to apply. The revised genera can be rec- ognized on the basis of their gross morpho- logical characters, although not the same characters as before. We hope that this treat- ment will assist those wishing to become fa- miliar with the new generic concepts. Acknowledgments We thank Drs. A. Love and J. McNeill for their careful review and criticism of the in- itial draft of this manuscript and for the nu- merous discussions concerning the taxonomy of the Triticeae in general. Literature Cited Atkins, R. J. 1983. A taxonomic study of Leymus arn- higuus and L. salinus (Poaceae). Unpublished thesis. Department of Biology, Utah State Univ. Atkins, R. J., M. E. Barkworth, and D. R. Dewey. In press. A taxonomic study of Leymus ambiguus and L. salinus (Poaceae). Syst. Bot. BowDEN, W. M. 1959. The taxonomy and nomenclature of the wheats, barleys, and ryes and their wild relatives. Canadian J. Bot. 37:657-684. Boyle, W. S., and A. H. Holmgren. 1955. A cytogenet- ic study of natural and controlled hybrids be- tween Agropyron trachycaiilum and Hordeum jubatum. Genetics 40:539-545. Brown, W. V., and G. A. Pratt. 1960. Hybridization and introgression in the grass genus Elyjnus. Amer. J. Bot. 47:669-676. Church, G. L. 1967a. Taxonomic and genetic relation- ships of eastern North American species of Elymus with setaceous glumes. Rhodora 69: 121-162. 1967b. Artificial hybrids of Elymus virginicus with E. canadensis, interruptus, riparius, and weigandii. Amer. J. Bot. 43:410-417. Dewey, D. R. 1964. Natural and synthetic hybrids of Agropyron spicatum X Sitanion hystrix. Bull. Torrey Bot. Club 91:396-405. 1967. Synthetic hybrids of Elymus canadensis X Sitanion hystrix. Bot. Gaz. 128:11-16. 1969a. Hybrids between tetraploid and hexaploid crested wheatgrasses. Crop Sci. 9:787-791. 1969b. Synthetic hybrids of Agropyron albicans X A. dasystachyum, Sitanion hystrix, and Elijmiis canadensis. Amer. J. Bot. 56:664-670. 1975. The origin of Agropyron smithii. Amer. J. Bot. 62:524-530. 1982. Genomic and phylogenetic relationships among North American perennial Triticeae. Pages 51-88 in J. R. Estes et al., eds.. Grasses and grasslands, Univ. of Oklahoma Press, Norman. 1983a. New nomenclatural combinations in the North American perennial Triticeae (Gramineae). Brittonia 35:30-33. 1983b. Historical and current taxonomical per- spectives of Agropyron, Elymus, and related gen- era. Crop Sci. 23:637-642. Dewey, D. R., and A. H. Holmgren. 1962. Natural hybrids of Elymus cinereus and Sitanion hystrix. Bull. Torrey Bot. Club 89:217-228. Gould, F. W. 1968. Grass Systematics. New York: McGraw-Hill Book Co. Hitchcock, A. S. 1951. Manual of the grasses of the United States, rev. A. Chase. USDA Misc. Publ. 200. Hitchcock, C. L. 1969. Gramineae. Pages 383-725 in C. L. Hitchcock, A. Cronquist, and M. Ownbey, eds.. Vascular plants of the Pacific Northwest Pt. 1. Univ. of Washington Press, Seattle. Holmgren, A. H., and N. H. Holmgren. 1977. Poaceae. Pages 175-465 in A. Cronquist et al., eds., Inter- mountain flora. Columbia Univ. Press, New York. HoLUB, J. 1973. New names in phanerogamae 2. Folia Geobot. Phytotax. 8:155-179. JozwiK, F. X. 1966. A biosystematic study of the slender wheatgrass complex. Unpublished dissertation. Univ. of Wyoming. Knowles, R. p. 1955. A study of variability in crested wheatgrass. Canadian J. Bot. 33:534-546. Krause, E. H. L. 1898. Florische Notizen II. Graser. Bot. Centrabl. 73:332-343. 572 Great Basin Naturalist Vol. 43, No. 4 Mackey, J. 1975. The boundaries and subdivisions of the genus Triticum. Paper read at Twelfth Int. Bot. Congr., Leningrad. Mackey, J., C. J. Humphries, and T. G. Tutin. 1980. Triticeae. Pages 185-206 in Flora Europaea Vol. 5. University Press, Cambridge. Morris, R., and E. R. Sears. 1967. The cytogenetics of wheat and its relatives. Pages 19-87 in K. S. Qui- senberry and L. P. Reitz, eds.. Wheat and wheat improvement, Amer. Soc. Agron. Inc. Madison, Agron. Ser. 13. Sarkar, R. 1956. Crested wheatgrass complex. Canadian J. Bot. 34:328-345. Schulz-Schaeffer, J., p. W. Allerdice, and G. C. Creel. 1963. Segmental allopolyploidy in tetra- ploid and hexaploid Agropyron species of the crested wheatgrass complex (Section Agropyroii). Crop Sci. 3:525-530. Stebbins, G. L., and L. A. Snyder. 1956. Artificial and natural hybrids in the Gramineae, tribe Hordeae. V. Amer. J. Bot. 43:305-312. Stebbins, G. L., and A. Vaarama. 1954. Artificial and natural hybrids in the Gramineae, tribe Hordeae. VII. Hybrids and allopollyploids between Ehjmtis glancus and Sitanion spp. Genetics 39:378-395. Stebbins, G. L., J. 1. Valencia, and R. M. Valencia. 1946. Artificial and natural hybrids in the Gra- mineae, tribe Hordeae I. Elymiis, Sitanion, and Agropyron. Amer. J. Bot. 33:338-351. Taylor, R. J., and G. A. McCoy. 1973. Proposed origin of tetraploid species of crested wheatgrass based on chromatographic and karyotypic analyses. Amer. J. Bot. 60:576-583. TsvELEV, N. N. 1976. Zlaki SSSR. Nauka, Leningrad. Tutin, T. G., V. H. Heywood, D. H. Valentine, S. M. Walters, and D. H. Webb. 1980. Flora Eu- ropaea. Vol. 5. Monocotyledones. Cambridge Univ. Press, Cambridge, U.K. ZoHARY, D., AND D. Imber. 1963. Genetic dimorphism in fruit types of Aegilops speltoides. Heredity 18: 223-231. REPRODUCTIVE ATTRIBUTES OF SOME ROCKY MOUNTAIN SUBALPINE HERBS IN SUCCESSIONAL CONTEXT David J. Schiinpfi'2 and Robert L. Bayn, ]i}-^ Abstract.— Selected reproductive attributes of herbaceous plant species were compared among three stages of a sere in the subalpine zone of Utah's Wasatch Mountains: herbaceous meadow, aspen grove, spruce-fir forest. No suc- cessional trends in seed size or inferred mode of seed dispersal were detected. We ascribe the deviation of these find- ings from those of most other studies to differences in climate, life-form composition, or community age between our sere and those of other studies. A variety of flower colors were found in the meadow stage, grading into a pre- dominance of white flowers under conifers. Animal vectors of pollen, capable of effecting plant outcrossing, were most abundant in the meadows and an order of magnitude less abundant under aspen. Attempts to develop inductive general- izations about ecological succession have in- cluded studies of reproductive characteristics of plants. Among the earliest were those of Salisbury (1942), who found British woodland species to have heavier seeds than those of species in open vegetation. This was largely due to woody species having heavier seeds than herbs, but forest herbs were also heavier seeded than meadow herbs. Salisbury sug- gested that low light intensity on forest floors favored individuals with stored materials in the seed sufficient to fabricate a greater light-intercepting surface. These results have been generally corroborated for grassland (Hayashi 1976, Werner and Piatt 1976) as well as forest (Opler et al. 1977, Abrahamson 1979) seres. The statistical significance of these increases in seed size during succession has not been established. Marino (1980) found no significant difference in seed size of foredune herbs and shrubs from those under dune forest, but those of the intermediate slack dune stage were significantly smaller. The mode of seed dispersal, as inferred from diaspore morphology, has been sug- gested to change toward a greater frequency of animal dispersal as forest succession pro- ceeds (Dansereau 1957, Dansereau and Lems 1957, Harper et al. 1970, Johnston and Odum 1956, Pijl 1972, Opler et al. 1977). Hayashi (1976) did not detect this trend in a grassland seje. Several of these authors, as well as others, suggest that animal dispersal adapta- tions may be more common among woody species than among herbaceous species. If so, the successional changes in dispersal mode may be due to changes in the proportions of various life forms. Statistical tests of these trends are lacking. Most of the above studies were conducted in temperate deciduous forest or tropical moist forest biomes. Our study examined some plant reproductive attributes in stands along a sere terminating in subalpine co- niferous evergreen forest. Seed sizes, dis- persal modes, flower color frequencies, and abundance of animal pollen vectors most likely to effect floral outcrossing were com- pared among the stages of this sere. Study Area and Methods Field work was conducted in the subalpine zone of the Wasatch Mountains in extreme northern Utah. The presumed sere involves vegetative colonization of well-drained, herb- dominated meadows by clones of quaking as- pen, Populus tremuloides Michx. The climax forest is dominated by subalpine fir, Abies lasiocarpa (Hook.) Nutt., and Engelmann spruce, Picea engelmannii Parry. No other 'Department of Biology and Ecology Center, Utah State University, Logan, Utah 84322. 'Present address: Department of Biology, University of Minnesota, Duluth, Minnesota 55812. 'Present address: Bio-Resources, Inc., P.O. Box 3447, Logan, Utah 84321. 573 574 Great Basin Naturalist Vol. 43, No. 4 woody species are significant community components. For further discussion of com- munity dynamics and a site description, see Schimpfetal. (1980). For each stage of succession, we selected abimdant herbaceous species producing sub- stantial numbers of seed, thus excluding vege- tative remnants of earlier stages. Species were included if they produced at least 100 seeds from a total of 10 or more mother plants within a 0.2-ha rectangular plot during the summers of 1976, 1977, or 1978. There was one such plot for each stage in the ex- ample of the sere known as Big Meadow (Schimpf et al. 1980): meadow, aspen, fir, spruce-fir. The species inclusion criteria proved unworkable for the fir and spruce-fir plots, owing to lower population densities and seed production per plant; these plots were treated as a single 0.4-ha conifer plot to determine species inclusion and for all data analyses. Species which qualified for in- clusion were all counted equally in the analy- ses of reproductive attributes, rather than being weighted by abundance. Total weight was determined for a sample of 100 seeds (or caryopses or achenes) for each species in a stage. Samples had been heated overnight at 100 C after collection to stop respiration. All seeds and caryopses were inspected microscopically, and all achenes sectioned transversely to confirm seed devel- opment before being weighed to the nearest 0.1 mg, in equilibrium with atmospheric hu- midity. Size was expressed as mean mg per seed. Baker (1972) reported that mean seed sizes of a group of plant species are often not normally distributed. The distribution of mean sizes in each stage was tested for nor- mality with the Kolmogorov-Smirnov meth- od (Sokal and Rohlf 1969), using both un- transformed and log-transformed variates. Transformed mean seed sizes were compared among stages by analysis of variance. Pairs of stages were compared for their intrinsic vari- ation in mean sizes by the F-ratio of varian- ces (log-transformed data) (Lewontin 1966). Flower color was recorded in the field for each species. The presumed mode of dis- persal was classified as animal, wind, or other from inspection of diaspore structure. The frequency distributions of flower color and dispersal mode categories were compared among stages by chi-square tests. Chi-square was also employed to test for differences in frequency of animal dispersal modes among different plant life forms. Insects suspected to be capable of effecting floral outcrossing were sorted from the gen- eral collections of insects associated with the herbaceous layer. Suspected pollinators in- cluded all adults in the following families: Syrphidae, Bombyliidae, Colletidae, Halic- tidae, Andrenidae, Megachilidae, Apidae, Sphingidae, Nymphalidae, Lycaenidae. These collections were obtained by D-Vac sampling of from 130 to 310 randomly chosen O.SO-m^ or 0.25-m2 plots that had been quickly cov- ered with an insect-tight cage (Southwood 1978). Samples were taken at regular inter- vals throughout the 1977 and 1978 growing seasons in nearby examples of the succession- al stages similar to those at Big Meadow. Re- sults were expressed as m^ sampled per pol- linator caught. Results The 46 herbaceous taxa studied included 26 species in the meadow plot, 22 taxa in the aspen plot (including two varieties of one species), and 23 species in the conifer plot (Table 1). Mean seed sizes fail to exhibit a normal distribution within a stage, based on the Kolmogorov-Smirnov test. Following log transformation, all three stages show a nor- mal distribution. The mean and standard de- viation of transformed variates is depicted for each stage in Figure 1. None of the F values are significant at the .05 level; the means and variances of seed sizes in the three stages are statistically indistinguishable. No successional trends in community seed size are apparent. There are also no discernible patterns of size change within the set of those species that occupy two or more stages (Table 1), based on sign tests (Sokal and Rohlf 1969). White- flowered species become increasingly fre- quent and the red- and blue-flowered species less frequent through successional time, al- though this trend was not significant at the 0.05 level (Table 2). Modes of dispersal have similar frequencies in all three successional stages (Table 3), suggesting that there is no trend in this attribute. October 1983 ScHiMPF, Bayn: Subalpine Herbs 575 Pollen vector abundance was notably lower during 1977 than during 1978 (Table 4). In both summers, abundance is an order of magnitude greater in the meadows than in the aspen understory, with intermediate val- ues in conifer understory. The most abundant pollinator families in the meadows were Ha- lictidae, Apidae, and Syrphidae, each about Table 1. Reproductive attributes of the herbaceous taxa. Taxon Mean seed size. mg Flower color Dispersal Meadow Aspen Conifer mode 0.166 3.017 0.157 2.513 3.359 0.197 3.324 White Orange Green Other Wind Animal Adnlleu millefolium L. ssp. lanulosa (Nutt.) Piper Agoseris aurantiaca (Hook.) Greene var. aurantiaca Agropijron tmdiijcaulum (Link) Malte var. glaiicwn (Pease & Moore) Malte Agropijwn tmchijcaulum (Link) Malte var. latiglume (Scribn. & Smith) A. A. Beetle Androsace filifomiis Retz. Aquilegia coerulea James var. ochroleuca Hook. Arabis drwnmondii Gray Arnica cordifolia Hook. var. cordi folia Arnica parnji Gray Aster engelnumnii (Eat.) Gray Aster integrifolius Nutt. Bromiis carinatus Hook. & ,\rn. Castilleja miniata Dougl. var. miniata Cknjtonia lanceolata Pursh var. lanceolata Collomia linearis Nutt. Delphinium nuttallianum Pritz. var. nuttallianum Descurainia richardsonii (Sweet) Schulz var. sonnei (Robins.) C.L. Hitchc. Draba stenoloba Ledeb. var. nana (Schulz) C.L. Hitchc. Epilobium brachi/carpnm Presl Epilobitim lactiflorwn Haussln. Erigeron speciosus (Lindl.) D.C. var. macranthus (Nutt.) Cronq. Eriogoniim heracleoides Nutt. Erysimum asperum (Nutt.) D.C. var. purshii Durand Erijthronium grandiflorum Pursh Galium bifolium Wats. Geranium viscosissimum Fisch. & C.A. Meyer var. nervosum C.L. Hitchc. Gi7ifl aggregata (Pursh) Spreng. Hackelia micrantha (Eastw.) J.L. Gentry Hieracium albiflorum Hook. Hieracium scouleri Hook. Hijdrophtjllum capitatum Dougl. var. capitatum Ligusticum filicinum Wats. Lupinus argenteiis Pursh var. rubricaulis (Greene) Welsh Madia glomerata Hook. Osmorhiza chilensis Hook. & Arn. Osmorhiza occidentalis (Nutt.) Torr. Pedicidaris racemosa Dougl. var. alba (Pennell) Cronq. Poa nervosa (Hook.) Vasev var. wheeleri (Vasey) C.L. Hitchc. Polygonum douglasii Green var. douglasii Potentilla arguta Pursh var. convallaria (Rydb.) Th. Wolf Rudbeckia occidentalis Nutt. var. occidentalis Senecio crassidus Gray Senecio serra Hook. Stipa lettennanii Vasey Trisetum spicatutn (L.) Richter Viola nutiallii Pursh var. major Hook. 2.807 3.125 Green Other 0..359 White Other 1.318 White Other 0.255 Pink Wind 1.080 Yellow Wind 1.462 Yellow Wind 3.091 White Wind 1.932 Blue Wind 6.905 Green Animal 0.383 Red Wind 0.740 0.823 0.782 White Other 2.141 Pink Other 0.601 Purple Other 0.206 0.205 Yellow Other 0.084 Yellow Other 0.115 0.125 White Wind 0.802 Pink Wind 0.263 0.294 Blue Wind 2.786 Y'ellow Other 0.719 Yellow Other 5.125 5.992 6.660 Yellow Other 2.427 3.260 3.338 White Animal 11.065 Pink Other 1.369 Red Other 3.621 4.058 Blue Animal 0.430 White Wind 0.814 0.926 Yellow Wind 4.752 White Other 6.501 5.932 White Other 27.706 26.478 Blue Other 2.664 Yellow Other 7.746 4.673 White Animal 12.666 13.953 White Other 1.976 White Other 0.470 0.580 Green Other 2.039 White Other 0.276 Yellow Other 2.070 1.657 Yellow Other 2.633 2.855 Yellow Wind 0.745 Yellow Wind 1.105 Green Animal 0.165 0.172 Green Animal 3.944 3.478 Yellow Animal 576 Great Basin Naturalist Vol. 43, No. 4 10 "O QJ 0) if) D) E 0.5 Q3 02 Oil Meadow N=26 Aspen N=22 F=0.97 (means) F= 1.23 (variances,A/M) F=0.87 (variances, C/ A) Conifer N = 23 Fig. 1. Seed sizes of species in each stage of succes- sion, depicted on log-scale. Dots represent means of log- transformed species means for each stage, with bars ex- tending one standard deviation above and below these. N is the number of species in a stage. None of the F val- ues are statistically significant. 12-15 m^ per individual averaged over the two years. So few pollinators were caught in the forest understories that we cannot recog- nize their numerically dominant families with confidence. Discussion Several explanations can be offered for the lack of a successional increase in seed size (Fig. 1). Unlike almost all seres for which seed size has been studied, ours does not in- clude a pioneer stage. The meadows we stud- ied have apparently been unforested for cen- turies and are being slowly invaded vegeta- tively by forests due to climatic change (Schimpf et al. 1980). Thus, we do not expect the meadows to . be dominated by species with light, highly vagile diaspores to the ex- tent that recently deforested sites often are. Likewise, our study site differs consid- erably from the deciduous forest and tropical moist forest sites of other studies with respect to environmental conditions. Soil at our site dries quickly after snowmelt, and summer rain is far less than potential evaporation. This is especially marked in the meadows, where evaporative potential is more than twice that under the conifer canopy (Schimpf et al. 1980). Because low moisture avail- ability has been correlated with greater seed size, both interspecifically (Baker 1972) and intraspecifically (Schimpf 1977), the dryness of the meadows may offset the dim illumina- tion of the spruce-fir understory as a force se- lecting for larger seeds. Marion (1980) found equally large seeds in the most xeric and most shaded stages of a sere. Some reports of increases in seed size with succession appear to be equivocal. Werner and Piatt (1976) found greater seed sizes of herbaceous Solidago species in climax than in serai ecosystems, but this is confounded by the location of the climax stand in a drier cli- mate than that of the serai stand. The signifi- cance of an interspecific successional increase in herb seed size (Abrahamson 1979) is due to the presence of a single large-seeded climax species; no significant increase can be shown if the sizes are first normalized by log trans- formation. Perhaps the most noteworthy seed size increases during succession are those as- sociated with shifts in life form composition. A number of recent community-level stud- ies of flower color in western North America concur that white flowers are relatively more Table 2. Number of species in each flower color cat- egory in each successional stage. Table 3. Number of species in each dispersal mode Color Meadow Aspen C Dnifer category in each successional stage. category Type of dispersal Animal Wind Meadow 4 7 Asp en 6 5 Red, orange, or pink Blue or purple Yellow 6 4 7 2 3 7 0 1 7 Conifer 5 7 White 6 6 12 Other 15 11 11 X2 = 10.70, .05.25 October 1983 ScHiMPF, Bayn: Subalpine Herbs 577 Table 4. Abundance of pollinators capable of effecting floral outcrossing. Total area D-Vac sampled over the course of the summer and area sampled per pollinator caught is expressed for each successional stage. Meadow Aspen Conifer Year m2 sampled m^ per pollinator m2 sampled m^ per pollinator m2 sampled m^ per pollinator 1977 1978 155 97 5 2 59 .36.5 59 19 3,3.5 .32.5 8 6 frequent under dense forest canopies than in better illuminated layers of vegetation (Baker and Hurd 1968, Daubenmire 1975, Moldenke 1976, Ostler and Harper 1978, del Moral and Standley 1979), but do not take a successional perspective. The similar pattern in our stands (Table 2) leads us to believe that a shift to- ward white-flowered species may be a wide- spread successional trend, but this also needs to be tested with seres including pioneer stages. We resist the temptation to propose functional interpretations of this pattern on the basis of human visual perceptions, which differ from those of pollen vectors, especially in the ultraviolet region (Kevan 1978, Gold- smith 1980). A spruce canopy acts as a neu- tral filter in the visible range, even when sun flecks are not considered (Federer and Tan- ner 1966). Therefore, the potential visibility of various colors (on a relative scale) prob- ably changes minimally along our sere. Though we might expect the conifer under- story to be bathed in radiation somewhat en- riched in ultraviolet relative to visible (Vezina and Boulter 1966), white flowers may be the least UV-reflective (Guldberg and Atsatt 1975); thus the importance of signals in the ultraviolet may not change much dur- ing succession. The similar frequencies of dispersal modes in all stages (Table 3) perhaps simply reflects the lack of change in life form composition in the lower strata of the sere. Shrubs and woody vines, surmised to have high propor- tions of species possessing adaptation for ani- mal dispersal, are commonly thought to be most important in intermediate stages of seres, though we are not aware of any rigor- ous tests of this hypothesis. It is interesting that Thompson and Willson (1978) demon- strated more rapid vertebrate removal of fleshy fruits when experimentally provided at a forest edge than when placed beneath a closed forest canopy, implying that temper- ate frugivores frequent intermediate serai stages, perhaps in response to vegetation physiognomy. The differences among successional stages in abundance of pollinators reported in Table 4 are large, but nonetheless underestimated. Several strong-flying vectors were not sam- pled by the D-Vac technique, but were com- mon to abundant nectar feeders in meadows. These were the sphinx moth Hyles lineata (Fabricius) and the hummingbirds Selas- phorus platycercus (Swainson) and S. rufus (Gmelin), which visited primarily Gilia, Del- phinium, and Geranium flowers. We casually observed essentially no moth activity in the forests, but expect some associated with the bloom of Aquilegia there. Smith (1982) quan- titatively recorded hummingbird feeding in aspen understory but only transient flights through spruce-fir stands. The level of herbaceous productivity im- doubtedly affects the abundance of associated vectors. Lower pollinator densities in 1977 (Table 4) are associated with reduced herb aboveground productivity following an ex- ceptionally dry winter (Schimpf et al. 1980). Within years, differences among stages in vector abundance do not correlate well with differences in herbaceous aboveground phytomass; meadow standing crop is four times that of herbs under conifers, and two to three times that of aspen understory (Reese 1981). Vectors respond, of course, to pollen and nectar rather than to total primary pro- duction, but we lack the requisite data to evaluate floral resource levels. The apparent low density of pollinators in aspen understory awaits elucidation. Acknowledgments We thank James A. MacMahon and Ivan G. Palmblad for comments on an earlier draft, and Linda Finchum, Karin Cowper, 578 Great Basin Naturalist Vol. 43, No. 4 and Avis Hedin for typing the manuscript. This work was supported by NSF Grant DEB 78-05328 to James A. MacMahon. Literature Cited Abrahamson, W. G. 1979. Patterns of resource alloca- tion in wildflower populations of fields and woods. American Journal of Botany 66:71-79. Baker, H. G. 1972. Seed weight in relation to environ- mental conditions in California. Ecology 53:997-1010. Baker, H. G., and P. D. Hurd, Jr. 1968. Intrafloral ecol- ogy. Annual Review of Entomology 13:385-414. Dansereau, p. 1957. Biogeography: an ecological per- spective. Ronald Press, New York. Dansereau, P., and K. Lems. 1957. The grading of dis- persal types in plant communities and their eco- logical significance. Contribution 71, Institut Botanique, Universite de Montreal, Quebec. Daubenmire, R. 1975. An analysis of structural and fimctional characters along a steppe-forest cate- na. Northwest Science 49:122-140. del Moral, R., and L. A. Standley. 1979. Pollination of angiosperms in contrasting coniferous forests. American Journal of Botany 66:26-35. Federer, C. a., and C. B. Tanner. 1966. Spectral distri- bution of light in the forest. Ecology 47:555-560. Goldsmith, T. H. 1980. Hummingbirds see near ul- traviolet light. Science 207:786-788. Guldberg, L. D., and p. R. Atsatt. 1975. Frequency of reflection and absorption of ultraviolet light in flowering plants. American Midland Naturalist 93:35-43. Harper, J. L., P. H. Lovell, and K. G. Moore. 1970. The shapes and sizes of seeds. Annual Review of Ecology and Systematics 1:327-356. Hayashi, I. 1976. Secondary succession of herbaceous communities in Japan. Japanese Journal of Ecolo- gy 27:191-200. Johnston, D. W., and E. P. Odum. 1956. Breeding bird populations in relation to plant succession on the Piedmont of Georgia. Ecology 37:50-62. Kevan, p. G. 1978. Floral coloration, its colorimetric analysis and significance in anthecology. Pages 51-78 in A. J. Richards, ed.. The pollination of flowers by insects. Linnean Society Symposium Series, No. 6, London. Lewontin, R. C. 1966. On the measurement of relative variability. Systematic Zoology 15:141-142. Marino, M. L. 1980. Sand dune succession: a com- parison of plant life history characteristics. Un- published dissertation. Michigan State Univ., East Lansing. Moldenke, a. R. 1976. California pollination ecology and vegetation types. Phytologia 34:.305-361. Opler, p. a., H. G. Baker, and G. W. Frankie. 1977. Recovery of tropical lowland forest ecosystems. Pages 379-421 in J. Cairns, Jr., K. L. Dickson, and E. E. Herricks, eds.. Recovery and restora- tion of damaged ecosystems. University Press of Virginia, Charlottesville. Ostler, W. K., and K. T. Harper. 1978. Floral ecology in relation to plant species diversity in the Wasatch Mountains of LUah and Idaho. Ecology 59:848-861. PijL, L. van der. 1972. Principles of dispersal in higher plants. 2d ed. Springer-Verlag, Berlin. Reese, G. A. 1981. Diversity and production of her- baceous vegetation in a northern Utah subalpine chronosequence. LInpublished thesis. Utah State Univ., Logan. Salisbury, E. J. 1942. The reproductive capacity of plants. G. Bell and Sons, London. ScHiMPF, D. J. 1977. Seed weight of Amaranthus retw- flexus in relation to moisture and length of grow- ing season. Ecology 58:450-453. ScHiMPF, D. J., J. A. Henderson, and J. A. MacMahon. 1980. Some aspects of succession in the spruce-fir forest zone of northern Utah. Great Basin Natu- ralist 40:1-26. Smith, K. G. 1982. Avian resource partitioning along a montane sere. Unpublished dissertation. Utah State LIniv., Logan. SoKAL, R. R., and F. J. RoHLF. 1969. Biometry. W. H. Freeman, San Francisco. SouTHwooD, T. R. E. 1978. Ecological methods, with particular reference to the study of insect popu- lations, 2d ed. Chapman and Hall, London. Thompson, J. N., and M. F. Willson. 1978. Distur- bance and the dispersal of fleshy fruits. Science 200:1161-1163. Vezina, p. E., and D. W. K. Boulter. 1966. The spec- tra! composition of near ultraviolet and visible radiation beneath forest canopies. Canadian Jour- nal of Botany 44:1267-1284. Werner, P. A., and W. J. Platt. 1976. Ecological rela- tionships of co-occurring goldenrods {Solidago: Compositae). American Naturalist 110:959-971. APPLICABILITY OF THE UNIVERSAL SOIL LOSS EQUATION FOR SOUTHEASTERN IDAHO WILDLANDS' Mark E. Jensen- Abstract.— In 1981, 20 sediment-collecting tanks and troughs were installed on range and timbered sites of the Caribou National Forest. Measured erosion losses from the first year of study were contrasted to Universal Soil Loss Equation (USLE) estimates utilizing three different vegetative factors. State of Idaho C factors, National Rangeland C factors, and the Vegetative Management (VM) factors were studied. The erosion estimates of all three USLE tests were significantly different than measured soil losses. All equations overestimated the measured mean soil loss, 0.52 megagrams/ha/yr (0.23 tons/ac/yr), by 33, 3,000, and 2,000 percent, respectively. The soil erodibility factor (K), Rangeland C, and VM showed significant relationships to soil loss. The K and VM factors accounted for 88 percent of the variability in sediment loss in multiple regression models. Erosion equations suitable for use on this study area are presented. Soil scientists are frequently required to provide land managers with estimates of soil erosion rates on specific sites. The ongoing preparation of Land Use Plans for forests managed by the U.S. Forest Service, Inter- mountain Region, has increased the need for realistic approaches to estimating erosion rates. The Universal Soil Loss Equation (USLE) is the dominant method used in mak- ing soil erosion estimates within the region (Wischmeier 1968). However, questions have been raised as to the validity of this equation when applied to wildlands (U.S. Department of Agriculture 1982). The USLE was devel- oped for agricultural lands where overland flow and erosion processes comparable to those described by Horton (1933) are oper- able. Such erosion processes are usually not encountered on wildlands with good vegeta- tive cover and snowmelt runoff. Accordingly, it seems likely that the USLE parameters will require modification for use on wildlands to insure that they will give reasonable erosion estimates. A description of the USLE factors used in this study is presented in Table 1 . The primary objective of this study was to con- trast the actual surface erosion rates of some southeastern Idaho wildlands to estimates de- rived by the USLE. A further objective was to determine which of the USLE parameters showed the strongest relationships to mea- sured soil loss. Such information will improve the usefulness of the USLE on wildlands. Study Area The Caribou National Forest is in south- eastern Idaho, covering an elevational range of 1,490 to 2,930 m. (Fig. 1). The forest lies primarily within the Middle Rocky Mountain physiographic province, with some inclusion of the Basin Range physiographic province (Fenneman 1931). The geology is rather com- plex, ranging from Precambrian metamor- phics in the Bannock and Portneuf Ranges, I DAHO WYOMING Fig. 1. The Caribou National Forest in southeastern Idaho. 'Contributed from U.S. Department of Agriculture, Forest Service, Caribou National Forest, Pocatello, Idaho 83201. 'Soil Scientist, Humboldt National Forest, Elko, Nevada 89801. 579 580 Great Basin Naturalist Vol. 43, No. 4 Table 1. A description of the Universal Soil Loss Equation factors used in this study. The Universal Soil Loss Equation Model is A = RtKLSCP Where: A = The estimated average soil loss per unit area in tons/acre for the time period selected for Rt, usu- ally 1 year. Rt = The rainfall factor, usually expressed in units of the rainfall-erosivity index, EI, and evaluated from an iso-erodent map. K = The soil-erodibility factor, usually expressed in tons/acre/EI units for a specific soil in cultivated continuous fallow tilled up and down slope. Val- ues for K in this study were determined from the soil erodibility nomograph. L = The slope length factor, the ratio of soil loss from the field slope length to that from a 22.1 m length on the same soil, gradient, cover, and management. S = The slope gradient factor, the ratio of soil loss from a given field gradient to that from a 9 per- cent slope with the same soil, cover, and manage- ment. In this study, the L and S factors were computed together, using the topographic factor (LS) nomograph. C = The vegetative factor, the ratio of soil loss from land managed under specified conditions to that from the fallow condition on which the factor K is evaluated. Three methods for determining this factor were studied. They are: (1) State C factors that are determined by the Rt values for a site; (2) National Rangeland C factors determined by the canopy cover, vegetation type, and ground cover on a site, and (3) Vegetative Management factors determined by the canopy cover, ground cover, and percent of bare ground with fine roots on a site. P = The erosion control factor, not usually ap- plied to wildlands. Notes (a) These factors take on dimensionless values when computing A. (b) Source = Warrington, 1980. Jurassic-Triassic sedimentaries in the Bear River and Webster Ranges, to Cretaceous sedimentaries in the Caribou Range. Bailey (1980) has classified the vegetation of the forest as belonging to the Rocky Mountain Forest Province— Douglas Fir For- est section and the Intermountain Sagebrush Province— Sagebrush- Wheatgrass section. The climate is a semiarid steppe regime with a wide range in mean annual precipitation. The lower elevations receive 330 mm of pre- cipitation per year, and higher elevations commonly experience 1,524 mm annual pre- cipitation. Approximately 60 percent of the precipitation is in the form of snow. Most soil erosion is observed to occur during the spring snowmelt period. Methods In the summer of 1981, 20 erosion plots were installed on the Caribou National For- est. Plots were equipped with erosion tanks and troughs to catch surface erosion losses in- duced by soil creep and sheet erosion. The data presented is from the 1981-1982 erosion year. The erosion plot construction tech- niques used were comparable to those em- ployed in Montana by Packer and Williams (1976). Erosion plots were .015 hectare in size, with dimensions of 2.4 by 10.1 m. The long axis of each plot was oriented up and down slope. Plots were constructed from 2.5 by 15.2 cm cedar boards on the top and sides. A metal trough, 15.2 cm deep by 25.4 cm wide, was placed on the downhill side to catch sediment. Water and sediment collect- ed from the plots were stored in sealed 907 liter tanks. Tanks were connected to the troughs by 15.2 cm diameter, steel reinforced hose. Sediment was removed from the troughs in early summer after the spring snowmelt. Minimal soil erosion occurred dur- ing the summer months. The accumulated sediment was oven dried to determine ero- sion loss weights. The USLE factors were computed using the procedures outlined in the WRENS doc- trine (Warrington 1980). Rainfall factor val- ues (Rt) were taken from an iso-erodent map developed for Idaho by the Soil Conservation Service (1977). This factor is a water drop impact indicator, with only a small com- ponent for runoff. The dominant erosion agent operable on the study sites, however, is snowmelt runoff. Adequate factors for this erosion parameter have not been developed, which necessitated the use of the Rt factor in this study. Length and percent slope at each site were used to determine the LS factors. Soil profile descriptions and lab analyses of the A horizon were made for each site. Soil organic matter was determined by the Walk- ley-Black Method, and particle size analysis was determined by the Hydrometer Method (Black 1965). Soil erodibility factors (K) were determined by the soil erodibility nomograph October 1983 Jensen: Universal Soil Loss Equation 581 (Warrington 1980). Soil classification fol- lowed procedures outlined in Soil Taxonomy (Soil Conservation Service 1975). The cropping management factor (C) is important in estimating erosion on wildlands (Dissmeyer 1980). Three different methods for determining the vegetative factor were tested in this study. They are: (1) the Vegeta- tive Management factor— VM (Warrington 1980), (2) the National C factor for range- lands-Range C (USD A 1977), and (3) state- developed C factors for Idaho— State C (Soil Conservation Service 1977). The vegetative information collected to compute these fac- tors utilized the Range Site Analysis pro- cedures of the Forest Service, Intermountain Region (U.S. Department of Agriculture 1969). The statistical methods employed fol- lowed Zar (1974). Sites were selected to sample over a wide range in vegetative and soil conditions. Results Site descriptions for the erosion plots are presented in Table 2. Soils of the order Molli- sols and sage-grass vegetative types (i.e., Ar- temisia vaseyana-Agropyron spicatum, A. vaseyana-Stipa comata, and A. vase- yana-Symphoricarpos oreophilus-Agropyron spicatum habitat types) were dominant (Hironaka 1981). Table 2. Site descriptions of the erosion plots. Site number Soil classification Vegetative type Elevation (m) Aspect Percent slope 1 Loamy skeletal, mixed family of Sage-Grass 2,620 the Typic Cryoborolls 2 Fine loamy, mixed family of the Mountainbrush 1,950 Argic Cryoborolls 3 Fine loamy, mixed family of the Sage-Grass 1,980 Argic Cryoborolls 4 Loamy skeletal, mixed family of Sage-Grass 1,950 the Argic Cryoborolls 5 Fine loamy, mixed family of the Sage-Grass 1,830 Cryic Pachic Paleborolls 6 Fine loamy, mixed family of the Sage-Grass 1,800 Argic Cryoborolls 7 Loamy skeletal, mixed family of Sage-Grass 2,620 the Argic Cryoborolls 8 Loamy skeletal, mixed, mesic Sage-Grass 1,830 family of the Typic Argixerolls 9 Fine loamy, mixed family of the Aspen 2,100 Argic Pachic Cryoborolls 10 Fine loamy, mixed, mesic family of Sage-Grass 1,650 the Typic Argixerolls 11 Coarse loamy, mixed, mesic family Juniper-Forb 1,610 of the Typic Xerorthents 12 Fine loamy, mixed family of the Sage-Grass 2,130 Argic Cryoborolls 13 Loamy skeletal, mixed family of Mountainbrush 2,070 the Typic Cryoborolls 14 Loamy skeletal, mixed family of Fir-Pinegrass 2,350 the Typic Cryorthents 15 Fine loamy, mixed family of the Fir-Pinegrass 2,200 Typic Cryorthents 16 Loamy skeletal, mixed family of Sage-Grass 1,950 the Argic Cryoborolls 17 Fine loamy, mixed family of the Sage-Grass 2,130 Typic Cryoboralfs 18 Loamy skeletal, mixed family of Sage-Grass 1,650 the Argic Cryoborolls 19 Fine loamy, mixed family of the Pine-Pinegrass 2,040 Mollic Cryoboralfs 20 Fine, mixed family of the Argic Sage-Grass 1,950 Cryoborolls w 15 E 50 W 60 s 60 NW 30 NW 21 S 35 w 25 E 20 W 45 s 23 SE 40 S 50 w 30 N 35 SE 56 SE 42 SW 60 N 35 S 30 582 Great Basin Naturalist Vol. 43, No. 4 Table 3. The Universal Soil Equation predictions for the study plots" C Factor used Mean Standard deviation Minimum Maximum n State C Range C VM 0.72 (0.32) 16.02(7.15) 11.51 (5.14) 0.52 (0.23) 0.54 (0.24) 35..39 (15.8) 31.81 (14.2) 1.37 (0.61) 0.02 (0.01) 0.76 (0..34) 0.20 (0.09) 0.02 (0.01) 1.86 (0.83) 153.6 (68.6) 138.4 (61.8) 5.8 (2.6) 20 20 20 Measured rate 20 "The first number provided is in units of megagrams/ha/yr. The second number provided is in units of tons/ac/yr. Note; The erosion predictions derived by all three C factor methods were found to be significantly different than the measured erosion losses by use of a Wilcoxon Paired Sample Test at the 95 percent confidence level. The USLE was tested for each site using the three vegetative factors (Table 3). Ero- sion loss estimates from all three USLE tests were significantly different from measured losses as determined by the Wilcoxon paired sample test. The USLE that utilized the State C factor provided the most reasonable esti- mates; it overestimated the mean erosion loss of the sites by 33 percent. USLE predictions that utilized the VM and Range C factors overestimated the mean loss by 2,L35 and 3,010 percent, respectively. These factors also yielded high standard deviations for mean losses and large ranges in predicted erosion rates. To determine how an improvement in the accuracy of the USLE might be made, simple linear regression analysis was performed on the data (Table 4). The percent variability in measured soil loss explained by the USLE factors were 80 percent for VM, 51 percent Table 4. Linear regression relationships between soil loss (A) and the USLE and site variables. Line equation r- 0.002 0. 27 0. 12 0. 02 0. 51 0. 80 0. 17 0. 23 0. 09 0. 17 0. 08 (1) ' Slopes of these equations found to be significantly different than 0 by use of a T test at the 95% confidence level. (2) Sample size = 20 in all cases. (3) A is in units of tons/ac/yr. The product A may be multiplied by 2.24 to obtain units of megagrams/ha/yr. USLE variables Rt A = = 0.01 + 0.0033 Rt K a° = -0.48 + 4.12 K LS a : = -0.19 -H 0.05 LS State C A : = 0.37 -1- 18.85 S-C Range C A" = - 0.21 -1- 4.02 R-C VM A" = -0.19 + 6.38 VM Site variables % Canopy cover A" = 0.67 - 0.0096 C % Vegetation 1& Utter A" = 0.83 - 0.0105 VL % Bare ground A : = -0.06 + 0.0114 BG % Pavement A° = 0.04 + 0.0146 P % Rock A = 0.06 + 0.0.351 R Production A = 0.58 - 0.00018 Prod. for Range C, 27 percent for K, 12 percent for LS, and 0 for Rt. The K, VM, and Range C factors showed significant linear relationships to the measured soil loss on the erosion plots. Of the site factors studied, percent canopy cover and percent vegetation plus litter gave significant negative correlation to soil loss. Percent pavement (i.e., rocks less than 1.9 cm in diameter) had a positive correlation to soil loss. Production, percent bare ground, and percent rock on the sites did not show signifi- cant linear relationships to measured soil loss. Table 5 shows Pearson Correlation Coefficients for two soil variables and the USLE factors. Percent clay has a strong posi- tive correlation to soil loss (A) and soil erod- ibility (K). The organic matter content in the soil showed a strong negative correlation to these factors. Since certain factors of the USLE were not found to have significant relationships to soil loss, equations using cor- related variables were developed (Table 6). Stepwise multiple regression analyses in- dicate that 80 percent of the variability in soil erosion loss is attributable to the VM fac- tor of a site. Considering the K factor with VM accounts for 88 percent of the variability in soil erosion loss. Adding the Rt and LS fac- tors does little toward improving predictions. This relationship is important since the VM Table 5. Pearson correlation coefficients between the USLE factors and soil clay content and organic matter. USLE factor % Clay in the A horizon % Organic matter in the A horizon Rt K LS State C Range C VM Soil loss (A) 0.15 0.65 0.39 -0.45 0.74 0.88 0.88 0.03 -0.57 -0..32 0.14 -0.66 -0.73 -0.70 Sample size = 20 in all cases. October 1983 Jensen: Universal Soil Loss Equation 583 and K factors can be easily determined through soil profile description and relatively simple vegetative analyses. The Rt factor is variable over wildlands of the Intermountain Region and will probably never be quantified for snowmelt situations. The LS factor also presents a problem for field determinations. An absence of uniformity and the benchy na- ture of slopes within the region make it diffi- cult to determine a site's contributing slope length and steepness. An accurate assessment of these variables is needed to derive the LS factor. Actual values for soil loss from plots were used to test the accuracy of different USLE formulations (Table 7). USLE estimates that utilized the State C factor showed a poor correlation with measured soil loss. However, USLE estimates that used the VM and Range C factors showed a high correlation with measured losses; yet they overestimated ac- tual rates. The new equations derived in this paper can be used to scale down USLE esti- mates when the designated vegetative factors are used to predict erosion on western wildlands. Discussion Erosion estimates generated by the USLE were not representative of actual soil losses on erosion plots. The three equations tested significantly overestimated erosion as shown by actual field measurements. This is con- sistent with the findings of Patric (1982) in his review of erosion research on forested lands. Patric suggests that the USLE tended to overestimate erosion losses on forested sites if limitations of the equation on such lands are not considered. Patric also points out that sediment yields of no more than 0.56 megagrams per ha per year provide an index Table 6. Stepwise multiple regression relationships between soil erosion loss (A) and the VM and K factors. A (tons/ac/yr) = - 0.19 + 6.4 X VM factor; r^ = 0.80, standard error = 0.28, n = 20 A (tons/ac/yr) = - 0.55 -I- 5.8 X VM factor + 2.4 X K factor; r- = 0.88, standard error = 0.23, n = 20 Note: (1) The inclusion of the Rt and LS factor increase the H value to 0.89. (2) The product A may be multiplied by 2.24 to obtain units of megagrams/ha/yr. of soil loss from relatively undisturbed forest watersheds. The mean erosion loss on plots considered in this study (i.e., 0.52 megagrams per ha per year) suggests that erosion rates on the Caribou National Forest are comparable to those on other wildlands. The Rt and LS factors present problems when using the USLE to estimate soil losses. These factors showed no significant relation- ship to measured soil losses in this study. This suggests that Rt and LS factors contribute little when the USLE is applied to western wildlands. More information is needed con- cerning the relationships these factors have to determining soil erosion losses on wild- lands with snowmelt runoff. The K factor showed a significant linear relationship to measured soil losses in this study. Laflen (1982) raised questions about the quality of the estimate that the K factor provides for use on wildlands, because the soils of such areas differ from agricultural soils. Steep slopes, high rock fragment con- tent, and high organic matter content of wildland soils contributed to differences in soil erodibility not addressed by Wischmeir (1969) in his early efforts to develop the K factor concept. The correlation between soil loss and the K factor can be improved with a thorough understanding of soil variables. Per- cent clay and organic matter in the A hori- zon were correlated with soil losses and most of the USLE factors considered in this study. Future applications of the USLE to inter- mountain wildlands should address these soil factors. USLE predictions, using the three different vegetative factors, gave erosion estimates higher than observed rates. Equations pre- sented in Table 7 offer a means of reducing estimates to more reasonable levels. Equa- tions that utilize the VM and the K factors (Table 6) provide the land manager with a Table 7. Regression equations correlating USLE esti- mates with measured soil losses when three different vegetative factors are used. Vegetative factor used Correction equation State C y = 0.02 + 0.62 x; r' = 0.06, n = 20 Range C y = -0.05 + 0.04 x; r' = 0.97, n = 20 VM y = 0.002 + 0.04 x; r- = 0.99, n = 20 Note: (1) y = measured soil loss (tons/ac/yr). (2) X = USLE estimated soil loss (tons/ac/yr). 584 Great Basin Naturalist Vol. 43, No. 4 simple approach for predicting erosion loss. These equations are effective because the two USLE factors that showed the greatest sensitivity in predicting soil loss are used in the construction. Conclusions The information collected during the first year of this study will assist those who use the USLE for predicting soil erosion on wild- lands. The results presented will be refined as the study continues. Further research is needed to quantify the relationships between USLE factors and soil erosion on wildlands in the Intermovmtain Region. Specifically, more work should be directed toward developing Rt factors for snowmelt runoff situations. The VM factor offers an effective means for pre- dicting soil erosion. It is particularly useful to the land manager since it allows for the test- ing of different management objectives against their effects on soil loss. Acknowledgments I thank Norm Bare for his assistance in this work. I also wish to thank the U.S. Depart- ment of Agriculture Intermountain Forest Experiment Station, Logan, Utah, for the ero- sion tanks and troughs used in this study. Literature Cited Bailey, R. G. 1980. Description of the ecoregions of the United States. Misc. Pub. No. 139L U.S. Depart- ment of Agriculture, Forest Service. Washington, D.C. 77 pp. Black, C. A. 1965. Methods of soil analysis. Agronomy Series 9. American Society of Agronomy. Madi- son, Wisconsin. 1569 pp. DissMEYER, G. E., A.ND G. R. FosTER. 1980. A guide for predicting sheet and rill erosion on forested land. Tech. Pub. SA-TPll. U.S. Department of Agri- culture, Forest Service, Southeastern Area. At- lanta, Georgia. 40 pp. Fenneman, N. M. 1931. Physiography of western United States. McGraw-Hill, New York. 5.34 pp. HoRTo.N, R. E. 19.33. The role of infiltration in the hy- drologic cycle. Trans. .\mer. Cieophvs. Un. 14:446-460. HiRONAKA, M., AND M. A. FosBERC. 1981. Nouforest habitat tvpe workshop— two. Agricultural Experi- ment Station. Univ. of Idaho. Moscow, Idaho. 87 pp. Laflen, J. M. 1982. Special problems of the USLE: soil erodibility (K). Pages 63-73 in Proc. of the work- shop on estimating erosion and sediment yield on rangelands. Agricultiual Reviews and Manuals, Western Series 26. U.S. Department of .Agricul- ture. Oakland, California. Packer, P. E., and B. D. Williams. 1976. Logging and prescribed burning effects on the hvdrologic and soil stability behavior of Larch /Douglas Fir For- ests in the Northern Rockv Mountains. Pages 465-479 in Proc. Montana Tall Timbers fire ecol- ogy conference symposium 14. Tall Timbers Re- search Station. Tallahassee, Florida. Patric, J. H. 1982. A perspective on soil loss from forest- ed lands. National Bulletin 190-2-18. U.S. Depart- ment of Agriculture, Soil Conservation Service. Washington, D.C. 16 pp. Soil Conservation Service. 1975. Soil taxonomy. Agri- cidtural Handbook 436. U.S. Department of Agri- culture, Soil Conservation Service. Washington, D.C. 743 pp. 1977. Erosion inventory instructions for the PSV and Point Data collection for the state of Idaho. U.S. Department of Agriculture, Soil Con- servation Service. Washington, D.C. 80 pp. U.S. Department of Agriculture. 1969. Range envi- ronmental analysis handbook. U.S. Department of Agriculture, Forest Service. Intermoimtain Re- gion. Ogden, Utah. 1977. Procedure for computing sheet and rill ero- sion on project areas. Tech. Release 41 (Rev. 2). L'.S. Department of .Agriculture, Soil Con- servation Service. Washington, D.C. 17 pp. U.S. Department of Agriculture, Agricultural Research Service. 1982. Proceedings of the workshop on estimating erosion and sediment yield on rangelands. Agricultural Reviews and Manuals. Western Series 26, U.S. Department of Agriculture, Oakland, California. 228 pp. Warrington, G. E. 1980. Surface erosion. Chapter 4 in An approach to water resources evaluation of non-point silviciiltural sources. EPA-6001 i-80- 012. U.S. Environmental Protection .Agency. Athens, Georgia. 861 pp. WiscHMEiER, W. H., AND D. D. Smith. 1968. A universal soil-loss equation to guide conservation farm planning. Trans. Int. Congr. Soil Sci. 1:418-425. WiscHMEiER, W. H., AND J. V. Mannering. 1969. Rela- tion of soil properties to its erodibility. Soil Sci. Soc. Am. Proc. 33:131-137. Zar, J. R. 1974. Biostatistical analysis. Prentice-Hall Inc., Englewood Cliffs, New Jersey. 592 pp. WINTER STONEFLIES (PLECOPTERA) OF NEW MEXICO Gerald Z. Jacobi' and Richard W. Baumann- Abstract.— Twenty-two species of winter emerging Plecoptera were collected in New Mexico from 1979 to 1982. Distributional records are given for 13 that are new state records, including 2 new species, and 9 previously reported species. Winter stoneflies, usually defined, mean species in the families Capniidae (Nebeker and Gaufin 1968) and Nemouridae (Baumann et al. 1977). For this study, we have ex- panded the list to include additional cold lot- ic species (Baumann 1979) in the Taeniop- terygidae and Perlodidae. These emerged on snow or ice, or prior to peak spring runoff when air and water temperatures were below 12 C and 8 C, respectively. New state records, including two pre- viously undescribed species, follow: Capnia barbata Prison, C. coloradensis Claassen, C. vernalis Newport, C. wanica Prison, Iso- capnia vedderensis (Ricker), Mesocapnia arizonensis (Baumann & Gaufin), M. iverneri (Baumann & Gaufin), Utacapnia logana (Nebeker & Gaufin), Utacapnia poda (Nebe- ker & Gaufin), Doddsia occidentalis (Banks), Taenionema pacificum (Banks), Taenionema sp. A, and Taeniopteryx sp. A. Previously recorded species (Stewart et al. 1974, Stark et al. 1975, and Baumann et al. 1977) are: Capnia confusa Claassen, C. graci- laria Claassen, C. fibula Claassen, Eu- capnopsis brevicauda (Claassen), Mesocapnia frisoni (Baumann and Gaufin), Prostoia besa- metsa (Ricker), Zapada cinctipes (Banks), Z. haysi (Ricker), and Skwala parallela (Prison). In addition to distributional data for new state records, recent distributional data are given for previously reported species. All specimens were collected by G. Z. Jacobi un- less otherwise noted. Capnia barbata Frison Capnia barbata Frison, 1944. This species was listed as being restricted to the Southern Rocky Mountain Zone (Nebeker and Gaufin 1967). It had been col- lected in Arizona and Colorado but not in New Mexico (Baumann et al. 1977). Here it is reported from seven counties in New Mexico, which include the northern Sangre de Cristo, central Manzano, south central Sacramento, and southwestern Black and Mogollon moun- tain ranges. Grant Co., Cherry Creek, Pinos Altos, 2,012 m, 6-XI-80, 3 $ (dried); Little Cherry Creek, Hwy 255, 2,012 m, 25-III-81, 3 ? ; Sapello River, Hwy 15, 1,767 m, 25-III-81, G. Z. J. and L. R. Smolka, 50 ? . Lincoln Co., Rio Bonito, Mill Creek Picnic Area, 2,164 m, 14-III-80, 2$ 29 ? ; Rio Ruidoso, 2,188 m, 14-III-80, 1 ? ; Eagle Creek, Hwy 127, 2,179 m, 14-III-80, 2 ? ; Eagle Creek, Hwy 117, 2,164 m, 14-III-80, IS 8?; Nogal Creek, Nogal, 1,975 m, 14-III-80, 1 ? . Three Rivers, Three Rivers Cmpgd., 1,859 m, 7-III-82, 1$ 1 ? . Rio Arriba Co., Canjilon Creek, north of Ghost Ranch, 2,102 m, 22-III-82, 1 ,5 ; Brazos Riv- er, Hwy 84-64 bridge, 2,256 m, 22-III-82, G. Z. J. and L. R. Smolka, 6$ 4 $ . San Miguel Co., Dalton Creek Canyon, 2,195 m, 28-11-79 reared to 2-IV-79, 1$ 2 ? ; 28-11-79, 3 ^ 11 ? 5n; 3-III-80, 26 -p :3 ■rH rO to X3 M U u (U o X OJ l^ l-{ OJ £ d) Mh 14 13 12 11 10 Fig. 10. A hypothetical phylogeny of the species of Micwrhopala. Filled squares represent ancestral characters; open squares represent derived characters. Numbers refer to characters in Table 1. its narrow form and in its parallel-sided pro- thorax. However, it is easily distinguished from that genus by antennal segments 8-11 that are fused, appearing as a single segment. Within the genus Micwrhopala this species most closely resembles M. excavata, from which it differs by the more slender form and the parallel-sided prothorax. Male.— Length 3.4-4.5 mm, 2.3-2.9 times as long as wide; color black or less commonly metallic blue. Head distinctly reticulate dorsally, not or indistinctly reticulate laterally and ventrally; frons arcuately, transversely angled or nar- rowly rounded below antennae; mesal im- pression of vertex margined laterally by a contiguous row of deep punctures; eye sepa- rated from oral fossa by a distance less than the width of antennal segment 3, margined behind by contiguous, confused punctures. Antennae black, sometimes with a metallic blue tint. Pronotum 0.7-0.8 times as long as wide, 0.7-0.9 times as wide as elytra at humeri, not or but slightly narrowed anteriorly; lateral margins appearing arcuate, sinuate, or bi- sinuate in lateral aspect; anterior margin with a small, thin, mesal piece of cuticle that October 1983 Clark: Revision of Microrhopala 617 does not extend laterally; minute reticulation distinct; punctures deep, mostly separated by much less than the diameter of a puncture. Elytra 1.7-2.2 times as long as wide, usual- ly parallel sided; punctures separated by less than the diameter of a puncture, arranged in regular rows; striae 5 and 6 with apical punc- tures often larger than basal punctures; cos- tae variable, well developed to completely absent; interstriae 9 undulate to slightly serrate. Hind femora usually wider than middle femora. Female.— Externally similar to male but averaging larger, 3.8-4.9 mm long. Variation.— The slenderness of the body, the extent of metallic coloration, and the de- gree to which the interstriae are elevated are all variable. Although no geographic trends are apparent, they may be discovered after more specimens are collected. Distrirution.— Florida to North Carolina (Fig. 5). USA: Florida: Marion Co., Polk Co., Putnam Co., Seminole Co., Sumpter Co., Volu.sia Co. Georgia: Chari- ton Co. North Carolina: Moore Co. Biology.— This species feeds on Pityopsis graminifolia from April to August. It has also been reported from Lupinus diffusus, which is a very unusual and perhaps erroneous record. Notes.— The above treatment was based on the holotype of M. floridana and on 25 other specimens. Acknowledgments Appreciation is extended to the following people and institutions for their kind assis- tance and loans of specimens: Donald Azuma, Academy of Natural Sciences of Philadelphia; Nicole Berti, Museum National d'Histoire Naturelle, Paris; Robert L. Blinn, University of Missouri— Columbia; Lee H. Herman, American Museum of Natural His- tory; Charles L. Hogue, Los Angeles County Museum of Natural History; David H. Kavanaugh, California Academy of Sciences; L. L. Pechuman, Cornell University; Laurent LeSage, Canadian National Collection; Ole Martin, Zoologisk Museum, Copenhagen; Al- fred F. Newton, Jr., Museum of Comparative Zoology; Carl A. Olson, University of Ari- zona; R. D. Pope, British Museum (Natural History); Edward G. Riley, Louisiana State University; Hans Silfverberg, Universitets Zoologiska Museum, Helsinki; Charles A. Triplehorn, Ohio State University; J. Reese Voshell, Jr., Virginia Polytechnic Institute and State University; Larry E. Watrous, Field Museum of Natural History; and Rich- ard E. White, Systematic Entomology Labo- ratory, USDA. Special thanks is given to the Department of Zoology, Brigham Young Uni- versity, and particularly to Dr. Stephen L. Wood, for their support and encouragement. Literature Cited Baly, J. S. 1864. Descriptions of genera and species of Hispinae. Ann. Mag. Nat. Hist. (3)14:261-271. 1885. Hispidae. Biologia Centrali-Americana 6(2): 1-124. Barber, H. S., and J. C. Bridwell. 1940. Dejean Cata- loge names (Coleoptera). Bull. Brooklyn Ent. Soc. 35:1-12. CouPER, W. 1865. Descriptions of new species of Cana- dian Coleoptera. Canadian Nat. and Geol. 2: 60-63. Crotch, G. R. 1873. Materials for the study of the Phytophaga of the United States. Proc. Acad. Nat. Sci. Philadelphia 25:19-83. Dejean, P. F. M. A. 1837. Catalogue des Coleopteres, ed. 3. Paris. Douglass, J. R. 1929. Coleoptera of Kansas. J. Kansas Ent. Soc. 2:1-15, 26-.38. Fabricius, J. C. 1798. Supplementum Entomologia Sys- tematica. Hafniae. Systema eleutheratorum, vol. 2, 687 pp. Kiliae. Gemmincer, M., and E. Harold. 1876. Catalogus co- leopterorum hucusque descriptorum synony- micus et systematicus, vol. 12. Munich. Hendrickson, G. O. 1930. Biologic notes on Micro- rhopala vittata Fabr. Canadian Ent. 62:98-99. Horn, G. H. 1883. Miscellaneous notes and short studies of North American Coleoptera. Trans. Amer. Ent. Soc. 10:269-312. LeConte, J. L. 1859a. Catalogue of the Coleoptera of Fort Tejon, California. Proc. Acad. Nat. Sci. Philadelphia 11:69-90. 1859b. Coleoptera of Kansas and eastern New Mexico. Smithsonian Contribution to Knowledge 11:1-58. McCauley, R. H. 1938. A revision of the genus Micro- rhopala in North America, north of Mexico. Bull. Brooklyn Ent. Soc. .33:145-169. Mannerheim, C. G. v. 1843. Beitrag zur Kaferfauna der Aleutischen Inseln, der Insel Sitka und Neu-Cali- forniens. Soc. Imp. Nat. Moscou (Moskov. Obshch. Isp. Prirody Otd. Biol. Biul.) 16:175-314. Melsheimer, F. V. 1846. Descriptions of new species of Coleoptera of the United States. Proc. Acad. Nat. Sci. Philadelphia 3:158-181. 1853. Catalogue of the described Coleoptera of the United States. Smithsonian Institution, Washington. 618 Great Basin Naturalist Vol. 43, No. 4 Newman, E. 18.38. Entomological notes. Ent. Mon. Mag. 5:372-402. 1841. Entomological notes. Entomologist 1: 73-78. Olivier, A. G. 1808. Entomologie, ou histoire naturelle des insectes, avec leurs caracteres generiques et specifiques, leur description, leur synonym ie, et leur figure enluminee. Coleopteres, vol. 6. Paris. Say, T. 1823. Descriptions of coleopterous insects col- lected in the late expedition to the Rocky Moun- tains, performed by order of Mr. Calhoun, secre- tary of war, under the command of Major Long. J. Acad. Nat. Sci. Philadelphia 3:403-462. ScHAEFFER, C. F. A. 1906. On new and known genera and species of the familv Chrysomelidae. Mus. Brooklyn Inst. Sci. Bull. 1:221-253. ScHWARZ, E. A. 1878. The Coleoptera of Florida. Proc. Amer. Philo. Soc. Philadelphia 17:35.3-471. Smith, J. B. 1885. Some new species of Hispini. Entomo- logia Americana 1:94. Van Dyke, E. C. 1925. Notes and descriptions of new species of west American Hispinae. Pan-Pacific Ent. 1:170-173. Weise, J. 1910. Beitrag zur Kenntnis der amerikanische Hispinen. Arch. Naturg. 76:67-127. 1911. Coleopterorum catalogus, chrysomelidae; Hispinae 35:1-94. FLORA OF THE STANSBURY MOUNTAINS, UTAH Alan C. Taye' Abstract.— The Stansbury Mountains of north central Utah rise over 2000 m above surrounding desert valleys to a maximum elevation of 3362 m on Deseret Peak. Because of the great variety of environmental conditions that can be found in the Stansburys, a wide range of plant species and vegetation types (from shadscale desert to alpine mead- ow) exist there. This paper presents an annotated list of 594 vascular plant species in 315 genera and 78 families. The largest families are Asteraceae (98 species), Poaceae (71), Brassicaceae (33), Fabaceae (27), and Rosaceae (26). Elymiis flcwescens was previously unreported from Utah. Statistical comparison of the Stansbury flora with neighboring mountain floras indicates that the Wasatch Mountains lying 65 km to the east have probably been the primary source area for development of the Stansbury flora. Many lowland species, especially those inhabiting sandy areas, apparently have migrated to the area from the south. The high mountain ranges of the Great Ba- sin are botanically interesting for their isolat- ed montane floras. Surrounded by desert, these islandlike ranges have characteristics in common with oceanic islands (Harper et al. 1978). One of these ranges, the Stansbury Mountains of north central Utah, is particu- larly interesting in supporting a vegetational zonation and flora that are transitional be- tween the Great Basin ranges and the Wasatch Mountains. Geography and Geology The Stansbury Mountains of Tooele Coun- ty, Utah, situated near the eastern edge of the Great Basin about 65 km west of Salt Lake City and the Wasatch Front, are located be- tween 40° 20' and 40° 45' N latitude and 112° 29' and 112° 44' W longitude. The range is bounded on the west by Skull Valley, on the east by Tooele and Rush valleys, on the north by the Great Salt Lake and Stans- bury Island, and on the south by the Onaqui Mountains. The range has a length of 45 km and a width of 21 km at its widest point, and occupies an area of about 909 km 2. The ele- vation ranges from 1280 m (4200 ft) in the valleys to 3362 m (11,031 ft) at the summit of Deseret Peak. Structurally, the Stansbury Mountains are a "gigantic eastward tilted fault block" (Rig- by 1958). The western escarpment rises abruptly from the floor of Skull Valley and is dissected by steep-walled canyons. The east- ern side of the range is generally less rugged except in the vicinity of Deseret Peak, where Pleistocene glacial activity has produced sheer canyon walls and several well-defined horns formed from coalescing glacial cirques. At least 17 cirque basins, two of which con- tain small lakes, occur in the range. Skirting the base of the range are terraces, wave-cut cliffs, spits, and other features produced by Lake Bonneville. Pediment surfaces, bajadas, and alluvial fans are present on the western and eastern edges of the range (Rigby 1958). The core of the range is composed of the Cambrian Tintic Quartzite. Younger Paleo- zoic sedimentary strata, which overlay and flank the quartzite throughout the range where not eroded away, compose the bulk of the northern and southern portions of the range. Lesser amounts of sedimentary and ig- neous formations of Tertiary age are also present as are Quaternary glacial, aeolian, and lacustrine deposits (Rigby 1958). Climate and Soils The climate for the area is classified as cold semiarid or steppe by Trewartha (1968). The city of Tooele, located 16 km to the east of the Stansburys at an elevation of 1545 m (5070 ft), has average January and July tem- peratures of -1.7 C and 24.7 C, respectively, with an average annual temperature of 10.6 C. The high and low temperatures at Tooele 'U.S. Army Intelligence Center and School, Fort Huachuca, Arizona 85613. 619 620 Great Basin Naturalist Vol. 43, No. 4 for 1979 were 35.6 C and -22.2 C (U.S. De- partment of Commerce 1980). Temperature data are not available for the higher portions of the range. Precipitation in the Great Basin is strongly influenced by the orographic effect, with mountains receiving greater amounts than the valleys (Houghton 1969). Tooele receives an average annual precipitation of 41.43 cm, with the largest amounts falling in the months of March (4.67 cm), April (5.59 cm), and May (4.16 cm). The driest months are July (1.78 cm), August (2.36 cm), and Sep- tember (1.83 cm) (U.S. Department of Com- merce 1980). A precipitation station located at 2820 m (9250 ft) on the lee side of Deseret Peak received an average of 140.28 cm (more than three times the valley station average) over the three-year period from 1974 to 1976 (Soil Conservation Service 1979). Soils are diverse in the study area. The En- tisol, Aridisol, and MoUisol soil orders and six soil associations have been mapped in the Stansburys by the Soil Conservation Service (1973). Types of soils range from the strongly alkaline, light-colored soil typical of the val- ley greasewood community to the strongly acidic, dark-colored soil of the montane spruce-fir community (Wilson et al. 1975). Vegetation Eight somewhat distinct vegetation zones or communities, discussed by Billings (1951) and N. Holmgren (1972), are present in the Stansbury Mountains. In order of increasing elevation, they are the shadscale, sage- brush-grass, juniper-pinyon, Douglas fir-white fir, upper sagebrush- grass, Engel- mann spruce-subalpine fir, limber pine-bristlecone pine, and alpine zone. The vegetational zonation in the Stansbury Mountains is transitional between the Wasatch type and the Basin Range type (Bil- lings 1951). Gambel oak {Qtiercus gambelii), a dominant species in the central and south- em Wasatch Mountains, and common on the opposite side of Tooele Valley in the Oquirrh Mountains, is conspicuously absent from the Stansburys. The oak habitat is dominated in- stead by a well-developed Utah juniper {Juni- pertis osteosperma) woodland. Blue spruce (Picea pungens), a component of the Douglas fir-white fir-blue spruce zone in the Wasatch range (N. Holmgren 1972), is also apparently absent from the Stansburys. Bristlecone pine {Pinus longaeva) is a major component of the Great Basin subalpine co- nifer community (Billings 1951, N. Holmgren 1972), and its presence in the Stansburys marks its northern and eastern limits of distri- bution in the Bonneville Basin. Botanical Exploration Captain Howard Stansbury, for whom the mountains and island are named, collected the types of Cowania mexicana var. stanshu- riana, Heuchera rubescens, and Perityle stans- biirii from nearby Stansbury Island in 1850 (Stansbury 1852). Marcus E. Jones made the first known collections from the Stansbury Mountains in 1891 and 1903 (Jones 1965) and collected the type for Phacelia incana from nearby Dugway Valley (Welsh 1982). T. H. Kearney et al. (1914) prepared extensive spe- cies lists for the plant communities of Tooele Valley in their study on the relationship of vegetation to soil moisture and salt content. One new species, Eriogonum kearneyi, was discovered (Tidestrom 1913). S. Flowers col- lected in the range in 1928 and 1930, and B. Maguire visited there in 1943. M. E. Lewis (pers. comm. 1979) prepared a preliminary species list (with 150 species) for the Stans- burys in 1957. In the past two decades a number of bota- nists have collected in the Stansbury Moun- tains, including B. Albee, L. C. Anderson, M. E. Barkworth, E. M. Christensen, W. P. Cot- tam, K. T. Harper, A. H. Holmgren, R. Kass, R. M. Lanner, E. Neese, K. H. Thorne, R. K. Vickory, Jr., and S. L. Welsh. Their collec- tions have contributed to this checklist. For this study, I visited the range from 1978 to 1981 and made over 1400 collections. Discussion of the Flora The diversity of climatic and edaphic habi- tats in the Stansbury Mountains is reflected in the large number of plant species occur- ring in this range. A total of 594 species from 315 genera and 78 families are listed follow- ing this discussion. Of this number, 494 spe- cies from 264 genera and 71 families are pre- sumably native to the range. Though October 1983 Taye: Stansbury Mountains Flora 621 occupying only 0.43 percent of Utah's land area, the Stansbury range has 19.2 percent of the state's 2575 native species (from Welsh et al. 1981). A statistical summary of the flora is presented in Table 1. The number of montane species expected to occur above 2286 m (7500 ft) in elevation on Great Basin mountain ranges can be pre- dicted from the species-area equation in Har- per et al. (1978). With an area of 140 km2 above 2286 m (Behle 1978), the Stansbury Mountains would be expected to have 225 montane species. The number I found was 385. The unexpectedly high number of spe- cies is probably due to the presence of an al- pine zone on this relatively narrow mountain range. Environmental heterogeneity and fa- vorability are more important than area in the determination of floral diversity (Harper et al. 1978). Statistical comparison of 11 mountain floras (listed in Table 2) in the eastern Great Basin (Taye 1981) shows the Stansbury flora to be most similar to the floras of Mount Timpanogos, northern Wasatch, and central Wasatch— 62.0, 61.2, and 60.8 percent sim- ilarity, respectively, using So^rensen's index of similarity (Fig. 1). Thus the Wasatch range is perhaps the primary source area for devel- opment of the Stansbury flora. This might be expected because of the close proximity (65 km) of the Stansbury Mountains to the flo- ristically rich Wasatch Mountains and of the finding by Harper et al. (1978) that Great Ba- sin mountains are dominated by species from the Rocky Mountain floristic element. The floristic relationship between the Wasatch and Stansbury ranges is also evident from the several montane species found in both ranges but not known to occur west of the Stans- bury s (Table 3). Though the Stansbury Mountains and cen- tral Wasatch Mountains rise to comparable heights (Table 2), the Stansbury flora has con- siderably fewer alpine species. Apparently missing are Polygonum viviparum, Salix arc- tica, Silene acaulis, Smelowskia calycina, and many others. Persistent snowdrifts, conducive Table 1. Statistical summary of the vascular plants of the Stansbury Mountains. Ind Families G igenous enera Species Families Introduced Genera Species Lycopodiophyta Eqiiisetophyta Polypodiophyta Pinophyta Magnoliophyta Magnoliopsida 1 1 1 2 56 1 1 4 5 204 1 3 4 10 382 0 0 0 0 7 0 0 0 0 43 0 0 0 3 70 Liliopsida 10 49 94 0 8 27 Totals: 71 Grand totals: Families Genera Species Largest families 264 (native + 494 78 315 594 introduced species) 7 51 100 Asteraceae 87 + 11 Scrophulariaceae 19 + 3 Poaceae 44 4- 27 Boraginaceae 17 + 2 Rosaceae 24+2 Polygonaceae 16 + 1 Cyperaceae Fabaceae 22+0 21+6 Apiaceae Chenopodiaceae 15 + 1 15 + 9 Brassicaceae 20 + 13 Largest genera (native + Onagraceae introduced species) 15 + 0 Carex 17 + 0 Artemisia 7 + 0 Astragalus Eriogonum 12 + 0 10 + 0 Cryptantha Poa 7 + 0 7 + 4 Erigeron 8 + 0 Ribes 7 + 0 622 Great Basin Naturalist Vol. 43, No. 4 to the growth of many alpine species (Billings 1978), are present throughout the summer but they are few and small in size. Altither- mal extinctions (Billings 1978), limited alpine habitat, or failure to reach the Stansburys are possible explanations for their absence there. A list of species occurring above 3050 m (10,000 ft) in elevation is presented in Table 4. Timberline is generally located from 3200 m (10,500 ft) to 3290 m (10,800 ft), but the south slope of Deseret Peak is nearly treeless to an elevation of 2865 m (9400 ft). The influence of the Great Basin floristic division (N. Holmgren 1972) on the Stans- bury flora is seen in the presence of many of the valley and foothill species. Most of these desert species have apparently migrated northward from the Mojave Desert during the warmer postglacial period of the last 10,000 years (Reveal 1979, Wells 1980), and many of them apparently reach their north- em or eastern limit of distribution in the study area (Table 3). The Stansbury flora, in comparison with the flora of the more mesic Wasatch Mountains, has a greater number of species from many characteristically desert genera including Astragalus, Camissonia, Cryptantha, Eriogonum, Phacelia, and Tetra- dymia. A number of species from these and other genera are partially or wholly restrict- ed to sandy areas at the base of the range (Table 5). Plant migration to the Stansbury Moun- tains from northern and western routes has probably been extremely limited because of past and present barriers in the Bonneville Basin. Lake Bonneville, a large freshwater lake which occupied most of northwestern Utah during the Pleistocene (Morrison 1965), and the present Great Salt Lake-Great Salt Lake Desert have undoubtedly restricted the migration of most plant species. One species that appears to have reached the Stansburys from the north is Elymiis flavescens, a spe- cies disjunct from the Snake River Plains of Idaho (Cronquist et al. 1977) and previously unreported from the state of Utah. The impact of humans on the Stansbury flora can be seen by the large number (100) of cultivated or adventive species. Most of these species are limited to low elevations where even the vegetation has been marked- ly altered in some areas. Especially common are Agropyron cristattim, Bromus tectorwn, Halogeton glomeratus, and Salsola iberica. Only a few introduced species such as Dac- tylis glomerata, Poa pratensis, and Taraxacum officinale are present at elevations greater than 2400 m (7874 ft). None of the species are endemic to the study area, though several are restricted to somewhat larger areas. Astragalus eurekensis is endemic to central Utah (Welsh 1978b); Astragalus lentiginosus var. pohlii is a very narrow endemic of Rush Valley and Skull Valley (Welsh and Barneby 1981); Eriogonum grayi is endemic to alpine areas of north cen- tral Utah (Reveal 1973); and Sphaeromeria di- versifolia is apparently restricted to the cen- tral and southern Wasatch Mountains and west to the Quinn Canyon Range of Nevada (Holmgren et al. 1976). Table 2. Floras of the eastern Great Basin (after Harper et al. 1978). Native + introduced species Maximum elevation (m) Source of information Mountain islands 1 Stansbury Mountains 2 East Tintic Mountains 3 Raft River Mountains 4 Deep Creek Mountains 5 Jarbidge Mountains 6 Ruby Mountains 7 Wheeler Peak Mainland areas 8 Northern Wasatch Mountains 9 Central Wasatch Mountains 10 Mount Timpanogos 11 Wasatch Plateau 494 + 100 3362 This report 162 + 48 2505 Nebeker 1975 303 + 23 3015 Preece 1950 569 -1- 47 3688 McMillan 1948, Welsh 1978a 478 + 22 3288 Lewis 1975 524 + 24 .3471 Lewis 1971 389 + 23 3981 Lewis 1973 767 + 528 3042 A. Holmgren 1972 911 -1- 228 3502 Arnow et al. 1980 538 -(- 90 3581 AUred 1975 826 + 86 3440 Lewis 1980 October 1983 Taye: Stansbury Mountains Flora 623 Fig. 1. Plexus diagram of floristic similarities among mountain ranges in the eastern Great Basin. High values cor- respond to high similarity. Comparisons were made using Sdrensen's index of similarity. Data are from Taye (1981). Acknowledgments Many individuals kindly offered assistance during the preparation of this checkhst. I am especially grateful to Dr. Stanley L. Welsh, curator of the herbarium of Brigham Young University, for guiding and encouraging the completion of this study. Drs. Kimball T. Harper, Elizabeth Neese, and Samuel R. Rushforth of Brigham Young University kind- ly reviewed and commented upon a portion of the manuscript. Assistance with plant identifications and herbarium research was given by Kaye H. Thome, assistant curator of the Brigham Young University herbarium; Dr. Mary E. Barkworth and Dr. Leila Shultz, director and curator, respectively, of the In- termountain Herbarium at Utah State Uni- 624 Great Basin Naturalist Vol. 43, No. 4 versity; Lois Amow and Beverly Albee, cura- tor and assistant curator, respectively, of the Garrett Herbarium at the University of Utah; and Sherel Goodrich and Mont E. Lewis of the U.S. Forest Service. Bill Wall of the U.S. Forest Service graciously shared the Stans- bury Guard Station with me for three sum- mers and helped collect many of the plant specimens. Special thanks are expressed to Professor Arthur H. Holmgren, retired cura- tor of the Intermountain Herbarium, for his sustained assistance and interest in this study. I also thank Dee Applegate for faithfully typ- ing the manuscript and Mathew Chatterly for preparing the figure. Finally, deep apprecia- tion is expressed to my parents for their love and support throughout the course of this study. Annotated List of Vascular Plants The following list of families, genera, and species is arranged in alphabetical order Table 3. Plants with distributional limits in the Stansbury Mountains and vicinity. Northern limit of distribution Astragalus ceramicus (Puddle Valley) Astragalus eurekensis Astragalus mollissimus Ceanothus maiiinii Cyinopterus fendleri Echinoceretis triglochidiatus var. melancicanthus (Stansbury Island) Eriogoniim kearneyi Eriogoniim umbellatum var. subaridum Flaveria campestris Geranium parnji Lijcium andersonii (Puddle Valley) Finns longaeva Stephanomeria pauciflora Eastern limit of distribution Chamaebatiaria millefolium (Oquirrh Mtns.) Erigeron argentatus Euphorbia ocellata var. arenicola Ribes velutinum var. velutinum (Oquirrh Mtns.) Southern limit of distribution Elijmus flavescens Western limit of distribution Castilleja applegatei var. viscida Eriogonum grayi Geranium parnji Mertensia arizonica var. leonardii Mertensia brevistyla within the divisions of Cronquist et al. (1972). Nomenclature generally follows that of Welsh et al. (1981). Volumes 1 and 6 of the Table 4. Nonarboreal species occurring above 3050 m in elevation in the Stansbury Mountains. Achillea millefolium Agropyron tracln/caulum Androsace septentrionalis Antennaria corymbosa Arabis holboellii var. secunda Arenaria kingii Artemisia tridentata var. vaseyana Astragalus kentrophyta var. implexus Astragalus tenellus Carex atrata var. erecta Carex haydeniana Castilleja applegatei var. viscida Castilleja rhexifolia Cirsium eatonii Cymopterus hendersonii Cyinopterus longipes Cystopteris fragilis Draba stenoloba Erigeron compositus Erigeron eatonii Erigeron leiomerus Eriogonum grayi Eriogonum umbellatum var. desereticum Festuca ovina var. brevifolia Geum rossii var. turbinatum Haplopappus macronema Heuchera rubescens Ivesia gordonii Juniperus communis var. depressa Lathyrus lanzwertii Lesquerella occidentalis var. cinerascens Leucopoa kingii Lewisia pygmaea Linum perenne Lupinus argenteus Luzula spicata Oxyria digyna Penstemon liumilis Phlox pulvinata Poa fendleriana Poa secunda Potentilla ovina Ribes montigenum Saxifraga rhomboidea Sedum lanceolatitm Selaginella watsonii Senecio fremontii Senecio streptanthifolius Sitanion hystrix Solidiigo parryi Swertia radiata Syniptioricarpos oreophilus Synthyris pinnatifida Thlaspi montanum Trisetum spicatum Valeriana acutiloba var. pubicarpa Zigadenus elegans October 1983 Taye: Stansbury Mountains Flora 625 Intemiountain Flora (Cronquist et al. 1972, 1977) were used for the nomenclature of the vascular cryptogams, gymnosperms, and monocots. Other helpful sources were Arnow et al. (1980), Hitchcock and Cronquist (1973), Welsh (1978b), Welsh and Moore (1973), and Welsh and Reveal (1977). Synonyms are not listed imless in recent use. Introduced species are preceded by an asterisk (°). A representa- tive collection number (my own unless other- wise noted) is cited for each species, and all specimens cited are deposited in the her- barium at Brigham Young University (BRY) unless otherwise indicated. A number of specimens are deposited in the Garrett Her- barium at the University of Utah (UT) and/ or the Intemiountain Herbarium at Utah State University (UTC). Frequency of most species is estimated based on the foll/Dwing scale from Thome (1967): rare, 1 to 3 collections or observation stations; infrequent, 4 to 7 sta- tions; frequent, 8 to 12 stations; common, more than 12 stations. This list should not be considered complete since many additional species remain to be discovered. Division Lycopodiophyta Selaginellaceae Selaginella watsonii Underw. Watson Spikemoss. Frequent; open rocky slopes at high elevations. 507. Division Equisetophyta Equisetaceae Equisetum arvense L. Field Horsetail. Rare; streamside in North Willow Canyon. 437. Equisetum hyemale L. Common Scouring Rush. Frequent; streamside at low and middle elevations. 311 (UT). Equisetum laevigatum A. Br. Smooth Scouring Rush. Frequent; streamside at low elevations. 850. Division Polypodiophyta Polypodiaceae Cystopteris fragilis (L.) Bernh. Brittle Blad- der Fern. Common; shaded and open mesic sites from middle elevations to alpine. 943. Pellaea breweri D. C. Eat. Brewer Cliff- brake. Rare; limestone outcrops at middle elevations. 649. Pohjstichum lonchitis (L.) Roth. Mountain Holly Fern. Rare; base of quartzite cliff in Douglas fir community. Taye & Herrick 1441. Woodsia oregana D. C. Eat. Oregon Woodsia. Rare; dry, rocky slopes in juniper zone. Taye & Herrick 1430. Table 5. Species restricted (or most common) to sandy areas in the Stansbury Mountains and vicinity. Abronia fragrans Agropi/wn dasystachyum AmarantJitis bhtoides Astragalus ceramicus Astragalus gey eri Astragalus molUssimus Camissonia parvula Camissonia scapoidea ssp. brachycarpa Chenopodium leptophyllum Cryptantha circumscissa Cryptantlia fendleri Cryptantha kelseyana Cryptantha pterocarya Cymopterus fendleri Elymus flavescens Erigeron argentatus Eriogonum cernuum Eriogonum liookeri Eriogonum kearneyi Eriogonum microthecum var. laxiflorum Eriogonum umbellatum var. subaridum Euphorbia ocellata var. arenicola Cilia inconspicua Cilia leptomeria Cilia polycladon Layia glandulosa Leptodactylon pungens Lupinus pusillus var. intermontanus Lygodesmia dianthopsis Malacothrix sonchioides Mentzelia albicaulis Nama densum Nicotiana attenuate Oenothera pallida Orobanche corymbosa Phacelia ivesiana Psoralea lanceolata Rumex venosus Sporobolus cryptandrus Stephanomeria exigua Stipa comata Streptanthella longirostris Tiquilia nuttallii Toicnsendia florifer Tripterocalyx micranthus Vulpia octoflora 626 Great Basin Naturalist Vol. 43, No. 4 Division Pinophyta Division Magnoliophyta Cupressaceae Juniperus communis L. var. depressa Pursh. Common Mountain Juniper. Frequent; meadows and open slopes at high elevations. 983. Juniperus osteosperma (Torr.) Little. Utah Juniper. Common; a dominant on dry slopes at low and middle elevations. 1006. Juniperus scopulorum Sarg. Rocky Moun- tain Juniper. Common; near streams and on mesic slopes from low to middle elevations. 863. Class Magnoliopsida Aceraceae Acer glabrum Torr. Rocky Mountain Maple. Common; streamside and mesic slopes at middle elevations. 1105. Acer grandidentatum Nutt. Bigtooth Maple. Infrequent; canyon bottoms at low elevations. 99 (UT and UTC). Acer negundo L. Boxelder. Common; streamside from low to middle elevations. 862. Pinaceae Abies concolor (Cord. & Glend.) Lindl. White Fir. Common; a dominant along streams and on mesic slopes at low and middle elevations. 356 (UT and UTC). Abies lasiocarpa (Hook.) Nutt. Subalpine Fir. Common; a dominant at moderately high elevations. 347 (UT and UTC). Picea engelmannii Parry. Engelmann Spruce. Common; a dominant at moderately high elevations. 350 (UT and UTC). "Picea pungens Engelm. Blue Spruce. Rare; apparently planted in South Willow Canyon. 578 (UTC). "Pinus contorta Dougl. Lodgepole Pine. Rare; apparently planted in South Willow Canyon. 1183. Pinus flexilis James. Limber Pine. Com- mon; a dominant on dry slopes from middle elevations to timberline. 355 (UT and UTC). Pinus longaeva D. K. Bailey. Bristlecone Pine. Infrequent; locally a dominant on lime- stone slopes at moderately high elevations. 589. Pinus monophylla Torr. & Frem. Singleleaf Pinyon. Infrequent; locally a dominant from low to middle elevations. 1031. "Pinus ponderosa Laws. Ponderosa Pine. Rare; apparently planted in South Willow Canyon. 688 (UTC). Pseudotsuga menziesii (Mirb.) Franco var. glauca (Beissn.) Franco. Douglas Fir. Com- mon; a dominant along streams and on dry to mesic slopes from low to high elevations. 574. Amaranthaceae Amaranthus blitoides Wats. Prostrate Pig- weed. [A. graecizans L.] Collected only from sandy site in juniper zone. 1301. Anacardiaceae Rhus trilobata Nutt. var. trilobata. Squaw- bush. Frequent; dry slopes of foothills. 859. Apiaceae Angelica pinnata Wats. Small-leaf Angel- ica. Infrequent; streamside at middle eleva- tions. 1179. Berula erecta (Huds.) Cov. Cutleaf Water- parsnip. Infrequent; along streams in the foothills. 1095. "Conium maculatum L. Poison Hemlock. Infrequent; mesic, sometimes disturbed sites at low elevations. 498. Cymopterus fendleri Gray. Chimaya. Rare; locally frequent in sandy areas of Skull Val- ley. 777. Cymopterus hendersonii (Coult. & Rose) Cronq. [Pteryxia hendersonii (Coult. & Rose) Math. & Const.] Infrequent; meadows and open slopes above 3000 m. 501. Cymopterus longipes Wats. Longfoot Springparsley. Frequent; open slopes from the foothills to near timberline. 982. Cymopterus purpurascens (Gray) Jones. Purple Springparsley. Infrequent; dry foot- hills. 766 (UTC). Heracleum spondylium L. ssp. montanum (Schleich.) Briq. Cow Parsnip. [H. lanatum October 1983 Taye: Stansbury Mountains Flora 627 Michx.] Common; streamside at middle ele- vations. 1116. Ligusticum filicinum Wats. Fernleaf Lo- vage. Infrequent; open slopes at middle ele- vations. Taye & Wall 1217. Lomatium dissectum (Nutt.) Math. & Const. Fernleaf Lomatium. Open slopes from low to middle elevations. 874. Lomatium grayi Coult. & Rose. Common; rocky slopes in juniper zone. 772. Lomatium nuttallii (Gray) Macbr. Thread- leaf Lomatium. Frequent; open slopes at middle elevations. Mill Fork, 135 (UT and UTC). Orogenia linearifolia Wats. Indian Potato. Frequent; foothills in springtime. 369. Osmorhiza chilensis H. & A. Sweetroot. Frequent; streamside and forest understory at middle elevations. 1103. Osmorhiza depauperata Phil. Sweetroot. Frequent; streamside and forest understory at middle elevations. 1104. Osmorhiza occidentalis (Nutt.) Torr. Sweetanise. Frequent; streamside and forest understory at middle elevations. 221 (UT and UTC). Apocynaceae Apocynum androsaemifolium L. Spreading Dogbane. Infrequent; open slopes at middle elevations. 1184. Apocynum cannabinum L. Indian Hemp. Rare; on road embankment in South Willow Canyon. 572 (UTC). Asclepiadaceae Asclepias asperula (Dene.) Woodson. Milk- weed. Rare; dry slopes in juniper zone. 1121. Asclepias speciosa Torr. Showy Milkweed. Infrequent; disturbed sites at low elevations. 593 (UTC). Asteraceae Achillea millefolium L. ssp. lanulosa (Nutt.) Piper. Common Yarrow. Common; meadows and mostly open slopes from the foothills to timberline. 1292. Agoseris aurantiaca (Hook.) Greene. Or- ange Dandelion. Frequent; open slopes from the foothills to subalpine. 417 (UTC). Agoseris glauca (Pursh) Raf. Mountain Dandelion. Open slopes from the foothills to subalpine. 693. Ambrosia acanthicarpa Hook. Burweed. Common; sandy areas at low elevations. 686. Ambrosia psilostachya DC. Western Rag- weed. Collected only from streamside in the foothills near Delle Ranch. Taye & Dillman 1417. Antennaria corymbosa E. Nels. Flattop Pussytoes. Frequent; meadows and open slopes near timberline. 969. Antennaria dimorpha (Nutt.) T. & G. Low Pussytoes. Infrequent; juniper zone. 771. Antennaria microphylla Rydb. Rosy Pussy- toes. [A. rosea Greene] Infrequent; locally common on open slopes from middle eleva- tions to subalpine. 931. "Arctium minus (Hill) Bernh. Common Bur- dock. Infrequent; streamside and shaded mes- ic sites from the valleys to middle elevations. 1187. Arnica cordifolia Hook. Heartleaf Arnica. Common; meadows and forest understory from middle elevations to subalpine. 223 (UT and UTC). Arnica longifolia D. C. Eat. Longleaf Ar- nica. Infrequent; locally common in mesic meadows at middle elevations. Taye & Wall 1172. Arnica mollis Hook. Hairy Arnica. In- frequent; meadows and streamside at middle elevations. 624. Artemisia arbuscula Nutt. Low Sagebrush. Open rocky slopes at middle elevations. 658. Artemisia dracunculus L. Tarragon. Fre- quent; open slopes from the foothills to middle elevations. 1266. Artemisia ludoviciana Nutt. var. incompta (Nutt.) Cronq. Louisiana Sagebrush. Locally common in subalpine meadows. 661. Artemisia ludoviciana Nutt. var. ludovi- ciana. Common; open slopes from the foot- hills to middle elevations. 1227. Artemisia nova A. Nels. Black Sagebrush. [A. arbuscula Nutt. var. nova (A. Nels.) Cronq.] Rocky slopes at low elevations. 1015. Artemisia spiciformis Osterhout. Locally common on open slopes just below tim- berline. 1259. Artemisia spinescens D. C. Eat. Bud Sage- brush. Infrequent; dry valley sites. 891. 628 Great Basin Naturalist Vol. 43, No. 4 Artemisia tridentata Nutt. ssp. tridentata. Big Sagebrush. Common; open slopes from the valleys to middle elevations. 1016. Artemisia tridentata Nutt. ssp. vaseyana (Rydb.) Beetle. Open slopes at middle eleva- tions. 1241. Aster chilensis Nees ssp. adscendens (Lindl.) Cronq. Everywhere Aster. Common; dry to mesic sites from the foothills to sub- alpine. 718. Aster engelmannii (D. C. Eat.) Gray. En- gelmann Aster. Infrequent; open forest un- derstory at middle elevations. 692. Aster glaucodes Blake. Infrequent; dry slopes at middle elevations. 727. Balsamorhiza hookeri Nutt. var. hispidula (Sharp) Cronq. Hooker's Balsamroot. Seen only in the foothills near South Willow Can- yon. 411 (UTC). Bakamorhiza sagittata (Pursh) Nutt. Ar- rowleaf Balsamroot. Common; open slopes in the foothills. 378. Brickellia grandi flora (Hook.) Nutt. Tas- selflower. Rocky sites in juniper zone. Taye & Herrick 1427. Brickellia microphylla (Nutt.) Gray. Little- leaf Brickellia. Rocky slopes at low eleva- tions. 750. "Centaiirea cyanus L. Bachelor's Buttons. Rare; near guard station in South Willow Canyon. 171 (UT). "Centatirea maculosa Lam. Rare; roadside near mouth of South Willow Canyon. 670. "Centatirea repens L. Russian Knapweed. Rare; streamside near mouth of Box Canyon. 1148. Chaenactis douglasii (Hook.) H. & A. Hoary Chaenactis. Frequent; juniper zone. 842. Chrysothamnus nauseosus (Pallas) Britt. var. alhicaidis (Nutt.) Rydb. Rubber Rabbit- brush. Frequent; valleys and foothills. 752. Chrysothamnus nauseosus (Pallas) Britt. var. consimilis (Greene) Hall. Collected only from the foothills. 1019. Chrysothamnus nauseosus (Pallas) Britt. var. gnaphaloides (Greene) Hall. [C.nauseosus ssp. hololeucus (Gray) H. & C] Collected only from the foothills. 1265. Chrysothamnus nauseosus (Pallas) Britt. var. turhinatus (Jones) Blake. Locally com- mon in sandy soil of Skull Valley. Anderson 5118. Chrysothamnus viscidiflorus (Hook.) Nutt. var. puberulus (D. C. Eat.) Jeps. Douglas Rabbitbrush. Locally common in clay soil in Skull Valley. Anderson 5116. Chrysothamnus viscidiflorus (Hook.) Nutt. var. viscidiflorus. Common; open slopes from the foothills to middle elevations. 307. °Cichorium intybus L. Common Chicory. Rare; streamside in the foothills. 1146. Cirsium eatonii (Gray) Robins. Eaton Thistle. Frequent; open and wooded slopes from middle elevations to subalpine. Taye & Wall 1443. Cirsium scariosum Nutt. Elk Thistle. Lo- cally frequent in Skull Valley. 1274. Cirsium undulatum (Nutt.) Spreng. Wavy- leaf Thistle. Frequent; dry foothills. 1001. "Cirsium vulgare (Savi) Ten. Bull Thistle. Disturbed sites at low to middle elevations. 259 (UTC). Conyza canadensis (L.) Cronq. Horseweed. Infrequent; streamside at low elevations. 682. Crepis acuminata Nutt. Tapertip Hawks- beard. Open slopes from the foothills to middle elevations. 435. Crepis atraharha Heller. Slender Hawks- beard. Open slopes at middle elevations. 605. Crepis modocensis Greene. Low Hawks- beard. Infrequent; juniper zone. 527 (UTC). Crepis occidentalis Nutt. Western Hawks- beard. Open slopes from the foothills to middle elevations. 867. Crepis runcinata T. & G. var. glauca (Nutt.) Babe. & Stebbins. Meadow Hawks- beard. Rare; near spring in Skull Valley. 1376. Erigeron argentatus Greene. Fleabane. In- frequent; sandy areas in foothills. 495. Erigeron compositus Pursh. Fernleaf Flea- bane. Infrequent; locally common in mead- ows and on open rocky slopes above 2800 m. 964. Erigeron divergens T. & G. Spreading Fleabane. Rare; streamside near mouth of Spring Canyon. 1132. Erigeron eatonii Gray. Eaton Fleabane. Common; open slopes and meadows from the foothills to alpine. 430. Erigeron engelmannii A. Nels. Engelmann Fleabane. Valleys and foothills. 419. Erigeron leiomerus Gray. Smooth Daisy. Infrequent; rocky slopes above 2800 m. 665. October 1983 Taye: Stansbury Mountains Flora 629 Erigeron lonchophyllus Hook. Spearleaf Fleabane. Rare; near spring in Skull Valley. 1377. Erigeron speciosus (Lindl.) DC. Showy Fleabane. Common; open and wooded slopes from middle elevations to subalpine. 650. Eupatoriwn occidentale Hook. Western Eupatorium. Rare; base of quartzite cliff in Douglas fir community in Muskrat Canyon. Taye & Herrick 1440. Flaveria campestris J. R. Johnst. Rare; lo- cally common near springs in Skull Valley. Anderson & Thome 5114. Gnaphaliiim pahistre Nutt. Lowland Cud- weed. Rare; streamside near mouth of Spring Canyon. 1131. 'Grindelia squarrosa (Pursh) Dun. Gum- weed. Common; disturbed sites at low eleva- tions. 258 (UT and UTC). Haplopappus acatdis (Nutt.) Gray. Cushion Goldenweed. Infrequent; open rocky slopes from the foothills to middle elevations. 532a. Haplopappus lanceolatus (Hook.) T. & G. Lanceleaf Goldenweed. Locally frequent in moist soil near spring in Skull Valley. Ander- son & Thome 5115. Haplopappus macronema Gray. Whitestem Goldenweed. Frequent; meadows and open slopes from middle elevations to alpine. Taye & Wall 1212. Haplopappus rydbergii Blake. Rydberg Goldenweed. Frequent; foothills to middle elevations, mostly on limestone outcrops. 999. Helianthella uniflora (Nutt.) T. & G. One- flower Helianthella. Locally common on open slopes at middle elevations. 513. Helianthus annuus L. Common Sunflower. Common; valleys and foothills, often in dis- turbed areas. 1161. Heliomeris multiflora Nutt. Showy Gold- eneye. [Viguiera multiflora (Nutt.) Blake] Frequent; meadows and wooded slopes at middle elevations. 729. Heterotheca villosa (Pursh) Shinners. Hairy Golden Aster. [Chrysopsis villosa (Pursh) Nutt.] Dry rocky slopes in juniper zone. Taye & Herrick 1429. Iva axillaris Pursh. Poverty Sumpweed. In- frequent; valleys and foothills near drainages. 1048. Lactuca pulchella (Pursh) DC. Blue Let- tuce. Infrequent; dry to mesic sites from the foothills to middle elevations. 602. Layia glandulosa (Hook.) H. & A. White- daisy Tidytips. Infrequent; sandy areas of the valleys and foothills. 778. Leucelene ericoides (Torr.) Greene. Rare; locally common on south-facing road em- bankment near Clover Creek in juniper zone. 457. Lygodesmia dianthopsis (D. C. Eat.) Tomb. Skeletonweed. [L. grandiflora (Nutt.) T. & G.] Frequent; foothills, mostly in sandy areas. 481. Machaeranthera canescens (Pursh) Gray. Hoary Aster. Frequent; dry slopes of juniper zone. 305. Madia glomerata Hook. Cluster Tarweed. Rare; near little-used road above Hickman Pass in Douglas fir community. 659. Malocothrix sonchioides (Nutt.) T. & G. Desert Dandelion. Infrequent; sandy areas in valleys and foothills. 796. Microseris nutans (Geyer) Schultz-Bip. Nodding Microseris. Open slopes at middle elevations. 462. "Onopardum acanthium L. Scotch Cotton Thistle. Rare; roadside in Skull Valley. 1422. Perityle stansburii (Gray) Macbr. [Laph- amia stansburii Gray] Frequent; on rock out- crops in foothills. 545. Petradoria putnila (Nutt.) Greene. Rock Goldenrod. Frequent; open slopes from the foothills to subalpine. 618. Rudbeckia occidentalis Nutt. Western Coneflower. Frequent; moist sites at middle elevations. 1180. Senecio eremophilus Rich. Desert Ground- sel. Frequent; open and wooded slopes from middle elevations to subalpine. 699. Senecio fremontii T. & G. var. blitoides (Greene) Cronq. Dwarf Mountain Butter- weed. Wooded and open slopes near tim- berline. Neese 9666. Senecio integerrimus Nutt. var. exaltatus (Nutt.) Cronq. Columbia Groundsel. Fre- quent; open and wooded slopes from the foothills to middle elevations. 392. Senecio multilobatus T. & G. Lobeleaf Groundsel. Common; juniper zone. 551. Senecio streptanthifolius Greene. Cleftleaf Groundsel. Frequent; open and wooded slopes from middle elevations to timberline. 526. Solidago canadensis L. Canada Goldenrod. Mesic sites at middle elevations. 310 (UT and UTC). 630 Great Basin Naturalist Vol. 43, No. 4 Solidago parryi (Gray) Greene. Parry Gold- enweed. [Haplopappus parryi Gray] In- frequent; open slopes near timberline. 704. Solidago sparsiflora Gray. Slender Gold- enrod. Open, rocky slopes from the foothills to middle elevations. 691. "Sonchus asper (L.) Hill. Prickly Sowthistle. Collected only from moist sandy soil in the foothills at Sand Spring. 683 (UTC). Sphaeromeria diversifolia (D. C. Eat.) Rydb. [Tanacetum diversifolium D. C. Eat.] Frequent; on rock outcrops from middle ele- vations to subalpine. 719. Stephanomeria exigua Nutt. Small Wirelet- tuce. Infrequent; sandy areas in the valleys and foothills. 1190. Stephanomeria pauciflora (Torr.) A. Nels. Wireiettuce. Rare; dry foothills of Salt Moun- tain. 1155. "Taraxacum officinale Weber. Common Dandelion. Frequent; dry to mesic, mostly disturbed sites from low to middle elevations. 879. Tetradymia canescens DC. Gray Horse- brush. Locally common in juniper zone. 214 (UT and UTC). Tetradymia glabrata Gray. Littleleaf Horsebrush. Frequent; valleys and foothills. 895. Tetradymia nuttallii T. & G. Nuttall Horsebrush. Frequent; valleys and foothills. 405. Tetradymia spinosa H. & A. Cottonthorn Horsebrush. Rare; valleys. 893. Townsendia florifer (Hook.) Gray. Showy Townsendia. Frequent; sandy areas of valleys and foothills. 480. °Tragopogon dubius Scop. Yellow Salsify. Rare; disturbed sites at low elevations. 436. Wyethia amplexicaulis Nutt. Mulesear Wyethia. Locally common on open slopes from the foothills to middle elevations. 882. Xanthium strumarium L. Common Cockle- bur. Rare; valleys and foothills in disturbed sites. 1269. Xanthocephalum sarothrae (Pursh) Shin- ners. Broom Snakeweed. [Gutierrezia sa- rothrae (Pursh) Britt. & Rusby] Common; dry slopes from the valleys to middle elevations. 306. Berberidaceae Mahonia repens (Lindl.) G. Don. Oregon Grape [Berberis repens Lindl.] Common; from dry slopes in the foothills to dense co- nifer forest at higher elevations. 963. Betulaceae Alnus incana (L.) Moench. Mountain Al- der [A. tenuifolia Nutt.] Rare; locally fre- quent in North Willow Canyon near stream. 215 (UT and UTC). Boraginaceae Amsinkia retrorsa Suksd. Rigid Fiddleneck. Known from one collection from South Wil- low Canyon. Maguire 21807a (UTC). Amsinkia tessellata Gray. Tessellate Fid- dleneck. Infrequent; dry slopes at low eleva- tions. 781. Cryptantha circumscissa (H. & A.) Johnst. Matted Cryptantha. Infrequent; locally com- mon in sandy areas of valleys and foothills. 775. Cryptantha fendleri (Gray) Greene. Sandy areas of valleys and foothills. 809. Cryptantha flavoctilata (A. Nels.) Payson. Dry slopes of valleys and foothills. 795. Cryptantha humilis (Gray) Payson. Dwarf Catseye. Common; rocky slopes below 2200 m. 794. Cryptantha kelseyana Greene. Collected only from a sandy area in the foothills near Condie Meadows. 915. Cryptantha pterocarya (Torr.) Greene. Winged Cryptantha. Collected only from a sandy area in Blue Canyon. 790. Cryptantha torreyana (Gray) Greene. Dry slopes in the foothills. 471. "Cynoglossum officinale L. Hound's Tongue. Frequent; disturbed sites at low ele- vations. 118 (UTC). Hackelia floribunda (Lehm.) Johnst. Many- flowered Stickseed. Frequent; meadows and open slopes from middle elevations to sub- alpine. 1205. Hackelia patens (Nutt.) Johnst. Spreading Stickseed. Frequent; dry slopes from low to middle elevations. 92 (UT and UTC). "Lappula echinata Gilib. European Stick- seed. Infrequent; disturbed sites in valleys and foothills. 784. Lappula occidentalis (Wats.) Greene. Western Stickseed. [L. redowskii (Hornem.) Greene] Frequent; dry slopes of valleys and foothills. 888. October 1983 Taye: Stansbury Mountains Flora 631 Lithospermum ruderale Dougl. Wayside Gromwell. Infrequent; dry foothills. 428. Mertensia arizonica Greene var. leonardii (Rydb.) Johnst. Arizona Bluebells. Infrequent; streamside and in open forest understory at middle elevations. 1057. Mertensia brevistyla Wats. Shortstyle Blue- bells. Infrequent; dry foothills. 461 (UTC). Mertensia oblongifolia (Nutt.) G. Don var. nevadensis (A. Nels.) L. O. Williams. Oblong- leaf Bluebells. Frequent; open slopes from the foothills to middle elevations. 13 (UT). Tiquilia nuttallii (Hook.) A. Richards. [Coldenia nuttallii Hook.] Rare; locally fre- quent in sandy areas of Skull Valley. 954. Brassicaceae "Alyssum alyssoides L. Pale Alyssum. Lo- cally common in juniper zone. 1388. ° Alyssum desertorum Stapf. Desert Alys- sum. Locally common in disturbed areas at low elevations. 398 (UTC). Arabis drummondii Gray. Drummond's Rockcress. Infrequent; open forest and mead- ows from middle elevations to subalpine. 576. Arabis glabra (L.) Bemh. Tower Mustard. Infrequent; open forest and meadows at middle elevations. 872. Arabis liolboellii Hornem. var. secunda (Howell) Jeps. Holboell's Rockcress. Com- mon; open slopes from the foothills to tim- berline. 944. Arabis lignifera A. Nels. Rockcress. Col- lected only from the foothills near Blue Can- yon. 807. "Barbarea vulgaris R. Br. Yellowrocket Wintercress. Rare; streamside in East Hick- man Canyon. 1294. "Camelina microcarpa Andrz. False Flax. Infrequent; juniper zone. 805. "Capsella bursa-pastoris (L.) Medicus. Shepherd's Purse. Rare; streamside in North Willow Canyon. 374. "Cardaria draba (L.) Desv. Whitetop. Com- mon; disturbed sites at low elevations. 1309. Chlorocrambe hastata (Wats.) Rydb. Rare; subalpine conifer community on Deseret Peak. Neese 9709. "Chorispora tenella (Pallas) DC. Locally common in distm-bed sites at low elevations. 764. Descurainia pinnata (Walt.) Britt. Western Tansymustard. Frequent; beneath juniper and on open slopes of foothills. 887. ° Descurainia sophia (L.) Webb. Flixweed. Frequent; disturbed sites at low elevations. 396. Draba cuneifolia Nutt. Wedgeleaf Draba. Infrequent; dry slopes from low to middle elevations. 803. Draba stenoloba Ledeb. Slender Draba. In- frequent; open slopes and moist meadows from middle elevations to timberline. 506. Erysimum asperum (Nutt.) DC. Wallflowr. Frequent; open slopes from the foothills to timberline. 885. Hutchinsia procumbens (L.) Desv. Locally common in the valleys near springs. 1303. Lepidium montanum Nutt. var. mon- tanum. Mountain Pepperweed. Infrequent; sandy areas of Skull Valley. 951. "Lepidium perfoliatum L. Clasping Pepper- weed. Frequent; disturbed sites at low eleva- tions. 892. Lesquerella occidentalis Wats. var. cine- rascens Maguire & Holmgren. Western Blad- derpod. Infrequent; rocky slopes, usually near timberline and above. 453. "Malcolmia africana (L.) R. Br. Frequent; disturbed sites at low elevations. 595 (UTC). "Nasturtium officinale R. Br. Watercress. [Rorippa nasturtium-aquaticum (L.) Schinz & R. Keller] Frequent; along streams at low ele- vations. 849. Physaria chambersii Rollins. Twinpod. In- frequent; dry slopes in juniper zone. 763. Rorippa curvipes Greene. Yellowcress. Col- lected only from edge of South Willow Lake in moist soil. Taye & Wall 725 (UTC). "Sisymbrium altissimum L. Tumbling Mus- tard. Infrequent; disturbed sites at low eleva- tions. 925. Stanleya pinnata (Pursh) Britt. Prince's Plume. Frequent; valleys and on dry slopes in the foothills. 998. Streptanthella hngirostris (Wats.) Rydb. Frequent; sandy areas of valleys and foothills. 793. Streptanthus cordatus Nutt. Collected only from a juniper community near Johnson Pass. 770. Thelypodium integrifolium (Nutt.) Endl. var. integrifolium. Rare; locally frequent in Skull Valley in greasewood community. 1194. 632 Great Basin Naturalist Vol. 43, No. 4 Thelypodium sagittatum (Nutt.) Endl. var. vermicularis Welsh & Reveal. Rare; grease- wood community in Skull Valley. 1333. "Thlaspi arvense L. Field Pennycress. Lo- cally common in disturbed sites. 633 (UTC). Thlaspi montanum L. var. montanum. Wild Candytuft. Frequent; meadows and open slopes from middle elevations to tim- berline. 962. Cactaceae Echinocereus triglochidiatus Engelm. var. melanacanthus (Engelm.) L. Benson. Hedge- hog Cactus. Frequent; dry slopes in juniper zone, often growing from limestone or quart- zite outcrops. 361. Opuntia polyacantha Haw. Prickly Pear. Common; valleys to middle elevations on dry slopes. 897. Campanulaceae "Campanula rapunculoides L. Creeping Bellflower. Rare; abandoned ranch at mouth of Big Hollow. 744. Capparidaceae Cleome serrulata Pursh. Rocky Mountain Beeplant. Frequent; valleys and foothills in generally disturbed sites. 958. Caprifoliaceae Sambucus caerulea Raf. Blue Elderberry. Frequent; near streams, in mesic forest, and on dry open slopes from the foothills to middle elevations. 1122. Sambucus racemosa L. Red Elderberry. In- frequent; meadows and open slopes from middle elevations to subalpine. 988. Symphoricarpos oreophilus Gray. Mountain Snowberry. Common; forest understory and open slopes from the foothills to subalpine. 858. Caryophyllaceae Arenaria kingii (Wats.) Jones. King's Sand- wort. Common; open slopes from the foot- hills to alpine. 425. °Cerastiuni fontanum Baumg. Mouse-ear Chickweed. [C. vulgatum L.] Collected only from streamside in Davenport Canyon. 56 (UT and UTC). "Holosteum umbellatum L. Jagged Chick- weed. Infrequent; locally common in rocky foothills. 1315. Lychnis drummondii (Hook.) Wats. Drum- mond Campion. [Silene drununondii Hook.] Infrequent; forest understory and open slopes from middle elevations to subalpine. 1238. Sagina saginoides (L.) Britt. Arctic Pearl- wort. Rare; moist subalpine sites. 555. Silene douglasii Hook. Douglas Campion. Frequent; open forest understory and open slopes from middle elevations to subalpine. 1244. Stellaria jamesiana Torr. Sticky Chick- weed. Frequent; forest understory and open slopes at middle elevations. 125 (UT and UTC). Stellaria umbellata Turcz. Umbrella Star- wort. Rare; moist sites at middle elevations. Taye& Wall 1168. Celastraceae Pachistima myrsinites (Pursh) Raf. Moun- tain Lover. Frequent; shaded sites from the foothills to subalpine. 880. Chenopodiaceae Allenrolfea occidentalis (Wats.) Kuntze. lo- dinebush. Locally common in Skull Valley in saline soil. 1195. Atriplex canescens (Pursh) Nutt. Fourwing Saltbush. Frequent; valleys and lower foot- hills. 714. Atriplex confertifolia (Torr. & Frem.) Wats. Shadscale. Common; a dominant in the valleys and lower foothills. 1005. "Atriplex hortensis L. Garden Orach. Rare; collected only from a roadside in Tooele Val- ley. 1278. "Atriplex rosea L. Tumbling Orach. Rare; collected only from a roadside in Rush Val- ley. 749. Atriplex tridentata Kuntze. Three-toothed Saltbush. [A. nuttallii Wats.] Locally com- mon in Skull Valley in saline soil. 1198. "Bassia hyssopifolia (Pallas) Kuntze. Five- hook Bassia. Locally common in Skull Valley in saline soil. 1197. October 1983 Taye: Stansbury Mountains Flora 633 Ceratoides lanata (Pursh) J. T. Howell. Winterfat. [Eurotia lanata (Pursh) Moq.] In- frequent; valleys and lower foothills. 900. " Chenopodiurn album L. Lambsquarters. In- frequent; disturbed sites at low elevations. 597 (UTC). " Chenopodium botrys L. Jerusalem-oak. Rare; rocky sites at low elevations. 1160. Chenopodium fremontii Wats. Fremont Goosefoot. Common; open and wooded slopes from the valleys to middle elevations. 492. " Clienopoditim glaucum L. Oakleaf Goose- foot. Rare; dry pond in Skull Valley. 1273. Chenopodium hybridum L. Mapleleaf Goosefoot. Rare; beneath juniper in sandy soil. 1033. Chenopodium leptophyllum Nutt. Slimleaf Goosefoot. Rare; collected only ^rom a sandy area in Skull Valley. 1411. Grayia spinosa (Hook.) Moq. Spiny Hop- sage. Infrequent; valleys and foothills. 779. " Halogeton ghmeratus C. A. Mey. Haloge- ton. Common; disturbed sites in valleys. 959. Kochia americana Wats. Graymolly. In- frequent; valleys. 890. "Kochia scoparia (L.) Schrad. Belvedere Summer Cypress. Infrequent; disturbed sites at low elevations. 748. Salicornia europaea L. Marshfire Pickle- weed. [S. rubra A. Nels.] Locally common at Big Spring in saline soil. 1308. Salicornia pacifica Standi, var. utahensis (Tidestr.) Munz. Utah Pickleweed. [S. utah- ensis Tidestr.] Locally common at Big Spring in saline soil. 1340. "Salsola iberica Sennen & Pau. Russian Thistle. [S. kali L.] Common; disturbed sites at low elevations. 300 (UT). Sarcobatus vermiculatus (Hook.) Torr. Greasewood. Common; a dominant in lower portions of valleys. 1276. Suaeda occidentalis Wats. Western Seep- weed. Locally common near Big Spring in Skull Valley. 1424. Suaeda torreyana Wats. Bush Seepweed. Frequent; saline valleys. 1189. Convolvulaceae "Convolvulus arvensis L. Field Morning- glory. Infrequent; disturbed sites at low ele- vations. 95 (UT and UTC). Cressa truxillensis H. B. K. Locally com- mon near Big Spring in saline soil. 1404. Cornaceae Cornus stolonifera Michx. Red-osier Dog- wood. Frequent; along streams at middle ele- vations. 1046. Crassulaceae Sedum debile Wats. Stonecrop. Infrequent; shaded and open rocky sites from middle ele- vations to timberline. 1245. Sedum lanceolatum Torr. Lanceleaf Stone- crop. Rocky sites from middle elevations to alpine. 323 (UTC). Cuscutaceae Cuscuta denticulata Engelm. Desert Dod- der. Parasitic on Artemisia tridentata and Chrysothamnus nauseosus in Skull Valley, 1421. Elaeagnaceae "Elaeagnus angustifolia L. Russian Olive. Not collected but seen cultivated at Willow Springs and Delle Ranch. Euphorbia seed Spurge. Euphorbia cola (Parish) mon on sand Euphorbia Spurge. Dry hills. 997. Euphorbiaceae glyptosperma Engelm. Ridge- Dry slopes in the foothills. 679. ocellata Dur. & Hilg. var. areni- Jeps. Spurge. Rare; locally com- dunes in Skull Valley. 1203. serpyllifolia Pers. Thymeleaf slopes in the valleys and foot- Fabaceae Astragalus argophyllus Nutt. var. martinii Jones. Silver-leaved Milkvetch. Collected only from an open mid-elevation slope in East Hickman Canyon. 648. Astragalus beckwithii T. & G. Beckwith Milkvetch. Common; dry slopes of valleys and foothills. 782. Astragalus ceramicus Sheld. Painted Milk- vetch. Rare; sandy foothills. 488. 634 Great Basin Naturalist Vol. 43, No. 4 Astragalus cibarius Sheld. Browse Milk- vetch. Common; open slopes in the foothills. 886. Astragalus convallarius Greene var. con- vallarius. Lesser Rushy Milkvetch. Frequent; open slopes from the foothills to medium ele- vations. 647. Astragalus eurekensis Jones. Eureka Milk- vetch. Common; valley benches and foothills. 767. Astragalus geyeri Gray. Geyer Milkevetch. Infrequent; locally common in sandy areas. 814. Astragalus kentrophyta Gray var. implexus (Canby) Bameby. Mountain Kentrophyta. In- frequent; locally common on open rocky slopes near timberline and above. 965. Astragalus lentiginosus Dougl. var. pohlii Welsh & Bameby. Infrequent; greasewood communities in Skull Valley. 1332. Astragalus mollissimus Torr. var. thomp- sonae (Wats.) Bameby. Woolly Locoweed. Infrequent; sandy foothills. 792. Astragalus tenellus Pursh. Pulse Milkvetch. Infrequent; meadows and open rocky slopes from middle elevations to timberline. 586. Astragalus utahensis (Torr.) T. & G. Utah Milkvetch. Frequent; open slopes at low ele- vations. 788. Hedysarum boreale Nutt. var. boreale. Northern Sweetvetch. Open slopes from the foothills to middle elevations. 947. Lathyrus brachycalyx Rydb. var. brachyca- lyx. Rydberg Sweetpea. Infrequent; rocky foothills. 774. Lathyrus lanzwertii Kellogg var. lan- zwertii. Lanzwert Sweetpea. Infrequent; open to wooded subalpine slopes. 1451. Lathyrus pauciflorus Fern. var. utahensis (Jones) Peck. Utah Sweetpea. Open to wood- ed slopes from the foothills to middle eleva- tions. 64 (UT and UTC). Lupinus argenteus Pursh var. rubricaulis (Greene) Welsh. Silvery Lupine, frequent; meadows and open slopes from middle eleva- tions to timberline. 184. Lupinus pusillus Pursh var. intermontanus (Heller) C. P. Sm. Dwarf Lupine. Infrequent; locally common in sandy areas. 815. Lupinus sericeus Pursh var. sericeus. Silky Lupine. Open to wooded slopes at middle elevations. 1284. °Medicago lupilina L. Black Medick. In- frequent; disturbed sites at low elevations. 869. "Medicago sativa L. Alfalfa. Disturbed sites (and cultivated) at low elevations. 257 (UT). "Melilotus alba Medicus. White Sweet- clover. Disturbed sites at low elevations. 669 (UTC). "Melilotus officinalis (L.) Pallas. Yellow Sweetclover. Disturbed sites at low eleva- tions. 98 (UT and UTC). Psoralea lanceolata Pursh. var. stenos- tachys (Rydb.) Welsh. Dune Scurfpea. Fre- quent; sandy areas at low elevations. 479. °Trifolium fragiferum L. Strawberry Clo- ver. Collected only from streamside in South Willow Canyon. 1041b. "Trifolium repens L. White Clover. Fre- quent; wet places from the foothills to sub- alpine. 853. Vicia americana Muhl. var. americana. American Vetch. Infrequent; dry foothills. 460. Fumariaceae Dicentra uniflora Kellogg. Steer's Head. Rare; open woodland on mid-elevation slope in Mining Fork in early spring. 22 (UT and UTC). Gentianaceae Centaurium exaltatum (Griseb.) Wight. Western Centaury. Seen but not collected by K. Thome at springs in Skull Valley. Swertia radiata (Kell.) Kuntze. Green Gen- tian. [Frasera speciosa Dougl.] Frequent; mesic open and wooded slopes from middle elevations to alpine. 948. Geraniaceae "Erodium cicutarium (L.) L'Her. Heronsbill. Frequent; disturbed sites at low elevations. 27 (UT). Geranium parryi (Engelm.) Heller. Parry Geranium. Infrequent; locally common on rocky (quartzite) slopes between 2200 m and 3000 m in elevation. Taye & Wall 1213. Jones and Jones (1943) state that the petal color for this species is pale to deep rose- purple. Plants collected from Stansburys, and October 1983 Taye: Stansbury Mountains Flora 635 also from the Canyon Range 80 km to the south (S. Goodrich, pers. comm. 1981), have white petals and thus are possibly worthy of taxonomic distinction. Geranium richardsonii Fisch. & Trautv. Richardson Geranium. Frequent; near streams at middle elevations. 1204. Geranium viscosissimum Fisch. & Mey. Sticky Geranium. Frequent; mesic, mostly open slopes at middle elevations. 936. Grossulariaceae Rihes aureum Pursh. Golden Currant. In- frequent; valleys and foothills in drainage bottoms. 818. Ribes cereum Dougl. Wax Currant. Com- mon; open slopes, near streams, and in forest understory from the foothills to subalpine. 877. Ribes inerme Rydb. Whitestem Goose- berry. Rare; streamside in South Willow Can- yon. 1234. Ribes montigenum McClatchie. Alpine Prickly Currant. Common; forest understory, meadows, and open slopes from middle ele- vations to alpine. 960. Ribes velutinum Greene var. velutinum. Desert Gooseberry. Infrequent; dry rocky foothills. 1003. Ribes viscosissimum Pursh. Sticky Currant. Frequent; forest understory at middle eleva- tions. 1030. Ribes wolfii Rothr. Wolf's Currant. Fre- quent; forest understory at middle elevations. 991. Hydrophyllaceae Hydrophyllum occidentale (Wats.) Gray. Western Waterleaf. Frequent; mesic, shaded to open sites from the foothills to middle ele- vations. 531. Nama densum Lemmon. Matted Nama. Rare; sandy area of Skull Valley. 837. Phacelia crenulata Torr. Scorpionweed. Rare; juniper community on Salt Mountain. 1153. Phacelia hastata Dougl. Infrequent; open rocky areas from middle elevations to sub- alpine. 575. Phacelia incana Brand. Seen only in rocky foothills near Big Spring. 1014. Phacelia ivesiana Torr. Infrequent; sandy areas at low elevations. 791. Phacelia linearis (Pursh) Holz. Threadleaf Scorpionweed. Infrequent; dry slopes from low to middle elevations. 789. Juglandaceae "Juglans nigra L. Black Walnut. Rare; aban- doned ranch in Big Hollow. 741. Lamiaceae Agastache urticifolia (Benth.) Kuntze. Gi- ant Hyssop. Frequent; open and wooded slopes at middle elevations. 175 (UT and UTC). "Marrubium vulgare L. Common Horehound. Infrequent; disturbed sites at low elevations. 302. Mentha arvensis L. Field Mint. Collected only from streamside near mouth of Spring Canyon. 1137. °Mentha piperita L. Peppermint. Collected only from streamside at mouth of South Wil- low Canyon. 1186. "Mentha spicata L. Spearmint. Collected only from marshy area in the foothills. Hardy 130. Monardella odoratissima Benth. In- frequent; open rocky slopes from middle ele- vations to subalpine. 228 (UT and UTC). "Nepeta cataria L. Catnip. Infrequent; dry to moist, sometimes disturbed sites at low elevations. 735. Linaceae Linum perenne L. Wild Blux Flax. [L. lew- isii Pursh] Frequent; open slopes from the foothills to near timberline. 937. Loasaceae Mentzelia alhicaulis Dougl. White Blazing Star. Rare; sandy areas in foothills. 484. Mentzelia laevicaulis (Dougl.) T. & G. Blazing Star. Infrequent; dry slopes in foot- hills. 1267. Malvaceae Iliamna rivularis (Dougl.) Greene. Moun- tain Hollyhock. Infrequent; open and wood- ed slopes at middle elevations. 1109. 636 Great Basin Naturalist Vol. 43, No. 4 "Malva neglecta Wallr. Cheeseweed. Dis- turbed sites at low elevation. 96 (UT and UTC). Sidalcea neomexicana Gray. New Mexico Checkermallow. Rare; near spring in Skull Valley. 1378. Sphaeralcea grossulariifolia (H. & A.) Rydb. Gooseberryleaf Globemallow. Com- mon; dry slopes of valleys and lower foothills. 860. Moraceae "Morns alba L. White Mulberry. In- frequent; cultivated at ranches. Taye & Dill- man 1416. Nyctaginaceae Abronia fragrans Nutt. Snowball Sand Ver- bena. Frequent; sandy areas at low eleva- tions. 838. Mirabilis linearis (Pursh) Heimerl. Narrow- leaved Four-o'clock. [Oxybaphus linearis (Pursh) Robins.] Rare; dry valley benches. 1047. Tripterocalyx micranthus (Torr.) Hook. Sandpuff. Rare; sand dunes in Skull Valley. 1391. Oleaceae "Syringa vulgaris L. Common Lilac. Rare; abandoned ranch in Big Hollow. 743. Onagraceae Camissonia boothii (Dougl.) Raven ssp. alyssoides (H. & A.) Raven. [Oenothera alys- soides H. & A.] Locally common in the foot- hills. 928. Camissonia minor (A. Nels.) Raven. [Oeno- thera minor (A. Nels.) Munz] Known from one collection from the foothills. Flowers 1222 (UT). The lectotype was collected by S. Watson from Stansbury Island in 1869 (Ra- ven 1969). Camissonia parvula (Nutt.) Raven. [Oeno- thera contorta (Dougl.) Kearney var. flexuosa (A. Nels.) Munz] Infrequent; sandy areas of foothills. 810. Camissonia scapoidea (T. & G.) Raven. [Oenothera scapoidea T. & G.] Infrequent; valleys and foothills. 957. Circaea alpina L. Enchanter's Nightshade. Rare; understory of Douglas fir community in South Willow Canyon. 1106. Epilobiimi alpinwn L. Alpine Willowherb. [£. hornemannii Reichenb.] Rare; wet places at middle elevations. Taye & Wall 1164. Epilobium angustifolium L. Fireweed. Common; streamside at middle elevations. 237 (UT). Epilobium brachycarpum Presl. Autumn Willowherb. [£. paniculatum Nutt.] Fre- quent; dry slopes in the foothills. 304. Epilobium canum (Greene) Raven. Garret Firechalice. [Zauschneria garrettii A. Nels.] Frequent; dry rocky slopes in juniper zone. 690. Epilobium ciliatum Raf. Frequent; stream side from low to middle elevations. 261. Gaura parviflora Dougl. Lizard Tail. Rare; roadside in Skull Valley. 1420. Gayophytum ramosissimum Nutt. Hair- stem Groundsmoke. Collected only from a sandy area in the foothills. 1035. Oenothera caespitosa Nutt. Tufted Evening Primrose. Frequent; dry foothills. 866. Oenothera liookeri T. & G. Hooker Eve- ning Primrose. Rare; streamside near mouth of Spring Canyon. 1141. Oenothera pallida Lindl. Pale Evening Primrose. Infrequent; dry, sometimes sandy areas in the foothills. 477. Orobanchaceae Orobanche corymbosa (Rydb.) Ferris. Flat- topped Broomrape. Rare; apparently para- sitic on Artemisia spinescens and Artemisia tridentata in Skull Valley. 950. Orobanche fasiculata Nutt. Clustered Broomrape. Rare; apparently parasitic on Ar- temisia tridentata in juniper woodland. 1075. Papaveraceae Argemone munita Dur. & Hilg. Prickly Poppy. Frequent; valleys and foothills, usual- ly in disturbed sites. 912. "Papaver orientale L. Oriental Poppy. Rare; abandoned ranch in Big Hollow. 745. Plantaginaceae "Plantago lanceolata L. Buckhorn Plantain. Collected only from roadside in South Wil- low Canyon. 1110. October 1983 Taye: Stansbury Mountains Flora 637 "Plantago major L. Common Plantain. Moist sites in the foothills. 626. Polemoniaceae CoUomia grandiflora Dougl. Large-flow- ered Collomia. Collected only from juniper community in Whiterocks Canyon. 883. Collo7nia linearis Nutt. Narrowleaf Col- lomia. Infrequent; dry open slopes at middle elevations. 868. Gilia aggregata (Pursh) Spreng. Scarlet Cilia. Frequent; open and wooded slopes from the foothills to middle elevations. 932. Gilia inconspicua (Smith) Sweet. Shy Gilia. [G. sinuata Dougl.] Frequent; dry slopes at low elevations. 483. Gilia leptotneria Gray. Infrequent; sandy areas at low elevations. 845. Gilia polycladon Torr. Rare; sandy area in Skull Valley. 956. Leptodactylon pungens (Torr.) Nutt. Prickly Phlox. Frequent; dry foothills in rocky or sandy soil. 797. Leptodactylon watsonii (Gray) Rydb. Prickly Gilia. Infrequent; limestone outcrops in the foothills. 546. Microsteris gracilis (Hook.) Greene. Locally frequent on dry slopes from low to middle elevations. 802. Phlox hoodii Rich. Hood's Phlox. Frequent; valley benches and foothills. 493. Phlox longifolia Nutt. Longleaf Phlox. Common; dry foothills. 395. Phlox pulvinata (Wherry) Cronq. Cushion Phlox. [P. caespitosa Nutt.] Infrequent; rocky slopes near timberline and above. 971. Polemonium foliosissimum Gray. Leafy Ja- cob's Ladder. Meadows at middle elevations. 993. Polemonium pulcherrimum Hook. var. del- icatum (Rydb.) Cronq. Skunkleaf. [P. delica- tum Rydb.] Infrequent; rocky subalpine slopes. 985. Polygonaceae Eriogonum brevicaule Nutt. var. laxifolium (T. & G.) Reveal. Shortstem Wild Buck- wheat. Frequent; dry slopes from low to middle elevations, often growing from cracks in rock outcrops. 432. Eriogonum cernuum Nutt. var. cernuum. Nodding Buckwheat. Frequent; valleys and foothills, usually in sandy areas. 672. Eriogonum grayi Reveal. Frequent; rocky slopes near timberline and above. 534. Eriogonum heracleoides Nutt. Wyeth Buck- wheat. Freqvient; open slopes from middle elevations to subalpine. 614. Eriogonum hookeri Wats. Rare; sandy areas in the foothills. 1150. Eriogonum kearneyi Tidestr. Infrequent; locally common in sandy areas at low eleva- tions. 675. Eriogonum microthecum Nutt. var. laxiflo- rum Hook. Slenderbush Buckwheat. Collect- ed only from a sandy area in juniper zone. Taye et al. 565 (UTC). Eriogonum ovalifolium Nutt. var. ovalifo- lium. Cushion Buckwheat. Frequent; valleys and foothills, often in sandy areas. 839. Eriogonum racemosum Nutt. Redroot Buckwheat. Infrequent; open slopes at middle elevations. 1078. Eriogonum umbellatum Torr. var. desereti- cum Reveal. Sulfurflower. Open slopes from middle elevations to timberline. 732. Eriogonum umbellatum Torr. var. suh- aridtim S. Stokes. Frequent; sandy areas of foothills. 1037. Oxyria digyna (L.) Hill. Mountain Sorrel. Frequent; rocky, often moist sites above 2600 m. Taye & Wall 1446. Polygonum aviculare L. Prostrate Knot- weed. Collected only from roadside in Tooele Valley. 596 (UTC). Polygonum douglasii Greene. Douglas Knotweed. Frequent; open and wooded slopes at middle elevations. 1084. Polygonum sawatchense Small. Sawatch Knotweed. Collected only from a sandy area in juniper zone. 1036. "Rumex crispus L. Curly Dock. Frequent; mesic, often disturbed sites from the foothills to middle elevations. 557. Rumex salicifolius Weinm. Willow Dock. Streamside at middle elevations. 558. Rumex venosus Pursh. Wild Begonia. In- frequent; locally common in sandy areas at low elevations. 832. Portulacaceae Claytonia lanceolata Pursh. Lanceleaf Springbeauty. Common; moist soil at middle elevations in springtime. 381. 638 Great Basin Naturalist Vol. 43, No. 4 Lewisia pygmaea (Gray) Robins. Least Lewisia. Infrequent; locally common in meadows near timberline. 966. Montia perfoliata (Donn) Howell. Miner's Lettuce. [Claytonia perfoliata Donn] In- frequent; streamside at low to middle eleva- tions. 864. Primulaceae Androsace septentrionalis L. Northern Rock Jasmine. Frequent; rocky slopes and meadows above 2600 m, usually in moist soil. 145 (UTC). Glaux maritima L. Sea Milkwort. In- frequent; near springs and streams at low ele- vations. 829. Primula parryi Gray. Parry's Primrose. In- frequent; moist subalpine sites. 207 (UT and UTC). Pyrolaceae Pyrola secunda L. Sidebells Wintergreen. Infrequent; conifer understory at middle ele- vations. Taye & Wall 1214. Ranunculaceae Aconitum columbianuni Nutt. Columbia Monkshood. Frequent; streamside at middle elevations. Taye & Wall 1169. Actaea rubra (Ait.) Willd. Western Bane- berry. Frequent; meadows and forest under- story from middle elevations to subalpine. 1074. Aquilegia caerula James. Colorado Colum- bine. Infrequent; streamside, in meadows, and in open forest understory from middle elevations to subalpine. 992. Aquilegia formosa Fisch. Red Columbine. Streamside at middle elevations. 559. Delphinium nuttallianum Pritz. Nuttal Larkspur. [D. nelsonii Greene] Common; open slopes and forest understory from the valley benches to 2900 m. 394. Delphinium occidentale Wats. Duncecap Larkspur. Meadows and open forest under- story at middle elevations. 180 (UT and UTC). Ranunculus cymbalaria Pursh. Shore But- tercup. Common; wet areas from the valleys to subalpine. 825. Ranunculus glaberrimus Hook. Sagebrush Buttercup. Rare; sagebrush community in North Willow Canyon. Taye & Taye 387. Ranunculus inamoenus Greene. In- frequent; moist meadows and streamside from middle elevations to subalpine. 190 (UTC). Ranunculus juniperinus Jones. Locally common in the foothills in early spring. Taye & Kass 1323. "Ranunculus testiculatus Crantz. Bur But- tercup. Common; disturbed sites at low ele- vations. 1314. Thalictrum fendleri Engelm. Fendler Meadowrue. Common; meadows and forest understory at middle elevations. 986. Rhamnaceae Ceanothus martinii Jones. Rare; sagebrush community in juniper zone at southern end of range. 525. Ceanothus velutinus Dougl. Snowbrush. Infrequent; open and wooded slopes at middle elevations. 940. Rosaceae Amelanchier alnifolia Nutt. Serviceberry. Common; mesic, open and wooded slopes from the foothills to middle elevations. 400. Cercocarpus ledifolius Nutt. Curl-leaf Mountain Mahogany. Common; a dominant on dry slopes in upper juniper zone. 1386. Cercocarpus ledifolius Nutt. X C. mon- tanus Raf. Rare; seen only in juniper zone in Vickory Canyon. 464. Cercocarpus montanus Raf. Birchleaf Mountain Mahogany. Rocky slopes at middle elevations. 463 (UTC). Chamaebatiaria millefolium (Torr.) Maxim. Fern Bush. Infrequent; dry rocky slopes be- low 2300 m in elevation, often growing from limestone outcrops. 1114. Cowania mexicana D. Don var. stansbu- riana (Torr.) Jeps. Stansbury Cliffrose. Com- mon; dry slopes at low elevations. 911. Crataegus douglasii Lindl. var. rivularis (Nutt.) Sarg. Douglas Hawthorn. [C. rivularis Nutt.] Rare; along Clover Creek at southern end of the range. 1384. Fragaria vesca L. var. bracteata (Heller) Davis. Woods Strawberry. Infrequent; mead- October 1983 Taye: Stansbury Mountains Flora 639 ows and open forest understory at middle ele- vations. 541 (UTC). Geum macrophyllum Willd. var. per- incisum (Rydb.) Raup. Largeleaf Avens. Fre- quent; streamside at middle elevations. 497. Getim rossii (R. Br.) Ser. var. turbinatum (Rydb.) Hitchc. Alpine Avens. Frequent; lo- cally common in subalpine meadows and on rocky alpine slopes, often near persistent snow patches. 1067. Geum trifhrum Pursh var. ciliatum (Pursh) Fassett. Prairie Smoke. Rare; open, mesic north-facing slope at 1950 m in Davenport Canyon. 512. Holodiscus dumosus (Nutt.) Heller. Bush Oceanspray. Frequent; dry rocky slopes from low to middle elevations. 1011. Ivesia gordonii (Hook.) T. & G. Gordon Ivesia. Infrequent; open rocky slopes from middle elevations to alpine. Taye & Wall 1448. Petrophytum caespitosum (Nutt.) Rydb. Tufted Rockmat. [Spiraea caespitosa Nutt.] Infrequent; locally common on limestone outcrops from low to middle elevations. 1185. Physocarpus malvaceus (Greene) Kuntze. Mallow Ninebark. Frequent; open slopes and forest understory at middle elevations. 949. Potentilla diversifolia Lehm. Varileaf Cinquefoil. Collected only from a cirque meadow in Antelope Canyon. 976. Potentilla glandulosa Lindl. Gland Cinque- foil. Common; meadows, open forest under- story, and on rocky slopes from middle eleva- tions to subalpine. 990. Potentilla gracilis Dougl. Soft Cinquefoil. Meadows and open slopes from middle eleva- tions to subalpine. 625. Potentilla ovina Macoun. Sheep Cinque- foil. [P. wyomingensis A. Nels.] Infrequent; meadows and open rocky slopes from sub- alpine to alpine. 970. Primus virginiana L. var. melanocarpa (A. Nels.) Sarg. Chokecherry. Common; stream- side and on mesic slopes from the foothills to middle elevations. 994. Purshia tridentata (Pursh) DC. Antelope Bitterbrush. Common; dry foothills. 811. Purshia tridentata (Pursh) DC. X Cowania mexicana D. Don. Rare; rocky foothills near Big Spring. 1018. Rosa nutkana Presl. Bristly Nootka Rose. Mesic, open and wooded slopes at middle elevations. 1285. "Rosa rubiginosa L. Sweetbrier. Rare; streamside near abandoned power station at mouth of South Willow Canyon. 903. Rosa woodsii Lindl. Wood's Rose. Com- mon; drainage bottoms and other generally mesic sites at middle elevations. 147 (UT and UTC). Rubus idaeus L. ssp. melanolasius (Dieck) Focke. Wild Red Raspberry. Frequent; forest understory and on rocky slopes from middle elevations to subalpine. 201. Rubus parviflorus Nutt. Western Thimble- berry. Rare; one streamside collection from Dry Lake Fork. Neese 9662. " Sanguisorba minor Scop. Burnet. Locally frequent in disturbed sites at low elevations. 1142. Rubiaceae Galium aparine L. Catchweed Bedstraw. Open forest understory and along streams from the foothills to middle elevations. 117 (UT and UTC). Galium bifolium Wats. Twinleaf Bedstraw. Open forest understory at middle elevations. 509 (UTC). Galium multiflorum Kellogg. Shrubby Bedstraw. Rare; dry rocky slopes in the foot- hills near Timpie. 1009. Galium triflorum Michx. Sweetscented Bedstraw. Frequent; streamside and open for- est understory at middle elevations. 1107. Salicaceae Populus angustifolia James. Narrowleaf Cottonwood. Common; a dominant along streams from the foothills to middle eleva- tions. 401. °Populus fremontii Wats. Fremont Cotton- wood. Rare; abandoned ranch in Big Hollow. 739 (UTC). "Populus nigra L. Lombardy Poplar. Culti- vated at several ranches. 738 (UTC). Populus tremuloides Michx. Quaking As- pen. Common; a dominant along streams at low elevations and on mesic slopes to moder- ately high elevations. 399. Salix amygdaloides Anderss. Peachleaf Willow. Streamside in the foothills. 1335. 640 Great Basin Naturalist Vol. 43, No. 4 Salix exigiia Nutt. Sandbar Willow. Com- mon; streamside from low to middle eleva- tions. 443. Salix rigida Muhl. Yellow Willow. [S. lutea Nutt.] Common; streamside from low to middle elevations. 438. Salix scouleriana Barratt. Scouler Willow. Locally frequent along streams at middle ele- vations. 444. Santalaceae Comandra umbellata (L.) Nutt. Bastard Toadflax. Locally frequent in dry, sometimes sandy areas in the foothills. 914. Saxifragaceae Heuchera parvifolia Nutt. Littleleaf Alum- root. Open forest understory and open rocky areas from middle elevations to subalpine. 533. Heuchera nibescens Torr. Red Alumroot. Streamside, open forest understory, and in open rocky areas from middle elevations to subalpine. 1100. Lithophragjna glabra Nutt. Fringecup Woodland Star. [L. hulhifera Rydb.] Moist sites at middle elevatons. 385a. Lithophragma parviflora (Hook.) Nutt. Smallflower Woodland Star. Locally frequent at middle elevations in open forest under- story and other shaded sites. 870. Mitella stauropetala Piper. Miterwort. Common; streamside and open forest under- story at middle elevations. 878. Saxifraga debilis Engelm. Pygmy Saxi- frage. Rare; below melting snowpatch in cirque of Big Creek Canyon. Taye & Wall 1170. Saxifraga odontoloma Piper. Brook Saxi- frage. [S. arguta D. Don] Infrequent; stream- side at middle elevations. 1206. Saxifraga rhomboidea Greene. Dia- mondleaf Saxifrage. Infrequent; near melting snow on cirque walls and in subalpine-alpine meadows. 503. Scrophulariaceae Castilleja applegatei Fern. var. viscida (Rydb.) Owenby. Sticky Indian Paintbrush. [C. viscida Rydb.] Frequent; open rocky slopes from middle elevations to alpine. 529. Castilleja chromosa A. Nels. Desert Indian Paintbrush. Common; open slopes from the valleys to middle elevations. 407. Castilleja linariifolia Benth. Narrowleaf In- dian Paintbrush. Locally frequent on open mesic slopes at lower elevations. Ill (UT and UTC). Castilleja rhexifolia Rydb. Splitleaf Indian Paintbrush. Infrequent; open slopes from sub- alpine to alpine. 579. Castilleja sulphurea Rydb. Sulphur Indian Paintbrush. Infrequent; openings in conifer forest at middle elevations. 1112. Collinsia parviflora Dougl. Blue-eyed Mary. Mesic slopes from low to middle eleva- tions. 30 (UT). Mimulus breweri (Greene) Rydb. Brewer's Monkeyflower. Rare; near subalpine spring in Dry Lake Fork. 613. Mimulus floribundus Dougl. Rare; near mid-elevation spring in Big Creek Canyon. Taye & Wall 1165. Mimulus guttatus Fisch. Yellow Mon- keyflower. Common; streamside from low to middle elevations. 857. Mimulus lewisii Pursh. Lewis Mon- keyflower. Frequent; streamside at middle elevations. 153 (UT and UTC). Orthocarpus tolmiei H. & A. Tolmie Owl Clover. Locally frequent on open slopes at middle elevations. 206 (UT and UTC). Penstemon cyananthus Hook. Wasatch Penstemon. Frequent; open and wooded slopes from middle elevations to subalpine. 606. Penstemon humilis Nutt. Low Penstemon. Common; rocky slopes from middle eleva- tions to alpine. 588. Penstemon whippleanus Gray. Whipple Penstemon. Infrequent; moist sites and co- nifer understory from middle elevations to subalpine. 195 (UT and UTC). Scrophularia lanceolata Pursh. Lanceleaf Figwort. Known from one roadside collection in drainage bottom of East Hickman Canyon. 635 (UTC). Synthyris pinnatifida Wats. Featherleaf Kittentails. Infrequent; locally common on rocky subalpine and alpine slopes. 535. "Verbascum thapsus L. Flannel Mullein. Frequent; disturbed sites at low elevations. 199 (UT and UTC). October 1983 Taye: Stansbury Mountains Flora 641 °Verbascum virgatum Stokes. Wand Mul- lein. Rare; disturbed sites at low elevations. 713 (UTC). Veronica americana Schwein. American Brooklime. Frequent; streamside at middle elevations. 1049. ° Veronica biloba L. Bilobed Speedwell. Dis- turbed sites at low elevations. 112b (UTC). Veronica peregrina L. Purslane Speedwell. Rare; streamside near mouth of Spring Can- yon. 1128. Veronica serpyllifolia L. Thyme-leaved Speedwell. Collected only from streamside in North Willow Canyon. 439. Solanaceae Lycium andersonii Gray. Anderson Wolf- berry. Infrequent; dry rocky .foothills at northern end of the range. 1328. Nicotiana atteniiata Torr. Coyote To- bacco. Locally frequent in disturbed and sandy sites at low elevations. 1088. Physalis virginiana Mill. Virginia Ground- cherry. [P. longifolia Nutt.] Seen only at roadside near South Mountain. 600 (UTC). Tamaricaceae "Tamarix ramosissima Ledeb. Salt Cedar Tamarisk. Rare; wet valley sites. 1144. Ulmaceae Celtis reticulata Torr. Netleaf Hackberry. Rare; near streamchannels in the foothills. 1140. "Uhnus pumila L. Siberian Elm. Rare; near abandoned power station at mouth of South Willow Canyon. 904. Urticaceae Urtica dioica L. ssp. gracilis (Ait.) Seland. Stinging Nettle. Common; streamside and open forest understory at middle elevations. 181 (UTC). Valerianaceae Valeriana acutiloba Rydb. var. pubicarpa (Rydb.) Cronq. Sharpleaf Valerian. In- frequent; rocky open slopes near timberline and above. 984. Valeriana occidentalis Heller. Western Va- lerian. Infrequent; meadows at middle eleva- tions. 446. Verbenaceae Verbena bracteata Lag. & Rodr. Bracted Vervain. Frequent; dry to moist, usually dis- turbed sites at low elevations. 301. Violaceae Viola adunca Sm. var. adunca. Mountain Blue Violet. Common; open forest understory and streamside at middle elevations. 590. Viola adunca Sm. var. bellidifolia (Greene) Harr. Rare; rock ledges of cirque in Dry Lake Fork. 191 (UT). Viola nephrophylla Greene. Bog Violet. Collected only from streamside at Condie Meadows. 826. Viola nuttallii Pursh. Yellow Prairie Violet. Mesic forest openings at middle elevations. 447. Viola purpurea Kellogg. Goosefoot Violet. Frequent; open slopes and open forest under- story from the foothills to subalpine. 372. Viscaceae Phoradendron jtiniperinum Engelm. Juni- per Mistletoe. Infrequent; parasitic on Juni- perus osteosperma in the foothills. 360. Class Liliopsida Cyperaceae Carex atrata L. var. erecta W. Boott. Black Sedge. Infrequent; subalpine meadows and open slopes near timberline. 702. Carex aurea Nutt. Golden Sedge. Locally common near streams in the foothills. 820. Carex douglasii F. Boott. Douglas Sedge. Collected only from streamside in Davenport Canyon. 61 (UT and UTC). Carex geyeri F. Boott. Elk Sedge. In- frequent; forest understory and open mesic slopes at middle elevations. 516. Carex haydeniana Olney. Cloud Sedge. In- frequent; locally common in subalpine mead- ows. 1246. 642 Great Basin Naturalist Vol. 43, No. 4 Carex hoodii F. Boott. Hood Sedge. Com- mon; meadows and open slopes from middle elevations to subalpine. 511. Carex lenticularis Michx. Locally common in wet areas at middle elevations. 412. Carex microptera Mackenzie. Smallwing Sedge. Meadows and streamside at middle elevations. 413. Carex nebrascensis Dewey. Nebraska Sedge. Common; streamside at low to middle elevatons. 440. Carex nova Bailey. Collected only from edge of South Willow Lake. Taye & Wall 724. Carex pachystachya Cham. Chamisso Sedge. Collected only from cirque in Dry Lake Fork. 288 (UT and UTC). Carex petasata Dewey. Collected only from an open, mesic mid-elevation slope in Davenport Canyon. 517. Carex phaeocephala Piper. Dunhead Sedge. Locally common in subalpine mead- ows. 703. Carex praegracilis W. Boott. Collected only from streamside in Davenport Canyon. 424. Carex raynoldsii Dewey. Raynold's Sedge. Locally common in meadows and along streams at middle elevations. 552. Carex rossii F. Boott. Ross Sedge. Conifer understory and meadows from middle eleva- tions to subalpine. 705. Carex vallicola Dewey. Valley Sedge. Meadows and open forest understory from middle elevations to subalpine. 580. Eleocharis palustris (L.) R. & S. Common Spikerush. [£. macrostachya Britt.] Locally common in wet places from the valleys to middle elevations. 906. Eleocharis rostellata (Torr.) Torr. Beaked Spikerush. Locally common near streams in the foothills. 823. Scirpus acutus Muhl. Hardstem Bulrush. Wet places at low elevations. 827. Scirpus maritimus L. Alkali Bulrush. Rare; pond in Skull Valley. 1272. Scirpus pungens Vahl. Common Three- square. Seen only near stream at Condie Meadows. 851. Iridaceae Iris missouriensis Nutt. Western Iris. Rare; near spring in Skull Valley. 1374. Sisyrinchium idahoense Bickn. Idaho Blue- eyed Grass. Rare; near springs and streams at low elevations. 847. uncaceae Juncus arcticiis Willd. Wiregrass. [/. bal- ticus Willd.] Common; wet places at low ele- vations. 423. Juncus articulatus L. Jointed Rush. Rare; streamside at mouth of South Willow Can- yon. 907. Juncus bufonius L. Toad Rush. Infrequent; streamside at low to middle elevations. 1129. Juncus ensifolius Wikstr. var. brunnescens (Rydb.) Cronq. Locally common along streams at low elevations. 416. Juncus ensifolius Wikstr. var. ensifolius. Collected only from streamside in South Wil- low Canyon. 218 (UT and UTC). Juncus ensifolius Wikstr. var. montanus (Engelm.) C. L. Hitchc. Streamside at middle elevations. 1045. Juncus longistylis Torr. Longstyle Rush. Locally common in meadows and along streams in the foothills. 822. Juncus torreyi Gov. Torrey Rush. In- frequent; locally abundant in wet places at low elevations. 684. Luzula spicata (L.) DC. Spike Woodrush. Infrequent; locally common in meadows and on rocky slopes from subalpine to alpine. 967. Juncaginaceae Triglochin maritima L. Shore Arrowgrass. Rare; wet area in foothills at Condie Mead- ows. 819. Liliaceae Allium acuminatum Hook. Tapertip On- ion. Common; dry slopes and open forest un- derstory from the valleys to middle eleva- tions. 603. Allium nevadense Wats. Nevada Onion. Infrequent; valleys and foothills. Taye & Kass 1324. Calochortus nuttallii T. & G. Sego Lily. Common; open slopes from the valleys to subalpine. 476. The state flower of Utah. October 1983 Taye: Stansbury Mountains Flora 643 Disporum trachycarpum (Wats.) Benth. & Hook. Fairy Bells. Infrequent; forest under- story at middle elevations. 1182. Erythronium grandiflorum Pursh. Glacier Lily. Common; moist slopes from low to middle elevations in springtime. 873. Fritillaria atroptirpurea Nutt. Leopard Lily. Open slopes from middle elevations to near timberline. 961. Fritillaria pudica (Pursh) Spreng. Yellov^^ Bell. Locally common in springtime on moist slopes. 380. Smilacina stellata (L.) Desf. Starry Solo- mon-plume. Low to middle elevations in open forest understory and along streams. 817. Veratrum californicum Dur. False Helle- bore. Infrequent; locally common in wet places from middle elevations to subalpine. 242 (UT and UTC). Zigadenus elegans Pursh. Mountain Death Camas. Infrequent; subalpine meadows and open conifer understory near timberline. 654. Zigadenus paniculatus (Nutt.) Wats. Foot- hill Death Camas. Common; dry, mostly open slopes from the valleys to middle eleva- tions. 553. Orchidaceae Corallorhiza striata Lindl. Striped Coral- root. Infrequent; dense conifer forest at middle elevations. 1059. Goodyera oblongifolia Raf. Rattlesnake Plantain. Rare; rich understory of conifer- deciduous forest in South Willow Canyon. 264 (UT). Habenaria dilatata (Pursh) Hook. White Bog Orchid. Rare; wet area below South Wil- low Lake. Taye & Wall. 722. Habenaria unalescensis (Spreng.) Wats. Alaska Rein Orchid. Rare; near springs and in aspen understory at middle elevations. 1060. Poaceae °Agropyron cristatum (L.) Gaertn. Crested Wheatgrass. Common; disturbed areas and a dominant in portions of the valleys. 97 (UT and UTC). Agropyron dasystachyum (Hook.) Scribn. Thickspike Wheatgrass. Infrequent; sandy areas at low elevations. 831. ° Agropyron elongatum (Host) Beauv. Tall Wheatgrass. Locally common in disturbed areas at low elevations. 639. ° Agropyron intermedium (Host) Beauv. In- termediate Wheatgrass. Locally common in disturbed areas at low elevations. 629. Agropyron smithii Rydb. Western Wheat- grass. Collected only from a sandy area in a juniper community. 638. Agropyron spicatum (Pursh) Scribn. & Smith. Bluebunch Wheatgrass. Common; dry slopes from the foothills to middle elevations. 470. Agropyron trachycaulum (Link) Malte. Slender Wheatgrass. Common; streamside, and on open and wooded slopes from middle elevations to alpine. 609. Agrostis exarata Trin. Spike Redtop. Fre- quent; wet places from the foothills to sub- alpine. 1125a. "Agrostis stolonifera L. Redtop Bentgrass. Common; streamside from low to middle ele- vations. 734. " Alopercurus pratensis L. Meadow Foxtail. Rare; pasture near guard station in South Willow Canyon. 1063. Aristida purpurea Nutt. Three-awn. [A. longiseta Steud.] Infrequent; valleys and foot- hills. 601. " Arrhenatherum elatius (L.) Presl. Tall Oat- grass. Rare; pasture near guard station in South Willow Canyon. 640 (UTC). "Bromus brizaeformis Fisch. & Mey. Rattle- snake Chess. Rare; pasture near guard station in South Willow Canyon. 94 (UT and UTC). Bromus carinatus H. & A. California Brome. [B. marginatus Nees; B. polyanthus Scribn.] Common; streamside and open to wooded slopes from middle elevations to sub- alpine. 646. Bromus ciliatus L. Fringed Brome. In- frequent; open woods and meadows at middle elevations. 709. "Bromus commutatus Schrad. Hairy Chess. Seen only near streamside at mouth of South Willow Canyon. 1053. "Bromus inermis Leys. Smooth Brome. Dis- turbed sites at low elevations. 103 (UT and UTC). "Bromus japonicus Thunb. Japonese Chess. Disturbed sites at low elevations. 1398. "Bromus tectorum L. Cheatgrass. Common; a dominant species in disturbed areas, also occurring in undisturbed areas. 434. 644 Great Basin Naturalist Vol. 43, No. 4 Catabrosa aquatica (L.) Beauv. Brookgrass. Frequent; wet places from the foothills to subalpine. 908. Cinna htifolia (Trev.) Griseb. Drooping Woodreed. Common; streamside at middle elevations. 238. "Dactylis glomerata L. Orchard Grass. Common; streamside and open forest under- story from the foothills to middle elevations. 60 (UT and UTC). Danthonia intermedia Vasey. Timber Oat- grass. Rare; locally common in a cirque meadow in Dry Lake Fork. 352 (UT and UTC). Distichlis spicata (L.) Greene. Saltgrass. Locally common in the valleys. 1120. Echinochloa crusgalli (L.) Beauv. Barnyard Grass. Rare; streamside at mouth of Spring Canyon. 1124. Elymus cinereus Scribn. & Merr. Great Ba- sin Wildrye. Common; valleys to middle ele- vations, often in dry drainage bottoms. 1223. Elymus flavescens Scribn. & Smith. Yellow Wildrye. Infrequent; locally common in sandy areas of Skull Valley and nearby foot- hills. 637. Elymus glaucus Buckl. Blue Wildrye. Streamside and open forest understory at middle elevations. 1260. "Elymus junceus Fisch. Russian Wildrye. Infrequent; disturbed sites at low elevations. 861. Elymus triticoides Buckl. Creeping Wild- rye. Rare; near spring in Skull Valley. 1380. "Festuca arundinacea Schreb. Reed Fescue. Seen only at streamside in Davenport Can- yon. 518. Festuca ovina L. var. brevifolia (R. Br.) Wats. Alpine Fescue. Frequent; meadows and rocky slopes above 2800 m. 581. Glyceria striata (Lam.) Hitchc. Fowl Mannagrass. Frequent; wet places at middle elevations. 441. Hikiria jamesii (Torr.) Benth. Galleta. In- frequent; dry slopes at low elevations. 910. Hordeum brachyantherum Nevski. Mead- ow Barley. Infrequent; wet places at low to medium elevations. 340 (UT). Hordeum jubatum L. Foxtail Barley. Fre- quent; mostly in disturbed sites at low eleva- tions. 591. "Hordeum murinum L. Infrequent; dis- turbed sites at low elevations. 905. Leucopoa kingii (Wats.) W. A. Weber. Spikegrass. [Hesperochloa kingii (Wats.) Rydb.] Common; dry meadows and rocky slopes from middle elevations to alpine. 450. "Lolium multiflorum Lam. Italian Ryegrass. Rare; pasture near guard station in South Willow Canyon. 112 (UT). Melica bulbosa Geyer. Oniongrass. Locally frequent on open slopes at middle elevations. 607. Muhlenbergia asperifolia (Nees & Meyen) Parodi. Scratchgrass Muhly. Rare; streamside at Condie Meadows. 1093. Oryzopsis hymenoides (R. & S.) Ricker. In- dian Ricegrass. Common; dry slopes from the foothills to middle elevations. 1032. Phletim alpinum L. Alpine Timothy. In- frequent; moist subalpine sites. Taye & Wall 1178. "Fhleum pratense L. Timothy. Collected only from streamside in South Willow Can- yon. 268 (UT). Phragmites australis (Cav.) Trin. Common Reed. [P. communis Trin.] Infrequent; valleys and foothills near springs. 1134. Poa ample Merr. Big Bluegrass. Collected only from streamside in South Willow Can- yon. 114 (UT and UTC). "Poa annua L. Annual Bluegrass. Seen only at streamside in South Willow Canyon. 415 (UT and UTC). "Poa bulbosa L. Bulbous Bluegrass. In- frequent; disturbed sites at low elevations. 397 (UTC). Poa canbyi (Scribn.) Howell. Canby Blue- grass. Collected only from mouth of Daven- port Canyon. 24 (UT). "Poa compressa L. Canada Bluegrass. Rare; streamside in the foothills. 1127. Poa fendleriana (Steud.) Vasey. Mutton Grass. Common; dry meadows and mostly open slopes from middle elevations to alpine. 451. Poa leptocoma Trin. Bog Bluegrass. Mesic slopes and meadows from middle elevations to subalpine. Taye & Wall 1175. Poa nervosa (Hook.) Vasey. Wheeler Blue- grass. Freqvient; open forest understory and meadows from middle elevations to sub- alpine. 989. "Poa pratensis L. Kentucky Bluegrass. Fre- quent; disturbed to pristine, generally mesic sites from the foothills to subalpine. 695 (UTC). October 1983 Taye: Stansbury Mountains Flora 645 Poa reflexa Vasey & Scribn. Nodding Blue- grass. Locally common in subalpine mead- ows. 1253. Poa secunda Presl. Sandberg Bluegrass. [P. sandbergii Vasey] Common; open slopes from the valleys to alpine. Arnow (1981) cites P. secunda as being the correct name for this species. °Polypogon monspeliensis (L.) Desf. Rabbit- foot Grass. Frequent; streamside at low ele- vations. 685. " Puccinellia distans (L.) Pari. European Al- kaligrass. Rare; streamside at middle eleva- tions. 643. ° Puccinellia fasiculata (Torr.) Bickn. Alka- ligrass. Locally common at Big Spring in Skull Valley. 1373. "Secale cereale L. Rye. Infrequent; valley benches (cultivated) and occasionally higher. 598. Sitanion hystrix (Nutt.) Smith. Squirreltail. Common; dry, mostly open slopes from the foothills to alpine. 499. Sphenopholis obtusata (Michx.) Scribn. Prairie Wedgescale. Rare; streamside near mouth of Spring Canyon. 1125b. Sporobolus airoides (Torr.) Torr. Alkali Sacaton. Infrequent; valleys and foothills. 1023. Sporobolus cryptandrus (Torr.) Gray. Sand Dropseed. Frequent; valleys and foothills, usually in sandy areas. 549. Stipa comata Trin. & Rupr. Needle-and- Thread Grass. Frequent; valleys and foothills in sandy areas. 844. Stipa lettemiannii Vasey. Letterman Nee- dlegrass. Rare; cirque meadow in Dry Lake Fork. 351 (UT and UTC). Stipa nelsonii Scribn. Columbia Needle- grass. [S. Columbiana Macoun] Frequent; meadows and mostly open slopes from middle elevations to subalpine. 1113. Trisetum spicatum (L.) Richter. Spike Trisetum. Common; meadows, streamside, and open forest understory from middle ele- vations to alpine. 582. °Triticum aestivum L. Wheat. Locally com- mon (cultivated) at mouth of East Hickman Canyon. 599 (UTC). "Vulpia myuros (L.) C. C. Gmelin. Foxtail Fescue. [Festuca myuros L.] Rare; streamside near mouth of Spring Canyon. 1138. Vidpia octoflora (Walt.) Rydb. Six-weeks Fescue. [Festuca octoflora Walt.] Infrequent; valleys and foothills, usually in sandy areas. 808. Ruppiaceae Ruppia maritima L. Widgeongrass. Locally common in Big Spring in Skull Valley. 1401. Typhaceae Typha latifolia L. Common Cattail. In- frequent; streamside in the foothills. 1145. Zannichelliaceae Zannichellia palustris L. Horned Pond- weed. Rare; in ponds and streams at low ele- vations. 1372. Literature Cited Allred, K. W. 1975. Timpanogos flora. Unpublished thesis. Brigham Young Univ., Provo, Utah. 178 pp. Arnow, L. 1981. Poa secunda Presl versus P. sandbergii Vasey, (Poaceae). Syst. Bot. 6:412-421. Arnow, L., B. Albee, and A. Wyckoff. 1980. Flora of the central Wasatch Front, Utah. 2d ed. rev. Univ. of Utah Printing Service, Salt Lake City. 663 pp. Behle, W. H. 1978. Avian biogeography of the Great Basin and Intermountain Region. Great Basin Nat. Mem. 2:55-80. Billings, W. D. 1951. Vegetational zonation in the Great Basin of western North America. Int. Union Biol. Sci. Ser. B, 9:101-122. 1978. Alpine phytogeography across the Great Basin. Great Basin Nat. Mem. 2:105-117. Cronquist, a., a. H. Holmgren, N. H. Holmgren, and J. L. Reveal. 1972. Intermountain flora. Vol. 1. Hafner Publ. Co., New York. 270 pp. Cronquist, A., A. H. Holmgren, N. H. Holmgren, J. L. Reveal, and P. K. Holmgren. 1977. Inter- mountain flora. Vol. 6. Columbia Univ. Press, New York. 584 pp. Harper, K. T., D. C. Freeman, W. K. Ostler, and L. G. Klikoff. 1978. The flora of Great Basin moun- tain ranges: diversity, sources, and dispersal ecol- ogy. Great Basin Nat. Mem. 2:81-10.3. Hitchcock, C. L., and A. Cronquist. 1973. Flora of the Pacific Northwest. Univ. of Washington Press, Seattle. 750 pp. Holmgren, A. H. 1972. Vascular plants of the Northern Wasatch. 4th ed. Privately printed by the author, Logan, Utah. 202 pp. Holmgren, A. H., L. M. Shultz, and T. K. Lowrey. 1976. Sphaeromeria, a genus closer to Artemisia than to Tanacetum (Asteraceae: Anthemideae). Brittonia 28:255-262. 646 Great Basin Naturalist Vol. 43, No. 4 Holmgren, N. H. 1972. Plant geography of the Inter- mountain Region. Pages 77-161 in A. Cronquist, A. H. Holmgren, N. H. Holmgren, and J. L. Re- veal, eds., Intermountain flora. Vol. 1. Hafner Publ. Co., New York. Houghton, J. G. 1969. Characteristics of rainfall in the Great Basin. Desert Research Institute. Univ. of Nevada System, Reno. 205 pp. Jones, G. N., and F. F. Jones. 1943. A revision of the perennial species of Geranium of the United States and Canada. Rhodora 45:5-26, 32-53. Jones, M. E. 1965. Botanical exploration of Marcus E. Jones, 1876-1919: an autobiographical account. Leafl. W. Bot. 10:189-236. Kearney, T. H., L. J. Briggs, H. L. Shantz, J. W. McLane, and R. L. Piemeisel. 1914. Indicator significance of vegetation in Tooele Valley, Utah. J. Agric. Res. 1:365-417. Lewis, M. E. 1971. Flora and major plant communities of the Ruby-East Humboldt mountains with spe- cial emphasis on Lamoille Canyon. Report to Humboldt National Forest, Elko, Nevada. 62 pp. 1973. Wheeler Peak Area species list. Report to Intermountain Region, U.S. Forest Service, Og- den, Utah. 17 pp. 1975. Plant communities of the Jarbidge Moun- tain complex, Humboldt National Forest. Report to Intermountain Region, U.S. Forest Service, Ogden, Utah. 22 pp. 1980. Plants of the Wasatch Plateau, Utah. Re- port to Intermountain Region, U.S. Forest Ser- vice, Ogden, Utah. 37 pp. McMillan, C. 1948. A taxononiic and ecological study of the flora of the Deep Creek Mountains of cen- tral western Utah. Unpublished thesis. Univ. of Utah, Salt Lake City. 99 pp. Morrison, R. B. 1965. Quaternary geology of the Great Basin. Pages 265-285 in H. E. Wright, Jr., and D. G. Frey, eds.. The Quaternary of the United States. Princeton Univ. Press, Princeton, New Jersey. Nebeker, G. T. 1975. Manual of the flora of the East Tintic Mountains, Utah. Unpublished thesis. Brigham Young Univ., Provo, Utah. 57 pp. Preece, S. J., Jr. 1950. Floristic and ecological features of the Raft River Mountains of north-western Utah. Unpublished thesis. Univ. of Utah, Salt Lake City. 103 pp. Reveal, J. L. 1973. Eriogonum (Polygonaceae) of Utah. Phytologia 25:169-217. 1979. Biogeography of the Intermountain Region: a speculative appraisal. Mentzelia 4:1-92. RiGBY, J. K. 1958. Geology of the Stansbury Mountains. Utah Geol. Soc. Guidebook 13:1-1.34. Soil Conservation Service. 1973. General soil map- Utah. Utah Agric. Expt. Sta., Logan. 1979. Mountain precipitation summary for Utah. Snow Survey Section, Federal Bldg., Salt Lake City, Utah. 168 pp. Stansbury, H. 1852. Exploration and survey of the val- ley of the Great Salt Lake of Utah, including a reconnaissance of a new route through the Rocky Mountains. Lippincott, Crambo, Philadelphia. 487 pp. Taye, a. C. 1981. A floristic and phytogeographic study of the Stansbury Mountains, Utah. Unpublished thesis. Brigham Young Univ., Provo, Utah. 172 pp. Thorne, R. F. 1967. A flora of Santa Catalina Island, California. Aliso 6(3): 1-77. TiDESTROM, I. 1913. Novitates florae utahensis. Proc. Biol. Soc. Washington 26:121-122. Trewartha, G. T. 1968. An introduction to climate. 4th ed. McGraw-Hill, New York. 408 pp. U.S. Department of Commerce. 1980. Climatological data annual summary, Utah, 1979. National Oce- anic and Atmospheric Administration, Asheville, North Carolina. 15 pp. Wells, P. V. 1980. Quaternary vegetational history of the Great Basin. Bull. Ecol. Soc. Amer. 57:106 (abstr.) Welsh, S. L. 1978a. Endangered and threatened plant species inventory for Deep Creek Mountains, Utah. Final unpublished report. Bureau of Land Management. 159 pp. 1978b. Utah flora: Fabaceae (Leguminosae). Great Basin Nat. 38:225-367. 1982. Utah plant types— historical perspective 1840 to 1981— annotated list and bibliography. Great Basin Nat. 42:129-195. Welsh, S. L., N. D. Atwood, S. Goodrich, E. Neese, K. H. Thorne, and B. Albee. 1981. Preliminary index of Utah vascular plant names. Great Basin Nat. 41:1-108. Welsh, S. L., and R. C. Barneby. 1981. Astragalus len- tiginosus (Fabaceae) revisited— a unique new va- riety. Isleya2(l):l-2. Welsh, S. L., and G. Moore. 1973. Utah plants, Tracheophyta. 3d ed. Brigham Young Univ. Press, Provo, Utah. 474 pp. Welsh, S. L., and J. L. Reveal. 1977. Utah flora: Brassi- caceae (Cruciferae). Great Basin Nat. 37:279-365. Wilson, L., M. E. Olsen, T. B. Hutchings, A. R. Southard, and A. J. Erickson. 1975. Soils of Utah. Agric. Expt. Sta. Bull. 492, Utah State Univ., Logan. 94 pp. NEW SYNONYMY AND NEW SPECIES OF AMERICAN BARK BEETLES (COLEOPTERA: SCOLYTIDAE), PART IX Stephen L. Wood' Abstract.— The following new synonymy in Scolytidae is proposed. Acorthylits Brethes (= Phacrijlus Schedl), Anuisa Lea (= Anaxi/leborus Wood), Cryptocttrus Schedl (= Hylopems Browne), Hylesinopsis Eggers (= Trypo- graphus Schedl, Chilodendron Schedl), Hypothenemtis Westwood (= Stylotenttis Schedl), Metahylesintis Eggers ( = Glochicoptenis Schedl), Pliloeotribus Latreille (= Neophloeotribtis Eggers), Pityophthorus Eichhoff (= Hypopityop- tlionts Bright), Scohjtoplatyptis Schaufiiss (= Spongoceriis Blandford, Taeniocenis Blandford, Stwphionocerus Samp- son), Scolytits Geoffroy (= Confusoscolyttis Tsai & Hwang), Styracoptinus Wood (= Afrotrypettis Bright), Stietis Mu- rayama (= Neohyorrhynchtis Schedl), Taphrorychus Eichhoff (= Pseudopoecilips Murayama), Webbia Hopkins ( = Pseudotcebbia Browne), Webbia dipterocarpi Hopkins (= Webbia 18-spinatus Sampson), Xyleborus Eichhoff ( = Anaeretiis Diiges). A neotype is designated for Anaerettts guanagitatensis Duges; this name becomes a junior syn- onym of Xyleborus volvulus (Fabricius). Species new to science are named from Mexico as follows: Cactopinus atkin- soni, burjosi, granulatus, setosus, Carphobius pilifer, Chaetophloeus corifinis, Chrarnesus exilis, exul, securus, tibialis, Cuemonyx equihuai, evidens, Cnesinus cornutus, nebulosus, parvicornis, Dendrosinus mexicanus, Liparthrum mexi- canum, pruni, Phloeotribus geminus, Pycnarthrum amersum, Scolytodes plumericolens, retifer. A review of the holotypes of the type-spe- cies of several obscure genera of Scolytidae has led to the detection of several previously unpublished or obscured synonyms. These are presented below in alphabetical order of the senior generic name. The synonymy of Web- bia dipterocarpi Hopkins is included. The continuing faunal survey of Mexico of Dr. T. H. Atkinson, Chapingo, Mexico, has resulted in the discovery of a number of spe- cies new to science. Twenty-two species found during that survey are presented be- low. They represent: Cactopinus (4), Car- phobius (1), Chaetophloeus (1), Chrarnesus (4), Cnemonyx (2), Cnesinus (3), Dendrosinus (1), Liparthrum (2), Phloeotribus (1), Pyc- narthrum (1), and Scolytodes (2). New Synonymy Acorthylus Brethes Acorthylus Brethes, 1922, Ann. Soc. Cien. Argentina 94:304 (Type-species: Acorthylus asperatus Brethes, monobasic) Phacrylus Schedl, 1938, Rev. Soc. Ent. Argentina 10:24 (Type-species: Phacrylus bosqui Schedl). New synonymy The Argentine species Acorthylus aspe- ratus Brethes has stood as an unidentifiable species since its description. However, the description clearly characterizes the 3-seg- mented antennal funicle with the middle seg- ment enlarged and almost equal in length to the scape. Because no other genus shares this character and because the description of the type species matches in every detail those of Argentine species placed by Schedl in his Phacrylus, it must be concluded that Pha crylus is a junior synonym of Acorthylus as indicated above. Amasa Lea Amasa Lea, 1894, Proc. Linn. Soc. New South Wales (2) 8:322 (Type-species: Amasa thoracicus Lea = To- jyiicus truncatus Erichson, monobasic) Anaxyleborus Wood, 1980, Great Basin Nat. 40:90 (Type-species: Tomicus truncatus Erichson, origi- nal designation). New synonymy When the name Anaxyleborus Wood (1980) was proposed, I overlooked the synon- ymy (Lea, 1904, Linn. Soc. New South Wales 29:106) of the type-species, Tomicus trun- catus Erichson, with Amasa thoracicus Lea. In view of this synonymy involving the type- species, Anaxyleborus automatically becomes a synonym of Amasa. 'Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. 647 648 Great Basin Naturalist Vol. 43, No. 4 Cryptocurus Schedl Cnjptocurus Schedl, 1957, Ann. Mag. Nat. Hist. (12) 10:869 (Type-species: Cnjptocurus spinipennis Schedl, monobasic) Hyloperus Browne, 1970, J. Nat. Hist. 4:.546 (Type-spe- cies: Hyloperus bicornis Browne, original desig- nation). New synonyrnij The male holotype of Cryptocurus spin- ipennis Schedl and the female holotype of Hyloperus bicornis Browne were examined and compared directly to one another and to other members of this species. They repre- sent opposite sexes of the same species. Con- sequently, Hyloperus becomes a junior syn- onym of the older name. Hylesinopsis Eggers Hylcsinopsis Eggers, 1920, Ent. Blatt. 16:40 (Type- species: Hylesinopsis duhius Eggers, monobasic) Trypographus Schedl, 1950, Rev. Francaise Ent. 17:21.3 (Type-species: Trypographus joveri Schedl, mon- obasic). Neiv synonymy Chilodendron Schedl, 195.3, Mem. Inst. Sci. Madagascar (E) 3:74 (Type-species: Chilodendron plm^icoUe Schedl, monobasic). New synonymy When Hylesinopsis Eggers (1920), Trypo- graphus Schedl (1950), and Chilodendron Schedl (1953) were named, there may have been some justification for the recognition of three genera. However, the subsequent dis- covery of additional species has closed the character gap used to distinguish them. I now see no justification for the recognition of more than one genus and place Tijpographus and Chilodendron in synonymy under Hylesi- nopsis as indicated above. Hypothenemus Westwood Hypothenemiis Westwood, 1836, Trans. Ent. Soc. Lon- don 1:34 (Type-species: Hypothenemus eruditus Westwood, monobasic) Stylotentus Schedl, 19.39, Rev. Zool. Bot. Afr. 32:380 (Type-species: Hypothenemus concolor Hage- dom, subsequent designation by Schedl, 1961, Rev. Ent. Moc^ambique 4:448). New synonymy Schedl (1936:380) established the genus Stylotentus on the basis of a peculiarity in the antennal club and fimicle. The club ap- pears to have fused with funicular segments 4 and 5; fimicular segments 1-3 are normal. The funicle in Hypothenemus is unstable, varying from three to five segments. Parital fusion of segments is a common feature, and occasionally the left and right antennae will bear different numbers of segments. Of the three specimens of concolor Hagedorn in my collection, the funicle of one is 5-segmented, one 3-segmented, and one 2-segmented (all four flagellar segments are fused). Two of the three specimens of ater (Eggers), also as- signed by Schedl to Stylotentus, have the fu- nicle 4-segmented. In view of the instability of this character in these species, and the var- iability of funicular segmentation in other Hypothenemus, Stylotentus is placed in syn- onymy under the senior name Hypo- thenemus. Metahylesinus Eggers Metahylesinus Eggers, 1922, Ent. Bliitt. 18:165 (Type- species: Pseudohylesinus togonus Eggers, automatic) Glochicopterus Schedl, 1954, Rev. Zool. Bot. Afr. 50:75 (Type-species: GlocJiicopterus haphiae Schedl, monobasic). New synonymy Following a study of Metahylesinus to- gonus (Eggers), of five other species currently assigned to Metahylesinus Eggers, and of Glochicopterus haphiae Schedl, I am unable to detect characters that separate these spe- cies into distinct genera. For this reason, Glochicopterus is placed in synonymy as in- dicated above. Phloeotribus Latreille Phloeotribus Latreille, 1797, Free, caract. gen. insects, p. 50 (Type-species: Hylesinus oleae Fabricius = Scohjtus scarahaeoides Bernard, monobasic) Neophloeotribus Eggers, 1943, Mitt. Miinchner Ent. Ges. 33:349 (Type-species: Phloeotribus nubilus Bland- ford, present designation). New synonymy Eggers (1943) proposed the subgeneric name Neophloeotribus for a group of species that included Phloeotribus nubilus Blandford and Phloeotribus suturalis Eggers. I designate Phloeotribus nubilus Blandford as the type- species of Neophloeotribus. At the present time, I see no need to subdivide Phloeotribus and therefore place Neophloeotribus in syn- onymy as indicated above. Pityophthorus Eichhoff Pityophthorus Eichhoff, 1864, Berliner Ent. Zeitschr. 8:.39 (Type-species: Bostrichus lichtensteini Ratzeburg) Hypopityophthorus Bright, 1981, Mem. Ent. Soc. Can- ada 118:14 (Type-species: Pityophthorus inops Wood). New synonymy October 1983 Wood: American Bark Beetles 649 Bright (1981) established the subgenus Hy- popityophtfiorus on the basis of degenerate sutures in the antennal club. The characters are exactly the same as those used by Black- man to characterize Pityophthoroides. Neo- tropical Pityophthorus exhibit varying de- grees of suture deterioration on an antennal club that increases in thickness. The trend is gradual and is best seen in the smallest spe- cies. Because Hypopityophthorus was pro- posed for the same group as Pityophthoroides, which is no more than a species group of in- definite extent, it is placed in synonymy un- der Pityophthorus. Scolytoplatypus Schaufuss Scolytopkitypits Schaufiiss, 1890, Beitrag zur Kaferfauna Madagascars 2:31 (Type-species: Scolytoplatypus pennims Schaufiiss, monobasic) Spongocerus Blandford, 1893, Trans. Ent. Soc. London 1893:431 (Type-species: Scolytoplatypus tycon Blandford, subsequent designation by Hopkins, 1914, Proc. U.S. Nat. Mus. 48:129). New synonymy Toeniocerus Blandford, 1893, Trans. Ent. Soc. London 1893:431 (Type-species: Scolytoplatypus mikado Blandford, subsequent designation by Hopkins, 1914, Proc. U.S. Nat. Mus. 48:431). Preoccupied by Kamp 1871 Strophionocerus Sampson, 1921, Ann. Mag. Nat. Hi,st. (9) 7:36. (Replacement name for Taeniocerus Bland- ford). New synonymy Blandford (1893) proposed for the genus Scolytoplatypus Schaufuss the subgeneric names Spongocerus and Taeniocerus. The lat- ter name was a homonym that was replaced by Strophionocerus. These names were un- necessary in the classification of the genus and have been ignored by subsequent work- ers treating the genus. Attention is called to them for the purpose of placing them in syn- onymy as indicated above. Scolytus Geoffroy Scolytus Geoffroy, 1762, Histoire abregee des insects, p. 309 (Type-species: Bostrichus scolytus Fabricius, .subsequent designation by International Commis- sion on Zoological Nomenclature) Arcluieoscolytus Butovitsch, 1929, Stettiner Ent. Zeit. 90:21, 23 (Species group name for Scolytus clavi- ger Blandford). No status Spinuloscolytus Butovitsch, 1929 Stettiner Ent. Zeit. 90:21, 24 (Species group name for Ips multi- striatus Marsham, Scolytus orientalis Eggers, Sco- lytus ecksteini Butovitsch). No status Tuholoscolytus Butovitsch, 1929, Stettiner Ent. Zeit. 90:21, 24 (Species group name for Eccoptogaster intricatus, Eccoptogaster carpini Ratzeburg, Sco- lytus koenigi Schevyrew). No status Pygmaeuscolytus Butovitsch, 1929, Stettiner Ent. Zeit. 90:21, 28 (Species group name for Scolytus kirschi Skalitzky, Scolytus fasciatus Reitter, Bostricnus pygmaeus Fabricius, Scolytus ensifer Eichhoff, Scolytus zuitzevi Butovitsch). No status Pinetoscolytus Butovitsch, 1929, Stettiner Ent. Zeit. 90:22, 48 (Species group name for Scolytus mora- witzi Semenov). No status Confusoscolytus Tsai & Hwang, 1962, Acta Ent. Sinica 11:4, 14 (Type-species: Eccoptogaster confusus Eggers). New synonymy Several mononominal designations within the genus Scolytus Geoffroy have been pub- lished that have caused confusion in the liter- ature treating Scolytidae. Five of these were published by Butovitsch (1929) in one paper, including Archaeo scolytus, Spinuloscolytus, Tuhuloscolytus, Pygmaeoscolytus, and Pine- toscolytus, as species-group names. As such they have no status in nomenclature. They were not intended to be genus-group names and should not be cited as such. The name Confusoscolytus Tsai & Hwang (1962) was presented as a subgenus of Sco- lytus and is of nomenclatural interest. How- ever, because the Scolytus species of Europe, Asia, and North America are not divisible into recognizable subgenera, Confusoscolytus has value no greater than a species-group and, therefore, must be treated as a synonym of Scolytus. Styracoptinus Wood Styracoptinus Wood, 1962, Great Basin Nat. 22:77 (Re- placement name for Styracopterus Blandford, preoccupied. Type-species: Styracopterus murex Blandford, automatic) Afrotrypetus Bright, 1981, Coleopt. Bull. 35:113 (Type- species: Afrotrypetus euphorbiae Bright, original designation. New synonymy During a visit with Dr. K. E. Schedl in 1965, I showed him a pair of specimens from the same series Bright (1981) later named as Afrotrypetus euphorbiae. We agreed that the species represented Styracoptinus. He had a vague recollection of having named it in an- other genus from a unique specimen that was deposited in another collection. Since then, other Styracoptinus have been named that more fully bridge the gap between murex and euphorbiae. For this reason Afrotrypetus must become a junior synonym of Styracoptinus. 650 Great Basin Naturalist Vol. 43, No. 4 Sueus Murayama Sueus Murayama, 1951, Bull. Facul. Agric. Yarnaguti Univ. 2:1 (Type-species: Sueus sphaerotrypoides Murayama = Hyorrhynchus niisimai Eggers, original designation) Neohyorrhynchus Schedl, 1962, Ent. Blatt. 58:202 (Type-species: Hyorrhynchus niisimai Eggers, monobasic). New synonymy The female holotypes of Sueus sphaerotry- poides Murayama and Hyorrhynchus niisimai Eggers were examined and compared di- rectly to my specimens. Because they repre- sent the same species, Neohyorrhynchus Schedl becomes an objective junior synonym of the older name. Taphrorychus Eichhoff Taphrorychus Eichhoff, 1878, preprint of Mem. Soc. Roy. Sci. Liege (2) 8:49, 204 (Type-species: Bos- trichus bicolor Herbst, subsequent designation by Hopkins, 1914, Proc. U.S. Nat. Mus. 43:130) Pseudopoecilips Murayama, 1957, Bull, Facul. Agric. Yamaguti Univ. 8:614 (Type-species: Pseudo- poecilips mikuniyamensis Murayama, original designation). New synonymy Following my examination of authentic specimens of all species currently assigned to Taphrorychus Eichhoff and of the three spe- cies of Pseudopoecilips named by Murayama, it was concluded that Pseudopoecilips fits well within the anatomical and biological limits of Taphrorychus. Accordingly, Pseudo- poecilips is placed in synonymy under Taph- rorychus as indicated above. Webbia Hopkins Webbia Hopkins, 1915, U.S. Department of Agriculture Bur. Ent. Tech Bull. 17(2):222 (Type-species: Webbia chpterocarpi Hopkins, original designation) Pseudowebbia Browne, 1961, Sarawak Mus. J. 10:308 (Type-species: Xyleborus trepanicauda Eggers, original designation). New synonymy In my review of the status of the genera of Scolytidae, the type specimens of the type species of Webbia Hopkins (W. dipterocarpi Hopkins) and Pseudowebbia Browne {Xyle- borus trepanicauda Eggers) were examined along with almost all the other species in these genera. It is apparent that dipterocarpi and trepanicauda both represent the same species group within the genus. For this rea- son, the junior name, Pseudowebbia, must be placed in synonymy. Webbia dipterocarpi Hopkins Webbia dipterocarpi Hopkins, 1915, U.S. Department of Agriculture Bur. Ent. Tech Bull. 17(2):223 (Holo- type, female; near Pagbilao, Philippine Islands; U.S. Nat. Mus.) Webbia 18-spinatus Sampson, 1921, Ann. Mag. Nat. Hist. (9)7:34 (Holotype, female; Penang, Bryant; British Mus. Nat. His.). New synonymy The female holotypes of Webbia diptero- carpi Hopkins and Webbia 18-spinatus Sampson were both compared directly to my specimens from the Philippines and Malaya. Because they are identical in all respects, it is concluded that they represent the same spe- cies. For this reason, 18-spinatus is placed in synonymy as indicated above. Xyleborus Eichhoff Xyleborus Eichhoff, 1864, Berliner Ent. Zeitschr. 8:37 (Type-species: Bostrichus monographus Fabricius, subsequent designation by Lacardaire, 1866, Hist. Gen. Coleopt. 7:381) Anaeretus Duges, 1887, Ann. Soc. Ent. Belgique 31:141 (Type-species: Anaeretus guanaguatensis Duges = Bostrichus volvulus Fabricius). New synonymy The genus Anaeretus Duges was estab- lished for guanaguatensis Duges and based on specimens deposited in the Museo Nacion- al de Historia Natural at Mexico City. The major portion of the Duges collection was later moved to the Universidad Nacional Au- tonoma de Mexico, also in Mexico City. Two unsuccessful searches were conducted for the types of Anaeretus guanaguatensis, first in 1974 by me, the second by W. F. Barr in February 1982. The specimens could not be found and are presumed to be lost. From the rather complete description of guanaguatensis, it is apparent that the type series was of either Xyleborus volvulus (Fab- ricius) or, possibly, X. affinis Eichhoff, both of which are common throughout Mexico ex- cept for the very dry northern areas. In order to remove ambiguity from the placement of Anaeretus in the classification of Scolytidae, I here designate as the neotype of Anaeretus guanaguatensis Duges the female lectotype of Xyleborus volvulus (Fabricius) that is in the Copenhagen museum. Correction Cnesinus equihuai Wood, emendation Cnesinus aquihuai Wood, 1982, Great Basin Nat. 42:226 (Holotype, female; between Cuetzalan and Pasa October 1983 Wood: American Bark Beetles 651 del Jardin, Puebla, Mexico; Wood Collection). Lapsus cdlmi A proofreading error occurred in the origi- nal spelling of the specific name of this spe- cies and in references to the name of the col- lector. Armando Equihua has been an enthusiastic student of the Scolytidae and an ardent collector of numerous rare forms, many of which have been named as new to science. New Taxa Cactopinus atkinsoni, n. sp. This species is distinguished from naiisutus Wood by the smaller size, by the less deeply excavated male frons with the upper margin more rounded, and by the jteeper lower declivity. Ma /e.— Length 1.3 mm (paratypes 1.3-1.5 mm), 2.4 times as long as wide; color black. Frons similar to nausutus except upper area of frons not as widely or as deeply exca- vated, upper margin much less acute; horn averaging smaller. Pronotum as in nausutus. Elytra as in nausutus except declivity steeper, more narrowly sulcate. Female.— Similar to male except frons about as in female nausutus. Type locality.— Estacion de Biologia, Cha- mela, Jalisco, Mexico. Type material.— The male holotype, fe- male allotype, and 18 paratypes were taken at the type locality on 28-V-1982, 80 m, S-497, Pachycerus, T. H. Atkinson and A. Equihua. The holotype, allotype, and paratypes are in my collection. Cactopinus burjosi, n. sp. This species is distinguished from niger Wood by the less distinctly concave male frons, by the larger strial punctures, by the presence of interstrial tubercles, and by the very different declivity as described below. Male.— Length 1.5 mm (paratypes 1.5-1.6 mm), 2.3 times as long as wide; color black. Frons largely hidden by pronotum, impres- sion apparently limited, if present; horns ba- sally contiguous and of about same size and form as cactophthorus Wood. Antennal club small, sutures weakly procurved, almost straight. Pronotum about as in niger except aspe- rities and tubercles in lateral areas larger. Elytra 1.4 times as long as wide; sides al- most straight and parallel on basal two-thirds, rather narrowly rounded behind; striae 1 weakly, others not impressed, punctures very coarse, deep, and poorly formed on basal half, decreasing posteriorly until obsolete by base of declivity, small granules between punctures on posterior half of disc gradually replace punctures posteriorly; interstriae about half as wide as striae, irregular, each armed by a uniseriate row of small tubercules except basal half of even-numbered inter- striae indistinctly punctured. Declivity oc- cupying posterior third, less abrupt and less strongly sulcate than in related species; striae and interstriae marked by small, acutely rounded tubercles, except on lower third of sulcus only obscure strial punctures in- dicated. Vestiture of sparse, confused, minute hairlike setae. Female.— Similar to male except frons sim- ilar to females of related species. Type locality.— Tepenene, Puebla, Mexico. Type material- The male holotype, fe- male allotype, and two paratypes were taken on l-X-1982, 1240 m, B-070, Neobuxbaumia mezealensis, A. Burjos and E. Saucedo. The holotype, allotype, and paratypes are in my collection. Cactopinus granulatus, n. sp. This species is distinguished from nausutus Wood by the smaller pronotal asperities and by the presence of discal tubercles on the striae and interstriae. Male.— Length 1.7 mm (paratypes 1.6-1.8 mm), 2.3 times as long as wide; color black. Frons as in nausutus except excavated area not quite as wide above eyes. Pronotum as in nausutus except asperities distinctly smaller. Elytra about as in nausutus except strial punctures larger, deeper; interstriae irregular on basal half, posterior half armed by irregu- larly placed small tubercles, a few similar tu- bercles on striae between punctures. Declivi- ty similar to nausutus, strial punctures 652 Great Basin Naturalist Vol. 43, No. 4 continue to apex, strial and interstrial tu- bercles absent except on interstriae 1 and 3. Vestiture sparse, short, of fine interstrial hair. Female.— Similar to male except frons as in female nausutus. Type locality.— Autlan, carr. Barra de Na- vidad km 163, Jalisco, Mexico. Type material.— The male holotype, fe- male allotype, and eight paratypes were taken at the type locality on 3- VII- 1982, S-751, Cactaceae, A. Equihua. The holotype, allotype, and paratypes are in my collection. Cactopinus setosus, n. sp. This species is distinguished from nausutus Wood by the stout body form, by the smaller strial punctures, and by the rather abundant elytral vestiture. Mfl/e.— Length 1.6 mm (paratypes 1.4-1.7 mm), 2.2 times as long as wide; color black. Frons similar to nausutus except upper ex- cavated area less strongly impressed; horn av- eraging slightly shorter, its apices usually blunt. Pronotum similar to nausutus except aspe- rities less numerous and smaller, median basal area more rounded and with fewer asperities. Elytra similar to nausutus except declivity less strongly, more narrowly sulcate; discal surface largely obscured by incrustation, ap- parently strial punctures smaller, not as deep, interstriae smooth, uniseriate punctures small; a few small granules on odd-numbered interstriae toward declivity; declivital strial punctures larger and deeper than on disc. Vestiture of rows of rather coarse, erect in- terstrial setae, each seta very slightly longer than distance between rows or between setae within a row. Female.— Similar to male except frons sim- ilar to female nausutus. Type locality.— Estacion de Biologia, Cha- mela, Jalisco, Mexico. Type material— The male holotype, fe- male allotype, and 13 paratypes were taken at the type locality on 28-V-1982, 80 m, S-498, Cactaceae, T. H. Atkinson and A. Equihua. The holotype, allotype, and paratypes are in my collection. Carphobius pilifer, n. sp. This species is distinguished from cupressi Wood by the much more abundant, longer vestiture throughout the body, by the finer pronotal punctures, and by other characters cited below. Female.— Length 2.9 mm (paratypes 2.8-3.0 mm), 2.3 times as long as wide; color very dark brown, elytra rather dark brown. Frons as in cupressi except epistomal pro- cess more conspicuous, vestiture more abun- dant and much coarser. Pronotum as in cupressi except punctures half as large, closer, not as deep; vestiture ob- scurely subplumose, appearing much coarser, longer, more abundant. Elytra as in cupressi except declivity steep- er, strial punctures smaller, interstriae wider, with punctures more numerous and confused, vestiture obscurely subplumose, longer, much more abundant. Type locality.— Tres Marias, Morelos, Mexico. Type material.— The female holotype and two female paratypes were taken at the type locality on 30- V- 1982, 2790 m, B-029, Cu- pressus lindleyi, A. Burjos and E. Saucedo. The holotype and paratypes are in my collection. Chaetophloeus confinis, n. sp. This species is distinguished from stru- thanthi Wood by the less strongly concave male frons, by the larger frontal granules in both sexes, and by the longer, more slender setae on the elytral declivity. Male.— Length 1.7 mm (allotype 1.7 mm), 1.7 times as long as wide; color dark brown. Frons shallowly concave to slightly above upper level of eyes; similar to struthanthi ex- cept concavity not as deep nor extending as high on vertex; long setae on upper margin shorter, not reaching middle of frons. Pronotum as in struthanthi except vestiture distinctly longer, surface without any reticulation. Elytra as in struthanthi except striae less distinctly impressed, punctures slightly larger, setae longer; longest setae at base of declivity six times as long as wide (in stru- thanthi not more than four times as long as wide.) October 1983 Wood: American Bark Beetles 653 Female.— Similar to male except frons con- vex, frontal tubercles larger, frontal vestiture normal. Tijpe locality.— Cuernavaca, Morelos, Mexico. Type material.— The male holotype was taken at the type locality on 28-VI-1982, 1500 m, AB-070 Phoradendron, by A. Burjos; the allotype from the same locality 4-VII-1982, 1519 m, SH-011, Phoradendron by E. Saucedo. The holotype and allotype are in my collection. Chramesus exilis, n. sp. This species is distinguished from gracilis Wood by the smaller size, by the finer, more slender (but not longer) vestiture, by the more slender pronotum, and by the less strongly impressed male frons. Male.— Length 1.7 mm (paratypes 1.6-1.8 mm), 2.3 times as long as wide, color very dark brown, vestiture pale. Frons moderately, concavely impressed on median two-thirds of lower two-thirds; sur- face reticulate, punctures not clearly evident; tubercles smaller and vestiture finer than in gracilis. Pronotum 0.94 times as long as wide; sur- face as in gracilis except granules more regu- larly present and vestiture finer. Female.— Similar to male except frons con- vex, a slight transverse impression just above epistoma; frontal tubercles present, but smalller. Type locality.— El Tuito, Jalisco, Mexico. Type material— The male holotype, fe- male allotype, and 30 paratypes were taken at the type locality on 28- V- 1982, 640 m, S-707, from Smilax by T. H. Atkinson and A. Equihua. The holotype, allotype, and paratypes are in my collection. Chramesus exul, n. sp. This species is distinguished from vitiosus Wood by the absence of pronotal reticulation and by the punctured male striae. Although it superficially resembles vitiosus, its true relationships are probably much closer to xyhphagus Wood. Male.— Length 1.3 mm (paratypes 1.1-1.5 mm), 1.5 times as long as wide; color very dark brown, with pale vestiture. Frons moderately, somewhat narrowly concave from epistoma to upper level of eyes, lateral margins weakly elevated, armed immediately below level of antennal in- sertion by a small tubercle; surface almost smooth, obscurely rugose-reticulate; vestiture fine, short, inconspicuous. Pronotum resembling xylophagus except more strongly arched, punctures closer, smaller, and deeper; vestiture short, rather stout (each at least six times as long as wide), moderately abundant. Elytra about as in xylophagus except setae in ground cover much stouter, erect setae of equal width and about twice as long as ground setae; each erect seta about three to four times as long as wide. Female.— Similar to male except frons con- vex, lateral tubercles absent. Type locality.— Nine km southeast of To- tolapan, Oaxaca, Mexico. Type material— The male holotype, fe- male allotype, and 10 paratypes were taken at the type locality, 21-VI-1967, 1000 m, No. 70, from an unidentified shrub, by me; 16 paratypes are from Estacion de Biologia, Chamela, Jalisco, 19-VIII-1982, 100 m, S-758, from a Leguminosae, by A. Equihua. The holotype, allotype, and paratypes are in my collection. Chramesus securus, n. sp. This species is distinguished from vitiosus Wood by the smooth, shining surface of the pronotum (between the small tubercles), by the more strongly arched elytral declivity, and by the more slender, erect interstrial setae. Male.— Length 1.5 mm (paratypes 1.5-1.7 mm), 1.5 times as long as wide; color dark reddish brown, vestiture pale. Frons as in vitiosus except lateral margin at level of antennal insertion more strongly, acutely elevated, with tubercle slightly above level of antennal insertion. Pronotum as in vitiosus except surface smooth, shining, punctures near median base very small. Elytra as in vitiosus except declivity begin- ning at middle of elytra, more strongly 654 Great Basin Naturalist Vol. 43, No. 4 arched, steeper, erect interstrial setae slightly stouter. Female.— Similar to male except frons con- vex, its lateral margins unarmed by tubercles. Type locality.— Estacion de Biologia, Cha- mela, Jalisco, Mexico. Type material— The male holotype, fe- male allotype, and six paratypes were taken at the type locality on 4-III-1982, 100 m, S-365, from a Leguminosae, by A. Equihua. The holotype, allotype, and paratypes are in my collection. Chramesus tibialis, n. sp. Although the Scolytodes-\ike protibia is unique in the genus, this species is somewhat remotely allied to incomptus Wood. This and other unique characters are described below. Male.— Length 1.6 mm (paratypes 1.6-1.8 mm), 2.1 times as long as wide; color very dark brown to almost black. Frons broadly, moderately concave from epistoma to slightly below upper level of eyes; lateral margins rather abrupt, neither acute nor armed, epistoma normal; surface shining, subreticulate at vertex, gradually be- coming minutely subrugose toward epistoma. Vestiture fine, moderately long, mostly on margins. Pronotum 0.91 times as long as wide; shape typical of genus; surface finely reti- culate; median basal area with fine, shallow punctures, these replaced by small, rounded tubercles anteriorly and laterally. Vestiture of fine, slender hair. Elytra 1.3 times as long as wide; sides al- most straight and parallel on basal two-thirds, broadly rounded behind; striae not im- pressed, punctures shallow, small; interstriae smooth, shining, about three times as wide as striae, imiseriate punctures largely replaced by small granules. Declivity steep, rather nar- rowly convex; sculpture about as on disc ex- cept surface rather dull, granules smaller. Vestiture of minute strial hair and erect in- terstrial hairlike setae, each seta shorter than distance between rows. Protibia with outer apical angle produced into dominant spine somewhat similar to Scolytodes, two minute socketed denticles on lateral margin above spine. Female.— Similar to male except frons convex. Type locality.— Urpanapan, Veracruz, Mexico. Type material.— The male holotype, fe- male allotype, and one male paratype were taken at Hidalgotitlan at the type locality, 27-IV-1982, S-442, from Olmeca recta, by T. H. Atkinson. The holotype, allotype, and paratype are in my collection. Cnemonyx equihuai, n. sp. This species is distinguished from liratus Wood by the very different frons as de- scribed below, by the less deep, oval pronotal punctures, and by the somewhat more broad- ly flattened lower declivity. Male.— Length 1.4 mm (paratypes 1.5 mm), 2.5 times as long as wide; color yellow- ish brown. Frons convex except median third con- cavely impressed on triangular area from epistoma to upper level of eyes, concave area glabrous and reticulate except lateral margins with a row of rather fine, moderately long setae, lower margin of concavity marked by a low, straight, acute carina. Pronotum about as in liratus except surface slightly shagreened, punctures oval, less strongly impressed. Elytra similar to liratus except on disc striae less distinctly impressed, punctures not as close, declivity much more broadly convex on lower half, not as steep, tubercles sim- ilarly placed but averaging smaller, particu- larly in lateral areas; vestiture stouter, about half as long. Female.— Similar to male except frontal impression very weak. Type locality.— Km 150 carr. Melaque— Puerto Vallarta, Jalisco, Mexico. Type material— The male holotype, fe- male allotype, and one male paratype were taken at the type locality on 6-III-1982, 300 m, S-383, Hura polyandra, A. Equihua. The holotype, allotype, and paratype are in my collection. Cnemonyx evidens, n. sp. This species is distinguished from vaga- bundus Wood by the larger size, by the reti- culate, more shallowly, more broadly im- October 1983 Wood: American Bark Beetles 655 pressed frons, by the more closely spaced in- terstrial punctures, and by the declivital sculpture. Male.— Length 1.8 mm (paratypes 1.7-1.9 mm), 2.5 times as long as wide; color very dark reddish brown. Frons very shallowly concave almost from eye to eye from epistoma to slightly above upper level of eyes; surface reticulate, punc- tures minute, obscure; epistoma shining, slightly elevated, a feeble, transverse carina on its lower margin; vestiture on median two-thirds of lower two-thirds except re- duced to almost absent on and near median line, consisting of abundant, stout, erect setae, each slightly longer than distance equal to width of scape. Pronotum similar to vagabundus, except pimctures slightly smaller. Elytra outline about as in vagabundus; striae weakly impressed toward declivity, punctures small, moderately deep; interstriae twice as wide as striae, feebly convex, almost smooth, shining, punctures fine, distinctly im- pressed, almost uniseriate, more closely spaced than those of striae. Declivity convex, rather steep; striae narrower and more deep- ly impressed than on disc, interstriae more strongly convex, 1 slightly, 7 and 9 more dis- tinctly elevated, 7 and 9 joining and contin- uing almost to 1; punctures on all interstriae largely replaced by fine, pointed tubercles, costal margin near apex finely serrate. Vesti- ture almost obsolete, consisting of very min- ute, rather stout interstrial setae. Female.— Similar to male except frontal impression slightly less extensive, frontal ves- titure slightly less abundant. Type locality.— Las Granjas, Morelos, Mexico. Type material— The male holotype, fe- male allotype, and six paratypes were taken at the type locality on 8- VI- 1982, in Ficus, by E. Martinez. The holotype, allotype, and paratypes are in my collection. Cnesinus comutus, n. sp. This species is distinguished from bicornis Wood by the smaller size, by the less exten- sively, less deeply impressed frons, and by very different armature of the epistoma. Female.— Length 2.8 mm (paratypes 2.8-2.9 mm), 2.7 times as long as wide; color dark reddish brown. Frons strongly, broadly impressed to upper level of eyes (otherwise about as in bicornis); epistoma on median fourth strongly elevated into an almost hornlike process, this process as high as wide and equal in length to com- bined width of four facets of eye, its apex armed by a pair of small, transversely ar- ranged tubercles. Pronotum about as in bicornis except grooves between longitudinal elevations on disc slightly wider and somewhat subreticulate. Elytra as in bicornis except ground vesti- ture slightly finer and shorter, erect setae very slightly stouter. Male.— Similar to female except epistomal armature absent; vestiture apparently slightly longer and more abundant. Type locality.— San Tlatotico, Morelos, Mexico. Type material.— The female holotype, male allotype, and four paratypes were taken at the type locality on 27 May 1982, 2110 m, S-675, from a Compositae, by A. Burjos and E. Saucedo. The holotype, allotype, and paratypes are in my collection. Cnesinus nebulosus, n. sp. This species is distinguished from carinatus Wood by the very different female frons and sculpture of the pronotum as described below. Female.— Length 2.4 mm (paratypes 2.4-2.7 mm), 2.3 times as long as wide; color dark reddish brown, vestiture pale except tan on declivity. Frons similar to carinatus except weak ca- rina poorly formed, area above carina broad- er and distinctly impressed, more coarsely, closely, uniformly punctured; vestiture long- er, more uniformly distributed, less special- ized; eyes separated by 2.0 times width of an eye. Pronotum similar to carinatus except rugae higher, shorter, much more tortuous. Elytra similar to carinatus except not im- pressed or sulcate on declivity, ground setae stouter, erect setae stouter and shorter, not 656 Great Basin Naturalist Vol. 43, No. 4 longer on declivity and present on declivital interstriae 1 and 2; vestiture pale on the disc and sides, tan on declivity. Male.— Similar to female except frons shal- lowly impressed on lower half, carina absent, frontal setae shorter. Type locality.— Pachuca, Hidalgo, Mexico. Type material- The female holotype, male allotype, and one female paratype were taken at the type locality on 2 April 1982, 2400 m, S-463, by A. Equihua. The holotype, allotype, and paratype are in my collection. Cnesinus parvicornis, n. sp. This species is distinguished from other members of the elegans group by the more extensive base and the more dorsal position of the epistomal tubercles, by the coarse, al- most oval pronotal punctures, and by the uni- formly rather short, almost scalelike elytral setae. Female.— Length 2.8 mm (paratypes 2.8-3.3 mm), 2.3 times as long as wide; color reddish brown. Frons moderately impressed on slightly more than lower half, impressed area partly filled by a low, triangular elevation arising on median half of episoma and extending dorsad almost to upper limits of impressed area; this elevated area armed by a pair of basally sep- arate, small tubercles in a slightly more dor- sal position than in related species; upper area convex, shining, impunctate in central area; vestiture of short, stout setae generally distributed except in upper impunctate area. Pronotum 1.0 times as long as wide; sur- face smooth, shining, punctures rather coarse, elongate-oval, separated transversely by di- ameter of a puncture, longitudinally by one- fourth that distance; glabrous, except at margins. Elytra 1.6 times as long as wide, 1.9 times as long as pronotum; sides straight and paral- lel on more than basal two-thirds, broadly rounded behind; striae narrowly impressed, punctures small, shallow, spaced by one and one-half diameters of a puncture; interstriae two to three times as wide as striae, weakly convex, almost smooth, shining, punctures al- most uniseriate, rather small, their anterior margins weakly subcrenulate. Declivity steep, convex, except shallowly sulcate on lower half between interstriae 3; sculpture about as on disc except interstrial tubercles not evident. Vestiture of minute strial hair and erect interstrial setae, these one-ranked on interstriae 1, 3-ranked on others, middle rank pale tan and half as long as distance be- tween rows and slightly longer than pale marginal rows; all setae of uniformly short length throughout. Male.— Similar to female except frontal elevation feeble, tubercle absent. Type locality.— Ruinas de Xochicalco, Morelos, Mexico. Type material.— The female holotype, male allotype, and nine paratypes were taken at the type locality on 21 February 1982, 1200 m, S-323, by T. H. Atkinson and A. Equihua. The holotype, allotype, and paratypes are in my collection. Dendrosinits mexicanus, n. sp. This species is distinguished from globosus Eichhoff by the shallowly concave, more coarsely pvmctured frons and by the much more coarsely, deeply punctured pronotum. Male.— Length 3.5 mm (paratypes 3.0-3.9 mm), 2.3 times as long as wide; color black, with dark vestiture. Frons very shallowly, broadly concave from epistoma to vertex; surface smooth, shining, and densely, rather coarsely punc- tured, except impunctate along epistomal margin and on median line on lower half; vestiture mostly pale, rather abundant, much longer than in globosus, setae equal in length to almost one-third distance between eyes. Antennal club slightly wider than in globosus. Pronotum as in globosus except punctures distinctly larger and deeper and anterolateral areas always with two clusters of asperities (usually three in each cluster). Elytra as in globosus except vestiture more slender. Female.— Similar to male in all respects ex- cept for segmentation of abdominal terga. Type locality.— Estacion de Biologia, Cha- mela, Jalisco, Mexico. Type material.— The male holotype, fe- male allotype, and six paratypes were taken on 5-II1-1982, 60 m, S-372, by A. Equihua. October 1983 Wood: American Bark Beetles 657 The holotype, allotype, and paratypes are in my collection. Liparthrum mexicanum, n. sp. This species is distinguished from thevetiae Wood by the presence of six crenulations on the base of each elytron, by the much more closely set interstrial scales, and by the more slender pronotum. Male.— Length 0.9 mm (paratypes 0.9-1.0 mm), 2.4 times as long as wide; color brown. Frons as in thevetiae. Pronotum 0.9 times as long as wide; more narrowly rounded in front and asperities dis- tinctly larger than in thevetiae. Elytra 1.5 times as long as wide; about as in thevetiae except strial punctures more deeply impressed, interstrial scales shorter, wider, much closer, spaced within a row by length of a scale; slender interstrial setae as long as scales, usually alternating with them on disc but not on declivity. Female.— Similar to male except pronotal asperities mostly reduced, those on anterior margin absent. Type locality.— Cuernavaca, Morelos, Mexico. Type material- The male holotype and two paratypes were taken at the type locality on 27-X-1982, 1670 m, B-077, by A. Burjos and E. Saucedo. The allotype and three para- types are from Jesu. Sta. Ma. Chihuappa, Tlaclizapan, Morelos, 3-XII-1982, 1000 m, B-122, by the same collectors. The holotype, allotype, and paratypes are in my collection. Liparthrum pruni, n. sp. This species is distinguished from alboseto- sum (Bright) by the smaller, shallower strial punctures, by the shorter, stouter interstrial scales, and by other characters cited below. Male.— Length 1.1 mm (paratypes 1.0-1.3 mm), 2.2 times as long as wide; color black. Frons convex, about as in albosetosum. Pronotum about as in albosetosum except more strongly convex, asperities distinctly larger. Elytra about as in albosetosum except strial punctures much smaller, not as deep, inter- strial setae shorter, each about as wide as long, spaced within a row by distances equal to about one and one-half times length of a scale. Female.— Similar to male except pronotal asperities smaller. Type locality.— Aranza, Michoacan, Mexico. Type material— The male holotype, fe- male allotype, and 14 paratypes were taken at the type locality on lO-VII-1982, S-756, Pruniis serotina, by A. Equihua. The holotype, allotype, and paratypes are in my collection. Ten specimens that probably belong to this species are from El Tuito, Jalisco, Mexico, 28-V-1982, 640 m, S-710, T. H. Atkinson and A. Equihua. Phloeotribus geminus, n. sp. This species is distinguished from demissus Blandford by the more coarsely punctured pronotum, with no granules on the disc, by the more slender pronotal and elytral vesti- ture, and, in the male, by the presence of a transverse, epistomal carina and a pair of small tubercles on the lateral margin of the frons at the level of the antennal insertion. The Acatlan series was erroneously reported as demissus in my monograph. Male.— Length 2.1 mm (paratypes 1.7-2.1 mm), 2.0 times as long as wide; color very dark brown to almost black, vestiture pale. Frons more narrowly but as deeply im- pressed as in demissus, impression ending slightly below upper level of eyes; epistoma armed on median third by a low, acute, trans- verse carina; lateral margins at level of an- tennal insertion armed by a pair of small tu- bercles as in many other species of this genus. Segments of antenna club much more strong- ly produced than in demissus, each about nine times as wide as long. Pronotum as in demissus except surface smooth, shining, punctures larger, more sharply, more strongly impressed, with no granules on disc. Elytra about as in demissus except inter- strial granules smaller, interstriae 9 slightly more strongly, acutely elevated in declivital area, apical margin from level of striae 3 to suture more strongly serrate, vestiture slightly more slender and very slightly longer. 658 Great Basin Naturalist Vol. 43, No. 4 Female.— Similar to male except frons con- vex, tubercles absent; pronotal and elytral vestiture more slender. Type locality.— Acatlan, Jalisco, Mexico. Type material— The male holotype, fe- male allotype, and four paratypes were taken at the type locality 3- VII- 1965, 1300 m. No. 158, from Ficus, by me. Eight paratypes are from Estacion de Biologia, Chamela, Jalisco, l-VII-1982, 110 m, S-731, by A. Equihua. The holotype, allotype, and paratypes are in my collection. Pycnarthrum amersum, n. sp. This species is distinguished from brosimi Wood by the larger size and stouter form, by the coarser vestiture, and by the evenly con- vex declivity. Male.— Length 2.0 mm (paratypes 2.0-2.3 mm), 2.0 times as long as wide; color brown, vestiture pale. Frons similar to brosimi except more strongly flattened over larger area; eyes sepa- rated by 1.8 times width of an eye (1.0 in brosimi). Pronotum similar to brosimi except vesti- ture much coarser. Elytra resembling brosimi except declivity convex, not impressed, interstriae without tu- bercles, erect interstrial bristles much stouter and strongly confused on 2, less confused on 3, minute ground setae stouter; discal striae 1 impressed, punctures on 1 and 2 slightly larger, deeper. Female.— Similar to male except frons convex. Type locality.— Tenacatita, Jalisco, Mexico. Type material— The male holotype, fe- male allotype, and six paratypes were taken at the type locality on 4-II-1983, 40 m, S-883, Brosimum alicastrum, T. H. Atkinson and N. Bautista. The holotype, allotype, and paratypes are in my collection. Scolytodes plumericolens, n. sp. This species is distinguished from plu- meriae W^ood by the smaller size, by the more slender body form, and by numerous other characters, some of which are treated below. Female.— Length 1.5 mm (paratypes 1.5-1.7 mm), 2.3 times as long as wide; color almost black. Frons resembling plumeriae except some- what more strongly convex, surface punc- tured throughout (without an impunctate area), vestiture much less abundant, finer, ending well below upper level of eyes on a narrower area. Pronotum 1.0 times as long as wide; sur- face uniformly reticulate, punctures con- spicuously smaller than in plumeriae. Elytra 1.4 times as long as wide; about as in plumeriae except minute interstrial punc- tures almost uniseriate; very minute, erect in- terstrial hair present. Type locality.— Estacion de Biologia, Cha- mela, Jalisco, Mexico. Type material— The female holotype and two female paratypes were taken at the type locality on 2- VII- 1982, 90 m, S-736, Fhi- meria rubra, A. Equihua. The holotype and paratypes are in my collection. Scolytodes retifer, n. sp. This species is distinguished from ficivorus Wood by the larger size, by the reticulate elytra, and by the very different female frons as described below. Female.— Length 2.0 mm (paratypes 1.8-2.2 mm), 2.2 times as long as wide; color brown to dark brown. Frons flattened on an ovate area from eye to eye from epistoma to vertex (stronger and more extensive than in ficivorus); oval area on central third of lower half impunctate, glaborous, reticulate, remaining areas closely, finely punctured and setose; vestiture con- sisting of abundant, long, subplumose, yellow hair, longest setae equal in length to more than half distance between eyes. Pronotum and elytra strongly reticulate, very similar to reticiilatus Wood except all punctures much smaller and anterior margin of pronotum neither costate nor serrate. Sub- glabrous, a very few hairlike setae on odd- numbered interstriae. Male.— Similar to female except frons con- vex, of uniform sculpture, setae sparse, inconspicuous. October 1983 Wood: American Bark Beetles 659 Type locality.- Texeal, Mpiotepoztlan, taken on l-XI-1982, 1710 m, B-082, Ceiba, Morelos, Mexico. A. Burjos. Type material- The female holotype, The holotype, allotype, and paratypes are male allotype, and eight paratypes were in my collection. PLANT COMMUNITY VARIABILITY ON A SMALL AREA IN SOUTHEASTERN MONTANA James G. MacCracken^'^, Daniel W. Ureslc', and Richard M. Hansen' Abstract.— Plant communities are inherently variable due to a number of environmental and biological forces. Canopy cover and abovegroimd biomass were determined for understory vegetation in plant communities of a prairie grassland— forest ecotone in southeastern Montana. Vegetation units were described using polar ordination and stepwise discriminant analysis. Nine of a total of 88 plant species encountered and cover of litter were the most useful variables in distinguishing among vegetation luiits on the study area and accounted for nearly 100 percent of the variation in the data. Seven vegetation units were different (P < 0.05) after all 10 variables had been entered into the analysis. Some plant communities were represented by two or three different vegetation units, indicating that some plant communities were variable and nonuniform in botanical composition over a relatively small area. This variability will influence management practices for these areas. Multiple-use management will benefit by recog- nition of inherent plant community variation. Mueller-Dombois and Ellenberg (1972) de- fined plant communities as concrete defin- able units of vegetation that can be recog- nized and are obvious to the eye. Plant commimities are often named after species that contribute to their unique structure or composition, or they are named after a unique environmental condition. Some exam- ples from southeastern Montana include sage- brush-grassland, pine forest, and riparian commimities. However, plant communities are variable and can be a mosaic of finer units of vegetation. Poore (1955) termed these vegetation abstractions noda, and they are presumably analagous to Whittaker's (1967) ecological groups. The variability within plant communities at any time is due to a number of environ- mental and biological forces. Environmental influences include the geology of an area, soil communities, climate, solar radiation, and fire. Biological influences can be soil mi- crobes, grazing animals, intra- and inter- specific competition, genetics, successional patterns, and evolution. These forces create a dynamic process of vegetation patterning. Within a person's lifetime, however, plant commimities are relatively stable, barring ca- tastrophic events. Variations within plant communities have long been recognized. Gleason (1926) stated that no two plant communities are exactly alike even though they contain the same spe- cies. Whittaker (1970) noted that plant com- munities are often less than discrete units, with no absolute boundaries among commu- nities. Other plant ecologists have come to similar conclusions (Curtis and Mcintosh 1950, Cottam 1949, Goodall 1953). Mueller- Dombois and Ellenberg (1972), however, sug- gested that plant communities can be indi- vidualists as well as continua. One aspect of current vegetaton ecology is the study of community variability and how that relates to the consequences of land management and the effects of human technology. Plant community variability can create problems for land managers regardless of the source of variability. Successful management of vegetation for livestock grazing, wildlife habitat, water yield, soil conservation, etc., requires knowledge of plant community vari- ability. Different vegetation units will not re- spond similarly to management. Practices recommended for one situation may be un- successful in another, even though the plant community appears to be the same. Many hectares of native rangeland are being ma- nipulated primarily to increase the number of livestock supported, while still maintaining a viable ecosystem. 'Department of Range Science, Colorado State University, Fort Collins, Colorado 80523. 'Present address: University of Alaska, Agricultural Experiment Station, Palmer Research Center, P.O. Box AE, Palmer, Alaska 99645. "USDA, Forest Service, Rocky Mountain Forest and Range Experiment Station, SDSM&T, Rapid City, South Dakota 57701. 660 October 1983 MacCracken et al.: Plant Community Variability 661 Much rangeland in the western United States, including southeastern Montana, is without acceptably published information on vegetation characteristics. We believe that recent quantitative practices in plant ecology can and should be applied to management problems at the local level. The purposes of this paper are (1) to pre- sent a method of assessing plant community variation, (2) to illustrate the variability with- in plant communities on a small study site, and (3) to identify potential consequences of plant community variation for management practices. Study Area and Methods The study was conducted on, about 11,300 ha of rangelands along the northern edge of the Black Hills in southeastern Montana. The study area was immediately west of Alzada, Carter County. Elevation ranged from 1036 to 1128 m and average annual precipitation is approximately 37 cm. Soils included alluvial clayey deposits in bottom areas and shale at higher elevations. Surface deposits of bentonite clay were nu- merous. Bentonic soils are characterized by a shallow A horizon and are saline or sodic (Bjugstad et al. 1981). Most of the area was in private ownership and grazed by both sheep and cattle on a rest rotation system. Mule deer {Odocoileus he- mionus), white-tailed deer (O. virginianus), and pronghom {Antilocapra americana) were present on the study area. Southeastern Montana is classified as a wheatgrass-needlegrass (Agropyron-Stipa) prairie by Kiichler (1964). Garrison et al. (1977) classified the study area as plains grassland with ponderosa pine {Pinus ponde- rosa) forest. Plant names follow those given by Scott and Wasser (1980). Four plant communities were recognized on the study area. A sagebrush-grassland community occupied a majority of the area. This community was dominated by big sage- brush {Artemisia tridentata) and buffalo grass {Buchhe dactyloides). A riparian community, primarily wooded stream bottoms, was the next most abundant plant community. Major plants there were boxelder maple {Acer ne- gundo) and snowberry {Symphoricarpos spp.). A pine forest community existed at higher elevations, consisting of ponderosa pine and western wheatgrass {Agropyron smithii). Iso- lated portions of the study area were open grassland. The most abundant plants there were western wheatgrass and needleleaf sedge {Carex eleocharis). These subjective classifications were made to facilitate design of an adequate sampling scheme. Four sample sites were selected in both the sagebrush and riparian commimities. Two sample sites were studied in the pine forest and two in the grassland community. These sites were judged to be representative of their respective plant communities, and en- compassed the range of perceived variability within these communities. The number of sampling sites established in each plant com- inimity was based on the total area occupied by that community, and/or the observed variability within each community. Canopy cover and aboveground biomass of plant species were estimated in each sam- pling site during summers of 1979 and 1980. Three parallel 50-m line transects were sys- tematically established approximately 30.5 m apart at each site. Canopy cover was mea- sured using 50 plots (2x5 dm) system- atically spaced at 1-m intervals along each transect (Daubenmire 1959). Six hundred plots were observed in both riparian and sagebrush areas and 300 in grassland and pine forest communities each year of the study. We assessed the adequacy of our sample size using the formula presented by Johnson and Lay cock (1972), with a degree of precision needed to estimate plant species within 15 percent of their mean with 95 percent confidence. Aboveground biomass at peak growth was estimated by clipping 20 plots at 5-m inter- vals along two of the transect lines at each site. All plants, excluding shrubs, were clipped at ground level, air dried for two weeks, oven dried at 60 C for 24 hours, then weighed to the nearest one-tenth gram. Individual transects of each year were grouped into similar vegetation units using multidimensional polar ordination (Bray and Curtis 1957), as described by Mueller- Dombois and EUenberg (1974). Ordination axes endpoints were chosen using guidelines and criteria suggested by Mueller-Dombois 662 Great Basin Naturalist Vol. 43, No. 4 100 r 80 c (D U O Q. 50 20 20 40 50 60 Percent Dissimilarity 80 100 Fig. 1. Location of vegetation transects along ordination axes and grouping of transects into vegetation groups. R = riparian, P = pine forest, G = grassland, and S = sagebrush. and EUenberg (1974) and Newsome and Dix (1968). For this analysis transects were ordi- nated based on canopy cover estimates. Such an approach provided for the assessment of variability among transects and sampling sites within a plant community. Gauch et al. (1977) found that polar ordination was sub- ject to less distortion than other ordination procedures when sampling is clustered and outlier samples are included. The vegetation groups were then analyzed with stepwise discriminant analysis for three reasons (Cooley and Lohnes 1962, Klebenow 1969, Klecka 1975). Since polar ordination is somewhat subjective and based on sample similarities with axes endpoints, some vegeta- tion groups may be erroneous. Most cluster- ing techniques do in fact derive non- significant groups (Strauss 1982). Dis- criminant analysis maximizes differences among groups and was used to determine if vegetation groups were significantly different from one another. Green (1980) suggested that multivariate tests were so powerful in detecting differences that a nonsignificant re- sult may be more meaningful than a signifi- cant result. Discriminant analysis also selects the set of variables (plant species) that are the most useful in differentiating among groups. This property is desirable in that many plant species are encountered that added little to explaining variation within and among plant communities. Discriminant analysis also generates classification functions from the most useful variables. These func- tions can be used to determine the vegetation groups to which nonsignificant groups were most similar. The classification functions could also be used to assign samples from fu- ture surveys to the vegetation group they most nearly resemble (MacCracken and Han- sen 1982). Aboveground biomass was analyzed by testing for differences between vegetation units (as determined by ordination and dis- criminant analysis), categories (grasses and forbs), and years, using a three-way analysis of variance test followed by Duncan's new multiple range test. Differences were consid- ered significant at a = 0.05. Results The number of plots needed to estimate canopy cover of plants with the degree of precision stated was 1025 and 1098 in 1979 and 1980, respectively. Our observation of 1800 plots per year was more than adequate. Ordination arranged the 72 transects in such a manner that eight groups could be delineated based on the proximity of tran- sects from similar sample sites (Fig. 1). Sage- brush and pine communities were each rep- resented by two groups, riparian commu- nities by three, and the grassland community October 1983 MacCracken et al.: Plant Community Variability 663 by one. Discriminant analysis indicated that the two sagebrush groups were not distinct (P > 0.05). As a result, both sagebrush groups were combined for final analysis, resulting in seven groups defined at this point as seven vegetation units. Eighty-eight plant species were encoun- tered along the transects; of these, 9 plant species and percent cover of litter were the most useful variables in discriminating be- tween vegetation units and accounted for nearly 100 percent of data variation (Table 1). Those discriminating variables, in order of significance, were smooth brome (Bromus in- ermis), litter, starry cerastium {Cerastium ar- vense). Rocky Mountain juniper (Juniperus scopulorum), snowberry (Symphoricarpos spp.), red threeawn (Aristida longiseta), big Table 1. Mean percent canopy cover of plant species, bare ground, and litter of seven vegetation units in south- eastern Montana. Estimates were taken during summers 1979 and 1980. Categories Vegetation units Riparian Grassland Pine Sagebrush 1 2 3 1 2 8 8 4 23 16 8 20 29 22 44 28 53 48 15 64 81 67 41 16 44 64 2 16 3 16 4 3 7 10 8 • • " Bare ground Litter + Total cover Grasses Agropyron S7nithii + Agropijron spp. Aristida longiseta + Boiiteloiia gracilis Bromus inermis-'t- B. japoniciis B. tectoriim Btichloe dactyloides Carex spp. Calanwvilfa longifolia Elymus macounii Hordeiim jubattim Koeleria cristata Miihlenbergia richardsonus Panictim capillare Phleum pratense Poa spp. Schedonnardits panicttlatus Stipa viridula Unidentified FORBS Achillea millefolium Cerastium arvense + Geum aleppicum Lactuca serriola Plantago spinulosa Rumex crispus Sphaeralcea coccinea Taraxicum officinale Thlaspi arvense Vicia americana Shrubs Artemisia tridentata + Juniperus scopulorum + Opuntia polycantha + Phlox hoodii Sarcobatus vermiculatus Symphoricarpos spp. -t- Rosa spp. -I- 21 27 1 14 1 1 1 1 5 17 5 59 1 • 1 4 2 1 1 . 1 o 1 • e 1 3 20 5 3 14 5 1 1 • 2 1 11 1 1 2 • 2 + Indicates plant species and variables most useful in discriminating between the seven vegetation units. 664 Great Basin Naturalist Vol. 43, No. 4 sagebrush, western wheatgrass, rose {Rosa spp.), and plains prickly pear {Opuntia poly- acantlm). Smooth brome, litter, and starry ce- rastium alone accounted for 94 percent of data variation. However, the remaining seven variables contributed significantly to the sep- aration of vegetation units. Discriminant function classification coefficients ranged from -0.01 to 56.42 (Table 2). Differences among the seven vegetation units (P < 0.05) arose as each variable was entered into discriminant analysis. These dif- ferences changed slightly as each variable was considered, but the seven units were dis- tinct (P < 0.05) after all 10 variables had been considered. Transects from each site combined into the same vegetation unit for each year, indicating that differences in plant canopy cover were not significant between years. Generally, transects from one or more sampling sites combined to produce a vegeta- tion unit. Nevertheless, there was some mix- ing of transects from the four riparian sites sampled among the three Riparian vegetation units. Differences were detected in aboveground biomass among units and plant taxa (Table 3). The Riparian 3 unit had more grass (P < 0.01) than the Riparian 1 and 2 units, sage- brush, grassland, and both pine units. Grass biomass was also greater (P < 0.01) in the Riparian 2 unit, sagebrush and grassland units than in both pine forest units. Still, forb bio- mass between units was similar (P > 0.05). Grass biomass was higher than forb biomass (P < 0.01) in all riparian units. Grass and forb biomass in other units were similar (P > 0.05). No year differences were observed for total biomass (P > 0.05) for any vegetation unit or category. Some plant species were common to all units, but others were in- dicative of a particular vegetation unit. Western wheatgrass was most abundant in the Grassland and Riparian 2 units. Red threeawn and starry cerastium were confined to pine forest areas. Smooth brome and snow- berry occurred exclusively in riparian units, as did combined wheatgrasses. Common tum- blegrass (Shedonnardis paniculatus), and plains prickly pear were useful in dis- tinguishing the grassland unit, and big sage- brush was dominant in the sagebrush unit. Discussion Vegetation units as defined in this study represent areas that are the most similar in vegetative composition. Variation inherent in sampling methods has been reduced to a minimum by the quantitative techniques used, and accurately describes these vegeta- tion units at a refined level. The methods used illustrate the variation from site to site within some plant communities. Discriminant analysis indicated that relatively few plant species accounted for the majority of varia- tion attributable to differences in plant cover among the vegetation units. Table 2. Discriminant function coefficients for the 10 variables most useful in distinguishing between vegetation units in southeastern Montana. Vegetation units Riparian Grassland Pine 1 2 Sagebrush Variables 1 2 3 Litter 1.63 0.92 1.19 1.34 4.42 4.49 0.82 Constant -32.87 -28.74 -57.49 -22.11 174.09 -185.15 -12.31 Grasses Agropyron smithii 0.08 0.97 -0.31 0.09 -1.01 -1.03 -0.04 Aristida longiseta 15.39 7.47 11.84 12.61 56.42 46.68 8.13 Brovius inermis -0.66 -0.43 1.01 -0.54 -1.71 -1.74 -0.32 FORBS Cerastium arvense 12.89 6.88 9.66 10.73 34.38 49.26 6.06 Shrubs Artemisia trideritata 0.19 0.14 0.13 -0.01 0.48 0.51 0.88 Juniperus scopulortim 11.86 6.49 8.82 9.89 36.15 33.83 6.00 Rosa spp. 0.01 1.39 -0.50 -0.06 -1.72 -1.76 -0.16 Symphoricarpos spp. 0.53 0.61 -0.04 0.07 -0.12 -0.12 0.01 Opuntia polyacantha -1.54 -2.55 -0.44 0.17 -1.81 -1.94 -0.53 October 1983 MacCracken et al.: Plant Community Variability 665 Polar ordination arranged transects along a moisture gradient for both axes. Vegetation units representing areas of high soil moisture (based on plant species presence) fell into the bottom left quadrant and xeric vegetation units fell into the upper right quadrant (Fig. 1). Many studies have shown strong correla- tions between plant community composition and soil moisture regimes in the western United States (Dahl 1963, Galbraith 1971, Marks and Harcombe 1981, Monk 1960, Marks and Harcombe 1975, Harniss and West 1973). Marks and Harcombe (1981) in- terpreted an ordination axis as representing a soil moisture gradient even through they did not measure soil moisture directly. Some plant commimities in southeastern Montana are relatively homogenous. The sagebrush-grass and grassland communities were not different in plant cover among the sites sampled within each type. We did sub- jectively divide the sagebrush-grass transects into two groups based on ordination results; however, discriminant analysis did not detect any differences (P > 0.05) in plant cover among the two groups. Polar ordination when used as a clustering technique can pro- duce nonsignificant groups. Current studies in plant community classification often use an ordination or clustering technique to de- fine plant community types (Marks and Har- combe 1981, Thilenius 1972, Severson and Thilenius 1976). Rarely are the groups that result from these techniques tested for signifi- cance (Strauss 1982). Riparian and pine forest communities are relatively heterogenous in southeastern Mon- tana. The variation and factors producing dif- Tabije 3. Mean kilograms per hectare of grasses and forbs occurring in seven vegetation units in southeastern Montana. Estimates were taken during summers of 1979 and 1980. Vegetation units Riparian Grassland Pine Plant species Sagebrush 1 2 48 59 125 5 4 6 6 • 30 21 14 2 2 8 4 3 5 17 115 66 86 Grasses Agmpijron smithii Agropyron spp. Aristida longiseta Bouteloua gracilis Bromus inennis B. japonicus B. tectorum Buchloe dactyloides Carex spp. Cahmovilfa longifolia Elymus macounii Hordeitm jubattnn Koeleria cristata Muhlenbergia richardsonus Pcinicwn capillare Phleum prutense Poa spp. Schedonnardus paniculatus Stipa viridula Others Total Grass Forbs Achillea millefolium Cerastium arvense Geum aleppicum Lactuca serriola Plantago spinulosa Rumex crispus Sphaeralcea coccinea Taraxicum officinale Thlaspi arvense Vicia americana Total Forb 75 24 192 109 1 14 11 5998 13 22 » 25 O 137 263 179 3 2 50 106 156 802 7 • 1 20 22 21 7 6 25 1 3 7 222 6 1 1 7 3 5 2 78 1 12 1 3 • 2 6 29 30 36 86 2 20 10 e 18 10 20 11 16 666 Great Basin Naturalist Vol. 43, No. 4 ferences among sites in riparian and pine for- est communities are recognizable and inter- pretable. For example, riparian communities (i.e., hardwood forests along stream bottoms) were divided into three distinct vegetation units in this study (Table 1 and 3, Fig. 1). In general, hardwood forests, occurring as woody draws and stringer woodlands, are de- clining on the northern Great Plains (Boldt et al. 1978). Declining woodlands are represent- ed by trees of old age, decadence, and ad- vanced stages of breakup. Reproduction is poor and ground cover is primarily her- baceous. In contrast, "healthy" woody draws are characterized by thrifty, moderately dense stands of trees, and a vigorous shrub understory (Boldt et al. 1978). The Riparian 1 unit was representative of a healthy area. ShRib cover averaged 27 percent, and her- baceous vegetative growth averaged approx- imately 117 kg/ha. The Riparian 3 unit rep- resented a declining woodland. Shrub cover averaged 1 percent, and herbaceous growth averaged 817 kg/ha, primarily because of the invasion of smooth brome from nearby hay meadows. The decline of hardwood forests on the northern Great Plains has been attributed to a number of environmental and biological factors (Boldt et al. 1978). Two vegetation units were recognized in the pine forest community. These units are more easily interpreted than those of riparian sites. The Pine 1 unit had a relatively dense stand of trees (Table 4). Understory cover and aboveground biomass were lower than in the Pine 2 unit, but percent ground litter was higher in this unit. The Pine 2 unit had a rel- atively more open stand of trees, with greater growth of understory vegetation and less ground Htter. The difference in tree density between the two units was perhaps due to moisture regimes as related to aspect of the sites. Management Implications Results of this study show that some plant communities in southeastern Montana are variable in botanical composition, being composed of distinct and differing vegetation units. This variability can be attributed to a number of environmental or biological fac- tors. Different vegetation units within a plant community will respond differently to man- agement practices. This site-specific vari- ability, once recognized, will influence man- agement decisions. For example, consider an area of riparian community in southeastern Montana in which a rancher wishes to con- vert part to hay meadows. If all three Ri- parian units were present, the decadent woodland would probably be most easily converted. The healthy woodland would be valuable as wildlife habitat, to trap winter snows to fill stock ponds downstream, and as shading areas for livestock. The Riparian 2 unit, an intermediate unit between healthy and decadent stands, could be slated for im- provement toward a healthy stand. Boldt et al. (1979) presented treatments aimed at im- proving decadent woodlands on the northern Great Plains. The Pine 1 unit could be thinned to increase forage production for livestock, water yield, and timber production. By simply recognizing the inherent vari- ability in plant communities, a number of management options became apparent. On federal lands, where multiple use manage- ment is law, this approach should be readily utilizable. Table 4. Density (no/ha) of trees in pine and riparian vegetation units in southeastern Montana. Vegetation units Pine Riparian Tree 12 12 3 Pinus ponderosa Qiierciis macrocarpa Juniperus scopulorum Fraxiniis pennsylvanicas Acer negundo Primus virginiana Salix amygdaloides Total 172 192 88 452 20 52 184 256 600 524 5 1144 976 496 1472 872 260 352 1484 October 1983 MacCracken et al.: Plant Community Variability 667 Recognition of some vegetation units de- fined in this study may not be easy, especially the Riparian 2 unit. However, discriminant classification functions can be used for that purpose. Using estimates of mean percent canopy cover of discriminator species multi- plied by discriminant function coefficients (Table 2), a composite score can be derived by adding the products for any sample. The function producing the largest score indicates the vegetation unit from which the sample came. Since only ground litter and nine plant species were important in distinguishing among the vegetation units on the study area, only these variables need be measured in fu- ture surveys (MacCracken and Hansen 1982). This should greatly reduce field effort and as- sociated costs. The application of these classi- fication functions beyond the immediate study area is questionable. However, they may be suitable for portions of southeastern Montana where the same plant communities occur and environmental and biological forces similar to those at work here operate. Ideally, each vegetation unit defined in this study should be managed on an individ- ual basis, using practices known to benefit those units whether management be for live- stock, wildlife, water, or minerals. This would require intensive management to achieve desired results. Nevertheless, it is possible to classify existing areas based on unit dominance and manage for that imit. Acknowledgments The authors wish to thank L. E. Alexander, D. Ohgmiller, V. Todd, and M. Loring for their assistance in this study and the Fosters of Wyotana Ranch, G. Brimmer, and the Carlton Grazing Association for access to their properties. Literature Cited Bray, J. R., and J. T. Curtis. 1957. An ordination of the upland forest communities in southern Wiscon- sin. Ecol. Monog. 27:325-349. BjuGSTAD, A. J., T. Yamamoto, and D. W. Uresk. 1981. Upland shrub establishment on coal and ben- tonite clay mine spoils. Pages 104-122 in L. H. Stetter, E. S. Depuit, and S. A. Mikol, eds., Pro- ceedings of Shrub Establishment Conference, Laramie, Wyoming. BoLDT, C. E., D. W. Uresk, and K. E. Severson. 1978. Riparian woodlands in jeopardy on Northern High Plains. Pages 184-189 in R. R. Johnson and J. F. McCormick, eds., USDA, Forest Service, Gen. Tech. Rep. WO-12. 410 pp. 1979. Riparian woodland enclaves in the prairie draws of the Northern High Plains: a look at problems, a search for solutions. Pages 31-.32 in Great Plains Agriculture Council, Forestry Com- mittee, eds. Riparian and wetland habitats of the Great Plains: proceedings of the 31st annual meeting. Great Plains Agric. Counc. Publ. 91. 88 pp. Cooley, W. W., and p. R. Lohnes. 1962. Multivariate procedures for the behavioral sciences. Wiley and Son, New York. 374 pp. Cottam, G. 1949. The phytosociology of an oak woods in southwestern Wisconsin. Ecology 30:271-287. Curtis, J. T., and R. P. McIntosh. 1950. The inter- relations of certain analytical and synthetic phy- tosociological characters. Ecology 31:434-455. Dahl, B. E. 1963. Soil moisture as a predictive to forage yield for the sandhills range type. J. Range Man- age. 18:94-96. Daubenmire, R. 1959. A canopy-coverage method of vegetation analysis. Northwest Sci. 33:43-64. Galbraith, a. F. 1971. The soil-water regime of the shortgrass prairie ecosystem. Unpublished dis- sertation. Colorado State Univ., Fort Collins. 127 pp. Garrison, G. A., A. J. Bjugstad, D. A. Duncan, M. E. Lewis, and D. R. Smith. 1977. Vegetation and environmental features of forest and range eco- systems. USDA, Forest Service, Agric. Handbook 475. 68 pp. Gauch, H. G., Jr., R. H. Whittaker, and T. R. Wentworth. 1977. A comparative study of re- ciprocal averaging and other ordination tech- niques. J. Ecol. 65:157-174. Gleason, H. a. 1926. The individualist concept of the plant association. Bull. Torrey Botan. Club. 53:7-26. Goodall, D. W. 1953. Objective methods for the classi- fication of vegetation: 1. The use of positive in- terspecific correlation. Australian J. Bot. 1:39-62. Green, R. H. 1980. Multivariate approaches in ecology: the assessment of ecologic similarity. Ann. Rev. Ecol. Syst. 11:1-14. Harniss, R. O., and N. E. West. 1973. Vegetation pat- terns of the National Reactor Testing Station, southeastern Idaho. Northwest Sci. 47:30-43. Johnson, W. M., and W. A. Laycock. 1962. Kind, num- ber, and selection of livestock for grazing studies and animal measurements suited for evaluating results. Pages 137-142 in USDA, Forest Service, Misc. Publ. 940. Klebenow, D. a. 1969. Sage grouse nesting and brood habitat in Idaho. J. Wildl. Manage. 33:649-662. Klecka, W. R. 1975. Discriminant analysis. Pages 434-462 in N. H. Nie, C. H. Hull, J. G. Jenkins, K. Steinbrenner, and D. H. Bent, eds., Statistical package for the social sciences. Edition 2. McGraw-Hill Book Company, New York. 675 pp. Great Basin Naturalist Vol. 43, No. 4 KucHLER, A. W. 1964. Potential natural vegetation of the coiinterminous United States. Amer. Geo. Soc. Special Publ. .36. 1 16 pp. MacCracken, J. G., AND R. M. Hansen. 1982. Her- baceous vegetation of habitat used by blacktail jackrabbits and nuttall cottontails in southeastern Idaho. Amer. Midland Nat. 107:180-184. Marks, P. L., and P. A. Harcombe. 1975. Community diversity of coastal plain forests in southern east Texas. Ecology 56:1004-1008. 1981. Forest vegetation of the Big Thicket, south- east Texas. Ecol. Monog. 51:287-305. Monk, C. D. 1960. A preliminary study on the relation- ships between the vegetation of a mesic ham- mock community and a sandhill community. Quart. J. Florida Acad. Sci. 23:1-12. MuELLER-DoMBOis, D., AND H. Ellenberg. 1974. Aims and methods of vegetation ecology. Wiley and Sons, New York. 547 pp. Newsome, R. D., and R. L. Dix. 1968. The forest of the Cypre.ss Hills, Alberta and Saskatchewan, Can- ada. Amer. Midland Nat. 80:118-185. PooRE, M. E. D. 1955. The use of phytosociological methods in ecological investigations. J. Ecol. 43:226-651. Scott, T. G., and C. H. Wasser. 1980. Checklist of North American plants for wildlife biologists. Wildl. Soc, Wa.shington, D.C. 58 pp. Severson, K. E., and J. F. Thilenius. 1976. Classifica- tion of quaking aspen stands in the Black Hills and Bear Lodge Mountains. USDA, Forest Ser- vice, Res. Paper RM-166. 24 pp. Strauss, R. E. 1982. Statistical significance of species clusters in association analysis. Ecology 63:6.34-639. Thilenius, J. F. 1972. Classification of deer habitat in the ponderosa pine forest of the Black Hills, South Dakota. USDA, Forest Service, Res. Paper RM-91. 28 pp. Whittaker, R. H. 1967. Gradient analysis of vegetation. Biol. Rev. 42:207-264. 1970. Communities and ecosystems. MacMillan Co., London. 162 pp. NEW LEAFHOPPER SPECIES OF COELIDIA WITH A REVISED KEY AND NOTES ON HOMONYMY AND DISTRIBUTION (HOMOPTERA: CICADELLIDAE, COELIDIINAE) Mervin W. Nielson' Abstract.— Five new species of CocUdio are described and illustrated. These include panatnensis and simplex from Panama and retrorsa, cochloea, and tortiila from Brazil. A revised key is also presented for 13 of the 14 species for which males are known. Coelidia gladia is proposed as a new name for Coelidia spangbergi Nielson, 1982 nee Coelidia spangbergi Linnavuori, 1956 and Coelidia spangbergi Metcalf, 1964. The nominate genus Coelidia Germar of the subfamily Coehdiinae was treated in Part IV of my revision of the tribe Coelidiini (Nielson 1982). In that work* nine species were included in a conceptually restricted group that formerly encompassed over 200 species represented in all zoogeographical re- gions of the world. In this paper five new species are described with a revised key to 13 of 14 species for which males are known. A new name is proposed for Coelidia spang- bergi Nielson, 1982, preoccupied by Coelidia spangbergi Linnavuori, 1956, and Coelidia spangbergi Metcalf, 1964. The genus Coelidia is characterized as hav- ing a large elevated crown that is usually broader than the width of the eyes, carinate laterally, and produced distally beyond the anterior margin of the eyes. The clypeus has an incomplete median longitudinal carina in some species, including the type species, ve- nosa Germar. The clypeal carina is the pri- mary tribal character that separates Teruliini from Coelidiini. It is absent in all genera of the latter tribe except Clypeolidia Nielson and 4 of 14 known species of Coelidia, where it is present but incomplete, i.e., does not reach the transclypeal suture from its ante- rior origin. The genitalic characters of Coelidia in- clude a pair of prominent processes on the caudal margin of the male pygofer, usually very long styles, and an elongate aedeagus that usually has 1-2 distal processes or a re- curved extension of the shaft. The present distribution of the genus is Neotropical. Coelidia venosa is the only widespread species and it ranges from Brazil to Colombia. Four species occur in Brazil, four in Colombia, and four in Panama. One species is common to Brazil and Colombia and one is common to Colombia and Pan- ama, suggesting that Colombia is the center of the southern (Brazil) and northern range (Panama) of the genus. Key to Males of Coelidia^ 1. Aedeagus with 1-2 prominent distal or subdistal processes 2 — Aedeagus without such processes, if present, only about as long as wide 11 2(1). Aedeagus with 1 distal process 3 — Aedeagus with 2 distal processes 7 3(2). Style short, length about equal to arms of connective (Fig. 3) 4 — Style long, length much greater than arms of connective (Fig. 8) 5 'Department of Zoology and Life Science Museum, Brigham Young University, Prove, Utah 84602. ■Includes 13 of the 14 known species; stalii (Spangberg), known only from ? is not keyed. 669 670 Great Basin Naturalist Vol. 43, No. 4 4(3). Aedeagus with short distal process, process about 1/4 length of shaft (Fig. 862, Nielson 1982); pygofer with ornate caudodorsal process (Fig. 858, Nielson 1982) venosa Germar — Aedeagus with long distal process, process about 1/2 length of shaft (Fig. 4); pygofer with broad, simple caudodorsal process (Fig. 1) retrorsa, n. sp. 5(3). Pygofer with long, narrow, sharply pointed caudoventral process (Figs. 864 and 898, Nielson 1982) 6 — Pygofer with a long, narrow, but distally enlarged caudoventral process (Fig. 7) panamensis, n. sp. 6(5). Pygofer with very long caudoventral process, process extending distally beyond apex of caudodorsal process (Fig. 898, Nielson 1982); aedeagus with short distal process, process 2-3 times as long as wide in lateral view (Fig. 902, Nielson 1982) attenuata Nielson — Pygofer with short caudoventral process, process not reaching apex of caudo- dorsal process (Fig. 864, Nielson 1982); aedeagus with long distal process, process 5-8 times as long as wide in lateral view (Fig. 868, Nielson 1982) germari Nielson 7(2). Aedeagus with 1 distal process and 1 subdistal process, processes unequal in length and in configuration (Figs. 873, 876, 882, Nielson 1982) 8 — Aedeagus with 2 distal processes, processes nearly equal in length and in configuration (Fig. 17) tortula, n. sp. 8(7). Aedeagus with long subdistal process, apex reaching to about midlength of shaft in lateral view (Figs. 876 and 882, Nielson 1982) 9 — Aedeagus with short subdistal process, apex not reaching midlength of shaft in lateral view (Fig. 873, Nielson 1982) atra Walker 9(8). Style in dorsal view very narrow at distal 2/3, narrower than aedeagal shaft (Figs. 879 and 884, Nielson 1982) 10 — Style in dorsal view broad at distal 2/3, as broad as or broader than aedeagal shaft (Figs. 877, Nielson 1982) nigra (Spangberg) 10(9). Aedeagus with very broad subdistal process, process broader than aedeagal shaft in dorsal and lateral views (Figs. 881 and 882, Nielson 1982) gladia, n. name — Aedeagus with very narrow subdistal process, process narrower than aedeagal shaft in lateral view (Fig. 888, Nielson 1982) gorgonensis Nielson 11(2). Aedeagus and style narrowed distally (Figs. 21 and 23) 12 — Aedeagus and style greatly enlarged distally (Figs. 891 and 894, Nielson) hulhata Nielson 12(11). Pygofer with ornate caudodorsal process, process enlarged basally with slender curved distal process (Fig. 19) cochloea, n. sp. — Pygofer with long caudodorsal process, process narrow, fingerlike (Fig. 25) simplex, n. sp. Coelidia retrorsa, n. sp. wings flavous anteriorly, becoming spotted (Figs. 1-6) ^jj-Ji fuscous markings posteriorly. Length: $, 10.00 mm. Head small, much narrower than pro- General color deep ochraceous with fus- notum, anterior margin obtusely angulate; cous costa and 5 narrow longitudinal pale crown produced beyond anterior margin of flavous stripes on pronotum; veins of fore- eyes, broad, width greater than width of eyes. October 1983 Nielson: New Leafhopper Species 671 Figs. 1-6. Coelidia retrorsa: 1, Male pygofer, lateral view. 2, Connective and right style, dorsal view. 3, Style, lateral view. 4, Aedeagus, dorsal view. 5, Aedeagiis, lat- eral view. 6, Plate, ventral view. elevated above level of eyes, carinate later- ally, foveate on either side of middle, lateral margins parallel; eyes large, elongate-ovoid, occupying less than 2/3 of entire dorsal area of head; pronotum very large, scutellum large; forewing elongate, apex broadly an- gulate, venation typical, appendix well devel- oped; clypeus long and broad with an in- complete median longitudinal carina, extending from anterior margin to about 2/3 length of clypeus; clypellus long, narrowed basally, expanded distally. S . Pygofer in lateral view very narrow with long, narrow caudoventral process and moderately long, broad caudodorsal process; 10th segment long and narrow, without ven- tral processes; aedeagus asymmetrical, long, slightly tubular, distal part recurved and ex- tending to about midlength of shaft, shaft narrowed along recurved portion, wrinkled and enlarged subapically, becoming slightly hooked distally; gonopore medial on shaft; connective Y-shaped with short stem and long arms; style very short, about as long as Figs. 7-12. Coelidia panamensis: 7, Male pygofer, lat- eral view. 8, Connective and right style, dorsal view. 9, Style, lateral view. 10, Aedeagus, dorsal view. 11, Ae- deagus, lateral view. 12, Plate, ventral view. arms of connective; plate long, profusely setose. ? . Unknown. Holotype ( $ ), BRAZIL: Amazon, Tonan- tins, no date, no collector, (NR). Remarks: This species is similar in general habitus and some male genital characteristics to venosa Germar but can be distinguished by the long recurved portion of the aedeagus and by the gonopore that is medial on the shaft. Coelidia panamensis, n. sp. (Figs. 7-12) Length: $ 8.40 mm. General color deep fuscous with 5 narrow flavous longitudinal lines and broad flavous band on lateral margins of pronotum, veins of forewing with flavous spots. Head small, much narrower than pro- notum, anterior margin obtusely angled; crown distinctly produced beyond anterior margin of eyes, broad, width about equal to width of eyes, elevated above level of eyes, foveate medially, lateral margins carinate; eyes large, semiglobular, occupying less than 2/3 of entire dorsal area of head; pronotum 672 Great Basin Naturalist Vol. 43, No. 4 and scutellum very large; forewing elongate, rounded distally, venation typical, appendix well developed; clypeus long and broad, without median longitudinal carina; clypellus long and narrow, expanded distally. $ . Pygofer in lateral view moderately broad with very long caudoventral process, process narrow at basal 2/3, enlarged at dis- tal 1/3 with small ventral spine, caudodorsal margin with long narrow process, process abruptly pointed distally; aedeagus partially asymmetrical, long, narrow, tubular through- out, recurved distally, distal portion very short and narrow; gonopore subapical; con- nective broadly Y-shaped with short stem and long arms; style very long, about as long as aedeagus, narrow throughout; plate long and narrow, setose along outer margin at distal half. ? . Unknown Holotype ( <5 ), PANAMA: San Bias, near Punta Escoces, 77°42'W: 8°48'N., 2-II-79. Carohne Ash (USNM). Remarks: Coelidia panamensis is similar in male genital characteristics to attenuata Nielson but can be easily separated by the caudoventral process of the pygofer, which is enlarged distally and bears a ventral spine. Coelidia tortula, n. sp. (Figs. 13-18) Length: $ , 10.00 mm. General color fusco-piceous, except for flavous apex on forewing, veins with ochraceous spots. Head much narrower than pronotum, ante- rior margin obtusely angled; crown produced beyond anterior margin of eyes, broad, broader than width of eyes, elevated above level of eyes, foveate medially, carinate later- ally, eyes semiglobular, large, occupying less than 2/3 of entire dorsal area of head; pro- notum and scutellum large; forewing elon- gate, apex rounded, venation typical, appen- dix well developed; clypeus long and broad, without median longitudinal carina; clypellus long and narrow, apex expanded. S . Pygofer in lateral view moderately broad, with long caudoventral process and shorter caudodorsal process, both processes except for length about equal in width and Figs. 13-18. Coelidia tortula: 13, Male pygofer, lateral view. 14, Connective and right style, dorsal view. 15, Style, lateral view. 16, Aedeagus, dorsal view. 17, Ae- deagus, lateral view. 18, Plate, ventral view. similar in configuration, aedeagus asymmet- rical, long, broad, twisted subapically in dor- sal view, with 2 narrow, curved, distal pro- cesses; gonopore near apex of shaft basad of distal processes; connective broadly Y- shaped, stem short, arms long; style long, nar- row, about as long as aedeagus; plate elon- gate, profusely setose. ? . Unknown. Holotype ( $ ), BRAZIL: Amazon, Fon- teboa, no date, no collector (NR). Remarks: This species has no apparent close relatives but is nearest to atra Walker. It can be distinguished from all known spe- cies of Coelidia by the aedeagus with a twisted shaft and the distal processes, which are nearly of equal length and configuration and arise from the apex of the shaft. Coelidia cochloea, n. sp. (Figs. 19-24) Length: $ , 8.60 mm., ? , 9.00-9.70 mm. General color ochraceous, forewings with piceous pigmentation along basal 2/3 of cos- ta and with broad, smoky, fuscous, oblique October 1983 Nielson: New Leafhopper Species 673 Figs. 19-24. Coelidia cochloea: 19, Male pygofer, lat- eral view. 20, Connective and right style, dorsal view. 21, Style, lateral view. 22, Aedeagiis, dorsal view. 23, Aedeagiis, lateral view. 24, Plate, ventral view. band subapically, distal 1/4 and middle of fore wing translucent. Head small, much narrower than pro- notum, anterior margin obtusely angled, crown produced slightly beyond anterior margin of eyes, broad, slightly broader than width of eyes, elevated above level of eyes, slightly carinate laterally; eyes large, semi- globular, occupying less than 2/3 of entire dorsal area of head, pronotum and scutellum large; forewing elongate, rounded distally, venation typical, appendix well developed; clypeus long and broad, without median lon- gitudinal carina; clypellus long and narrow, margins expanded distally. S . Pygofer in lateral view broad, with long, bladelike caudoventral process and large ornate caudodorsal process, which is enlarged basally, slightly twisted basally and abruptly curved at distal half; aedeagus nearly symmetrical, simple, broad at basal 2/3 and narrowed at distal 1/3 in dorsal view, sinuate in lateral view, apex slightly hooked; gonopore subapical; connective broadly Y-shaped; style long, about as long as aedeagus, narrow at distal 2/3; plate elon- Figs. 25-30. Coelidia simplex: 25, Male pygofer, later- al view. 26, Connective and right style, dorsal view. 27, Style, lateral view. 28, Aedeagus, dorsal view. 29, Ae- deagus, lateral view. 30, Plate, ventral view. gate, profusely setose at distal half along out- er marginal area. $ . Seventh sternum large, about 2X as long as preceding segment, caudal margin produced along middle. Holotype (S), BRAZIL: Para, Belen Mo- cambo, 05-III-1977, T. Pimentel (OSU). Al- lotype ? , same data as holotype except col- lector, A. Y. Harada (OSU). Paratypes. BRAZIL: Manaos, 1 $ , 19-XI., no year, no collector (author's collection), Prata, 1 $ , 19-VII., no year, no collector (OSU), Para, Sta. Isabel, 1 ? , 13-VII-1973, B. Masca- renhas (USNM). Remarks: Coelidia cochloea is most closely related to simplex Nielson, and can be sepa- rated by the pygofer with the ornate caudo- dorsal process, which is enlarged basally and narrowed distally. Coelidia simplex, n. sp. (Figs. 25-30) Length: ^ , 8.60 mm. General color piceous except for narrow translucent apex on forewing, ochraceous 674 Great Basin Naturalist Vol. 43, No. 4 spots on veins, 5 narrow longitudinal lines on pronotum, 3 similar ones on scutellum, and broad ochraceous band on lateral margins of pronotum. Head small, much narrower than pro- notum, anterior margin obtusely angled; crown produced beyond anterior margin of eyes, broad, about as broad as width of eye, carinate laterally; eyes large, semiglobular, occupying nearly 2/3 of entire dorsal area of head; pronotum and scutellum large; fore- wing (right one missing on holotype) elon- gate, obtusely rounded distally, appendix well developed; clypeus long and broad, without median longitudinal carina; clypellus long, narrow, lateral margins expanded distally. S . Pygofer in lateral view broad, with very long, slender, acuminate caudoventral process and long, fingerlike caudodorsal pro- cess; aedeagus nearly asymmetrical, simple, long, narrow, tubular, broadly sinuate in lat- eral view, apex slightly recurved; gonopore near middle of shaft; connective broadly Y- shaped; style very long, longer than ae- deagus, very narrow, tapered distally; plate elongate, with numerous setae along outer margin. $ . Unknown. Holotype ( S ), PANAMA: Barro Colorado, Canal Zone, forest, 3- VI- 1976. H. Wolda (USNM). Remarks: This species is similar to cochlea Nielson in characters of the aedeagus but can be distinguished by caudal processes on the pygofer. The caudoventral process is very long, acuminate, and reaches the apex of the slender, fingerlike caudodorsal process. Coelidia gladia, n. name Coelidia spangbergi Nielson 1982 is a ju- nior homonym of Coelidia spangbergi Linna- vuori, 1956, and Coelidia spangbergi Metcalf, 1964, and must be replaced. In my revision of the tribe Teruliini (Niel- son 1979), Docalidia metcalf i Nielson was proposed as a new name for Coelidia spang- bergi Metcalf, 1964, nee Coelidia spangbergi Linnavuori, 1956. Metcalf (1964) proposed Coelidia spangbergi as a new name for Jassus flavicosta Spangberg, 1878, nee Jassus flavi- costa Stal, 1862. Coelidia spangbergi Linna- vuori was made a junior synonym of Stalo- lidia dissoluta (Jacobi) by Nielson (1979). All the Spangberg names originally assigned to Coelidia are either synonyms or homonyms and thus are no longer valid. Acknowledgments I appreciate the loan of specimens for this study provided by the following institutions and individuals: Dr. Per Inge Persson, Natur- historiska Riksmuseum, Stockholm (NR), Dr. J. P. Kramer, U.S. National Museum of Natu- ral History, Washington, D.C., (USNM), Dr. C. H. Triplehorn, Ohio State University, Co- lumbus (OSU), and Dr. H. Wolda, Smithso- nian Research Institute, Washington, D.C., (USNM). I am also indebted to Mr. Joel Floyd for preparing the illustrations. Literature Cited Metcalf, Z. P. 1964. General catalogue of the Homop- tera. Fasc. VI. Cicadelloidea Pt.II. Coelidiidae. U.S. Department of Agriculture, Washington, D.C. 182 pp. Nielson, M. W. 1979. A revision of the subfamily Coeli- diinae (Homoptera; Cicadellidae). III. Tribe Teruliini. Pacific Insects Monogr. 35. 329 pp., 1282 figs. 1982. A revision of the subfamily Coelidiinae (Homoptera: Cicdellidae). IV. Tribe Coelidiini. Pacific Insects Monogr. 38. 318 pp., 1104 figs. EYE FLUKE (DIPLOSTOMUM SPATHACEUM) OF FISHES FROM THE UPPER SALMON RIVER NEAR OBSIDIAN, IDAHO Richard Heckmann' Abstract.— Following a preliminary survey (1981) of diplostomatosis in fish from the Salmon River near Obsidian, Idaho, an extensive survey was conducted during the summer of 1982. From the initial sampling site on the Salmon River, 98 percent of .384 sculpin, Cottus bairdi, 8 percent of 317 salmonids, and 13 percent of 16 Dace and suckers were infected with Diplostomiiin spathaceum. Upriver from the initial sampling site and from three drainages enter- ing the Salmon River 28 percent of 185 sculpin and 1 percent of 70 salmonids were infected with D. spathaceum. The number of worms per eye was greater for sculpin (1 to 100+) than for salmonids (1 to 18) from the same area. The metacercariae of D. spathaceum occupy the vitreous body-retina area of infected fish. There is a prominent pa- thology associated with the infection, including detachment of the retina. Sculpin represent an indicator species for the range of diplostomatosis. The high infection rate of sculpin is associated with their bottom-dwelling character- istic and with their feeding habits. Following a preliminary study of the eye fluke of fishes from the Upper* Salmon River during the summer of 1981, an extensive sur- vey was conducted on the incidence of Di- plostomum spathaceum of fishes from the same locality during 1982. Diplostomum spathaceum (Rudolf i 1819) (Diplostomidae), the fish eye fluke that causes the disease diplostomatosis (diplos- tomatiasis), has been reported in many areas of North America and other parts of the world. Extensive surveys have been con- ducted in Utah concerning incidence, life his- tory, and pathology (Heckmann 1978). Di- plostomatosis, which is due to the presence of the metacercarial stage of this parasite in fish, causes cataracts of the lens and damage to the vitreous body and the retina of the eye. Diplostomum spathaceum is a digenetic trematode that has numerous synonyms in the literature (McDonald 1969). The life cycle of D. spathaceum. includes the adult parasite that lives in the intestinal tract of a piscivorous bird. The eggs from the adult trematode are passed in fecal deposits from the definitive host. The eggs embryo- nate in water and release a free-swimming miracidium in two to three weeks. The mira- cidium has approximately 24 hours in which to locate and infect the first intermediate host, which is a species of snail. In the snail the mother and daughter sporocysts develop in liver tissue. The daughter sporocysts re- lease free-swimming cercariae in approx- imately 6 weeks after miracidial penetration of the snail. The cercariae have from 24 to 48 hours to penetrate the second intermediate host. Fish are the most common second inter- mediate hosts; however, infections in am- phibians, reptiles, and mammals have also been reported (Ferguson 1943). Once the cer- cariae have penetrated the second inter- mediate host, they lose their forked tails and migrate to the lens tissue, where the metacer- cariae develop in 50 to 60 days (Erasmus 1958). When infected lens tissue is eaten by a bird, the adult fluke develops in the gut with- in five days (Oliver 1940). To date, 15 species of snails, 70 species of fish, and 37 species of birds have been reported worldwide as hosts for D. spathaceum (Palmieri et al. 1977). Currently there are several studies under- way throughout this country to determine the correct binominal name for the fish eye fluke. Consensus is that the metacercariae in- habiting the orbit of fish eyes in the Upper Salmon River is D. spathaceum,. Hoffman (1970) states that the genus Diplostomum in- cludes metacercarial stages in the eyes of fish. He lists two species for the eyes; D. spa- thaceum found in the lens with a distinct hindbody and D. huronense found in the vit- reous chamber, a worm less than three times 'Department of Zoology, Brigham Young University, Prove, Utah 84602. 675 676 Great Basin Naturalist Vol. 43, No. 4 as long as broad with a distinct hindbody. In both cases gulls are listed as the primary de- finitive host, with Lymnaea snails as the pre- ferred first intermediate host. According to the above characteristics, the eye fluke we have studied for this report should be D. huronense, but Dubois (1935) and Dubois and Mahon (1959) consider D. huronense to be a synonym for D. spathaceum. Thus, we will consider the fish eye fluke of the Upper Salmon River to be D. spathaceum. Beginning in June and ending in Septem- ber, sampling was conducted in a series of four periods in 1982. One of the major objec- tives of this study was to determine the num- ber of metacercariae in fish from (a) the bank and (b) midwater of the Upper Salmon River and streams draining into the river. Materials and Methods Four collection trips were scheduled and completed to the Upper Salmon River area. During each trip, fish were collected by one of three methods: electrofishing, hook and line, and nets. Where possible, samples were obtained from fishermen. Each fish was ex- amined for eye flukes by removing the soft tissue from the orbit of the eye, placing the contents in a petri dish, and examining the sample with a dissecting microscope. Sam- ples of eyes that contained numerous meta- cercariae (80 to 100+ per eye) were fixed in 10 percent formalin for sectioning and stain- ing to determine the pathology of the in- fection. Fish were sampled from the Upper Salmon River near Obsidian, Idaho, and from four other locations upriver that are identi- fied in Tables 1-3. Results The results of fish samples taken from the Upper Salmon River area are found in Tables 1, 2, and 3. From these data it is apparent that the mottled sculpin (Cottus bairdi) is the most susceptible to the eye fluke. The sculpin was used as a primary indicator species for other areas and feeder streams (Table 4). Whitefish (Prosopium williamsoni) (Table 2) carried the second highest number of meta- cercariae within the eye orbit. Chinook salmon (Oncorhynchus tshawytscha) are rela- tively free of the fish eye fluke. These tables also show that the infection in fish reached a peak toward the end of the summer. Fish sampled from upriver sites contained a lower number of metacercariae (Decker Flat) to no worms for fish from feeder streams and the headwaters of the Salmon River (Tables 2 and 4). Fish inhabiting slow- moving water and pools in the main river are more susceptible to cercarial invasion than those in fast water (Table 1). As expected, larger fish of the same species in general car- ry a greater number of worms than smaller fish (Table 1). Histological examination of the infected fish indicated a vitreous body-retina location for the worms (Fig. 1). The metacercariae cause a detachment of the retina from the outer vascular and fibrous coats (choroid, sclera). Thus, heavily infected fish (40-1- worms) are blind. The eye fluke found in fish in Utah inhabits the lens. The pathological effects of Diplostomum spatheceum upon the fish host are many. Ex- amination of those fish blinded with cataract and containing a heavy burden or larval metacercariae revealed stunted growth (length, girth, and weight), abnormal feeding behavior (lack of response to visual stimuli), and decreased vital acuity (Palmieri et al. 1977). Ashton et al. (1969) reported that lar- vae migrate to the eye via vascularvenous channels and showed that the lens, vitreous, or cortex of the eye may be proliferated with metacercariae. In older fish, chronic in- fections and pronounced subacute in- flammatory reactions in the vitreous in- volving heterophils, eosinophils, and macrophages with ingested lens material occurred. Visual acuity for infected fish can be slightly hampered or lost due to worm bur- den. In addition to visual loss and con- comitant pathogenesis, fish show retarded growth and a change in food habits. Fish- ermen consider the fluke as one of the rea- sons for a decrease in number of fish caught on artificial lures. Discussion Due to the unique nature and location of this fluke within the eye of the fish and due to its associated pathogenicity, much time October 1983 Heckmann: Eye Fluke of Fishes 677 Table 1. Summary of all samples from the mottled sculpin, Cotttis hairdi, checked for the eye fluke, Diplostomum spathaceum during 1982, Salmon River, Idaho. Date of Number sample Number of Size with Eye flukes Location in 1982 fish class ° eye fluke per eye Salmon River 28 June 9 M 9 6 (Side channel) 27 July 15 M and L 14 23 29 July 10 M 10 16 25 Aug 3 L 3 48 6 M 6 18 12 S 12 5 lOct 8 L 8 7 4 M 4 5 2 S 2 2 Salmon River 28 June 0 High water No sample (Middle channel) 27 July 32 M and L 31 39 25 Aug 12 L 12 88 + 9 M 9 43 10 S 10 11 lOct 1 XL 1 100 + » 14 L 14 81 + 11 M 11 35 + 6 S 6 27 + Salmon River 28 June 0 High water No sample (Main channel) 29 July 2 M 2 10 25 Aug 4 L 4 56 14 M 14 7 4 S 4 5 lOct 5 L 5 7 4 M 3 6 2 S 1 6 Frenchman Creek 28 July 12 M 0 0 Headwaters 28 July 10 M 0 0 (Salmon River) Salmon River 25 Aug 9 L 8 3 (Decker Flat) 15 M 13 3 7 S 4 1 Beaver Creek 26 Aug 12 M 0 0 7 S 0 0 Frenchman Creek 26 Aug 2 L 0 0 10 M 0 0 3 S 0 0 Headwaters 26 Aug 6 L 0 0 (Salmon River) 12 M 0 0 12 S 0 0 Salmon River 2 Oct 3 L 3 2 (Decker Flat) 21 M 21 2 2 S 2 1 Beaver Creek 2 Oct 8 L 0 0 12 M 0 0 2 S 0 0 Frenchman Creek 2 Oct 4 L 0 0 10 M 0 0 4 S 0 0 Headwaters 2 Oct 2 M 0 0 (Salmon River) 'The sculpin were divided into four size classes based on total length (TL). XL: greater than 115 mm TL, L: 95 to 115 mm TL, M: 85 to 94 mm TL, S: 65 to 84 mm TL. * "When there is more than 100 metacercariae in the orbit of a fish eye, a plus ( + ) designation is used. 678 Great Basin Naturalist Vol. 43, No. 4 Fig. 1. Figure A represents normal tissue found in the vitreous-retina area of a fish eye. B,C,D,E show infected eyes of a sculpin in which the Diplostomum metacercariae occupy the vitreous-body (V)-retina (R) area of fish. Note the detachment (arrow) of the retina due to metacercarial invasion (B,C). and money have been spent in an attempt to control and ultimately eradicate it. The greatest damage caused by this fluke is blind- ness and death in a variety of game fish throughout the world and specifically in Utah and Idaho. The survey completed on the potential hosts from the ichthyofauna of the Upper Salmon River and drainages is quite exten- sive. During 1982, 384 sculpin were obtained from the main Salmon River near Obsidian, Idaho, of which 98 percent were infected with the eye fluke (1 to 100-1- worms per eye); 185 sculpins were sampled upriver from the first collection site and from drainages into the river, of which 28 percent were in- fected (1 to 3 worms per eye); 317 salmonids and 16 Dace and Suckers were sampled from October 1983 Heckmann: Eye Fluke of Fishes 679 Tabue 2. Summary of all samples from fish representing the family Salmonidae checked for the fish eye fluke, Diplostomimi spathaceum, 1982, Salmon River, Idaho. Location Date of sample in 1982 Species of fish Number of fish Size of flsh Number with eye flukes Eye flukes per eye Salmon River (Side channel) 28 June Chinook Oncorhynchus tshawytscha 35 Fingerling 1 1 Steelhead Salmo gairdneri 12 Catchable 10-14 inches (TL)° 1 1 Salmon River (Main channel) 29 June Whiteflsh Prosopium williainsoni 3 10-16 inches (TL) 2 18 Steelhead Salmo gairdneri 4 10-14 inches (TL) 0 0 Rainbow Trout Salmo gairdneri 1 12 inches (TL) 0 0 30 June Whiteflsh Prosopim williamsoni 2 10-14 inches (TL) 2 15 Steelhead Sahno gairdneri 3 10-16 inches (TL) 0 0 1 Aug Whiteflsh Prosopium williamsoni 1 12 inches (TL) 0 0 27 Aug Rainbow Trout Salmo gairdneri 5 10-15 inches (TL) 4 2 Salmon River (Main channel) 27 Aug Steelhead Salmo gairdneri 6 10-14 inches (TL) 1 1 Chinook Oncorhynchus tshawytscha 17 Fingerling 2 1 Salmon River (Main channel) 28 July Brook Trout Salvelinus fontinalis 1 15 inches (TL) 0 0 Rainbow Trout Salmo gairdneri 3 9-14 inches (TL) 1 1 Salmon River (Side channel) 29 July Chinook Oncorhynchus tshawytscha 12 Fingerling 0 0 Holding tank (Salmon River water) 29 July Chinook Oncorhynchus tshawytscha 12 Fingerling 0 0 Salmon River (Middle channel) 25 Aug Chinook Oncorhynchus tshawytscha 15 Fingerling 0 0 Salmon River (Side channel) 25 Aug Chinook Oncorhynchus tshawytscha 32 Fingerling 0 0 Salmon River (Main channel) 25 Aug Chinook Oncorhynchus tshawytscha 14 Fingerling 1 1 Salmon River (Middle channel) 25 Aug Rainbow Trout Salmo gairdneri 7 9-15 inches (TL) 3 3 Salmon River (Side channel) 25 Aug Rainbow Trout Sabno gairdneri 2 10-12 inches (TL) 2 1 Salmon River (Side channel) 25 Aug Chinook Oncorhynchus tshawytscha 5 Spawners 26-36 inches (TL) 0 0 Salmon River (Side channel) 26 Aug Chinook Oncorhynchus tshawytscha 6 Spawners 26-42 inches (TL) 0 0 •TL: total length of fish. 680 Great Basin Naturalist Vol. 43, No. 4 Table 2 continued. Location Date of sample in 1982 Species of fish Number of fish Size of fish Number with eye flukes Eye flukes per eye Holding Tank (Salmon River water) 26 Aug Chinook Oncorhynchus tshawijtscha 26 Fingerlings 1 1 Salmon River (Middle channel) lOct Chinook Oncorhynchus tshcnv ytscha 19 Fingerlings 1 1 Salmon River (Main channel) Chinook Oncorhynchus tshaw ytscha 6 Fingerlings 0 0 Salmon River (Side channel) Chinook Oncorhynchus tshawytscha 25 Fingerlings 0 0 Holding Tank (Salmon River water) lOct Chinook Oncorhynchus tshaw ytscha 20 Fingerlings 1 1 Salmon River (Middle channel) lOct Rainbow Trout Sahno gairdneri 3 7-12 inches (TL) 1 3 Beaver Creek 28 July Brook Trout Salvelinus fontinalis 3 6-10 inches (TL) 0 0 Frenchman Creek 28 July Brook Trout Salvelinus fontinalis 3 5-9 inches (TL) 0 0 Frenchman Creek 28 July Chinook Oncorhynchus tshaw ytscha 1 Fingerling 0 0 Salmon River (Decker Flat) 25 Aug Rainbow Trout Sabno gairdneri 2 6-7 inches (TL) 0 0 Brook Trout Salvelinus fontinalis 2 5-6 inches (TL) 0 0 Beaver Creek 26 Aug Brook Trout Salvelinus fontinalis 7 4-10 inches (TL) 0 0 Frenchman Creek 26 Aug Brook Trout Salvelinus fontinalis 2 5-8 inches (TL) 0 0 Headwaters 26 Aug Brook Trout Salvelinus fontinalis 1 5 inches (TL) 0 0 Salmon River (Decker Flat) 2 Oct Chinook Oncorhynchus tshaw 'ytscha 26 Fingerlings 1 1 Beaver Creek 2 Oct Rainbow Trout Salnio gairdneri 4 8-10 inches (TL) 0 0 Brook Trout Salvelinus fontinalis 7 3-8 inches 0 0 Frenchman Creek 2 Oct Brook Trout Salvelinus fontinalis 19 7-12 inches (TL) 0 0 •TL; total length of fish. the first site, of which 81 percent (1 to 18 worms per eye) and 13 percent (1 worm per eye) were infected, respectively; and up river only 1 percent of the 70 salmonids was in- fected with 1 worm per eye. I have checked most of the fish species in that part of the Salmon River for metacercariae. The sculpin, Cottus bairdi, appears to be an excellent in- dicator host for the eye fluke. The fluctuation in numbers of metacercariae per infected fish correlates with sporocyst stages in the snail, which will be the topic for another paper. Diplostomatosis has been reported in Russia, Germany, Finland, Ireland, Mexico, Italy, Af- rica, England, Scotland, and the United States (Hoffman 1970, Davies 1972). Diplostomatosis is considered to be specific for freshwater fish. Dogiel (1962 and 1934) showed that lampreys and salmon become in- fected with the eye fluke during spawning October 1983 Heckmann: Eye Fluke of Fishes 681 migrations to fresh water. It is possible that salmon fry become free of Diplostomum after they return to the sea (Dogiel 1962). Direct contact between the fish and its parasite is required for cercarial penetration. Thus, the fish must swim into the infected areas since cercariae have a limited swim- ming ability. Slyczynska-Jurewuz (1959) uti- lized cages to show that fish have a greater tendency to get diplostomatosis as they move closer to the shore. This is due to the pre- ferred habitat of snails. The maximum rate of infection occurs during the months of June and July, coinciding with the peak of cerca- rial discharge (Kamenskii 1964). The peak in- fection occurred during August and Septem- ber for the current study. Snails prefer warm, clean, slow-moving water with vegetation in whi(5h to live (Ma- con 1950). This was also observed for snails infected with sporocysts from the Upper Salmon River. Lymnaeidae are generally found in water with at least 15 parts per mil- lion of bound carbon dioxide and with a pH of 7 or above (Pennak 1953). These snails are known to eat both plant and animal material but prefer vegetation when available. They live approximately one and a half years and have been known to estivate up to 3 years (Pennak 1953). Lymnaeids usually are found in less than 4.5 feet of water and can live without free oxygen (Cheatum 1934). Young snails are more susceptible to miracidial pen- etration than older snails, which appear to have some type of resistance (Cort et al. 1957). Fish and other cold-blooded vertebrates seem to have a fairly low resistance to meta- zoan parasites; thus, extensive damage to host tissue is not uncommon (Snieszko 1969). There continues to be debate concerning the general pathologic effects of D. spathaceum infecting the fish lens. Visual perception of infected fish varies from total blindness (Fer- guson 1943a) to impaired vision (Ghittino 1974). Pathologic effects to the eye by the para- site are characterized by inflammation, vas- cular disturbances, exophthalmia, destruction of lens tissue, necrosis, ulceration of the cor- nea, and eventual loss of the lens. Secondary damage can occur through the development of Saprolegnia within the necrotic tissue (Pal- mieri et al. 1976). Diplostomum spathaceum, causes several diseases of the eye region in a variety of fish. First signs of an infection are a number of lo- calized swellings or red patches on the fins, body, or eye area where cercariae penetrate and cause rupture of the surface blood ves- sels. In certain reported cases, mass entry of cercariae through the skin or gills causes ob- struction of the blood vessels in the gills, re- sulting in asphyxia, shock, and damage to the nervous system. Once the ultimate site loca- tion is found, metacercariae penetrate the iris, retina, and lens capsule by means of an- terior spines and secretions of the anterior penetration glands and encyst in these tissues or within the vitreous body or crystalline lens of the infected fish, causing immediate hem- orrhaging of the local area. The worms may Table 3. Summary of all samples from Dace and Suckers checked for the fish eye fluke, Diplostomum spathaceum, 1982, Salmon River, Idaho. Location Date of sample in 1982 Species of fish Nimiber of fish Size of fish Number with eye flukes Eye pe flukes r eye Salmon River (Side channel) 28 June Dace (Rhinichthijes) 4 2-4 inches (TL) 1 1 Salmon River (Main channel) 1 July Sucker (Catostomus) 1 14 inches (TL) 0 0 Salmon River (Main channel) 27 July Dace (Rhinichthyes) 1 3 inches (TL) 0 0 Salmon River (Side channel) 28 July Dace (Rhinichthyes) 4 3-4 inches (TL) 1 1 Salmon River (Side channel) 25 Aug Dace (Rhinichthyes) 6 4-5 inches (TL) 0 0 °TL: total length of fish. 682 Great Basin Naturalist Vol. 43, No. 4 Table 4. Range of eye fluke infection; from initial sampling area along the Upper Salmon River to the headwaters of the Salmon River: Cottus bairdi (Sculpin) indicator species. Location of sample Miles from initial sample site Species Date in 1982 Total fish Number infected "Number of eye flukes per eye Salmon River (Initial site) 0 Sculpin 27 July 32 31 39 Frenchman Creek 22 Sculpin 28 July 12 0 0 Headwaters (Salmon River) 23 Sculpin 28 July 10 0 0 Salmon River (Initial site) 0 Sculpin 25 Aug 31 31 46 + Salmon River (Decker Flat) 9 Sculpin 25 Aug 31 25 2 Beaver Creek 18 Sculpin 26 Aug 19 0 0 Frenchman Creek 22 Sculpin 26 Aug 15 0 0 Headwaters (Salmon River) 23 Sculpin 26 Aug 30 0 0 Salmon River (Initial site) 0 Sculpin lOct 32 32 61 + Salmon River (Decker Flat) 9 Sculpin 2 Oct 26 26 2 Beaver Creek 18 Sculpin 2 Oct 22 0 0 Frenchman Creek 22 Sculpin 2 Oct 18 0 0 Headwaters (Salmon River) 23 Sculpin 2 Oct 2 0 0 'When there is more than 100 metacercariae in the orbit of a fish eye, a plus ( + ) designation is used. icaudum (Cort and Brooks 1928) in snail inter- Stay viable from 10 months to two years or mediate hosts of different sizes. J. Parasitol. longer, causing chronic blindness due to par- 43:221-234. asitic cataract, keratoglobus, herniation, and Da vies, R. B. 1972. The life cycle and ecology of Di- tumor formation. During this time fish can- plostomum spathaceum RndoUi (1891), in North r , 11 1 1 . Park, Colorado. Unpublished thesis, Colorado not feed normally, and they stop growing or ^^^^^ ^niv., Fort Collins, Colorado. die. DoGiEL, V. 1962. General parasitology. Oliver and Boyd. Edinburg and London. DoGiEL, v., AND G. Petruschewsky. 1934. Die Wirkung Acknowledgments des Aufenthaltsortes auf die Parasitenfauna des ,^ 1111 Leches Wahrend Seiner Verschidensen Lebens- Thanks are given to the Idaho Fish and perioden. Arch. Hydrobiol. 26(4):659-673. Game Commission for their financial support Dubois, G. 1935. Contributions a I'etude de quelques of the project, especially to Grant Christen- parasites de I'ordre des Strigeatoidea. Rev. Suisse sen, who made final arrangements for the t-> <- ~ i m imwi c^ j ^ i^ „. ' 11 1 Dubois, G., and J. Mahon. 1959. Etude de quelques study. Amie Miller and Gary Gadwa helped trematodes Nor-Americaines (avec note sur la po- with the collections of fish during each trip. sition systematique de Parorchis Nicoll 1907) sui- vie d'une revision des genres Galactosomum Looss 1899 et Ochetosoma Braun 1901. Bull. Soc. Literature Cited Neuchatel. Sci. Nat. 82:191-229. Erasmus, D. A. 1958. Studies on the morphology, biolo- Ashton, N., N. Brown, and D. Easty. 1969. Trematode gy and development of a strigeid cercariae (cer- cataract in the freshwater fish. J. Small Anim. caria X Bayllis). Parsitology 48:312-335. Pract. 10:471-478. Ferguson, M. S. 1943. Development of eye flukes of Cheatum, E. P. 1934. Limnological investigations on ^s^es in the lenses of frogs, turtles, birds and respiration, annual migratory cycle, and other re- mammals. J. Parasitol. 29:136-142. lated phenomena in freshwater pulmonate snails. Chitting, P 1974. Rilievi cliniei e pathologiei su un Am. Microscop. Soc. Trans. 53:348-407. ^f^° ^' Cattarata Verminosa in trotelle iridee Cort, W. W., K. L. Hussey, and D. J. Ameel. 1957. d allevamento. Pisc.coltura e ittiopatholog.a X(2): Variations in infestations of Diplostomum flex- 59-61. October 1983 Heckmann: Eye Fluke of Fishes 683 Heckmann, R. A., and J. R. Palmieri. 1978. The eye fluke disease (diplostomatosis) in fishes from Utah. Great Basin Nat. 38:473-477. Hoffman, G. L. 1970. Parasites of North American freshwater fishes. Univ. of California Press, Berkeley. 486 pp. Kame.nskii, I. V. 1964. Diplostomatoz foreli v rybhoze "Snhodaya," TR vses inst. Geomintol. 11: 194-198.' Macon, T. T. 1950. Ecology of freshwater miillusca in the Engli.sh Lake District. J. Anim. Ecol. 19(2): 124-146. McDonald, M. 1969. Catalogue of helminths of uater- foiil (Anatidae). Bureau of Sport Fisheries and Wildlife. Special Scientific Report-Wildlife 126. Oliver, L. 1940. Development of Diplostomuin flex- icatidum (Cort and Brooks) in the chicken by feeding precocious metacercariae obtained from the snail intermediate host. J. Parasitol. 26(1): 85-86. Palmierl J. R., R. A. Heck.mann, and R. S. Evans. 1977. Life history and habitat analysis of the eye fluke Diplostomujii spatliaceuni (Trematoda: Diplo- stomatidae) in Utah. J. Parasitol. 63:427- 429. Palmierl J. R., R. A. Heckmann, and R. S. Evans. 1976. Life cycle and incidence of Diplostomiim spatha- ceimi Rudolphi (1819) (Trematoda: Diplostom- atidae) in Utah. Great Basin Nat. 36:86- 96. Pen'nak, R. W. 1953. Freshwater invertebrates of the United States. Ronald Press Co., New York. 769 pp. Slyezynska-Jurewuz, E. 1959. Expansion of cercariae of Diplostomum spatliaceiim Rudolfi 1819, a com- mon parasite of fishes in the littoral zone of the Lake Polskie. Arch. Hydrobiology 6:105-116. EVALUATION OF VARIETIES IN STANLEY A PINNATA (CRUCIFERAE) Robert W. Lichvar' Abstract.— Stanleya pinnata var. gibberosa Rollins is a narrow endemic from southwestern Wyoming. This taxon is based upon the morphological characters of crooked petals and all leaves bipinnate. During field work in 1980, a population near the type locality of this variety was located that had plants with both bipinnate and entire leaves. Due to this unusual population, hirther field and herbarium studies were done to resolve the taxonomy of var. gibbe- rosa. These further studies showed that this taxon is based upon young individuals of var. bipinnuta. Rollins (1939), in his monograph of the genus Stanleya, treated four varieties under S. pinnata (Pursh) Britt. He felt that this polymorphous species treated with just four varietal entities was rather unsatisfactory be- cause such a large range of variation had to be included. During field work in 1980 to evaluate the threats to, and distribution of Stanleya pin- nata var. gibberosa Rollins for proposed pro- tection under the Endangered Species Act, an unusual population of S. pinnata was dis- covered near the type locality for var. gibbe- rosa. Var. gibberosa was characterized by Rollins (1939) as having a crooked petal and all leaves bipinnate. The unusual population from near Ft. Bridger, Wyoming, had plants with both bipinnate and entire leaves and straight and crooked petals. Another popu- lation located in 1982 near the Owl Creek Range in central Wyoming showed the same variable characteristics as those from Ft. Bridger. These two populations were studied further. Methods Both field and herbarium studies were un- dertaken to sample the range of variation for Stanleya pinnata var. gibberosa. The closest variety morphologically, var. bipinnata (Greene) Rollins, was used for comparison. Field observations included flowering dates, habitats, leaf and petal shapes, and variations within the populations. Herbarium studies were done at the Rocky Mountain Her- barium (RM), Laramie, Wyoming, the New York Botanical Garden (NY), Bronx, New York, and the Gray Herbarium (GH), Cam- bridge, Massachusetts. All the characters used to separate varieties of S. pinnata were stud- ied. The following set of specimens were most pertinent to the study: at RM: Rollins 2320, 2351, 2382, 2388, 3077; Dorn 2942; Lichvar 2859, 4196, 5174; Nelson 3562, 7375; Porter 3367, 7768; Freytag 16; Berth s.n.; Goodding 1925; and Osterhout 1094: at GH: Rollins 2320, 2351, 2382, 2388, 3077, 57265, 79155: and at NY: Rollins 57265. Results and Discussion The questionable status of Stanleya pin- nata var. gibberosa, a narrow endemic, was first apparent after a visit to a population at Ft. Bridger, Wyoming, on 11 June 1980. This population had individuals with two different types of leaves and a wide range of plant heights. Those plants that were smaller in stature had all leaves bipinnate, and those in- dividuals larger in stature had bipinnate, pin- nate, and entire leaves. A continuum of these characters existed in this population between the two extremes of all bipinnate or all simple leaves, however. These observations, combined with those of Dorn (1979), stating that the key character of a crooked petal had broken down due to it being found in other varieties of the species, warranted further field and herbarium studies. The main morphological characters that have been used to separate var. gibberosa and var. bipinnata are: 'Wyoming Natural Heritage Program, The Nature Conservancy, 160.3 Capitol Avenue, Room 325, Cheyenne, Wyoming 82001. 684 October 1983 Lichvar: Stanleya pinnata 685 Petals straight or nearly so; leaves bipinnate to entire var. bipinnata Petals strongly crooked between blade and claw; all leaves bipinnate .. var. gibberosa Rollins (1939) stated that these two vari- eties each had a separate unified range in certain parts of Wyoming and Colorado. Based on his interpretation of ranges of these taxa, all the specimens that are used in this study had been previously identified accord- ing to the ranges given by Rollins. Specimens of both varieties were com- pared (Table 1) and showed that three speci- mens of var. gibberosa had crooked petals and five specimens had straight petals. Four collections of var. bipinnata had crooked pet- als and four had either all straight or a mix- ture of crooked and straight petals. This vari- ation confirmed observations made by Dorn (1979). The same group of herbarium specimens was also compared for pubescence and leaf shape. Three specimens of var. gibberosa had scant pubescence, three had dense, and two had a combination of dense and scant hairs. In var. bipinnata, five specimens had scant hairs and three had scant to dense hairs. Complete overlap occurs in the pubescence. Var. gibberosa had five specimens with some upper leaves entire and three with all leaves bipinnate. Var. bipinnata had five specimens with entire upper leaves and three with bi- pinnate upper leaves. The leaf characters of these two varieties overlap. Also, three other characters were recorded from herbarium sheets, the flowering dates, fruit shapes, and habitat types. All three of these features showed a continuous overlap. The combination of petal and leaf shapes in these two varieties showed similar overlap. Var. gibberosa had four specimens with straight petals and entire leaves at the sum- mit of the plants, the combination for var. bi- pinnata. Var. bipinnata had two specimens with crooked petals and all leaves bipinnate, the combination for var. gibberosa. These two specimens. Porter 3367 and Nelson 7375, are from southeastern Wyoming and are far outside the supposed range of var. gibberosa. Field studies showed the same kind of overlap. Lichvar 5174 was collected in 1982 in the upper edge of the Wind River Basin along the south flank of the Owl Creek Mountains, Wyoming. This population of Stanleya pinnata included a series of plants that ranged from all bipinnate leaves to ones with a mixture of pinnate and entire leaves. The plants in this population also had flowers that had both straight and crooked petals. The individuals that had a combination of all bipinnate leaves and crooked petals were small in stature and were young in age. As the individuals in this population grew in stature and older in age, the leaf shapes Table 1. Analysis of herbarium specimens of two varieties of Stanleya pinnata. Character Var. gibberosa Var. bipinnata Petal shape 3 crooked ° 5 straight Leaf shape 3 all bipinnate 5 entire Pubescence 3 scant 3 dense 2 with dense and scant Flowering dates 24 May to 29 June Fniit shape Torulose Habitat type disturbed soil of canal banks to limy bluff Combination of 4— straight petals with some entire leaves petal and leaf 1— crooked petals and all leaves bipinnate shapes 3— crooked petals and some entire leaves 'The number represents the number of herbarium specimens with that character. 4 crooked 4 with a mixture of crooked or straight 3 all bipinnate 5 entire 5 scant 3 scant to dense 2 May to 3 July Torulose gumbo swales to limy gravelly ridge 2— straight petals and some leaves entire 2— crooked petals with all leaves bipinnate 2— straight petals with all leaves bipinnate 1— plants in fruit with some leaves entire 1— plants in fruit with all bipinnate leaves 686 Great Basin Naturalist Vol. 43, No. 4 ranged from pinnate to entire and had a mix- ture of crooked and straight petals. The smaller individuals in the population were of the var. gihherosa aspect and the larger indi- viduals were both the var. bipinnata and var. pinnata aspects. This population expressed the same type of variability as the one at Ft. Bridger but was 180 miles (288 km) outside the previously known distribution of var. gibberosa. Conclusions A reevaluation of the status of Stanleya pinnata var. gibberosa can now be made be- cause of the availability of more collections and further field observations. The small stat- ure of var. gibberosa with nearly all or all bi- pinnate leaves is not necessarily unique to this taxon. Several specimens from well out- side the supposed range of var. gibberosa are identical. Also, at least two populations exist with highly variable morphology encompass- ing both var. gibberosa and var. bipinnata. Both populations have a continuum of var. gibberosa type individuals that are small in stature and young in age with all leaves bi- pirmate and crooked petals to older individ- uals with pinnate leaves plus a mixture of crooked and straight petals. Other genera in Cruciferae have similar variation in leaf shapes. Within Lepidium densiflorum Schrad. there are several differ- ent shapes of leaves. The basal leaves range from entire to pinnatifid, but no varietal dis- tinctions have been based upon these various leaf shapes. If the Ft. Bridger and the Owl Creek Range populations represent two varieties at each location that are hybridizing and back- crossing with the parents, then one would ex- pect several different types of plants with various age groups for each. Instead, these two populations have a continuum of charac- ters that are associated with variously aged individuals. It appears that var. gibberosa is based on young individuals with a small stature. Therefore, it is concluded that var. gibberosa is synonymous with var. bipinnata. Further study may show both are synonymous with var. pinnata. Acknowledgments Reed Rollins is thanked for openly dis- cussing his views of the taxonomy of Stanleya pinnata and Robert Dorn for reviewing the manuscript and making valuable comments. Literature Cited Dorn, R. D. 1979. Vascular Plants. In T. Clark and R. Dorn, eds.. Rare and endangered vascular plants and vertebrates of Wyoming. Privately published. Rollins, R. C. 1939. The Cruciferous genus Stanleya. Lloydia 2:109-127. SOME ASPECTS OF THE PRESETTLEMENT VEGETATION OF THE PICEANCE BASIN, COLORADO William L. Baker' Abstract.— Eight plant associations, not previously described in Colorado, and representative of part of the pre- settleinent vegetation spectrum in the oil shale region of northwestern Colorado, are described and illustrated, based on an inventory of relatively imdisturbed vegetation remnants. The Piceance Basin is underlain by oil shale estimated to contain 1.2 trillion barrels of oil (Murray and Haun 1974). Interest in the potential development of this resource has resulted in extensive study of the current vegetation of the basin. Most of this research consists of reports (Ferchau* 1974, Keam- merer 1977, Keammerer and Stoecker 1975) and theses (Tiedeman 1978, Vories 1974), though two published studies are available (Tiedeman and Terwilliger 1978, Ward et al. 1974). A few regional studies (James and Marr 1966, Marr and Buckner 1974, Marr et al. 1973) contain some quantitative data or general description of the Piceance Basin. Some additional reports pertain to the Roan and Parachute Creek areas south of the Piceance Basin (Ferchau 1973, Keammerer 1974, Keammerer and Keammerer 1980, Keammerer and Peterson 1981, Thome Eco- logical Institute 1973). Graham (1937) pro- vides a general overview of major vegetation types in the Uinta Basin, including this gener- al area of Colorado. None of these studies characterizes poten- tial vegetation, presettlement vegetation, or habitat types (Daubenmire 1952), concentrat- ing instead on existing vegetation, much of which has been altered by over 100 years of domestic hvestock grazing and agriculture. Knowledge of both potential and existing vegetation is essential if land managers are to be able to effectively rehabilitate disturbed lands. The goal of this paper is to discuss and present data on eight plant associations rep- resentative of presettlement vegetation in the Piceance Basin, based on a study of relatively undisturbed remnants. A qualitative overview of the vegetation in this area, and the im- pacts of grazing on this vegetation are dis- cussed in Baker (1982). This information was gathered as part of a general botanical inven- tory of the Piceance Basin (Peterson and Baker 1982). Study Area The Piceance Basin is located in Rio Blanco and Garfield counties in northwestern Colorado. It is an approximately 2850 km^ (1100 mi^) saucer-shaped basin bounded on the south by the Roan Plateau, on the west by the Douglas Creek drainage, on the north by the White River, and on the east by the Grand Hogback. The Piceance Basin is often considered to be a part of the eastern Uinta Basin. The entire study area is underlain by the Eocene Green River Formation, a kerogen- bearing marlstone (Donnell 1961), which out- crops as large cliff exposures around the mar- gin of the basin, and also occurs scattered throughout the central part of the basin as narrow bands, or tongues, in a Uinta Forma- tion matrix. The Eocene Uinta Formation, predominately a brown sandstone, is the pri- mary exposed surface rock over much of the central part of the basin. Climatic data are available from Craig, Colorado (Gale Research Co. 1980), 75 km northeast of the study area, at about the same elevation as the central part of the basin. Mean annual precipitation there is 338 mm. 'Colorado Natural Heritage Inventory, 1550 Lincoln Street, Suite 110, Denver, Colorado 80203. 687 688 Great Basin Naturalist Vol. 43, No. 4 distributed fairly evenly throughout the year, with a slight peak from afternoon convective thundershowers in August. Mean January temperature is -7.7 C, with mean July tem- perature 19.4 C. The study area is vegetationally similar to otlier parts of the Uinta Basin. Atriplex con- fertifolia stands occur at the lowest eleva- tions on slopes, with Arternisia tridentata stands, sometimes mixed with Sarcobatus ver- miculatus, occupying draws and creek bot- toms. Junipenis osteospenna-Piniis edtiHs woodlands alternate with Artemisia triden- tata openings on uplands below 2300 m, with Agrapijron spicatwn var. inerme grasslands formerly occupying flat ridges and uplands, and southerly-facing slopes. Above 2300 m, these grasslands are interrupted on slopes by a mixed shrub vegetation dominated by Qiiercus gambelii, Amelanchier iitahensis, Pnintis virginiana, Rosa ivoodsii, Cercocarpus montanus, and Symphoricarpos oreophilus, and occasional patches of Pseudotsuga men- ziesii or Populns tremidoides forests on the most protected northerly-facing slopes. The study area has been extensively grazed by cattle and sheep since the late 1800s. Many of the valley bottoms have been converted to agriculture. Mining of oil shale is currently limited to two 5000-acre Federal prototype lease tracts. Methods From May to August 1982, a reconnais- sance survey was conducted to locate rela- tively ungrazed and unlogged vegetation remnants in the study area. Every section of approximately 1 100 sections in the study area was searched. Methods of locating such rem- nants are similar to methods used by Dauben- mire (1970). Remnant areas were recognized by the following general features: (1) absence of obvious physical signs of grazing, such as cattle and sheep trails and terraced slopes, bedding areas, excessive amounts of trampled and broken shrub stems, compacted soils, scat, and logging or woodcutting signs, such as stumps, access roads, cutting debris, etc.; (2) absence or low coverage of exotic plant species (e.g., Bromus tectorum, Poa pratensis, Clwrispora tenella); (3) low coverage of plant species known to increase when domestic grazing occurs (e.g., Chrysothamnus spp., Gutierrezia sarothrae, Artemisia frigida); (4) presence of at least remnants of a soil crypto- gam layer on relatively flat sandy to silty soils. On rocky sites, slopes, or talus, presence of large crustose lichens on exposed rock sur- faces is suggestive of lack of recent heavy use, because rocks turned or dislodged by cattle and sheep hoof action cannot maintain large lichen growths; (5) presence of healthy, large native plants, generally abundant grass cover, with individual grass plants having many flower stalks, standing litter, and live centers; (6) general absence of plant pedestal- ing, excessive rilling and gullying, and other signs of excessive or accelerated erosion. Some additional features are specific to par- ticular associations. Generally, a combination of these factors made identification of rem- nants relatively simple. Remnants generally occurred on steep or inaccessible slopes far from water, or in areas excluded from graz- ing by accidents of fencing. Observations on effects of grazing on each association are based on a qualitative comparison of several sites and fenceline contrasts. Remnants located were sampled quan- titatively using a .1 hectare (20 X 50 m) plot method widely used in gradient analysis (e.g., Peet 1981). Shrub and herb percent canopy cover were sampled along the center line of the plot using 25 consecutive .5 X 2 m quad- rats. Plots were located in areas of visually homogenous vegetation. Tree size class struc- ture was sampled by tallying stem within the plot in 2-inch size classes, with diameter measured at breast height, or below the ma- jor point of branching (on Juniperus and Pinus). This study concentrates on eight plant as- sociations sampled and characterized based on 27 stands. Classification follows the meth- ods of Daubenmire (1970). The entire spec- trum of presettlement vegetation in the Pi- ceance Basin could not be sampled quantitatively, partly because of time con- straints, and partly because sufficient rem- nants could not be located that were free of disturbance effects, to characterize the origi- nal composition of all the associations. A pre- liminary qualitative classification of the origi- nal vegetation of the basin, based on October 1983 Baker: Colorado Vegetation 689 inference and reconnaissance data, and com- parison with literature from adjoining areas is in Baker (1982). Nomenclature follows Kartesz and Kartesz (1980). Voucher specimens are deposited at the Colorado State University Herbarium (CS). Soil types cited with each association were not sampled in each plot, but are based on recent soil maps (Tripp et al. 1982). Results and Discussion Table 1 summarizes shrub and herb per- cent cover. Table 2 summarizes tree size class data. Each of the associations is illus- trated in Figure 1 and discussed below. 1. Juniperus osteospenna-Pinus edtilis/ Agropyron spicatum var. inenne This association occupies gently sloping ridge tops and crests of low hills and mesas, often southerly-facing, between 1700 and 2150 m in elevation. It most often occurs on Uinta Formation sandstone, but may also oc- cur on Green River Formation marlstone. Sampled stands occur exclusively on the Rentsac soil series, a Lithic Ustic Torrior- thent, common in the basin (Tripp et al. 1982). The association has a savannalike appear- ance (Figure la), with widely spaced trees in a dense grass matrix and few shrubs present. Oryzopsis hyinenoides may codominate in some stands, but is most commonly a minor species or is absent entirely. A moderately developed soil cryptogam layer occurs in ex- cellent condition stands. The association is at the lower, drier eleva- tional end of the pinyon-juniper zone in this area. Juniperus appears to be slightly better adapted to these sites than Piniis, having more stems on most sites (Table 2). A few sites may lack Finns entirely. Both species generally have good reproduction and com- monly have a few large, old stems on most sites. The association degrades on relatively flat sites, under heavy domestic grazing, to a sim- ilar community with Haplopappus acaitlis dominant in the understory. This community has been described by Vories (1974, Associ- ation 14). On more sloping sites Artemisia tridentata, Gutierrezia sarothrae, Chry- sothamnus spp., and Bromus tectorum be- come dominants. This association has not been reported from other parts of Colorado, but Shute and West (no date) mention a similar association from the Uinta Basin near Price, Utah, where Agropyron spicatum var. inemie dominates the vmderstory of pinyon-juniper woodlands on "level mesa tops, deep wind-deposited (or sandy if shallow over sandstone) soils . . ." (p. 26). Though compositional data are not pro- vided, this is a similar environmental position and similar dominants. Data in Isaacson (1967, Table 5, plots 147-149, 192) also ap- pear to represent this association. These data were collected at unspecified localities in the Uinta Basin of Colorado and Utah. It appears likely that this association occurs in scattered localities across the Uinta Basin. This associ- ation is related to the Juniperus os- teosperma/ Agropyron spicatum association common in western Wyoming (Wight and Fisser 1968), which also occurs in Moffat County, Colorado. That association occurs north of the range limit of Pinus edulis. The awned variety (var. spicatum) of Agropyron spicatum is rare in the Piceance Basin, and never occurs mixed with A. spicatum var. inerme. 2. Juniperus osteosperma-Pinus edulis/ Amelanchier utahensis-Cercocarpus mon- tanus marlstone barren This association occurs on generally south- erly-facing slopes of white marlstone of the Green River Formation, from 1975 to 2450 m in elevation. These sites have soils mapped as a complex of Torriorthents and Rock Out- crops (Tripp et al. 1982). A well-developed tree and shrub layer are always present, but almost no herbaceous layer occurs. The association has conspicuous expanses of open, bare, exposed, partly de- composed white marlstone (Fig. lb), alternat- ing with clumps of Amelanchier and Cerco- carpus. Ephedra viridis is often present. Pinus edulis is often more abundant than Juniperus osteosperma on these sites, but old stems of both species commonly occur. Pinus edulis generally has more seedlings and sap- lings in this association than in any other as- sociation in the basin (Table 2). 690 Great Basin Naturalist Vol. 43, No. 4 Table ]. Percent cover and constancy ot shrubs and herbs. Plant association ninnbers correspond to those in the text. 1 = Jimipenis osteosperma-Pinus edulis/Agwpyron spkattim var. inerme, 2 = Jiinipertis osteospermci-Pinus edulis/ Amelanchier utahensis-Cercocarpus inontantis inarlstone barren, 3 = Pinits ediilis/AnielancIiier iita- hensis-AntofitaplnjIos patiila-Cercocarpu.s monianiis / Carex pitijophila, 4 = Pseudotsitgct menziesii/Amekinchier uta- hennis-Quercus gaiubelii-Sijmphoricarpos oreophiliis/ Carex geijeri-Poa fendleriana, 5 = Artemisia tridentata ssp. wyoniingensis-Symphoricarpos oreopliiltii/ Ehjinus cinereus, 6 = Atriplex confertifoUa/ Agropyron spicatnm var. in- erme-Oryzopsis hymenoides, 7 = Agropyron spicatnm var. inerme Great Basin grassland, 8 = Agropyron spicatnm var. inerme-Oryzopsis hymenoides Great Basin grassland. Table entries are percent canopy cover, followed by per- cent constancy, tr = trace quantities (less than .5 percent average cover); 100 percent is abbreviated to 99.9 percent. Plant association number 1 2 3 4 5 6 7 8 Number of stands 4 3 3 3 3 4 4 3 Shrubs Artemisia tridentata ssp. uyomingensis .9 99.9 tr 33.3 tr 99.9 19.1 99.9 .8 99.9 tr 50.0 tr 99.9 Atriplex confertifoUa tr 25.0 5.4 99.9 tr 25.0 Chrysothamnus viscidiflorus tr 75.0 tr 33.3 tr 66.6 tr 33.3 1.9 99.9 tr 75.0 1.0.50.0 1.7 66.6 CItrysotliamniis naiiseosus tr 75.0 3.0 66.6 tr 66.6 Symphoricarpos oreophiliis tr 99.9 tr 66.6 tr 66.6 12.4 99.9 28.6 99.9 tr 25.0 tr 25.0 Tetradymia canescens tr 50.0 tr 33.3 tr 75.0 Ceratoides lanata tr 25.0 tr 50.0 Amelanchier utahensis tr 25.0 11.5 99.9 11.7 99.9 3.7 66.6 tr 99.9 tr 50.0 Cercocarpiis montaniis tr 25.0 15.1 99.9 16.9 99.9 tr 33.3 tr 33.3 Querciis gambelii 13.6 99.9 Ephedra liridis 1.3 33.3 Purshia tridentata tr 50.0 tr 33.3 tr 33.3 tr 33.3 Mahonia repens tr 66.6 tr 66.6 tr 99.9 Gutierrezia sa roth rae tr 99.9 tr 33.3 tr 50.0 tr ,50.0 Clnysotliamntis parry i tr 25.0 Arctostaphylos patida 24.8 99.9 Pachistima myrsinites tr 33.3 .6 66,6 Artemisia frigida tr 50.0 .9 99.9 tr 75.0 .8 99.9 Rosa uoodsii tr ,33.3 .8 ,33.3 Ribes inerme tr 66.6 CeanotJius martinii tr ,33.3 Ribes cereum tr 25.0 Graminoids Carex pityoplnki .6 99.9 tr 66.6 4.2 99.9 .8 99.9 Carex geyeri 7.8 99.9 1.9 66.6 Carex sp. tr 66.6 Bromiis tectonim tr 75.0 tr ,33.3 tr 25.0 tr 33.3 Stipa comata 1.4 75.0 1.6,33.3 tr 25.0 .6 25.0 tr 66.6 Poa sandbcrgii tr 75.0 tr 25.0 Poa fendleriana tr 75.0 3,2 99.9 tr 33.3 Agropyron spicatnm var. inerme 16.0 99.9 tr 33.3 tr 66.6 tr .33.3 13.8 99.9 22.1 99.9 11.8 99.9 Oryzupsis h ymenoides 5.1 99.9 tr 33.3 tr 66.6 tr 99.9 tr 66.6 5.6 99.9 1.2 99.9 9.8 99.9 Kocleria cristata 1.8 99.9 tr 99.9 tr ,33.3 tr 25.0 2.4 75.0 Oryzopsis micrandia 1.0 66.6 Bromiis ciliatus tr ,33.3 1.5 99.9 Elymiis cinereus tr 33.3 20.5 99.9 tr ,33.3 Stipa Columbiana 1.8 99.9 Poa pratensis 1.9 66.6 Agropyron snntliii tr ,33.3 .5 99.9 tr 25.0 Agropyroi^ trachycaidum tr 66.6 5.6 99.9 Poa interior .8 99.9 Bromiis carinatits tr 66.6 Stipa lettermanii tr 33.3 tr 66.6 Poa nevadcnsis tr 33.3 Sitanion hystrix tr 66.6 tr 25.0 FoRBS Eriogoniim itmbelhititm tr 50.0 tr 33.3 tr 66.6 tr ,33.3 Machaeranthera grindelioides .8 99.9 tr 66.6 tr 99.9 tr 50.0 October 1983 Baker: Colorado Vegetation 691 Table 1 continued. Plant association number Number of stands Opuntki polyacantha tr 50.0 Clidniae.'itice fendleri tr 50.0 PItlox lioodii .9 99.9 Phijsaria acutifolia tr 75.0 tr 99.9 tr 66.6 tr 75.0 tr 75.0 tr 66.6 TciidxaniDi officinale tr 25.0 tr .33.3 1.5 99.9 Cliiliiim colomdense .6 99.9 tr 25.0 tr 25.0 1.1 ,33.3 Haphpappus acatilis .7 75.0 Cnjptantha scricea tr 99.9 tr 33.3 tr .33.3 tr 66.6 tr 75.0 .5 75.0 tr .33.3 Comimmdra umbelhita tr 50.0 tr .33.3 tr 25.0 Astr(ig(dus clicimaelence tr 50.0 tr 50.0 tr 50.0 tr 33.3 Hijmenopcippus filifolitis .5 25.0 tr ,33.3 tr 25.0 tr 25.0 Senccio multilobattis tr 99.9 tr 66.6 tr ,33.3 tr 25.0 Arahis spp. tr 25.0 tr ,33.3 tr .33.3 tr 25.0 tr 33.3 Eriogoninn lonclioplujlltiin tr 75.0 1.4 99.9 1.5 99.9 1.4 99.9 Cirsitiin spp. tr 75.0 tr ,50.0 1.4 75.0 .9 66.6 Stephanomeria ten iiifolia tr 50.0 tr 25.0 tr 50.0 tr 66.6 Leptodacttjhm pungens tr 33.3 tr 25.0 tr 25.0 tr ,33.3 Astragalus con vallariiis trJ5.0 tr 33.3 tr 33.3 tr 25.0 tr 25.0 tr 33.3 Senccio werneriifolius tr 50.0 Arabis lignifera tr 75.0 tr 33.3 Ipomopsis aggregata tr 50.0 tr 33.3 tr 25.0 tr .33.3 Penstcmon osterhoutii tr 50.0 tr 25.0 tr 25.0 Eriogoniini alatwn tr 25.0 tr 25.0 Phlox aiistroniontana .6 99.9 2.166.6 tr 66.6 Streptantliiis cordattis tr 50.0 tr 66.6 tr 66.6 Draba sp. tr .33.3 Cryp ta ntha fla voctda ta tr 99.9 tr 33.3 tr 25.0 Descurainea sp. tr 33.3 tr 25.0 Penstemon caespitostis tr 25.0 tr 66.6 tr 99.9 tr ,33.3 tr 25.0 Lithospenniim ruderale tr ,33.3 .5 99.9 Frasera speciosa tr ,33.3 tr 33.3 Achillea millefolium var. lantilosa tr 66.6 tr 66.6 1.5 99.9 tr 33.3 Astragalus miser .8 ,33.3 1.4 66.6 tr 33.3 Caidanth us crassicaulis tr 25.0 tr .33.3 tr 66.6 tr 25.0 Galitim boreale tr 66.6 .9 99.9 tr 25.0 Erigeron speciosus 1.8 99.9 tr 99.9 Clematis occidentalis var. dissecta tr 33.3 Penstemon strictus tr 33.3 tr 66.6 Balsamorliizd sagittata tr 66.6 tr 66.6 tr 66.6 tr 66.6 Castilleja linariifolia tr 66.6 tr 99.9 Crepis occidentalis tr 33.3 tr 66.6 Calochortus gii nnison ii tr 33.3 tr 99.9 Cirsium calcareum tr .33.3 tr .33.3 tr 66.6 tr25.0 Artemisia Itichviciana tr 25.0 tr 66.6 tr .33.3 tr 50.0 tr 50.0 1.8 99.9 Viguiera multiflora tr 33.3 Chenopoditim sp. tr 33.3 tr 66.6 Crepis acuminata tr 33.3 Oenothera sp. 1.5 99.9 Descurainea pinnata tr 33.3 Collomia linearis tr 66.6 Androsace septentrionalis tr ,33.3 tr ,33.3 Microsteris gracilis ssp. humilis tr 33.3 Penstemon icatsonii tr 66.6 .5 33.3 Ligitsticum porteri tr 66.6 Geranium fremontii .9 99.9 Lupinus caudatus 1.1 66.6 Composite sp. tr 33.3 tr 66.6 692 Great Basin Naturalist Vol. 43, No. 4 Table 1 continued. Plant association number 1 2 3 4 5 6 7 8 Number of stands 4 3 3 3 3 4 4 3 Potentilki gracilis var. piildierrima tr 33.3 Tragopogon duhius tr 99.9 Astrcigdht.s httosiis tr 25.0 tr 50.0 Penstemon sp. tr 25.0 tr 25.0 tr 25.0 tr 33.3 Arteviisid chacinuiilus tr 75.0 tr 25.0 tr 66.6 Mentzelid huntilis tr 50.0 tr 25.0 .8 99.9 Astnigdhis kentrophyta tr 75.0 tr 25.0 1.5 25.0 tr 33.3 Linum Icwisii tr 25.0 .7 75.0 tr 33.3 Euphorbia rohusta tr 50.0 tr 33.3 tr 25.0 tr .50.0 tr 66.6 Phacelia heteroph ijUa tr 33.3 tr 25.0 tr 33.3 Penstemon frcmontii tr 25.0 tr .33.3 tr .50,0 tr 25.0 Lescpierclla sp. tr 25.0 Ertjsimiiin aspeniiu tr 66.6 tr 66.6 tr 25.0 tr 50.0 Astragalus spatulatus tr 25.0 tr 25.0 Hechjsarum horeale tr 33.3 tr 25.0 1.1 .33.3 Phlox longifolia tr 25.0 Hifmenoxys acaulis tr 25,0 Chaenactis douglasii tr 50,0 Oenothera caespitosa tr 50.0 tr 66.6 The association has not been reported or named from other areas, though data in Keammerer and Peterson (1981:24) suggest an association very similar to this in composi- tion occurs on the Naval Oil Shale Reserve directly adjoining the southern boundary of this study area. It is likely the as.sociation is restricted to Green River Formation expo- sures in the Piceance Basin /Roan Plateau area of Colorado. 3. Pinus edulis/AmeJanchier iitahensis- Arctostaphylos patula-Cercocarpus inon- taniis/Carex pityophila This association occurs on flat to gently sloping ridge top exposures of Green River Formation marlstone, from 2100 to 2450 m in elevation. All stands are mapped as occur- ring on the Rentsac soil series, a Lithic Ustic Torriorthent common in the basin (Tripp et al. 1982). This association has a very dense shrub lay- er, and a depauperate herbaceous layer. It is the only association in the basin containing Arctostaphylos patula and Ceanothus mar- tinii. The association always has very sharp boimdaries. One can step across a line onto adjoining Uinta Formation sandstone, where Arctostaphylos does not occur, Junipenis os- teosperrna is co-dominant with Pmus edulis, and the herbaceous layer is much better de- veloped. Pinus edulis is the only tree present on most sites. Junipenis osteospenna may have a few stems or seedlings on some sites (Table 2). The association appears to be very fire sus- ceptible, possibly due to the high shrub den- sity. About half the range of the association in the basin has bvirned in the 100 years, and is in a postburn stage dominated by the three shrubs dominant in the understory of the ma- ture stage. This association has not been reported from other areas. It may be restricted to midelevation exposures of Green River For- mation in the Piceance Basin. 4. Pseudotsuga menziesii/Amelanchier utahensis-Quercus gambelii-Symphoricarpos oreophilus / Carex geyeri-Poa fendleriana This as.sociation typically occurs on the brows of northerly-facing slopes of draws, 1900-2600 m in elevation. It occurs on either Uinta Formation sand.stone or Green River Formation marlstone. Slopes generally do not exceed about 30 degrees. Soils are highly variable from stand to stand, ranging from Cryoborolls and Haploborolls to Torri- orthents (Tripp et al. 1982). This association is more open and less pro- tected than the Pseudotsuga men- ziesii/ Symphoricarpos oreophilus / Carex geyeri-Poa fendleriana association also found in the basin (Baker 1982) on steeper slopes October 1983 Baker: Colorado Vegetation 693 and in more mesic locations. The association has a dense shrub layer, with patches of Carex often densest under shrubs, and with Poa in the openings. Quercus seldom reaches tall shrub stature in this association, most commonly occurring as a low, often trailing shrub. Amelanchier may be absent from some low elevation stands on the driest sites. Quercus may also be absent from an occa- sional stand. Pseudotsuga jnenziesii may be the only tree in some stands. Junipenis scopulonmi is more commonly present, and may co- dominate on the most open, driest sites. Both trees appear to reproduce well in this associ- ation. /. scopuloriim is excluded from the name because of only moderate constancy. The association characteristically is criss- crossed with game trails, which result in much bare ground. Grazing of domestic ani- mals generally reduces Carex and Poa and re- sults in an increase in the exotic Poa pratensis and bare ground. The association has not been described or named previously, but data in Vories (1974, Association 1), Keammerer (1974:23, 1977:43), Marr et al. (1973, plot 43), and Fer- chau (1973:23) probably represent the associ- ation. The association has not been located outside the Piceance Basin/Roan Plateau area of Colorado. 5. Artemisia tridentata ssp. wyo- mingensis-Symphoricarpos oreophilus/Ely- nms cinereus This association occurs near the heads of draws on alluvium, from 2200 to 2600 m in elevation. Soils are mapped in several series, all CryoboroUs (Tripp et al. 1982). The asso- ciation grades downstream into an Artemisia tridentata ssp. tridentata / Elymtis cinereus as- sociation. The association typically occupies only the upper 1-2 km and headwaters area of intermittent stream drainages. The association is characterized by Arte- misia scattered through a matrix of tall Elymus clumps. Symphoricarpos often grows under or interwoven with the Artemisia stems and is not readily visible (Figure le). Grazing by domestic animals decreases Elymus cinereus, which results in an increase in exotic species (e.g., Poa pratensis, Bromiis tectorum), and the density of shrubs. This association apparently has not been described previously. It has been observed by this author in scattered localities in western Colorado, including the Danforth-Gray Hills area north of Meeker, and in northern Eagle county, always in essentially the same envi- ronmental position near the heads of draws at midelevations. 6. Atriplex confertifolia/ Agropyron spica- tum var. inerme-Oryzopsis hymenoides This association occurs on moderately steep to steep talus slopes of Uinta Formation sandstone, or occasionally on tongues of Green River Formation, from 1850 to 2075 m in elevation. It often occurs on southerly- facing slopes, but may also occur on other as- pects. These sites have soils mapped as a complex of Torriorthents and Rock Outcrops (Tripp et al. 1982). The association has a grassland appearance (Figure If), but consistently contains 5-6 per- cent cover of Atriplex confertifolia. The asso- ciation is characteristically sparse, with only 20-25 percent total cover, and much exposed bare soil. Grazing by domestic animals generally de- creases perennial grasses, and results in in- creases in Artemisia tridentata, Gutierrezia sarothrae, and bare ground. Only very rarely does Atriplex confertifolia become dense un- der heavy grazing pressure. This association apparently has not been described previously. Tiedeman and Terwilli- ger (1978, p. 212) describe a soil-vegetation unit in the Piceance Basin similar to this asso- ciation, but with the Agropyron identified as A. trachycaulum. This author has seen only A. spicatum var. inerme on dry slopes mixed with Atriplex, A. trachycaulum being found on more sheltered mesic north-facing slopes and in draws with a mixture of Amelanchier, Arteiyiisia, Symphoricarpos, and other shrubs. A related association, Atriplex confer- tifolia/Oryzopsis hymenoides also occurs in the Piceance Basin and in the Roan Plateau area (Baker 1982), where it is very common on more directly south-facing slopes. This lat- ter association also occurs across the northern Great Basin to California. 7. Agropyron spicatum var. inerme Great Basin grassland. 694 Great Basin Naturalist Vol. 43, No. 4 This association occurs in two settings in hymenoides, which co-dominates in associ- the Piceance Basin: (1) from 1950 to 2450 m ation 8, occurs in this type with generally less in elevation on generally south-facing slopes, than 10 percent of the cover of Agropyron often on steep talus, on either Uinta Forma- spicatum var. inerme, and is most often ab- tion sandstone or Green River Formation sent entirely. Where it does have significant marlstone, on soils mapped as a complex of cover in this association, it has low frequen- Torriorthents and Rock Outcrops (Tripp et cy, occurring as occasional small, dense al. 1982); (2) from 2450 to 2700 m in eleva- patches. tion on broad ridge tops and plateaus that are After observing numerous stands lacking often gently south or southwest facing, occur- the undisturbed characteristics cited in the ring on either Uinta Formation sandstone or methods section, five successional stages were Green River Formation marlstone, on soils recognized, based on increasing amounts of mapped as the Starman-Vandamore complex, grazing impact to this association: (1) Ag- which is a complex of Lithic and Typic Cry- ropyron spicatum var. inerme dominated cli- orthents (Tripp et al. 1982). max, (2) Agropyron spicatum var. in- This association is a rather sparse grassland erme-Koeleria cristata, (3) Koeleria cristata with 15-30 percent total cover. Oryzopsis dominated, with occasional small patches of Table 2. Tree diameter size distribution. Tree diameters are in inches measured at breast height (DBH). Seedhngs are less than 1 in DBH and less than 1 ni tall. Saplings are less than 1 in DBH and greater than 1 m tall. Entries are number of stems per size class. Species codes are JUOS = Junipenis osteospenna, JUSC = Jiiniperus scopttlonim, PIED = Pinus edulis, PSME = Pseudotsiiga menziesii. Stand No. Species code Seedlings Saplings 1-3 3-5 5-7 7-9 Junipenis osteosperma-Piniis edulis /Agropyron spicatum var. inerme 1 JUOS 0 0 0 15 0 PIED 0 0 0 12 2 2 JUOS 2 0 12 2 1 PIED 4 0 14 2 0 3 JUOS 2 0 2 3 8 0 PIED 4 0 10 11 JUSC 0 0 2 0 3 0 4 JUOS 11 4 6 4 4 0 PIED 3 1 2 2 0 1 Junipenis osteospertna-Pinus edulis / Amelanchier utahensis-Cercocarpus montanus marlstone barren 5 JUOS 4 0 4 2 0 0 PIED 16 12 10 8 4 10 6 JUOS 0 4 2 5 0 0 PIED 4 8 9 14 6 5 7 JUOS 0 1 2 0 0 0 PIED 5 10 15 4 3 3 Pinus edulis /Amelanchier utahensis-Arctostaphijlos patula-Cercocarpus montanus /Carex pitijophila 8 PIED 3 1 3 3 1 2 9 JUOS 1 1 0 10 1 PIED 4 0 3 2 2 2 10 JUOS 2 0 0 0 0 0 PIED 6 6 12 13 3 1 Pseudotsuga menziesii/ Amelanchier utaliensis-Quercus gambelii-Symphoricarpos oreuphilus/Carex geijeri- Poa fendleriana 11 PSME 23 11 13 11 8 10 12 PSME 18 16 12 2 2 2 JUSC 18 16 6 6 2 2 13 PSME 10 2 4 0 2 8 JUSC 14 4 4 12 12 0 PIED 2 0 0 0 0 0 October 1983 Baker: Colorado Vegetation 695 Agropyron, (4) Gutierrezia sarothrae- Chrysothamntis spp.-Cryptantha sericea dominated, with patches of Koeleria, (5) bare ground. On steep slopes, stages 2 and 3 may not occur, and Artemisia frigida, Machaeran- thera grindelioides, and other weedy species may mix with stage 4 species. Photographs il- lustrating each of these stages are in Baker (1982). Hanson and Stoddart (1940) discuss reasons Agropyron spicatiim var. inerme is easily damaged by domestic grazing. Formerly this very likely was one of the most common vegetation types in the Pi- ceance Basin. Now, perhaps 100-200 acres remain in stage 1 or 2, with most of the for- mer range in stage 3. Hanson and Stoddart (1940) indicate that the association has been similarly depleted in northeastern Utah and southeastern Idaho by overgrazing domestic livestock, though they do not describe succes- sional stages. This association is known to have occurred in the northern Great Basin in the Cache Val- ley area of northeastern Utah and south- eastern Idaho (Hull and Hull 1974, Hanson 1939, Hanson and Stoddart 1940), approx- imately 400 km northwest of the Piceance Basin, where it has similarly been described as representative of presettlement conditions in that area. The association has not been de- scribed in Colorado. Data in Keammerer and Stoecker (1975:13), and in Tiedeman and Terwilliger (1978:200) appear to represent the Koeleria successional stage of this associ- Table 2 cont inued. Inches 9-11 11-13 13-15 15-17 17-19 19-21 21-23 23-25 25-27 27 + 2 3 3 0 1 1 1 1 2 1(30 in) 1 1 0 0 0 0 0 0 0 0 2 0 1 2 0 1 0 2 0 1(34 in) 2 1 1 1 1 1 1 0 0 1(36 in) 8 0 0 1 1 0 0 0 1 1(36 in) 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 3 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 2 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 4 0 0 0 2 0 0 0 0 1(40 in) 0 0 2 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 1 1 0 1 0 0 0 0 3 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 4 5 5 3 1 1 0 0 0 0 2 2 2 2 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 4 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 696 Great Basin Naturalist Vol. 43, No. 4 ation. These latter authors suggest that, if un- disturbed, this type "would reach a stable plant community dominated by needle and thread" (p. 201). We could not locate any remnant areas dominated by Stipa comata. Stipa comata is commonly a minor com- ponent of the higher elevation version of cli- max Agropyron spicatum var. inenne grass- lands. Where it occurs most commonly, there is always abundant evidence of domestic live- stock use, suggesting it may increase tempo- rarily at the expense of the more palatible A. spicatum var. inenne, and then also decline as impact increases. Data in Vories (1974, As- sociation 3) appear to represent stage 4 of the successional series. This association is undoubtedly related to the Agropyron spicatum associations typical of the Palouse region in the Columbia River Basin of Washington, Idaho, and northern Utah and Nevada, which are represented in the northern Rocky Mountains as montane grasslands, and also occur in central and northern Colorado. Typical A. spicatum var. spicatum, with long divaricate awns, is rare in the Piceance Basin and never forms grass- lands or mixes with A. spicatum var. inerme. This appears to be the case in northeastern Utah also, where grasslands of the two vari- eties do occur, but generally are geographi- cally separated (Hull and Hull 1974). Passey and Hugie's (1963) data suggest that A. spica- tum var. inerme grasslands may be limited to the northern Great Basin, not extending into the typical Palouse region, where A. spica- tum var. spicatum dominates. 8. Agropyron spicatum var. inerme- Oryzopsis hymenoides Great Basin grass- land This association occurs on southerly-facing steep talus slopes of Uinta Formation sand- stone, from 1975 to 2200 m in elevation. Soils are mapped as a complex of Torriorthents and Rock Outcrops (Tripp et al. 1982). This association is also a sparse grassland, with 15-25 percent total cover. Oryzopsis hymenoides has half or more of the cover of Agropyron spicatum var. inerme, and high frequency. Grazing by domestic animals decreases perermial grasses and results in an increase in Artemisia frigida, Artemisia tridentata, and Chrysothamnus spp., along with the exotic Bromus tectorum. This association does not follow the same successional sequence as oc- curs with association 7. This association apparently has not been described previously. Vories (1974, Associ- ation 25) describes a community that may represent a poor condition example of this as- sociation, but the species of Agropyron is not identified. Ward et al. (1974, Type II-E) mention this combination of co-dominants but give no additional details. Ferchau (1974, p. 2) mentions a "Wheatgrass-Ricegrass" type that may represent this association, but the species of Agropyron is not identified. This association is apparently restricted to the Pi- ceance Basin/Roan Plateau area of Colorado. Current data, from this study and from Colorado Natural Heritage Inventory files, suggest associations 2, 3, 4, 6, and 8 are re- stricted to the Piceance Basin/ Roan Plateau area of Colorado, though additional informa- tion is needed from similar exposures of Green River Formation in northeastern Utah. Associations 1, 5, and 7 are more wide rang- ing, occurring in scattered localities in north- ern Utah and northwestern Colorado. Baker (1982) lists an additional three associations, not quantitatively sampled, that are appar- ently restricted to the Piceance Basin/ Roan Plateau area of Colorado. With the exception of association 4, restricted associations are ei- ther found on dry southerly-facing slopes and ridge tops (associations 6 and 8), or on rocky exposures of Green River Formation marl- stone (association 2 and 3). These more ex- treme environments in the basin have unique plant associations, in addition to being pri- mary habitat for most of the basin's rare plant taxa (Peterson and Baker 1982). More mesic areas tend to contain wide-ranging as- sociations (e.g., association 5) and lack rare taxa. Although the concept of climax and the validity of the plant association continue to be subjects of ecological debate, many of the remaining fragments of the presettlement vegetation spectrum are being subjected to inadvertent loss during development and land use. Much of the forest vegetation of the western United States is being or has been studied and classified (cf. Pfister 1982) based October 1983 Baker: Colorado Vegetation 697 ^ ^ £ P -^ 2 2 C =1- is o * S =:i =" c > •3- "5 ~ ^ 2 ^ "S "a. ■^ s •■5. -S "^^ ^ ^ ^■"3, , "^ S !». i^ = ■- S- "> = 5 S ^ ^ sj ^ 5ft ■~ ~S o "— i* ^' 2 "^ a. 6c J3 "" 2~ c <■ -^ ^. X — s -c ^ _c "" 'v^ 1 = -2 "S C-T3 ■i; "S -S 2 C :^ -^ ^ 5D « ^ ■*** ^ ^ !/i « ^ ^ ■S s 2^2 ^ 5b Si s -o ^ /Agrop inus ed s-Quer 2 .S -2 a, -^ 2 ^ ■^ ^ — . ^ _~ qj ••^ ^ ^ - — ' »^ -c O ^ ^' -2 a.^ 2 ij ~ § 5i )^ *^ ■S ^ ^ o C Bs ^ := ma- one ^ 5 a.i: = ■^ > ? p ^ "3. § o 2 ? S-, "2 S T? ^' -i 2 ^ '— i =^ E ~ S ^ 5 ~S i^ Sue ^ S S^ _ g S a. ^3S~A o 2 -2 2 T2 ^ d -c "^ -Sa s tso ■7 ~ B •^ ^ ■•: 03 .^ u :^ 0 0 m :^ <; (? ^ iO =ft JO < jj 2 o:i c ^ ^ 1 0 S3 -2 c ^ ^ c OJ "^ c ~ ^ -£; -c: ax 3" S 698 Great Basin Naturalist Vol. 43, No. 4 on potential or presettlement plant associ- ations, but a similar effort is needed to cata- log and describe plant associations on non- forested and lower elevation sites prior to the loss of the remaining opportunities for study. Although in many areas, such as the Piceance Basin, it may be too late to comprehensively describe and classify the presettlement vege- tation spectrum, land managers charged with rehabilitating disturbed lands cannot begin to effectively achieve this goal without as much information as can now be provided on pre- disturbance conditions. Acknowledgments Part of this study was completed during a contract with the Bureau of Land Manage- ment, Craig District Office, Craig, Colorado. I am grateful to Karen Wiley-Eberle, Vernie Armstrong, and Curt Smith of BLM for as- sistance with logistics. Field crews assisted in locating remnant vegetation areas. I appreci- ate the efforts of Dr. Dieter Wilken of the Colorado State University Herbarium, who verified all plant specimens. Warren Keammerer, K. Vories, John Marr, Thome Ecological Institute, EXXON, TOSCO, ARCO, and TRW all supplied access to un- published data and reports. This study could not have been completed without Tamara Naumann, who volunteered assistance in field sampling. I am also grateful for the support and encouragement of J. Scott Peterson. Literature Cited Baker, W. L. 1982. Natural vegetation of the Piceance Basin, Colorado. In J. S. Peterson and W. L. Bak- er, eds.. Inventory of the Piceance Basin, Colo- rado: threatened and endangered plants, plant as- sociations, and the general flora. Report prepared for the Craig District Office, Bureau of Land Management, by Colorado Natural Heritage In- ventory, Denver. Appendix D. 113 pp. Daubenmire, R. 1952. Forest vegetation of northern Idaho and adjacent Washington, and its bearing on concepts of vegetation classification. Ecol. Monogr. 22:.301-330. 1970. Steppe vegetation of Washington. Wash- ington Agric. Expt. Sta. Tech. Bull. 62. 131 pp. DoNNELL, J. R. 1961. Oil-.shale resources of the Piceance Creek Basin between the Colorado and White rivers, northwestern Colorado. Pages 835-890 in U.S. Geol. Surv. Bull. 1082-L. Ferchau, H. 1973. Vegetative inventory, analysis, and impact study of the Parachute Creek area, Gar- field county, Colorado. Pages 1-77 in Colony en- vironmental study, Thome Ecological Institute, Boulder, Colorado. Part II, Vol. 1, Chap. VI. 1974. An ecological analysis of the vegetation, Pi- ceance Creek Basin, Rio Blanco and Garfield counties, Colorado. In Regional oil shale studv. Report to the State of Colorado by Thorne Eco- logical Institute, Boulder. Vol. 14. 19 pp. Gale Research Company. 1980. Climates of the states: Colorado. Gale Res. Co., Book Tower, Detroit, Michigan. 2d ed. p. 104. Graham, E. H. 1937. Botanical studies in the Uinta Ba- sin of Utali and Colorado. .\nn. Carnegie Mu- seimi 26:1-432. Hanson, W. R. 19.39. The ecology of Agwpyron inenne on protected and heavily grazed range land in Cache Valley, Utah. Unpublished thesis. Utali State Univ., Logan. 33 pp. Hanson, W. R., and L. A. Stoddart. 1940. Effects of grazing upon bunch wheat grass. J. .\mer. Soc. Agron. 32:278-289. Hull, A. C, Jr., and M. K. Hull. 1974. Presettlement vegetation of Cache Valley, Utali and Idaho. J. Range Management 27:27-29. Isaacson, H. E. 1967. Ecological provinces within the pinyon-jimiper type of the Great Basin and Colo- rado Plateau. Unpublished thesis. Utah State Univ., Logan. James, J. W., and J. W. Marr. 1966. Route 40 mountain environmental transect: Colorado and Utah. Re- port prepared for the Institute of Arctic and Al- pine Research, Univ. of Colorado, Boulder. 218 pp. + appendices. Kartesz, J. T., AND R. Kartesz. 1980. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Univ. of North Carolina Press, Chapel Hill. 498 pp. Keammerer, W. R. 1974. Vegetation of Parachute Creek Valley. Pages 4-91 in Environmental inventorv and impact analysis of a proposed utilities corri- dor in Parachute Creek Valley, Colorado. Unpub- lished report prepared for Colony Development Operation, Atlantic Richfield Co., Operator, Denver. 1977. Final report: vegetation baseline studies, oil .shale tract C-b. Unpublished report. Stoecker- Keammerer and Assoc, Ecological Consultants, Boulder, Colorado. 183 pp. Keammerer, W. R., and D. B. Keammerer. 1980. Pre- liminary vegetation studies on the Naval Oil Shale Reserve. Unpublished report prepared for TOSCO Corp., Denver, Colorado, by Stoecker- Keammerer and Assoc, Ecological Consultants, Boulder. 14 pp. Keammerer, W. R., and S. J. Peterson. 1981. Vegeta- tion studies on the Naval Oil Shale Reserve. Un- published report prepared for TRW Energy Sys- tems Group, McLean, Virginia, by Stoecker- Keammerer and Assoc, Ecological Consultants, Boulder, Colorado. 77 pp. Keammerer, W. R., and R. E. Stoecker. 1975. Vegeta- tion and wildlife studies along proposed corridors for oil .shale tract C-b. Unpublished report pre- pared for Shell Oil Co., Operator, Denver, Colo- rado, by Stoecker-Keanimerer and Assoc, Eco- logical Con.sult;ints, Boulder, Colorado. 86 pp. Marr, J. W., and D. L. Buckner. 1974. Colorado to Wvoming pipeline corridor study. Unpublished October 1983 Baker: Colorado Vegetation 699 report prepared for Colony Development Operation, At- lantic Richfield Co., Denver, (Colorado, hv Tliorne Ecological Institute, Boulder. 79 pp. Marb, J. W'., D. L. Bi'CKNER, AND C. MuTEL. 1973. Eco- logical analysis of potential oil products corridors in Colorado and Utah. Unpublished report pre- pared for Colony Development Operation, Atlan- tic Richfield Co., Denver, Colorado. 96 pp. -I- appendices. Mlrray, D. K., and ]. D. Haun. 1974. Introduction to the geology of the Piceance Creek Basin and vi- cinity, northwestern Colorado. Pages 29-39 in Guidebook to the Energy Resources of the Pi- ceance Creek Basin, Colorado. Rocky Mountain .Assoc, of Geologists, Denver, Colorado. Passey, H. B., and V. K. Huc;ie. 1963. Variation in blue- bunch wheatgrass in relation to environment and geographic location. Ecology 44:158-161. Peet, R. K. 1981. Forest vegetation of the Colorado Front Range: composition and dynamics. Vegeta- tio 45:3-75. Peterson, J. S., and VV. L. Baker. 1982. Inventory of the Piceance Basin, Colorado: threatened and en- dangered plants, plant associations, and the gen- eral flora. Unpublished report prepared for the Craig District Office, Bureau of Land Manage- ment, by Colorado Natural Heritage Inventory, Denver. 5 volimies. Pfister, R. D. 1982. Designing succession models to meet management needs. Pages 44-53 hi J. E. Means, ed.. Forest succession and stand devel- opment research in the northwest. Proceedings of the symposium held 26 March 1981 at Corvallis, Oregon. Forest Research Laboratory, Oregon State Univ., Corvallis. Shute, D., and N. E. West. No date. The application of ECOSYM vegetation cla.ssification to rangelands near Price, Utah. Unpublished reports 14 and 16. In J. A. Henderson and L. S. Davis, eds., Ecosys- tem classification and data storage .system for natural resources management. Department of Forestry and Outdoor Recreation, Utah State Univ., Logan. 53 pp. Thorne Ecological Institute. 1973. Environmental setting of the Parachute Creek Valley: an ecologi- cal inventory. Pages 36-40, map in Unpublished report prepared for Colony Development Oper- ation, Atlantic Richfield Co., Operator, Denver, Colorado, by Thorne Ecological Institute, Boidder. TiEDEMAN, J. A. 1978. Phyto-edaphic classification of the Piceance Basin. Unpublished dissertation. Colo- rado State Univ., Fort Collins. 281 pp. TiEDEMAN, J. A., AND C. Terwilliger, Jr. 1978. Phyto- edaphic classification of the Piceance Basin. Col- orado State Univ., Range Science Department Science Series 31. 265 pp. Tripp, W. P., L. W. Williams, D. K. Alstatt, J. J. Rawinski, and C. F. Spears. 1982. Soil survey of Rio Blanco county area, Colorado. U.S. Dept. of Agric, Soil Conservation Service, and U.S. Dept. of the Interior, Bureau of Land Management. 219 pp. + maps. VoRiES, K. C. 1974. A vegetation inventory and analysis of the Piceance Basin and adjacent drainages. Unpub!i.shed thesis. Western State College, Gun- nison, Colorado. 243 pp. Ward, R. T., W. Slauson, and R. L. Dix. 1974. The natural vegetation in the landscape of the Colo- rado Oil Shale region. Pages 30-66 in C. W. Cook, ed.. Surface rehabilitation of land distur- bances resulting from oil .shale development. Fi- nal Report, Phase I, to the Colorado Department of Natural Resources. Environmental Resources Center, Colorado State Univ., Fort Collins, Tech. Rep. Series 1. Wight, J. R., and H. G. Fisser. 1968. Jtiniperus os- teosperma in northwestern Wyoming: their distri- bution and ecology. Univ. of Wyoming, Agric. Expt. Sta. Sci. Monogr. 7. 27 pp. NEW VARIETY OF OPUNTIA BASILARIS (CACTACEAE) FROM UTAH Stanley L. Welsh' and Elizabeth Neese- Abstract.— Described as a new variety is Optintia basihiris Engelm. & Bigel. var. lieilii Welsh & Neese. Work leading to a treatment of the flora of Utah has drawn attention to the presence of a segment of the variation within Opuntia has- ilaris that is beyond the circumscription of previously described infraspecific taxa (Ben- son, 1982). The plants stand apart from the remainder of the complex, being situated on saline soils of the southern end of the San Ra- fael Swell and the north end of the Henry Moimtains. The remainder of the species is far to the south and southwest of this area. The variety is named in honor of Kenneth Heil, enthusiastic student of the Cactaceae. Opuntia hasilaris Engelm. & Bigel. var. heilii Welsh & Neese. Similis var. hasilaris sed in articulis coloris (non violaceis) glo- chidis stramineis et ambitis differt. Joints spatulate to obovate, rounded to truncate apically, yellowish (rarely bluish) green; areoles lacking spines, 8-22 mm apart; glochids straw colored; flowers 4.5-6 cm long, violet; ovaries and fruit areolate, with glochids and often with spinules; fruit dry, ca 2 cm long and 1.5 cm wide; seeds ca 7.3 mm long, pale tan. Type: USA Utah. Wayne Co., T29S, RIOE, S23 (NWI1/2), Blue Benches SW of Hanks- ville, N of Henry Mts., 1464 m, sandy clay, Mancos Shale Formation, 1 July 1978, E. Neese 5938 (Holotype BRY). Additional specimens: Emery Co., T25S, RIOE, SI (SWI1/2), 8.8 km WNW of Goblin Valley Campground, 1479 m, salt desert shrub community, Curtis Formation, soil powdery silty sand, 19 May 1982, E. Neese & K. Mutz 11715 (BRY); do, T26S, R9E, S4, San Rafael Swell, Keesle Country, near Delta Mine, 1586 m, ephedra-atriplex community, sandy soil and rocky outcrops, 3 June 1980, J. G. Harris 833 (BRY). This variety is similar to var. hasilaris, differing in subtle modifications of joint out- line, in color, and in glochid color. It is isolat- ed from the type variety by 200 km and from var. aurea by 100 km. It is similar in pad col- or to var. aurea, but differs in pad outline and in flower color. Literature Cited Benson, L. 1982. The cacti of the United States and Canada. Stanford Univ. Press, California. 1044 pp. 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. -Life Science Museum, Brigham Young University, Provo, Utah 84602. 700 VEGETATIVE TYPES AND ENDEMIC PLANTS OF THE BRYCE CANYON BREAKS Robert A. Graybosch' and Hayle Biichaiian- Abstract.— The scenic Bi yce Canyon "breaks" constitute harsh and inhospitable habitats for plant life. The eroded pink cliffs and talus slopes are sites of some of the most rapid natiual erosion on earth. This paper divides the plant life on the breaks of tiie main Bryce Canyon amphitheater into four vegetative types. A checklist of all plant species found in the main amphitheater is included. Many of the rare and endangered endemic species of the park are found in the Pinus longaeva vegetative type. Recommendations for managing the breaks to protect endemics are presented. The breaks community is one of the six major plant communities of, Bryce Canyon National Park as described by Buchanan (1960). It consists of a relatively narrow band of imvegetated or sparsely vegetated bad- lands formed by the red beds of Claron (Wasatch) formation along the eastern edge of the Pausaugunt Plateau. This paper repre- sents the results of a study of the main am- phitheater of Bryce Canyon to define the vegetative units. Intricate erosional formations can be viewed from Sunrise, Sunset, Inspiration, and Bryce viewpoints. The diverse topography and beauty of the formations attract thou- sands of park visitors to the viewpoints and rim trails, although relatively few of them venture into the main amphitheater. The ma- jority of hikers and horse riders who traverse the breaks remain on well-maintained trails, and thus have relatively little impact on the plant communities. The flora of Bryce Canyon is rich in spe- cies endemic to the High Plateaus of south- em Utah (Welsh and Thorn 1979, Buchanan and Graybosch 1981). Several of these threat- ened and endangered plant species have lim- ited populations in the main Bryce Canyon amphitheater. Preservation of such species depends on recognition of preferred habitats and provision of means to protect them from visitor impact. Description of the Study Area Bryce Canyon National Park is located on the eastern edge of the Pausaugunt Plateau in south central Utah. The Pausaugunt Plateau occupies a position midway between 37° and 39° north latitude 10 miles west of the 110th meridian. The boundaries of the main am- phitheater circumscribe the drainage system of Bryce Wash, an intermittent tributary to the Paria River, an area 4.5 km-. The bound- aries differ somewhat from those considered by Lindquist (1977) and Buchanan and Gray- bosch (1981). Elevations within the study area range from 2,200 to 2,530 m (7,250-8,300 ft). The study area lies along the western border of the Kaiparowits Basin, the flora of which was reported by Welsh et al. (1978). The geologic stratigraphy of the study area is reported by Brox (1961), Anderson and Rowley (1975), Doelling (1975), and Lind- quist (1980). The Claron limestone, a Ter- tiary deposit, is divisible into Red Eocene beds and White Oligocene beds, which differ somewhat in presence or absence of pigmen- tation in the form of iron and manganese oxides, and in amounts of sand and con- glomerates in the limestone. The Claron for- mation is characterized by a rapid rate of erosion, largely a function of creep resulting from winter freeze-thaw activity and wash- 'Departnient of Genetics, Iowa State University, Ames, Iowa 50011. -Department of Botany, Weber State College, Ogden, Utah 84408. 701 702 Great Basin Naturalist Vol. 43, No. 4 away by summer thunderstorm runoff. Freeze-thaw cycles are most pronounced on south-facing slopes. Soil development is limited. Climatic features of the study area may be inferred from weather records kept at park headquarters, 2 km from the main am- phitheater. The average annual precipitation is 41 cm, falling largely in the form of winter snow and late summer thunderstorms. Mean January maximum temperature is 20 C; mean July maximum temperature is 27 C. The mean January and July minimum temper- atures are -13 and 7.6 degrees C, respective- ly. The main amphitheater is generally more xeric than these values indicate (Buchanan 1960), with higher temperatures and a great- er evaporative capacity of the air prevailing. Water availability to plants is decreased on the predominantly dry substrates, which have low infiltration rates and high runoff. Ravines and north-facing slopes within the main am- phitheater are probably more hydric than the adjacent plateau forest at park headquarters. Methods The vegetation of the study area was sur- veyed by means of 100 X 100 m 2 plots. Plots were subjectively placed in areas of homo- geneous vegetation among the clifflike for- mations. Within each plot, density and basal area of all mature trees (over 5 cm dbh) and density only of juvenile trees (less than 5 cm dbh) were recorded. An importance value was formulated for mature trees by summing the relative values of density and basal area. Understory vegetation was surveyed through the use of four 10 m line intercepts, placed at 2 m intervals. Importance values for under- story herbs and shrubs were determined through the summation of the relative values of. density, dominance, and frequency. Under- story species present in plots but failing to contact the survey lines were recorded as being present and this information was used in determining the relative frequency values. Using the importance values of dominant species, plots of similar composition were grouped together to define vegetative types. The mean importance values for all species in each type were determined, and com- parisons were made between types for all species having a mean importance value of greater than 1.0 (all trees, five shrubs, and seven herbs) using Sorensen's Index of Sim- ilarity (Sorensen 1948). All species present in the study area were recorded and assigned to vegetative types when possible. Voucher specimens are on file in the herbarium of Bryce Canyon National Park. Nomenclature employed is that of Welsh et al. (1981). Results and Discussion A complete list of species occurring in the main amphitheater is presented in the Ap- pendix. Two species, Puccinellia nuttalliana and Schizachyriiim scopariwn, are additions to the flora of Bryce Canyon as reported by Buchanan and Graybosch (1981). Additions to the flora of the Kaiparowits Basin, based on Welsh et al. (1978), are noted in the Appendix. Data to be presented below allows the di- vision of the breaks (as it occurs in the main amphitheater) into the following vegetative types. Some of these are variable and are fur- ther divided into phases. Each shall be dis- cussed in turn. 1. Pinus ponderosa—Arctostaphylos patula type (Pipe-Arpa type) 2. P. ponderosa—A. patula type, Cercocarpus montanus phase (Pipo-Arpa type, Cemo phase) 3. P. ponderosa—Pseudotsuga menziesii type (Pipo-Psme type) 4. Mixed coniferous type, Picea pungens phase (MC type, Pipu phase) 5. Mixed coniferous type, Abies concolor phase (MC type, Abco phase) 6. Pinus longaeva type (Pilo type) 7. Washes, clay and talus slopes Table 1 summarizes the dominant species of each vegetative type in terms of mean im- portance values. A similarity matrix is given in Table 2, comparing the various types and phases. Description of Types and Phases 1. Pipo-Arpa Type Occurrence: Canyon bottoms and south- facing slopes between elevations of 2,200 and October 1983 Graybosch, Buchanan: Bryce Canyon Plants 703 Table 1. Dominant species (in terms of mean importance value) in each vegetative type and phase. Types and phases are given the numerical designation employed in the text. Type or phase Species Trees Ahic's concolor mean s.d. Junipenis scopiilonini Piccd ptingens Finns flexilis Finns kmgaeiii Finns pondemsa Fscndotsnga mcnziesii Shrubs Aver ghibrnm Anielanchier utahensis Arctostaphylos patnhi Ceanotluis maiiinii Cercocaiyns monta n ns Jnniperns commnnis Mdhonid repens Purshici tridentdta Rihes cerenm Xdnthocephdhim sarotliide Grasses and forbs Astragahts kentropliytd Cirsinni drizonicttm Clemdtis colwnbidnd Crijtantha dbdta Cymopteris pnrptirens Elyntns sdlina Eriogoniini pdngnicense 81.7 33.3 29..3 27.2 13.0 24.2 25.8 37.9 24.2 21.4 27.2 29.7 2.3 89.7 25.3 8.1 36.3 29.7 7.6 17.2 22.5 3.0 44.9 23.6 23.1 31.5 6.6 48.4 1.0 4.0 128.3 5.9 19.2 71.3 158.8 172.8 102.1 .35.2 35.5 8.6 43.9 24.2 45.6 36.0 29.6 16.8 1.0 67.6 25.9 28.8 4.4 48.1 25.1 24.9 3.2 19.4 19.1 19.8 7.9 29.8 25.9 23.9 20.2 1.3 1.8 31.3 4.7 5.1 85.6 14.1 70.9 57.9 19.7 76.8 32.0 18.6 58.1 40.7 40.5 43.7 12.2 4.5 7.0 8.9 2.0 2.2 13.0 7.3 7.9 9.2 4.5 7.3 5.4 138.0 23.6 30.1 2.2 13.8 49.7 50.2 46.8 7.2 2.1 19.1 34.4 4.4 6.9 25.9 46.8 7.2 35.7 15.9 46.7 50.3 89.6 1.5 27.3 26.2 .39.6 40.5 60.1 3.3 1.0 1.2 1.4 16.1 2.5 2.9 1.7 5.6 4.4 46.1 1.9 1.0 4.9 1.7 1.0 1.7 6.4 ,3.9 4.0 1.4 6.7 6.5 6.6 4.7 1.9 2.1 4.0 4.4 3.3 6.6 2.7 3.5 4.6 3.5 34.4 4.0 .36.6 13.7 34.8 9.9 32.9 19.7 3.9 7.2 7.4 5.3 10.5 11.8 10.4 704 Table 1 continued. Great Basin Naturalist Vol. 43, No. 4 Speci Type or phase 1 2 3 4 5 6 5.9 2.6 2.0 1.0 6.4 11.0 6.2 7.1 2.6 8.8 7.9 13.8 14.2 19.4 17.9 14.7 6.7 1..3 2.0 8.9 3.1 3.2 6.7 7.0 4.4 1.5 4.4 2.5 1.3.6 1.7 4.8 11.1 1.0 10.7 10.2 4.2 10.1 12.9 3.2 9.6 Haplopappiis armerioidea llipit(uo})(i))piis filifolitis Ivesia sahulosa Liniiiti kinf^ii Litlio.spcnniiiti iiiultifloruui Mdchdcninthcra orindcUoide.' Oeiwtlicni hmdiijvarpa ( 'Inizoptiis liipnemndes 2,400 m, continuing to lower elevations out- side the study area. Occurs on substrates de- rived from both the Claron formation and Quarternary alluvium. Vegetation: The dominant tree is P. pon- derosa, with Jimiperus scopulorum and Piniis flexilis as common associates. Thickets of Quercus gamheUi occur, although in- frequently. The shrub layer is dominated by A. patiiki, with Mahonia repens and Ceo- nothiis martinii of secondary importance. The most common herbs are Elymiis salina and Oryzopsis hynienoides. In contrast to the ponderosa pine forests of the adjacent plateau, Piirshia tridentata is rare. A total of 65 species occurs in this type, .several being restricted to it. There are largely taxa (i.e., Mahonia fremontii, Streptanthus cordatus, and Euphorbia fendleri) that are nearing their upper elevational limits in the study area. Relation to other types and phases: Closest resemblance is seen between this type and the Pipo-Psme type and the Pipo-Arpa type, Cemo phase. It forms ecotones with all other types, as well as with a woodland of Jimi- perus osteospertna, Pinus edulis, and Quercus gambelii at the lower elevational limits of the study area. 2. Pipo-Arpa type, Cemo Phase Occurrence: Adjacent to washes, on allu- vium, between elevations of 2,200 and 2,300 m. Vegetation: The dominant trees are P. pon- derosa and /. scopulorum, with an understory dominated by Cercocarpus montanus. Arne- lanchier utahensis is an additional frequently encountered shrub. Arctostaphylos patula is infrequent and variable in importance values. Herbaceous vegetation is uncommon. Relation to other types and phases: This particular phase exists only in narrow bands adjacent to washes and is generally sur- rounded by the Pipo-Arpa type in its typical manifestation. Possibly recognized as a dis- tinct type if more widespread, it is most closely related to additional types dominated by P. ponderosa. The importance values of P. ponderosa and /. scopulorum are nearly iden- tical in plots assignable to this phase and those in the typical Pipo-Arpa type (Table 1). This phase probably exists as a result of peri- odic flooding, with subsequent alteration of the physical features of the substrate. This provides a microhabitat that evidently favors C. montanus over A. patula, but does not in- fluence the nature of the canopy. 3. Pipo-Psme Type Occurrence: North-facing slopes of the Cla- ron formation between elevations of 2,285 and 2,380 m. Soil development is extensive due to limited winter freeze-thaw activity. Vegetation: Dominance is shared by P. ponderosa and P. menziesii; Pinus flexilis is occasionally encountered. The principle October 1983 Graybosch, Buchanan: Bryce Canyon Plants 705 shnib is A. patuki, with M. repens a common associate. In several plots, however, the den- sity of trees was of sufficient magnitude to exclude most understory species. A signifi- cant increase in the abundance of more mesophytic species (i.e., Acer gkihnim. Cle- matis Columbiana) is noted when this type is compared to the Pipo-Arpa type. A total of 38 species was found in this type. Relation to other types and phases: Most closely related to the Pipo-Arpa type, this type also shows affinities with the phases of the MC type. Ecotones are formed with these communities. MC Type Occurrence: Canyon bottoms and steep north-facing slopes; substrates occupied are derived from both Red and White members of the Claron formation. Elevational distribu- tion is between 2,285 and 2,450 m; the low- est elevation corresponds to the furthest ex- tension of hoodoos. Ill-defined and somewhat polymorphic, this type is best described in terms of its two phases. Fifty-five species oc- cur within this type. 4. MC Type, Pipu Phase Vegetation: This phase is recognized by the consistent dominance of P. pungens in either pure stands or in mixed associations with ad- ditional conifers, the most common of these being P. ponderosa, P. menziesii, P. flexilis, and /. scopulorum. The understory varies from sparse along washes (where both C. montanus and C. ledifolius are frequent) to dense in plots not subjected to inundation. In the latter, A. patuki, M. repens, Juniperus communis, and A. glabrum predominate. Relation to other types and phases: This phase is most similar to the MC type, Abco phase. The differences between the two lie largely in the paucity of herbs and grasses in the Abco phase. Both are characterized by a high diversity of conifers, the primary differ- ence between the two canopies being the identity of the dominant tree. Ecotones occur with the Pipo-Arpa type, the Pipo-Arpa type, Cemo phase, and the Pipo-Psme type. On upper slopes this phase gradually thins to relatively isolated individual trees. 5. MC Type, Abco Phase Vegetation: Consistent dominance by A. concolor, and lack of dominance by P. pun- gens, is the hallmark of this phase. Most of the other conifers are common. Table 1 in- dicates P. ponderosa as having the second highest mean importance value. However, this phase is present on the White limestone where P. ponderosa is infrequent. In such stands, P. menziesii and P. pungens are the most common associates. Dominant shrubs are largely those of the Pipu phase, although P. tridentata is more common. Relation to other types and phases: This phase is most closely allied to the MC type, Pipu phase. The two are separated spatially; the Abco phase occurs only on north-facing slopes and canyons east and north of the Wall-of-Windows, with an isolated stand in the Queen's Garden. It is possible that the Pipu phase represents an early serai stage of the Abco phase, although no juveniles of A. concolor were found in plots assignable to the Pipu phase. More likely, the two phases rep- resent points along a moisture gradient. P. pungens seems able to exist in situations that are too dry to allow growth of A. concolor. The two phases do not form ecotones, except in the Queen's Garden. Here, A. concolor is found in a nearly pure stand in the shade of some isolated hoodoos. On more exposed sites at higher elevations in the same canyon it is absent, the area being dominated by P. pun- gens. Ecotones are recognizable between this phase and both the Pipo-Psme and Pipo-Arpa types. 6. Pilo Type Occurrence: This type is well defined only on badlands of the Claron formation, espe- Table 2. Matrix comparing vegetative types and phases through use of Index of Similarity (Sorensen 1948). Consuh text for information on the numbering svstem. 1 2 3 4 5 1 2 58 3 70 40 4 51 31.5 61 5 .32.5 31 42 60 6 .33 9.6 32 .30 11 706 Great Basin Naturalist Vol. 43, No. 4 cially on the ridgeline dividing the drainages of Bryce Wash and Campbell Canyon. There is no soil development; the substrate is gener- ally clay-limestone overlaid by gravels or larger particles. The type ranges in elevation from 2,200 to 2,400 m. Bailey (1970) reports the lowest elevational record of P. longaeva as being 2,200 m. Vegetation: P. longaeva in open stands is the usual appearance of this type. Pinus pon- derosa and P. flexilis may also occur, all trees being twisted and stunted. It should be noted that P. longaeva does not attain the wide girth that one usually associates with the spe- cies (the largest specimen had a dbh of 22 cm). Because of this it is doubtful that it reaches the extreme ages of 4,000-5,000 years that have been reported (Cronquist et al. 1972). LaMarche (1969) mentions that the oldest reported bristlecone pine in Bryce Canyon is 1,560 years of age; the number of trees and their exact location was not given. Shrubs are uncommon; only A. patula is of any real abundance. Understory vegetation generally covers less than 10 percent of the plots. The forb component differs from that of all other types; Linum kingii, Ivesia sahu- losa, and Eriogonum panguicense var. pan- guicense are the most common. A total of 48 species was recorded for this type, several of which are typical of subalpine zones. Based on the distributions given by Cronquist et al. (1972), Dixon (1935), Ellison (1954), Harring- ton (1954), and Welsh and Moore (1973), these are: Agropyron scribneri, Aqiiilegia sco- pulorum. Aster glaucodes, Erigeron simplex, Monardella odoratissima, P. longaeva, Poten- tilla fruticosa, Senecio attratus, and Silene pettersonii. Relation to other types and phases: As seen from the similarity matrix, this type has little in common with any others, the highest IS being 33. Ecotones are formed only with the Pipo-Arpa type, which extends up narrow washes draining the badlands. 7. Washes, Clay, and Talus Slopes Much of the main amphitheater is devoid of vegetation or contains only infrequently encountered individuals of numerous species scattered about the eroding Claron forma- tion. Structured plant communities tend to cluster in canyon bottoms, on the slopes of the wider canyons, or on rolling badlands. Species found on these barren areas have been recorded and are cited in the Appendix. Most of these slopes are not easily accessible; hence, the list may be incomplete. Plots were not used to survey such areas; if any pattern exists in the distribution of species, it has not been determined. The White beds form only vertical cliffs in the study area; plots were placed only on the Red member. Table 3. Distribution of trees as a function of moisture availability. Types and phases are given in order of most xeric to most niesic. A = Pilo type; B = Pipo-Arpa type; C = Pipo-Arpa type, Cemo pfiase; D = Pipo-Psme type; E = MC type, Pipu pfiase; F = MC type, Abco pfiase. Mean i.v. ABCDEF ABCDEF ABCDEF ABCDEF ABCDEF ABCDEF ABCDEF 0 1-fO If -20 2f-30 3f-4() 4f-5() 5f-6() 61-70 71-80 81-90 91-100 101-110 111-120 121-1.30 131-140 141-1.50 1.51-160 161-175 Species + + + + + + + + -I- -I- 4- -I- -I- + + + + + + + + + + + + + + Abco Jusc Pipu Pifl Pilo Pipo Psnie October 1983 Graybosch, Buchanan: Bryce Canyon Plants 707 Environmental Factors Influencing Distribution of Types At this point, it can only be speculated as to which environmental parameters are most critical in determining distribution of species. Moisture availability and solar insolation may be critical, but variation in substrates may also warrant consideration. Based on topogra- phy, substrate, and exposure, it is believed that moisture availability in communities in- creases in the order presented in Tables 3 and 4. Mean importance values for all trees and the most common shrubs are plotted as a function of increasing moisture availability. Certain trees cluster at a given end of the spectrum; A. concolor, P. pungens, and P. menziesii are common only in moist situa- tions. Pinus longaeva is restricted to the most arid sites. Juniperus scopulorum and P. flexilis occur at somewhat constant levels through- out, although both diverge from this pattern by decreasing in a given area. Pinus pon- derosa, although uncommon in the most xeric area, generally decreases with increasing moisture levels. For shrubs the pattern is similar though more complex. Acer glabrum, J. communis, and M. repens increase with moisture levels; A. patula decreases, C. martinii remains con- stant. However, A. utahensis and C. mon- tanus both demonstrate substantial increases in the Pipo-Arpa type, Cemo phase, appar- ently at the expense of A. patula and M. repens. From this it can be concluded that gradients of moisture availability are in- volved in sorting species into communities. However, moisture is not always the factor of paramount importance. Differences in sub- strate may be responsible for part of the pat- tern, especially in the increased abundance of A. utahensis and C. montanus in one given type. Only a complete ecological survey of the area will provide an answer to this question. Distribution of Endemic Species Species endemic to southern Utah and found within the main amphitheater are list- ed in Table 5, along with their ecological dis- tribution within the study area. Also given for each species is its current status (Federal Register, 15 Dec. 1980, vol. 45, No. 242). No status is given for E. panguicense var. pan- guicense, which, although restricted in range (Reveal 1965), is apparently not rare (Welsh et al. 1975, Welsh and Thome 1979). Status is defined by the following categories. Category 1. Information is presently on hand to support listing as en- dangered or threatened species. Category 2. Information is available that indicates a probable appropri- ateness of listing, but sufficient information is not yet available to support listing as endan- gered or threatened. Table 4. Distribution of most abundant shrubs as a huiction of moisture availability. Tvpes and phases given as in Table 3. Mean i.v. ABCDEF ABCDEF ABCDEF ABCDEF ABCDEF ABCDEF ABCDEF 0 1-10 11-20 21-30 31-40 41-50 51-60 61-70 71-80 81-90 91-100 101-110 111-120 121-130 131-140 + + + + + -I- + -t- + -t- + -I- -I- + + + + Acgl Amut Arpa Cema -I- Cemo Juco Mare 708 Great Basin Naturalist Vol. 43, No. 4 Category 3c. Taxa proven to be more abun- dant or widespread than pre- viously considered. Not under consideration at the present time. From Table 5 it may be seen that 9 of the 11 species listed occur in the Pilo type, 3 are foimd in the Pipo-Arpa type, and 2 occur in the MC type. Two species, Oxijtropis jonesii and Psoralea pariensis, were not found in a recognizable vegetative type. Species that occur only in the Pilo type have the narrow- est geographic distributions, although Eriogo- nimi panguicense var. panguicense is an ex- ception. It evidently has broader ecological tolerance inasmuch as it has been observed in several additional portions of Bryce Canyon. The narrow distributions of most of these species is no doubt a fimction of the uncom- mon occiu-rence of P. longaeva communities or other similar habitat throughout the High Plateaus. Within Bryce Canyon, most of these en- demics are restricted to the Claron forma- tion. In this study, however, Draba sub- alpina, Lesqiierella rubicundiila, and Toivns- endia minima were observed on Quarternary alluvium, a formation from which they have not previously been reported. Oxytropis jo- nesii, P. pariensis, and E. panguicense are not limited to the Claron formation throughout their geographic range. The remainder of the endemics have been reported only from the Claron formation. The majority of the vast number of endem- ic species found in southern Utah are restrict- ed to substrates derived from a specific geo- logic formation (Welsh 1979). Welsh notes that most of these taxa are found in areas of exposed parent material; soil development provides a barrier between plant and sub- strate. In the main amphitheater, soils are well defined only in the Pipo-Psme type. It is significant that no endemic species occur in this type. The distribution of endemic species in Utah is not a random one; fine-textured sub- strates support more species than coarser Table 5. Distribution of endemic species in the main amphitheater. Species Distribution in' Status' Utah (counties) Geologic distribution' Ecologic distribution in main amphitheater Castilleja levealii 1 Cniptantlid ochrolcuca 1 Dmha suhalpina .3c Eriof>pntim panguicense — var. panguicense Lesijuerelhi luhicundula 2 Lotuatiuni luininiuni 1 Oxiitropis joncsii 3t Penstemon bracteatus 1 Psoralea pariensis 1 Silene pettersonii var. ntinor 1 Townsendia minima 2 Garfield Garfield R; W R;\V Garfield, Iron, R; W; QA Kane, Millard Garfield, Iron, R; W Kane, Sevier, Washington C;arfield R; W; QA Garfield, Iron, R; W Kane Emerv, Garfield, W; Flagstaff limestone, L'intah Green River shale Garfield R Garfield, Kane R; alluvium and sandy alluvium Garfield, Iron R Garfield, Kane R; VV; QA Pilo type; wa.shes, clay- limestone slopes Pilo type; clav-liniestone slopes Pipo-Arpa type, Pilo type, MC type Pilo type, clay-limestone slopes Pipo-Arpa type, Pilo type, clay-limestone slopes Pilo type, clav-limestone slopes Sand-limestone slopes Pilo type, clav-limestone slopes clav-limestone slopes Pilo type, clav-limestone slopes Pilo type, Pipo-Arpa type, MC type 'See text for discussion. -'From: Reveal 1965, Welsh et al. 1975, Welsh 1978a, Welsh and Thome 1979. 'Refers to the entire range of species. From: Welsh 1978a, Welsh and Thorne 1979, personal observations. White beds, QA = Quarternary alluvium. 'Reveal (1965) defines the typical substrate of the species only by the designation "clay slopes. " With its w substrates other than those derived from the Claron formation. R = Red beds of Claron formation: W = ide geographic range it probably occurs on October 1983 Graybosch, Buchanan: Bryce Canyon Plants 709 ones, and desert and foothill vegetation is richer in endemic species than montane com- mimities (Welsh 1978b, 1979). Based on these observations, Welsh has developed a "pre- dictive model for establishing priority areas for the study of endangered and threatened plants of Utah," in which the highest priority is assigned to fine-textured soils supporting pinyon-juniper or desert shrub vegetation. A similar model may now be established for en- demics of the Claron formation. Outcrops of this formation supporting communities of bristlecone pine are most likely to contain endemic species. Based on this assumption, populations of these species have been found at several locations in Bryce Canyon. Species generally restricted to such habitat conditions are likely to be less widely distributed than those capable of invading other communities on the Claron limestone. Higher priority for listing as threatened or endangered species should be assigned to those taxa concentra- ting in such habitat. The protection of sites containing populations of bristlecone pine promises to be the most productive strategy for ensuring the continued survival of these plants. Stands of P. longaeva within Bryce Canyon National Park are of critical importance to botanical science. In addition to representing the preferred habitat of several endemic spe- cies, they are of interest as subalpine vegeta- tion found at atypical elevations. Additional investigation of such areas is likely to be fruitful in studies of the population biology of endemic species and the environmental fac- tors that govern plant distribution. Within the main amphitheater, the heavy use by visitors does not seem to present any danger to plant populations. The only activi- ties are hiking and horseback riding, both re- stricted to established trails. Few seem to stray from the trails because the steep topog- raphy makes getting lost or injured a high probability. By preventing expansion of the existing trail system, park officials can likely maintain species populations at the present levels. Summary An investigation on plant community structure in the main amphitheater of Bryce Canyon National Park has shown that the vegetative community previously referred to as the "breaks" is divisible into discrete vege- tative types. Although most of the area con- sists of sparsely vegetated cliffs and slopes, well-defined communities are found on gentle lower slopes, rolling badlands, and canyon bottoms. Four major types are recog- nized, some being further divided into phases. Each has been characterized as to canopy, understory vegetation, and relation- ship to other types. A checklist of all species found in the area is given in the Appendix. The ecologic distribution of several en- demic species of southern Utah is given. The Pinus longaeva type is shown to be the rich- est in rare plants. This habitat is also unique in that it contains species normally found at subalpine elevations. It is predicted that en- demic species of the Claron formation will tend to cluster in similar habitat. Protection of bristlecone pine communities is urged as the simplest means of providing protection for these rare plants. Appendix Checklist of species and their ecological distribution in the main amphitheater, Bryce Canyon National Park. I = Pipo-Arpa type, II = Pipo-Psnie type. III = MC type, IV = Pilo type, V = washes, clay and talus slopes. ° = ad- ditions to the flora of the Kaiparowitz Basin. Species Vegetative type I II III IV V + + -1- + + + -1- + + + + ACERACEAE Acer glubrwn Torr. Apiaceae (Umbelliferae) Cijniopterus pitrpureus Wats. Lonuitium minimum (Mathias) Mathias 710 Great Basin Naturalist Vol. 43, No. 4 Appendix continued. Vegetative type Species III IV Apocynaceae Apocijninii androsaemifoUum L. ASCLEPIDACEAE Asclepias (ispertihi (Decne) Woodson Asterac:eae (C^ompositae) Aster glaucodes Blake Chn/wtlunnnus luitiseosus (Pallas) Britt. C. pel try i (Gray) Greene Cirsiiim arizonictim (Gray) Petrak Erigewn simplex Greene" Haplopappus annerioides (Nutt.) Gray Htp)ienopcippus filifolius Hook. lltpnetwxijs acatilis (Piirsh) Parker //. richardsonii (Hook.) Cockerell Leucelene ericoides (Torr.) Greene Macluieranthera grindelioides (Nutt.) Shinners Petiadoria pwnila (Nutt.) Greene ^enecio attratus Greene" S. multdohatus T. & G. ex Gray Solkkigo sparsiflom Gray Stephanomerki teniiifolia (Torr.) Hall Tetradymia canescens DC. Townsendia exscapa (Richards) T. C. Porter T. tninima Eastwood Xantliocephalum sarothrae (Pursh) Shinners ( = Gtitierrezia sarothrae Pursh) Berberujaceae Mahonia fremontii (Torr.) Fedde (= Berheris fremontii Torr.) M. repens (Lindl.) G. Don. (= B. repens Lindl.) Betulaceae Betiila oeeidentalis Hook. Boraginaceae Crijptuntha abata Johnst. C. ochroleuca Higgins" Lithospennum multifloriim T. & G. Brassicac:eae (Cmciferae) Arabis penduUna Greene Descurainia sopliia (L.) Webb, ex Engler & Prantl. Draba subalpina Goodnui. & Hitchc. Lesqiierella rubicundula Rollins Pliysaria chambersii Rollins Streptantlius corchitus Nutt. ex T. & G. Thlaspi arvense L. Caprifoliac:eae Santbuctis caerulea Raf. Symphoricarpos oreopliihts Gray Caryophyllac;eae Arenaria fendleri (Rydb.) Fern. iiilene petiersonii Maguire var. minor Hitchc. & Maguire Cor.naceae Corntis stolonifera Michx.° CUPRESSACEAE Jiinipenis communis L. /. osteospenna (Torr.) Little J. scopidorum Sarg. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + October 1983 Graybosch, Buchanan: Bryce Canyon Plants 711 Appendix continued. Vegetative type Species Ekicaceae Aictostaphylos patiila Greene Elaeagnaceae Shephenlia cauadensis (L.) Nutt. El phorbiac.eae Euphorbia femllch T. & G. E. ItiiuLi Engelnni. ex Ives" Fabaceae (Legiuninosae) Astragalus convallariiis Greene" A. kcntropliyta Gray A. Dwfficarpas (Nutt.) Gray Oxi/tropis jonesii Barneby Psoralea pariensis Welsh & Atwood" Fac.aceae Qucrciis gamhclii Xutt. Gentianaceae GentUmella tenella (Rottb.) Borner° (= Gentiaiui tenella Rottb.) Suertia racliata (Kellogg) Kuntze" Lamiaceae (Labiatae) Monardella odoratissima Benth. Ll\A( EAE Linuni kingii Wats. L. perenne L. var. lewisii (Pursh) Eat. & Wright Ona(;raceae Calijlopluis lav andulue folia (T. & G.) Raven ( = Oenothera lavandiilaefolia T. & G.) Oenthera brachijearpa (Gray) Britt. Pinaceae Abies eoneolor (Gord. & Glend.) Lindl. ex Hildebr. Piceu pungens Engelmn. Pinus edidis Engelmn. & Wisliz. P. flexilis James ex Long P. tongaeca D. K. Bailey P. ponderosa Dougl. ex Lawson Pseudotsuga menziesii (Mirb.) Franco PoACiEAE (Gramineae) Agropijron eristatuvi (L.) Gaertn. A. scribneri Vasev" A. traehijcauhim (Linke) Malte Calamagrostis scopulorum Jones Ehjuais salina Jones Onjzopsis hijmenoides (R. & S.) Ricker ex Piper Poa contpressa L. ° Puccinellia niittalliana (Schult.) Hitchc. ex Jeps. Schizaehi/rium scoparium (Michx.) Nash ex Small" Sitanion hystrix (Nutt.) J. G. Smith Stipa eoliimbiana Macoun" POLEMONIACEAE Phlox austromontana Gov. PoLYC;ONACEAE Eriogoniim corymboswn Benth. £. panguicense (Jones) Reveal var. panguicense Ranunculaceae Aquilegia scopulorum Tidestr. Clematis columbiana (Nutt.) T. & G. {= C. pseudoalpina (Kuntze) A. Nels.) Ill IV + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + 712 Great Basin Naturalist Vol. 43, No. 4 Appendix continued. Vegetative type Species I II III IV V + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + Rhamnaceae Ceonothtts martinii Jones ROSACEAE Amekinchier titdhcnsis Koehn Cercocarpiis Icdifolius Niitt." C. niontanus Raf. Holodiscus chiniosus (Niitt.) Heller Ivesia sahulosii (Jones) Keck Potentilhi fruticosa L.° Purshia tridentatii (Piirsh) DC. Salicaceae Populus angustifolia James ex Torr. P. tremuloides Michx. Salix exigiia Nutt. Saxifragaceae Ribes cereiim Dougl. Scrophulariaceae Castilleja linariaefolia Benth. ex DC. C. revealii N. Holmgren Pedicularis centranthera Gray ex Torr. Penstemon bracteatus Keck Literature Cited Anderson, J. J., and P. D. Rowley. 1975. Cenozoic stratigraphy of southwestern high plateaus of Utah. Pages 1-51 in Cenozoic geology of south- western high plateaus of Utah. Geol. Soc. Amer., Special Paper 160. Bailey, D. K. 1970. Phytogeography and taxonomy of Pintis, subsection Balfotirianac. Ann. Missouri Bot. Card. 57:210-249. Brox, G. S. 1961. The geology and erosional devel- opment of northern Bryce Canyon National Park. Unpublished thesis, Univ. of Utah, Salt Lake City. Buchanan, H. 1960. The plant ecology of Bryce Canyon National Park. Unpublished di.ssertation, Univ. of Utah, Salt Lake City. Buchanan, H., and R. Graybosch. 1981. Revised check- list of the vascular plants of Bryce Canyon Na- tional Park, Utah. Great Basin Nat. 41:109-120. Cronquist, a., a. H. Holmgren, N. H. Holmgren, and J. L. Reveal. 1972. Intermountain flora. Vol. 1. Hafner Publ. Co., New York. Dixon, H. 19.35. Ecological studies on the high plateaus of Utah. Bot. Gaz. 97:272-320. Doelling, H. H. 1975. Geology and mineral resources of Garfield Co., Utah. Utah Geol. and Min. Survey, Bull. 107. Ellison, L. 1954. Subalpine vegetation of the Wasatch Plateau, Utah. Ecol. Mon. 24:89-184. Harrington, H. D. 1954. Manual of the plants of Colo- rado. Sage Books, Denver. LaMarche, V. C. 1969. Environment in relation to age in bristlecone pines. Ecology 50:53-59. LiNDQUisT, R. C. 1977. The geology of Bryce Canyon National Park. Bryce Canyon Natural History As- soc, Bryce Canyon, Utah. 1980. Slope processes and forms at Bryce Canyon National Park. Unpublished dissertation. Univ. of Utah, Salt Lake City. Reveal, J. L. 1965. Notes on three LItah Eriogonums. Proc. Utah Acad. Sci. 42:287-292. Sorensen, T. 1948. A method of establishing groups of equal amplitude in plant sociology based on sim- ilarity of species content. Kong. Dan Vidensk. Biol. Skr. 5:1-34. Welsh, S. L. 1978a. Utah flora: Fabaceae (Legumi- nosae). Great Basin Nat. .38:225-367. 1978b. Problems in plant endemism on the Colo- rado Plateau. Great Basin Nat. Mem. No. 2: 191-196. 1979. Endangered and threatened plants of Utah: a case study. Great Basin Nat. Mem. No. 3:69-80. Welsh, S. L., and G. Moore. 1973. Utah plants: Tracheophyta. Brigham Young University Press, Provo, Utah. Welsh, S. L., N. D. Atwood, and J. L. Reveal. 1975. Endangered, threatened, extinct, endemic, and rare or restricted Utah vascular plants. Great Ba- sin Nat. .35:327-376. Welsh, S. L., N. D. Atwood, and J. R. Murdoch. 1978. Kaiparowitz flora. Great Basin Nat. 38:125-179. Welsh, S. L., and K. H. Thorne. 1979. Illu,strated man- ual of proposed endangered and threatened plants of Utah. U.S. Fish and Wildlife Service, Denver, Colorado. Welsh, S. L., N. D. Atwood, S. Goodrich, E. Neese, K. H. Thorne, and B. Albee. 1981. Preliminary index of Utah vascular plant names. Great Basin Nat. 41:1-108. SEASONAL GROWTH OF THE TUI CHUB, GILA BICOLOR, IN PYRAMID LAKE, NEVADA Joseph L. Kennedy' .\bstrac:t.— Tui chubs collected from November 1975 through November 1977 from Pyramid Lake, Nevada, were analyzed for seasonal growth patterns. Major growth in length occurred during the fall and early winter, and major reproductive development occurred during the late spring and earlv suiumer. The tui chub, Gila bicolor, is the most abundant fish in Pyramid Lake, Nevada, and is the major food source of the threatened Lahontan cutthroat trout, Salmo clarki hen- shawi. The tui chub is found in the drainage of western Nevada and eastern Cahfornia from the San Joaquin system to southern Ore- gon and the Cokimbia River (LaRivers 1962). The tui chub is an opportunistic feeder that utihzes algae, benthic invertebrates, zoo- plankton, and fish (Snyder 1917, Kimsey 1954, LaRivers 1962, Langden 1978). The tui chub spawns in late June or early July; dur- ing this time they are found in large numbers along the shore (Snyder 1917, Kucera 1978). Aerial surveys have shown that large schools of tui chub are also found in the open water during the spawning season. Preliminary growth studies of the tui chub suggested that growth in length may not oc- cur during the time of year when food and temperature are optimum. Although this is unusual, the timing of the reproductive cycle and the pekk occurrence of some food items suggested the possibility of an atypical growth pattern. Methods and Materials From November 1975 through November 1977, 2,400 tui chubs were collected. All fish were measured to the nearest millimeter in fork length and weighed to the nearest gram in body weight. Age was determined from scales obtained from the left side of the fish above the lateral line. Validity of the scale-based ages and cal- culations was established using criteria sug- gested by VanOosten (1979) and Hile (1941). The seasonal growth pattern was portrayed by plotting the length achieved at various in- tervals throughout the year against the date of scale collection. The length achieved from time of annulus formation was estimated by subtracting length at time of formation of the last annulus from the length at time of cap- ture (Gerking 1966). The length at last annu- lus formation was determined by extrapola- tion of the body length-scale radius relationship. The body-scale relationships and length- weight relationships were calculated accord- ing to Tesch (1971). Condition factors (K) were calculated according to Carlander (1969). Results and Discussion Seasonal Growth in Length The Pyramid Lake tui chub population was composed of two morphological forms: coarse rakered with gill raker counts of 9 to 15 and a fine-rakered form with raker counts of 20 to 40. The coarse-rakered form was found inshore or on the bottom. The fine- rakered form was found in the upper 20 m, both offshore and inshore, but was not com- monly found offshore on the bottom (Vigg 1978). Their food habits differed, but their growth rates and patterns were similar (Ku- cera et al. 1978). The young fish showed almost continuous growth throughout their first two years of life. The fish collected inshore and offshore on the bottom had a slight decrease in Western Montana College, Dillon. Montana 59725. 713 714 Great Basin Naturalist Vol. 43, No. 4 110- 100- ?AgeO c £ a; u a o c ^ 60- 50- 40 30 20-1 ^ 10 1.. o i-u 0 J JASONDJ FMAM Month Fig. Seasonal growth curves for tui chub age-groups O through IV, collected in Pyramid Lake, Nevada. growth in late April and May, but the pelagic chubs collected at the station did not have an interruption in growth until their second year (171 mm FL). Age-groups II through IV showed the same general pattern of growth, with peaks in the fall and winter and a seasonal low during the late spring and early summer. These age groups never experienced a period of rapid growth in length but had a long, continuous growing season that lasted from late summer through the winter. The peak growth for age- groups II through IV was in late fall-early winter (Figure 1). The young-of-the-year fish achieved most of their growth in length dur- ing the first summer of life. This 0 age-group attained a length of 48.5 mm in July and 98.5 mm by September. Total group growth for the first year was 121.9 mm. Annulus forma- tion by II through V began in May, but some did not form an annulus and resume growth until as late as August. The beginning of growth occurred after the major portion of the energy required for reproductive tissue development had been expended, and rela- tively late in the summer from the standpoint of optimal environmental conditions. The growth curves for all age-groups, other than 0 and 1, showed growth during the time of year when the water temperature was cool- ing or at a stable, cold temperature (6 C) and little growth in lengths during the spring and summer when warmer temperatures occurred. Annulus formation also reflected the differ- ent growth patterns for young and adult fish. The coarse-rakered young-of-the-year fish formed their annulus/check mark in late April and May. This interruption in scale growth was very narrow and formed in a very short period of time. It also did not have the characteristic crowding of circuli that normally accompanied annulus formation. The fine-rakered chubs did not have an inter- ruption in scale growth during the first year. The adult fish (older than II) formed their an- nulus from late June through August. The an- nulus was more diffuse, had the characteristic crowding of circuli, and occurred over a longer time period. This was the case for both the fine-rakered form and the coarse- rakered form. The later annulus formation October 1983 Kennedy: Tui Chub 715 and the lack of growth in length during the summer months is explained by the timing of the reproductive cycle and probably intense competition from young-of-the-year fish. The tui chub began to accumulate repro- ductive tissue during March and April with a major increase in May and June (Kucera 1978). This was followed by spawning in July. The peak in reproductive tissue, in- dicated by the gonadal somatic index, coin- cided with the formation of the annulus. In- terestingly, the annulus formed by adult fish was actually a combination of an annulus and reproductive check marks. The annu- lus/check mark formed by the young-of-the- year coarse-rakered group occurred when the lake was rapidly warming. The rapidly changing water temperature aaid the accom- panying change in seasonal distribution pat- tern probably combined to stress the young fish and induce annulus formation. Following spawning, the adults began to grow, but the expected rapid growth period did not occur. The onset of adult growth in length coincided with the seasonal low of macroinvertebrates and periphyton (Rob- ertson 1978). Also, the spring peaks of zoo- plankton had started to decline, possibly from the feeding pressure exerted by the young-of-the-year chubs (Kennedy et al. 1977). But total zooplankton, dominated by Diaptonius sicilis, still numbered 18 to 41 or- ganisms per liter during this time (Lider and Langden 1978). This density of zooplankton was equal to or greater than that which Noble (1975) found during peak growth peri- ods for yellow perch. The growth rates continued to increase throughout the fall and winter, and peaked when periphyton and macroinvertebrates peaked. Growth during the winter months has also been reported for bluegills (Gerking 1966, Krumholz 1948), but I have not found reports of fish species having major growth periods during the winter months. Pyramid Lake may also be unique with the peak abun- dance of macroinvertebrates and periphyton occurring during the winter. In addition, the total zooplankton numbers are still relatively high during the winter months (Lider and Langden 1978). The young-of-the-year and the one-year- old fish grew during the warmer summer months, when the adults did not grow in length, and continued to grow during the fall and winter, pausing only briefly during early spring. The much longer growth period of the younger fish was also reflected in the larger increments of growth by the younger fish (I, 123 mm; II, 48 mm; III, 43 mm; IV, 37 mm; V, 35 mm; and VI, 47 mm). Seasonal Growth in Weight As with growth in length, the adults and young fish had different patterns of seasonal growth. The young fish (O, I) increased in weight throughout the year in much the same pattern as growth in length. For adults (some II and all older fish) growth in weight includ- ed both somatic and reproductive tissue and must be examined with this in mind. The adult fish increased in weight from August af- ter spawning through the following June. There was a significant decrease in weight following spawning, as would be expected. Close examination of the data showed that the mean weight following spawning was higher than the mean weight preceding the rapid increase in reproductive tissue (March), and this probably represents a slight amount of somatic growth that occurred during the spawning season. Summary The tui chub shows remarkably synchrony with its environment. The species has evolved to utilize the food available during the fall and winter for much of its somatic growth and to utilize the abundant zooplank- ton populations in spring and summer for re- productive tissue and growth by young fish. The differential utilization of food allows this species to maximize the energy going into the population and minimize the competition between size or age groups. This adaptation to the particular environment stresses the im- portance of not introducing exotic species that might compete with the chub or disrupt the timing of their seasonal growth or repro- ductive cycles. Acknowledgments The Pyramid Lake Painted Tribe initiated the ecological study of Pyramid Lake from 716 Great Basin Naturalist Vol. 43, No. 4 which tliis study developed. The research was funded by the Bureau of Indian Affairs, con- tract H50C 14209487. I wish to thank the en- tire crew of W. F. Sigler and Associates, es- pecially Denise Robertson, our age and growth specialist. Literature Cited C^ARLANDER, K. D. 1969. Handbook of freslivvater tisheiv biology. Vol. 1. Iowa State Univ. Press, Ames, Iowa. 752 pp. Gerkinc, S. D. 1966. Lengtli of the growinir seasons of the bluegill sunfish in northern Indiana. Verb. In- ternal. Verein. Lininol. 16:1056-1064. IIiLE, R. 1941. Age and growth of the cisco Coregoniis arteilti (LeSiieur) in the lakes of the northeastern highlands, Wisconsin. Bull. U.S. Bur. Fish 43(19;35):209-.3I7. Kennedy, Joseph L., E. L. Lider, and S. Robertson. 1977. The limnology of Pyramid Lake. Pages 3.37-354 in D. C. Green, ed., Desertic terminal lakes. UWRL Pub.. Utah State Univ. KiMSEY, J. B. 1954. The life history of the tui ehul), Siph- (iteles bicolor (Girard), from Eagle Lake, Califor- nia. California Fish and Game 40(4):.395-410. Krumholz, L. a. 1948. Variations in size composition of fish populations in recently stocked ponds. Ecolo- gy 29(4): 401-414. KicERA, P. A. 1978. Reproductive biology of the Tui cliub, Gila bicolor, in Pyramid Lake, Nevada. Great Basin Nat. 38: 203-207. KicEBA, P., G. Workman, D. Robertson, S. Vice;, R. W'halev, and R. Lancden. 1978. Life history of the Tui chub. Chapter 4 in W. F. Sigler and J. L. Kennedy, eds., Pyramid Lake ecological study. W. F. Sigler and Associates, Inc., Reno, Nevada. La.\c:den, R. W. 1978. Food habits of the Tui chub (Gila bicolor) in Pyramid Lake. Unpublislied tiiesis, Humbolt State Univ. 43 pp. LaRivers, I. 1962. Fislies and fisheries of Nevada. Ne- vada State Fish and Game Comm. 782 pp. Lhjer, E. L., a.nd R. Langden. 1978. Benthic ecology. Chapter 10 in W. F. Sigler and J. L. Kennedy, eds.. Pyramid Lake ecological studv. W. F. Sigler and .Associates, Inc., Reno, Nevada. Noble, Ric:hard L. 1975. Growth of young yellow perch [Perca flincscerxs) in relation to zooplankton pop- ulations. Trans. Amer. Fish. Soc. 104(4):731-741. Robertson, S. 1978. Benthic Ecology. Chapter 10 in W. F. Sigler and J. L. Kennedy, eds.. Pyramid Lake ecological study. W. F. Sigler and Associates, Inc., Reno, Nevada. Snyder, J. O. 1917. The fishes of the Lahontan system of Nevada and northeastern California. U.S. Bur. Fi.sh. Bull. 1915-1916(.35):3I-86. Tesc;h, F. W. 1971. Age and growth. Pages 98-131 in W. Bicker, ed.. Methods of assessment of fish production in fresh waters. Blackwell Sci. Pub!., Oxford. VanOosten, J. 1929. Life history of the lake herring {Corcgoniis artedii LeSueur), of Lake Huron as re- vealed by its scales, with critique of the scale method. Bull. U.S. Bur. Fish. 44:265-428. Vu;(;, S. 1978. Vertical di.stribution of adult fish in Pyra- mid Lake, Nevada. Great Basin Nat. 38:417-428. BIRD DISTRIBUTIONAL AND BREEDING RECORDS FOR SOUTHEASTERN IDAHO, UTAH, AND ADJACENT REGIONS Clavton M. White'. Herbert H. Frost', Dennis L. Shirlev', G. Merrill Webb', and Richard D. Porter' Abstract.— New distributional records or the status for .33 species of birds that have occurred within Utah, adja- cent southeastern portions of Idaho, or along the border of states siurounding lUah are reviewed. Four species, the Cattle Egret {Biibuhtis ibis). Common Moorhen [Gallinula chloropus). Great-tailed Crackle (Qtiiscaltis mexicaniis), and Common Crackle {Quiscalufi (jtiiscuhi), represent new Utali breeding records established within the past decade, and thev are commented upon. One other, the Black-tailed Cnatcatcher {Polioptiki melaniira), may be breeding in Utah. The Mockingbird (Miiiiiifi poh/^lottos) may likewise be breeding in southeastern Idaho adjacent to Utah. In 1972 the Birds of Idaho appeared (Bur- leigh 1972). It more or less represented the accumulation of a host of published and un- published accounts plus observations from the more than 20 years of Burleigh's own field work in Idaho. Unfortunately, Burleigh resided in northern Idaho, where most of the data come from; and a quick review of the book will reveal the spotty nature of data from southern Idaho. For Utah, Behle and Perry (1975) and Hayward et al. (1976) brought together and updated most Utah rec- ords. Currently, Behle (pers. comm.) is in the final stages of bringing together his life's work on the birds of Utah. With all this re- cent material and the summation of Behle's work at hand, it seemed appropriate to re- cord new data for adjacent regions in both states in cases where our new information clarified distribution or added new knowl- edge. Most of the Idaho data were gathered during an intensive study of raptors in and about the Raft River region of Cassia County, southeastern Idaho, 1976-1980 (Thurow et al. 1980). Some of these data are given to correct the misimpressions left by Burleigh. Much of the Utah material represents infor- mation accumulated since about 1974 by graduate students and faculty at Brigham Young University, but after the cutoff date for the Birds of Utah by Hayward et al. (1976). Where specimens were available their catalogue numbers are given in parentheses for either the Monte L. Bean Life Science Museum, Brigham Young University (BYU), or the Museum of Natural History, University of Utah (UU). Hereafter, reference to Bur- leigh, Hayward et al., or Behle and Perry will refer to the above references unless indicated by a date. Species Accounts: Cattle Egret {Buhulciis ibis). The Cattle Egret was recorded in Utah as occasional by both Behle and Perry and Hayward et al. During the summer of 1980, an estimated 25 pairs were reported nesting along with the Snowy Egret {Egretta thula) on a small Utah Lake island at the mouth of Provo Bay (Utah Division of Wildlife Resources personnel — UDWR; see also Kingery 1981c). On 28 July 1981 David Ng, a zoology graduate student, reported seeing 8-10 adults and 15 immature Cattle Egrets at this same heronry. Sub- sequently we visited the island on 11 August and found 29 nests, 14 with eggs, containing clutches of two to five eggs, as follows: 1/5, 2/4, 9/3, and 2/2. One of the 3 clutch nests had one egg pipping. At least 2 nests had eggs estimated to be no more than three-four days old and were laid in freshly built nests. Several clutches were laid in nests that had been previously used earlier in the season by 'Department of Zoology, Brigham Young University, Provo, Utah 84602. Division of Wildlife Resources, 1115 North Main Street, Springville, Utah 84663. 1063 East 400 North, Orem, Utah 84057. '.325 North 300 West, R.R. #1, Mapleton, Utah 84663. 717 718 Great Basin Naturalist Vol. 43, No. 4 Snowy Egrets, based on the fouling of the nests. At me remaining 15 nests young Cattle Egrets ranged from newly hatched to those moving about in trees and nearly capable of sustained flight. Three of the largest young were collected and these weighed .340 g (BYU 7617), 316 g (BYU 7618), and 340 g (UU 22,603). Palmer (1962) gives adult weights as 300-400 g. Their stomachs were full of orthopterans; one contained 13 heads. In 1982 some 50 nests were found. R. Isham (1975) studied the same island colony in 1973 and did not find this species breeding there with Snowy Egrets. Several of the Snowy Egret nests studied by Isham were marked with metal tags, and these same trees or nests were used by Cattle Egrets in 1981. Their es- tablishment in Utah, Colorado, and Nevada has been summarized by Kingery (1980c); and, according to Rogers (1982b), they may now be breeding about 80 km N of the Utah localities at Lake Walcott, Power County, Idaho. Cooper's Hawk (Accipiter cooperi). A spe- cies listed as an imcommon breeder for south- em Idaho by Burleigh and by Levy (1962), the Cooper's Hawk was found by us to be rather common in the canyons of the western slope of the Black Pine Mountains over- looking the Raft River Valley (see Thurow et al. 1980). We found them in all canyons that we were able to travel up. They nested both in aspen and conifer trees on south- and north-facing slopes. We suspect they were more common than we found them since we only explored canyons that had roads. Common Black Hawk {ButeogaUus anthra- cinus). Most of the data on this species comes from the 1960s, when it apparently moved into extreme southern Utah as a breeder about 1961-1962 (Hayward et al.). Only one specimen exists for Utah. Here we report a specimen that came to Brigham Young Uni- versity (BYU 7619) after it had been shot by himters. An adult female (wt. 866 g), it was reportedly found along the Virgin River N of Littlefield, Arizona, and 6-7 km S of the Utah border. The bird, with tail practically shot off, came to Stelline Ure, a Salt Lake City raptor rehabilitator, on 3 September 1980 and died within the day. It had been originally taken to Cedar City, where it ap- parently remained for two-three days before being taken to Salt Lake City. Broad-winged Hawk {Buteo platypterus). There were at least two individuals reported for Utah, one near Salt Lake City in 1970 and one in Provo in 1975. Behle and Perry considered it hypothetical, and Hayward et al. assign it an uncertain status. Recent re- cords indicate it to be a rare to casual mi- grant. Steve Hoffman (pers. comm.), Office of Endangered Species, U.S. Fish and Wild- life Service, and a hawk bander, recorded 17 individuals during fall migrations over a six- year period as follows (some of the records have been listed by Kingery 1980a): 1 ad 18 Sep 1977 Pilot Mt., Box Elder Co., UtaJT 2 ad- 19 Sep 1979 Goshute Mts., Elko Co., 1 im Nevada (just west of the Utah border) 1 ad 22 Sep 1979 Goshen Mts., Elko Co., Nevada (just west of the Utah border) 1 ad 24 Sep 1979 Wellsville Mts., Box Elder Co., Utali 2 ad 28 Sep 1979 Goshute Mts., Nevada 1 inim 30 Sep 1979 Wellsville Mts., Utali 1 inim 4 Oct 1979 Goshute Mts., Nevada 1 ad .5 Oct 1979 Goshute Mts., Nevada 1 ad 18 Sep 1980 Goshute Mts., Nevada 1 imm 2.3 Sep 1981 Goshute Mts., Nevada 3 imui .5 Oct 1981 Goshute Mts., Nevada 1 iiiini 21 Sep 1982 Goshute Mts., Nevada 1 imni 23 .Sep 1982 Goshute Mts., Nevada One of the above immatures was actually trapped but escaped before it could be taken from the trap and banded. On 5 May 1982 an immature was seen in the Cub Creek area of Dinosaur National Monument, Uintah Coun- ty, Utah, by members of the Utah Field Or- nithologists (UFO). This hawk has recently been reported about 160 km N of the Pilot Mts. — Wellsville Mt. area in Idaho (Rogers 1982a). Red-tailed Hawk {Buteo jamaicensis har- lani). Accounts by Behle and Perry and Hay- ward et al. give the impression that this race is rare within Utah. This race of the red-tail has recently been discussed by Mindell (1983) and an analysis of plumage characteristics given. Mindell outlines some of the diffi- culties of correctly identifying this form, es- pecially the immature, and the widespread nature of intergradation with the race ca- lurus. Despite field identification difficulties with typical harlani, several interesting sight records have occurred in Utah County in the past five years, and two specimens brought to Brigham Young University are worth report- ing. Since 1978 an adult bird of this form, presumably the .same individual, occupied a October 1983 White et al.: Bird Records 719 tree perch as part of a winter territory along Interstate Highway 15 near American Fork. It usually arrived by mid-December and was gone by mid-March. It could be approached to within 50 m and gave excellent opportu- nity for observation, especially of the tail. The one previous specimen for the state reported by Worthen (1973) was an imma- ture and thus not easily separable from im- matures of dark (black)-phased cahinis. Be- cause of this problem, its proper identity is open to question. In addition to that speci- men, we now have received two adults. One (BYU 7621) is a male in dark phase and has a typical harlani tail coloration. It weighed 965 g and was found injured at the Lehi air- port on 28 December 1981. The other, also a male (BYU 7622) from Provo, found 17 Janu- ary 1983 (wt. 948 g), has a nearly immaculate breast (light phase?), dark back, and a mot- tled whitish tail with a rusty tip almost iden- tical to the tail of an adult Ferruginous Hawk {Buteo regalis). The tail of this specimen is more like that of a harlani x cahirus inter- grade, although the breast is certainly unlike such intergrades (Mindell pers. comm.). The coloration of the tail and whiteness of the breast may be the result of intergradation with krideri. The sight records in Kingery (1982b), if correct, may show this race to be a regular visitor to Utah; and this may be a re- cent event, within the past two decades. Ferruginous Hawk {Buteo regalis). The en- tire species account in Burleigh misrepresents the status of this hawk, not only within Idaho but in southeastern Idaho in particular. We have not attempted to review the literature for all of Idaho but will restrict our remarks to southeastern Idaho. The reports of Power et al. (1975), Power and Craig (1976), and Thurow et al. (1980) put into perspective the density of this species in but two counties of southeastern Idaho, where as many as 50-75 pairs may nest in "good" food years (perhaps half that many in "poor" years). Porter (1951) was the first to discuss the species in Cassia County and suggested that it was probably a common summer resident. Burleigh and, earlier, Jollie (1952) were both convinced of quite the contrary, however, based on their limited experience in that re- gion and so it is thus published. The species has been shown to be rather cvclic (Thurow et al. 1980), and it may be that both Burleigh and Jollie (1952) visited the region in a low prey year. The available findings, however, vindicate Porter's initial impressions and show the species to be a common breeder in southeastern Idaho with as many as 20-30 pairs in an average year in a small area of Cassia County alone (Thurow et al. 1980). Mr. Jack Pierce (pers. comm.), who has been a resident of Malta, Cassia County, Idaho, for at least 50 years, remembers that two pairs nested on his property for "as long as I can remember." Although there is a general be- lief that the Rough-legged Hawk {Buteo la- gopus) is the only large buteo to "hover" dur- ing hunting bouts, we observed the Ferruginous Hawk to use stationary hovering during foraging on numerous occasions. Rough-legged Hawk {Buteo lagopus). This hawk reportedly leaves northern Utah by 16 April for its arctic breeding grounds (Behle and Perry), and in southern Idaho it is said to leave by "the last of April" (Burleigh: 66). We found a freshly shot individual (BYU 7616) on 4 June 1978 about 8 km E of Malta, Idaho. The bird was an immature male in ex- tremely worn plumage but in good physical condition (wt. 948 g). The tips of the primary feathers on the right wing were shot off. We saw another individual 5 km S of Malta on 11 May 1979 as it fed on a dead rabbit on the roadside. An individual, presumably the same, was seen in the above area again on 11 June 1979. Based on the tail coloration (see Cade 1955), this latter individual was also an immature from the previous summer's hatch. Pat Benson (pers. comm.) also had "summer" records from Gray's Lake, Idaho. Porter (1951) reported no late spring or early sum- mer sightings of this species in southeastern Idaho, and most of his data were from this same valley. American Kestrel {Falco sparverius). De- spite this species being listed as an uncom- mon breeder in that portion of southern Idaho encompassing Raft River Valley by Burleigh and by Levy (1962), we, however, found them to be common breeders there. They nested in deciduous trees (three pairs in Malta alone), in buildings (e.g., near Six Mile Canyon, two pairs), in cliffs (at least three pairs), in an abandoned silo (two pairs), or in juniper trees (many pairs, in hollows and in 720 Great Basin Naturalist Vol. 43, No. 4 Black-billed Magpie [Pica pica] nests). We saw kestrels enter squirrel holes that were in the cut banks of heavily eroded washes in at least two different localities. We suspected them of nesting in such holes because of their territorial behavior and because they carried food into them. Thus, casual observations, in the course of an intensive study of Ferru- ginous Hawks, suggested that as many as 25-30 pairs of kestrels bred in the valley. Nests in junipers were both at the edge of the stands of forest (juniper-sagebrush eco- tone), as are Ferruginous Hawks, and deeper within the forest itself. In this latter aspect they were quite unlike that reported by McArthur (1977), who had none of the 20 nesting boxes placed throughout his study area in juniper forest in Millard County, Utah, occupied. This may have been because kestrels were naturally rare there as breeders. McArthur did, however, have some utiliza- tion of boxes when placed on poles in open salt desert scrub areas. Likewise, Craig (1979), who worked about 120 km N of Raft River, had good utilization of nest boxes when placed in deciduous trees with large open areas around them or at the edge of woodlots. Merlin {Faico columbarius). Burleigh's dis- cussion of this species pertains mainly to ex- amples from the central or northern part of Idaho. A nest in a juniper was recently re- ported by Craig and Renn (1977) for the Snake River Plain somewhat north of Cassia County, Idaho. Closer to our study area there are nest records for Bannock County just to the northeast of Cassia County (Stanley A. Temple, pers. comm.). These consisted of egg clutches taken some 70 yrs ago and were ap- parently from nests in a riparian region. Be- tween May and July 1977 a pair of territorial Merlins were repeatedly seen near Bridge, Cassia County. They frequented an area of abandoned buildings surrounded by cotton- wood and box elder trees that contained dis- used Black-billed Magpie nests. Although no eggs were found, the actions of the pair sug- gested a breeding attempt. A territorial pair was not seen in subsequent years. On 5 Au- gust 1980 White found two young that ap- peared to be fledged about two weeks earlier 27 km NE of Montpelier, Bear Lake County, Idaho. The date and apparent age of the young suggested they were not too distant from their nest. The habitat was riparian, but there were no large trees other than a few scattered conifers on the hillsides that could harbor nests. If a nest was nearby, it may have been in a magpie nest in willows (Salix). Gyrfalcon (Falco rusticolus). Behle and Perry record this species as hypothetical for Utah and provide one sight record. Hayward et al. list it as of uncertain status but provide four additional sight records for northern Utah, all by reliable persons. Three other ob- servations for scattered locations should be placed on record. Howard Brinkerhoff (pers. comm. 1980), a falconer from the Uinta Ba- sin, saw one there "about three years ago " in the early part of a particularly cold winter. Joe Terry, a local falconer, Gerald Richards, a Provo biologist, and Steve Chindgren, Salt Lake Tracy Aviary, (pers. comm.) had a wild Gyrfalcon attracted to their trained Gyrfal- con used in falconry while in Cache Valley, Utah, on 27 October 1978. Lastly, Steve Chindgren (pers. comm.) showed White a photograph of a Gyrfalcon taken on 3 Febru- ary 1983 west of Kaysville, Utah. Chindgren and Larry Barker were hunting with a trained Northern Goshawk {Accipiter gentilis) when the falcon appeared. They watched it for nearly three hours, during which time it attempted to kill prey three different times. The falcon was identified as an immature fe- male because of size and plumage character- istics. To date, most individuals have been gray color phase, although one was called white phase. White, who has had more than 20 years' experience with Gyrfalcons on their arctic breeding grounds, has no doubt as to the identification of the falcon in the photos. Although the species does occasionally es- cape from falconers and may be seen in the wild, it seems doubtful that enough birds could escape at such a temporal and geo- graphic distribution to account for the Utah sightings. Based on the photograph and accu- mulation of records, we recommend that the species be removed from the hypothetical category and be considered an occasional winter visitant. Common Moorhen (GaUinula chloropus). Recorded as either a rare permanent resident (Behle and Perry) or of casual occurrence October 1983 White et al.: Bird Records 721 (Hayward et al.), this species has now estab- hshed another small breeding population, this time at Utah Lake. Hayward suspected their breeding as early as 1969 but found no defi- nite evidence. Webb found adults and three immatures on 31 July 1980 and saw them again on 11 November 1980 at Powell Slough near the Orem sewage ponds. On 18 July 1981 two immatures and on 29 August 1981 one immature was again seen at the same locality. Several observations of adults with broods along Interstate Highway 15 in the Provo Bay area of Utah Lake in May 1983 may indicate a spread of nesting to the more southern area. They were first sus- pected of breeding in southern Utah (Wash- ington County) as early as 1964 (see Hay- ward et al.). Whooping Crane {Grus anlericana). Until 1976 (see Behle 1981), this species was not heretofore recorded in Utah, although Utah may have been within its historical range. The species is being introduced into Gray's Lake, Caribou County, Idaho (Drewien and Bizeau 1978) where they are being fostered by the Sandhill Crane (Gnis canadensis). The migratory route normally takes cranes from this region through Colorado into New Mexi- co to winter. On 13 April 1983 Webb and Shirley saw two individuals (one adult and one immature) in a flock of approximately 400 Sandhill Cranes near Stewart Lake Waterfowl Man- agement Area, Uintah County, Utah. Both wore colored leg bands indicating that they were part of the cross-fostered flock from Gray's Lake. Single birds were also seen in the spring of 1981 and 1982 in the same area (UDWR personnel). Other records were for Ouray National Wildlife Refuge, near Vernal, Uintah County, for a summering immature also from the Gray's Lake population (King- ery 1976); Hyrum, Cache County, 25 Sep- tember 1981 (Kingery 1982a), and Jensen, Uintah County, 15 February 1981 (Kingery 1981a). Mountain Plover {Charadrius montanus). Although known for Utah, this species is rare enough to record recent sightings and speci- mens. Behle and Perry record it as a rare transient, and Hayward et al. give some six separate records for scattered portions of the state. Behle (1981) does not give any records for northeastern Utah. A series of recent rec- ords were gathered by Billy Green and A. Ray Johnson (field notes) in Uintah County, Utah (E of Bonanza), and in Rio Blanco County, Colorado. Five individuals were seen between 9 May and 20 June 1979 in Kennedy Basin on or near the Utah-Colorado border, and one was collected (BYU 7075) on 20 June 1979 about one km E of the Colorado border. The individual was a male with testes six mm long, and, although no definite evidence was found, they were suspected of being breeders rather than migrants. This species probably breeds marginally into Utah in the Uinta Basin. Hudsonian Godwit {Limosa liaemastica). Based on one April 1968 sight record, this species was considered hypothetical in Utah (Behle and Perry). However, two additional birds in alternate plumage were seen on 5 June 1976 at The Barrens, near Amalga, Cache County, Utah (Sordahl 1981) and sev- en were reported near Randlett, Uintah County, 2 May 1981 (Kingery 1981b). A specimen (BYU 7615) was collected on 15 May 1982 at Pelican Lake, Uintah County, Utah. The bird was a male in alternate plu- mage with gonads measuring 10 x 4.5 mm, weighed 256 g, and had heavy subcutaneous fat. The bird was accompanying a flock of 60 Marbled Godwit (Limosa fedoa). Snowy Owl (Nyctea scandiaca). Although this species is an occasional or rare winter visitant, there are only three extant speci- mens recorded by Behle and Perry and four mentioned by Hayward et al. On 25 Febru- ary 1982 we received a dead bird (BYU 7609) originally found alive "a few miles " northeast of the Salt Lake City International Airport in January. It was turned over to personnel of the Utah Division of Wildlife Resources and then taken to Hogle Zoological Gardens where it subsequently died. When prepared, we found 19 porcupine quills in the forearm and hand of the right wing. They appeared to have been imbedded in the bird for long enough to have healed but may have been the reason that the bird was debilitated and caught. The amount of dark pigmentation and ventral spots suggests that it was an im- mature, although it lacked any of the first- year gray-colored feathers that are often re- tained from the juvenile plumage. 722 Great Basin Naturalist Vol. 43, No. 4 Northern Hawk Owl {Surnia uhda). On 11 February 1976 one was brought to Brigham Yoimg University (BYU 5895) by a local resi- dent, Alice Chipman. It was found dead on the road "a few miles" from the Sundance Ski Resort in Provo Canyon, about 40 km NE of Provo, Utah County, Utah. The habitat there is mixed coniferous and deciduous for- est. It was some time before the owl was re- ceived by us and circumstances surrounding the finding were never adequately deter- mined. It is unknown how long it lay dead before being found. The skull was intact, al- though it was crushed, apparently by a ve- hicle. The legs, wings, back, and neck were too dry for proper specimen preparation and the sex was not determined. The plumage was badly worn but appeared to be that of an adult rather than an immature in that it lack- ed the more reddish brown underparts and broadly white-tipped tail (Bent 1938). A specimen taken in Alaska on 9 August shows a decidedly reddish brown cast which is de- scribed for the immature, but the amount of spotting on the upper parts agrees with pre- sumably adult Alaskan specimens collected on 8 January and 7 February. Ridgway (1914) made no distinction between adult and im- mature birds based on plumage. The speci- men shows traces of "hunger streaks" or "shock marks" across the rectrices, character- istic of feather growth in birds undergoing physiological stress, such as hunger, at the time the feather is growing. This might in- dicate a bird of the year raised in a food stress situation. The specimen appears to be the first record in western U.S. south of the Brookings, South Dakota (Serr 1978), Nampa, Idaho (Rogers 1974), and Pocatello, Idaho (Rogers 1978b) regions. We were unable to locate any winter records for Wyoming. Although this owl appears as an "invasion" species within different areas of its normal winter range, a perusal of Ajnerican Birds for 1975-76 did not show any unusual southward movement of this species that year. Further, most of the winter records are of birds in more open deciduous woods or prairie habi- tat rather than conifer habitat. Scissor-tailed Flycatcher {Tyrannus forfi- catus). There are four separate observations of this species in the literature, all based on sight records, and the species is considered accidental in Utah (Hayward et al.). The cir- cumstances svirrounding the following addi- tional observation seems noteworthy. On 29 May 1982 there was a large high pressure area over a considerable portion of Utah and Nevada. At Elberta, Utah County, Utah, winds from the south at 40-50 km/h lasted most of the day, but by evening they shifted within 10 min to the north and the temper- ature dropped 10-15 C within the same time period. These winds lasted throughout the night. The following morning, 30 May, was cahn, clear, and unseasonably cool. Martin Dobson and Jvidy Wray, two zoology gradu- ate students working on a bird project, found this flycatcher foraging along the fence row adjacent to Utah Route 68, eight km N of El- berta. They approached it to within 10 m in a vehicle and watched it forage for about 45 min as it moved south to north. They re- turned to camp to get cameras; but when they, along with several other people, re- turned an hour later to the location of the bird, it could not be found. Then, one week later, on 6 June, A. Ray Johnson (pers. comm.) was traveling along the same high- way about 17 km N of the previous observa- tion and saw a scissor-tail, perhaps the same individual, foraging along the fence row. He approached to within 25 m and watched it for about one min. He judged the bird to be an adult. Steller's Jay {Cyanocitta stelleri). Behle (1958) did not find this species in the Raft River Mountains, extreme northwestern Box Elder County, Utah, during his extensive studies there. He did, however, indicate that Clarence Cottam found them there and re- ported them to him. In light of Behle's find- ings, one observation should be placed on re- cord. Rosey Rosa saw one on 30 November 1950 at Standrod, a ranch area on the north slope of the Raft River Mountains just a few miles south of the Idaho border, and Porter saw one there from December 1950 through 28 January 1951. Behle and Perry indicate that the northern race annectens moves into northern Utah in winter. This observation may represent an individual of that race. American Crow {Corvus brachyrhynchos). The breeding distribution of this species for Utah has been most recently discussed by Richards and White (1963). Nowhere in Utah October 1983 White et al.: Bird Records 723 are breeding crows as common as they are in adjacent regions, as for example along the Humbolt River Valley, northern Nevada. Some nesting records come from extreme southwestern Utah, while the rest are from central and eastern Utah. Their status is poorly known for the northwestern part of Utah (Box Elder County) and adjacent Idaho. Burleigh does not describe their nesting dis- tribution for that region of Idaho. Levy (1950) called the species a common summer resident in the south central Idaho region he covered, although most of that region was well northward into southern Idaho. Over a straight line 56 km distance from 14 km N of Malta to Clear Creek, Box Elder County, Utah, we found five nesting pairs. The spac- ing of pairs was rather regular, and they oc- curred in the central part of the Raft River Valley. Although one nest was in a juniper tree in a cultivated riparian situation, the others were in deciduous trees in partially cultiv^ed or manipulated areas. This distri- bution contrasted markedly with the Com- mon Raven (Corvus corax), which nested al- most exclusively in juniper trees at the edges of the valley (the juniper-sagebrush ecotone), or on tall electric power transmission pylons that ran through the center of the valley. Three were on cliffs. In that same distance we found 15 raven nests. Four fresh crow eggs were found on 28 April, and at two nests, young 5-7 days old and 10-12 days old were found on 29 May. By contrast, ravens were starting to fledge by 1 June. Thus, the spatial placements of nests, nesting chronology, and density of ravens was notably different from crows, with only limit- ed overlap in these variables. In addition to the Clear Creek, Utah, nest a second nest was found along Grouse Creek, 20 km N of the town of Grouse Creek (ca. 10 km SW of Lynn). Both nests were in willows in a ri- parian situation. These are the only two nests thus far reported for that region (western Box Elder County) of Utah. Since Behle (1958) worked in the Lynn and Clear Creek areas and did not find them breeding, nor did he have reports from early investigations, and since Porter also failed to see them in the Raft River area during his studies in the early 1950s, they may be recently established there. Black-tailed Gnatcatcher {Polioptila mela- niira). This species is listed as hypothetical for Utah based on a single December 1969 record in St. George, Washington County (Behle and Perry). There are, however, nu- merous Nevada records adjacent to Washing- ton County. On 3 April 1982 members of a Brigham Young University ornithology field trip to Beaver Dam Wash, Washington County, watched a pair as they foraged about three to four km N of Lytle's Ranch. They pursued the pair for about IV4 hours and had many close observations. Members of the group, David Ng, Tod DeLong, Ed Robey, and David Fischer, have had experience with gnatcatchers in a variety of habitats and areas. The black crown was particularly evi- dent. Some of the observers had earlier on the trip seen the Blue-gray Gnatcatcher {Po- lioptila caendea). Then, on 2 June 1982, A. Ray Johnson and Dan Landeen watched a pair for 8 to 10 min in the same region of the Beaver Dam Wash as on the earlier date. It is doubtful that both parties found the same pair, and it seems likely that a small breeding population exists along Beaver Dam Wash. Members of the Weber State University or- nithology class saw a pair in Beaver Dam Wash about 1.6 km E of Terry Ranch in Josh- ua tree habitat on 14 May 1983 (via David Fischer, pers. comm.). Because of the varia- bility in the extent of black on the head, ex- amples of this species should be examined in the hand to verify these observations. Mockingbird (Mimus polyglottos). This species was not recorded as breeding in Idaho by Burleigh and apparently only straggles into that state. Stephens and Rey- nolds (1983) list it as an occasional erratic vis- itor to southwestern Idaho. Since it had not been recorded breeding in southern Idaho and its status in northern Utah is not clearly defined and breeding records are rare there (Hayward et al.), the following observations are of value. Steve Hoffman (pers. comm.) saw adults feeding young in late April 1974, 16 km N of the Utah-Idaho border on the W side of the Sublette Hills, Oneida County, Idaho. By 1977, they were reported another 160 km northward at Atomic City, Bingham County, Idaho (Rogers 1978a). On 28 June 1947 Porter saw five together at Locomotive Springs, Box Elder County. 724 Great Basin Naturalist Vol. 43, No. 4 On 16 June 1953 Porter (Porter, Bushman, and Behle unpubl. ms.) found a nest with three young at Dugway Proving Grounds, Tooele County, Utah (see photograph, page 109 in Hayward et al.). The habitat was desert scrub with an occasional juniper tree. The nest was about 0.7 m above the ground in a 2 m fourwing saltbush {Atriplex canes- cens). The Dugway area was used again in 1966. The species was attempting to breed north of Tooele as early as 1934 (Woodbury et al. ms.). Shirley observed two singing males on territories 5 km SE of Gold Hill, Tooele County, Utah on 25 May 1982. Bendire's Thrasher {Toxostoma bendirei). Mainly a species of the southern half of Utah, there are scattered sight records in north cen- tral and northwestern Utah (Behle and Per- ry). A. Ray Johnson and Billy Green (field notes) took photographs of a pair and recent- ly fledged young in the northeastern portion of the state at Coyote Wash, 17 km NE Bo- nanza, Uintah County, Utah (see Kingery 1980c). They were seen through the period 31 May — 4 June 1980. This may represent the northern extreme of the breeding range. The species was reported again in 1981 from Randlett, Uintah County, Utah (Kingery 1981c). Ovenbird (Seiurus aurocapillus). Called hypothetical by Behle and Perry and Hay- ward et al. because of the lack of appropriate documentation (specimen or photograph), the species is now represented by a specimen found by Lloyd Gunther, formerly of the U.S. Fish and Wildlife Service, in Brigham City, Box Elder County, Utah on 20 September 1977 (BYU 5860). The specimen weighed 15 g and is thought to be an adult female. When prepared it was freeze dried, so sex and skull ossification condition could not be examined. Canada Warbler {Wilsonia canadensis). This species was not listed by Behle and Per- ry, and only the date and location of collec- tion was mentioned by Hayward et al. The specimen (BYU 5390) is an adult male in al- ternate plumage, based on color and mark- ings, and was well preserved as a mummy ex- cept for the loss of the right eye and portions of the right side of the face. This bird was found dead at Callao, Tooele County, Utah, along with numerous other dead and dried birds at the base of a cottonwood tree. They perished in an unseasonable cold and snowy spell between 30 April and 23 May 1975 (see Whitmore et al. 1977 for a discussion of mor- tality during this period). Many of the speci- mens were in a similar state of preservation due to the extremely arid conditions of Cal- lao. The date of 31 May 1975 given by Hay- ward et al. was the date the bird was found and probably represents at least a week after the species actually arrived in Callao, per- haps as much as three weeks after arrival. Northern Cardinal {Cardinalis cardinalis). This species had not been heretofore record- ed for Utah. An adult male was seen at the feeder of Merlin Killpack in Ogden, Utah, on 10 March 1983. The bird (BYU 7620) was trapped at the feeder for observation and to photograph, but died before it could be re- leased. It weighed 44.5 g and was in good condition. There is no indication that the car- dinal was an escapee from captivity. Indigo Bunting {Passerina cyanea). Bur- leigh does not list this species for Idaho, and Stephens and Reynolds (1983) list it as acci- dental for southwestern Idaho. On 18 May 1979 at the mouth of Six Mile Canyon, Raft River Valley, Idaho, we found a male in al- ternate plumage accompanied by a chestnut- colored female, which we took to be also of this species rather than the more tan-colored female of the Lazuli Bunting {Passerina amoena). The male was seen to interact on two occasions with a male Lazuli Bunting. Both times the Lazuli was seen chasing the Indigo Bunting. The pair was seen briefly as they flew across the road on 19 May but not thereafter. Sage Sparrow {Amphispiza belli). Behle and Perry indicated that this species occurs normally in Utah between March and the end of November and leaves the state in mid- winter, although it has also been known to occur in extreme southwestern Utah during winter (Hayward et al.) This species, in fact, occurs over a much wider portion of western Utah throughout the entire year, though more scarce and spotty in winter, and should be considered a permanent resident. Porter et al. (unpublished ms) commonly found loose flocks of 3 to 5 and an occasional single spe- cies flock of up to 50 individuals from De- cember through March in Dugway Valley, October 1983 White et al.: Bird Records 725 Tooele County, in 1952-1954. They occu- pied salt scrub habitat where greasewood (Sarcobatiis) was the principal plant. Lark Bunting {Calamospiza nielanocorys). This species occurs near the western edge of its breeding range in western Utah and is rare there (see Porter and Egoscue 1954), with a poorly documented breeding distribution. Behle and Perry list it as an uncommon tran- sient for most of Utah outside of the Uinta Basin, where it is a regular breeder. Hayward et al. list four other scattered or isolated pre- sumed Utah breeding records outside the Uinta Basin based on time of collection or re- ported egg clutches. Burleigh stated that it reaches the extreme western portion of its breeding range in southern Idaho, that it is a local and uncommon summer resident, and he lists a few summer records of birds in breeding condition. One mentioned by Levy (1962) from the Caribou Basin, some 160 km NE of Raft River, had testes in breeding con- dition on 28 May. Stephens and Reynolds (1983) list it as an accidental for southwestern Idaho (they consider 114°W Long, about 32 km W of our study area, as the eastern limits of the region covered). On 24 May 1979 we saw a male in Raft River Valley, and on 19 June 1979 found a pair, the male still court- ing, near the mouth of Six Mile Canyon, Raft River Valley, Cassia County, Idaho. Then, on 12 July 1979 we found three other pairs in the Black Pine Valley (the valley ex- tends from Box Elder County, Utah, into Oneida County, Idaho, and about 19 km SE of the Six Mile Canyon). One male with food in its bill as though feeding young was about 12 km W of Snowville, Utah; directly N about 5 km was another foraging pair, and about IV2 km farther N into Idaho was a third male also gathering food. The habitat is a mixed greasewood-sagebrush-grass com- mimity. Harris' Sparrow {Zonotrichia querula). The temporal stay of this species in Utah is given by Behle and Perry as late October to the end of April. It reportedly arrives later than most wintering finches and departs later than many. Its spring departure may in fact be correlated with weather. In 1983 unsea- sonably cold weather with intermittent snow prevailed throughout the spring with a heavy snow fall during the period of 10-13 May and again on 16 May. On 14 May a bright- pink-billed adult male in alternate plumage was seen in Mapleton, Utah County, Utah, by Porter. It remained until 18 May. Two other individuals also occurred in Pleasant Grove during the same period, arriving on 8 May and departing on 20 May (UDWR per- sonnel). The late date suggests that the spe- cies may stay two to three weeks longer than previously recorded, but these later depar- tures may be dictated by weather conditions near normal departure time. Bobolink (Dolichonyx oryzivorus). Because recent data on the Bobolink are scanty and nesting information poor (Hayward et al.), observations from the past few years are of interest. Shirley counted six territorial males on 12 June 1982 in a grassy pasture W of In- terstate Highway 15 near Springville, Utah County, Utah. This small nesting population has been observed at the same location for several years. It is unknown whether success- ful nesting takes place. Each year the grass hay is cut part way through the nesting sea- son, which may destroy the nests. Additional sightings by Webb in 1981 and 1982 of a breeding pair the first week in June near Midway, Heber Valley, may indicate nesting in that area. Great-tailed Grackle [Quiscalus mexi- canus). This species is not mentioned by ei- ther Behle and Perry or Hayward et al. for Utah. The range, habits, and comparison with a close species, the Boat-tailed Grackle {Quiscalus major) has recently been reviewed by Pruitt (1975), and the former is shown to range to about central Arizona and New Mexico. The first Utah record was of a male in worn plumage seen on 1 July 1977 at Mapleton, Utah, by Porter (unpublished ms). The most distinctive feature about the bird, other than the tail, was the dis- proportionately small head and neck. Then, on 6 June 1978, Webb saw this species along the Virgin River S of St. George, Washington County, Utah. In separate sightings during the week of 13-17 May 1980, as many as 8 were observed near the sewage ponds at Washington, not far from St. George, by Webb and Steve Hedges. Hedges supplied a photograph verifying a state record (Kingery 1980b). On 9 October 1980 Webb then ob- served a group of 10 (males, females, and 726 Great Basin Naturalist Vol. 43, No. 4 immatures) S of Washington. During May 1982 breeding pairs were again seen at the Washington sewage ponds and at Ivin's Res- ervoir, 12 km W of St. George. Territorial males were observed during the spring of 1983 at Ivin's Reservoir, and on 7 May 3 males and 2 females were seen flying up Ma- gotsu Creek 1.6 km W of Veyo, Utah. During the 1982 Christmas bird count, 5 were seen on the east shore of Utah Lake not far from Provo by Webb. They are also reported to have reached Bicknell, Wayne County, Utah, by 18 April 1981 (Kingery 1981b). The species has been expanding its range beyond that shown by Pruitt (1975) into sev- eral western states. It was recorded to breed for the first time in California's lower Colo- rado River Valley in 1969 after having been seen first in 1964 (Small 1974). For Colorado the first breeding occvirred in Monte Vista in 1973 (Stepney 1975), when eight nests were foimd, and in Nevada it occurred in Ruby Valley and Sunnyside in 1981 (Kingery 1981b). Common Crackle {Quiscalus quiscuki). This species is listed as accidental by Hay- ward et al. and as a rare transient by Behle and Perry, based on scattered records throughout the year. They were first found breeding in 1977 at Vernal (Behle 1981). Then on 22-23 May 1981 three nests were found on the grounds of the Dinosaur Mu- seum of Natural History, Vernal, Uintah County, Utah, by Steve Hedges. Again on 15 May 1983, four nesting pairs were observed (see Kingery 1981b). It may also be breeding in the area of Utah Lake based on several April 1983 records and an adult male found dead on 22 April 1983 in SpringviUe (BYU 7623) that had testes in a breeding condition (12 X 8 mm). The bird weighed 125 g. Scott's Oriole {Icterus parisoriim). Al- though there are scattered records for this species throughout Utah, neither Behle and Perry nor Hayward et al. mention the 1936 observations of Twomey (1942) for Powder Springs, Uintah County, Utah, nor the adja- cent Rio Blanco County, Colorado, records. Twomey 's map shows Powder Springs to be in Colorado; his description of the location, page 359, places it in Utah. The 7.5 min USGS Cliff Ridge quadrangle for Utah matches Twomey's description with a Pow- der Springs Wash, but a Powder Springs lo- cation does not appear on the appropriate (Mellen Hill) 7.5 min quadrangle for Colo- rado to match Twomey's map. Behle (1981) subsequently rightly mentions the Powder Springs location in Utah. In 1979 A. Ray Johnson and Billy Green (field notes and pers. comm.) found this species in June and July 10-12 km from Twomey's Colorado observa- tion, and in August (11-14) four to five indi- viduals were seen directly west of the Colo- rado sightings some 12-16 km into Uintah County, Utah (see Kingery 1980c). It appears that a limited but consistent population oc- curs in that region that lies between Rangley, Colorado, and Ouray, Utah, and probably has been there at least in the 40-year period since Twomey. It is interesting to note that the Colorado Field Ornithologists (Reddall 1976) opted to drop it from the state list because of what they considered unconvincing details prior to 1974, and then added it in 1975 based on an early May individual from Jeffer- son County. Although Burleigh does not list them for Idaho, Steve Hoffman (pers. comm.) observed several in the Sublette Hills, ap- proximately 8 km S of Holbrook, Oneida County, Idaho, just to the east of the Raft River Valley during the breeding season of 1974-75. He believed them to be nesting. Acknowledgment In addition to the graduate students men- tioned in the text, we also thank M. Ralph Browning, U.S. National Museum, for com- ments on the manuscript, and Tom L. Thu- row, Dan Johnson, Richard Howard, William Mader, and T. Craig White for help in the field. Steve Hoffman, U.S. Fish and Wildlife Service, Albuquerque, New Mexico, supplied us with several pertinent records. A. Ray Johnson and Billy Green were funded on a contract from Burns and McDonnell Engi- neering Company through H. D. Smith. Literature Cited Behle, \V. H. 1958. The birds of the Raft River Moun- tains, northwestern Utah. Univ. Utah Biol. Ser. 11(6): 1-40. 1981. The birds of northeastern Utah. Occas. Publ. No. 2, Utah Museum Nat. Hist., Univ. Utah. October 1983 White et al.: Bird Records 727 Behle, W. H., and M. L. Perry. 1975. Utah biid.s: guide, check-list and occurrence chart.s. Utah Mus. Nat. Hi.st., Univ. Utah, Salt Lake City. Bent, A. C. 1938. Life histories of North American birds of prey. Part 2. Bull. U.S. Nat. Mus., 170. Burleigh, T. D. 1972. Birds of Idalio. Caxton Printers, Caldwell, Idaho. Cade, T. J. 1955. Variation of the common rough-legged hawk in North America. Condor 57:313-346. Craig, T. H. 1979. The raptors of the Idaho National Engineering Laboratory Site. U.S. Dept. Energy, Idaho Operations Office, IDO- 12089. Craig, T. H., and F. B. Renn. 1977. Recent nestings of the merlin in Idaho. Condor 79:392. Drewien, R. C. and E. G. Bizeau. 1978. Cross-fostering whooping cranes to sandhill crane foster parents. Pages 201-222 in S. A. Temple, ed.. Endangered birds — management techniques for preserving threatened species. LIniv. Wisconsin Press, Madi- son, Wisconsin. Hayward, L. C, C. Cottam, A. M. Woodbury, and H. H. Frost. 1976. Birds of Utah. Great Basin Nat. Mem., 1. IsHAM, R. S. 1975. The spatial distribution of the nests of the black-crowned night heron (Nijcticorax nyc- ticorax) and the snowy egret (Leucopharyx thula) in central Utah. Unpublished thesis, Brigham Young Univ., Provo, Utah. 52 pp. JoLLiE, M. 1952. Comments on the check-list of the birds of Idaho. Condor 54:172-173. Kingery, H. E. 1976. Mountain West. Amer. Birds 30:982-985. 1980a. Mountain West. Amer. Birds 34:184-187. 1980b. Mountain West. Amer. Birds 34:800-803. 1980c. Mountain West. Amer. Birds 34:914-918. 1981a. Mountain West. Amer. Birds 35:321-323. 1981b. Mountain West. Amer. Birds 35:846-849. 1981c. Mountain West. Amer. Birds 35:963-966. 1982a. Mountain West. Amer. Birds 36:201-203. 1982b. Mountain West. Amer. Birds 36:315-317. Levy', S. H. 1950. Summer birds in southern Idaho. Mur- relet 31:2-8. 1962. Additional summer southern Idaho bird notes. Murrelet 43:279-292. McArthur, L. B. 1977. Utilization of nest boxes by birds in three vegetational communities with spe- cial reference to the American kestrel {Falco sparverins). Unpublished thesis, Brigham Young Univ., Provo, Utah. Mindell, D. p. 1983. Harlan's hawk {Biiteo jamaicensis harlani): a valid subspecies. Auk 100:161-169. Palmer, R. S. (ed). 1962. Handbook of North American birds, vol. 1. Yale Univ. Press, New Haven, Connecticut. Porter, R. D. 1951. The status of rough-legged hawks in Idaho. Condor 53:257-258. Porter, R. D., and H. J. Egoscue. 1954. The lark bunt- ing in Utah. Wilson Bull. 66:219-221. Power, L. R., and T. H. Craig. 1976. Status of nesting ferruginous hawks in the Little Lost River valley and vicinity, southeastern Idaho. Murrelet 57:46-47. Power, L. R., R. Howard, C. H. Trost. 1975. Popu- lation status of the ferruginous hawk in south- eastern Idaho and northern Utah, Pages 153-157. in J. R. Murphy, C. M. White, and B. E. Harrell, eds.. Population status of raptors. Raptor Re- search Foundation, Raptor Report No. 3, Vermil- lion, South Dakota. Pruitt, J. 1975. The return of the great-tailed grackle. Amer. Birds 29:985-992. Reddall, J. 1976. Colorado field ornithologists' official records committee report 1972 through 1975. Western Birds 7:81-97. Richards, G. L., and C. M. White. 1963. Common crow nesting in Utah. Condor 65:530-531. Ridgway, R. 1914. The birds of Middle and North America. Bull. U.S. Nat. Mus. No. 50, part VI. Rogers, T. H. 1974. Northern Rocky Mountain — Inter- mountain region. Amer. Birds 28:80. 1978a. Northern Rocky Mountain — Inter- mountain region. Amer. Birds 32:231-235. 1978b. Northern Rocky Mountain — Inter- mountain region. Amer. Birds 32:378. 1982a. Northern Rocky Mountain — Inter- mountain region. Amer. Birds 36:875-877. 1982b. Northern Rocky Mountain — Inter- mountain region. Amer. Birds 36:998-1000. Serr, E. M. 1978. Northern Great Plains region. Amer. Birds 32:367. Small, A. 1974. The birds of California. Collins Books, New York. SoRDAHL, T. A. 1981. Phenology and status of the shore- birds in northern Utah. Western Birds 12:173-180. Stephens, D. A., and T. D. Reynolds. 1983. Birds of southwestern Idaho. Great Basin Nat. 44. In press. Stepney, P. H. R. 1975. First recorded breeding of great- tailed grackle in Colorado. Condor 77:208. Thurow, T. L., C. M. White, R. P. Howard, and J. F. Sullivan. 1980. Raptor ecology of Raft River Valley, Idaho. EGG-2054, E. G. and G. Idaho, Inc., Idaho Falls, Idaho. Twomey, a. C. 1942. The birds of the Uinta Basin, Utah. Annals of the Carnegie Mus. 28:341-490. Whitmore, R. C, J. A. MosHER, AND H. H. Frost. 1977. Spring migrant mortality during unseasonable weather. Auk 94:778-781.' Worthen, G. L. 1973. Harlan's hawk from LItah: first record for the Great Basin. Wilson Bull. 85:79. BIRDS OF SOUTHWESTERN IDAHO Daniel A. Stephens'-^ and Timothy D. Reynolds^-^^ .\bstract.— Based on personal birding experience, interviews with local birders, written comments and responses to a preliminary check-list, and a review of state and federal agency records and the ornithological literature, a list of 333 species of birds known to occur in southwestern Idaho is presented. The preferred habitat, season of use, breed- ing status, and relative abvmdance of each species are given. Idaho has extreme geologic, topographic, and chmatic diversity. The diversity of habi- tats and microhabitats is reflected by the abundance of vertebrate species within the state. In this regard, southwestern Idaho (south of the Salmon River and west of the 114° Meridian; Fig. 1) is no exception. With- in this region, elevation ranges from around 350 m ASL in Hells Canyon on the Snake River to over 3,600 m ASL in the White Cloud Peaks of the Sawtooth Mountains. Al- though sagebrush steppe and coniferous for- ests are dominant vegetation types in south- western Idaho, numerous pond, marsh, lake, reservoir, agricultural, alpine, and riparian habitats are present. As the varied habitat suggests, there is a concomitant diversity in the avifauna. Here, typically northern taiga species such as the Boreal Owl {Aegolius fu- nereus) can be found less than 200 km from strictly desert species like the Black-throated Sparrow {Amphispiza bilineata). The objec- tive of this study was to develop an accurate check-list of the birds occurring in south- western Idaho, documenting the breeding status, abundance, season of use, and pre- ferred habitat for each species. Material and Methods Although many regional bird check-lists are generally a combination of birds known to occupy an area as well as birds suspected to be in the region, our list (Table 1) includes only those species for which there are re- liable records. Data were gathered from a series of interviews with local birders, re- viewing wildlife records from various state and federal agencies, an extensive review of the ornithological literature, and personal birding experience in southwestern Idaho. Fig. 1. The southwe,stern Idaho study area (.shaded), south of the Salmon River and west of the 114° longitude. 'Department of Biology, Boise State University, Boise, Idaho 83725. 'Present address: Biology Department, Central Washington University, Ellensbiirg, Washington 98926. 'Present address: Biology Department, Idaho State University, Pocatello, Idaho 8.3209. 728 October 1983 Stephens, Reynolds: Idaho Birds 729 Table 1. Birds of southwestern Idaho. Introduced species are preceded by an asterisk (°) Taxa Abundance', season and breeding status- Preferred habitat' References Gaviiformes Gaviidae Red-throated Loon. Gavia stellata V6 Arctic Loon. G. artica M6 Common Loon. G. irnmer B4, M3, W4 Yellow-billed Loon. G. adamsii 7 PODICIPEDIFORMES Podicipedidae Pied-bill Grebe. Podihjmbus podiceps 32, W4 Horned Grebe. Podiceps auritus M3, W4 Red-necked Grebe. P. grisegena M5 Eared Grebe. P. nigricoUis 33, M2, W4 Western Grebe. Aechmophoriis 32, M2, W4 occidentalis 23, 7, 36 MC, JD, RK, AL 32 7, 13, 23, MC, DJ, TDR, DAS Pelecaniformes Pelecanidae American White Pelican. Pelicanus S4, M3 erijthrorhijnchos Brown Pelican. P. occidentalis V6 Phalacrocoracidae Double-crested Cormorant. B3, W4 Phalacrocorax auritus 7,33 CiCONIIFORMES Ardeidae American Bittern. Botaurus lentiginosus Least Bittern. Ixobrychus exilis Great Blue Heron. Ardea herodias Great Egret. Casmerodius albus Snowy Egret. Egretta thula Cattle Egret. Bubulcus ibis Green-backed Heron. Butorides straitus Black-crowned Night-Heron. Nycticorax nycticorax Threskiomithidae Wliite Ibis. Eudociinus albus White-faced Ibis. Plegadis chihi Ciconiidae Wood Stork. Mijcteria americana Anseriformes Anatidae Fulvous Whistling-Duck. Dendrocygna bicolor Tundra Swan. Cygnus columbianus Tmmpeter Swan. C. buccinator Greater White-fronted Goose. Anser albifrons Snow Goose. Chen caerulescens Ross' Goose. C. rossii Canada Goose. Branta canadensis Wood Duck. Aix sponsa 33 w M6 w R2 w 34, M3, W5 w 33 w V5 w, f V4 w R3 w, d V6 w 33 w V6 d, w M2, W4 w V5 w M3 w M3 w, f M4 w Rl w, f B4, M3, W4 w, d USFWS AL, IDF&G, USBLM, USFWS 7, WB 7, DJ 7, JFD, DT, USFWS 'See text for Abundance Code. 'Breeding and Seasonal Use Code: R = Breeder and year-round resident. B = Summer breeder M = Migrant W = Winter visitor S = Summer visitor; no breeding records V = Erratic visitor; no breeding records ^Habitat Code. (Multiple habitats listed for a particular species are given in descending order of occurrence.): w = on or near water or marsh g = grassland or steppe j = juniper woodland d = deciduous woodland or riparian c = coniferous woodland a = alpine f = agricultural 730 Great Basin Naturalist Vol. 43, No. 4 Table 1 continued. Taxa Abundance', season and breeding status^ Preferred habitat' References Green-winged Teal. Anas crecca American Black Duck. A. rubripes Mallard. A. platyrhynchos Northern Pintail. A. acuta Blue-winged Teal. A. discors Cinnamon Teal. A. cyanoptera Northern Shoveler. A. clypeata Gadwall. A. strepera Eurasian Wigeon. A. penelope American Wigeon. A. americana Canvasback. Aythya valisineria Redhead. A. americana Ring-necked Duck. A. collaris Greater Scaup. A. marila Lesser Scaup. A. affinis Harlequin Duck. Histrionicus histrionicus Oldsquaw. Clangula hyemalis White-winged Scoter. Melanitta fiisca Common Goldeneye. Bitcephala clangula Barrow's Goldeneye. B. islandica Bufflehead. B. albeola Hooded Merganser. Lophodytes cucullatus Common Merganser. Mergus merganser Red-breasted Merganser. M. serrator Ruddy Duck. Oxyura pmaicensis Falconiformes Cathartidae Black Vulture. Coragyps atratus Turkey Vulture. Cathartes aura Accipitridae Osprey. Pandion haliaetus Black-shouldered Kite. Elanus caeruleus Bald Eagle. Haliaeetus leucocephahis Northern Harrier. Circus cyaneus Sharp-shinned Hawk. Accipiter striatus Cooper's Hawk. A. cooperii Northern Goshawk. A. gentilis Broad-winged Hawk. Buteo platypterus Swainson's Hawk. B. sivainsoni Red-tailed Hawk. B. jamaicensis Ferruginous Hawk. B. regalis Rough-legged Hawk. B. lagopus Golden Eagle. Aquila chrysaetos Falconidae American Kestrel. Falco sparverius Merlin. F. colwnbarius Peregrine Falcon. F. peregrinus Gyrfalcon. F. rusticolus Prairie Falcon. F. mexicaniis B3, Ml, W3 w 7 w THR Rl w, f B3, Ml, W,3 w B4, M3 w B2, M2, W6 w B4, M2, W4 w R3 w V5 w 7, 8. TDR, DT B4, Ml, W2 w B4, M2, W4 w B2, M2, W4 w B4, M3, W4 w M6, W5 w MC, LR, DT B4, M2, W3 w S6, M6 w 7, 39, MC, BH, DJ, AL M6, W5 w 7, 26, 42, RK, AL, IDF&G V6 w 7, 32, USFWS M2, W2 w M4, W3 w M2, W3 w M4, W4 w B3, M2, W2 w M4, W5 w B3, M2, W4 w 7 THR B3, W6 g'C, j.f B3, M3, W6 w, c V6 g 35 B5, M3, W3 w, c, ,d R2 W, g: .f-j R3 g,d, f,j,c R3 &h d, c R3 c,d, j V6 g 6, 7, MC B3 &d. f R2 &d. c,j,f B3, W6 g.j W2 gJ R3 g.j. d R2 g.f- d,j R4 d,g, f, c B?5, M5, W5 g'W , c, a 7, 14, JFD, TDR, CT, IDF&G USFWS, USBLM W6 f^g. w 7,JD B2, W3 gj' a, c Galliformes Phasianidae "Gray Partridge. Perdix perdix "Chukar. Alectoris chukar "Ring-necked Pheasant. Phasianus colchicus R3 R2 Rl g,f g'j f>g October 1983 Stephens, Reynolds: Idaho Birds 731 Table 1 continued. Taxa Abundance', season Preferred and breeding status- habitat' References R4 c R3 c R6 a, c 13 R3 c, d R3 K Spnice Grouse. Dendragapiis canadensis Blue Grouse. D. ol)sctirus White-tailed Ptarmigan. Lagopus leucurus Ruffed Grouse. Bonasa umbellus Sage Grouse. Centrocercus urophasianus Sharp-tailed Grouse. Tijmpanuchus phasianeUus "Wild Turkey. Meleagris gallopavo "Northern Bobwhite. Colinus virginianus "Gambel's Quail. Callipepla gambelii "California Quail. C. californica Mountain Quail. Oreortyx picttis Gruiformes Rallidae Yellow Rail. Coturnicops noveboracensis Black Rail. Laterallus jamaicensis Virginia Rail. RaUtis Umicola Sora. Porzana Carolina Common Moorhen. Gallintila chloropus American Coot. Fiilica americana Gniidae Sandhill Crane. Grus canadensis Charadriiformes Charadriidae Black-bellied Plover. Phivialis squatarola Snowy Plover. Charadrins alexandriniis Semipalmated Plover. C. semipahnaUis Killdeer. C. vociferiis Mountain Plover. C. montanus Haematopodidae American Oystercatcher. Haematopus palliatus Recurvirostridae Black-necked Stilt. Himantopus mexicanus American Avocet. Rectirvirostra americana Scolopacidae Greater Yellowlegs. Tringa melanoleuca Lesser Yellowlegs. T. flavipes Solitary Sandpiper. T. solitaria Willet. Catoptrophorus semipalnmtus Spotted Sandpiper. Actitis macidaria Upland Sandpiper. Bartramia longicauda Long-billed Curlew. Numenins americanus Marbled Godwit. Limosa fedoa Ruddy Turnstone. Arenaria interpres Red Knot. Calidris canutus Sanderling. C. alba Semipalmated Sandpiper. C. pusilla Western Sandpiper. C. mauri Least Sandpiper. C. minutilla R4 R4 R4 R5 R2 R4 7 7 B3, W6 B3, W5 V6 B2, Ml, W2 B4, M2 M5 V5 M3 Bl, Ml, W3 V6 B3 B3, M2 M3 M3 M4 B3, M3 B2, M3 B4, M4 B2 M4 7 M5 M4 M3 M2 M2 d, c f.g-d g.d 2, 31, MC, LR g,d,f d, c, f w 28 w AL g. w 36 7, 14, 28, MC, DAS, IDF&G, USFWS 13, JD, LR IDF&G 38, DAS 13 13, MC, RK, AL, JM, TDR, DAS 732 Great Basin Naturalist Vol. 43, No. 4 Table 1 continued. Taxa Abundance', season and breeding status- Preferred habitat' References Baird's Sandpiper. C. bairdii M3 Pectoral Sandpiper. C. mehinotos M5 Dunlin. C. alpina M5 Stilt Sandpiper. C. hiniantopiis M4 Buff-breasted Sandpiper. Tryngites M6 siibruficoUis Long-billed Dowitcher. Limnodronius M.3 .icolopaceus Common Snipe. Callinago gaUinago B2, W4 Wilson's Phalarope. Phalawpits tricolor 83, M2 Red-necked Phalarope. P. lobatus M3 Laridae Pomarine Jaeger. Stercorarius 7 pomarinus Franklin's Gull. Lurtts pipixcan Bonaparte's Gull. L. Philadelphia Ring-billed Gull. L. delawarensis California Gull. L. californiciis Herring Gull. L. argentatus Thayer's Gull. L. thayeri Glaucous-winged Gull. L. glaucescens Glaucous Gull. L. hyperboreiis Sabine's Gull. Xema sabini Caspian Tern. Sterna caspia Common Tern. S. hirundo Forster's Tern. S. forsteri Least Tern. S. antillarum Black Tern. Chlidonias niger Alcidae Ancient Murrelet. Synthliboramphus V6 antiquus COLUMBIFORMES Cohnnbidae °Rock Dove. Coliimba livia Rl d, f, j, c Band-tailed Pigeon. C. fasciata V4 d, g, j, c Mourning Dove. Zenaida macroura BL ML W3 g, f, d, j, c CUCULIFORMES Cuculidae Black-billed Cuckoo. Coccyzus B4 d erythropthalmiis Yellow-billed Cuckoo. C. americanus B5 d Strigiformes Tytonidae Common Barn-Owl. Tyto alba R2 Strigidae Flammulated Owl. Otiis flammeolus B5 Western Screech-Owl. O. kennicottii R3 Great Horned Owl. Bubo virginianus R2 Snowy Owl. Nyctea scandiaca W5 Northern Hawk-Owl. Siirnia alula 7 Northern Pygmy-Owl. Glaucidium R3 gnoma Burrowing Owl. Athene cunicularia B2 Barred Owl. Strix varia 7 Great Gray Owl. S. nebulosa R4 Long-eared Owl. Asio otus R3 Short-eared Owl. A. flammeus R2 Boreal Owl. Aegolius funereus R5 Northern Saw-whet Owl. A. acadicus R4 g.f.j-d 2,7, 15, AL,JM, IDF&G 2, JB, JM, DAS, TDR, USFWS 15 7,24 B?4, M3 w M4 w Rl f, w Bl, W4 f, w M4, W4 f, w 7 w MC 7 f 27 W6 w 4, USFWS V6 w 7, USFWS B,3, M4 w M5 w 7, USFWS B4, M3 w V6 w 5 B4, M3 w 13 7, 13, 28, AL, USFWS c, j, d 7, 18, LP, TR d, f g. j. d, c, f g 2, 7, 22, DAS 7 & t, d &f LP, USBLM c d, f w, f MC, DAS d, j, c October 1983 Stephens, Reynolds: Idaho Birds 733 Table 1 continued. Taxa Abundance', season and breeding status- Preferred habitat' References Caprimuu;iformes Caprimulgidae Lesser Nighthawk. Chordeiles acutipennis Common Nighthawk. C. minor B2 Common Poorwill. Philaenoptihis B3 nuttallii Apodiformes Apodidae Black Swift. Cypseloides niger 7 Vaux's Swift. Chaetura vaiixi B3 White-throated Swift. Aenmatttes B3 saxdtalis Trochilidae Black-chinned Hummingbird. B3, W6 Archilochus alexandri Anna's Hummingbird. Cahjpte anna M5, , W5 Calliope Hummingbird. Stelluki B2 calliope Broad-tailed Hummingbird. Selasplwnis B4 pUitijcercus Rufous Hummingbird. S. ritfus B3, M4 CORACIIFORMES Alcedinidae Belted Kingfisher. Cenjle alcijon PiCIFORMES Picidae Lewis' Woodpecker. Melanerpes lewis Red-headed Woodpecker. M. erijthrocephalus Yellow-bellied Sapsucker. Sphyrpicits variiis Williamson's Sapsucker. S. thyroicleiis Downy Woodpecker. Picoides pubescens Hairy Woodpecker. P. villosus White-headed Woodpecker. P. albolawattts Three-toed Woodpecker. P. tridactylus Black-backed Woodpecker. P. arctictis Northern Flicker. Colaptes auratus Pileated Woodpecker. Dryocopus pileatus Passeriformes Tyrannidae Olive-sided Flycatcher. Contopus borealis Western Wood-Pewee. C. sordiduhis Willow Flycatcher. Empidonax traillii Least Flycatcher. E. minimus Hammond's Flycatcher. E. hammondii Dusky Flycatcher. E. oberholseri Gray Flycatcher. E. wrightii Western Flycatcher. E. difficilis Black Phoebe. Sayornis nigricans Say's Phoebe. S. saya R2 B2, W4 V6 B3 B4 R2 R3 R4 R4 R5 R2 R3 B.3, M3 THR g, d, j, f, c g. J. ^ w, c. d w, g, c. a c,d d c, d d,] c, d BH, AL, LM, TDR, DAS w, d c, d d c, d d, c d, c c c d, c, g, f c, d 7, 25, MC 2, 7, 16, 17, DJ B3, M3 c, d B2 d 7 d MC B2 c B3 d, c, j B3 g'j B5 d, c 7, 46, TR, IDF&G, USFWS, USBLM 7 7 B2 g.d 734 Great Basin Naturalist Vol. 43, No. 4 Table 1 continued. Taxa Abundance' and breedin , season g status- Preferred habitat' References B3 B2 B3 V6 J. g d,g,f g 40 Ash-throated Flycatcher. Mijiarchus cinerascens Western Kingbird. Tyrannus verticalis Eastern Kingbird. T. tyrannus Scissor-tailed Flycatcher. T. forficatus Alaudidae Homed Lark. Eremophilia alpestris Hirundinidae Purple Martin. Progne subis Tree Swallow. Tachycineta bicolor Violet-green Swallow. T. thahissina Northern Rough-winged Swallow. Stelgklopteryx senipennis Bank Swallow. Riparia ripariu Cliff Swallow. Hirundo pyrrhonuta Barn Swallow. H. nistica Corvidae Gray Jay. Perisoreus canadensis Steller's Jay. Cyanocitta stelleri Blue Jay. C. cristata Scrub Jay. Aphelocoma coerulescens Pinyon Jay. Gymnorhinus cyanocephalus Clark's nutcracker. Nucifraga Columbiana Black-billed Magpie. Pica pica American Crow. Corvus brachyrhynchos Common Raven. C. corax Paridae Black-capped Chickadee. Parus atricapillus Mountain Chickadee. P. gambeli Chestnut-backed Chickadee. P. rufescens Plain Titmouse. P. inornatus Aegithalidae Bushtit. Psaltriparus minimus Sittidae Red-breasted Nuthatch. Sitta canadensis White-breasted Nuthatch. S. carolinensis Pygmy Nuthatch. S. pygmaea Certhiidae Brown Creeper. Certhia americana Troglodytidae Rock Wren. Salpinctes obsoletus Canyon Wren. Catherpes mexicanus Bewick's Wren. Thryomanes bewickii House Wren. Troglodytes aedon Winter Wren. T. troglodytes Marsh Wren. Cistothorus palustris Cinclidae American Dipper. Cinclus mexicanus Muscicapidae Golden-crowned Kinglet. Regulus satrapa Ruby-crowned Kinglet. R. calendula Rl M6 w, d B2, M 1 c, d, w B2, Ml c, d, w B2, M2 w, c, d B2, M2 w, f, g Bl, Ml g, w, c, f Bl, Ml f. w, g R3 c R2 c, d V4 d R4 j>g R4 J. g R2 Rl Rl R2 R2 R2 7 R5 R3 R2 R3 R4 R3 B2, W5 B3, W4 7 B2 R3 B2, W3 R3 B4, W3 R2 c, a g. f> d, j d, f, g, j, c d, g, j, c, a c, d d,j c, d c c, d g-j d d cd MC, LP 39 2, 7, 13, 15, USBLM MC c, d c, d October 1983 Stephens, Reynolds: Idaho Birds 735 Table 1 continued. Taxa Abundance', season and breeding status- Preferred habitat' References Blue-gray Gnatcatcher. Polioptila B4 caenilea Western Bluebird. SUilia mexiccma Mountain Bluebird. S. currucoides Town.sends Solitaire. Mifadestes townsendi Veery. Catharus fttscescens Swainson's Thnush. C. ttstuhitus Hermit Thnish. C. gitttatus Wood Thni.sh. Hijlocichki mustelinu American Robin. Turdus migratorius Varied Thnish. Ixoreiis naeviiis Mimidae Gray Catbird. Dunietella carolinensis Northern Mockingbird. Mimus pohjglottos Sage Thrasher. Oreoscoptes inontanus Motacillidae , Water Pipit. Anthtis spinoletta Bombycillidae Bohemian Waxwing. Bombijcilla garrulus Cedar Waxwing. B. cedrorum Laniidae Northern Shrike. Laniiis excitbitor Loggerhead Shrike. L. Itidovicianus Stumidae "European Starling. Sturntis vtdgaris Rl Vireonidae Solitary Vireo. Vireo solitarius B3 Warbling Vireo. V. gilvus B2 Red-eyed Vireo. V. olivaceus B4 Emberizidae Golden-winged Warbler. Vennivora V6 chrijsoptera Tennessee Warbler. V. peregrina M5 Orange-crowned Warbler. V. celata B3, W5 Na.shville Warbler. V. ruficapUla B3 Virginia's Warbler. V. virginiae V5 Yellow Warbler. Dendroica petechia B2 Magnolia Warbler. D. magnolia 7 Yellow-mmped Warbler. D. coronata B2, W4 Black-throated Gray Warbler. D. B3 nigrescens Townsend's Warbler. D. townsendi B4, M3 Blackbumian Warbler. D. fusca 7 Blackpoll Warbler. D. striata 7 Black-and-white Warbler. Mniotilta M6 varia American Redstart. Setophaga ruticiUa B4, M3 Ovenbird. Seiurus atirocapiUiis M6 Northern Waterthrush. S. B5, M5 noveboracensis MacGillivray's Warbler. Oporornis B3 tolmiei Common Yellowthroat. Geothh/pis B3 trichas Wilson's Warbler. Wihonia ptisilla B3, M3 Yellow-breasted Chat. Icteria virens B2 Western Tanager. Piranga ludoviciana B2, M2, W5 d,j B3 c B2, Ml, W5 c, d, j R3 j, c, d, a B3 d B2, M3 c, d B.3, M3, W5 c, d 7 Rl c, d, f, g, j B4, M4, W5 c, d B3 d V4 g.d B2 g B?4, M2, W4 g, w, a, f W2 c, d, j, f B3, W2 c, d, j, f W3 g.M.j B3, W4 &M.j f, d, g, c, j c, d d d d d d, c j.d d c, d c, d i c, d d d c, d d d, c, w d d c, d TR 7, JB, MC, LM 7, 14, MC 7 7 MC 7, 13, 39, USFWS JD, JM 7, JB, MC, DAS, DT 736 Great Basin Naturalist Vol. 43, No. 4 Table 1 continued. Taxa Abundance', season and breeding status- Preferred habitat' References Rose-breasted Gosbeak. Pheucticiis ludovicianus Black-headed Grosbeak. P. mekinocephalus Blue Grosbeak. Cuiraca caenilea Lazuli Bunting. Passerina ainoena Indigo Bunting. P. cijanea Green-tailed Towhee. Pipilo chlorurus Rufous-sided Towhee. P. enjthrophthalmus American Tree Sparrow. Spizella arborea Chipping Sparrow. S. passerina Clay-colored Sparrow. S. pallida Brewer's Sparrow. S. breiveri Vesper Sparrow. Pooecetes graminetis Lark Sparrow. Chondestes grammacus Black-throated Sparrow. Amphispiza bilineata Sage Sparrow. A. belli Lark Bvniting. Calamospiza melanocorijs Savannah Sparrow. Passerculus sandwichensis Baird's Sparrow. Amniodramus bairdii Grasshopper Sparrow. A. savannanim Fox Sparrow. Passerella iliaca Song Sparrow. Melospiza melodia Lincoln's Sparrow. M. Uncolnii Swamp Sparrow. M. georgiana White-throated Sparrow. Zonotrichia albicollis Golden-crowned Sparrow. Z. atricapilla White-crowned Sparrow. Z. leucophnjs Harris' Sparrow. Z. qtienda Dark-eyed Junco. Junco Injemalis Lapland Longspur. Calcarius lapponicus Snow Bunting. Plectrophenax nivalis Bobolink. Dolichonyx onjzicorus Red-winged Blackbird. Agelaius phoenicexis Tricolored Blackbird. A. tricolor Western Meadowlark. Sttirnella neglecta Yellow-headed Blackbird. Xanthocephalus xanthocephalus Brewer's Blackbird. Euphagus cijanocephalus Common Crackle. Quiscalus quiscula Bronzed Cowbird. Molothrtts aeneus Brown-headed Cowbird. M. ater Orchard Oriole. Icterus spiirius Northern Oriole. /. galbula Fringillidae Rosy Finch. Leiicosticte arctoa Pine Grosbeak. Pinicola eniicleator Purple Finch. Carpodacus piirpiiretis Cassin's Finch. C. cassinii House Finch. C. mexicanus Red Crossbill. Loxia citrvirostra V6 B3 d d 7, RK, AL, JM B4 d-g B2, M2 c, d V6 d 7, .30, MC, US] B3 j'd, g B2, M3, W4 d, g. j W3 B2, M2, VV5 c, d, j 7 &d MC Bl g B2 g't' B2 g.j B4 g B3 g V6 g 7,35 B2, W5 & w 7 g 7,24 B5, M5 g 7, 28, MC, DAS, USBLM B4, M4, W5 c, d Rl w, d B3, W5 d, a W6 d 37 M5, W5 g 7, 41, MC, AL, LM, DT M5, W6 g.d 7, 9, 10, 13, 26, 29, MC, DT Rl g, a, f, d, c W4 &d Rl d, c, g W5 g.c 2, 7, DJ, DT W3 g B4, M4 g.d.f Rl w, f, g, d 7 w 7, LM Bl, W2 g.f Bl, W4 w, f Rl g, f, j, c, d V6 d MC, GEAS 7 7 B2, W6 &f 7 7 B2 d R3 a, g R3 c V6 d, c 3,7 R2, M3 c, d R2 d, c, f R2 c October 1983 Stephens, Reynolds: Idaho Birds 737 Table 1 continued. Taxa Abundance', season and breeding status- Preferred habitat' References White-winged Crossbill. L. leucoptera Common Redpoll. Cardiielis flammea Hoary Redpoll. C. horneincinni Pine Siskin. C. pinus Lesser Goldfinch. C. psaltria American Goldfinch. C. tristis Evening Gro.sbeak. Coccothraiistes vespertinus Passeridae "House Sparrow. Passer domesticus W6 c 7, 11, 12 W4 &d 7 27 B2, M2, W3 c, d B?5 d 7, 34, DAS R2 d,g,f R2 d, c Rl &d,f This information was first compiled as a pre- liminary check-list which was then sent to 30 local birders and other ornithologists with birding expertise and experience in south- western Idaho for review and modification. The cover letter accompanying each draft asked reviewers to assess the abundance for each species listed, and to include personal observation data (date, location, habitat, and number of individuals observed) for all spe- cies tentatively classified as occasional, rare, accidental, vagrant, or hypothetical. The abundance code, though qualitative by neces- sity, had a quantitative basis. Assuming a competent birder rigorously searched the proper habitat at the correct time of year, abundance categories were: 1. Abimdant— more than 40 individuals per day 2. Common— 10-39 individuals per dav 3. Uncommon— less than 10 individuals per day 4. Occasional— 1-5 total records per year 5. Rare— 3-10 total records for southwestern Idaho 6. Vagrants— accidental species for which there are 2 or fewer reliable records supported by a photograph, specimen, or detailed and accurate field notes 7. Hypothetical species— unverified sight records not .supported by documentation Results Table 1 presents the results of our in- vestigations. The sequence of presentation and nomenclature follows the 34th supple- ment to the American Ornithologists' Union Check-list of North American birds (1982). The authority for the rare, vagrant, and hy- pothetical species are given where known. Initials correspond to contributors credited below or the authors (DAS, TDR); numbers correspond to literature accounts cited at the end of the paper. Acknowledgments The following persons, organizations, and agencies were extremely helpful in donating their time, records, and expertise in helping generate this list of birds of southwestern Idaho: J. Bamett, W. Belknap, J. Carson, M. Collie, J. F. Dixon, J. Doremus, Golden Eagle Audubon Society, B. Hammond, D. Jones, R. Kuntz, A. Larson, H. Larson, J. Marks. L. Mohler, L. Powers, T. Rich, T. H. Rogers, L. Reichert, B. Sturges, D. Taylor, C. Trost, E. Yensen, Idaho Department of Fish and Game, U.S. Bureau of Land Management, U.S. Forest Service, and the U.S. Fish and Wildlife Service. We sincerely thank each contributor for their valuable input. We gratefully acknowledge the skills of B. Donahue, C. Levesque, and M. Reynolds in typing and proofreading the manuscript. Literature Cited 1. AOU Committee on Classification and Nomenclature. 1982. Thirty-fourth supplement to the American Ornithologists' Union Check- List of North American Birds. Supplement to Auk99:l-16CC. 2. Burleigh, T. D. 1971. Birds of Idaho. Caxton Print- ers, Caldwell, Idaho. 3. Cruickshank, a. D., ed. 1972. Christmas bird count. Am. Birds 26:476-477. 4. 1970. Christmas bird count, .'\udubon Field Notes 24:412. 5. Davis, W. B. 1934. Bird notes from Owyhee Co., Idaho. Murrelet 15:69-72. .6 19.36. Broad-winged Hawk in Idaho. Condor 38:86. 7. Heckathorn, J. 1978. Birds of southwest Idaho. Golden Eagle Audubon Society, Boise, Idaho. 8. Heilbrun, L. H., and H. Downing, eds. 1981. Christmas bird count: Montana, Wvoming, Colo- rado, Idaho, Utah. Am. Birds 35:375-376, 673. 9. 1980. The eightieth Audubon Christmas bird count. Am. Birds 34:615. 738 Great Basin Naturalist Vol. 43, No. 4 10. Heilbrun, L. H., and P. R. Julian, eds. 1976. The seventy-sixth Audubon Christmas bird count. Am. Bi'rds. 30: 176-552. 11. Jewett, S. G. 1912. Some birds of the Saw-tooth Mountain, Idaho. Condor 14:191-194. 12. KiNGERY, H. E., ed. 1972. The fall migration, Great Basin Central Rocky Mountain Region. Am. Birds 26:99. 13. Larrison, E. J., J. L. Tucker, and M. T. Jollie. 1967. Guide to Idaho birds. J. Idaho Acad. Sci. 5:1-220. 14. Levy, S. H. 1950. Summer birds in southern Idaho. Murrelet 31:2-8. 15. 1962. Additional summer southern Idaho bird notes. Murrelet 43:10-14. 16. Marshall, W. H. 1945. Winter bird observations in the Boise National Forest, Idaho. Condor 47:170-172. 17. Merriam, C. H. 1891. Results of a biological recon- naissance of south central Idaho. N. Amer. Fauna 5:1-108. 18. 1892. The Dwarf Screech Owl {Megascops flainneolus idahoensis Merriam). Auk 9:169-171. 19. Newhouse, V. F. 1960. Birds of selected irrigated river valleys of west central Idaho. Murrelet 41:1-6. 20. Powers, L. R. 1969. Sight record of the Blue Gros- beak in Idaho. Murrelet 50:20-21. 21. Powers, L. R., and C. H. Trost. 1977. Early re- cords of avifauna from a 1907 scientific expedi- tion into Idaho. Murrelet .58:87-88. 22. Rogers, T. H., ed. 1975. The winter season North- em Rocky Mountain-Intermountain Region. Am. Birds 29:718. 23. 1976. The fall migration Northern Rocky Mountain-Intermountain Region. Am. Birds 30:98. 24. 1977. The nesting season Northern Rocky Mountain-Intermountain Region. Am. Birds 31:201-203. 25. 1977. The winter season Northern Rocky Mountain-Intermountain Region. Am. Birds 31:354. 26. 1978. The fall migration Northern Rocky Mountain-Intermountain Region. Am. Birds .32:233-235. 27. 1978. The winter season Northern Rocky Mountain-Intermountain Region. Am. Birds 32:378-379. 28. 1978. The spring migration Northern Rocky Mountain-Intermountain Region. Am. Birds 32:10.34-10.35. 29. 1979. The winter season Northern Rocky Mountain-Intermountain Region. Am. Birds .33:.300. .30. 1979. The spring migration Northern Rocky 31. .32. 33. 34. .35. 36. 37. .38. Mountain-Intermountain Region. Am. Birds .33:792. 1979. The nesting season Northern Rocky Mountain-Intermountain Region. Am. Birds 33:882. 1980. The spring migration Northern Rocky Mountain-Intermountain Region. Am. Birds .34:182-183. 1980. The winter season Northern Rocky Mountain-Intermountain Region. Am. Birds 34:291. 1980. The spring migration Northern Rocky Mountain-Intermountain Region. Am. Birds 34:800. 1980. The nesting season Northern Rocky Mountain-Intermountain Region. Am. Birds 34:912-914. 1981. The autumn migration Northern Rocky Mountain-Intermountain Region. Am. Birds 35:205-206. 1981. The winter season Northern Rocky Mountain-Intermountain Region. Am. Birds 35:321. 1981. The spring migration Northern Rocky Mountain-Intermountain Region. Am. Birds ,35:844. .39. Seidensticker, J. C, and R. E. Welch, compilers. 1972. The birds of Big Creek and Chamberlain Basin-Idaho Primitive Area. USDA, Forest Ser- vice, Payette National Forest, McCall, Idaho. 40. Thornburg, F. 1956. Scissor-tailed flycatcher in Idaho. Condor 58:72-73. 41. Wyman, L. E. I9I2. White-throated Sparrow in Idaho. Auk 29:247. 42. YocuM, C. F. 1950. Sight record of Old-squaw in Idaho. Murrelet 31:47. NEW RECORDS FOR THE VASCULAR FLORA OF WYOMING AND MONTANA Robert W. Lichvar', Robert D. Dorn', and Erwin F. Evert' Abstract.— Apparent first state records of 11 taxa for Wyoming and one taxon for Montana are listed. A range ex- tension is noted for the genus Shoshonea. Recent field work from 1980 and 1982 has added one additional record to the flora of Montana and 11 additional records for Wyoming. The recently described genus and species Shoshonea pulvinata Evert and Con- stance, thought to have been a highly re- stricted endemic, is reported here well out- side its previously known rangp. Apiaceae Shoshonea pulvinata Evert and Constance, WY, Fremont Co., T6N R5E S34, 2286 m (7500 ft), calcareous ridge, 26 July 1982, R. Lichvar 5382 RM. This recently described genus and species by Evert and Constance (1982) was known only from a restricted range near Cody, Wyoming. This collection, Lichvar 5382, is a distance of 55.8 km (90 mi) southeast of the Cody sites. ASTERACEAE Adenocaulon bicolor Hook., WY, Crook Co., T51N R60W S20, 1646 m (5400 ft), wet ravine, 28 July 1982, R. Dorn 3798, COLO, NY, RM. Same county, T51N R61W Sll SEi/4, 1646 m (5400 ft), wet ravine, 12 Sept. 1982, E. Evert 4968, RM. Brassicaceae Lepidium sativum L., WY, Laramie Co., T14N R67W S28 SEV4SW1/4, 1890 m (6200 ft), disturbed area, 21 Aug. 1982, R. Dorn 3824, RM. Caryophyllaceae Gypsophila scorzonerifolia Ser., WY, La- ramie Co., T14N R67W S17 N1/4, 1890 m (6200 ft), disturbed roadside, 5 Sept. 1982, R. Dorn 3825, RM. Cyperaceae Carex alopecoidea Tuckerm., WY, Crook Co., T51N R60W S33 SWV4, 1798 m (5900 ft), wet meadow, 27 July 1982, R. Dorn 3783, RM. C. deweyana Schw. ssp. deweyana, WY, Crook Co., T51N R60W S33 SW1/4, 1798 m (5900 ft), mossy bank, 27 July 1982, R. Dorn 3785, NY, RM. C. peckii Howe, WY, Crook Co., T51N R60W S20, 1646 m (5400 ft) wet ravine, 28 July 1982, R. Dorn 3796, RM. C. rosea Schk. ex Willd., WY, Crook Co., T51N R60W S33 SW1/4, 1798 m (5900 ft), wet meadow, 27 July 1982, R. Dorn 3781, RM. Fabaceae Astragalus coltonii Jones var. moahensis Barneby, WY, Uinta Co., T14N R113W S31, 2195 m (7200 ft), sagebrush, 4 June 1980, R. Dorn 3447, RM. Same location and date, R. Lichvar 2780, RM. T13N, 113W, S18 SE1/4, 5 mi N 25° W of Lonetree, E side Hickey Mtn., S. Goodrich, D. Atwood 17169, 17193, 30 June 1983, BRY. T14N R113W S27, 18 mi N 12° E of Lonetree, N of Sage Cr. Mtn., S. Goodrich, D. Atwood, 17212, 1 July 1982, 'Wyoming Natural Heritage Program/The Nature Conservancy, 1603 Capitol Avenue, Room 325, Cheyenne, Wyoming 82001. 'P.O. Box 1471, Cheyenne, Wyoming 82003. '1476 Tyrell, Park Ridge, Illinois 60068. 739 740 Great Basin Naturalist Vol. 43, No. 4 BRY. Sweetwater Co., Cedar Mtn., T13N RlllW S3 NEV4, 2195 m (7200 ft), sage- brush, 11 June 1980, R. Lichvar 2852, KM. Same location, 8 July 1981, R. Lichvar 4580, RM. This species was discussed by Barneby (1964) as possibly being collected in Wyo- ming by R. Rollins, no. 177, in 1932. This specimen was distributed by RM as Hedysa- rum sp. The authors were unable to relocate this specimen at RM; it is presumed filed un- der an Astragalus. Barneby felt that this spec- imen was probably mislabeled. These collec- tions confirm its existence in Wyoming. Onagraceae Circaea lutetiana L., WY, Crook Co., T52N R60W S21 EV2, 1280 m (4200 ft), stream bank, 26 July 1982, R. Dorn 3773, COLO, NY, RM. POLEMONIACEAE Leptodactylon caespitosum Nutt., MT, Carbon Co., T9S R27E S28 and 29 line, 1463 m (4800 ft), barren red slope, 24 June 1982, R. Dorn 3728, COLO, NY, RM. POLYGONACEAE Polygonum scandens L., WY, Crook Co., T54N R63W Sll NEi/4, 1432 m (4700 ft), edge of beaver pond, 25 July 1982, R. Dorn 3762, RM. Same location and date, R. Lich- var 5375, RM. SCROPHULARIACEAE Veronica arvensis L., WY, Crook Co., T54N R63W SI SW1/4, 1463 m (4800 ft), dis- turbed area in Ponderosa Pine, 25 July 1982, R. Dorn 3761, NY, RM. Same location and date, R. Lichvar 5336, RM. Same location and date, E. Evert 4438, RM. Literature Cited Barneby, R. 1964. Atlas of North American Astragalus. Memoirs of the New York Botanical Garden, vol. 13. Evert, E., and L. Constance. 1982. Shosl^onea pulvi- nata, a new genus and species of Umbelliferae from Wyoming. Systematic Botany 7(4):471-475. RANGE EXTENSIONS FOR TWO DWARF MISTLETOES {ARCEUTHOBIUM SPP.) IN THE SOUTHWEST Robert L. Mathiasen' and Kenneth H. Jones' Abstract.— A second small population of white fir dwarf mistletoe, Arcetithobimn ahietiniim f. sp. concoloris, is reported from the Santa Catalina Mountains in southern Arizona. The discovery of a second population of this mis- tletoe in southern Arizona supports the contention that it once had a more southern distribution but has survived past climatic changes in only a few locations in the southwestern United States. The distribution of the Western spnice dwarf mistletoe, Arceuthohium micwcarptim, is extended to the Sacramento Mountains of south central New Mexico, a range extension of approximately 170 miles. The mistletoe is restricted to an area of about 300 acres, but its potential for further spread is high. The possible implications of this range extension to the biosystematics of Picea spp. and the evolution of A. microcarptim are discussed. White fir dwarf mistletoe, Arceuthobium abietinum Engelm. ex Munz f. sp. concoloris Hawksw. & Wiens, is a serious pathogen of white fir, Abies concolor (Gord. & Glend.) Lindl., in the western United States (Scharpf 1964, Scharpf and Parmeter 1967). Hawks- worth and Wiens (1972) hst the distribution of this dwarf mistletoe from southern Wash- ington southward through the Cascade Range and Sierra Nevada to the San Bernardino Mountains in southern California, with four isolated populations known: the Charleston Mountains and Spring Creek Mountains, Ne- vada; Kane County in southwestern Utah; and Grand Canyon National Park, Arizona. More recently it has been reported from the Chiricahua Mountains of southeastern Ari- zona, nearly 300 miles south of the Grand Canyon populations (Mathiasen 1976). Western spruce dwarf mistletoe, Arceutho- bium microcarpum (Engelm.) Hawksw. & Wiens, severely parasitizes Engelmann and blue spruce, Picea engelmannii Parry and P. pungens Engelm., in the southwestern United States (Hawksworth and Graham 1963, Hawksworth and Wiens 1972). In Arizona it is known from the Kaibab Plateau, the San Francisco Peaks and Kendrick Peak, the White Mountains, and the Pinaleno Moun- tains (Hawksworth and Wiens 1972). In New Mexico it has only been reported from the Mogollon Mountains in the west central part of the state (Hawksworth and Wiens 1972). This paper reports isolated populations of A. abietinum from the Santa Catalina Moun- tains in south central Arizona and of A. mi- crocarpum from the Sacramento Mountains in south central New Mexico. In 1979 a very small population of A. abietinum was discovered parasitizing white fir in the Santa Catalina Mountains of Pima County, Arizona (Fig. 1). The population oc- curs in an area of about 10 acres in Marshall Gulch (T. 12 S., R. 16 E., Section 6) near the center of the mountain range at elevations of 2,310 to 2,340 m. The predominant tree spe- cies in the infested stand are white fir, Douglas fir (Pseudotsuga menziesii [Mirb.] Franco), and southwestern white pine (Pinus strobiformis Engelm.). Douglas fir dwarf mis- tletoe {Arceuthobium douglasii Engelm.) also occurs in the stand on Douglas fir. Douglas fir dwarf mistletoe has rarely been collected on white fir and can easily be distinguished from white fir dwarf mistletoe using shoot morphology (Hawksworth and Wiens 1972). This is the second report of a small, isolated population of A. abietinum from southern Arizona. The first was reported in the Chiri- cahua Mountains, approximately 100 miles southeast of the Santa Catalina population (Mathiasen 1976). The Santa Catalina popu- lation of A. abietinum is nearly 260 miles south of the Grand Canyon population of this taxon, and no other populations of A. abieti- num are known between these localities (Mathiasen 1976). 'School of Forestry, Northern Arizona University, Flagstaff, .\rizona 86011. 'Sacramento Mountain Laboratory, New Mexico State University, Siinspot, New Mexico 88349. 741 742 Great Basin Naturalist Vol. 43, No. 4 Fig. 1. Distribution of Abies concolor and Arcetithobiiim abietiniim (1-4) in Arizona and New Mexico. Numbers 3 and 4 represent the A. abietinitm populations in the Santa Catalina and Chiricahua mountains, respectively. The discovery of a second population of A. abietinum in southern Arizona supports the suggestion that A. abietinum had a more widespread southern distribution in the past and only relict populations now exist where climatic conditions have remained favorable for its survival in the southwestern United States (Mathiasen 1976). Both of the southern Arizona populations of A. abietinum occur on mesic, north-facing slopes of narrow can- yons at approximately the same elevational range, and near the lower elevational limits of white fir in the Southwest. The occurrence of an extremely isolated population of the white fir true mistletoe, Phoradendron den- sum Torr. ex Trel. subsp. pauciflorum (Torr.) Wiens, parasitizing white fir in the Santa Catalina Mountains, may indicate this mis- tletoe had a more southern past distribution also. Phoradendron densum subsp. pauciflo- rum has not been found on white fir in other mountain ranges in Arizona, but it does occur in the Sierra San Pedro Martir in Baja Cali- fornia, which is farther south than the Santa Catalina populations (Wiens 1964). There- fore, it may be possible that both of these white fir parasites migrated to southern Ari- zona along a southern route when Baja Cali- fornia and the present mainland of Mexico were continuous (Hamilton 1961). However, A. abietinum has not been reported from Baja California, but may not have survived there for the same reasons it has not been re- ported from more localities in the south- western United States. The discovery of addi- tional relict populations of A. abietinum and P. densum subsp. pauciflorum in Mexico or in the mountain ranges with white fir popu- lations between the Grand Canyon and southern Arizona would provide more evi- dence concerning the past distribution of these taxa and their probable migration routes into the Southwest. In 1980, a small population of A. micro- carpum was discovered parasitizing Picea engelmannii in the Sacramento Mountains of Otero County, New Mexico (Fig. 2). This population is approximately 170 miles east and slightly south of the nearest known popu- lation of A. microcarpum in the Mogollon Mountains of west central New Mexico. Our surveys over the last two years indicate the infestation of A. microcarpum in the Sacra- mento Mountains is restricted to three small populations totaling about 300 acres occur- ring on northern exposures in the general vi- cinity of upper Hay Canyon (T. 17 S., R. 12 E., Sections 33-34; T. 18 S. R. 12 E., Sections October 1983 Mathiasen, Jones: Dwarf Mistletoes 743 Fig. 2. Distribution of Picea engehnannii and Arceuthohium microcarpum (1-6) in Arizona and New Mexico. Numbers 5 and 6 represent the A. microcarpum populations in the Pinaleno and Sacramento mountains, respective- ly. Number 7 represents the Chiricahua Mountain population oi Picea engehnannii var. mexicana. 3-4) at elevations ranging from 2,760 to 2,840 m. Our examinations of spruce stands in adjacent Spring, Hubbell, Wills, and Sac- ramento canyons have not detected addition- al populations of A. microcarpum. These can- yons are similar in elevation and topography to Hay Canyon, and their north-facing slopes are dominated by a geographically restricted mixed conifer habitat type {Picea engel- mannii/Acer glabrum HT) described by Moir and Ludwig (1979) in which Engelmann spruce and Douglas fir are co-dominant over- story species. In addition, all the areas sur- veyed in the Sacramento Mountains are well below the upper elevational limit of 3,100 m reported for A. microcarpum (Acciavatti and Weiss 1974, Hawksworth and Wiens 1972, Mathiasen and Hawksworth 1980). There- fore, the restriction of A. microcarpum to this small an area is anomalous because condi- tions for its spread into adjacent spruce stands appear to be favorable. The apparent confinement of this parasite to Hay Canyon may indicate it has arrived relatively recently in the Sacramento Mountains and did not ac- company P. engelmannii at the time of this tree's migration into this range. Certainly if A. microcarpum had migrated there with P. engelmannii, additional populations should have survived elsewhere in the Sacramento Mountains and in the other mountain ranges with large P. engelmannii populations in southwestern New Mexico (Little 1971) (Fig. 2). Therefore, the possibility of a more recent introduction of A. microcarpum into the Sac- ramento Mountains would appear more plau- sible than the relict population hypothesis, which seems more applicable to the disjunct populations of A. ahietinum in southern Arizona. A more recent introduction of A. micro- carpum to the Sacramento Mountains may then be an example of long-range dis- semination of a dwarf mistletoe by an avian vector. Hawksworth and Wiens (1972) cite examples of dwarf mistletoe distributions that might be best explained by seed dis- semination by avian vectors, and certain in- festation patterns of dwarf mistletoes in con- ifer stands are best explained by this means also (Hudler et al. 1979). One factor that must be considered when assessing the possi- bilities of long-range dissemination of dwarf mistletoes by birds is that Arceuthohium spp. are dioecious, and a male and female plant must become established in an isolated area for a successful infestation to occur. How ever, investigations of small satellite dwarf 744 Great Basin Naturalist Vol. 43, No. 4 mistletoe infection centers several meters from larger infestations have demonstrated that both male and female plants can become independently established in satellite centers (Hudler et al. 1979, Ostry 1978). In addition, Hudler et al. (1979) reported that they be- lieve birds are the most likely agents of long- distance dispersal of dwarf mistletoes. There- fore, the possibility of the establishment of the Sacramento Mountains population of A. microcarpum by avian vectors should be considered. The occurrence of A. microcarpttin in southern New Mexico may be of interest in relation to recent studies of the taxonomic relationships of Picea spp. in the south- western United States and northern Mexico. Engelmann spruce populations in the Sacra- mento Mountains (32° 48' N), the Pinaleno Mountains (32° 30' N), and the Chiricahua Mountains (31° 32' N) have been considered as the three most southern populations of this species in the United States (Little 1950, 1971, Daubenmire 1972, Taylor et al. 1975). However, Taylor and Patterson (1980) have shown that the P. engelmannii population from the Chiricahua Mountains differs slightly morphologically and chemically from more northern populations of this taxon and that the recently described Picea mexicana Martinez (Martinez 1961) is in their opinion not sufficiently different from P. engelmannii to warrant separation at the specific level. Therefore, they have reduced P. mexicana to a variety of P. engelmannii (P. engelmannii Parry var. mexicana (Martinez) Taylor & Pat- terson) and have included the Chiricahua Moimtains population as representative of this combination (Fig. 2). Although Taylor and Patterson (1980) did not include samples of the P. engelmannii population from the Sacramento Mountains in their study, they did sample the P. engelmannii population in the Pinaleno Mountains and concluded it was representative of P. engelmannii var. engel- mannii. We consider the spruce populations in the Hay Canyon vicinity to be morpho- logically representative of P. engelmannii var. engelmannii also (Daubenmire 1972, Jones 1977). In addition, the occurrence of A. microcarpum in the Sacramento Mountains may have taxonomic significance regarding the classification of this spruce population. Arceuthobium microcarpum has not been re- ported on spruce in the Chiricahua Moun- tains (Hawksworth and Wiens 1972), but it is present on P. engelmannii in the Pinaleno Mountains, only 60 miles to the northwest, and is here reported on P. engelmannii from the Sacramento Mountains. Because dwarf mistletoes are relatively host specific para- sites, the absence of A. microcarpum from the Chiricahua Mountains may be the result of the close phylogenetic affinities of that spruce population to the Mexican popu- lations of Picea reported by Taylor and Pat- terson (1980). Hawksworth and Wiens (1972) reported that dwarf mistletoes have not been found parasitizing Picea chihuahuana Marti- nez in northern Mexico and there have been no reports of dwarf mistletoes on the Mexi- can populations of P. engelmannii var. mexi- cana (Hawksworth, F. C, pers. comm., 1982). Therefore, these taxa of Picea may have diverged from their northern relatives to the extent that they are less susceptible or immune to parasitism by extant species of Ar- ceuthobium including A. microcarpum. If this is the case, then parasitism of the spruce pop- ulations in the Sacramento and Pinaleno mountains would help demonstrate their phy- letic affinities to other P. engelmannii popu- lations in the Southwest. The apparent ab- sence of A. microcarpum from the Chiricahua Mountains does not indicate conclusively that P. engelmannii var. mexicana is less suscep- tible or immune to this mistletoe. Most vari- etal taxa of principal hosts of dwarf mis- tletoes are susceptible to parasitism when they occur within the mistletoes' geographic range (Hawksworth and Wiens 1972), and the absence of A. microcarpum from the Chiricahua Mountains may simply be a result of its geographic isolation from the spruce population there. Hawksworth and Wiens (1972) suggested that A. microcarpum may have had a more northern distribution in the past, but has be- come isolated as relict endemic populations near the southern limits of its principal host ranges. They hypothesized that A. micro- carpum may have evolved in a more northern area (possibly from the ancestor of A. laricis or A. tsugense since these taxa resemble A. microcarpum morphologically, our addition), migrated southward, and become isolated in October 1983 Mathiasen, Jones: Dwarf Mistletoes 745 the Southwest. Recent studies of the dwarf mistletoe population parasitizing bristlecone pine, Pinus aristata Engelm., as a principal host on the San Francisco Peaks in north cen- tral Arizona have shown this mistletoe is A. microcarpum (Crawford and Hawksworth 1979, Mathiasen and Hawksworth 1980). However, this population differs morphologi- cally and physiologically from other A. mi- crocarpum populations (Mathiasen and Hawksworth 1980), and rarely parasitizes Pinus strobiformis and Abies lasiocarpa var. arizonica (Merriam) Lemm. (Hawksworth and Wiens 1972, Mathiasen and Hawksworth 1980). Therefore, A. microcarpum may have evolved in the Southwest from an ancestral species with a broader host range, including species of Pinus and Picea. The A. micro- carpum population parasitizing Pinus aristata on the San Francisco Peaks would then be an extant population with close phyletic affi- nities to this hypothesized ancestral species and may indicate a relatively recent evolu- tion of A. microcarpum with its radiation and specialization onto P. engelmannii and P. pungens in the southwestern United States. A more recent evolutionary origin for A. micro- carpum would mean this species may still be migrating into areas where spruce popu- lations are available for colonization. This may better explain its absence from the ex- tensive spruce-fir forests of the central and northern Rocky Mountains than the hypoth- esis of a more northern evolutionary origin for A. microcarpum. It seems probable that A. microcarpum is not restricted to its present geographic range by climatic factors or host susceptibility, but that it evolved and special- ized on Picea shortly before its potential host populations became isolated into small scat- tered populations in the higher mountain ranges of the Southwest. At present its fur- ther migration is severely hampered by its short dissemination range and hence its in- ability to spread over the long distances nec- essary for its migration into the central Rocky Mountains or into Mexico. However, the Sacramento Mountains population of A. microcarpum may indicate that this parasite can occasionally make an extremely large ex- tension from its otherwise limited distribu- tion. If it is occasionally possible for A. mi- crocarpum to make a large jump, by whatever means of dissemination, then, given enough time, A. microcarpum may eventually spread into the more northern and southern spruce populations of North America. Literature Cited AcciAVATTi, R. E., AND M. J. Weiss. 1974. Evaluation of dwarf mistletoe on Engelmann spruce, Fort Apache Indian Reservation, Arizona. Plant. Dis. Reptr. 58:418-419. Daubenmire, R. 1972. On the relation between Picea pungens and Picea engelmannii in the Rocky Mountains. Canadian J. Rot. 50:733-742. Hamilton, W. 1961. Origin of the Gulf of California. Geol. Soc. Amer. Bull. 72:1307-1318. Hawksworth, F. G., and D. P. Graham. 1963. Dwarf mistletoes on spruce in the Western United States. Northwest Sci. 37:31-38. Hawksworth, F. G., and D. Wiens. 1972. Biology and classification of dwarf mistletoes (Arceuthobium). U.S. Department of Agriculture, Forest Service, Agricultural Handbook 401. 234 pp. Hudler, G. W., N. Oshima, and F. G. Hawksworth. 1979. Bird dissemination of dwarf mistletoe on ponderosa pine in Colorado. Amer. Midi. Nat. 102:27.3-280. Jones, J. R. 1977. How to tell Engelmann from blue spruce in the Southwest. U.S. Department of Ag- riculture, Forest Service, Gen. Tech. Rept. RM- 34. 10 pp. Little, E. L. 1950. Southwestern trees. U.S. Depart- ment of Agriculture, Forest Service, Agricultural Handbook 9. 109 pp. 1971. Atlas of United States trees. Vol. 1. Con- ifers and important hardwoods. U.S. Department of Agriculture, Forest Service, Misc. Pub. 1146. 9 pp. + illus. Martinez, M. 1961. Una nueva especie de Picea en Mexico. Anal, del In.st. de Biol. .32:137-142. Mathiasen, R. L. 1976. Southern range extension of the white fir dwarf mistletoe in Arizona. Great Basin Nat. 36:461-462. Mathiasen, R. L., and F. G. Hawksworth. 1980. Tax- onomy and effects of dwarf mistletoe on bristle- cone pine on the San Francisco Peaks, Arizona. U.S. Department of Agriculture, Forest Service, Res. Pap. RM-224. 10 pp. MoiR, W. H., and J. A. Ludwig. 1979. A classification of spruce-fir and mixed conifer habitat types of Ari- zona and New Mexico. U.S. Department of Agri- culture, Forest Service, Res. Pap. RM-207. 47 pp. OsTRY, M. E. 1978. Vectors of eastern dwarf mistletoe. Unpubli.shed thesis. Univ. of Minnesota. 141 pp. ScHARPF, R. F. 1964. Dwarf mistletoe on true firs in Cal- ifornia. U.S. Department of Agriculture, Forest Service, For. Pest Leafl. 89. 7 pp. ScHARPF, R. F., and J. R. Parmeter. 1967. The biology and pathology of dwarf mistletoe, Arceuthobium campylopodum f. abietinum, parasitizing true firs {Abies spp.) in California, U.S. Department of Agriculture, Forest Service, Tech. Bull. 1362. 42 pp. 746 Great Basin Naturalist Vol. 43, No. 4 Taylor, R. J., S. Williams, and R. Daubenmire. 1975. Taylor, R. J., and T. F. Patterson. 1980. Biosystemat- Interspecific relationships and the question of in- ics of Mexican spruce species and populations, trogression between Picea engelmannii and Picea Taxon 29:421-440. ptingens. Canadian J. Bot. 53:2547-2555. Wiens, D. 1964. Revisions of the acataphyllous species oi Phoradendron. Brittonia 16:11-54. FIRST NEST RECORDS FOR THE PLAIN TITMOUSE AND BLUE-GRAY GNATCATCHER IN WYOMING Scott L. Findholf Abstract.— Nests of Plain Titmice and Blue-gray Gnatcatchers are reported from Wyoming for the first time. Both species probably breed fairly commonly in southwestern and south central portions of the state. In Wyoming the Plain Titmouse {Parus in- omatus) and Blue-gray Gnatcatcher (Poliop- tila caerulea) are considered uncommon, per- ipheral species and occur primarily in southwestern and south central portions of the state (Oakleaf et al. 1982). Nest records have not been previously reported for either species. The first confirmed nesting record of the Plain Titmouse in Wyoming occurred on 4 Jime 1981 when S. D. Fitton located a nest near Powder Rim, about 113 km southwest of Rawlins, Sweetwater Co. Evidence of a nest included the presence of two adult titmice that were observed entering and leaving a nest cavity several times. The nest cavity was not examined for the presence of eggs or young. During 1982 four nests of this species were found in Wyoming. The first nest was located by S. D. Fitton on 17 May near Little Fire- hole Canyon, 13 km southeast of Green Riv- er, Sweetwater Co. On 29 May a second nest was discovered by S. D. Fitton south of Pow- der Rim, about 22 km southeast of the 1981 nesting locale and approximately 1 km north of Colorado. Both nests contained an unde- termined nimiber of young that were audible from the nest cavities. Also, adult titmice were observed defending the nest sites. Six days later I located a Plain Titmouse nest containing two recently hatched nestlings in Firehole Canyon, 18 km southeast of Green River, Sweetwater Co. On the following day I found another nest containing an undeter- mined number of nestlings near Minnies Gap. approximately 55 km south of Green River, Sweetwater Co., and less than 1 km from Utah. All nests found during 1981 and 1982 were in cavities of Utah juniper (Juniperus osteosperma) trees. Several family groups containing recently fledged young titmice were also located in southwestern and south central Wyoming. This indicates that Plain Titmice probably breed rather commonly throughout these portions of the state. The first possible breeding evidence for the Blue-gray Gnatcatcher in Wyoming was re- ported by White and Behle (1960). On 4 July 1959 a family group of gnatcatchers were ob- served at mile 371 on the Green River, Sweetwater Co. Also, one adult and one juve- nile were collected at the same locale. Addi- tional details of this possible breeding record are lacking. The first nest of the Blue-gray Gnatcatcher in Wyoming was not located until 10 June 1982, when I observed a pair of gnatcatchers constructing a nest in Firehole Canyon, 18 km southeast of Green River, Sweetwater Co. The nest was located in a Utah juniper tree about 6 m from the ground. When the nest was rechecked 10 days later, it contained six eggs. Since the nest site was not revisited, I am uncertain how many eggs hatched or how many young fledged. On 28 June 1982 S. D. Fitton found the second Wyoming nest near Powder Rim, about 113 km southwest of Rawlins, Sweet- water Co. Three nestlings were present in the nest and about ready to fledge. This nest was also in a Utah juniper tree. Although addi- tional nests or family groups containing 'Wyoming Natural Heritage Program, The Nature Conservancy, 1603 Capitol Avenue, Room 325, Cheyenne, Wyoming 82001. Present address: Wyo- ming State Training School, Lander, Wyoming 82520. 747 748 Great Basin Naturalist Vol. 43, No. 4 recently fledged young were not found, sev- eral pairs of Blue-gray Gnatcatchers were ob- served in southwestern and south central Wyoming and suspected of breeding. This suggests that the Blue-gray Gnatcatcher may also breed fairly commonly in these portions of the state. The breeding habitat of the Plain Titmouse and Blue-gray Gnatcatcher consists mainly of Utah juniper woodlands interspersed with open areas containing big sagebrush {Arte- misia tridentata), other shrubs, and grasses. Both the Plain Titmouse and Blue-gray Gnatcatcher also breed in adjacent areas of northeastern Utah (Behle 1981), northwestern Colorado (Kingery and Graul 1978), and southeastern Idaho (Burleigh 1972). The lack of observers in southwestern and south cen- tral Wyoming has probably accounted for the paucity of nesting records of both species. I thank S. D. Fitton for allowing me to use his observations in this paper. Literature Cited Behle, W. H. 1981. The birds of northeastern Utah. Utah Museum of Nat. Hist. Occas. Pub. 2, Univ. of Utah, Salt Lake City. 136 pp. Burleigh, T. D. 1972. Birds of Idaho. Caxton Printers, Ltd., Caldwell, Idaho. 467 pp. Kingery, H. E., and W. D. Graul. 1978. Colorado bird distribution iatiiong study. Colorado Div. Wild!., Denver, Colorado. 58 pp. Oakleaf, B., H. Downing, B. Raynes, M. Raynes, and O. K. Scott. 1982. Wyoming avian atlas. Game and Fish Dept., Cheyenne, Wyoming. 87 pp. White, C. M., and W. H. Behle. 1960. Birds of Flam- ing Gorge reservoir basin. Pages 185-208 in Eco- logical studies of the flora and fauna of Flaming Gorge reservoir basin, Utah and Wyoming. Univ. of Utah. Dept. of Anthropology, Anthropological Paper 48, Salt Lake City, Utah.' A NEW SPECIES OF THELESPERMA (ASTERACEAE) FROM WYOMING Robert D. Dorn' Abstract.— TJielesperma ptibescens Dorn is described as new to science from Uinta County, Wyoming. In southwest Wyoming, I encountered what appeared to be Thelesperma margina- tum Rydb. but the leaves were conspicuously pubescent. This was very unusual since spe- cies of Thelesperma in this region are all glabrous or nearly so. Further study revealed that these plants also had .a different caudex from T. marginatum. These major differences support specific status for these plants. Thelesperma pubescens Dorn, sp. nov. Perennis; radicibus crassis lignosis; caudi- cibus ramosis; caulibus glabris 3-12 cm altis; foliis basalibus plerumque pinnatidivisis pu- bescentibus; capitulis 1 vel 2; involucris 5-9 mm altis; ligulis nuUis; disci corollis luteis ca 5 mm longis; pappi nullis; achaeniis glabris ca 4 mm longis (Fig. 1). Perennial from a thick woody taproot and branched caudex that bears a dense series of persistent old leaf bases (absent in very young plants); stems 3-12 cm high, glabrous, mostly leafless; leaves mostly basal, 1-5 cm long, pinnately divided into mostly 3-5, usually linear segments, rarely simple, conspicuously pubescent; heads 1 or rarely 2; involucre 5-9 mm high, inner bracts much broader and longer than outer and with broad scarious margins; ray flowers lacking; disk corollas yellow, about 5 mm long, dilated at base, with reddish brown longitudinal veins that split at each corolla sinus and pair and meet with an adjacent vein at tip of corolla lobes; pappus lacking; achenes glabrous, angled, about 4 mm long, each subtended by a longer membranous bract. Type.— USA. Wyoming: Uinta Co., Hickey Mountain, T13N, R114W, S13 E1/2, 8,400 ft, rocky ridge, 20 August 1982, Dorn 3823 (Holotype RM; Isotypes to be distributed). Other specimens.— Same location as holotype, 8 July 1982, Dorn 3752 (RM); Wyoming: Uinta Co., Sage Creek Mountain, T14N, R113W, S34 NEV4, 8,200 ft, rocky outcrop, 30 July 1979, Aldrich 592 (RM). Thelesperma pubescens differs from T. marginatum in having conspicuously pu- bescent leaves and a thicker, more branched caudex with a dense series of persistent old leaf bases. The latter characteristic is com- mon in desert plants. The caudex in T. mar- ginatum is generally simple or few-branched, slender, somewhat rhizomatous, and lacks the persistent old leaf bases. Another related spe- cies, T. subnudum Gray, is similar to T. mar- ginatum except ray flowers are usually pres- ent. The two are often considered varieties of the same species. These species are quite scattered in their distribution so it is difficult to say from which of these two T. pubescens was derived. The location of T. pubescens is between the distributions of the other two species. Thelesperma marginatum is found to the north, and T. subnudum is found to the south. It is also possible, but unlikely, that these two species were both derived from T. pubescens. 'Box 1471. Cheyenne, Wyoming 82003. 749 750 Great Basin Naturalist Vol. 43, No. 4 Fig. 1. nelespemia pubescens Dorn: lower center, entire plant, scale bar = 1 cm. Upper left, flower, scale bar 2 mm. Upper right, leaf tip, scale bar = 1 mm. INDEX TO VOLUME 43 The genera and species described as new to science in this volume appear in bold type in this index. A bouquet of daisies {Erigeron, Compositae), p. 365. A bibliography of Colorado vegetation description, p. 45. A comparative study of coyote food habits on two Utah deer herds, p. 432. A list of Utah spiders, with their localities, p. 494. A mammalian humerus from the Upper Jurassic of Colorado, p. 551. A new species of Penstemon (Scrophulariaceae) from the Uinta Basin, Utah, p. 429. A new species of Thelespemia (Asteraceae) from Wyoming, p. 749. A re-evaluation of the postglacial vegetation of the Laramie Basin, Wyoming-Colorado, p. 377. A review of the genus Soliperla (Plecoptera: Peltoperlidae), p. 30. A revision of the genus Microrhopala (Coleoptera: Chrysomelidae) in America north of Mexico, p. 597. A small carnivore survey technique, p. 438. A vascular flora of the San Rafael Swell, Utah, p. 79. Aculeata Hymenoptera of Sand Mountain and Blow Sand Mountains, Nevada, p. 403. Adkins, Betty S., Deborah L. EUiott-Fisk, and Jeanine L. Spaulding, article by, p. 377. Agropyron arizonicum (Gramineae: Triticeae) and a natural hybrid from Arizona, p. 13L Allred, Dorald M., and B. J. Kaston, article by, p. 494. Alpine and subalpine wetland plant communities of the Uinta Mountains, Utah, p. 523. Andersen, Perron L., John R. Crellin, Craig R. Nichols, and Peter M. Schantz, article by, p. 65. Anderson, Loran C, article by, p. 358. Applicability of the Universal Soil Loss Equation for southeastern Idaho wildlands, p. 579. Atkins, Riley J., Mary E. Barkworth, and Douglas R. Dewey, article by, p. 561. Baker, William L., articles by, p. 45, 687. Bakewell, George, Joseph M. Chopek, and Gary L. Burkholder, article by, p. 477. Barkworth, Mary E., Douglas R. Dewey, and Riley J. Atkins, article by, p. 56L Bats of the Colorado oil shale region, p. 554. Baugh, Thomas M., and James E. Deacon, article by, p. 592. Baumann, Richard W., and Gerald Z. Jacobi, article by, p. 585. Bayn, Robert L., Jr., and David J. Schimpf, article by, p. 573. Bechtel, R. C, R. W. Rust, and L. M. Hanks, article by, p. 403. Bird distributional and breeding records for southeastern Idaho, Utah, and adjacent regions, p. 717. Birds of southwestern Idaho, p. 728. Black, Hal L., article by, p. 456. Bond, Carl E., and Jack E. Williams, article by, p. 409. Briggs, George M., and James A. MacMahon, article by, p. 523. Brotherson, Jack D., article by, p. 137. Brotherson, Jack D., and Samuel R. Rushforth, article by, p. 73. Buchanan, Hayle, and Robert A. Graybosch, article by, p. 701. Burkholder, Gary L., George Bakewell, and Joseph M. Chopek, article by, p. 477. Cactopinus atkinsoni, p. 651. Cactopinus burjosi p. 651. Cactopinus granulatus, p. 651. Cactopinus setosus, p. 652. Caire, William, Robert B. Finley, Jr., and Dallas E. Wilhelm, article by, p. 554. Campbell, Thomas M. Ill, and Tim W. Clark, article by, p. 438. 751 752 Great Basin Naturalist Vol. 43, No. 4 Campos, E. G., and R. B. Eads, article by, p. 168. Carphobius pilifer, p. 652. ChaetopJiloeus confinis, p. 652. Chopek, Joseph M., George Bakewell, and Gary L. Burkholder, article by, p. 477. Chramesus exilis, p. 653. Chramesus exul, p. 653. Chramesus securus, p. 653. Chramesus tibialis, p. 654. Clark, Shawn M., article by, p. 597. Clark, Tim W., and Thomas M. Campbell III, article by, p. 438. Cnemonyx equihuai, p. 654. Cnemonyx evidens, p. 654. Cnesinus cornutus, p. 655. Cnesinus nebulosus, p. 655. Cnesinus parvicornis, p. 656. Coelidia cochloea, p. 672. CoeUdia panamensis, p. 671. CoeUdia retrorsa, p. 670. Coelidia simplex, p. 673. Coelidia tortula, p. 672. Collins, Patrick D., Kimball T. Harper, and Burton K. Pendleton, article by, p. 385. Comparative life history and floral characteristics of desert and mountain floras in Utah, p. 385. Comparative successional roles of trembling aspen and lodgepole pine in the Southern Rocky Mountains, p. 447. Crellin, John R., Ferron L. Andersen, Craig R. Nichols, and Peter M. Schantz, article by, p. 65. Daily and yearly movement of the Devil's Hole pupfish Cyprinodon diabolis Wales in Devil's Hole, Nevada, p. 592. Deacon, James E., and Thomas M. Baugh, article by, p. 592. Deer mouse, Peromysciis maniculatus, and associated rodent fleas (Siphonaptera) in the arctic-alpine life zone of Rocky Mountain National Park, Colorado, p. 168. Dendrosinus mexicanus, p. 656. Despain, Del W., and Grant A. Harris, article by, p. 421. Dewey, Douglas R., Mary E. Barkworth, and Riley J. Atkins, article by, p. 561. Differential utilization of bat boxes by house wrens {Troglodytes aedon), p. 456. Dom, Robert D., article by, p. 749. Dom, Robert D., Robert W. Lichvar, and Erwin F. Evert, article by, p. 739. Eads, R. B., and E. G. Campos, article by, p. 168. Elliott-Fisk, Deborah L., Betty S. Adkins, and Jeanine L. Spaulding, article by, p. 377. Erigeron awapensis, p. 365. Erigeron canaani, p. 366. Erigeron carringtonae, p. 366. Erigeron goodrichii, p. 366. Erigeron maguirei var. harrisonii, p. 367. Erigeron untermannii, p. 367. Erigeron wahwahensis, p. 368. Evaluation of a program to control hydatid disease in central Utah, p. 65. Evaluation of Draba oligosperma, D. pectinipila, and D. juniperina complex (Cruciferae), p. 441. Evaluation of varieties in Stanleya pinnata (Cruciferae), p. 684. Evert, Erwin F., Robert W. Lichvar, and Robert D. Dorn, article by, p. 739. Eye fluke {Diplostomum spathaceum) of fishes from the upper Salmon River near Obsidian, Idaho, p. 675. Findholt, Scott L., article by, p. 747. Finley, Robert B., Jr., William Caire, and Dallas E. Wilhelm, article by, p. 554. First nest records for the Plain Titmouse and Blue-gray Gnatcatcher in Wyoming, p. 747. Flora of the Lower Cretaceous Cedar Mountain Formation of Utah and Colorado, part I. Paraphyllanthoxylon titahense, p. 394. Flora of the Stansbury Mountains, Utah, p. 619. Floristics of the upper Walker River, California and Nevada, p. 93. Food of larval Tui chubs, Gila bicolor, in Pyramid Lake, Nevada, p. 175. Frost, Herbert H., Clayton M. White, Dennis L. Shirley, G. Merrill Webb, and Richard D. Porter, article by, p. 717. Galat, David L., and Nancy Vucinich, article by, p. 175. Goodrich, Sherel, article by, p. 531. Goodrich, Sherel, and Stanley L. Welsh, article by, p. 375. Graybosch, Robert A., and Hayle Buchanan, article by, p. 701. Yanks, L. M., R. W. Rust, and R. C. Bechtel, article by, p. 403. Hansen, Richard M., James G. MacCracken, and Daniel W. Uresk, article by, p. 660. October 1983 Index 753 Haplopappus armerioides var. gramineus, p. 371. Haplopappus crispus, p. 359. Haplopappus crispus and H. zionis (Asteraceae): new species from Utah, p. 358. Harper, Kimball T., Patrick D. Collins, and Burton K. Pendleton, article by, p. 385. Harris, Grant A., and Del W. Despain, article by, p. 421. Harris, James G., article by, p. 79. Hart, E. Blake, and Michael Trumbo, article by, p. 492. Heckmann, Richard, article by, p. 675. Helm, William T., William F. Sigler, Paul A. Kucera, Steven Vigg, and Gar W. Workman, article by, p. 1. Holomuzki, Joseph R., article by, p. 475. Hy7nenoxys lapidicola, p. 373. Influence of cryptogamic crusts on moisture relationships of soils in Navajo National Monument, Arizona, p. 73. Jacobi, Gerald Z., and Richard W. Baumann, article by, p. 585. Jensen, James A., and Donald R. Prothero, article by, p. 551. Jensen, Mark E., article by, p. 579. Jones, Kenneth H., and Robert L. Mathiasen, article by, p. 741. Kaston, B. J., and Dorald M. Allred, article by, p. 494. Kennedy, Joseph L., article by, p. 713. Kramer Palouse natural area, p. 421. Kucera, Paul A., William F. Sigler, William T. Helm, Steven Vigg, and Gar W. Workman, article by, p. 1. Lavin, Matt, article by, p. 93. Lichvar, Robert W., articles by, p. 441, 684. Lichvar, Robert W., Robert D. Dom, and Erwin F. Evert, article by, p. 739. Life history of the Lahontan cutthroat trout, Salmo clarki henshawi, in Pyramid Lake, Nevada, p. 1. Liparthrum mexicanum, p. 657. Liparthrum pruni, p. 657. MacCracken, James G., Daniel W. Uresk, and Richard M. Hansen, article by, p. 660. MacMahon, James A., and George M. Briggs, article by, p. 523. Maser, Chris, and Ronald S. Rohweder, article by, p. 425. Mathiasen, Robert L., and Kenneth H. Jones, article by, p. 741. Microrhopala rileyi, p. 609. Milton, N. M., and T. L. Purdy, article by, p. 457. Neese, Elizabeth, and Stanley L. Welsh, articles by, p. 373, 429, 700. New generic concepts in the Triticeae of the intermountain region: keys and comments, p. 561. New Haplopappus variety in Utah (Compositae), p. 371. New leafliopper species of Coelidia with a revised key and notes on homonymy and distribution (Homoptera: Cicadellidae, Coelidiinae), p. 669. New records for the vascular flora of Wyoming and Montana, p. 739. New species of Hymenoxys and Perityle (Compositae) from Utah, p. 373. New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), part IX, p. 647. New taxa in Thelesperma and Townsendia (Compositae) from Utah, p. 369. New variety of Opuntia basilaris (Cactaceae) from Utah, p. 700. New variety of Stephanomeria tenuifolia (Compositae) from Utah, p. 375. Nichols, Craig R., Ferron L. Andersen, John R. Crellin, and Peter M. Schantz, article by, p. 65. Nielson, Mervin W., article by, p. 669. Notes on reproduction of the side-blotched lizard Uta stansburiana stansburiana in southwest Idaho, p. 477. Observations on alpine vegetation near Schoolroom Glacier, Teton Range, Wyoming, p. 483. Parker, Albert J., and Kathleen C. Parker, article by, p. 447. Parker, Kathleen C, and Albert J. Parker, article by, p. 447. Pederson, Jordan C, article by, p. 445. Pederson, Jordan C, and R. Gary Tuckfield, article by, p. 432. Pendleton, Burton K., Patrick D. Collins, and Kimball T. Harper, article by, p. 385. Penstemon flowersii, p. 429. Perityle specuicola, p. 373. Phloeotribus geminus, p. 657. Plant and soil relationships in two hydrothermally altered areas of the Great Basin, p. 457. 754 Great Basin Naturalist Vol. 43, No. 4 Plant community variability on a small area in southeastern Montana, p. 660. Plasticity and polymorphism in seed germination of Mimulus guttatus (Scrophulariaceae), p. 470. Porter, Richard D., Clayton M. White, Herbert H. Frost, Dennis L. Shirley, and G. Merrill Webb, article by, p. 717. Predatory behavior of larval Ambystoma tigrintim nebuloswn on Limnephilus (Trichoptera) larvae, p. 475. Presence of maxillary canine teeth in mule deer in Utah, p. 445. Pronghom responses to hunting coyotes, p. 88. Prothero, Donald R., and James A. Jensen, article by, p. 551. Purdy, T. L., and N. M. Milton, article by, p. 457. Pycnarthrum amersum, p. 658. Pyrah, Grant L., article by, p. 131. Range extensions for two dwarf mistletoes {Arceuthobiwn spp.) in the southwest, p. 741. Reproductive attributes of some Rocky Moimtain subalpine herbs in successional context, p. 573. Reynolds, Timothy D., article by, p. 88. Reynolds, Timothy D., and Daniel A. Stephens, article by, p. 728. Rohweder, Ronald S., and Chris Maser, article by, p. 425. Rushforth, Samuel R., and Jack D. Brotherson, article by, p. 73. Rust, R. W., L. M. Hanks, and R. C. Bechtel, article by, p. 403. Schantz, Peter M., Ferron L. Andersen, John R. Crellin, and Craig R. Nichols, article by, p. 65. Schimpf, David J., and Robert L. Bayn, Jr., article by, p. 573. Scolytodes plumericolens, p. 658. Scolytodes retifer, p. 658. Seasonal growth of the Tui chub, Gila bicolor, in Pyramid Lake, Nevada, p. 713. Shaw, Richard J., and John R. Spence, article by, p. 483. Shirley, Dennis L., Clayton M. White, Herbert H. Frost, G. Merrill Webb, and Richard D. Porter, article by, p. 717. Sigler, William F., William T. Helm, Paul A. Kucera, Steven Vigg, and Gar W. Workman, article by, p^ 1^ / jj [J (i t] Smith, Frank J., and Stanley L. Welsh, article by, p. 371. Soliperla sierra, p. 36. Soliperla tillamook, p. 41. Some aspects of the presettlement vegetation of the Piceance Basin, Colorado, p. 687. Spaulding, Jeanine L., Deborah L. Elliott- Fisk, and Betty S. Adkins, article by, p. 377. Species composition, distribution, and phytosociology of Kalsow Prairie, a mesic tall-grass prairie in Iowa, p. 137. Spence, John R., and Richard J. Shaw, article by, p. 483. Stark, Bill P., article by, p. 30. Status and life history notes on the native fishes of the Alvord Basin, Oregon and Nevada, p. 409. Stephanomeria tenuifolia var. uintahensis, p. 375. Stephens, Daniel A., and Timothy D. Reynolds, article by, p. 728. Stokes, W. L., G. F. Thayne, and W. D. Tidwell, article by, p. 394. Taye, Alan C, article by, p. 619. Thayne, G. F., W. D. Tidwell, and W. L. Stokes, article by, p. 394. Thelospemia pubescens, p. 749. Thelospertna subnudwn var. alpinum, p. 369. Tidwell, W. D., G. F. Thayne, and W. L. Stokes, article by, p. 394. Townsendia jonesii var. lutea, p. 369. Townsendia montana var. caelilinensis, p. 370. Trumbo, Michael, and E. Blake Hart, article by, p. 492. Tuckfield, R. Gary, and Jordan C. Pederson, article by, p. 432. Uresk, Daniel W., James G. MacCracken, and Richard M. Hansen, article by, p. 660. Utah flora: Compositae (Asteraceae), p. 179. Utah flora: Salicaceae, p. 531. Vegetative types and endemic plants of the Bryce Canyon Breaks, p. 701. Vickery, Robert K., Jr., article by, p. 470. Vigg, Steven, William F. Sigler, William T. Helm, Paul A. Kucera, and Gar W. Workman, article by, p. 1. Vucinich, Nancy, and David L. Galat, article by, p. 175. Webb, G. Merrill, Clayton M. White, Herbert H. Frost, Dennis L. Shirley, and Richard D. Porter, article by, p. 717. October 1983 Index 755 Welsh, Stanley L., articles by, p. 179, 365, 369. Welsh, Stanley L., and Elizabeth Neese, articles by, p. 373, 429, 700. Welsh, Stanley L., and Frank J. Smith, article by, p. 371. Welsh, Stanley L., and Sherel Goodrich, article by, p. 375. White, Clayton M., Herbert H. Frost, Dennis L. Shirley, G. Merrill Webb, and Richard D. Porter, article by, p. 717. Wilhelm, Dallas E., Robert B. Finley, and William Caire, article by, p. 554. Williams, Jack E., and Carl E. Bond, article by, p. 409. Winter food habits of cougars from northeastern Oregon, p. 425. Winter stomach contents of South Dakota badgers, p. 492. Winter stoneflies (Plecoptera) of New Mexico, p. 585. Wood, Stephen L., article by, p. 647. Workman, Gar W., William F. Sigler, William T. Helm, Paul A. Kucera, and Steven Vigg, article by, p. 1. The Great Basin Naturalist VOLUME 43, 1983 Editor: Stephen L. Wood Published at Brigham Young University, by Brigham Young University NOTICE TO CONTRIBUTORS Manuscripts intended for publication in the Great Basin Naturalist or Great Basin Natural- ist Memoirs must meet the criteria outUned in paragraph one on the inside front cover. They should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Natu- ralist, 290 Life Science Museum, Provo, Utah 84602. Three copies of the manuscript are re- quired. 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 Council of Biology Editors Style Manual, Fourth Edition (AIBS 1978) 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 article. All manuscripts receive a critical peer review by specialists in the subject area of the manu- script under consideration. Authors may recommend one or two reviewers for their article. Accepted manuscripts less than 100 printed pages in length will automatically be assigned to the Great Basin Naturalist. Those larger than 100 printed pages 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. The original illustrations or glossy photoprints of them should accompany the manuscript. Illustrations should be prepared for reduction by the printer to either single-column (2% inches) or double-column (5V^ inches) width, with the length not exceeding IV2 inches. Costs Borne by Contributor. Contributors to the Great Basin Naturalist should be prepared to donate from $10 to $40 per printed page toward publication of their article (in addition to reprint costs). Excessive or complex tables requiring typesetting may be charged to the author at cost. Authors publishing in the Great Basin Naturalist Memoirs may be expected to contrib- ute $40 per printed page in addition to the cost of the printed copies they purchase. No re- prints are furnished free of charge. Reprint Schedule for the Great Basin Naturalist 100 copies, minimum cost for 2 pages, $26. Each additional 2 pages, $6. Each additional 100 copies, $4 for each 2 pages. Examples: 300 copies of 10 pages = $82; 200 copies of 13 pages = $86. 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. No. 3 The endangered species: a symposium. $6. No. 4 Soil-plant-animal relationships bearing on revegetation and land reclamation in Nevada deserts. $6. No. 5 Utah Lake monograph. $8. No. 6 The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. $60. No. 7 Biology of desert rodents. $8. TABLE OF CONTENTS Alpine and subalpine wetland plant connnunities of the Uinta Mountains, Utah. George M. Briggs and James A. MacMahon 523 Utah flora: Salicaceae. Sherel Goodrich 531 A mammalian humerus from the Upper Jurassic of Colorado. Donald R. Prothero and James A. Jen.sen .' 551 Bats of the Colorado oil shale region. Robert B. Finlev, Jr., William Caire, and Dallas E. Wilhelm '. 554 New generic concepts in the Triticeae of the Intermountain Region: kev and comments. Mary E. Barkworth, Douglas R. Dewey, and Riley J. Atkins 561 Reproductive attributes of some Rocky Mountain subalpine herbs in successional context. David J. Schimpf and Robert L. Bayn, Jr 573 Applicability of the imiversal soil loss equation for southeastern Idaho wildlands. Mark E. Jensen 579 Winter stoneflies (Plecoptera) of New Mexico. Gerald Z. Jacobi and Richard W. Baumann 585 Daily and yearly movement of the Devil's Hole pupfish Cyprinodon diaboUs Wales in Devil's Hole, Nevada. Thomas M. Baugh and James E. Deacon 592 A revision of the genus Microrhoptda (Coleoptera: Chrysomelidae) in America north of Mexico. Shawn M. Clark 597 Flora of the Stan.sbury Mountains, Utah. Alan C. Taye 619 New .synonymy and new species of American bark beetles (Coleoptera: Scolytidae), part IX. Stephen L. Wood 647 Plant community variability on a small area in southeastern Montana. James G. MacCracken, Daniel W. Uresk, and Richard M. Hansen 660 New leafhopper species of Coelidia with a revised key and notes on homonymy and distribution (Homoptera: Cicadellidae, Coelidiinae). Mervin W. Nielson 669 Eye fluke (Diplostomum spadiaceum) of fishes from the upper Salmon River near Obsidian, Idaho. Richard Heckmann 675 Evaluation of varieties in Stanleija pinnata (Cruciferae). Robert W. Lichvar 684 Some aspects of the presettlement vegetation of the Piceance Basin, Colorado. William L. Baker 687 New variety of Opuntia basilaris (Cactaceae) from Utah. Stanley L. Welsh and Elizabeth Neese 700 Vegetative types and endemic plants of the Bryce Canyon Breaks. Robert A. Graybcsch and Hayle Buchanan 701 Seasonal growth of the Tui chub, Gila bicolor, in Pyramid Lake, Nevada. Joseph L. Kennedy 713 Bird distributional and breeding records for southeastern Idaho, Utali, and adjacent regions. Clayton M. White, Herbert H. Frost, Dennis L. Shirley, G. Merrill Webb, and Richard D. Porter .' 717 Birds of southwestern Idaho. Daniel A. Stephens and Timothy D. Reynolds 728 New records for the vascular flora of Wyoming and Montana. Robert W. Lichvar, Robert D. Dorn, and Erwin F. Evert 739 Range extensions for two dwarf mistletoes {Arceuthobitim spp.) in the southwest. Robert L. Mathiasen and Kenneth H. Jones 741 First nest records for the Plain Titmouse and Blue-gray Gnatcatcher in Wyoming. Scott L. Findholt '. 747 A new species of Thelesperina (Asteraceae) from Wyoming. Robert D. Dorn 749 Index 751 ACMP NOV b 1984 lOOCAMBRiDGt STREET CHARLESTOWISI, MASS. 3 2044 072 231 251